Effects of bird predation on arthropod abundance and tree growth across an elevational gradient

Considerable uncertainty surrounds the conditions under which birds can cause trophic cascades. In a three‐year experiment, we studied the direct and indirect effects of insectivorous birds on arthropod abundance, herbivory, and growth of striped maple Acer pensylvanicum saplings in a northern hardwood forest of central New Hampshire, USA. We manipulated bird predation by erecting exclosures around saplings and directly manipulated herbivory by removing herbivores. We also examined how climate modifies these interactions by replicating the experiment at three locations along an elevational gradient. Effects of bird predation were variable. Overall, mean arthropod biomass was 20% greater on saplings within bird exclosures than on controls (p<0.05). The mean biomass of leaf‐chewing herbivores, primarily Lepidoptera larvae, was 25% greater within exclosures but not statistically different from controls. To a lesser degree, mean herbivore damage to foliage within exclosures exceeded that of controls but differences were not significant. We also did not detect significant treatment effects on sapling shoot growth. The high understory vegetation density relative to bird abundance, and low rate of herbivory during the study (mean 5% leaf area removed, controls), may have limited the ability of birds to affect sapling growth. Climate effects operated at multiple scales, resulting in a complex interplay of interactions within the food web. Regional synchrony of climatic conditions resulted in annual fluctuations in herbivore abundance and tree growth that were shared across elevations. At the same time, local environmental variation resulted in site differences in the plant, herbivore, and bird communities. These patterns resulted in a mosaic of top–down strengths across time and space, suggesting an overall pattern of limited effects of birds on plant growth, possibly interspersed with hotspots of trophic cascades.     

The direct and indirect effects of insectivory by birds in two contrasting Neotropical forests

A goal among community ecologists is to predict when and where trophic cascades occur. For example, several studies have shown that forest birds can limit arthropod abundances on trees, but indirect effects of bird predation (i.e. decreased arthropod damage to trees) are not always observed and their context is not well understood. Because productivity is one factor that is expected to influence trophic cascades, we compared the extent to which birds indirectly limit herbivore damage to trees in two lowland Neotropical forests that differed in seasonality of leaf production and rainfall. We compared the effects of bird predation on local arthropod densities and on damage to foliage through a controlled experiment using bird exclosures in the canopy and understory of two forests. We found that birds decreased local arthropod densities and leaf damage in the canopy of the drier site during periods of high leaf production, but not in the wetter forest where leaf production was low and sporadic throughout the year. Birds had no effect on arthropod abundances and leaf damage in the understory where leaf production and turnover rates were low. In support of these experimental interpretations, although we observed that arthropod densities were similar at the two sites, bird densities and the rate at which birds captured arthropods were greater at the drier, seasonally productive site. The influence of top-down predation by birds in limiting herbivorous insects appears to be conditional and most important when the production and turnover of leaves are comparatively high.Electronic Supplementary Material Supplementary material is available for this article at     

The direct and indirect effects of insectivory by birds in two contrasting Neotropical forests

A goal among community ecologists is to predict when and where trophic cascades occur. For example, several studies have shown that forest birds can limit arthropod abundances on trees, but indirect effects of bird predation (i.e. decreased arthropod damage to trees) are not always observed and their context is not well understood. Because productivity is one factor that is expected to influence trophic cascades, we compared the extent to which birds indirectly limit herbivore damage to trees in two lowland Neotropical forests that differed in seasonality of leaf production and rainfall. We compared the effects of bird predation on local arthropod densities and on damage to foliage through a controlled experiment using bird exclosures in the canopy and understory of two forests. We found that birds decreased local arthropod densities and leaf damage in the canopy of the drier site during periods of high leaf production, but not in the wetter forest where leaf production was low and sporadic throughout the year. Birds had no effect on arthropod abundances and leaf damage in the understory where leaf production and turnover rates were low. In support of these experimental interpretations, although we observed that arthropod densities were similar at the two sites, bird densities and the rate at which birds captured arthropods were greater at the drier, seasonally productive site. The influence of top-down predation by birds in limiting herbivorous insects appears to be conditional and most important when the production and turnover of leaves are comparatively high. Figure legends were missing in the original article published under Plant Animal Interactions, Oecologia (2005) 143: 106–166. The complete article is repeated here. The online version of the original article can be found at     

Where Is the Extended Phenotype in the Wild? The Community Composition of Arthropods on Mature Oak Trees Does Not Depend on the Oak Genotype

Through a series of common garden experiments, it has been shown that heritable phenotypic differences between individual trees can affect arthropod communities. However, field studies under heterogeneous environmental conditions remain rare. In the present study, we investigated the genetic constitution of 121 mature oak host trees at different trophic levels from 10 sites across Bavaria, southern Germany and their associated insect communities. A total of 23,576 individuals representing 395 species of beetles and true bugs were evaluated. In particular, we determined whether the composition of arthropod communities is related to the oak genotype and whether the strength of the relationships decreases from lower to higher trophic levels, such as for phytophagous, xylophagous, zoophagous, and mycetophagous species. The genetic differentiation of oaks was assessed using eight microsatellite markers. We found no significant influence of the oak genotype on neither the full beetle and true bug community nor on any of the analyzed trophic guilds. In contrast, the community composition of the insects was highly related to the space and climate, such that the community similarity decreased with increases in spatial distance and climatic differences. The relationship with space and climate was much stronger in beetles than in true bugs, particularly in mycetophagous species. Our results suggest that spatial processes override the genetic effects of the host plant in structuring arthropod communities on oak trees. Because we used neutral markers, we cannot exclude the possibility that trait-specific markers may reveal a genetic imprint of the foundation tree species on the composition of the arthropod community. However, based on the strength of the spatial patterns in our data set, we assume that genetic differences among oaks are less important in the structuring of arthropod communities. Future whole-genome studies are required to draw a final conclusion.     

Predation on sentinel prey increases with increasing latitude in Brassica‐dominated agroecosystems

In natural ecosystems, arthropod predation on herbivore prey is higher at lower latitudes, mirroring the latitudinal diversity gradient observed across many taxa. This pattern has not been systematically examined in human‐dominated ecosystems, where frequent disturbances can shift the identity and abundance of local predators, altering predation rates from those observed in natural ecosystems. We investigated how latitude, biogeographical, and local ecological factors influenced arthropod predation in Brassica oleracea‐dominated agroecosystems in 55 plots spread among 5 sites in the United States and 4 sites in Brazil, spanning at least 15° latitude in each country. In both the United States and Brazil, arthropod predator attacks on sentinel model caterpillar prey were highest at the highest latitude studied and declined at lower latitudes. The rate of increased arthropod attacks per degree latitude was higher in the United States and the overall gradient was shifted poleward as compared to Brazil. PiecewiseSEM analysis revealed that aridity mediates the effect of latitude on arthropod predation and largely explains the differences in the intensity of the latitudinal gradient between study countries. Neither predator richness, predator density, nor predator resource availability predicted variation in predator attack rates. Only greater non‐crop plant density drove greater predation rates, though this effect was weaker than the effect of aridity. We conclude that climatic factors rather than ecological community structure shape latitudinal arthropod predation patterns and that high levels of aridity in agroecosystems may dampen the ability of arthropod predators to provide herbivore control services as compared to natural ecosystems.     

Contrasting cascades: insectivorous birds increase pine but not parasitic mistletoe growth

1. Intraguild predation occurs when top predators feed upon both intermediate predators and herbivores. Intraguild predators may thus have little net impact on herbivore abundance. Variation among communities in the strength of trophic cascades (the indirect effects of predators on plants) may be due to differing frequencies of intraguild predation. Less is known about the influence of variation within communities in predator-predator interactions upon trophic cascade strength. 2. We compared the effects of a single predator community between two sympatric plants and two herbivore guilds. We excluded insectivorous birds with cages from ponderosa pine Pinus ponderosa trees parasitized by dwarf mistletoe Arceuthobium vaginatum. For 3 years we monitored caged and control trees for predatory arthropods that moved between the two plants, foliage-feeding caterpillars and sap-feeding hemipterans that were host-specific, and plant damage and growth. 3. Excluding birds increased the abundance of ant-tended aphids on pine and resulted in an 11% reduction in pine woody growth. Mutualist ants protected pine-feeding aphids from predatory arthropods, allowing aphid populations to burgeon in cages even though predatory arthropods also increased in cages. By protecting pine-feeding aphids from predatory arthropods but not birds, mutualist ants created a three-tiered linear food chain where bird effects cascaded to pine growth via aphids. 4. In contrast to the results for tended aphids on pine, bird exclusion had no net effects on untended pine herbivores, the proportion of pine foliage damaged by pine-feeding caterpillars, or the proportion of mistletoe plants damaged by mistletoe-feeding caterpillars. These results suggest that arthropod predators, which were more abundant in cages as compared with control trees, compensated for bird predation of untended pine and mistletoe herbivores. 5. These contrasting effects of bird exclusion support food web theory: where birds were connected to pine by a linear food chain, a trophic cascade occurred. Where birds fed as intraguild predators, the reticulate food webs linking birds to pine and mistletoe resulted in no net effects on herbivores or plant biomass. Our study shows that this variation in food web structure occurred between sympatric plants and within plants between differing herbivore guilds.     

Disturbance,species loss and compensation: Wildfire and grazing effects on the avian community and its food supply in the Serengeti Ecosystem,Tanzania

An important question in biodiversity studies is whether disturbances in ecosystems will cause a net loss of species or whether such losses can be compensated by replacement of other species. We use two natural disturbances, fire and grazing, to examine the response of bird and arthropod communities in grasslands of Serengeti, Tanzania. Both burning and grazing by migrant ungulates take place at the end of the rains in June–July. We documented the communities before disturbance, then 1, 4 and 20 weeks after disturbance on three replicate plots and compared them with three undisturbed plots. Birds were recorded by observation, arthropods from pitfall, tray trap and sweepnet samples. We expected that as the grass biomass was reduced by either disturbance, bird communities would change with concomitant change in arthropod food abundance. Alternatively, bird communities would change not with the absolute amount of food but with the greater accessibility of food as the grass structure changed from long to short grass. Results showed first that both bird species richness and abundance increased after both types of disturbance, but burnt sites showed a greater increase than that for grazed sites. Second, there was a change in bird species composition with disturbance. The functionally equivalent athi short‐toed lark (Calandrella athensis) was replaced by the red‐capped lark (Calandrella cinerea). Third, the abundance of most groups of arthropods was lower on disturbed sites than those on undisturbed sites, and the reduction of arthropod numbers was greatest on burnt sites. These results imply that bird abundance did not occur through an increase in arthropod abundance but rather through a change in the grass structure making food more accessible; and the higher predation could have caused the lower arthropod abundance. In addition, some bird species replaced others thus functionally compensating for their loss.     

Bird predation on herbivorous insects: indirect effects on sugar maple saplings

Insectivorous birds have been shown to have direct effects on abundances of herbivorous arthropods, but few studies have tested the indirect effects of birds on plant performance through consumption of herbivorous insects. In a 3-year study at the Hubbard Brook Experimental Forest, New Hampshire, we tested whether bird predation indirectly affects leaf herbivory levels and leaf and shoot biomass production of understory sugar maple (Acer saccharum) saplings. Trees were randomly assigned to one of four treatments: an insecticide application to reduce herbivory levels, exclosures that prevented bird access, addition of Lepidoptera larvae, and controls. Trees sprayed with an insecticide supported significantly fewer Lepidoptera larvae than other treatments throughout the study. Also, trees in exclosures supported more Lepidoptera larvae than controls during one count each year, and pooled across all counts during the second year. As predicted, the mean proportion of leaf area consumed varied significantly among treatments and was least in the insecticide treatment, followed by controls, exclosures, and Lepidoptera additions. Significant differences among treatments in herbivory levels, however, did not lead to differences in leaf or shoot biomass production. Thus, bird predation decreased Lepidoptera abundances and decreased herbivory levels, but did not increase biomass production during the following year. Over 85% of the herbivores in our study were Homoptera nymphs that were not folivorous and are not important bird prey items, potentially dampening the indirect effects of bird predation on biomass production. A comparison of these results with previous studies suggests that the indirect effects of bird predation on plant biomass production may depend on the plant species, abundance and composition of the herbivore community, and primary productivity of the ecosystem.     

Land rehabilitation and the conservation of birds in a degraded Afromontane landscape in northern Ethiopia

The few remaining Afromontane forest fragments in northern Ethiopia and the surrounding degraded, semiarid matrix form a habitat mosaic of varying suitability for forest birds. To evaluate the effect of recent land rehabilitation efforts on bird community composition and diversity, we studied bird species distributions in ten small forest fragments (0.40–20.95 ha), five grazing exclosures (10-year-old forest restoration areas without wood extraction and grazing livestock) and three grazed matrix sites during the rainy season (July–October 2004) using 277 one-hour species counts. Based on the distribution pattern of 146 bird species, sites were assigned to one of three bird communities (birds of moist forest, dry forest or degraded savanna), each occupying a well-defined position along an environmental gradient reflecting decreasing vegetation structure and density. All three communities were representative of the avifauna of Afrotropical Highland open forest and woodland with a high proportion of invasive and competitive generalist species (31%). Apart from these, exclosures shared more species with forest fragments (20%) than did the grazed matrix (5%), indicating local ecosystem recovery. By increasing habitat heterogeneity, exclosures have the potential to enhance landscape connectivity for forest birds and are, therefore, an effective instrument for conserving species in a fragmented landscape. However, 52 bird species (36%) occurred exclusively within forest patches and many forest birds that use exclosures are unlikely to maintain viable populations when forest fragments disappear, particularly as forest fragments may be a critical resource during the hot dry season. This highlights the high conservation value of small isolated forest fragments for less tolerant, forest-limited and/or biome-restricted species.     

Responses of belowground communities to large aboveground herbivores: Meta‐analysis reveals biome‐dependent patterns and critical research gaps

The importance of herbivore–plant and soil biota–plant interactions in terrestrial ecosystems is amply recognized, but the effects of aboveground herbivores on soil biota remain challenging to predict. To find global patterns in belowground responses to vertebrate herbivores, we performed a meta‐analysis of studies that had measured abundance or activity of soil organisms inside and outside field exclosures (areas that excluded herbivores). Responses were often controlled by climate, ecosystem type, and dominant herbivore identity. Soil microfauna and especially root‐feeding nematodes were negatively affected by herbivores in subarctic sites. In arid ecosystems, herbivore presence tended to reduce microbial biomass and nitrogen mineralization. Herbivores decreased soil respiration in subarctic ecosystems and increased it in temperate ecosystems, but had no net effect on microbial biomass or nitrogen mineralization in those ecosystems. Responses of soil fauna, microbial biomass, and nitrogen mineralization shifted from neutral to negative with increasing herbivore body size. Responses of animal decomposers tended to switch from negative to positive with increasing precipitation, but also differed among taxa, for instance Oribatida responded negatively to herbivores, whereas Collembola did not. Our findings imply that losses and gains of aboveground herbivores will interact with climate and land use changes, inducing functional shifts in soil communities. To conceptualize the mechanisms behind our findings and link them with previous theoretical frameworks, we propose two complementary approaches to predict soil biological responses to vertebrate herbivores, one focused on an herbivore body size gradient, and the other on a climate severity gradient. Major research gaps were revealed, with tropical biomes, protists, and soil macrofauna being especially overlooked.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Native Birds and Alien Insects: Spatial Density Dependence in Songbird Predation of Invading Oak Gallwasps

Revealing the interactions between alien species and native communities is central to understanding the ecological consequences of range expansion. Much has been learned through study of the communities developing around invading herbivorous insects. Much less, however, is known about the significance of such aliens for native vertebrate predators for which invaders may represent a novel food source. We quantified spatial patterns in native bird predation of invading gall-inducing Andricus wasps associated with introduced Turkey oak (Quercus cerris) at eight sites across the UK. These gallwasps are available at high density before the emergence of caterpillars that are the principle spring food of native insectivorous birds. Native birds showed positive spatial density dependence in gall attack rates at two sites in southern England, foraging most extensively on trees with highest gall densities. In a subsequent study at one of these sites, positive spatial density dependence persisted through four of five sequential week-long periods of data collection. Both patterns imply that invading galls are a significant resource for at least some native bird populations. Density dependence was strongest in southern UK bird populations that have had longest exposure to the invading gallwasps. We hypothesise that this pattern results from the time taken for native bird populations to learn how to exploit this novel resource.     

Herbivore damage increases avian and ant predation of caterpillars on trees along a complete elevational forest gradient in Papua New Guinea

Signals given off by plants to alert predators to herbivore attack may provide exciting examples of coevolution among organisms from multiple trophic levels. We examined whether signals from mechanically damaged trees (simulating damage by herbivores) attract predators of insects along a complete elevational rainforest gradient in tropical region, where various predators are expected to occur at particular elevational belts. We studied predation of artificial caterpillars on trees with and without ‘herbivorous’ damage; as well as diversity and abundances of potential predators at eight study sites along the elevational gradient (200–3700 m a.s.l.). We focused on attacks by ants and birds, as the main predators of herbivorous insect. The predation rate decreased with elevation from 10% d^?1 at 200 m a.s.l. to 1.8% d^?1 at 3700 m a.s.l. Ants were relatively more important predators in the lowlands, while birds became dominant predators above 1700 m a.s.l. Caterpillars exposed on trees with herbivorous damage were attacked significantly more than caterpillars exposed on trees without damage. Results suggest that relative importance of predators varies along elevational gradient, and that observed predation rates correspond with abundances of predators. Results further show that herbivorous damage attracts both ants and birds, but its effect is stronger for ants.     

Structural complexity of arthropod guilds is affected by the agricultural landscape heterogeneity generated by fencerows

Intensive farming imposes harsh conditions impeding the persistence of most arthropod species within crop fields. Hence, arthropods surviving the unfavourable conditions prevailing within crop fields may disperse towards nearby uncropped margins, such as fencerows. Here, we evaluate the influence of landscape heterogeneity on the abundance of different guilds, particularly herbivores and their natural enemies. Said heterogeneity mostly derives from fencerow network density. Hence, we developed an approach based on fitting linear‐mixed models to elucidate the effects of landscape heterogeneity and field position (fencerows and crop interiors) on arthropod diversity. Mixed models were fitted to arthropod data obtained by pitfall trap samplings in 36 crop fields. Arthropod communities were structurally and functionally more complex along fencerows than within nearby crop interiors. Arthropods abundance was modulated by landscape heterogeneity, increasing the abundance of natural enemies as the landscape heterogeneity increased. On the contrary, herbivores abundance decreased as landscape heterogeneity increased. Consequently, the ratio between herbivores and natural enemies also decreased as landscapes became more heterogeneous. Natural enemies with larger body sizes, mostly carabid beetles, were more sensitive to landscape homogenisation. Our study reveals that, despite the coarse‐grained landscapes in the Rolling Pampa, fencerow density appears as a key factor for structuring complex arthropod guilds in intensively farmed agricultural mosaics. In landscapes with higher density of fencerows, arthropods tend to concentrate along them, thus increasing the community structural complexity as well as the predation pressure over herbivores. This structural complexity of upper trophic levels enhances the ‘top‐down’ regulation of herbivore populations, consequently decreasing the probability of pest outbreaks within crop fields.     

From genes to geography: a genetic similarity rule for arthropod community structure at multiple geographic scales

We tested the hypothesis that leaf modifying arthropod communities are correlated with cottonwood host plant genetic variation from local to regional scales. Although recent studies found that host plant genetic composition can structure local dependent herbivore communities, the abiotic environment is a stronger factor than the genetic effect at increasingly larger spatial scales. In contrast to these studies we found that dependent arthropod community structure is correlated with both the cross type composition of cottonwoods and individual genotypes within local rivers up to the regional scale of 720,000 km(2) (Four Corner States region in the southwestern USA). Across this geographical extent comprising two naturally hybridizing cottonwood systems, the arthropod community follows a simple genetic similarity rule: genetically similar trees support more similar arthropod communities than trees that are genetically dissimilar. This relationship can be quantified with or without genetic data in Populus.     

Goose grazing influences the fine-scale structure of a bryophyte community in arctic wetlands

Herbivores can shape plant communities, especially in the Arctic. We tested the role of geese for structuring bryophyte communities at fine spatial scales in the arctic tundra by excluding them from 4 × 4 m areas. We surveyed the presence and absence of bryophyte species in quadrats (10 × 10 cm) divided into 25 cells outside and inside these exclosures, after 5 and 11 years of treatment. Species richness per cell (4 cm²) was higher in the presence of geese, especially after 11 years of treatment, while geese had little effect on richness at larger scales (i.e. quadrat and whole exclosure). The slope of the species–area relationship within quadrats was consequently shallower outside exclosures. Our results further suggest that the community outside the exclosures was more variable in space and time than that inside the exclosures. We conclude that goose foraging activity promotes the coexistence of bryophyte species at the centimetre scale.     

Poachers and Forest Fragmentation Alter Seed Dispersal,Seed Survival,and Seedling Recruitment in the Palm Attalea butyraceae,with Implications for Tropical Tree Diversity1

We examined the interaction between a palm and two bruchid beetles along with several mammal species to explore how poachers and habitat fragmentation may indirectly alter the spatial pattern of seed dispersal, seed predation, and seedling recruitment in central Panama. The large, stony endocarps of Attalea butyraceae decay slowly and bear distinctive scars when opened by rodents or beetles. We determined the final distance between endocarps and reproductive trees (which we call an ecologically effective dispersal distance), the predation status of each endocarp, and the distance between seedlings and reproductive trees. The 68 focal trees were divided among 14 sites and four levels of anthropogenic disturbance. Levels of disturbance included full protection from poachers, light and heavy pressure from poachers, and small island habitat fragments. Ecologically effective seed dispersal distances were greatest for protected sites, intermediate for lightly poached sites, and shortest for heavily poached sites and habitat fragments. Seed predation by rodents increased with distance to the nearest reproductive Attalea and was greatest for fully protected sites, intermediate for lightly poached sites, and least for heavily poached sites and habitat fragments. Seed predation by beetles reversed the patterns described for seed predation by rodents. Total seed predation by beetles and rodents combined was independent of distance, greatest for fully protected sites, and lower for poached sites and habitat fragments. Seedling densities were always greatest close to reproductive trees; however, the increase in seedling densities close to reproductive trees was minimal for fully protected sites, clearly evident for poached sites, and pronounced for habitat fragments. Increased seedling recruitment near conspecific trees may in time reduce tree diversity where humans disrupt mammal communities.     

Control of arthropod abundance, richness, and composition in a heterogeneous desert city

There is a demand for mechanistic studies to explore underlying drivers behind observed patterns of biodiversity in urban areas. We describe a two-year field experiment in which we manipulated bottom-up (resource availability) and top-down (bird predation) forces on arthropod communities associated with a native plant, Encelia farinosa, across three land-use types—urban, desert remnant, and outlying natural desert—in the Phoenix metropolitan area, Arizona, USA. We monitored the trophic structure, richness, and similarity of the arthropod communities on these manipulated plants over a two-year period. We predicted that (1) increased water resources increase plant productivity, (2) increased productivity increases arthropod abundances, and (3) in the urban habitat, top-down forces are greater than in other habitats and limit arthropod abundances. We also predicted that urban remnant habitats are more similar to urban habitats in terms of arthropod richness and composition. Strong interannual differences due to an unusual cold and dry winter in the first year suppressed plant growth in all but urban habitats, and arthropod abundances in all habitats were severely reduced. In the following year, arthropod abundances in desert and remnant habitats were higher than in urban habitats. Water had positive effects on plant growth and arthropod abundance, but these water effects emerged through complex interactions with habitat type and the presence/absence of cages used to reduce bird predation. Plants grew larger in urban habitats, and phenology also differed between urban and desert habitats. The results from caging suggest that bird predation may not be as important in cities as previously thought, and that arthropods may retard plant growth. As expected, desert communities are strongly bottom-up regulated, but contrary to predictions, we did not find evidence for strong top-down control in the city. Remnant habitats were intermediate between desert and urban habitats in terms of diversity, richness, evenness, arthropod composition and phenology, with urban habitats generally lowest in terms of diversity, richness, and evenness. Our study shows that control of biodiversity is strongly altered in urban areas, influenced by subtle shifts in top-down and bottom-up controls that are often superseded by climatic variations and habitat type.     

Ecological assembly rules on arthropod community inhabiting mistletoes

1. The colonisation of a new habitat by a community is led by deterministic and stochastic processes at different spatio-temporal scales. Parasitic plants, such as mistletoe, represent a new habitat within forest canopy that is free to be colonised by many organisms. 2. This study investigates how ecological factors operating at forest and plant scales drive changes in both specialist (mistletoe-dwelling) and tourist (transient visitors) arthropod communities inhabiting European mistletoe, Viscum album subsp. austriacum, in a Mediterranean pine forest. The influence of elevation along a broad elevational gradient was tested by sampling arthropod communities dwelling in mistletoe plants and host pine branches and the effects of mistletoe plant size, distances to other mistletoes, and temporal variation in arthropod assemblages inhabiting mistletoes. 3. The diversity of the specialist community remained constant along the elevational gradient and over the summer period, while the tourist and pine-dwelling arthropod communities showed species turnover. Larger mistletoes were occupied by more species and individuals, whereas more isolated mistletoes presented the same equilibrium point as the more aggregated ones. Thus, mistletoe size is key to the composition of the arthropod community. 4. In conclusion, this study's findings indicate contrasting assembly rules for specialised and tourist arthropod communities associated with mistletoe. The specialist community was highly stable and followed a deterministic trophic sequence of colonisation as the assemblage rule: first, colonisation by the main specialist herbivore, Cacopsylla visci, and, second, by its predator Anthocoris visci. Meanwhile, the tourist community, being a subset of the arthropod assemblage of the pine, acts independent of mistletoe presence.     

Latitudinal variation in plant chemical defences drives latitudinal patterns of leaf herbivory

A long‐standing paradigm in ecology holds that herbivore pressure and thus plant defences increase towards lower latitudes. However, recent work has challenged this prediction where studies have found no relationship or opposite trends where herbivory or plant defences increase at higher latitudes. Here we tested for latitudinal variation in herbivory, chemical defences (phenolic compounds), and nutritional traits (phosphorus and nitrogen) in leaves of a long‐lived tree species, the English oak Quercus robur. We further investigated the underlying climatic and soil factors associated with such variation. Across 38 populations of Q. robur distributed along an 18° latitudinal gradient, covering almost the entire latitudinal and climatic range of this species, we observed strong but divergent latitudinal gradients in leaf herbivory and leaf chemical defences and nutrients. As expected, there was a negative relationship between latitude and leaf herbivory where oak populations from lower latitudes exhibited higher levels of leaf herbivory. However, counter to predictions there was a positive relationship between leaf chemical defences and latitude where populations at higher latitudes were better defended. Similarly, leaf phosphorus and nitrogen increased with latitude. Path analysis indicated a significant (negative) effect of plant chemical defences (condensed tannins) on leaf herbivory, suggesting that the latitudinal gradient in leaf herbivory was driven by an inverse gradient in defensive investment. Leaf nutrients had no independent influence on herbivory. Further, we found significant indirect effects of precipitation and soil porosity on leaf herbivory, which were mediated by plant chemical defences. These findings suggest that abiotic factors shape latitudinal variation in plant defences and that these defences in turn underlie latitudinal variation in leaf herbivory. Overall, this study contributes to a better understanding of latitudinal variation in plant–herbivore interactions by determining the identity and modus operandi of abiotic factors concurrently shaping plant defences and herbivory.