Life history of the Hawaiian blackhead triplefin, <Emphasis Type="Italic">Enneapterygius atriceps</Emphasis>(Blennioidei,Tripterygiidae)

Synopsis We described aspects of the life history of the small, cryptic, triplefin blenny, Enneapterygius atriceps, collected from the mouth of Kaneohe Bay, Oahu, Hawaii. We examined otolith microstructure to construct a von Bertalanffy growth curve, described a length–weight relationship, performed a detailed dietary analysis, used standard histological techniques to determine size at maturity and reproductive patterns, determined batch fecundity, and used an index of gonadal development to examine spawning seasonality. Enneapterygius atriceps is a remarkably short-lived species with low fecundity and a short, well-defined breeding season. Dietary analysis suggests this fish lives and feeds around the bases of dead coral heads. This study generated the first otolith-based growth curve for any member of the Tripterygiidae.     

Feeding ecology of the endangered Valencia toothcarp,Valencia hispanica (Actinopterygii: Valenciidae)

Patterns of food resource utilisation by Valencia toothcarp were studied in a small coastal lagoon in the Iberian Peninsula, its most northern population. Emphasis was placed on feeding activity, diet composition, seasonal and intraspecific variation in diet and feeding strategy. Important features of the overall feeding pattern of the Valencia toothcarp are decreased feeding activity during winter, concentration on benthic invertebrates and narrow dietary breadth, with a focus on gammarid amphipods. The feeding of V. hispanica was independent of seasonal variation in food availability, because its main prey were much the same throughout the year. There were no differences in seasonal niche width and niche overlap between seasons was high. There are also no differences in diets between sexes, although diet composition, niche width and niche overlap differ between juvenile and adult fish. The Valencia toothcarp population from Santes Creus lagoon was formed by specialist individuals, with narrow niche widths, that fed on four preferential prey types, but also consumed some occasional prey.     

Reproductive traits of the “ocellated killifish” Floridichthys polyommus Hubbs, 1936 (Pisces: Cyprinodontidae) inhabiting estuary of the Champotón River (Campeche,Mexico)

This study explores the reproductive biology and the influence of environmental factors on the reproductive traits of Floridichthys polyommus inhabiting the estuary of the Champoton River. The analysis included population structure, gonadal developmental stages, reproductive period, size at first sexual maturity, fecundity, sex ratio, and somatic indexes. A spatio‐temporal pattern in the distribution of the seven size classes was detected. The species showed a positive allometric growth. Sex ratio was 1.21:1 (females:males). Size at first maturity was 42.23 mm (females) and 47.8 mm (males). A higher absolute and relative fecundity was detected in Puente Champotón estuary (PCH) with lower salinities than those detected in the Delta (DE). The adaptive significance of this trait could be in response to the heavy osmotic constraint imposed by extreme salinities in the DE. Floridichthys polyommus prefers PCH for spawning, and as a result a higher number of ripe individuals was observed in this site, characterized by the highest nutrient levels, which can boost productivity and food availability for fish. Hepatosomatic index was negatively correlated with gonadosomatic index, suggesting energy transfer from the liver towards gamete production. Floridichthys polyommus exhibits an opportunistic strategy (early maturity, small oocytes, small clutches) suggesting that energy is allocated towards reproduction to compensate for the unpredictable hydrological conditions imposed by the estuary (particularly during the hurricane season). Floridichthys polyommus shows a reproductive strategy exhibited by fish living in unstable systems. A seasonal divergence was observed in the reproductive traits of F. polyommus, while differences between study sites were for fecundity.     

Studies on the life cycle and reproduction of the parasitic amphipodHyperia galba in the North Sea

The structure of aHyperia galba population, and its seasonal fluctuations were studied in the waters of the German Bight around the island of Helgoland over a period of two years (1984 and 1985). A distinct seasonal periodicity in the distribution pattern of this amphipod was recorded. During summer, when its hosts—the scyphomedusaeAurelia aurita, Chrysaora hysoscella, Rhizostoma pulmo, Cyanea capillata andCyanea lamarckii—occur in large numbers, supplying shelter and food, a population explosion ofH. galba can be observed. It is caused primarily by the relatively high fecundity ofH. galba which greatly exceeds that of other amphipods: a maximum of 456 eggs was observed. The postembryonic development is completed in the medusae infested; only then are the young able to swim and search for a new host. The smallest fréely-swimming hyperians obtained from plankton samples were 2.6 mm in body size. The size classes observed as well as moult increment and moulting frequencies in relation to different temperatures suggest that two generations are developed per year: a rapidly growing generation in summer and a slower growing generation in winter that shifts to a benthic mode of life and hibernation. For short periods, adult hyperians may become attached to zooplankters other than scyphomedusae. However, when releasing the progeny, they are dependent on the presence of these coelenterates. Apparently, a host specificity does not exist. During daytime, the hyperians seem to avoid a host change; only 0.2% of all the individuals sampled in the plankton during the day were not associated with medusae. The heavy infestation of medusae by this crustacean leads to a weakening and a progressive breakdown of these important predators on fish larvae.H. galba occupies a specific position in the marine food web which is discussed in detail.     

Comparisons between the influence of habitat type, season and body size on the dietary compositions of fish species in nearshore marine waters

Fish were collected from over bare sand in nearshore shallow waters at three sites that varied in the extent to which they were exposed to wave activity and which were located within a 45-km stretch on the lower west coast of Australia. Sampling was undertaken within a 4-6-week period in each season. The volumetric contributions of different prey to the stomach contents and the mouth characteristics of four species, i.e. Sillago bassensis and Sillago vittata (Sillaginidae), Spratelloides robustus (Clupeidae) and Pseudorhombus jenynsii (Bothidae), were determined. Overall, the dietary compositions of the four species differed significantly from each other and those of fish at both the three different sites and in four consecutive seasons were also significantly different. In comparison with S. bassensis, the morphologically similar S. vittata fed to a relatively greater extent on polychaetes than zooplankters, presumably reflecting in part its greater ability to extend its upper jaw downwards towards the benthos. S. robustus typically targeted calanoid copepods in the plankton, whereas P. jenynsii fed on larger benthic prey taxa, reflecting the large differences in mouth morphology and feeding behaviour of these species. Although the diets of S. bassensis and S. vittata were strongly influenced by habitat type and season, the former variable was slightly more important for both of these species. However, the reverse applied with S. robustus. Season strongly influenced the dietary composition of P. jenynsii at the one site at which it was regularly caught. The diets of the two Sillago species and P. jenynsii underwent pronounced size-related changes, which would help distribute the food resources among the individuals of the different size classes of each of these species. In contrast, all size classes of S. robustus fed predominantly or exclusively on calanoid copepods at all sites and in all seasons, except at the most sheltered site in winter when these zooplankters were not found in samples taken from the water column. The seasonal and habitat variations recorded in the diets of the fish species in this study imply that these species are able to feed opportunistically, a characteristic that would be of particular value to fish that live in nearshore waters where the relative abundance of the different prey types varies with habitat type and season.     

Maturity and fecundity of golden grey mullet (Liza aurata Risso, 1810) in Iranian waters of the Caspian Sea

The objective of the study was to determine some reproductive parameters in the golden grey mullet (Liza aurata Risso, 1810) during two periods half a decade apart while also trying from limited data to identify changes in fecundity over the past half century. Maturity and fecundity of golden grey mullet caught in Iranian waters of the Caspian Sea is not well documented and were thus studied during several seasonal cycles during 2002–2004 to establish basic data on reproduction in the area. The fork length (FL) of 462 sampled fish was between 22.1 and 51.9 cm. The fork length–weight relationship derived from all fish was W = 0.0118FL^2.964. Overall male to female ratio was 1 : 1.42. The reproductive season extended from October to December. Fifty percent of sexual maturity was at FL 26.0 cm. Mature gonads were present in 20% of fish at age 3, 63% at age 4, 88% at age 5, and 97% at age 6. Individual absolute fecundity of the golden grey mullet ranged from 113 386 to 1.47 million eggs, with a mean (±SD) of 451 963 (±274,114.2). An overlap exists between the spawning and fishing seasons. Therefore, the start of the fishing season should be delayed for 1 month in order for stocks to complete their spawning.     

Population Dynamics and Production of a Planktonic Marine Copepod,. Acartia clausii,in a Small Temperate Lagoon on San Juan Island,Washington

The population dynamics and production of Acartia clausii, a planktonic marine copepod, were studied during 1973 and 1974 in a small temperate lagoon. An approach which integrated laboratory and in situ experiments with time-series sampling of the field population was used to analyze seasonal changes in the parameters of population growth (development, growth, fecundity, and mortality) and to elucidate the processes which affect these changes. The objective of the study was to understand the factors which control the seasonal cycle of abundance. The cycles of abundance were similar in the two years of study and were not affected by differences in the cycles of tidal inflow, temperature, and food availability even though the latter two factors appreciably affected growth, development, and fecundity rates. The abundance cycle is controlled by an annually consistent pattern of copepodid and adult mortality believed to be due to predation by the three-spined stickleback, the dominant fish species in the lagoon. Cannibalism and periodic tidal stimulation of hatching of accumulated eggs in the sediment help to regulate population abundance within seasonal limits. Production by the lagoon population was 229 kg C for the entire study period; 84—88% of the annual production occurred from April through July. In the area of the lagoon greater than 3-m deep the mean daily productions during the peak months of each year were 70 and 55 mg C m⁻² for 1973 and 1974, respectively.     

Fast response of freshwater consumers to a new trophic resource: Predation on the recently introduced Asian bivalve Limnoperna fortunei in the lower Paraná river,South America

Abstract Comparison of Limnoperna fortunei numbers and biomass in screened (5, 15 and 40 mm) and unscreened cages deployed for 18 months in the lower Paraná delta indicates that predators harvest 26–79% (numbers), or 20–85% (biomass) of the mussel population. Predation impact decreases with mussel size. On average, 6 kg of whole live mussel × m⁻² × year⁻¹ (0.36 g of dry mussel tissue × m⁻² × day⁻¹) were eliminated from the unscreened cages. Cages with 15 and 40 mm screens lost between 1 and 2 kg × m⁻² × year⁻¹. Aquatic mammals, birds, and especially fish, are probably the main consumers of large mussels. Small L. fortunei are most probably eaten by fish and also by several invertebrates, including crustaceans, leeches and gastropods. It is suggested that L. fortunei intercepts a significant fraction of the organic carbon that the Paraná‐Uruguay rivers flush into the ocean, locally boosting numbers of benthophagous animals, deposit feeders and, indirectly, higher level predators. Our results indicate that only 15 years after its first introduction in South America this invasive species is very actively consumed by local predators, but predatory suppression of the mussel seems very unlikely. Comparisons with the effects reported for the zebra mussel (Dreissena polymorpha) in Europe and North America suggest that L. fortunei is consumed more actively and that its negative impact on the local fauna is more restricted. These differences are attributed to the fact that while D. polymorpha feeds chiefly on plankton, a limited resource, L. fortunei feeds on detrital particulate organic matter, whose supply in these large South American rivers largely exceeds consumption.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

Reproductive biology of Bagrus docmak in the Victoria Nile,Uganda

Aspects of the reproductive biology of Bagrus docmak in the Victoria Nile were investigated between November 2005 and October 2006. Macroscopic and histological analysis of the gonads confirmed it as an asynchronous batch spawner which spawns throughout the year with bimodal spawning peaks coinciding with rainfall seasons. The first spawning peak occurred from March to May, the second from September to November. The sex ratio did not significantly deviate from 1:1. Length at sexual maturity was 33.6 cm and 31.6 cm fork length (FL) for females and males, respectively. Batch fecundity ranged from 1 000 eggs in 34 cm FL fish to 43 000 eggs in 79 cm FL fish, and correlated linearly with FL (r = 0.72) and body weight (r = 0.79). Mean relative batch fecundity was 6 eggs g^?1 (SE 2). These results could guide research into the possibility of artificially inducing the fish to spawn, and its subsequent culture.     

Life-history phenotypes in a live-bearing fish Brachyrhaphis episcopi living under different predator regimes: seasonal effects?

Several key life-history attributes in a tropical live-bearing fish, Brachyrhaphis episcopi, have previously been shown to differ between populations that co-occur with large predatory fish (Characin sites) and those that do not (Rivulus sites). Here we show that differences between Characin and Rivulus localities are also repeatable over time; patterns observed in the wet season also persisted during the dry. Both sexes reached maturity at a smaller size at Characin sites. Although there was no difference in fecundity between larger females living in different predator communities, smaller females at Characin sites produced more offspring. Females also produced smaller offspring at Characin localities. These differences are remarkably similar to those reported in two other species of live-bearing fish, B. rhabdophora and Poecilia reticulata suggesting possible convergent adaptation in life-history strategies due to predator-mediated effects or correlates thereof. We also found seasonal changes in life-history traits that were independent of predator community. In the wet season, mature males were larger, females allocated more to reproduction, and offspring mass was also greater. The results of our study generate testable predictions using B. episcopi to further our understanding of life-history evolution.     

Aspects of the biology of the euryhaline Asian cichlid,Etroplus suratensis

Synopsis Aspects of the biology of the euryhaline Asian cichlid Etroplus suratensis in a coastal lagoon of Sri Lanka were studied for a period of 15 months. Comparisons are made from the catches from the fishery of manmade, inland freshwater reservoirs. Changes in the stock structure in the sublittoral region of the lagoon indicate that recruitment takes place twice a year, during the high rainfall/low salinity periods. The feeding habits of E. suratensis were different in the two habitats. In the lagoon it fed mainly on molluscs and in the freshwater reservoirs on macrophytes. Feeding chronology based on diurnal surveys indicate that it feeds mainly during the daylight hours. The dentition of E. suratensis is adapted for both tearing and crushing (pharyngeal teeth). Distinct differences in the mean relative intestinal length between populations from the coastal lagoon and inland reservoirs were evident and these differences are correlated to their respective feeding habits. Diurnal changes in feeding activity are associated with changes in the stomach pH. The egg-diameter distribution of mature fish indicate that E. suratensis is a single spawner, that it sheds its mature eggs all at once. It is inferred that E. suratensis breeds twice a year but that an individual female is capable of spawning only once a year.     

ECOLOGICAL SEPARATION BETWEEN WHITE AND PINK-BACKED PELICANS IN THE RUWENZOFU NATIONAL PARK, UGANDA

The number and distribution of the White Pelican Pelecanus onocrotalus roseus and the Pink-backed Pelican P. rufescens were studied in the Ruwenzori National Park, Uganda, by means of aerial surveys carried out at approximately weekly intervals between May 1968 and December 1969. The White Pelican is migratory and was present in the Park at highest numbers in August 1968 and May 1969 when 2510 and 3927 respectively were recorded. It does not breed within the Park. The Pink-backed Pelican is resident, with a population averaging about 1350 birds. Pelicans are distributed in discrete flocks on loafing grounds. Flocks of the White Pelican tend to be bigger than those of the Pink-backed. Forty-three loafing grounds were recognized in the Park, and it is shown that there is no difference between the species in the frequentation of these grounds. The areas favoured by both species are characterized by flat, open stretches close to water; steep slopes and heavily vegetated shore-lines are avoided. The feeding behaviour was studied by direct observation and through cine-photography. Only the White Pelican feeds communally in coordinated groups; this species also often feeds singly. The feeding sequence of both species is described, and is divided into four phases:searching, trapping, catching and swallowing. The searching phase lasts about six times longer in the Pink-backed Pelican than in the White. The White Pelican feeds further from the shore than does the Pink-backed. There is little difference between the species in the daily feeding pattern. Feeding takes place throughout the day but mostly from dawn to about 10.00 hrs and from 17.00 hrs until dusk. The rest of the day is spent mainly in preening, resting or bathing. The food of pelicans is exclusively fish. The White Pelican takes mainly large Tilupia, and Haplochromis and fish fry to a lesser extent. The Pink-backed Pelican feeds largely on fish fry, but Tilapia and Haplochromis are frequently taken and by weight are more important than the fry. However, the Tilapiu are smaller than those taken by the White Pelican. Ecological separation between the two species is achieved through these differences in their feeding behaviour. It is estimated that the White Pelican takes 1201 g and the Pink-backed Pelican 776 g of fish each day. The total amount of fish eaten by both species from Lake George during 1969 was calculated to be 591 709 kg. This is small (c. 3%) compared with the estimated total fish production of the lake but quite appreciable (12.7%) as a proportion of the total caught by man. It is concluded that the pelican populations are probably not important as a link in the chain whereby organic matter transferred from land to water by the Hippopotamus is returned to the land.     

Reproductive cycling and fecundity estimation in the upland bully

The upland bully is a prolific and widespread member of New Zealand freshwater fish communities that may mature at age 1 (or earlier). It begins spawning in spring, laying as many as eight clutches of eggs over the spring and summer, at periods as brief as 13 days. Ovaries contain two modes of maturing oocytes, so that oogenesis must be continual during the spawning season. This poses difficulties in fecundity estimation, which were solved by captive rearing. The ovoid eggs are about 2 mm long; egg size declines with clutch number during the summer, but increases with fish size. Clutch size varied, in 70 spawnings, from 121 to 880 eggs, with total seasonal fecundity up to >5000 eggs. The production of multiple clutches greatly increases seasonal fecundity and may explain the success of this species in sometimes unreliable or harsh environments.     

The effect of extreme northerly climatic conditions on the life history of the minnow, Phoxinus phoxinus (L.)

Over its exceptionally wide geographical range the minnow displays a remarkable variability in its life history. In southern England many fish mature after one year and few fish survive to their third birthday. Very few fish spawned for more than one season but a 65-mm female is estimated to spawn up to 3172 eggs over the extended spawning season. In the River Utsjoki in Finnish Lapland maturity was strongly size-dependent and delayed until the fish reached 5, 6 or even 7 years of age, with a maximum age of 13 years at a length of only 75 mm. Unlike some other small-sized species in unproductive environments, individual females continued to spawn successive clutches of eggs though over a much shorter spawning season. The maximum estimated egg production was 824 for a 65-mm fish. Temperature had a strong direct or indirect effect on growth, with significantly lower growth increments in cold summers. At a second Arctic site in eastern Lapland growth rates were higher and maturity earlier, yet clutch size was greatly reduced with a seasonal fecundity estimate of only 314. There were only small differences in size of eggs or larvae between the populations. Evidence from other species indicates that most of the observed changes over the range of the minnow are phenotypic responses to the contrasting environments. However, the different strategies displayed at the two Arctic sites could represent the results of selection for differences in pre- and post-reproductive survival.     

Biology of Oxyeleotris heterodon and its major prey,Ophieleotris aporos,two floodplain sleepers (Pisces: Eleotrididae) of the Sepik River fishery,northern Papua New Guinea

Synopsis Oxyeleotris heterodon attains 483 mm standard length (1.83 kg) and Ophieleotris aporos 245 mm (0.171 kg). Both species feature significantly in the local fishery in lowlands. Ophieleotris aporos is approximately one hundred times more abundant than Ox. heterodon in catches and landings which were influenced by river flood conditions in both species. Oxyeleotris heterodon is predominantly piscivorous but also eats Macrobrachium spp. prawns; fish prey consumed are almost exclusively Op. aporos. Ophieleotris aporos is omnivorous and feeds mainly on aquatic insects. Condition, fat deposition, feeding and reproduction in Op. aporos were related to the flood cycle with increases in the flood season; in Ox. heterodon variations in these factors related more to the availability of Op. aporos, seasonal changes being less marked. Ophieleotris aporos matures at a small size, has large gonads, small eggs and is highly fecund; Ox. heterodon exhibited opposite trends in these life-history characteristics by comparison. Reproductive styles adopted by the two species are interpreted in relation to the different predation pressures they experience. Oxyeleotris heterodon may not be adapted to high adult mortality and the fisheries implications of this are discussed.     

Reproductive strategy of Psectrogaster rhomboides Eigenmann & Eigenmann, 1889, a freshwater fish from Northeastern Brazil

Several environmental parameters related to the spawning season of the freshwater fish, Psectrogaster rhomboides were identified. The hypothesis predicts synchrony of the spawning season with the rainy period, presenting a seasonal reproductive strategy. This study investigated the total body length and body mass, sex ratio, body size at first sexual maturity, gonadosomatic index, condition factor, fecundity and reproductive period of P. rhomboides in the natural environment. Twelve monthly samplings (mean n = 20) were carried out from March 2009 to February 2010. Fish were captured using 4 cm diameter cast nets and environmental parameters such as rainfall, temperature, dissolved oxygen concentration, pH and electrical conductivity of the water were registered. The sampled population of P. rhomboides (n = 444) showed a sex ratio of 1 : 1. Females attained first sexual maturity at 15 cm total length and males at 15.3 cm total length. Spawning occurred from March to May, with high gonadosomatic index values during the rainy season. This species is a total spawner, with a mean fecundity of 6188 (±367) mature oocytes. P. rhomboides presents a seasonal reproductive strategy, providing a population increase and exploitation of the environmental resources during the rainy period. Rainfall (196.5 ± 4.9 mm) and the concentration of dissolved oxygen (7.82 ± 0.7018 mg L^?1) were influential factors during the spawning season. Gonadosomatic index and the condition factor were negatively correlated during the gonadal development cycle of this species. The monthly gonadosomatic index values show that spawning begins when the rainy season is at its maximum and ends synchronously (females r = 0.9171; males r = 0.889) with the end of the rainy season.     

Reproductive cycle of Triplophysa stenura (Herzenstein, 1888) (Balitoridae: Nemacheilinae) from the Yarlung Tsangpo River in the Tibetan Plateau,China

The reproductive biology of Triplophysa stenura, an indigenous cyprinid species from the Qushui County in the middle of the Yarlung Tsangpo River in the Tibetan Plateau, China, was investigated by bimonthly sampling from May 2014 to April 2015 using electrofishing. The gonadosomatic index (GSI) of T. stenura differed significantly between males and females over the seasonal trend, indicating that spawning extends from March (shortly after the ice melts) to May, as observed for indigenous species in response to environmental conditions on the cold plateau. Average oocyte diameter was 0.66 ± 0.13 mm, whereby the oocyte diameter distribution exhibited a clear seasonal pattern as the temporal pattern of GSI. Absolute fecundity of T. stenura was 3999 eggs/individual, which increased significantly with the increase in fish size and gonad weight, demonstrating the relationship of an exponential equation between them.     

Small-scale demographic variability of the biocolor damselfish, <Emphasis Type="Italic">Stegastes partitus</Emphasis>, in the Florida Keys USA

The demographic responses of reef fish to their environment can be complex and in many cases, quite strong. Growth, mortality, longevity, and even reproductive effort have been demonstrated to vary for the same species of reef fish over scales of 100s to 1,000s of kilometers due to physiological and ecological interactions. Though few studies have explicitly documented it, this sort of habitat-mediated demography can also exist at very local scales. Here we present the results of a 2-year study of the bicolor damselfish, Stegastes partitus, in the Florida Keys, USA. We measured density and distribution, calculated key demographic rates (growth, survival, and fecundity), and characterized the environment (resident fish assemblage, substrate type and complexity, and food availability) of populations living in two adjacent but different habitats, the continuous fore reef and patchy back reef. Fish on the fore reef had an elevated growth rate and asymptotic size, increased mortality, and higher fecundity than fish on the back reef. We identified four potential causative mechanisms for these differences: food availability; competition; intraspecific density-dependent effects; and predation risk. Our data did not support an effect of either food availability or intraspecific density-dependence, but rather suggested that demographic responses are affected by both competition and predation risk.