Nitrogen and phosphorus enrichment cause declines in invertebrate populations: a global meta-analysis

Human-driven changes in nitrogen (N) and phosphorus (P) inputs are modifying biogeochemical cycles and the trophic state of many habitats worldwide. These alterations are predicted to continue to increase, with the potential for a wide range of impacts on invertebrates, key players in ecosystem-level processes. Here, we present a meta-analysis of 1679 cases from 207 studies reporting the effects of N, P, and combined N + P enrichment on the abundance, biomass, and richness of aquatic and terrestrial invertebrates. Nitrogen and phosphorus additions decreased invertebrate abundance in terrestrial and aquatic ecosystems, with stronger impacts under combined N + P additions. Likewise, N and N + P additions had stronger negative impacts on the abundance of tropical than temperate invertebrates. Overall, the effects of nutrient enrichment did not differ significantly among major invertebrate taxonomic groups, suggesting that changes in biogeochemical cycles are a pervasive threat to invertebrate populations across ecosystems. The effects of N and P additions differed significantly among invertebrate trophic groups but N + P addition had a consistent negative effect on invertebrates. Nutrient additions had weaker or inconclusive impacts on invertebrate biomass and richness, possibly due to the low number of case studies for these community responses. Our findings suggest that N and P enrichment affect invertebrate community structure mainly by decreasing invertebrate abundance, and these effects are dependent on the habitat and trophic identity of the invertebrates. These results highlight the important effects of human-driven nutrient enrichment on ecological systems and suggest a potential driver for the global invertebrate decline documented in recent years.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Soil Nematode Responses to Increases in Nitrogen Deposition and Precipitation in a Temperate Forest

The environmental changes arising from nitrogen (N) deposition and precipitation influence soil ecological processes in forest ecosystems. However, the corresponding effects of environmental changes on soil biota are poorly known. Soil nematodes are the important bioindicator of soil environmental change, and their responses play a key role in the feedbacks of terrestrial ecosystems to climate change. Therefore, to explore the responsive mechanisms of soil biota to N deposition and precipitation, soil nematode communities were studied after 3 years of environmental changes by water and/or N addition in a temperate forest of Changbai Mountain, Northeast China. The results showed that water combined with N addition treatment decreased the total nematode abundance in the organic horizon (O), while the opposite trend was found in the mineral horizon (A). Significant reductions in the abundances of fungivores, plant-parasites and omnivores-predators were also found in the water combined with N addition treatment. The significant effect of water interacted with N on the total nematode abundance and trophic groups indicated that the impacts of N on soil nematode communities were mediated by water availability. The synergistic effect of precipitation and N deposition on soil nematode communities was stronger than each effect alone. Structural equation modeling suggested water and N additions had direct effects on soil nematode communities. The feedback of soil nematodes to water and nitrogen addition was highly sensitive and our results indicate that minimal variations in soil properties such as those caused by climate changes can lead to severe changes in soil nematode communities.     

Nitrogen Addition Regulates Soil Nematode Community Composition through Ammonium Suppression

Nitrogen (N) enrichment resulting from anthropogenic activities has greatly changed the composition and functioning of soil communities. Nematodes are one of the most abundant and diverse groups of soil organisms, and they occupy key trophic positions in the soil detritus food web. Nematodes have therefore been proposed as useful indicators for shifts in soil ecosystem functioning under N enrichment. Here, we monitored temporal dynamics of the soil nematode community using a multi-level N addition experiment in an Inner Mongolia grassland. Measurements were made three years after the start of the experiment. We used structural equation modeling (SEM) to explore the mechanisms regulating nematode responses to N enrichment. Across the N enrichment gradient, significant reductions in total nematode abundance, diversity (H' and taxonomic richness), maturity index (MI), and the abundance of root herbivores, fungivores and omnivores-predators were found in August. Root herbivores recovered in September, contributing to the temporal variation of total nematode abundance across the N gradient. Bacterivores showed a hump-shaped relationship with N addition rate, both in August and September. Ammonium concentration was negatively correlated with the abundance of total and herbivorous nematodes in August, but not in September. Ammonium suppression explained 61% of the variation in nematode richness and 43% of the variation in nematode trophic group composition. Ammonium toxicity may occur when herbivorous nematodes feed on root fluid, providing a possible explanation for the negative relationship between herbivorous nematodes and ammonium concentration in August. We found a significantly positive relationship between fungivores and fungal phospholipid fatty acids (PLFA), suggesting bottom-up control of fungivores. No such relationship was found between bacterivorous nematodes and bacterial PLFA. Our findings contribute to the understanding of effects of N enrichment in semiarid grassland on soil nematode trophic groups, and the cascading effects in the detrital soil food web.     

Changes in trophic state and aquatic communities in high Arctic ponds in response to increasing goose populations

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Effects of drought on avian community structure

Droughts are expected to become more frequent under global climate change. Avifauna depend on precipitation for hydration, cover, and food. While there are indications that avian communities respond negatively to drought, little is known about the response of birds with differing functional and behavioural traits, what time periods and indicators of drought are most relevant, or how response varies geographically at broad spatial scales. Our goals were thus to determine (1) how avian abundance and species richness are related to drought, (2) whether community variations are more related to vegetation vigour or precipitation deviations and at what time periods relationships were strongest, (3) how response varies among avian guilds, and (4) how response varies among ecoregions with different precipitation regimes. Using mixed effect models and 1989–2005 North American Breeding Bird Survey data over the central United States, we examined the response to 10 precipitation‐ and greenness‐based metrics by abundance and species richness of the avian community overall, and of four behavioural guilds. Drought was associated with the most negative impacts on avifauna in the semiarid Great Plains, while positive responses were observed in montane areas. Our models predict that in the plains, Neotropical migrants respond the most negatively to extreme drought, decreasing by 13.2% and 6.0% in abundance and richness, while permanent resident abundance and richness increase by 11.5% and 3.6%, respectively in montane areas. In most cases, response of abundance was greater than richness and models based on precipitation metrics spanning 32‐week time periods were more supported than those covering shorter time periods and those based on greenness. While drought is but one of myriad environmental variations birds encounter, our results indicate that drought is capable of imposing sizable shifts in abundance, richness, and composition on avian communities, an important implication of a more climatically variable future.     

A large trophic quilt

Plants and invertebrate herbivores are major constituents of terrestrial food webs. Identifying component species and tracing their interactions in highly diverse communities are a monumental task. Novotny et al. 2010 present the first broad conspectus of herbivore–plant interactions in a forest in Papua New Guinea. In more than 15 years, nearly 7000 feeding links were traced between about 200 plants and 1500 insect herbivores. Although staggering, these figures might represent only 15% of the total herbivore richness and interaction diversity in that lowland forest. Standardized comparisons also showed distinctive specialization and diversification patterns in different feeding guilds, restricting the possibility of using any single guild as surrogate for the entire assemblage.     

Trends in estuarine fish assemblages facing different environmental conditions: combining diversity with functional attributes

Changes in the Mondego estuary (Portugal) fish assemblage were documented with a long-term monitoring program between June 2003 and March 2010, during which several extreme environmental events occurred, including severe droughts and heavy precipitation. The structure and composition of the fish assemblage was analyzed based on a set of indicators: dominance, diversity, evenness, and composition in functional groups (ecological and feeding guilds). Higher species number, diversity, and evenness were observed in the dry periods. Variations in the relative abundance of the ecological guilds were also observed: in dry years, estuarine residents were the most abundant group, while in typical and rainy years, the marine estuarine-dependent species increased in abundance. Among the feeding guilds, the most abundant group was the invertebrate and fish feeders. Planktivorous and invertebrate feeders occurred in higher abundance in dry years, while the opposite was verified for omnivorous species. The mean trophic level of the fish assemblage increased during the drought period, constantly decreasing afterward, which could be attributed to an increase in predators in dry years. The cumulative responses of estuarine fish assemblages to ongoing climate changes and discrete extreme weather events confirm their importance as indicators of environmental changes.     

Avian Use of Wetlands in Reclaimed Minelands in Southwestern Indiana

We studied the use of mineland wetlands by birds and the relationship between avian communities and wetland characteristics. Data were collected from 20 wetlands in Pike County, Indiana, and included wetland size, depth, water conductivity and salinity, aquatic macroinvertebrate abundance, vegetation, and bird use. Principal component analysis showed that physical variables could be explained by two principal component scores and that wetlands could be grouped on the basis of size and conductivity. Principal component analysis could not reduce vegetation variables to fewer principal component scores, meaning that wetland vegetation characteristics were independent of one another and did not show any trend. Most wetlands had low invertebrate density, and wetlands with higher invertebrate density had low invertebrate diversity. Wetlands with similar habitat characteristics (physical, vegetative, and invertebrate) did not necessarily show similarities in bird assemblages. Bird similarity index values ranged from 0 to 59%, implying that each wetland has its own bird community. Stepwise multiple regression analysis (α= 0.05) relating bird use and habitat characteristics showed that bird species richness increased with the species richness of submergent vegetation and was correlated negatively with the species richness of emergent vegetation. There was no significant relationship between bird species richness or bird species diversity and wetland size. The number of species within different avian guilds correlated with different habitat characteristics. The species richness of submergent plants was a factor that correlated positively with the number of species of several guilds (dabblers, wading birds, and plunge divers). Wetland age was not a factor that determined bird use.     

Agroecological transition increases arthropod diversity and decreases herbivore abundance on field margins

1. In peri-urban areas, many farmers are transitioning from conventional agriculture to agroecological practices to reduce pesticide input and preserving ecosystem services such as natural pest control. Field margins represent a stable habitat for arthropods, but community structure depends on many factors, including management type and vegetation features.
2. We studied the effects of agroecological transitions and vegetation features on arthropods of horticultural field margins, focusing on three feeding guilds (herbivores, predators and parasitoids). We sampled arthropods using the beat-sheet method in five conventional fields and five under agroecological transition. We also measured vegetation height, richness, flower abundance and plant cover.
3. Our results showed that arthropod diversity was higher in agroecological fields whereas herbivore abundance was lower, with a consistent pattern across most taxonomic orders. Vegetation features displayed multiple effects among functional and taxonomic groups. Herbivores did not respond to most vegetation variables whereas predators correlated with several, with similar trends among orders.
4. We conclude that agroecological transitions and field margins with high vegetation richness and floral resources influence arthropod communities with potential benefits regarding pest regulation. These practices might be more effective if considered alongside other methods that enhance biodiversity and if they are consistent at a landscape scale.

     

Short-term seasonal changes in parasite community structure in northern leopard froglets (Rana pipiens) inhabiting agricultural wetlands

Parasite community structure can change seasonally with shifts in host habitat and in diet. However, anthropogenic activity may influence the natural changes in transmission dynamics of different parasite species. Effects of seasonal and agricultural activity on the parasite communities of newly metamorphosed northern leopard frogs (Rana pipiens) were investigated in July and September 2001 in 5 wetlands, 3 of which were exposed to pesticide runoff from surrounding agriculture. Nineteen parasite taxa were found. Component community richness was consistently high at the pristine reference wetland, whereas the communities at a managed reference wetland remained depauperate. Infracommunity richness increased throughout the season, but more so in frogs resident in agricultural wetlands. Digeneans using frogs as intermediate hosts dominated the communities, although many species were much lower in abundance in September, suggesting mortality of heavily infected frogs. Mean abundance of Haematoloechus spp. was positively related to that of odonate naiads in the frog diet, which appeared to reflect differential second intermediate host availability between reference and agricultural wetlands. Although virtually absent from wetlands in July just after frog metamorphosis, monoxenous nematodes were more prevalent and abundant at agricultural wetlands as the season progressed. Our results suggest that agricultural activity may further facilitate the transmission of monoxenous nematodes as frogs become more terrestrial.     

Experimental evidence that aboveground predators are sustained by underground detritivores

Detrital infusion into grazing food web is considered to be important in terrestrial communities, but there is hardly any experimental evidence showing that generalist predators aboveground are sustained by belowground detritivores. We established two types of experimental plots in the forest floor, one with sheets on the ground to prevent the emergence of belowground arthropods and the other without sheet, to test the hypotheses that 1) reduced input of detrital arthropods decreases the abundance and species richness of web spiders (major generalist predators in terrestrial ecosystems) and 2) lower number of spiders increases the abundance of herbivorous arthropods. We found that spiders were less abundant in plots where the emergence of detrital arthropods was reduced, while the abundance of herbivores did not significantly increase in these plots. These results provide empirical evidence that organisms moving from underground to aboveground may be important for the maintenance of aboveground predators, although the cascading effect of predator abundance on the grazing food chain was not detected in the present study.     

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.     

Seasonal fluctuations of resource abundance and avian feeding guilds across forest–farmland boundaries in tropical Africa

Seasonal fluctuations in climatic factors are expected to increase in future decades. However, little is known about the response of tropical species communities to seasonal fluctuations in climate and resource availability, particularly across different habitat types. We examined the relationship between spatio‐temporal fluctuations in the abundance of fruits and invertebrates and two avian feeding guilds, i.e. frugivores and insectivores, in forest and farmland habitats in western Kenya. Fruits and invertebrates fluctuated substantially throughout the year, but seasonal fluctuations were asynchronous between the two habitat types. Species richness and total abundance of frugivores and insectivores also fluctuated strongly and were closely related to the abundance of their respective resources. Frugivore species richness fluctuated anti‐cyclical in forest and farmland habitats, suggesting that several frugivorous species tracked fruit resources across habitat boundaries. In contrast, insectivorous bird richness fluctuated synchronously in the two habitat types, suggesting a lack of local‐scale movements across habitat boundaries. We conclude that bird communities strongly respond to seasonal fluctuations in resource availability, but responses differ between feeding guilds. While frugivores seem to respond flexibly to seasonal fluctuations, for instance by tracking fruit resources across habitat boundaries, insectivorous birds appear to be more susceptible to the expected increase in seasonal fluctuations in resource availability.     

Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.     

Environmental and plant community determinants of species loss following nitrogen enrichment

Global energy use and food production have increased nitrogen inputs to ecosystems worldwide, impacting plant community diversity, composition, and function. Previous studies show considerable variation across terrestrial herbaceous ecosystems in the magnitude of species loss following nitrogen (N) enrichment. What controls this variation remains unknown. We present results from 23 N-addition experiments across North America, representing a range of climatic, soil and plant community properties, to determine conditions that lead to greater diversity decline. Species loss in these communities ranged from 0 to 65% of control richness. Using hierarchical structural equation modelling, we found greater species loss in communities with a lower soil cation exchange capacity, colder regional temperature, and larger production increase following N addition, independent of initial species richness, plant productivity, and the relative abundance of most plant functional groups. Our results indicate sensitivity to N addition is co-determined by environmental conditions and production responsiveness, which overwhelm the effects of initial community structure and composition.     

Urbanization drives cross‐taxon declines in abundance and diversity at multiple spatial scales

The increasing urbanization process is hypothesized to drastically alter (semi‐)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno‐terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground‐ and web spiders, macro‐moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local‐scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape‐scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity.     

Simulated ungulate herbivory affects differently two herbivorous arthropod guilds in bilberry

Dense ungulate populations, both domestic and wild, accompanied by high grazing intensities, are known to have effects on the abundance and diversity of the community of arthropods hosted by plants. However, research combining ungulate herbivory with nutrient addition effects is scarce. By means of a nitrogen (N) addition and clipping experiment, we investigated the effects of both factors on the diversity, abundance and trophic structure of the arthropod community hosted by the bilberry, Vaccinium myrtillus, focusing on the response of herbivorous arthropods by considering leaf-chewers and sap-feeders separately. In addition, we assessed altitudinal variation on the arthropod community and also specifically on herbivores. Neither of the variables estimated for the arthropod community (total abundance, richness and biomass) were affected by simulated ungulate herbivory. However, clipping reduced the abundance of sap-feeders although had no significant effect on leaf-chewer insects. We therefore suggest that ungulates can compete with sap-feeders by reducing suitable food resources, or via changing plant nutritional state. Moreover, the lack of response on leaf-chewers could correspond with the absence of chemical defence in bilberry after repeated damage. Despite this effect on sap-feeder abundance, clipping did not cause a cascade effect on the trophic structure of the arthropod community associated with bilberry, at least in the short term. In turn, N fertilization at levels close to atmospheric N deposition did not cause any significant effect on the invertebrate community. Finally, the number of arthropod morphospecies and biomass decreased with altitude.     

Arthropods associated with the crown of Mauritia flexuosa (Arecaceae) palm trees in three different environments from Brazilian Cerrado

Canopy arthropods, mainly from palm trees, are little known in the Brazilian Cerrado. In order to describe the arthropod community structure associated with the crown of Mauritia flexuosa (Arecaceae), we sampled 150 palm trees in six "veredas" of the Federal District, Brazil, in wild, rural and periurban areas in the rainy season. The arthropods within abandoned bird nests, mammal refuges, leaves and organic matter were manually collected, preserved in ethanol 70% and separated by order, family, morphospecies and feeding guilds. Stem height and diameter of the palm crowns were measured and leaves and bird nests were counted. We collected 3,862 arthropods, from 15 orders, 45 families and 135 morphospecies. The most abundant orders were Coleoptera (28.6%), Blattodea (21.8%), Collembola (11.4%) and Hemiptera (10.2%). The families Blaberidae, Entomobryidae, Reduviidae, Oniscidae, Staphylinidae, Carabidae and Formicidae, represented 82.1% of all individuals collected. The majority of morphospecies was not abundant, 71 (52.6%) were represented by less than 1 individual/tree. Coleopterans accounted for the highest number of morphospecies (43.7%) followed by Araneae (20.0%). The analysis of the arthropod feeding guilds showed prevalence of predatory/hematophagous ones (36.0%). Arthropod richness and abundance presented smaller values for periurban environment. The number of bird nests presented positive correlation with abundance and richness; this was not found when considering the measurements of the palm trees. The importance of M. flexuosa for the maintenance of the arthropod fauna of the "veredas" in Cerrado biome is discussed.     

Multi‐trophic guilds respond differently to changing elevation in a subtropical forest

Negative relationships between species richness and elevation are common and attributed to changes in single environmental properties associated to elevation, such as temperature and habitat area. However, research has lacked taxonomic breadth and comprehensive elevation studies that consider multiple groups from different trophic levels are rare. We thus analysed 24 groups of plants, arthropods, and microorganisms grouped into six trophic guilds (predators, detritivores, herbivores, plants, bacteria and fungi) along a relatively short elevational gradient (~600 m) in a subtropical forest in south‐east China. The total species richness of all organisms was not related to elevation, nor was the richness of plants, herbivores or microorganisms. However, species richness and abundance in two major trophic guilds of arthropods changed with elevation, which was mediated by changes in elevation‐associated habitat properties. Specifically, deadwood mass increased with elevation, which increased detritivore richness indirectly via detritivore abundance, thus supporting the ‘more individuals hypothesis’. In contrast, lower predator richness at higher elevations was directly related to lower mean temperatures, which had no effect on abundance. Our study demonstrates that even along relatively short gradients, elevation can have strong direct and abundance‐mediated effects on species richness, but with effects varying from positive to negative signs depending on local resource availability and the characteristics of groups or trophic guilds. If elevation positively influences local environmental properties that benefit a given group, richness can increase towards higher elevations. Thus, the effect of global change in mountainous regions should be evaluated within the local environmental context using multi‐taxon approaches.