Light heterogeneity affects understory plant species richness in temperate forests supporting the heterogeneity–diversity hypothesis

One of the most important drivers for the coexistence of plant species is the resource heterogeneity of a certain environment, and several studies in different ecosystems have supported this resource heterogeneity–diversity hypothesis. However, to date, only a few studies have measured heterogeneity of light and soil resources below forest canopies to investigate their influence on understory plant species richness. Here, we aim to determine (1) the influence of forest stand structural complexity on the heterogeneity of light and soil resources below the forest canopy and (2) whether heterogeneity of resources increases understory plant species richness. Measures of stand structural complexity were obtained through inventories and remote sensing techniques in 135 1‐ha study plots of temperate forests, established along a gradient of forest structural complexity. We measured light intensity and soil chemical properties on six 25 m² subplots on each of these 135 plots and surveyed understory vegetation. We calculated the coefficient of variation of light and soil parameters to obtain measures of resource heterogeneity and determined understory plant species richness at plot level. Spatial heterogeneity of light and of soil pH increased with higher stand structural complexity, although heterogeneity of soil pH did not increase in conditions of generally high levels of light availability. Increasing light heterogeneity was also associated with increasing understory plant species richness. However, light heterogeneity had no such effects in conditions where soil resource heterogeneity (variation in soil C:N ratios) was low. Our results support the resource heterogeneity–diversity hypothesis for temperate forest understory at the stand scale. Our results also highlight the importance of interaction effects between the heterogeneity of both light and soil resources in determining plant species richness.     

Nematode Spatial and Ecological Patterns from Tropical and Temperate Rainforests

Large scale diversity patterns are well established for terrestrial macrobiota (e.g. plants and vertebrates), but not for microscopic organisms (e.g. nematodes). Due to small size, high abundance, and extensive dispersal, microbiota are assumed to exhibit cosmopolitan distributions with no biogeographical patterns. This assumption has been extrapolated from local spatial scale studies of a few taxonomic groups utilizing morphological approaches. Recent molecularly-based studies, however, suggest something quite opposite. Nematodes are the most abundant metazoans on earth, but their diversity patterns are largely unknown. We conducted a survey of nematode diversity within three vertical strata (soil, litter, and canopy) of rainforests at two contrasting latitudes in the North American meridian (temperate: the Olympic National Forest, WA, U.S.A and tropical: La Selva Biological Station, Costa Rica) using standardized sampling designs and sample processing protocols. To describe nematode diversity, we applied an ecometagenetic approach using 454 pyrosequencing. We observed that: 1) nematode communities were unique without even a single common species between the two rainforests, 2) nematode communities were unique among habitats in both rainforests, 3) total species richness was 300% more in the tropical than in the temperate rainforest, 4) 80% of the species in the temperate rainforest resided in the soil, whereas only 20% in the tropics, 5) more than 90% of identified species were novel. Overall, our data provided no support for cosmopolitanism at both local (habitats) and large (rainforests) spatial scales. In addition, our data indicated that biogeographical patterns typical of macrobiota also exist for microbiota.     

Comparative patterns in taxonomic and functional spider diversities between tropical vs. temperate forests

High diversity in tropical compared to temperate regions has long intrigued ecologists, especially for highly speciose taxa like terrestrial arthropods in tropical rainforests. Previous studies showed that arthropod herbivores account for much tropical diversity, yet differences in the diversity of predatory arthropods between tropical and temperate systems have not been properly quantified. Here, we present the first standardized tropical–temperate forest quantification of spider diversities, a dominant and mega‐diverse taxon of generalist predators. Spider assemblages were collected using a spatially replicated protocol including two standardized sampling methods (vegetation sweep netting and beating). Fieldwork took place between 2010 and 2015 in metropolitan (Brittany) and overseas (French Guiana) French territories. We found no significant difference in functional diversity based on hunting guilds between temperate and tropical forests, while species richness was 13–82 times higher in tropical versus temperate forests. Evenness was also higher, with tropical assemblages up to 55 times more even than assemblages in temperate forests. These differences in diversity far surpass previous estimates and exceed tropical–temperate ratios for herbivorous taxa.     

Diversity of mammals in the tropical–temperate Neotropics: hotspots on a regional scale

The tropical–temperate interface of the southern Neotropics harbours an interdigitating array of biomes (Puna, Monte, Chaco, Yungas). This topographic and climatically complex region needs urgent conservation efforts, as it is being transformed by human activities at an accelerating pace. We analyse georeferenced field records of mammal species in northwestern Argentina (provinces: Catamarca, Jujuy, Salta, Tucuman) in order to define biodiversity hotspots on the basis of 0.5°× 0.5° grid cells within northwestern Argentina according to total richness of mammal species, richness of megaspecies (species above 10 kg), and endemic species (species restricted to Argentina or neighbouring countries with shared biomes). The mammal fauna of northwestern Argentina is fairly well known (176 species). The biomes differ considerably in species richness (Puna low, Yungas high) and species composition. We found no significant difference between endemic and non-endemic species regarding cell occupancy or body size. Cell occupancy was not correlated to body size. Across grids, species richness, number of megaspecies as well as richness of endemics are all correlated to sampling effort. More than 50% of the species in the region are restricted to one or two biomes. Overall, the species turn-over between biomes in northwestern Argentina is high. Using a simple algorithm we identified 10 grid cells which covered 90% of the total number of recorded species, and contrast them with the protected areas. While the Puna and Yungas biomes are rather well protected, the arid and semiarid Monte and Chaco are in need of urgent attention in biodiversity conservation.     

Ecometagenetics confirm high tropical rainforest nematode diversity

The general patterns of increasing biodiversity from the poles to the equator have been well documented for large terrestrial organisms such as plants and vertebrates but are largely unknown for microbiota. In contrast to macrobiota, microbiota have long been assumed to exhibit cosmopolitan, random distributions and a lack of spatial patterns. To evaluate the assumption, we conducted a survey of nematode diversity within the soil, litter and canopy habitats of the humid lowland tropical rainforest of Costa Rica using an ultrasequencing ecometagenetic approach at a species-equivalent taxonomic level. Our data indicate that both richness and diversity of nematode communities in the tropical rainforests of Costa Rica are high and exceed observed values from temperate ecosystems. The majority of nematode species were unknown to science, providing evidence for the presence of highly endemic (not cosmopolitan) species of still completely undiscovered biodiversity. Most importantly, the greater taxonomic resolution used here allowed us to reveal predictable habitat associations for specific taxa and thus gain insights into their nonrandom distribution patterns.     

Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

Global patterns of plant diversity

Summary Using 94 data sets from across the globe, we explored patterns of mean community species richness, landscape species richness, mean similarity among communities and mosaic diversity. Climate affected community species richness primarily through productivity while other climatic factors were secondary. Climatic equability affected species richness only in temperate regions where richness was greatest at high levels of temperature variability and low levels of precipitation variability. Landscape species richness correlated positively with community species richness. A global gradient in mean similarity existed but was uncorrelated with community species richness. Mean similarity was least and mosaic diversity was greatest between 25 and 30° latitude. The most diverse landscapes (low mean similarity) correlated with warm temperatures, high elevations, large areas and large seasonal temperature fluctuations. The most complex landscapes (high mosaic diversity) correlated with large areas, high productivity and warm winters. We compared diversity measures among continents and found only one significant difference: Australian landscapes have greater mosaic diversity than African landscapes. Based on our analyses we propose two hypotheses: (1) for plants, biotic interactions are more important in structuring landscapes in warmer climates and (2) longer isolated landscapes have more clearly differentiated ecological subunits.     

Tropical nematode diversity: vertical stratification of nematode communities in a Costa Rican humid lowland rainforest

Comparisons of nematode communities among ecosystems have indicated that, unlike many organisms, nematode communities have less diversity in the tropics than in temperate ecosystems. There are, however, few studies of tropical nematode diversity on which to base conclusions of global patterns of diversity. This study reports an attempt to estimate nematode diversity in the lowland tropical rainforest of La Selva Biological Research Station in Costa Rica. We suggest one reason that previous estimates of tropical nematode diversity were low is because habitats above the mineral soil are seldom sampled. As much as 62% of the overall genetic diversity, measured by an 18S ribosomal barcode, existed in litter and understorey habitats and not in soil. A maximum-likelihood tree of barcodes from 360 individual nematodes indicated most major terrestrial nematode lineages were represented in the samples. Estimated 'species' richness ranged from 464 to 502 within the four 40 × 40 m plots. Directed sampling of insects and their associated nematodes produced a second set of barcodes that were not recovered by habitat sampling, yet may constitute a major class of tropical nematode diversity. While the generation of novel nematode barcodes proved relatively easy, their identity remains obscure due to deficiencies in existing taxonomic databases. Specimens of Criconematina, a monophyletic group of soil-dwelling plant-parasitic nematodes were examined in detail to assess the steps necessary for associating barcodes with nominal species. Our results highlight the difficulties associated with studying poorly understood organisms in an understudied ecosystem using a destructive (i.e. barcode) sampling method.     

The challenge of accurately documenting bee species richness in agroecosystems: bee diversity in eastern apple orchards

Bees are important pollinators of agricultural crops, and bee diversity has been shown to be closely associated with pollination, a valuable ecosystem service. Higher functional diversity and species richness of bees have been shown to lead to higher crop yield. Bees simultaneously represent a mega‐diverse taxon that is extremely challenging to sample thoroughly and an important group to understand because of pollination services. We sampled bees visiting apple blossoms in 28 orchards over 6 years. We used species rarefaction analyses to test for the completeness of sampling and the relationship between species richness and sampling effort, orchard size, and percent agriculture in the surrounding landscape. We performed more than 190 h of sampling, collecting 11,219 specimens representing 104 species. Despite the sampling intensity, we captured <75% of expected species richness at more than half of the sites. For most of these, the variation in bee community composition between years was greater than among sites. Species richness was influenced by percent agriculture, orchard size, and sampling effort, but we found no factors explaining the difference between observed and expected species richness. Competition between honeybees and wild bees did not appear to be a factor, as we found no correlation between honeybee and wild bee abundance. Our study shows that the pollinator fauna of agroecosystems can be diverse and challenging to thoroughly sample. We demonstrate that there is high temporal variation in community composition and that sites vary widely in the sampling effort required to fully describe their diversity. In order to maximize pollination services provided by wild bee species, we must first accurately estimate species richness. For researchers interested in providing this estimate, we recommend multiyear studies and rarefaction analyses to quantify the gap between observed and expected species richness.     

Estimating species richness and catch per unit effort from boat electro‐fishing in a lowland river in temperate Australia

Abstract Biodiversity estimates are typically a function of sampling effort and in this regard it is important to develop an understanding of taxon‐specific sampling requirements. Northern hemisphere studies have shown that estimates of riverine fish diversity are related to sampling effort, but such studies are lacking in the southern hemisphere. We used a dataset obtained from boat electro‐fishing the fish community along an essentially continuous 13‐km reach of the Murrumbidgee River, Australia, to investigate sampling effort effects on fish diversity estimates. This represents the first attempt to investigate relationships between sampling effort and the detection of fish species in a large lowland river in Australia. Seven species were recorded. Species‐specific patterns in catch per unit effort were evident and are discussed in terms of solitary and gregarious species, recreational fishing and the monitoring of rare and threatened species. There was a requirement to sample substantial lengths of river to describe total species richness of the fish community in this river reach. To this end, randomly allocated sampling effort and use of species richness estimators produced accurate estimates of species richness without the requirement for excessive levels of effort. Twenty operations were required to estimate species richness at this site, highlighting the need for comparable studies of river fish communities in lowland rivers elsewhere in Australia and the southern hemisphere.     

A nested-intensity design for surveying plant diversity

Managers of natural landscapes need cost-efficient, accurate, and precise systems to inventory plant diversity. We investigated a nested-intensity sampling design to assess local and landscape-scale heterogeneity of plant species richness in aspen stands in southern Colorado, USA. The nested-intensity design used three vegetation sampling techniques: the Modified-Whittaker, a 1000-m² multiple-scale plot (n = 8); a 100-m² multiple-scale Intensive plot (n = 15); and a 100-m² single-scale Extensive plot (n = 28). The large Modified-Whittaker plot (1000 m²) recorded greater species richness per plot than the other two sampling techniques (P < 0.001), estimated cover of a greater number of species in 1-m² subplots (P < 0.018), and captured 32 species missed by the smaller, more numerous 100-m² plots of the other designs. The Intensive plots extended the environmental gradient sampled, capturing 17 species missed by the other techniques, and improved species–area calculations. The greater number of Extensive plots further expanded the gradient sampled, and captured 18 additional species. The multi-scale Modified-Whittaker and Intensive designs allowed quantification of the slopes of species–area curves in the single-scale Extensive plots. Multiple linear regressions were able to predict the slope of species–area curves (adj R ² = 0.64, P < 0.001) at each Extensive plot, allowing comparison of species richness at each sample location. Comparison of species–accumulation curves generated with each technique suggested that small, single-scale plot techniques might be very misleading because they underestimate species richness by missing locally rare species at every site. A combination of large and small multi-scale and single-scale plots greatly improves our understanding of native and exotic plant diversity patterns.     

Diversity patterns of wet meadows along geochemical gradients in central Spain

Four measures of biodiversity (species number per site, total species number, mean similarity and mosaic diversity) and their relationships with soil chemical composition were studied in vascular plant communities in groundwater discharge ecosystems of central Spain. Species richness decreased with increasing salinity, alkalinity and halite concentration. Species richness was apparently controlled more by soil toxicity than by soil nutrient levels, although a positive correlation of Ca²⁺ with species richness was found after accounting for the effects of toxic compounds. All relationships were strictly monotonic. Six community types were identified based on their soil chemical characteristics: glycophyte, subglycophyte, tolerant, subalkalinophyte, alkalinophyte, and halocalcicole communities. Within community types, species richness showed very few significant relationships with soil characteristics. Mean species richness was lowest in the environmentally stressful communities. Total species richness was greatest in the ecotonal community type. Mean similarity, a measure of among-community diversity, and mosaic diversity, a measure of landscape complexity, differed among community types. Mean similarity was smaller (higher diversity) in species-poor community types, while mosaic diversity was greatest (greater complexity) in species-rich community types. The halocalcicole community type was richest in rare species.     

Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini

Aim  A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.
Location  The New World.
Methods  We test several proposed ecological, temporal, evolutionary and spatial explanations for latitudinal diversity gradients in the New World snake tribe Lampropeltini, which exhibits its highest biodiversity in temperate regions.
Results  We find that an extratropical peak in species richness is not explained by latitudinal variation in diversification rate, the mid-domain effect, or Rapoport's rule. Rather, earlier colonization and longer duration in the temperate zones allowing more time for speciation to increase biodiversity, phylogenetic niche conservatism limiting tropical dispersal and the expansion of the temperate zones in the Tertiary better explain inverse diversity gradients in this group.
Main conclusions  Our conclusions are the inverse of the predictions made by the tropical conservatism hypothesis to explain higher biodiversity near the equator. Therefore, we suggest that the processes invoked are not intrinsic to the tropics but are dependent on historical biogeography to determine the distribution of species richness, which we refer to as the 'biogeographical conservatism hypothesis'.     

Butterfly Communities of Urban Forest Fragments in Campinas, São Paulo, Brazil: Structure, Instability, Environmental Correlates, and Conservation

A comparative study of butterfly communities in 15 urban/suburban remnants of tropical semideciduous forest in Campinas (São Paulo state, SE Brazil; 24°S, 47°W), with areas from 1.0 to 252ha and widely varying histories and environments, shows that the most significant factors, besides area and sampling time, distinguishing the sites and influencing their diversity (80–702 species) and composition are connectivity, permanent water, vegetation, flowers, and human impact (negative, including pollution). The diversified Nymphalidae butterflies (38–213 species) and especially two fermented-bait-attracted groups (Satyrinae, 2–30 species, and Biblidini, 9–44 species), are among the more useful indicators of the quality and diversity of the environments in these fragments. Effective conservation of butterfly communities in tropical cities may be achieved by maintenance of arboreal green corridors along streets and watercourses between moderately large (>10ha) humid areas, not near to the most built-up or polluted city centre(s), and the inclusion within these areas of ponds or streams, diversified native forest, and open vegetation including abundant nectar-rich flowers.     

The effect of earthworms on plant diversity and soil properties under different landuses

Earthworms have significant effect on vegetation and soil physical and chemical properties. The relationship between earthworms and biodiversity is important in ecological researches. Therefore, the effects of earthworm on plant diversity and various indices of diversity and richness, as well as physical and chemical properties of soil in shrub, grass and bare vegetation types on exclosure and grazing sites of Miankaleh Mazandaran biosphere reserve is investigated in this work. Accordingly, sampling was carried out randomly-systematically in three different vegetative types (shrubs, grass and bare soil) at 0 to 20 cm depth. Each vegetative types was placed on three transects of 100 m, 10 plots at a distance of 30 m. Earthworm and soil were sampled from the center of each plot of 25 cm × 25 cm. The Simpson and Shannon-Wiener indexes and Margalef and Menhinick richness indices were used to evaluate diversity and richness of species at different grazing intensities. Stepwise regression was used to study the relationship between diversity and richness indices with different earthworm parameters. The results showed that there was significant difference of potassium and nitrogen in vegetation types of both exclosure and grazing sites at P ≤ .01. Furthermore, the organic carbon, organic matter and potassium in the exclosure site and nitrogen and soil moisture in the grazing site are increased by earthworm abundance.     

Biodiversity of marine nematodes in Australian sandy beaches from tropical and temperate regions

The universal occurrence and abundance of nematodes provides opportunities to investigate ecological factors that may influence biodiversity. Clarke and Warwick (2001) have proposed diversity indices average taxonomic distance(AvTD), variation in taxonomic distinctness (VarTD) for computing marine nematode biodiversity based on classification trees. Faith [Biological Conservation 61 (1992) 1] had previously proposed a diversity index based on phylogenetic trees, though not applied to nematodes. Clarke and Warwick (2001) also considered an index AvPD analogous to AvTD. These indices have been applied to five very large collections of free-living nematodes from three exposed sandy beaches in Australia. Two were from a beach close to Darwin in northern Australia, two from the temperate southeast coast of Australia and one from the south of the Australian mainland. The collections extend over a considerable range of latitude, from 12°26S to 38°33S and the controversial hypothesis that latitudinal gradients per se influence the biodiversity of marine nematode assemblages is examined. AvTD did not vary among collections and its value for any collection was indistinguishable from that of random samples of the same size drawn from the total species pool. VarTD showed no variation for three of the collections, but was slightly higher than expected for the mid-latitude beach, attributed to unevenness in the classification trees. AvTD and VarTD were not greatly affected by differences in sampling intensity. PD varied directly with the number of species found but observed PD did not differ from the PD of random samples of the same number of species taken from the total species pool. Thus, the observed variation in PD is wholly accounted for by variations in species richness. The number of species found increased with decreasing latitude. It appears that species richness by itself is an adequate index of biodiversity for the free-living nematodes of these sandy beaches, and more complex indices such as AvTD, VarTD and PD are unnecessary.     

Landscape patterns of understory composition and richness across a moisture and nitrogen mineralization gradient in Ohio (U.S.A.) Quercus forests

This study quantified relationships of understory vascular plant species composition and richness along environmental gradients over a broad spatial scale in second-growth oak forests in eastern North America. Species frequencies were recorded in 108 25 × 25 m plots in four study sites extending over 70 km in southern Ohio, U.S.A.. The plots were stratified into three long-term soil moisture classes with a GIS-derived integrated moisture index (IMI). In addition to the IMI, the environmental data matrix included eight soil and three overstory variables. Canonical correspondence analysis (CCA) indicated that variations in understory species composition were most strongly related to topographic variations in predicted moisture (IMI), N mineralization rate, nitrification rate, and soil pH. In addition, floristic variation at the regional scale was correlated with variations in soil texture, nitrification, pH, and PO₄ ^–, resulting from differences in the soil parent material complexes among sites. Species richness averaged 65 species/plot, and increased with moisture and fertility. Stepwise regression indicated that richness was positively correlated with N mineralization rate and nitrification rate, and inversely correlated with tree basal area. Greater richness on fertile plots was the largely the result of increasing forb richness. Forb richness per quadrat (2 m²) was most strongly and positively related to N mineralization rate. Conversely, richness of understory individuals of tree species was greatest on xeric, less-fertile plots. Our results describe general, broad-scale species-environment relationships that occurred at both the topographic scale (long-term moisture status and fertility) and the regional scale (geomorphological differences among the sites). Strong species richness-N mineralization correlations indicate an important link between below-ground processes and above-ground biodiversity. Because N availability was a strong correlate to vegetation patterns at a broad-scale, our results suggest that the increasing rates of atmospheric N deposition in the region could have a major impact on understory vegetation dynamics.     

Sampling for Regional Monitoring of Nematode Communities in Agricultural Soils

Regional assessment of nematode communities to monitor the condition or ecological health of agricultural soils requires sampling programs with measures of known reliability and the ability to detect differences over time. Numbers of fields sampled in a region, samples taken per field, and subsamples assayed per sample must be balanced with cost to provide the best sampling scheme. We used components of variance from statewide surveys in North Carolina (1992) and Nebraska (1993) to estimate number of (i) fields to be sampled; (ii) 20-core, composite soil samples to be obtained for each field; and (iii) subsamples to be assayed for each composite sample to detect a specified amount of change in index values within a geographic region. Variances for these three components were used to estimate the degree of reliability for five ecologically based indices (four measures of maturity and one of diversity) of nematode communities. Total variance for maturity and diversity indices, based upon communities of free-living nematodes, was greater in North Carolina than in Nebraska; the opposite was true for indices based strictly upon maturity of communities of plant-parasitic nematodes or of all nematodes in soil. Variability within samples was greater in North Carolina than in Nebraska, especially for maturity indices based only upon free-living nematodes. We identified two possible sampling strategies for a regional survey: Option 1, with two independent samples per field and a single subsample assayed per sample, which would provide a reliability ratio value ≥0.6 for most indices; and Option 2, with three independent samples per field and two subsamples assayed per sample, which would provide a reliability ratio value ≥0.7 for several indices. When cost was considered, Option 1 was the better strategy. Number of fields to be sampled within a region or state varied with the index chosen; with specific indices, however, a 10% change in mean index value could be detected with a sample of 50 to 100 fields.     

Functional diversity of avian communities increases with canopy height: From individual behavior to continental‐scale patterns

Vegetation complexity is an important predictor of animal species diversity. Specifically, taller vegetation should provide more potential ecological niches and thus harbor communities with higher species richness and functional diversity (FD). Resource use behavior is an especially important functional trait because it links species to their resource base with direct relevance to niche partitioning. However, it is unclear how exactly the diversity of resource use behavior changes with vegetation complexity. To address this question, we studied avian FD in relation to vegetation complexity along a continental‐scale vegetation gradient. We quantified foraging behavior of passerine birds in terms of foraging method and substrate use at 21 sites (63 transects) spanning 3,000 km of woodlands and forests in Australia. We also quantified vegetation structure on 630 sampling points at the same sites. Additionally, we measured morphological traits for all 111 observed species in museum collections. We calculated individual‐based, abundance‐weighted FD in morphology and foraging behavior and related it to species richness and vegetation complexity (indexed by canopy height) using structural equation modeling, rarefaction analyses, and distance‐based metrics. FD of morphology and foraging methods was best predicted by species richness. However, FD of substrate use was best predicted by canopy height (ranging 10–30 m), but only when substrates were categorized with fine resolution (17 categories), not when categorized coarsely (8 categories). These results suggest that, first, FD might increase with vegetation complexity independently of species richness, but whether it does so depends on the studied functional trait. Second, patterns found might be shaped by how finely we categorize functional traits. More complex vegetation provided larger "ecological space" with more resources, allowing the coexistence of more species with disproportionately more diverse foraging substrate use. We suggest that the latter pattern was driven by nonrandom accumulation of functionally distinct species with increasing canopy height.     

Ecological structure of a pelagic seabird community in the Southern Ocean

Summary The pelagic avifauna (excluding penguins) of the African sector (30°S–70°S, 20°W–40°E) of the Southern Ocean is described quantitatively, based on 3,005 10-min observation of seabirds during seven oceanic cruises in April 1979 – April 1980. The avifauna is characterized according to species richness, diversity, abundance and biomass. These indices are correlated with groups of birds ordered into principal diet and feeding-method classes. Birds eating either plankton, cephalopods or a mixed diet accounted for 51, 23 and 22% of the total avifaunal abundance and 22, 49 and 25% of the total biomass, respectively. Piscivores were represented poorly. Planktivores were especially abundant south of the Antarctic Convergence and, to a lesser extent, at the sub-Tropical Convergence. Cephalopod-eaters were most abundant north of the sub-Tropical Convergence. The greatest abundance of omnivores occurred where planktivores and cephalopod-eaters were least abundant. The distribution of the planktivores and cephalopodeaters is related tentatively to the availability of the birds' principal prey.