Eyal Shochat

Trophic Dynamics in Urban Communities

Abstract Human activities dramatically change the abundance, diversity, and composition of species. However, little is known about how the most intense human activity, urbanization, alters food webs and trophic structure in biological communities. Studies of the Phoenix area, situated amid the Sonoran Desert, reveal some surprising alterations in the control of trophic dynamics. Species composition is radically altered, and resource subsidies increase and stabilize productivity. Changes in productivity dampen seasonal and yearly fluctuations in species diversity, elevate abundances, and alter feeding behaviors of some key urban species. In urban systems—in contrast to the trophic systems in outlying deserts, which are dominated by limiting resources—predation by birds becomes the dominant force controlling arthropods on plants. Reduced predation risk elevates the abundance of urban birds and alters their foraging behavior such that they exert increased top-down effects on arthropods. Shifts in control of food web dynamics are probably common in urban ecosystems, and are influenced by complex human social processes and feedbacks.

Invasion, Competition, and Biodiversity Loss in Urban Ecosystems

The global decline in biodiversity as a result of urbanization remains poorly understood. Whereas habitat destruction accounts for losses at the species level, it may not explain diversity loss at the community level, because urban centers also attract synanthropic species that do not necessarily exist in wildlands. Here we suggest an alternative framework for understanding this phenomenon: the competitive exclusion of native, non-synanthropic species by invasive species. We use data from two urban centers (Phoenix and Baltimore) and two taxa (birds and spiders) to link diversity loss with reduced community evenness among species in urban communities. This reduction in evenness may be caused by a minority of invasive species dominating the majority of the resources, consequently excluding nonsynanthropic species that could otherwise adapt to urban conditions. We use foraging efficiency as a mechanism to explain the loss of diversity. Thus, to understand the effects of habitat conversion on biodiversity, and to sustain species-rich communities, future research should give more attention to interspecific interactions in urban settings.

Urbanization and Spider Diversity: Influences of Human Modification of Habitat Structure and Productivity

As a part of the Central Arizona–Phoenix Long-Term Ecological Research project, we determined how land-use alteration influenced spider and harvestman diversity. We sampled spiders in six habitat types (desert parks, urban desert remnants, industrial, agricultural, xeric- and mesic-residential yards) and tested how habitat type and productivity affected spider diversity and abundance. As expected, agricultural fields and mesic yards were more productive than the other, xeric habitats. These more productive habitats were characterized by higher abundances but lower spider diversity and were dominated by Lycosidae (wolf spiders), followed by Linyphiidae (sheet-web weavers). The increase in wolf spider abundance was positively correlated with habitat productivity and negatively correlated with the abundance of other predatory arthropods that might compete with, or prey upon, wolf spiders.

Linking Optimal Foraging Behavior to Bird Community Structure in an Urban‐Desert Landscape: Field Experiments with Artificial Food Patches

Urban bird communities exhibit high population densities and low species diversity, yet mechanisms behind these patterns remain largely untested. We present results from experimental studies of behavioral mechanisms underlying these patterns and provide a test of foraging theory applied to urban bird communities. We measured foraging decisions at artificial food patches to assess how urban habitats differ from wildlands in predation risk, missed‐opportunity cost, competition, and metabolic cost. By manipulating seed trays, we compared leftover seed (giving‐up density) in urban and desert habitats in Arizona. Deserts exhibited higher predation risk than urban habitats. Only desert birds quit patches earlier when increasing the missed‐opportunity cost. House finches and house sparrows coexist by trading off travel cost against foraging efficiency. In exclusion experiments, urban doves were more efficient foragers than passerines. Providing water decreased digestive costs only in the desert. At the population level, reduced predation and higher resource abundance drive the increased densities in cities. At the community level, the decline in diversity may involve exclusion of native species by highly efficient urban specialists. Competitive interactions play significant roles in structuring urban bird communities. Our results indicate the importance and potential of mechanistic approaches for future urban bird community studies.

Causes and Patterns of Mortality in Lesser Prairie-chickens Tympanuchus pallidicinctus and Implications for Management

Abstract Life-history studies of prairie grouse have focused on reproductive ecology, habitat use, movement patterns and survivorship, with only cursory or anecdotal references to mortality causes, or they have been of insufficient duration or scale to infer mortality patterns. Because mortality causes and patterns affect other life-history traits, their determination adds to our overall understanding of grouse demographics. As part of a long-term study on lesser prairie-chicken Tympanuchus pallidicinctus natural history in Oklahoma and New Mexico, we recovered 322 carcasses of radio-tagged birds captured on leks. We were able to determine the cause of death for 260 of these birds. Predation by raptors accounted for the largest number of mortalities (91), followed by collisions with fences (86), predation by mammals (76), collisions with power lines (4), and collisions with automobiles (3). Mortality causes differed considerably between study sites and between sexes, with all collisions more frequent in Oklahoma than in New Mexico, in females than in males, and in older than in young females. Although predation is a major cause of mortality, we argue that predator control may not be effective for grouse conservation. Moreover, in cases where top predators reduce mesopredator population densities, for example those of red foxes Vulpes vulpes, indiscriminate removal of predators may hasten the decline of grouse populations. Land managers striving to conserve prairie-chickens and other grouse species should attempt to reduce or eliminate collision mortality risks in addition to efforts to improve nesting or brood-rearing habitat. Collision risks should also be evaluated for potential release sites of translocated or captive-reared grouse.

Habitat Edge, Land Management, And Rates Of Brood Parasitism In Tallgrass Prairie

Bird populations in North Americas grasslands have declined sharply in recent decades. These declines are traceable, in large part, to habitat loss, but management of tallgrass prairie also has an impact. An indirect source of decline potentially associated with management is brood parasitism by the Brown-headed Cowbird (Molothrus ater), which has had substantial negative impacts on many passerine hosts. Using a novel application of regression trees, we analyzed an extensive five-year set of nest data to test how management of tallgrass prairie affected rates of brood parasitism. We examined seven landscape features that may have been associated with parasitism: presence of edge, burning, or grazing, and distance of the nest from woody vegetation, water, roads, or fences. All five grassland passerines that we included in the analyses exhibited evidence of an edge effect: the Grasshopper Sparrow (Ammodramus savannarum), Henslows Sparrow (A. henslowii), Dickcissel (Spiza americana), Red-winged Blackbird (Agelaius phoeniceus), and Eastern Meadowlark (Sturnella magna). The edge was represented by narrow strips of woody vegetation occurring along roadsides cut through tallgrass prairie. The sparrows avoided nesting along these woody edges, whereas the other three species experienced significantly higher (1.9-5.3x) rates of parasitism along edges than in prairie. The edge effect could be related directly to increase in parasitism rate with decreased distance from woody vegetation. After accounting for edge effect in these three species, we found evidence for significantly higher (2.5-10.5x) rates of parasitism in grazed plots, particularly those burned in spring to increase forage, than in undisturbed prairie. Regression tree analysis proved to be an important tool for hierarchically parsing various landscape features that affect parasitism rates. We conclude that, on the Great Plains, rates of brood parasitism are strongly associated with relatively recent road cuts, in that edge effects manifest themselves through the presence of trees, a novel habitat component in much of the tallgrass prairie. Grazing is also a key associate of increased parasitism. Areas managed with prescribed fire, used frequently to increase forage for grazing cattle, may experience higher rates of brood parasitism. Regardless, removing trees and shrubs along roadsides and refraining from planting them along new roads may benefit grassland birds.

Inherited microbial symbionts increase herbivore abundances and alter arthropod diversity on a native grass

Some microbial symbionts of plants are maternally inherited and thus functionally increase genetic and phenotypic variation within plant populations. This variation, coupled with that of the host plant and environment, may alter abundances, diversity, and trophic structure of associated plant and animal communities. Fungal endophytes in the genus Neotyphodium are vertically transmitted, asexual microbial symbionts of grasses that remain asymptomatic and rely upon their hosts for resources and transmission via seeds, often providing benefits to their hosts, including protection against herbivores. Endophyte infections may influence associated arthropod communities in agronomic grasses, but the long-term effects of endophytes and variation in host genotype and resource availability on arthropod communities in native grass populations are unknown. We conducted a long-term field experiment with four maternal genotypes of an infected (E+) native grass (Festuca arizonica) from whence the endophyte was experimentally removed (E-) and water availability was controlled, to test the effects of infection, plant genotype, and resources on abundances, biomass, diversity (richness and evenness), and trophic structure of the arthropod community. Generally, E+ grasses harbored more arthropods, including more herbivores, predators, and detritivores, suggesting that the effects of endophytes cascaded upward through trophic levels in terms of abundances, at least in early ontogeny of the host. That E+ plants harbored more herbivorous insects than E- plants suggests that infection does not increase but instead decreases resistance to herbivores, contrary to prevailing concepts of endophytes as defensive mutualists. Infection did not alter overall species richness of the arthropod community or richness of herbivores but reduced natural enemy richness, especially that of parasites, and increased richness of detritivores. Reduced richness and shifts in evenness of natural enemies on E+ plants suggest that endophytes may disproportionately affect diversity at higher trophic levels and may partially explain increases in abundances of herbivorous insects on E+ plants. Biomass of predators, detritivores, and omnivores increased on plants with supplemented water, and arthropod and herbivore biomass varied by plant genotype. Symbiont-mediated phenotypic variation interacts with variation from plant genotype and environmental factors to alter arthropod abundances and diversity, and these effects shift with ontogeny of the host.

Urban Food Webs: Predators, Prey, and the People Who Feed Them

A prevailing image of the city is of the steel and concrete downtown skyline. The more common ex‐ perience of urban residents, however, is a place of irrigated and fertilized green spaces, such as yards, gardens, and parks, surrounding homes and business‐ es where people commonly feed birds, squirrels, and other wildlife. Within these highly human-modified environments, researchers are becoming increasingly curious about how fundamental ecological phenom‐ ena play out, such as the feeding relationships among species. While food webs have long provided a tool for organizing information about feeding relation‐ ships and energy flows through natural habitats, they have not been applied to urban ecosystems until re‐ cently (Faeth et al. 2005).

Linking Foraging Decisions to Residential Yard Bird Composition

Urban bird communities have higher densities but lower diversity compared with wildlands. However, recent studies show that residential urban yards with native plantings have higher native bird diversity compared with yards with exotic vegetation. Here we tested whether landscape designs also affect bird foraging behavior. We estimated foraging decisions by measuring the giving-up densities (GUD; amount of food resources remaining when the final forager quits foraging on an artificial food patch, i.e seed trays) in residential yards in Phoenix, AZ, USA. We assessed how two yard designs (mesic: lush, exotic vegetation; xeric: drought-tolerant and native vegetation) differed in foraging costs. Further, we developed a statistical model to calculate GUDs for every species visiting the seed tray. Birds foraging in mesic yards depleted seed trays to a lower level (i.e. had lower GUDs) compared to birds foraging in xeric yards. After accounting for bird densities, the lower GUDs in mesic yards appeared largely driven by invasive and synanthropic species. Furthermore, behavioral responses of individual species were affected by yard design. Species visiting trays in both yard designs had lower GUDs in mesic yards. Differences in resource abundance (i.e., alternative resources more abundant and of higher quality in xeric yards) contributed to our results, while predation costs associated with foraging did not. By enhancing the GUD, a common method for assessing the costs associated with foraging, our statistical model provided insights into how individual species and bird densities influenced the GUD. These differences we found in foraging behavior were indicative of differences in habitat quality, and thus our study lends additional support for native landscapes to help reverse the loss of urban bird diversity.