Density dependence and density independence during the early life stages of four marine fish stocks

Recruitment variability caused by density-dependent and density-independent processes is an important area within the study of fish dynamics. These processes can exhibit nonlinearities and nonadditive properties that may have profound dynamic effects. We investigate the importance of population density (i.e., density dependence) and environmental forcing (i.e., density independence) on the age-0 and age-1 abundance of capelin (Mallotus villosus), northeast Arctic cod (Gadus morhua), northeast Arctic haddock (Melanogrammus aeglefinus), and Norwegian spring spawning herring (Clupea harengus) in the Barents Sea. We use statistical methods that explicitly account for nonlinearities and nonadditive interactions between internal and external variables in the abundance of these two pre-recruitment stages. Our results indicate that, during their first five months of life, cod, haddock, and herring experience higher density-dependent survival than capelin. The abundance of age-0 cod depends on the mean age and biomass of the spawning stock, a result which has implications for the management of the entire cod stock. Temperature is another important factor influencing the abundance at age-0 and age-1 of all four species, except herring at age-1. Between age-0 and age-1, there is an attenuation of density-dependent survival for cod and herring, while haddock and capelin experience density dependence at high and low temperatures, respectively. Predation by subadult cod is important for both capelin and cod at age-1. We found strong indications for interactions among the studied species, pointing to the importance of viewing the problem of species recruitment variability as a community, rather than as a population phenomenon.     

The influences of climate and hydrology on population dynamics of waterbirds in the lower Murrumbidgee River floodplains in Southeast Australia: Implications for environmental water management

Restoration of waterbird diversity and abundance is a key objective of river system management in Australia. Therefore, understanding the effects of climatic and hydrological variables on waterbird population dynamics is fundamental for successful river restoration programs. We investigated the population dynamics of waterbirds (total abundance) and seven functional waterbird groups in the floodplains of lower Murrumbidgee River. We found a general declining abundance trend from 1983 to 2007, except for the deep water foragers. We modelled the relative contribution of the climatic and hydrological factors to waterbird population decrease using the generalized additive model (GAM) framework after identifying the negative binomial distribution. Most of the seven functional groups were positively related to both annual rainfall and water usage, defined as the total water volume intercepted by the river reach, and the models indicated that rainfall was slightly more important. Temperature also played a role in waterbird abundance: the maximum summer temperature negatively influenced the abundance of dabbling ducks, shoreline foragers and fish eaters, while the minimum winter temperature positively affected the abundance of dabbling ducks and shoreline foragers. Overall, our results support the practice of providing environmental water for sustaining waterbird populations. However, environmental water provision is likely to be most effective when timed to coincide with antecedent rainfall.     

Diet of 0-group stages of capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>), herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) and cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during spring and summer in the Barents Sea

Recruitment of capelin in the Barents Sea fail when juvenile herring and cod are abundant and the potential for feeding competition of wild sympatric capelin and herring larvae and small cod juveniles were investigated. The frequency of gut evacuation after capture of capelin larvae were also studied in mesocosms. Small capelin larvae (<35 mm length) fed on small prey including phytoplankton, invertebrate eggs and nauplii, bivalves, other invertebrate larvae and small copepods. Calanus copepodites were only observed in large capelin larvae (>26 mm length). Calanus copepodites were the major food sources for contemporary herring larvae (25–35 mm length) and Calanus and euphausiids were the major prey for small juvenile herring (37–60 mm length) and cod (18–40 mm length). Capelin larvae reared in mesocosms evacuated the guts shortly after capture. Capelin larvae had a smaller mouth and fed on smaller prey than herring and cod of the same length. This implies that the small capelin larvae, in contrast to sympatric small herring and cod, are not tightly linked to the food chain involving Calanus and euphausiids. Thus, exploitative competition between capelin larvae and planktivorous fish that rely on Calanus and euphausiids in the Barents Sea may be relaxed.     

Allee Effect and the Uncertainty of Population Recovery

Recovery of depleted populations is fundamentally important for conservation biology and sustainable resource harvesting. At low abundance, population growth rate, a primary determinant of population recovery, is generally assumed to be relatively fast because competition is low (i.e., negative density dependence). But population growth can be limited in small populations by an Allee effect. This is particularly relevant for collapsed populations or species that have not recovered despite large reductions in, or elimination of, threats. We investigated how an Allee effect can influence the dynamics of recovery. We used Atlantic cod (Gadus morhua) as the study organism and an empirically quantified Allee effect for the species to parameterize our simulations. We simulated recovery through an individual‐based mechanistic simulation model and then compared recovery among scenarios incorporating an Allee effect, negative density dependence, and an intermediate scenario. Although an Allee effect significantly slowed recovery, such that population increase could be negligible even after 100 years or more, it also made the time required for biomass rebuilding much less predictable. Our finding that an Allee effect greatly increased the uncertainty in recovery time frames provides an empirically based explanation for why the removal of threat does not always result in the recovery of depleted populations or species. El Efecto Allee y la Incertidumbre de la Recuperación de Poblaciones     

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.     

Continued decline of an Atlantic cod population: how important is gray seal predation?

The continental shelf ecosystem on the Eastern Scotian Shelf (ESS) has experienced drastic changes. Once common top predators are a small fraction of their historical abundance, and much of the current community structure is now dominated by pelagic fishes and invertebrates. Embedded within this food web, Atlantic cod and gray seal populations have recently exhibited nearly opposite trends. Since 1984, cod populations have decreased exponentially at a rate averaging 17% per year, whereas gray seals have continued to increase exponentially at a rate of 12%. We reexamined the impact of gray seals on Atlantic cod dynamics using more than 30 years of data on the population trends of cod and gray seals while incorporating new information on seal diet and seasonal distribution. The closure of the cod fishery over 10 years ago allowed for a better estimation of natural mortality rates. We quantified the impact of seals on ESS cod by (1) estimating trends in seal and cod abundance, (2) estimating the total energy needed for seal growth and maintenance from an energetics model, (3) using estimates of the percentage of cod in the total diet derived from quantitative fatty acid signature analysis (QFASA) and of the size-specific selectivity of cod consumed (derived from otoliths collected from fecal samples), and (4) assuming a gray seal functional response. Uncertainties of the model estimates were calculated using the Hessian approximation of the variance-covariance matrix. Between 1993 and 2000, cod comprised, on average, < 5% of a gray seal's diet. Our model shows that, since the closure of the fishery, gray seals have imposed a significant level of instantaneous mortality (0.21), and along with other unknown sources of natural mortality (0.62), are contributing to the failure of this cod stock to recover.     

Impact of minimum winter temperatures on the population dynamics of Dendroctonus frontalis.

Predicting population dynamics is a fundamental problem in applied ecology. Temperature is a potential driver of short-term population dynamics, and temperature data are widely available, but we generally lack validated models to predict dynamics based upon temperatures. A generalized approach involves estimating the temperatures experienced by a population, characterizing the demographic consequences of physiological responses to temperature, and testing for predicted effects on abundance. We employed this approach to test whether minimum winter temperatures are a meaningful driver of pestilence from Dendroctonus frontalis (the southern pine beetle) across the southeastern United States. A distance-weighted interpolation model provided good, spatially explicit, predictions of minimum winter air temperatures (a putative driver of beetle survival). A Newtonian heat transfer model with empirical cooling constants indicated that beetles within host trees are buffered from the lowest air temperatures by approximately 1-4 degrees C (depending on tree diameter and duration of cold bout). The life stage structure of beetles in the most northerly outbreak in recent times (New Jersey) were dominated by prepupae, which were more cold tolerant (by >3 degrees C) than other life stages. Analyses of beetle abundance data from 1987 to 2005 showed that minimum winter air temperature only explained 1.5% of the variance in interannual growth rates of beetle populations, indicating that it is but a weak driver of population dynamics in the southeastern United States as a whole. However, average population growth rate matched theoretical predictions of a process-based model of winter mortality from low temperatures; apparently our knowledge of population effects from winter temperatures is satisfactory, and may help to predict dynamics of northern populations, even while adding little to population predictions in southern forests. Recent episodes of D. frontalis outbreaks in northern forests may have been allowed by a warming trend from 1960 to 2004 of 3.3 degrees C in minimum winter air temperatures in the southeastern United States. Studies that combine climatic analyses, physiological experiments, and spatially replicated time series of population abundance can improve population predictions, contribute to a synthesis of population and physiological ecology, and aid in assessing the ecological consequences of climatic trends.     

Habitat associations and diet of young-of-the-year Pacific cod (<Emphasis Type="Italic">Gadus macrocephalus</Emphasis>) near Kodiak,Alaska

The influence of environmental variables and habitat on growth and survival of juvenile gadoid species in the Atlantic has been clearly demonstrated; conversely, in the North Pacific little is known about the habitat and ecology of juvenile Pacific cod (Gadus macrocephalus Tilesius). The hypothesis that density of young-of-the-year (YOY) Pacific cod in nearshore habitats is predicted by shallow depth and the presence of eelgrass and macroalgae was tested in a variety of nearshore habitats adjacent to commercial fishing grounds near Kodiak Island, AK. From 10 to 22 August 2002, a beach seine and small-meshed beam trawl were used to capture YOY Pacific cod (n = 254) ranging from 42 to 110 mm fork length. Depth, water temperature, salinity, sediment grain size, and percent cover by emergent structure (i.e. tube-dwelling polychaetes, sea cucumber mounds, macroalgae) were measured prior to fishing. Density of YOY Pacific cod was highest in areas of moderate depth (15–20 m) and positively and linearly related to percent cover by sea cucumber mounds and to salinity. No previous studies have documented fish utilizing sea cucumber mounds as habitat. Furthermore, eelgrass and macroalgae were inconsequential to cod distribution. Diets consisted mainly of small calanoid copepods, mysids, and gammarid amphipods and were significantly related to cod length and depth. This work provides important information on previously undocumented factors that affect distribution and feeding of YOY Pacific cod, which ultimately influence growth and survival in this species.     

Effects of law enforcement and community outreach on mammal diversity in a biodiversity hotspot

Management activities such as law enforcement and community outreach are thought to affect conservation outcomes in protected areas, but their importance relative to intrinsic environmental characteristics of the parks and extrinsic human pressures surrounding the parks have not been explored. Furthermore, it is not clear which is more related to conservation outcomes—the management itself or local people's perceptions of the management. We measured objective (reports by park staff) and subjective (reports by local people) levels of community outreach and law enforcement based on responses to 374 questionnaires. We estimated mammal abundance and diversity of 6 protected areas based on data from 115 camera traps in Xishuangbanna, southwest China, a biodiversity hotspot with high hunting and land-conversion pressures. We then examined correlations among them and found that local people's perception of law enforcement was positively related to the local abundance of 2 large, hunted species, wild boar (Sus scrofa) (β = 15.22) and muntjac (Muntiacus vaginalis) (β = 14.82), but not related to the abundance of smaller mammals or to objective levels of enforcement. The subjective frequency of outreach by park staff to local communities (β = 3.42) and park size (β = 3.28) were significantly and positively related to mammal species richness, whereas elevation, human population density, and subjective frequency of law enforcement were not. We could not conclude that community outreach and law enforcement were directly causing increased mammal abundance and diversity. Nevertheless, the patterns we detected are some of the first empirical evidence consistent with the idea that biodiversity in protected areas may be more positively and strongly related to local perceptions of the intensity of park management than to either intrinsic (e.g., elevation, park size) or extrinsic (e.g., human population density) environmental factors.     

Testing the irruptive paradigm of large-herbivore dynamics

A dominant paradigm in understanding and managing large herbivores is that, after introduction to new range or release from harvesting, the herbivore population increases to peak abundance, crashes to a lower abundance, and then increases to a carrying capacity lower than peak abundance. However, support for the paradigm has been largely anecdotal. We first developed two mathematical models to better describe irruptive dynamics. The models differed in the form of the postcrash growth toward carrying capacity: the "Caughley model" included a time lag that generated dampening oscillations, and the "Leopold model" did not. We then evaluated which of four models (theta-logistic, delayed-logistic, Leopold, and Caughley) best described the dynamics of seven ungulate populations either introduced to new range (n = 5 populations) or released from harvesting (n = 2). The dynamics of six of the populations were best described by irruptive models (two by the Leopold, one by the Caughley, and three by the delayed-logistic), and one of the populations did not display irruptive dynamics (theta-logistic model). The limited data thus support the widespread existence of irruptive dynamics, and we encourage the consideration of irruptive models in studies of large-herbivore dynamics.     

Influence of increased cod abundance and temperature on recruitment of northern shrimp (Pandalus borealis)

Despite an increase in northern shrimp (Pandalus borealis) female biomass in the past years, the recruitment of the offshore population north and northeast of Iceland has remained very low. In this study, the influence of abiotic and biotic factors was studied in relation to shrimp recruitment. Two factors, cod (Gadus morhua) abundance and summer sea surface temperature (SST), were found to have a negative effect on offshore shrimp recruitment, explaining 71 % of the observed variation. Both cod abundance and temperature on the offshore shrimp grounds have increased in the past years, while recruitment has decreased and been at historically low levels since 2005. No significant relationship was found between recruitment and spawning biomass, indicating that recruitment variability is mainly driven by other factors. Cod abundance and summer SST are likely to affect different life stages of shrimp, as SST influences shrimp during its planktonic phase while cod abundance influences the demersal stage.     

Irruptive population dynamics in Yellowstone pronghorn.

Irruptive population dynamics appear to be widespread in large herbivore populations, but there are few empirical examples from long time series with small measurement error and minimal harvests. We analyzed an 89-year time series of counts and known removals for pronghorn (Antilocapra americana) in Yellowstone National Park of the western United States during 1918-2006 using a suite of density-dependent, density-independent, and irruptive models to determine if the population exhibited irruptive dynamics. Information-theoretic model comparison techniques strongly supported irruptive population dynamics (Leopold model) and density dependence during 1918-1946, with the growth rate slowing after counts exceeded 600 animals. Concerns about sagebrush (Artemisia spp.) degradation led to removals of >1100 pronghorn during 1947-1966, and counts decreased from approximately 700 to 150. The best models for this period (Gompertz, Ricker) suggested that culls replaced intrinsic density-dependent mechanisms. Contrary to expectations, the population did not exhibit enhanced demographic vigor soon after the termination of the harvest program, with counts remaining between 100 and 190 animals during 1967 1981. However, the population irrupted (Caughley model with a one-year lag) to a peak abundance of approximately 600 pronghorn during 1982-1991, with a slowing in growth rate as counts exceeded 500. Numbers crashed to 235 pronghorn during 1992-1995, perhaps because important food resources (e.g., sagebrush) on the winter range were severely diminished by high densities of browsing elk, mule deer, and pronghorn. Pronghorn numbers remained relatively constant during 1996-2006, at a level (196-235) lower than peak abundance, but higher than numbers following the release from culling. The dynamics of this population supported the paradigm that irruption is a fundamental pattern of growth in many populations of large herbivores with high fecundity and delayed density-dependent effects on recruitment when forage and weather conditions become favorable after range expansion or release from harvesting. Incorporating known removals into population models that can describe a wide range of dynamics can greatly improve our interpretation of observed dynamics in intensively managed populations.     

Decreased lipid storage in juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) during settlement in cold-water eelgrass habitat

We characterized the prey field and the lipid classes/fatty acids in the flesh of age 0 juvenile cod (Gadus morhua) during their late-summer/fall arrival and settlement into eelgrass (Zostera marina) in coastal Newfoundland. Examination of available prey demonstrated a high abundance of small zooplankton (Acartia, Microsetella and Oithona sp.) with no larger Calanus sp. prey. Breakpoint analysis showed significant changes in the accumulation of relative (mg g⁻¹ wet weight) and absolute (μg fish⁻¹) amounts of lipid with standard length at the time of settlement (~60 mm standard length). Settling juvenile cod showed an alternate lipid utilization strategy where they catabolized phospholipids (PL) to a greater extent than triacylgylcerols (TAG). Polyunsaturated fatty acids (PUFA) content in cod flesh decreased as fish grew indicating that nearshore zooplankton quality was not optimal for PL formation. The dramatic reduction in cod PL was likely due to both catabolism of muscle and a lack of dietary PUFA suitable for PL synthesis. However, juvenile cod continued to grow, leading to decreased lipid stores and suggesting that cod settling into eelgrass are under intense selection pressure for growth prior to the onset of winter, possibly as a means of escaping gape-limited predation. These data contrast better-studied freshwater and estuarine systems in which lipid storage is critical for successful overwintering.     

Changes in the timing of otolith zone formation in North Sea cod from otolith records: an early indicator of climate-induced temperature stress?

We examine the seasonal variation in otolith increment formation in southern North Sea cod as a means of monitoring how changes in sea temperature over the past 20 years have affected cod in the wild. Seasonal opaque zone formation was related to winter and early spring. Timing of opaque zone formation was not influenced by either temperature or fish length, but increasing age led to slightly earlier but slower opaque zone formation. In contrast, there was a clear shift in the timing of translucent zone formation with temperature. In warm years, translucent growth occurs up to 22 days earlier than in colder periods. Increasing age and smaller size-at-age resulted in an earlier transition from opaque to translucent edge formation. Translucent zone formation appears indicative of increasing metabolic stress, and the earlier onset provides direct evidence of the impact of increasing sea temperatures on wild North Sea cod stocks.     

Optimal exploitation of a fishery employing a non-linear harvesting function

A harvesting function is developed to described the rate of removal of fish from a fish population. The function incorporates the effects of both the handling or processing time of the catch and the competition, between boats in the fleet, for the fish.We will assume that the growth rate of the fish population can be modelled with a concave, dome shaped growth curve. With this assumption, it has been shown that if the rate of harvesting the fish is linearly related to both effort (which can be thought of as some measure of the number of boats in the fleet) and the population size, then the population will tend towards a single equilibrium level which is globally stable. This paper shows that the saturation effects due to the handling time may generate two equilibrium levels (one stable, one unstable) rather than a single globally stable equilibrium. The results of competition between boats are economically undesirable because of the decrease in efficiency. However, this competition may be beneficial to the exploited fish population.Using the harvesting model derived earlier, the steady state or long term optimal harvesting policies as well as the transition paths to these states are developed. The only constraint is on the maximum allowable effort which is effectively an upper limitation on the fleet size or number of man-hours of fishing.     

Importance of biotic and abiotic regulators of abundance of the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae) and assessment of the accuracy and precision of sampling methods for detecting abundance changes

We examined seasonal patterns of abundance for the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae). Amphipods were sampled with an epibenthic sled during the daytime high tide period from 18 May to 8 August 1988. Amphipod density increased from May to June and reached maxima in both late June and early August. Amphipod density was unrelated to any abiotic component measured in the intertidal community. These variables included sampling location, wave height, water column height, water temperature, salinity and cloud cover. Changes in abundance were related with sampling date and with the onset of capelin (Mallotus villosus) spawning activity in the intertidal. Capelin eggs are an important food item for amphipods. The accuracy of density estimates obtained with the epibenthic sled was assessed through comparison with densities obtained with a more efficient quadrat sampler. Sled samples consistently sampled ca. 1% of the amphipod population. We found that a large portion of the amphipod population burrowed into the sediment and was not effectively sampled by the sled. Sled sampler precision was roughly equivalent to that of quadrats with D (precision) ranging from 0.26 to 0.42 for sled samples and D=0.29 for quadrat samples. Although sample collection with the epibenthic sled was achieved more quickly and under a wider range of weather conditions than was possible with the quadrat sampler, the serious underestimate of amphipod density based on sled samples alone indicates that both sled and quadrat samples, obtained in concert, are required to obtain accurate measures of daily variation in C. laeviusculus abundance.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches oceanographique du Quebec)     

Nonlinear effects of climate and density in the dynamics of a fluctuating population of reindeer

Nonlinear and irregular population dynamics may arise as a result of phase dependence and coexistence of multiple attractors. Here we explore effects of climate and density in the dynamics of a highly fluctuating population of wild reindeer (Rangifer tarandus platyrhynchus) on Svalbard observed over a period of 29 years. Time series analyses revealed that density dependence and the effects of local climate (measured as the degree of ablation [melting] of snow during winter) on numbers were both highly nonlinear: direct negative density dependence was found when the population was growing (Rt > 0) and during phases of the North Atlantic Oscillation (NAO) characterized by winters with generally high (1979-1995) and low (1996-2007) indices, respectively. A growth-phase-dependent model explained the dynamics of the population best and revealed the influence of density-independent processes on numbers that a linear autoregressive model missed altogether. In particular, the abundance of reindeer was enhanced by ablation during phases of growth (Rt > 0), an observation that contrasts with the view that periods of mild weather in winter are normally deleterious for reindeer owing to icing of the snowpack. Analyses of vital rates corroborated the nonlinearity described in the population time series and showed that both starvation mortality in winter and fecundity were nonlinearly related to fluctuations in density and the level of ablation. The erratic pattern of growth of the population of reindeer in Adventdalen seems, therefore, to result from a combination of the effects of nonlinear density dependence, strong density-dependent mortality, and variable density independence related to ablation in winter.     

Effects of Harvesting Flowers from Shrubs on the Persistence and Abundance of Wild Shrub Populations at Multiple Spatial Extents

Abstract: Wildflower harvesting is an economically important activity of which the ecological effects are poorly understood. We assessed how harvesting of flowers affects shrub persistence and abundance at multiple spatial extents. To this end, we built a process‐based model to examine the mean persistence and abundance of wild shrubs whose flowers are subject to harvest (serotinous Proteaceae in the South African Cape Floristic Region). First, we conducted a general sensitivity analysis of how harvesting affects persistence and abundance at nested spatial extents. For most spatial extents and combinations of demographic parameters, persistence and abundance of flowering shrubs decreased abruptly once harvesting rate exceeded a certain threshold. At larger extents, metapopulations supported higher harvesting rates before their persistence and abundance decreased, but persistence and abundance also decreased more abruptly due to harvesting than at smaller extents. This threshold rate of harvest varied with species’ dispersal ability, maximum reproductive rate, adult mortality, probability of extirpation or local extinction, strength of Allee effects, and carrying capacity. Moreover, spatial extent interacted with Allee effects and probability of extirpation because both these demographic properties affected the response of local populations to harvesting more strongly than they affected the response of metapopulations. Subsequently, we simulated the effects of harvesting on three Cape Floristic Region Proteaceae species and found that these species reacted differently to harvesting, but their persistence and abundance decreased at low rates of harvest. Our estimates of harvesting rates at maximum sustainable yield differed from those of previous investigations, perhaps because researchers used different estimates of demographic parameters, models of population dynamics, and spatial extent than we did. Good demographic knowledge and careful identification of the spatial extent of interest increases confidence in assessments and monitoring of the effects of harvesting. Our general sensitivity analysis improved understanding of harvesting effects on metapopulation dynamics and allowed qualitative assessment of the probability of extirpation of poorly studied species.     

Life history traits predict relative abundance in an assemblage of forest caterpillars

Species in a given trophic level occur in vastly unequal abundance, a pattern commonly documented but poorly explained for most taxa. Theoretical predictions of species density such as those arising from the metabolic theory of ecology hold well at large spatial and temporal scales but are not supported in many communities sampled at a relatively small scale. At these scales ecological factors may be more important than the inherent limits to energy use set by allometric scaling of mass. These factors include the amount of resources available, and the ability of individuals to convert these resources successfully into population growth. While previous studies have demonstrated the limits of macroecological theory in explaining local abundance, few studies have tested alternative generalized mechanisms determining abundance at the community scale. Using an assemblage of forest moth species found co-occurring as caterpillars on a single host plant species, we tested whether species abundance on that plant could be explained by mass allometry, intrinsic population growth, diet breadth, or some combination of these traits. We parameterized life history traits of the caterpillars in association with the host plant in both field and laboratory settings, so that the population growth estimate was specific to the plant on which abundance was measured. Using a generalized least-squares regression method incorporating phylogenetic relatedness, we found no relationship between abundance and mass but found that abundance was best explained by both intrinsic population growth rate and diet breadth. Species population growth potential was most affected by survivorship and larval development time on the host plant. Metabolic constraints may determine upper limits to local abundance levels for species, but local community abundance is strongly predicted by the potential for population increase and the resources available to that species in the environment.     

Effects of Forest Fragmentation on a Dung Beetle Community in French Guiana

Abstract:  Fragmentation is the most common disturbance induced by humans in tropical forests. Some insect groups are particularly suitable for studying the effects of fragmentation on animal communities because they are taxonomically and ecologically homogenous. We investigated the effects of forest fragmentation on a dung beetle species community in the forest archipelago created in 1994–1995 by the dam of Petit Saut, French Guiana. We set and baited an equal number of pitfall traps for dung beetles on three mainland sites and seven island sites. The sites ranged from 1.1 to 38 ha. In 250 trap days, we captured 50 species in 19 genera. Diversity indices were high (2.18–4.06). The lowest diversity was on the small islands and one mainland site. Species richness and abundance were positively related to fragment area but not to distance from mainland or distance to the larger island. The islands had lower species richness and population than mainland forest, but rarefied species richness was relatively invariant across sites. There was a marked change in species composition with decreasing fragment that was not caused by the presence of a common fauna of disturbed-area species on islands. Small islands differed from larger islands, which did not differ significantly from mainland sites. Partial correlation analyses suggested that species richness and abundance of dung beetle species were positively related to the number of species of nonflying mammals and the density index of howler monkeys ( Alouatta seniculus ), two parameters positively related to fragment area.