Food web structure and body size: trophic position and resource acquisition

Ulrich Brose Body size is recognized as an important determinant of trophic structure as it affects individual energetic demands, population density, and the interaction between potential prey and predators. However, its relationship with trophic position remains unclear. It has been hypothesized that a positive relationship between body size and trophic position would be associated to some particular trophic structures, which would allow larger organisms to satisfy their energetic demand and sustain viable population sizes at higher trophic positions, where fewer resources are available. To test this hypothesis, we analyzed the diet of 619 killifishes from four species (Austrolebias cheradophilus, A. luteoflammulatus, A. viarius and Cynopoecilus melanotaenia), collected in temporary ponds occurring in the grasslands of Rocha, Uruguay. Trophic position, diet richness, number of energy sources, and evenness were estimated for 20 size classes, formed by consecutive groups of 31 individuals. Gape limitation and preference for the larger available prey were evaluated as explanations for observed patterns with an individual based model (IBM). In agreement with the hypothesis, killifishes presented a strong positive relationship between trophic position and body size (R²=0.86), associated with a trophic structure that could allow larger organisms to have access to more energy from the environment. This was reflected in a positive relationship between body size and 1) prey richness, 2) number of basal energy sources (i.e. plants, detritus, phytoplankton and terrestrial prey), and 3) evenness in prey use. IBM results showed that changes in trophic structure with body size are well explained by gape limitation, but not by size preferences. Our results suggest that the fulfilment of the greater energetic demands of larger organism will depend on community diversity, which typically increases with ecosystem size, indicating a novel connection between area, diversity, body size, and food chain length.     

Differences in population dynamics and potential impacts of a freshwater invader driven by temporal habitat stability

Understanding population dynamics and population regulation of invasive species is critical for predicting their effects on native ecosystems as well as for control strategies. Many species of gastropod in the genus Pomacea are successful aquatic invaders that have caused economic and ecological impacts in Southeastern Asia where their large fecundity and broad reproductive window helps them to colonize and take advantages of ephemeral agricultural habitats. We followed the population dynamics of P. insularum in permanent, stable freshwater systems (ponds and streams), and in ephemeral agricultural habitats in the upper Texas Gulf Coast region, USA. We found that although P. insularum has a large reproductive potential, its density, biomass and size structure in stable permanent systems did not change significantly from March to November, and densities averaged <2 m⁻². This same species, however, displayed very different population dynamics in ephemeral agricultural environments. We found high densities (>130 m⁻²), and no stable size structure through time. Differences in the stability and persistence of these two types of environments appear to drive these patterns. Stability and persistence of habitats can result in different predator communities and the risk of predation for snails. We suggest that such factors may cause the differences in population dynamics and structure observed. The ability of snails to escape population control and explode in ephemeral habitats could drive the types of impacts seen on agricultural crops.     

Body Size Variations in Caribou Ecotypes and Relationships With Demography

ABSTRACT In many vertebrates size is one of the most influential and variable individual characteristics and a strong determinant of reproductive success. Body size is generally density dependent and decreases when intraspecific competition increases. Frequent and long-distance movements increase energy expenditures and, therefore, may also influence body size, particularly in highly mobile species. Caribou (Rangifer tarandus, also known as reindeer) exhibit tremendous variation in size and movements and thus represent an excellent candidate species to test the relationships between body size, population size, and movements. We analyzed body measurements of adult female caribou from 7 herds of the Québec-Labrador Peninsula, Canada, and we related their morphology to population size, movements, and annual ranges. The herds represented 3 ecotypes (migratory, montane, and sedentary). Ecotypes and herds differed in size (length), shape (roundness), and movements. The sedentary ecotype was larger and moved 4 to 7 times less than the migratory ecotype in the 1990s. At the start of a demographic growth period in the early 1960s, migratory caribou from the Rivière-George (hereafter George) herd had longer mandibles than caribou of the sedentary ecotype. Mandible length in the George herd declined in the 1980s after rapid population growth, while individuals performed extensive movements and the herd's annual range increased. Migratory caribou then became shorter than sedentary caribou. After the George herd decline in the 1990s, mandible length increased again near levels of the 1980s. Caribou from the migratory Rivière-aux-Feuilles herd later showed a similar decline in mandible length during a period of population growth, associated with longer movements and increasing annual range. We hypothesize that the density-dependent effect observed on body size might have been exerted through summer habitat degradation and movement variations during herd growth. Our study has 2 important implications for caribou management: the distinctiveness of different populations and ecotypes, and the correlations between population trajectories and changes in body condition and habitat.     

Time‐limited environments affect the evolution of egg–body size allometry

Initial offspring size is a fundamental component of absolute growth rate, where large offspring will reach a given adult body size faster than smaller offspring. Yet, our knowledge regarding the coevolution between offspring and adult size is limited. In time‐constrained environments, organisms need to reproduce at a high rate and reach a reproductive size quickly. To rapidly attain a large adult body size, we hypothesize that, in seasonal habitats, large species are bound to having a large initial size, and consequently, the evolution of egg size will be tightly matched to that of body size, compared to less time‐limited systems. We tested this hypothesis in killifishes, and found a significantly steeper allometric relationship between egg and body sizes in annual, compared to nonannual species. We also found higher rates of evolution of egg and body size in annual compared to nonannual species. Our results suggest that time‐constrained environments impose strong selection on rapidly reaching a species‐specific body size, and reproduce at a high rate, which in turn imposes constraints on the evolution of egg sizes. In combination, these distinct selection pressures result in different relationships between egg and body size among species in time‐constrained versus permanent habitats.     

Effects of environmental factors on the ecology and survival of a widespread,endemic Cerrado frog

Understanding the mechanisms that affect habitat use by vertebrates is critical for understanding how species are distributed across landscapes and how they cope with habitat change. The Brazilian Savanna (the Cerrado) has vegetation ranging from grassland to woodland savannas and harbors a rich and diverse amphibian fauna impacted by accelerated habitat loss. Here, we test the influence of vegetation type (from grassy scrubland to woodland) and distance from breeding sites (ephemeral water bodies) on body size, abundance, and survival of the frog Physalaemus nattereri in a natural metapopulation system of south-central Brazil. We also test whether body size is a significant predictor of population abundance. We found that the abundance of P. nattereri varies according to the mean snout–vent length of each metapopulation (sampling unit), as well as a higher estimated mortality rate in woodlands compared with typical Cerrado. Furthermore, we found no difference in estimated mortality among sampling units located far or close to ephemeral water bodies. Thus, our results highlight variable responses of P. nattereri metapopulations to environmental factors, despite the observed high heterogeneity among sampled habitats and the importance of ephemeral water bodies for reproduction. These findings highlight that land cover and availability of breeding sites might not always interact to explain population persistence of Cerrado frogs.     

How do invasive species affect native species? Experimental evidence from a carrion blowfly (Diptera: Calliphoridae) system

1. The necrobiome is a unique microcosm in which various organisms interact and compete for access to an ephemeral resource, such as carrion, that ultimately determines the structure and composition of these assemblages. 2. Blowfly species exhibit different competitive abilities which, when associated with other types of behaviour, such as predation or cannibalism, influence coexistence. Knowledge of the effects of competition between native and invasive species on development and survival is essential to understanding the dynamics of insect communities and to assess biological invasions. 3. Laboratory experiments were performed to evaluate the effect of interspecific competition on the bionomics and survival of a native (Cochliomyia macellaria) and an invasive (Chrysomya rufifacies) blowfly species at different population densities. 4. The deleterious effect of competition on the larval parameters of C. macellaria increased proportionally with increases in the larval density of C. rufifacies. When exposed to increased densities of C. rufifacies, larvae of C. macellaria accelerated their development and, as a trade‐off for this strategy, surviving adults were smaller and had reduced wing size, which were likely to reduce dispersal and reproductive capacity. 5. Larval competition – both as species‐dependent and density‐dependent phenomena – influences morphological and biological traits of surviving individuals. The impact of the invasive species has consequences at the population level, such as displacement or local population depletion of native species, a phenomenon likely to occur in other systems involving insects and ephemeral resources.     

Seasonal dynamics in community structure,abundance, body size and sex ratio in two species of Neotropical annual fishes

Seven ephemeral pools on the coastal plain of southern Brazil were found to be inhabited by three annual and 22 non‐annual fish species. Two common annual species (Austrolebias minuano and Cynopoecilus fulgens) exhibited clear seasonal dynamics, with the appearance of young fishes in the austral autumn (May to June) and a decline in abundance over the seasonal cycle. The third annual species, Austrolebias wolterstorffii, was rare. No seasonal dynamics were observed in non‐annual fishes. The relative abundance of non‐annual fishes compared with annual fishes increased over the seasonal cycle, but they coexisted widely. The size structure of annual fishes suggested the presence of a single age cohort in most pools though a second age cohort was registered in one pool in August, coinciding with a large flooding. Strong sexual dimorphism in body size was found in C. fulgens throughout the seasonal cycle, while no sexual dimorphism in body size was found in A. minuano. Female‐biased sex ratios were recorded in both common annual fish species in the last three sampling dates (in spring), but not during the first two sampling dates (in winter). The natural lifespan of annual fishes was <8 months. Annual fishes disappeared before habitat desiccation in half of the pools, while non‐annual fishes were still present.     

Demography of two lemming species on Bylot Island, Nunavut, Canada

Lemmings play a key role in the tundra food web and their widely reported cyclic oscillations in abundance may have a strong effect on other components of the ecosystem. We documented seasonal and annual variations in population density, reproductive activity, survival, and body mass of two sympatric species, the brown (Lemmus trimucronatus) and collared lemmings (Dicrostonyx groenlandicus), over a 2-year period on Bylot Island, Nunavut, Canada. We live trapped and marked lemmings on two grids throughout the summer and we estimated demographic parameters using three different capture–recapture methods. All three methods are based on robust estimators and they yielded similar population density estimates. The density of brown lemmings declined markedly between the 2 years whereas that of collared lemmings was relatively constant. For brown lemmings, 2004 was a peak year in their cycle and 2005 a decline phase. Density of brown lemmings also decreased during the summer, but not that of collared lemmings. The recruitment of juvenile brown lemmings in the population increased during the summer and was higher in the peak year than in the year after, but no change was detected in collared lemmings. Survival rates of both species tended to be lower during the peak year than in the following year and body mass of brown lemmings was higher in the peak year than in the following year. We conclude that both changes in adult survival and juvenile recruitment occur during the population decline of brown lemmings.     

The effect of population density on growth and survival of a Neotropical annual killifish

We used a field experiment to test the effects of population density on the growth rate and survival of Austrolebias bellottii, a Neotropical annual killifish. Effects differed between the sexes: males at high densities achieved a smaller final size and experienced higher mortality while no such effects were observed in females. This sex-specific effect could be an indirect consequence of mate competition.     

Variation in the population dynamics of the intertidal pulmonate gastropod Salinator solida Martens (Gastropoda: Amphibolidae) at Towra Point,NSW, Australia

This paper describes variation in the population dynamics of the intertidal pulmonate gastropod solida among mangrove and saltmarsh habitats at Towra Point, NSW, Australia over a two-year period. In general, this paper describes the degree to which the density of the populations fluctuated and the size structure of the populations varied among heights on shore. The density of individuals in the upper mangrove forest was most variable through time and their population size structure was dominated by smaller individuals. The density of populations in the Sarcocornia high on the shore fluctuated least through time and were dominated by large individuals. Those populations found on the mid-shore in the Sarcocornia had population characteristics intermediate to those populations in the upper mangrove forest and the Sarcocornia high on the shore. Analysis of size frequency data indicated that the growth rates of individuals and their annual mortality decreased with increasing height on shore.     

The effect of prey morphology on the feeding behaviour and population growth of the predatory rotifer Asplanchna sieboldi: a case study using five species of Brachionus (Rotifera)

1. We investigated the numerical response, functional response and prey preference of Asplanchna sieboldi to five different prey brachionids. We also analysed the feeding behaviour of the predator in terms of encounters, attacks, capture and prey ingested per unit time. 2. The five prey species (Brachionus havanaensis, B. rubens, B. patulus, B. macracanthus and B. calyciflorus) differed in their body size and spine length. 3. The population growth rates of A. sieboldi ranged from 0.074 ± 0.03 to 0.431 ± 0.02 depending on prey type and density. There was a significant impact of the spine length rather than body size per se on the population growth rates of the predator. 4. The maximum number of prey consumed depended on both body size and spine length. In the functional response analyses, the plateau was reached at a prey density of 4–8 ind. mL^?1. 5. There was a significant impact of prey density on the prey preference of the predator.     

The role of sediment type in growth and fecundity of mud snails (Hydrobiidae)

Summary We test the hypothesis that body size and population density of the deposit-feeding gastropod, Hydrobia truncata, are greater in muddy than in sandy habitats as a result of faster growth on fine- compared to coarse-grained sediments. We refute this hypothesis using a combination of field measurements and laboratory experiments. Three out of three populations tested had higher maximal growth rates and two of three populations approached their asymptotic size more quickly on sand than on silt-clay fractions of natural sediment. Growth decreased with increasing snail density and was as high or higher on sand as on silt-clay at all densities. Two populations were more fecund on sand than on silt-clay, and fecundity of the third population was not affected by sediment type. We show that the smaller body sizes observed in snails from the sandiest habitat result from late recruitment of these snails, relative to the other populations.     

Phenotypic plasticity in sex allocation for a simultaneously hermaphroditic coral reef fish

Phenotypic plasticity can facilitate reproductive strategies that maximize mating success in variable environments and lead to differences in sex allocation among populations. For simultaneous hermaphrodites with sperm competition, including Serranus tortugarum a small coral reef fish, proportional male allocation (testis in total gonad) is often greater where local density or mating group size is higher. We tested whether S. tortugarum reduced male allocation when transplanted from a higher density site to a lower density site. After 4 months, transplants mirrored the sex-allocation patterns of the resident population on their new reef. Transplants had significantly lower male allocation than representatives from their source population, largely as a result of reduced testis mass relative to body size.     

Determinants of size at metamorphosis in an endangered amphibian and their projected effects on population stability

Roughly 80% of animal species have complex life cycles spanning a major habitat shift, and delayed life history effects play an important role in their population dynamics. Through their effect on size at metamorphosis, factors in the pre‐ metamorphic environment often have profound effects upon survival and fecundity in the post‐metamorphic environment. Here, we adopted a combined experimental and field observational approach to investigate the factors that determine size at metamorphosis in pond‐breeding amphibians, and to predict some of their downstream effects on population stability. We set up ecologically realistic mesocosm communities for the endangered California tiger salamander Ambystoma californiense to test the effects of larval density, prey density and hydroperiod on mean size at metamorphosis. We found significant effects for all three factors, with mean size at metamorphosis negatively correlated with larval density and positively correlated with prey density and hydroperiod. We also used six years of field survey data to identify the most informative model explaining mean size at metamorphosis and thus validate our mesocosm results. The optimal three‐term model identified terms that were roughly analogous to each of the mesocosm treatments and with similar effect sizes, providing strong field confirmation of our experimental results. The field data also provide correlations between each factor and the number of metamorphs recruited to the population, allowing us to predict the effect of each factor on population stability. Finally, we show that these populations of the endangered A. californiense are strongly resource limited, which has important implications for their management and recovery as an endangered taxon.     

Population cycles and changes in body size of the lynx in Alaska

The lynx Lynx canadensis is a common predator in the boreal forests of North America. Its population fluctuates during a 9- to 11-year cycle in synchrony with the population size of its main prey, the snowshoe hare Lepus americanus. Using adult museum specimens, we studied changes in skull (and hence body) size of the lynx in Alaska during the second half of the 20th century. The population cycle in Alaska averaged 9 years, similar to that reported in the neighbouring Yukon. Using harvest data of lynx as an estimate of population size, we found that skull size was negatively related to population size. This relationship was strongest not for the population density in the year of death (X), but for year X-3, a carry-over effect from the first year (or years) of life, indicating that conditions during the fast-growth years are determining body size. We suggest that the density-dependent effect is probably due to changes in food supply, either resulting from the adverse effects of competition or a possible diminished availability of food. Two skull parameters decreased significantly during the second half of the 20th century. We do not know the cause for the year effect and suggest that it might be due to a long-term change in the availability of prey. Canine size did not change during the study period, probably an indication that snowshoe hares maintained their status as the main prey of the lynx throughout the study period. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Diseases and reproductive success in a wild mammal: example in the alpine chamois

Density-dependent and climatic factors affect reproduction and dynamics of wild ungulates. Parasites can also decrease reproductive success through either a direct abortive effect or a negative impact on host growth and body condition. However, few studies have investigated the effect of parasitism on fecundity of ungulates in natural conditions. We studied three bacterial infections caused by Salmonella enterica serovar Abortusovis, Chlamydophila abortus and Coxiella burnetii. These bacteria are leading causes of reproductive failure in sheep, goat and cattle, which raises the question of their influence on population dynamics of wild ungulates. A long-term study of demography and epidemiology of an alpine chamois (Rupicapra rupicapra, L.) population (Les Bauges Reserve, France) and a generalized linear modeling approach were used to analyze the reproductive success of chamois according to population density, weather conditions and the prevalence of antibodies against the three bacteria in females. This approach enabled us to identify the confounding effect of weather and parasitism on fecundity in a natural population. After accounting for density, the prevalence of antibodies against the three bacteria explained 36% of the annual variation in reproductive success, and weather conditions explained an additional 31%. This study was, to our knowledge, the first to compare the decrease in fecundity due to bacterial infections and weather conditions in a population of wild mountain ungulates.     

Mycophagy and its influence on habitat use and ranging patterns in Callimico goeldii

Mycophagy has been documented in a number of species of marmosets and lion tamarins (Callitrichinae) but its effect on ranging behavior is not known. We present the results of 10 years of research on five groups of Goeldi's monkey (Callimico goeldii) at a field site in northwestern Bolivia. We studied the diet and ranging behavior of two of the groups. On average, groups contained 4.5 individuals (range 2.0–9.0), but they gradually decreased in size until only the breeding female remained in the home range. The annual diet was composed of fungi (31.1–34.9%), fruits (34.0–40.6%), prey (17.4–30.1%), and exudates (1.0–10.9%). They had large home ranges (114–150 ha) and over time individuals tended to shift their core areas of use. They used secondary and bamboo forest and forest with dense understories more than expected based on availability. We suggest that the large home ranges and shifting core areas used by C. goeldii are components of a foraging strategy to track patchy, low density, and ephemeral fungal fruiting bodies. Our results, along with data published on other callitrichines, indicate that groups of Leontopithecus, Callithrix, and Callimico that eat fungi have larger home ranges than those that do not. Mycophagy is one of the several factors that evidently affect home range size in callitrichines. Fungi are clearly an important food source for a number of populations, but additional studies are needed to determine why some eat fungi frequently while others do not. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

What if it gets crowded? Density‐dependent tortuosity in individual movements of a Neotropical mammal

Effects of density dependence on animal movements have received much attention in ecology, but it is still debated to what extent dispersal and movements in general are density dependent, and their potential contribution to population regulation processes. Here, we determine the occurrence and nature of density dependence in the movements of a Neotropical marsupial, the black‐eared opossum Didelphis aurita Wied‐Neuwied 1826. Using spool‐and‐line tracking devices, we estimated the tortuosity of fine‐scale movements of 149 individuals by their fractal dimension D. We evaluated the relative importance of population size, reproductive or climatic seasons and reproductive maturity of individuals as determinants of movement tortuosity, using a model selection approach. Population size was the most important determinant of movement tortuosity, with season (climatic seasons for females, reproductive seasons for males) and reproductive maturity as secondary but also important variables. We detected a positive density‐dependent effect on movement tortuosity, resulting in more intensive use of areas by individuals during periods of high population size. This positive association between movement tortuosity and population size is more likely to result from intraspecific competition, which forces individuals to explore their environment more intensively during high‐density periods. Therefore, despite being density dependent, movements in D. aurita apparently do not contribute to population regulation mechanisms.     

Microspatial heterogeneity in the distribution of ciliates in a small pond

Five transects of contiguous samples from the surface of a small pond and one transect from its bottom were collected in order to quantify microspatial heterogeneity in the distribution of ciliated protozoa. Examination of the frequency-abundance relations for these transects suggests that they can be approximated by negative binomial distributions with a commonk of 1.87. Contagiousness or crowding increases with population density.Mean patch size and mean interpatch distance were measured for 4 transects as 1.5 to 2 cm and 3 to 4 cm, respectively. This heterogeneity is suggested to arise from behavioral aggregation about discrete food sources and be very ephemeral.Blocking of adjacent contiguous samples was used to investigate the effect of sample size on the apparent correlation between the numbers of pairs of taxa. In all cases examined, taxa were relatively independent in their distribution at small sample sizes and became more negatively or positively associated as samples were combined. This may reflect that the small scale patches are essentially monospecific.     

Filling the gaps: phylogeography of the self-fertilizing Kryptolebias species (Cyprinodontiformes: Rivulidae) along South American mangroves

Mangrove killifishes of the genus Kryptolebias have been historically classified as rare because of their small size and cryptic nature. Major gaps in distribution knowledge across mangrove areas, particularly in South America, challenge the understanding of the taxonomic status, biogeographical patterns and genetic structuring of the lineages composing the self-fertilizing “Kryptolebias marmoratus species complex.” In this study, the authors combined a literature survey, fieldwork and molecular data to fill major gaps of information about the distribution of mangrove killifishes across western Atlantic mangroves. They found that selfing mangrove killifishes are ubiquitously distributed across the Caribbean, Central and South American mangroves and report 14 new locations in South America, extending the range of both the “Central clade” and “Southern clade” lineages which overlap in the Amazon. Although substantial genetic differences were found between clades, the authors also found further genetic structuring within clades, with populations in Central America, north and northeast Brazil generally showing higher levels of genetic diversity compared to the clonal ones in southeast Brazil. The authors discuss the taxonomic status and update the geographical distribution of the Central and Southern clades, as well as potential dispersal routes and biogeographical barriers influencing the distribution of the selfing mangrove killifishes in the western Atlantic mangroves.