Zooplankton community size structure and taxonomic composition affects size-selective grazing in natural communities

The body size of an individual zooplankton is well related to its grazing rate and to the range of particle sizes it can ingest, and since cladocerans and copepods feed differently, they follow different relationships. Based on these general patterns in individual organisms, we tested whether the size structure and taxonomic composition of more complex natural zooplankton communities are related to their in situ grazing rate and to the range of algal sizes they graze. We compared community grazing rates on individual algal taxa in two communities dominated by small cladocerans, three communities dominated by large cladocerans and three copepod-dominated communities. Small algae were usually grazed most intensively, but grazing rates were poorly related to algal size alone. The range in size of grazed algae increased with increasing mean zooplankton body size, but differed systematically with their taxonomic composition. Communities dominated by Ceriodaphnia or Holopedium grazed a narrower size range of algae [maximum greatest axial length dimension (GALD)=16–36 μm)] than communities with large biomasses of Bosmina or Daphnia (maximum GALD=28–78 μm). Copepod-dominated communities followed the same general relationship as cladocerans. Daphnia-dominated communities grazed the broadest range of algal sizes, and their total grazing rates were up to 2.4 times their grazing rates on small (<35 μm) “highly edible” algae, a difference of similar magnitude to those found in successful trophic cascade biomanipulations. Received: 31 March 1998 / Accepted: 19 October 1998     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Zooplankton community structure in southwestern Quebec lakes: the rOles of acidity and predation

The zooplankton community structure of 22 lakes with varyingacidity and fish biomass, located southwest of the mining andsmelting region of Rouyn/Noranda, Quebec, was examined in July1987. Lakes with dominant piscivorous fish communities couldbe discriminated from non-piscivorous communities using solelypH and Chaoborus abundance, demonstrating that modificationsin the zooplankton community occurred concurrently with changesin the fUh communities. Strong discontinuities in zooplanktonspecies distributions were discerned in lakes with pH values<5.3; abundance ranges for several species could be semi-quantitativelyrelated to lake morpho-metry or chemistry. Small shallow kettlelakes with elevated heat budgets had high biomasses of smallherbivorous organisms. The presence of >5 mg m^–3 ofadvanced Chaoborus instars (III and IV) was associated withreduced microcrustacean biomass in many lakes, particularlythose with low fish biomass. There was little evidence for size-selectivepredation by fish in these oligotrophic lakes. Fish biomass/effortcould be semi-quantitatively related to the biomass of Leptodorakindtii and the ratio of adult Diaptomus nunutus to copepoditestage IV.     

Ungulate community structure and ecological processes: body size,hoof area and trampling in African savannas

A wide range of bioenergetic, production, life history and ecological traits scale with body size in vertebrates. However, the consequences of differences in community body-size structure for ecological processes have not been explored. We studied the scaling relationships between body mass, shoulder height, hoof area, stride length and daily ranging distance in African ungulates ranging in size from the 5 kg dik-dik to the 5,000 kg African elephant, and the implications of these relationships on the area trampled by single and multispecies herbivore communities of differing structure. Hoof area, shoulder height and stride length were strongly correlated with body mass (Pearson's r >0.98, 0.95 and 0.90, respectively). Hoof area scaled linearly to body mass with a slope of unity, implying that the pressures exerted on the ground per unit area by a small antelope and an elephant are identical. Shoulder height and stride length scaled to body mass with similar slopes of 0.32 and 0.26, respectively; larger herbivores have relatively shorter legs and take relatively shorter steps than small herbivores, and so trample a greater area of ground per unit distance travelled. We compared several real and hypothetical single- and multi-species ungulate communities using exponents of between 0.1 and 0.5 for the body mass to daily ranging distance relationship and found that the estimated area trampled was greater in communities dominated by larger animals. The impacts of large herbivores are not limited to trampling. Questions about the ecological implications of community body-size structure for such variables as foraging and food intake, dung quality and deposition rates, methane production, and daily travelling distances remain clear research priorities.     

Zooplankton community structure and environmental conditions in a set of interconnected ponds

We studied the zooplankton community structure in a set of 33 interconnected shallow ponds that are restricted to a relatively small area (`De Maten', Genk, Belgium, 200 ha). As the ponds share the same water source, geology and history, and as the ponds are interconnected (reducing chance effects of dispersal with colonisation), differences in zooplankton community structure can be attributed to local biotic and abiotic interactions. We studied zooplankton community, biotic (phytoplankton, macrophyte cover, fish densities, macroinvertebrate densities), abiotic (turbidity, nutrient concentrations, pH, conductivity, iron concentration) and morphometric (depth, area, perimeter) characteristics of the different ponds. Our results indicate that the ponds differ substantially in their zooplankton community structure, and that these differences are strongly related to differences in trophic structure and biotic interactions, in concordance with the theory of alternative equilibria. Ponds in the clear-water state are characterised by large Daphnia species and species associated with the littoral zone, low chlorophyll-a concentrations, low fish densities and high macroinvertebrate densities. Ponds in the turbid-water state are characterised by high abundances of rotifers, cyclopoid copepods and the opposite environmental conditions. Some ponds show an intermediate pattern, with a dominance of small Daphnia species. Our results show that interconnected ponds may differ strongly in zooplankton community composition, and that these differences are related to differences in predation intensity (top-down) and habitat diversity (macrophyte cover).     

Zooplankton biomass and community structure in a Danube River floodplain system: effects of hydrology

1. Zooplankton density and biomass was examined in a Danube River floodplain section with highly variable hydrological dynamics. Temporal patterns were analysed to assess the effects of hydrological conditions on zooplankton community structure and the differential response of the two major zooplankton taxa, rotifers and crustaceans.
2. Calculated floodplain water age was used as an integrated parameter describing hydrological conditions and connectivity.
3. Total zooplankton biomass, crustacean biomass and crustacean species number were significantly positively related to water age. Rotifer biomass followed a hump-shaped relationship with water age, and rotifer species number decreased with increasing water age.
4. Rotifers dominated the community in periods of low to medium water ages. In periods of higher water ages the community was dominated by crustaceans.
5. We propose that the hydrological regime of floodplains is crucial for zooplankton biomass patterns and succession, through the alternation of washing-out effects, taxon-specific potential of reproduction and biological interactions. Flood events and high water levels reset the community to an early successional phase.     

Zooplankton spatial patterns in two lakes with contrasting fish community structure

Horizontal and temporal patterns in crustacean zooplankton communities were analyzed in two small, oligotrophic lakes which were morphologically and chemically similar, but had contrasting fish communities. Ranger Lake was dominated by two bass species and the planktivores numbered < 25 ind. ha^–1. Mouse Lake had no large piscivores and planktivores numbered > 1200 ind. ha^–1. There were significant differences in the distribution of zooplankton taxa and size classes between sampling stations. In Ranger Lake, the smallest size classes were more abundant at the deeper stations and the larger individuals were more abundant at the shallower stations. In Mouse Lake, the smaller individuals were more common at the shallow stations and the larger individuals were more common at the deeper stations. These differences suggest medium scale patterns induced by vectorial forces, but modified by species specific migration patterns. We tested the hypothesis that horizontal heterogeneity should be influenced by planktivore density and found that none of the taxa showed significant between-lake differences in the variance-mean regressions. We also tested the hypothesis that larger taxa should be more heterogeneous and we found that cladocerans were more heterogeneous than copepods and nauplii. In terms of sampling methodology our data suggest that the between-station variability was so high that a single mid-lake sample would certainly lead to completely unacceptable errors in the estimation of population densities and biomasses.     

Warming alters community size structure and ecosystem functioning

Global warming can affect all levels of biological complexity, though we currently understand least about its potential impact on communities and ecosystems. At the ecosystem level, warming has the capacity to alter the structure of communities and the rates of key ecosystem processes they mediate. Here we assessed the effects of a 4°C rise in temperature on the size structure and taxonomic composition of benthic communities in aquatic mesocosms, and the rates of detrital decomposition they mediated. Warming had no effect on biodiversity, but altered community size structure in two ways. In spring, warmer systems exhibited steeper size spectra driven by declines in total community biomass and the proportion of large organisms. By contrast, in autumn, warmer systems had shallower size spectra driven by elevated total community biomass and a greater proportion of large organisms. Community-level shifts were mirrored by changes in decomposition rates. Temperature-corrected microbial and macrofaunal decomposition rates reflected the shifts in community structure and were strongly correlated with biomass across mesocosms. Our study demonstrates that the 4°C rise in temperature expected by the end of the century has the potential to alter the structure and functioning of aquatic ecosystems profoundly, as well as the intimate linkages between these levels of ecological organization.     

Abundance, diversity and body size: patterns from a grassland arthropod community

1. The empirical relationships among body size, species richness and number of individuals may give insight into the factors controlling species diversity and the relative abundances of species. To determine these relationships, we sampled the arthropods of grasslands and savannahs at Cedar Creek, MN using sweep nets (90 525 individuals of 1225 species) and pitfall traps (12 721 individuals of 92 species). Specimens were identified, enumerated and measured to determine body size.
2. Both overall and within abundant taxonomic orders, species richness and numbers of individuals peaked at body sizes intermediate for each group. Evolution could create unimodal diversity patterns by random diversification around an ancestral body size or from size-dependent fitness differences. Local processes such as competition or predation could also create unimodal diversity distributions.
3. The average body size of a species depended significantly on its taxonomic order, but on contemporary trophic role only within the context of taxonomic order.
4. Species richness ( S _i) within size classes was related to the number of individuals ( I _i) as S _i =  I i^0·5. This relationship held across a 100 000-fold range of body sizes. Within size classes, abundance distributions of size classes were all similar power functions. A general rule of resource division, together with similar minimum population sizes, is sufficient to generate the relationship between species richness and number of individuals.
5. Smaller bodied species had slightly shallower abundance distributions and may, in general, persist at lower densities than larger species.
6. Our results suggest there may be fewer undescribed small arthropod species than previously thought and that most undescribed species will be smaller than arthropods.     

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.     

Wave stress and coral community structure in Hawaii

Summary The most significant factor determining the structure of Hawaiian reef coral communities is physical disturbance from waves. Sequential analysis of community structure off the west coast of the island of Hawaii shows that variation of wave energy and storm frequency clearly affects organization in time and space. Normal conditions of low wave stress maintain four well-defined reef zones; diversity is highest at intermediate depths and decreases in physically rigorous shallow areas and stable deep reef slopes. Intermediate level storm wave events cause variable effects within the reef zones, but the zonation pattern, as a whole, is maintained. Diversity increases in zones that are dominated by a single species largely through nonlethal fragmentation and transport, but decreases in the zone of most equitable species distribution. Conversely, severe infrequent storm disturbances that cause massive mortality to all coral species wipe out the pattern of community structure and return the entire community to a low diversity early successional stage.Hawaii Institute of Marine Biology Contribution No.616     

Connecting landscape structure and patterns in body size distributions

Understanding the interaction between community structure and landscape structure represents a pressing theoretical challenge of great applied importance considering the increasing structural modification of ecosystems through habitat loss and fragmentation. Dispersal ability and energetic demands coupled to body size determine the landscape structure experienced by an organism, which could essentially be fragmented for small individuals but continuous for large ones. Although discontinuities in species assemblages have been predicted and detected, no explicit association between habitat structure and body size distributions has been demonstrated. In this contribution, we propose that body size structure in local communities should reflect such different perceptions of landscape structure. To this end, we explore this association in a simple metacommunity located in the Atacama Desert, in northern Chile. Using graph theory we found that species of different size and trophic position (carnivores and herbivores) perceive the landscape at contrasting spatial scales. In each community (n = 31) we determined the observed and the expected body size distributions – in a random sample from the metacommunity of 18 727 individuals –, which allowed us to identify the body sizes at which an overrepresentation or underrepresentation of individuals occur. Such aggregations and discontinuities in body sizes were related, for carnivores, to patch location within the landscape, and to the internal banded vegetation pattern within patches for herbivores. Our study shows, for the first time, an empirical connection between the spatial distribution of communities, their local attributes, and the existence and locations of discontinuities and aggregations in body size distributions.     

Problems of body structure and size of the hypophysis.

The size of the hypophysis, especially of the anterior lobe, is related to body size while the nervous lobe is related to the brain weight. The anterior lobe of the hypophysis in domestic animals is smaller than that of their ancestors in the wild. The proportions of the body, the distribution of bulk, as they appear in different races--whippets and bulldogs--are related to the size of the anterior lobe. Same body weight implies that the anterior lobe is nearly twice as large in animals with a compact body structure. These intraspecific results can also be shown interspecificly by comparing the species of African Bovidae. Independent of the influence of body size, the subfamilies Alcelaphinae and Hippotraginae possess a larger anterior lobe. A tendency can be shown for a relation to body structure as short-legged species living on marshy grounds (Kobus) or soft sands (Addax) have larger anterior lobes.     

Phytoplankton size distribution and community structure: influence of nutrient input and sedimentary loss

This study examines the effects of nutrient availability andsedimentary loss on the dynamics of freshwater phytoplankton,from the physiological state at the cellular level to the competitionissue at the community level. We studied the separated and combinedresponses to nutrient inputs and mixing, of a phytoplanktoncommunity, in terms of size fractionated photosynthetic activityand biomass, exported biomass and species composition. The communitywas composed of two dominant species, which differed in theirsinking velocity. Experiments were conducted during 8 days in12 water columns of 50 L, in light and temperature controlledconditions. The nutrient input was found to be the predominantfactor affecting sedimentary fluxes and community composition,but mixing had also a significant effect on the community structureand sedimentary fluxes when combined with nutrient inputs. Inthis latter case, the large sinking species, Tribonema sp. (Xanthophyceae),developed, due to both a decrease of sedimentary loss by upwardadvection and the enhanced growth from nutrient inputs, as indicatedby the higher photosynthetic activity measured using chlorophylla in vivo fluorescence. The combined effect of both factorsreduced the difference between growth and loss rates and increasedthe viability of the sinking populations.     

Shape and size of the body vs. musculoskeletal stress markers

The objective of this paper is to assess the relationship between the degree of development of muscle attachment sites (musculoskeletal stress markers - MSM1) and the length and circumference measurements of long bones and the body build expressed with the reconstructed values of body height (BH) and body mass (BM). The bone material (102 male and 99 female skeletons) used in the study was collected in the medieval burial ground in Cedynia, Poland. The authors analyzed 10 musculoskeletal stress markers located on the scapula (2), humerus (2), radius (2), femur (2) and tibia (2). The frequency and the degree of expression of muscle attachment size was carried out using the scale prepared by Myszka (2007). The scale encompassed three degrees of expression of muscle attachment size. Only changes of robusticity type (nonpathological changes) were taken into account. The assessment of body build of individuals was carried out according to the method proposed by Vancata & Charvátová (2001). Body height was reconstructed from the length of the humerus and femur using eight equations. Body mass was reconstructed from the measurements of the breadth of the proximal and distal sections of the femur and tibia (mechanical method) using twenty one equations. The equations were developed for different reference populations. The same equations were used for men and women. The correlation between the MSM and the length and circumference measurements of the bones was analyzed using the principal components analysis and the Gamma correlation coefficient. The strength of the correlation between the reconstructed body build traits (BH, BM) and the moderate degree of musculoskeletal stress markers expression was studied based on the principal components method and the Pearson correlation coefficient. A linear correlation was found between musculoskeletal stress markers and the circumference measurements and the reconstructed body mass, but no relationship with body height and the length measurements of long bones was revealed. From previous research it is evident that the relationship between the MSM and metric skeletal traits does not occur in every population. Divergent findings necessitate further corroboration of results on diverse skeletal material.     

Animal community structure as a function of stream size

The species-area relationship of the island biogeography theory was calculated for macroinvertebrates in 22 coastal, adjacent streams. A z-value of 0.19 was obtained. The low z-value was probably a consequence of the short distances between streams as well as high dispersal rates. In addition, a cluster analysis based on the dissimilarity of species assemblages showed that stream size was of prime importance in categorizing the streams. To a smaller extent water quality affected the community structure in the streams.     

Effects of body size and temperature on population growth

For at least 200 years, since the time of Malthus, population growth has been recognized as providing a critical link between the performance of individual organisms and the ecology and evolution of species. We present a theory that shows how the intrinsic rate of exponential population growth, rmax, and the carrying capacity, K, depend on individual metabolic rate and resource supply rate. To do this, we construct equations for the metabolic rates of entire populations by summing over individuals, and then we combine these population-level equations with Malthusian growth. Thus, the theory makes explicit the relationship between rates of resource supply in the environment and rates of production of new biomass and individuals. These individual-level and population-level processes are inextricably linked because metabolism sets both the demand for environmental resources and the resource allocation to survival, growth, and reproduction. We use the theory to make explicit how and why rmax exhibits its characteristic dependence on body size and temperature. Data for aerobic eukaryotes, including algae, protists, insects, zooplankton, fishes, and mammals, support these predicted scalings for rmax. The metabolic flux of energy and materials also dictates that the carrying capacity or equilibrium density of populations should decrease with increasing body size and increasing temperature. Finally, we argue that body mass and body temperature, through their effects on metabolic rate, can explain most of the variation in fecundity and mortality rates. Data for marine fishes in the field support these predictions for instantaneous rates of mortality. This theory links the rates of metabolism and resource use of individuals to life-history attributes and population dynamics for a broad assortment of organisms, from unicellular organisms to mammals.