The effect of prey consumption on territorial defense by harriers: differential responses to neighbors versus floaters

Summary Food consumption may reduce fighting intensity of territory owners by decreasing resource value (additional food cannot be consumed) and/or increasing fighting costs (food in the digestive tract may raise injury risks). A territorial harrier's (Circus cyaneus, adult females) decision to reduce its level of aggression should depend upon whether or not the intruder was a competitor for individual prey items (as are smaller male floaters) or for the territory proper (as are female floaters and especially female neighbors). Accordingly, following meals, aggressive intensity of owners was strongly reduced towards male floaters (more were ignored), slightly reduced towards female floaters (more were called at than chased), and remained unchanged towards neighbors (virtually all were chased). Hence, alterations in aggressive behavior of territory owners following food consumption may depend upon the type of intruder and the resource under contest (a food item or a territory).     

The selective advantages of cannibalism in a Neotropical mosquito

We investigated the benefits of larval cannibalism in the Neotropical mosquito Trichoprosopon digitatum. The clutch size of the mosquito in the field was strongly correlated with adult female size, indicating a fitness advantage to being large. In controlled laboratory experiments, we compared the survivorship and eventual adult sizes of larvae that were given the opportunity to cannibalise conspecifics throughout their lifetimes with the survivorship and adult sizes of larvae that were prevented from cannibalising. Since the benefits of cannibalism are likely to depend on the context in which it occurs, the experiment was conducted at two levels of alternative food availability. When food availability was high most larvae survived to adulthood, females cannibalised more than males and there was no measurable advantage to cannibalism in terms of survival rate, emergence time or adult size. Larvae were significantly more cannibalistic when food availability was lower, although under these conditions no larvae survived to adulthood. Nevertheless, under low food an important fitness benefit to cannibalism was revealed: individuals which had the opportunity to cannibalise survived significantly longer as larvae than those which did not. This increased longevity is likely to provide an important advantage to mosquito larvae when they wait for the input of unpredictable food sources. Received: 7 October 1995/Accepted after revision: 13 April 1996     

Intrasexual competition,territoriality and acoustic communication in male strawberry poison frogs (<Emphasis Type="Italic">Oophaga pumilio</Emphasis>)

In many species male reproductive success is limited by access to females. Territoriality is one behavioural strategy which helps to acquire females. In the present study, we investigated the correlation between territory size and (1) female availability and (2) rate of intrusion by conspecific males in strawberry poison frogs, Oophaga pumilio. Males defended smaller territories in areas with a high female density and high rate of intrusion by conspecific males. Only males with high body condition values were able to establish territories in areas of high female density probably due to better fighting abilities. Moreover, dominant calling frequency was lower during agonistic interactions. Because only males with high body condition values were able to produce very low dominant frequencies, the acoustic properties appear to be an honest signal containing information about fighting abilities. Thus, the negative correlation between dominant frequency and mating success of males found in previous studies seems to be in part the result of intrasexual competition between males.     

Food availability and immune capacity in serin (Serinus serinus) nestlings

Parental feeding effort is an important determinant of chick development in birds. Quality and amount of food provided to each nestling, however, might affect the development of the immune system. Food availability around the nest site may account for part of the difference in body condition and immune capacity of nestlings. We tested this idea in the socially monogamous serin (Serinus serinus), investigating growth and immune capacity of nestlings in relation to food availability around the nest site. We used a set of immunological and serological assays, including antibody responses to sheep red blood cells (SRBCs), the T lymphocyte cell-mediated immune response (PHA), granulocyte:lymphocyte ratio (G:L), haematocrit and sedimentation rate. The results suggest that serins raised under poor environmental conditions were less immunocompetent. The immune response to PHA and SRBCs significantly increased with food availability. Haematocrit values were positively, sedimentation rate and the G:L ratio negatively correlated with food availability. Similar results were obtained when examining chick development in relation to the immunological and serological variables. However, there was a negative relationship between parental feeding rates and food availability, as well as between parental feeding rates and chick development. We conclude that body condition and immune capacity of nestlings are strongly correlated with the availability of food around the nest site.     

The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males

Summary Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m²) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.     

Effects of seasonality on the reproductive cycle of <Emphasis Type="Italic">Diadema</Emphasis> aff. <Emphasis Type="Italic">antillarum</Emphasis> in two contrasting habitats: implications for the establishment of a sea urchin fishery

Reproduction of Diadema aff. antillarum was examined between 2002 and 2005 at subtidal rocky bottoms around the Canary Islands. Two contrasting habitats (urchin barrens and grazing fronts) characterized by different levels of food availability were chosen, and factors thought to influence reproductive periodicity were monitored, including temperature, photoperiod, phytoplankton abundance and benthic food availability. Histological analyses showed that D. aff. antillarum had an annual reproductive cycle that was relatively synchronous across the studied sites and habitats. Photoperiod was the most significant factor that correlated with gonad periodicity; benthic food availability of 2 month lag was also correlated. However, some differences were detected between males and females in the timing of the onset of gametogenesis. Spawning was synchronized between both sexes from June to August. Results suggest that the optimum time of year to harvest urchin gonads would be between May and June when gonads were maximal in size but not full of gametes. This species may not provide optimal conditions for an industrial scale fishery, as sea urchins occurring in high density had small gonads and those producing larger volume or more marketable gonad tissue occurred in low densities where harvesting costs would exceed profit.     

Opposing selective pressures on hatching asynchrony: egg viability, brood reduction, and nestling growth

At least 19 hypotheses have been proposed to explain the evolutionary significance of avian hatching asynchrony, and hatching patterns have been suggested to be the result of several simultaneous selective pressures. Hatching asynchrony was experimentally modified in the black kite Milvus migrans by manipulating the onset of incubation during the laying period. Delayed onset of incubation reduced egg viability of first-laid eggs, especially when ambient temperature during the laying period was high. Brood reduction (nestling mortality by starvation or siblicide) was more commonly observed in asynchronous nests. The growth rate was slower in synchronous broods, probably due to stronger sibling rivalry in broods with high size symmetry. Last-hatched chicks in synchronous broods fledged at a small size/mass, while in control broods, hatching order affected growth rates, but not final size. Brood reduction, variable growth rates, and the ability to face long periods of food scarcity are probably mechanisms to adjust productivity to stochastic food availability in a highly opportunistic predator. The natural pattern of hatching asynchrony may be the consequence of opposing selective forces. Extreme hatching synchrony is associated with slow growth rates, small final size of last-hatched chicks, and low viability of first-laid eggs, while extreme hatching asynchrony is associated with high mortality rates. Females seem to facultatively manipulate the degree of hatching asynchrony according to those pressures, because hatching asynchrony of control clutches was positively correlated with temperature during laying, and negatively correlated with the rate of rabbit consumption. Received: 25 October 1999 / Revised: 30 May 2000 / Accepted: 25 June 2000     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Growth costs of territorial overlap: experiments with juvenile lizards (Anolis aeneus)

Summary Several theoretical models have suggested that increases in territory overlap with conspecifics should result in lower rates of resource accumulation by territorial residents. This assumption was tested using juvenile Anolis aeneus lizards, which exhibit variable degrees of territory overlap in both the field and laboratory. In a series of laboratory trials under controlled density and resource conditions, juvenile growth rates were negatively related to the degree of territory overlap. This effect aplied to juveniles of all but the lowest status groups, although higher status juveniles had more exclusive territories than did lower status juveniles. Focal studies indicated that food depletion in the zones of overlap was the most likely reason for the negative relationship between territory overlap and growth. It is suggested that juveniles in the field may tolerate extensive territory overlap in spite of growth penalties because of a general shortage of predator-safe homesites in undisturbed habitats.     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Gunnison’s prairie dog (<Emphasis Type="Italic">Cynomys gunnisoni</Emphasis>): testing the resource dispersion hypothesis

Few studies have experimentally tested the resource dispersion hypothesis (RDH). In this study, I tested whether space use and social organization of Gunnison’s prairie dog responded to changes in the dispersion and abundance of resources. Food manipulations were carried out during the reproductive and nonreproductive seasons across 2 years. Gunnison’s prairie dog adults responded to the experiments by decreasing territory size as food became patchier in space and time. Both males and females modified their home ranges, with no detectable difference between sexes, either prior to or during the experiments. As food became patchier in space and time, the spatial overlap of adults increased, whereas it decreased as food became more evenly dispersed. The average size of a group, defined as those individuals occupying the same territory, did not change significantly as a result of the experiments. Where changes in the composition and size of groups did occur, there was no indication that such changes were sex specific. Results from this study support critical components of the RDH and strongly suggest that patterns of space use and social structure in Gunnison’s prairie dogs are the result of individual responses to resource abundance and distribution.     

Correlation between nectar supply and aggression in territorial honeyeaters: causation or coincidence?

Summary A fundamental prediction of food-based economic models of territoriality is that animals will not defend territories if food is so abundant that defense will not improve access to food. Several studies of nectar-feeding birds support this prediction, with territoriality being rare or absent in years when nectar was particularly abundant. However, these results could potentially be an artefact of changes in bird density with nectar availability, and in at least some cases the correlations between territory defense and nectar availability could be purely coincidental. This paper reports the first experimental test of whether cessation of territory defense in nectar-feeding birds results from a direct response to abundance of nectar. New holland honeyeaters Phylidonyris novaehollandiae and white-cheeked honeyeaters P. nigra show pronounced changes in their levels of territorial aggression over the 7–8 months that they breed. These changes are predictable from economic considerations in that the birds are least aggressive in the months when nectar is extremely abundant. I tested whether the birds were responding to changes in nectar availability by providing sugar-water feeders at neutral locations that were easily accessible to territory holders, but far enough away from territories that intrusion rates were unaffected. I tested for responses at two time scales feeders were put out for 48-h periods in 1987, and were left out continuously from January to October 1988. The only effect was that territory holders visited feeders instead of flowers when floral nectar was scarce. They continued to defend their territories aggressively at those times, showed seasonal changes in aggressiveness similar to birds on a site without feeders, and did not shift their territories toward feeders. I conclude that the observed changes in aggressiveness are not responses to changes in nectar availability, and suggest alternative explanations.     

Territory quality and territorial behavior in two sympatric species of hummingbirds

Summary Changes in territorial behavior of blackchinned hummingbirds (Archilochus alexandri) in response to experimental changes in territory quality were investigated using artificial feeders and simultaneous, pair-wise observations of owners. Some of the responses of A. alexandri were similar to those documented by a previous study of the Anna's hummingbird (Calypte anna): A. alexandri defending rich territories spent more time on their territories, encountered a greater percentage of intruders, and chased more intruders than did A. alexandri defending poor territories. In contrast to C. anna, A. alexandri supplemented chases with energetically inexpensive threat vocalizations more extensively when territory quality was increased. This difference may be related to A. alexandri's more tenuous control of rich territories. When both species were observed on very poor territories, A. alexandri chased a greater percentage of intruders, consumed a greater proportion of available food, and obtained a greater net energy gain from their territories. When observed simultaneously on territories with ad lib food, both species defended highly exclusive territories but A. alexandri suffered higher intrusion pressure and spent more time and energy in defense. These interspecific differences in territorial behavior may facilitate sympatric coexistence of the two species through aggressive partitioning of flower patches according to productivity: the greater payoff received by C. anna on rich territories and A. alexandri on poor territories should favor a corresponding monopolization of rich areas by C. anna and poor areas by A. alexandri.     

Influence of seasonal variation in temperature, salinity and food availability on module size and colony growth of the estuarine bryozoan Conopeum seurati

Zooid size and colony growth of the estuarine bryozoan Conopeum seurati (Canu) (order: Cheilostomatida; suborder: Malacostegina) were examined over 15 mo at Avonmouth Dock, Avon, England. Data were analysed in conjunction with synchronous measurements of temperature, salinity and food availability. Zooid length, width and area were strongly temperature-dependent, while both food availability and colony growth rate had no significant effect on zooid length, width or area. Salinity and the interaction of temperature and salinity significantly influenced zooid length and area, suggesting that changes in zooid size may result from oxygen limitation in warm waters. The validity of a number of other mechanisms proposed to account for temperature-related changes in zooid size is discussed. The results support the use of zooid size as an indicator of both long-term trends and seasonal variations in temperature in Recent and fossil assemblages as long as data sets are large and the effects of other factors on zooid size are considered. Colony growth rate was found to be significantly influenced by both the amount of food available to the colonies and the combined effect of temperature and food availability, suggesting that growth rate increases as food increases, but that the former may be limited at low temperatures when metabolic rates are low. Received: 30 March 1999 / Accepted: 24 September 1999     

Do kin always make better neighbours?: The effects of territory quality

Summary The effects of territory quality on kin-biased territorial defence behaviour and fitness of juvenile rainbow trout (Oncorhynchus mykiss) were examined by manipulating food and predation risk levels in an artificial stream channel. Groups of related (full sibling) or unrelated fry were observed in the channel under one of four treatments: (1) high food-low predator (high territory quality); (2) high food-high predator; (3) low food-low predator; and (4) low food-high predator (low territory quality). Fish within kin groups always initiated fewer aggressive interactions, defended smaller territories, exhibited higher proportions of threat type territorial defence behaviours (as opposed to overtly aggressive behaviours) and had a higher mean weight increase than non-kin groups. Within both kin and non-kin groups, decreasing territory quality significantly increased the frequency of aggressive interactions, the size of territories and decreased growth. It is argued that while kin-biased territorial defence behaviours are always present, decreased availability of resources and increased predation risk serve to reduce the magnitude of this kin bias. However, even in low-quality territories, the benefit of associating with kin may increase the probability of overwintering survival (increased fitness) of the young fish by increasing body size.     

Relationships between spacing behavior and growth rates: A field study of lizard feeding territories

Summary Relationships between spacing behavior and growth rates were investigated in a field experiment with juvenile lizards, Anolis aeneus. The behavioral variable most closely related to juvenile growth was distance moved per unit time. This variable had a curvilinear relationship with growth, such that juveniles moving approximately 400 cm/h grew more rapidly than those traveling either larger or shorter distances per unit time. Daily fluctuations in arthropod abundance were also related to growth rates, with restricted growth during periods of low food availability. Temporal fluctuations in prey and distance traveled per unit time had independent effects on growth; together these two variables accounted for 43% of the variance in growth rate for the juveniles in this study.Territory size, overlap and social status appeared to affect growth indirectly, by influencing distance traveled per unit time. Optimal travel distances of around 400 cm/h were most likely when a juvenile had a relatively exclusive territory of about 0.5 m². High ranking juveniles were more apt to achieve this spacing pattern than were low ranking juveniles, but some high ranking juveniles had very large territories, extensive overlap with subordinates, supraoptimal travel distances and relatively low growth rates. Low ranking juveniles tended to fall into two groups: subordinates, with a small home range overlapping that of a more dominant individual and low travel distances, and floaters, with a large home range overlapping several more dominant individuals and high travel distances. Although a few low ranking juveniles achieved travel distances permitting high growth rates, most had either supra or suboptimal travel distances and relatively low growth rates.     

Environmental and genetic influences on mating strategies along a replicated food availability gradient in guppies (Poecilia reticulata)

Food availability is expected to influence the relative cost of different mating tactics, but little attention has been paid to this potential source of adaptive geographic variation in behavior. Associations between the frequency of different mating tactics and resource availability could arise because tactic use responds directly to food intake (phenotypic plasticity), because populations exposed to different average levels of food availability have diverged genetically in tactic use, or both. Different populations of guppies (Poecilia reticulata) in Trinidad experience different average levels of food availability. We combined field observations with laboratory “common garden” and diet experiments to examine how this environmental gradient has influenced the evolution of male mating tactics. Three independent components of variation in male behavior were found in the field: courtship versus foraging, dominance interactions, and interference competition versus searching for mates. Compared with low-food-availability sites, males at high-food-availability sites devoted more effort to interference competition. This difference disappeared in the common garden experiment, which suggests that it was caused by phenotypic plasticity and not genetic divergence. In the diet experiment, interference competition was more frequent and intense among males raised on the greater of two food levels, but this was only true for fish descended from sites with low food availability. Thus, the association between interference competition and food availability in the field can be attributed to a genetically variable norm of reaction. Genetically variable norms of reaction with respect to food intake were found for the other two behavioral components as well and are discussed in relation to the patterns observed in the field. Our results indicate that food availability gradients are an important, albeit complex, source of geographic variation in male mating strategies.     

Northern harriers on feeding territories respond more aggressively to neighbors than to floaters

Summary Territory owners often respond less aggressively towards intruding neighbors than towards intruding floaters, an observation termed the dear enemy phenomenon. Comparisons of territory owners' responses to intruding neighbors versus their responses to intruding floaters usually have been made for owners of multi-purpose and/or breeding territories. Here, I describe responses of female northern harriers Circus cyaneus (owners) on winter feeding territories towards three types of intruders (female neighbors, female floaters, and male floaters) and show that the dear enemy phenomenon does not occur. Owners' responses towards neighbors were more intense (mostly flights rather than calls) than responses towards female floaters, which in turn were more intense than responses towards male floaters. The greater intensity of owners' responses towards neighbors compared to owners' responses towards male and female floaters may be related to differences in the threat posed by each of the three intruder types in terms of fighting ability (RHP) and potential losses from intrusion. Hence, whether owners respond more aggressively towards neighbors or floaters, and whether the dear enemy phenomenon is observed, may depend upon the relative magnitude of threat presented by neighbors and floaters to owners in terms of fighting ability and potential losses from intrusion.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.     

Life history and reproduction of the amphipod Synchelidium trioostegitum (Crustacea, Oedicerotidae) on a sandy shore in Korea

The life history and reproductive strategy of the amphipod Synchelidium trioostegitum were studied on a sandy shore at Dolsando, South Korea. Samples were taken once a month for 1 year using a 0.3-mm sledge net on the bottom in 1 m of water at spring tide low water. The highest density of S. trioostegitum occurred from February through March. Ovigerous females were recorded virtually year-round, with a particularly high proportion in fall and early spring, indicating continuous recruitment with two dominant periods. The occurrence of ovigerous females was not correlated with environmental factors, such as temperature and salinity, and no significant difference between the body lengths of females and males was observed. The mean adult body length was greater in the early spring breeding period than in the fall. Brood size and embryo volume were positively correlated with the body length of ovigerous females. Brood size significantly decreased with increases in embryonic developmental stage. Embryo volume was significantly larger in the fall than in the early spring, but brood size was significantly smaller in fall, suggesting a strategy of using the same amount of reproductive energy during breeding periods. This type of reproductive effort is different from that of other Synchelidium amphipods having the same habitat and feeding regime. Our results suggest that interspecific competition for food and territory may be important in defining the reproductive strategy.