The influence of turbidity on growth and survival of fish larvae: a numerical analysis

Growth and survival through the early larval phase probably limit the production potential of many commercially important fish stocks. Attempts to increase the production of these stocks by restocking of juveniles have generally failed. Here, we analyse how enhanced concentrations of phytoplankton and zooplankton affect the survival of fish larvae during their early life stages. The analysis is developed for larvae feeding on copepod eggs and nauplii, with fish and invertebrates as major predators. A model of feeding and growth of fish larvae is applied to assess the benefit of enhanced phytoplankton and zooplankton abundance. The analysis shows that the shading effect of higher phytoplankton concentration may reduce predation rates on fish larvae substantially. This `top-down' effect may be more important for the cohort survivorship than the `bottom-up' mechanism in situations when larval food is sufficiently abundant. However, while increased algal biomass will improve recruitment at high zooplankton concentration, it may also reduce recruitment at low zooplankton concentrations and shallow mixing depths. Both the larvae and their vertebrate predators are dependent on light to detect their prey, and the longer reactive distance of the predators make them more susceptible than the larvae to reduced light levels and increased turbidity. We discuss the implications of reduced predation and increased zooplankton abundance on recruitment and production of fish larvae, and point at environmental conditions where changing algal biomass is likely to affect recruitment success.     

Role of feeding and prey distribution of summer and southern flounder in selection of estuarine nursery habitats

Diets of metamorphosing larvae and early juvenile summer and southern flounder were examined during the settlement period when their distributions overlapped and during a subsequent period when the two flounders were found in different habitats. Prey abundance on tidal flats was examined along an upstream transect. Though initially similar, diets of the two species diverged prior to the development of a segregated distribution. These diet differences reflected those found in the diets of the two species following segregation. Southern flounder ate more active epifaunal prey: mysids, amphipods and calanoid copepods and appeared to be an ambush predator. The summer flounder ate primarily infauna: polychaetes, and invertebrate parts and appeared to be an active forager. Distribution of prey within the study area suggested that mysid gradients may influence movement of southern flounder to nursery grounds. The coupling of biotic and abiotic gradients are probably important in creating nursery areas and guiding fishes to species specific habitat types.     

Seasonal change in the gastric evacuation rate of rainbow trout feeding on natural prey

Seasonal changes in the gastric evacuation rate ( R ) and gut contents of a wild population of rainbow trout Oncorhynchus mykiss feeding on natural prey at four water temperatures (2, 7, 9 and 12° C) were measured. The R and mass of the gut contents increased with water temperature, and prey items changed seasonally. These results suggest that the R of fish feeding on natural food depends primarily on water temperature, with their consumed prey being a secondary factor.     

Sediment dynamics and estuarine circulation in the turbidity maximum of the Elbe river

Data are presented from several campaigns concerning to the residual transport of water, salt and suspended particulate matter (SPM) at different depths and at different longitudinal and cross-sectional locations in the Elbe Estuary, Northern Germany. The intratidal behaviour of SPM is analysed and explained in terms of cyclic settling and resuspension. Superimposed on these local processes are advective horizontal transports of water and SPM in different vertical layers. Tidally averaged horizontal fluxes of salinity and SPM have been calculated at different depths. For the most part, they show upstream directed transport in the bottom layers, while in the upper water column SPM is carried towards the sea. Examples of the influence of vertical salinity gradients on SPM dynamics, of the lateral variation of residual transport of water and SPM, and the effects of a runoff peak are also presented.     

The influence of prey abundance on the feeding ecology of two piscivorous species of coral reef fish

Despite the potential importance of predation as a process structuring coral reef fish communities, few studies have examined how the diet of piscivorous fish responds to fluctuations in the abundance of their prey. This study focused on two species of rock-cod, Cephalopholis cyanostigma (Valenciennes, 1828) and Cephalopholis boenak (Bloch, 1790) (Serranidae), and monitored their diet in two different habitats (patch and contiguous reef) at Lizard Island on the northern Great Barrier Reef, Australia, over a 2-year period. The abundance of the rock-cods and the abundance and family composition of their prey were monitored at the same time. Dietary information was largely collected from regurgitated samples, which represented approximately 60% of the prey consumed and were unbiased in composition. A laboratory experiment showed that fish were digested approximately four times faster than crustaceans, leading to gross overestimation of the importance of crustaceans in the diet. When this was taken into account fish were found to make up over 90% of the diet of both species. Prey fish of the family Apogonidae, followed by Pomacentridae and Clupeidae, dominated the diet of both species of rock-cod. The interacting effect of fluctuations in prey abundance and patterns of prey selection caused dietary composition to vary both temporally and spatially. Mid-water schooling prey belonging to the families Clupeidae and to a lesser extent Caesionidae were selected for over other families. In the absence of these types of prey, apogonids were selected for over the more reef-associated pomacentrids. A laboratory experiment supported the hypothesis that such patterns were mainly due to prey behaviour. Feeding rates of both species of rock-cod were much higher in summer than in winter, and in summer they concentrated on small recruit sized fish. However, there was little variation in feeding rates between habitats, despite apparent differences in prey abundance. In summary, our observations of how the feeding ecology of predatory fish responded to variation in prey abundance provide potential mechanisms for how predation may affect the community structure of coral reef fishes.     

Effects of river flow on larval growth and survival of Japanese seaperch Lateolabrax japonicus (Pisces) in the Chikugo River estuary, upper Ariake Bay

The abundance and growth history of larval and juvenile Japanese seaperch Lateolabrax japonicus were investigated in the Chikugo River estuary, upper Ariake Bay, from 1990 to 2000. Growth during the larval period (up to 15 mm standard length, L _S, the size at recruitment into the estuary) was backcalculated using sagittal otolith microstructures by the biological intercept method. Growth rates in length declined at body sizes >14 mm L _S. High freshwater discharge through the Chikugo River was associated with high temperatures of the upper Ariake Bay where the larvae spend their planktonic life. Mean larval stage duration (days) from hatch to 15 mm ( D ₁₅) varied between 48·8 and 76·2 days and was inversely correlated with the estimated mean temperature history [mean daily temperature (° C) experienced by the larvae during the period from hatch to 15 mm, T ₁₅]. Mean abundance (number m⁻²) of larvae and juveniles was highest in years when T ₁₅, D ₁₅ and freshwater discharge were at intermediate levels. Although the abundance was not correlated with either of these variables, an exponential relationship between abundance and D ₁₅ was found when data collected during the highest river discharge years (1990, 1991 and 1998) were excluded. The increase in freshwater discharge through the Chikugo River probably had the potential to enhance or diminish Japanese seaperch recruitment in two ways: 1) it could increase recruitment probability by increasing temperature and larval growth and 2) high river flow also had the potential to decrease the probability of immigration into the river by increasing larval seaward dispersion, predation due to decreased turbidity and starvation due to decreased zooplankton prey abundance in the estuary.     

Spatial and temporal patterns in the occurrence of peritrich ciliates as epibionts on calanoid copepods in the Chesapeake Bay, USA

We investigated temporal and spatial patterns of distribution in two peritrich ciliates (i.e. Zoothamnium intermedium and Epistylis sp.) living as epibionts on calanoid copepods (i.e. Acartia tonsa and Eurytemora affinis) in Chesapeake Bay. Net tow samples collected along the main axis of the Bay were analyzed to estimate the occurrence of epibionts on copepods and to explore relationships among infestation prevalence, host abundance, and environmental variables. Zoothamnium intermedium and Epistylis sp. colonized populations of A. tonsa during spring and summer months, while only Z. intermedium colonized E. affinis during spring. Occurrence of epibionts on copepods showed high interannual variation, marked seasonality, and geographic heterogeneity. Extensive statistical analyses rejected simple scenarios of interactions between epibiosis, environmental variables, and host density, suggesting a more complex dynamics for the system. Analyses of epibiont colonies and zooids per host area (i.e. the sum of width and length of the body including antennae and swimming legs calculated assuming a cylindrical shape) were also performed. Overall, epibiont infestation prevalence (i.e. colonies/host area) and load (i.e. zooids/host area) were higher on copepodites than on adults for both host species, suggesting a preferential attachment to juveniles, or a higher predation pressure on adult stages. Infestation density and loads of both epibiont species were higher on the cephalothorax and abdomen of A. tonsa and E. affinis in comparison to the antennae and swimming legs, suggesting that ciliates can more easily colonize less active parts of the host.     

The evolutionary ecology of detritus feeding in the larvae of freshwater Diptera

Detritus (dead organic matter), largely of terrestrial origin, is superabundant in inland waters but because of its indigestible nature, would appear to be a poor food source for animals. Yet this unpromising material is widely used as food and indeed can be viewed as a defining characteristic of the freshwater environment. We here explore the relationships among animals, detritus and its associated micro-organism decomposers, taking a functional approach. We pose questions about interrelationships and attempt to arrive at new insights by disentangling them from an adaptive point of view. To do this we have been careful in selecting the habitats for detailed consideration. Rain pools on rock surfaces in tropical Africa and pools on peat moorland in the UK were chosen. Both examples have a relatively simple community structure and hence offer the prospect of achieving our aim. As model organisms for study we focus principally on the aquatic stages of selected holometabolous insects; that is, selected genera of the universally common midges, Ceratopogonidae and Chironomidae. We approach these case studies from an evolutionary ecology perspective and see detritus as a simple template upon which a beautiful complex of adaptations can evolve.     

Diet and Food Web Support of the White Perch, <Emphasis Type="Italic">Morone americana</Emphasis>, in the Hackensack Meadowlands of New Jersey

Synopsis I analyzed stomach contents of white perch, Morone americana, caught throughout the year in the Hackensack River in northern New Jersey. The dominant prey items were amphipods, with shrimp and fish being less important. Stable isotope analysis indicated that the primary producers of importance in the food web leading to white perch were benthic or planktonic microalgae and secondarily Phragmites australis, the dominant macrophyte in the Hackensack Meadowlands. Despite their large size, white perch do not generally appear to feed any higher in the food web than killifish, Fundulus heteroclitus.     

Spatial distribution and trophic ecology of dominant copepods associated with turbidity maximum along the salinity gradient in a highly embayed estuarine system in Ariake Sea, Japan

The present study aimed to investigate into the feeding ecology of the dominant copepods along a salinity gradient in Chikugo estuary. Copepod composition was studied from samples collected from stations positioned along the salinity gradient of the estuary. Copepod gut pigment concentrations were measured by fluorescence technique and hydrographical parameters such as temperature, salinity, transparency, suspended particulate matter (SPM); pigments such as chlorophyll-a (Chl-a), phaeopigment; and particulate nutrients such as particulate organic carbon (POC) and particulate organic nitrogen (PON) were measured. Two distinct zones in terms of nutrient and pigment concentrations as well as copepod distribution and feeding were identified along the estuary. We identified a zone of turbidity maximum (TM) in the low saline upper estuary which was characterized by having higher SPM, higher POC and PON but lower POC:PON ratios, higher pigment concentrations but lower Chl-a/SPM ratios and higher copepod dry biomass. Sinocalanus sinensis was the single dominant copepod in low saline upper estuary where significantly higher concentrations of nutrients and pigments were recorded and a multispecies copepod assemblage dominated by common coastal copepods such as Acartia omorii, Oithona davisae and Paracalanus parvus was observed in the lower estuary where nutrient and pigment concentrations were lower. Copepods in the estuary are predominantly herbivorous, feeding primarily on pigment bearing plants. However, completely contrasting trophic environments were found in the upper and the lower estuary. It was speculated from the Chl-a and phaeopigment values that copepods in the upper estuary receive energy from a detritus-based food web while in the lower estuary an algal-based food web supports copepod growth. Overall, the upper estuary was identified to provide a better trophic environment for copepod and is associated with higher SPM concentrations and elevated turbidity. The study demonstrates the role of estuarine turbidity maximum (ETM) in habitat trophic richness for copepod feeding. The study points out the role of detritus-based food web as energy source for the endemic copepod S. sinensis in the upper estuary, which supports as nursery for many fish species.     

The feeding strategy of dab Limanda limanda in the southern North Sea: linking stomach contents to prey availability in the environment

The feeding strategy of dab Limanda limanda was investigated within a predefined research area in the southern North Sea for 2 years. The aims were: 1) to establish the relationship between prey availability and dab abundance, distribution, body condition and prey selection, and 2) to determine the underlying functional processes of prey choice in dab. Temporal and spatial trends of prey availability in relation to dab abundances, distribution, condition and prey selection were analysed. Temporal trends in dab condition, percentage of empty stomachs and numbers of ingested prey were apparent. In particular in the winter period 2000–2001 dab were in poor condition and prey items were only rarely found in stomachs. Similarly prey availability decreased in the environment while the abundance of dab increased. Thus, temporal changes in the nutritive state were partly thought to be attributed to density‐dependent effects. The stomach composition of dab during the two consecutive summers was mainly dominated by Pariambus typicus and Ophiura albida while in the winter period only O. albida made a significant contribution to the stomach content. Analysis of prey characteristics indicated that only prey densities in the environment significantly influenced prey choice in dab, while other characteristics such as the position of the prey in the habitat, its palatability or mobility did not have significant effects. The feeding strategy of dab thus was highly opportunistic, although trends were apparent that suggested that buried living fauna was less likely to be ingested, which was also observed in previous studies.     

The nutritional ecology of larvae of Alsophila pometaria and Anisota senatoria feeding on early- and late-season oak foliage

The larvae of Alsophila pometaria (Harr.), feeding on the young foliage of oak, has a higher relative growth rate (RGR) and relative nitrogen accumulation rate (RNAR) than the larvae of Anisota senatoria (J. E. Smith), feeding on the mature foliage of oak. Although the young oak foliage is more efficiently digested by A. pometaria (higher AD's), it is not more efficiently assimilated and used for growth (no difference in ECI's). Thus, the higher growth rate of A. pometaria is due entirely to a higher consumption rate (RCR and RNCR). Young foliage is significantly higher in nitrogen and water than mature foliage, but phenol and tannin levels are comparable in young and old foliage. A. pometaria consumes the foliage of different oak species at the same rate, independent of nitrogen content, while A. senatoria increases its consumption rate in response to decreased nitrogen levels. As a result, the growth rate of A. pometaria is directly related to leaf nitrogen content, while the growth rate of A. senatoria is independent of leaf nitrogen. The two species of insects have digestive systems that are very similar biochemically, and that are well-designed for effective protein digestion. Tannins and phenols do not influence the nutrional indices of either species. We suggest that the major benefit of spring feeding is the availability of succulent, high-nitrogen foliage, and not the avoidance of high-tannin foliage. The spring feeder appears to have a feeding strategy that favors rapid growth at the expense of efficiency, while the late summer feeder has a strategy that favors efficiency over rate.

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Résumé Alimentées sur feuillage jeune de chêne, les chenilles d'Alsophila pometaria avaient un taux relatif de croissance (RGR) et un taux relatif d'accumulation d'azote (RNAR) plus élevés que les chenilles d'Anisota senatoria alimentées sur feuillage mûr de chêne. Bien que le jeune feuillage soit plus efficacement digéré par A. pometaria (AD plus élevé), il n'est pas assimilé et utilisé pour la croissance avec de meilleurs rendements (les ECI ne sont pas différents). Ainsi le taux de croissance plus élevé d'A. pometaria est dû entièrement à un taux de consommation plus important (RCR et RNCR). Le feuillage jeune est significativement plus riche en azote et en eau que le feuillage mûr, mais les niveaux de phénol et de tanins sont les mêmes. A pometaria consomme les feuilles de différentes espèces de chênes au même taux, indépendamment de la teneur en azote, tandis que A. senatoria accroît sa consommation en réponse à une diminution de la teneur en azote. Il en résulte que le taux de croissance d'A. pometaria dépend directement de la teneur en azote des feuilles, tandis que celui d'A. senatoria en est indépendant. Les systèmes digestifs des deux insectes sont biochimiquement semblables et sont efficaces pour la digestion des protéines. Les tanins et les phénols n'influent pas sur les indices nutritionnels de ces deux espèces. Nous estimons que le principal intérêt de l'alimentation printanière est la disponibilité en feuillage succulent, riche en azote, et non l'absence de feuilles à haute teneur en tanin. L'alimentation printanière semble correspondre à une strategie alimentaire qui favorise la croissance aux dépens de l'efficacité tandis que l'alimentation en fin d'été est une stratégie qui favorise l'efficacité sur la rapidité.

     

Spatial variation in abundance of prey and diet of trumpeter (Pelates sexlineatus: Teraponidae) associated with Zostera capricorni seagrass meadows

Abstract The diet of juveniles of Pelates sexlineatus was examined at six estuaries, separated by tens to hundreds of kilometres, and two sites within each estuary, separated by hundreds of metres to kilometres. Fish were collected in Zostera capricorni seagrass meadows along the coast of New South Wales (Australia). Spatial variability in diet was compared with the abundance of prey. Pelates sexlineatus had a broad diet (27 different prey) but generally preferred crustaceans (harpacticoid copepods, gammarid amphipods, ostracods and tanaids), although at some sites other prey items were important (e.g. polychaetes, nematodes and foraminiferans). Composition of the diet varied among estuaries and between sites. Proportional representation of the different size of prey eaten by the three sizes of juveniles (40‐ 54 , 55–74 and 75–94 mm standard length) was similar. Fish generally preferred prey smaller than 1 mm. Abundance of prey also varied at both spatial scales. At five of the 12 sites, there was a significant correlation between abundance of prey in the seagrass meadows and abundance of prey in the diet. Variation in the composition of the diet was partly explained by spatial variation in abundance of prey. When crustaceans were not abundant in the seagrass, P. sexlineatus had a broad diet, taking both benthic and planktonic prey items. It is concluded therefore that trophic linkages between P. sexlineatus and benthic invertebrates may vary greatly with spatial scales from hundreds of metres to hundreds of kilometres, and are strongly related to availability of prey in seagrass habitats.     

The feeding ecology of a carnivorous plant (Pinguicula nevadense): prey analysis and capture constraints

Summary The taxonomic composition and size of arthropods captured by Pinguicula nevadense, an endemic carnivorous plant of the high-mountain zone of the Sierra Nevada (southern Spain), are analysed. The actual prey of P. nevadense and the available arthropods trapped by mimic-traps are compared, in order to identify the capture constraints of the plant. The results show that P. nevadense captures various arthropod taxa. Winged insects, especially Nematocera, make up the main component of the diet. The range of prey sizes in all P. nevadense populations studied is similar. The taxonomic composition of arthropods trapped by the mimic-traps is similar to that of the actual prey of P. nevadense. However, the plant captures prey only below a specific size threshold. These size constraints appear to be the principal factor determining the actual prey of this carnivorous plant.     

Comparative ontogeny of the feeding apparatus of sympatric drums (Perciformes: Sciaenidae) in the Chesapeake Bay

The anatomy of the feeding apparatus in fishes, including both oral and pharyngeal jaw elements, is closely related to the ecology of a species. During ontogeny, the oral and pharyngeal jaws undergo dramatic changes. To better understand how such ontogenetic changes occur and relate to the feeding ecology of a species, ontogenetic series of four closely related members of the family Sciaenidae (Cynoscion nebulosus, Cynoscion regalis, Micropogonias undulatus, and Leiostomus xanthurus) were examined. Sciaenids were selected because as adults they exhibit considerable specialization of the feeding apparatus correlated with differences in foraging habitats. However, it is not clear when during ontogeny the structural specializations of the feeding apparatus develop, and thereby enable early life history stage (ELHS) sciaenids to partition their foraging habitats. A regression tree was recovered from the analysis and three divergences were identified during ontogeny. There are no measurable differences in elements of the feeding apparatus until the first divergence at 8.4 mm head length (HL), which was attributed to differences in average gill filament length on the second ceratobranchial. The second divergence occurred at 14.1 mm HL and was associated with premaxilla length. The final divergence occurred at 19.8 mm HL and was associated with differences in the toothed area of the fifth certatobranchial. These morphological divergences suggest that ELHS sciaenids may be structurally able to partition their foraging habitats as early as 8.4 mm HL. J. Morphol. 277:183–195, 2016. © 2015 Wiley Periodicals, Inc.     

The effects of turbidity, prey density and environmental complexity on the feeding of juvenile Murray cod Maccullochella peelii

Juvenile Murray cod Maccullochella peelii exhibited a type II functional response while preying on blackworms Lumbriculus variegatus, and the parameters of the type II model did not differ significantly between clear (0 NTU) and turbid (150 NTU) treatments. Further experiments showed that vision may not be necessary for prey detection and capture by juvenile M. peelii; consumption of inanimate prey was not significantly different between light and dark (<1 × 10(-4) μE m(-2) s(-1)) trials. These results imply that the sensory physiology of M. peelii is well adapted to a turbid visual environment. In addition, habitat complexity increased the food consumption rate of juvenile M. peelii, perhaps by relaxing innate predator avoidance behaviours that depress foraging in more open environments.     

Spotted hyaena Crocuta crocuta feeding ecology and selectivity of large herbivorous prey in the Namib desert

We have investigated the relationship between spotted hyaenas in the south Namib Desert and large herbivorous prey and have summarized an updated overview of predator‐prey relationships in this resource‐limited arid environment. Over the 52‐month study, we recorded the densities (#/km⁻², ±SE) of the four local large herbivorous prey species: gemsbok (1.229, ±0.50), springbok (1.352, ±0.48), ostrich (0.648, ±0.23), and greater kudu (0.343, ±0.00). A fecal analysis was performed on 146 collected spotted hyaena scats, and prey items were identified and hairs cross‐follicle analyzed to the species level. Spotted hyaena diet at the study area remained opportunistic with 240 identified prey items representing eight differing prey species being recorded, ranging from ostrich eggs to large ungulates. The Ivlev''s Electivity Index was used to determine which large herbivorous prey was most selected for. Although gemsbok had a higher representation of prey items in the sampled scats, all sampled large herbivorous prey species scored below 0 and are thus generally avoided in relation to their availability in the environment. If any prey preferences are expressed by spotted hyaena in the Namib, it can be presumed to be a nonsampled prey species. We therefore promote further detailed investigations into all other prey species present, and seasonal variations of prey densities and scat sampling, within the study environment.     

Kin recognition in Drosophila: the importance of ecology and gut microbiota

The animal gut commonly contains a large reservoir of symbiotic microbes. Although these microbes have obvious functions in digestion and immune defence, gut microbes can also affect behaviour. Here, we explore whether gut microbiota has a role in kin recognition. We assessed whether relatedness, familiarity and food eaten during development altered copulation investment in three species of Drosophila with diverse ecologies. We found that a monandrous species exhibited true kin recognition, whereas familiarity determined kin recognition in a species living in dense aggregations. Finally, in a food generalist species, food eaten during development masked kin recognition. The effect of food type on copulation duration, in addition to the removal of this effect via antibiotic treatment, suggests the influence of bacteria associated with the gut. Our results provide the first evidence that varied ecologically determined mechanisms of kin recognition occur in Drosophila, and that gut bacteria are likely to have a key role in these mechanisms.     

The feeding ecology of the dingo II. Dietary and numerical relationships with fluctuating prey populations in south-eastern Australia

The dietary and predator-prey relationships of Canis familiaris dingo were studied for 9 yr at a coastal site and for 1.5 yr at a montane site in south-eastern Australia. The percentage occurrences of items eaten were obtained from faeces, and the abundances of prey by counting water-birds, trapping small mammals, and tracking large and medium-sized mammals on specially prepared soil plots. Dingoes were also estimated by tracking. The diet was broad but predominantly mammalian (23 species). Dietary frequencies were grouped around three weight modes, 0.1, 1.25 and 16 kg, corresponding with bush rat (Rattus fuscipes), ringtail possum (Pseudocheirus peregrinus) and swamp wallaby (Wallabia bicolor). The amplitudes of those frequencies approximately doubled from mode to mode, viz. 4, 13.5 and 28.3%, respectively. Medium-sized mammals have been recognized as the staple prey because of their dependability. Their density estimates fluctuated least among prey-groups, and track records of dingoes in the mountains were significantly related to them. Other categories of prey were supplementary (large mammals), opportune (small mammals) and scavenged. Although there were general tendencies for dietary frequencies to follow prey abundances, significant functional and numerical responses were obtained only for water-birds (coot and swan). Their super-abundance in the mid-years of the coastal study and their highly clumped distribution were the likely causes. Predation was disproportionately severe on mammalian prey-classes after an extensive wildfire at the coastal site. Such predation may have suppressed populations of wallabies and kangaroo for 2 yr until the water-birds became super-abundant. The prevalence of wombats in the mountains may have induced heavy predation upon other less numerous large prey. Concepts of profitability in feeding appear to apply to the dingo more than those of optimization of time or energy. The decline in dingoes was correlated most with long-term declines in water-birds and medium-sized mammals. At the time, abundance estimates of wallabies and kangaroos were increasing and those species increasingly predominated in the diet. The ability to hunt co-operatively was apparently ineffectual in preventing decline in dingo numbers. It is suggested that pack size is related inversely to the level of temporal instability in the environment. Frequent wildfires may prevent staple (mediumsized) and supplementary (large) prey from being abundant simultaneously, a combination thought necessary for large pack size.