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Urbanization drives unique latitudinal patterns of insect herbivory and tree condition
Abstract
Urban landscapes are characterized by high proportions of impervious surface resulting in higher temperatures than adjacent natural landscapes. In some cities, like those at cooler latitudes, trees may benefit from warmer urban temperatures, but trees in many cities are beset with problems like drought stress and increased herbivory. What drives patterns of urban tree health across urbanization and latitudinal temperature gradients? In natural systems, latitude–herbivory relationships are well‐studied, and recent temperate studies have shown that herbivory generally increases with decreasing latitudes (warmer temperatures). However, the applicability of this latitude–herbivory theory in already‐warmed urban systems is unknown. In this study, we investigated how the interaction of urbanization, latitudinal warming and scale insect abundance affected urban tree health. We predicted that trees in warmer, lower latitude cities would be in poorer health at lower levels of urbanization than trees at cooler, higher latitudes due to the interaction of urbanization, latitudinal temperature and herbivory. To evaluate our predictions, we surveyed the abundance of scale insect herbivores on a single, common tree species Acer rubrum in eight US cities spanning 10° of latitude. We estimated urbanization at two extents, a local one that accounted for the direct effects on an individual tree, and a larger one that captured the surrounding urban landscape. We found that urban tree health did not vary with latitudinal temperature but was best predicted by local urbanization and herbivore abundance. We did not observe increased herbivore abundance in warmer, lower latitudes cities, but instead herbivore abundance peaked in the mid latitudes of our study. This study demonstrates that urban landscapes may deviate from classical theory developed in natural systems and reinforces the need for research reconciling ecological patterns in urban landscapes.
... Their primary distribution is in the United States, and they have been regularly recorded from Maryland to Florida and west to Texas (Ferris 1941, Johnson and Lyon 1976, Waltman et al. 2016. Gloomy scales are one of the most important ornamental tree pests in the mid-Atlantic and southeast United States (Frank 2019), with greatest densities in the midlatitudes of the eastern U.S. distribution (Just et al. 2018(Just et al. , 2019. The geographic range of gloomy scale is not likely limited by dispersal or host availability, since red maples (Acer rubrum L.; Sapindales: Sapindaceae) and other hosts are nearly ubiquitous. ...
... Gloomy scales are most tolerant of the thermal conditions in the middle of their primary distribution, near Raleigh, NC. This area is where densities are greatest and winter temperatures rarely descend below −1.0°C (Just et al. 2019, Just andFrank 2020). ...
... Within cities, gloomy scale body size, reproductive output, and survival is positively correlated with warmer temperatures and the amount of surrounding impervious surface cover (Dale and Frank 2014a, Youngsteadt et al. 2015, Dale et al. 2016). Gloomy scale is most abundant on host trees in the warmest parts of cities Frank 2014a, 2017) and those hosts with a high proportion of circumjacent impervious surface cover (Dale et al. 2016, Just et al. 2019. For example, gloomy scale density was 300 times greater on red maples with the warmest canopy temperatures versus those just 2.5°C cooler in Raleigh, NC (Dale and Frank 2014a). ...
Gloomy scale, Melanaspis tenebricosa (Comstock), is native to the eastern United States and feeds on deciduous trees. In natural areas, it is a background herbivore that typically remains at low densities. Gloomy scale generally responds positively to warming with greater egg production, size, survival, and abundance. In urban areas, which are warmer than surrounding natural areas, gloomy scale is pestiferous on planted trees, particularly red maple (Acer rubrum L.; Sapindales: Sapindaceae) but other native maples as well. They live on the bark and damage host trees by feeding from plant cells and tissues, which deprives the trees of energy and nutrients, reducing the trees’ growth and overall health. Gloomy scales are likely to expand their range beyond the Southeast and become pestilent in new areas with continued climatic warming and urbanization. Here we present a review of the biology, ecology, response to environmental conditions, host range and damage, and management of gloomy scale.
... These scale species develop invasive traits, proliferate, and become chronic urban tree pests due to the UHI effect [49][50][51]. Other scale species on the same trees under the same conditions do not display invasive phenotypes or become pests [52,53]. Melanaspis tenebricosa and P. quercifex are consistent with characterizations of sleeper species as they are innocuous native species that become pests with environmental change, here, the UHI effect. ...
... The strongest path indicated that impervious surface cover increased temperature which, in turn, increased scale density. Subsequent research on urban red maples corroborated these results [53,56,61,87]. ...
... In 1922, the range of M. tenebricosa was delimited to include states from Maryland in the north to Florida in south along the east coast of the USA and westward from Missouri to Texas. Recent surveys e.g., [53,85] have found them throughout the southeastern US, in agreement with the Metcalf (1922) map. We surveyed red maple street trees in Boston, MA, Queens, NY, Philadelphia, PA, Newark, DE, and Baltimore, MD, along the east coast of the US. ...
Sleeper species are innocuous native or naturalized species that exhibit invasive characteristics and become pests in response to environmental change. Climate warming is expected to increase arthropod damage in forests, in part, by transforming innocuous herbivores into severe pests: awakening sleeper species. Urban areas are warmer than natural areas due to the urban heat island effect and so the trees and pests in cities already experience temperatures predicted to occur in 50-100 years. We posit that arthropod species that become pests of urban trees are those that benefit from warming and thus should be monitored as potential sleeper species in forests. We illustrate this with two case studies of scale insects that are important pests of urban trees in parts of the US. Melanaspis tenebricosa and Parthenolecanium quercifex are geographically native to the US but take on invasive characteristics such as higher survival and reproduction and become disconnected from natural enemies on urban trees due to the urban heat island effect. This allows them to reach high densities and damage their host trees. Parthenolecanium quercifex density increases up to 12 times on urban willow oaks with just 2 °C of warming due to higher survival and adaptation to warmer temperatures. The urban heat island effect also creates a phenological mismatch between P. quercifex and its parasitoid complex, and so egg production is higher. Melanaspis tenebricosa density can increase 300 times on urban red maples with 2.5 °C of warming. This too is due to direct effects of warmer temperatures on survival and fecundity but M. tenebricosa also benefits from the drought stress incurred by warmer urban trees. These effects combine to increase M. tenebricosa density in forests as well as on urban trees at latitudes higher than its native range. We illustrate how cities provide a unique opportunity to study the complex effects of warming on insect herbivores. Studying pestilent urban species could be a pragmatic approach for identifying and preparing for sleeper species.
... We also measured in situ overwintering survival of M. tenebricosa at northern, mid, and southern locations within their geographic distribution collected throughout winter. Because M. tenebricosa densities are the greatest in the midlatitudes of their distribution (Metcalf 1922, Just et al. 2019, we predicted that survival would be highest at temperatures from these latitudes. Finally, because scale insect thermal tolerances can vary by season (e.g., Zhang et al. 2010, Wang et al. 2019a, we predicted that cold tolerance would decline in late winter when conditions become milder. ...
... These collection cities largely encompass the latitudinal extent of the known distribution of M. tenebricosa (Metcalf 1922), with Newark, Raleigh, and Gainesville located at the approximate northern, mid, and southern distribution extents, respectively. We collected in Raleigh more often because of greater M. tenebricosa densities at this latitude (Just et al. 2019) and practical considerations, such as proximity to the authors. In each city, we identified three sites with infested A. rubrum (n = 2-4 trees per site). ...
... In this study, we present the first assessment of M. tenebricosa physiological thermal tolerance, including tolerances at representative temperatures from throughout their geographic distribution. Given that the density of M. tenebricosa peaks in the mid latitudes of its distribution (Just et al. 2018(Just et al. , 2019, we predicted that temperatures representative of the midlatitudes (e.g., Raleigh, NC) would be the least lethal for M. tenebricosa and, thus, their thermal tolerance, measured here as upper and lower lethal thermal durations for a given temperature, would be the longest. Looking at our heat tolerance experiments, we found support for our prediction as Raleigh 2018 scales tolerated the Raleigh-derived temperatures best, providing the longest ULD 50 at 32.2°C (mean maximum July temperature) of 27.7 h (Table 3). ...
An insect species' geographic distribution is probably delimited in part by physiological tolerances of environmental temperatures. Gloomy scale (Melanaspis tenebricosa (Comstock)) is a native insect herbivore in eastern U.S. forests. In eastern U.S. cities, where temperatures are warmer than nearby natural areas, M. tenebricosa is a primary pest of red maple (Acer rubrum L.; Sapindales: Sapindaceae) With warming, M. tenebricosa may spread to new cities or become pestilent in forests. To better understand current and future M. tenebricosa distribution boundaries, we examined M. tenebricosa thermal tolerance under laboratory conditions. We selected five hot and five cold experimental temperatures representative of locations in the known M. tenebricosa distribution. We built models to predict scale mortality based on duration of exposure to warm or cold experimental temperatures. We then used these models to estimate upper and lower lethal durations, i.e., temperature exposure durations that result in 50% mortality. We tested the thermal tolerance for M. tenebricosa populations from northern, mid, and southern locations of the species' known distribution. Scales were more heat and cold tolerant of temperatures representative of the midlatitudes of their distribution where their densities are the greatest. Moreover, the scale population from the northern distribution boundary could tolerate cold temperatures from the northern boundary for twice as long as the population collected near the southern boundary. Our results suggest that as the climate warms the M. tenebricosa distribution may expand poleward, but experience a contraction at its southern boundary.
... Fifty per cent of scales die within 5 hours at −15.2 • C which frequently occurs at the northern edge of their range (Frank andJust 2020, Just and. However, the urban heat island effect allows gloomy scales to persist on urban trees north of their previously recorded range (Just et al. 2019, Frank andJust 2020). The frequency and rate of freezing define the geographic extent and severity of black pineleaf scale outbreaks in the Pacific Northwest, US (Edmunds 1973). ...
... Red maples are hosts for many scale insect species in the eastern US (Johnson andLyon 1976, Frank et al. 2013). A few species can reach high densities on urban trees in the southern US but only gloomy scales have been found to consistently increase with temperature making it the primary long-term pest of urban maples (Metcalf 1912, Dale and Frank 2014b, a, Dale and Frank 2017, Frank 2019, Just et al. 2019, Frank and Just 2020. Likewise, Parthenolecanium spp. ...
Higher temperatures and drought are key aspects of global change with the potential to alter the distribution and severity of many arthropod pests in forest systems. Scale insects (Hemiptera: Coccoidea) infest many tree species and are among the most important pests of trees in urban and rural forests, plantations and other forest systems. Infestations of native or exotic scale insects can kill or sicken trees with economic and ecosystem-wide consequences. Warming can have direct effects on the life history, fitness and population dynamics of many scale insect species by increasing development rate, survival or fecundity. These direct benefits can increase the geographic distribution of scale insects and their consequences for tree health. Warming and drought can affect scale insects indirectly by altering the quality of their host trees. Additive or interactive effects of warming and drought can change tree quality in such a way that it increases scale insect fitness and population growth. However, the effects are species- and context-dependent with some scale insect species negatively affected by drought-induced changes in tree quality. Warming and drought are often coincident in urban forests and predicted to co-occur in many parts of the world under climate change scenarios. The individual and interactive effects of these factors require further research to inform predictions and management of scale insect pests. Warming also indirectly affects scale insects by altering interactions with natural enemies. This includes changes in natural enemy phenology, community composition and abundance. In addition, warming can alter scale insect phenology or voltinism causing asynchrony with natural enemies or population growth too rapid for natural enemies to suppress. Direct and indirect effects of warming and drought on scale insects can increase the potential for some exotic species to become established and for some native species to become invasive. Unfortunately, much research on scale insects is confined to a few particularly important native or exotic pests which limits our ability to predict the effects of warming on many current or potential pests. More research is required to understand how warming and drought affect scale insects, scale insect management and the forest systems they inhabit.
... . High scale densities can worsen tree condition and reduce their aesthetic value Just et al. 2019). However, trees growing where less impervious surface cover is present, such as in yards and parks, host lower scale densities that tend not to cause damage or reduce growth (Dale et al. 2016;Just et al. 2018;Meineke and Frank 2018). ...
Scale insects are common tree pests in
urban ecosystems. Although severe scale infestations
can worsen tree condition, trees can tolerate moderate
scale densities. Scale insects are prey for many arthro�pod natural enemies that also feed on plant pests
throughout urban landscapes. Because scale-infested
trees support natural enemy communities, they may
support biological control services on nearby plants
and function analogously to banker plants in greenhouse production systems. In this study we tested if sentinel insect prey were more likely to be removed on shrubs below scale-infested trees compared to scale-uninfested trees. We conducted several biological control experiments from 2019 to 2021 using fruit flies, aphids, and caterpillars in potted and planted holly shrubs below scale-infested and scale-uninfested oak trees. We found that caterpillars in potted. shrubs and fruit flies in planted landscape shrubs were more likely to be removed underneath scale-infested trees compared to scale-uninfested trees. Caterpillars were also more likely to be removed from landscape Ilex vomitoria shrubs compared to I. cornuta shrubs. In all other experiments we found no effect of scale infestation status or shrub species on prey removal. Our results suggest that scale-infested trees can support biological control services in shrubs below them but that this effect can vary depending on prey and shrub species. The natural enemy communities in urban trees and shrubs appear to be linked and tolerating tree pests can favor conservation biological control services in urban landscapes.
... (QGIS Development Team 2022). We used the zonal statistics tool to calculate the percentage of impervious surface within a 20 meter buffer surrounding each sampled tree, which is more predictive of herbivorous insect density than impervious surface at larger spatial scales (Just et al. 2019). Although impervious surface data were not available at a smaller spatial scale, the zonal statistics tool allowed Urban environments have species-specific associations with invasive insect herbivores | 3 us to obtain an estimate of impervious surface within 20 m of each tree using 30 Â 30 meter data by computing an average impervious surface value based on weighted averages of the extent to which each 30 Â 30 meter pixel overlapped with the 20 meter buffer around a tree. ...
Urban forests are critically important for providing ecosystem services to rapidly expanding urban populations, but their health is threatened by invasive insect herbivores. To protect urban forests against invasive insects and support future delivery of ecosystem services, we must first understand the factors that affect insect density across urban landscapes. This study explores how a variety of environmental factors that vary across urban habitats influence density of invasive insects. Specifically, we evaluate how vegetational complexity, distance to buildings, impervious surface, canopy temperature, host availability and density of co-occurring herbivores impact three invasive pests of elm trees: the elm leaf beetle (Xanthogaleruca luteola), the elm flea weevil (Orchestes steppensis) and the elm leafminer (Fenusa ulmi). Insect responses to these factors were species-specific, and all environmental factors were associated with density of at least one pest species except for distance to buildings. Elm leafminer density decreased with higher temperatures and was influenced by an interaction between vegetational complexity and impervious surface. Elm flea weevil density increased with greater host availability, and elm leaf beetle density increased with higher temperatures. Both elm leaf beetle and elm flea weevil density decreased with greater leafminer density, suggesting that insect density is mediated by species interactions. These findings can be used to inform urban pest management and tree care efforts, making urban forests more resilient in an era when globalization and climate change make them particularly vulnerable to attack.
... Acer rubrum represents a significant proportion of the urban and natural forest species composition throughout the eastern U.S. and has traits that many urban trees are selected for, such as rapid growth, high aesthetic quality, dense canopies, and moderate tolerance to water stress [66]. Although previous work has found that the effects of local abiotic factors on urban Acer rubrum pests and tree condition persist across latitudes [45,51], the implications of our results for urban forest health may not translate to other tree species. Therefore, future work should investigate similar interactions with other common urban tree taxa to develop more comprehensive guidelines for urban forest managers. ...
Warmer temperatures and frequent drought directly affect urban tree health. Both abiotic conditions also affect tree health via increased density of some insect pests. Warming is predicted to benefit urban trees by increasing carbon sequestration and allocation to biomass. However, increased drought and pests are rarely considered despite often co-occurring with heat. To determine the combined effects of these abiotic and biotic factors, we manipulated water availability for established urban red maple trees across a gradient of warming and pest density and measured leaf-level processes and tree growth over two years. We find that water availability is a major determinant of tree growth, physiological processes, and resilience to urban stress factors. Maples performed better with more water, which also made them resistant to effects of temperature and pest density. However, when drought became too severe, leaf-level processes declined with warming. Tree basal area growth was unaffected after two years, but stem elongation increased with increasing water, temperature, and pest density. We discuss potential mechanisms driving these responses and the implications in the context of urban forest management. Urban forest designs that reduce drought and align species adaptations to local conditions are critical for designing more resilient and productive urban forests.
... Increased recognition of the high value of trees to urban environments has encouraged municipal tree planting programs around the world [3]. Although it has been reported that trees at higher, cooler latitudes may benefit from urban warming [4,5], urban trees, in general, are under continuous pressure from environmental problems typical to the urban environment. Paved surfaces modify the moisture dynamics of underlying soils [6], increasing the risk of water stress for trees, and therefore, their susceptibility to pests [7][8][9]. ...
Declining urban tree health can affect critical ecosystem services, such as air quality improvement, temperature moderation, carbon storage, and biodiversity conservation. The application of state-of-the-art remote sensing data to characterize tree health has been widely examined in forest ecosystems. However, such application to urban trees has not yet been fully explored—due to the presence of heterogeneous tree species and backgrounds, severely complicating the classification of tree health using remote sensing information. In this study, tree health was represented by a set of field-assessed tree health indicators (defoliation, discoloration, and a combination thereof), which were classified using airborne laser scanning (ALS) and hyperspectral imagery (HSI) with a Random Forest classifier. Different classification scenarios were established aiming at: (i) Comparing the performance of ALS data, HSI and their combination, and (ii) examining to what extent tree species mixtures affect classification accuracy. Our results show that although the predictive power of ALS and HSI indices varied between tree species and tree health indicators, overall ALS indices performed better. The combined use of both ALS and HSI indices results in the highest accuracy, with weighted kappa coefficients (Kc) ranging from 0.53 to 0.79 and overall accuracy ranging from 0.81 to 0.89. Overall, the most informative remote sensing indices indicating urban tree health are ALS indices related to point density, tree size, and shape, and HSI indices associated with chlorophyll absorption. Our results further indicate that a species-specific modelling approach is advisable (Kc points improved by 0.07 on average compared with a mixed species modelling approach). Our study constitutes a basis for future urban tree health monitoring, which will enable managers to guide early remediation management.
... Although latitude can be a useful integrator of several axes of global variation in climate, relatively few studies investigate the hypothesized underlying climatic drivers behind latitudinal patterns in herbivory (e.g., Adams & Zhang, 2009;Gao et al., 2019;Moreira et al., 2015;Zhang, Zhang, & Ma, 2016), while even fewer consider nutrient availability that can have different spatial patterns than climate variables (e.g., Lynn & Fridley, 2019;Moreira, Castagneryrol, Abdala-Roberts, & Berny-Mier y Teran JC, Timmermans BGH, Bruun HH, Covelo, F, Glauser G, Rasmann S, Tack AJM., 2018). In addition, it is possible that latitudinal variation in insect herbivore damage and plant resistance to herbivory are driven by resource availability, trade-offs in plant growth and defenses (Kim, 2014), and that they depend on herbivore specialization (Dyer & Forister, 2019), as well as land use or urbanization (Just, Dale, Long, & Frank, 2019). Though climate is surely a strong driver of latitudinal variation in herbivory, deviations from this expected pattern are likely due to unmeasured edaphic and plant-trait controls on consumption rates. ...
Patterns of insect herbivory may follow predictable geographical gradients, with greater herbivory at low latitudes. However, biogeographic studies of insect herbivory often do not account for multiple abiotic factors (e.g., precipitation and soil nutrients) that could underlie gradients. We tested for latitudinal clines in insect herbivory as well as climatic, edaphic, and trait‐based drivers of herbivory. We quantified herbivory on five dominant grass species over 23 sites across the Great Plains, USA. We examined the importance of climate, edaphic factors, and traits as correlates of herbivory. Herbivory increased at low latitudes when all grass species were analyzed together and for two grass species individually, while two other grasses trended in this direction. Higher precipitation was related to more herbivory for two species but less herbivory for a different species, while higher specific root length was related to more herbivory for one species and less herbivory for a different species. Taken together, results highlight that climate and trait‐based correlates of herbivory can be highly contextual and species‐specific. Patterns of insect herbivory on dominant grasses support the hypothesis that herbivory increases toward lower latitudes, though weakly, and indicates that climate change may have species‐specific effects on plant–herbivore interactions.
... The former mainly links to the socio-economic dimension of urbanization and the latter focuses more on urban biophysical and environmental factors. These two kinds of indices are more straightforward and easily obtained from urban census data or retrieved from remotely sensed data, which have been widely utilized to study urbanization and their relationships to, or impacts on urban biodiversity [72,[100][101][102][103]. Compared with most of the previous studies, our study directly used the urban land use proportion to quantify the urbanization degree and established a quantitative relationship to plant species diversity that can help easily understand and predict the influences of urbanization on plant diversity; therefore, this study can provide insight to optimize ecosystem services in future urban green space planning and design [104][105][106]. ...
Urbanization is one of the major causes for plant diversity loss at the local and regional scale. However, how plant species distribute along the urban–rural gradient and what the relationship between urbanization degree and plant diversity is, is not very clear. In this paper, 134 sample sites along two 18 km width transects that run across the urban center of Shanghai were investigated. We quantified the spatial patterns of plant diversity along the urban–rural gradient and measured the relationship between plant diversity and urbanization degree, which was calculated using a land use land cover map derived from high spatial resolution aerial photos. We recorded 526 vascular plant species in 134 plots, 57.8% of which are exotic plant species. Six spatial distribution patterns of species richness were identified for different plant taxa along the rural to urban gradient. The native plant species richness showed no significant relationship to urbanization degree. The richness of the all plants, woody plants and perennial herbs presented significant positive relationship with urbanization degree, while the richness of annual herbs, Shannon-Wiener diversity and Heip evenness all exhibited a negative relationship to urbanization degree. Urbanization could significantly influence plant diversity in Shanghai. Our findings can provide insights to understand the mechanism of urbanization effects on plant diversity, as well as plant diversity conservation in urban areas.
... Red maple is native to eastern North America and is commonly planted in urban areas (McPherson 2010, USDA NRCS 2019. Gloomy scale is a univoltine armored scale insect with a range extending from Florida to New York (Metcalf 1922, Nakahara 1982, Miller and Davidson 2005, with highest abundance at midlatitudes (e.g., North Carolina) (Just et al. 2019). First instar scales, called crawlers, emerge in June and July and walk short distances or are transported by wind (Beardsley and Gonzalez 1975, Miller and Davidson 2005. ...
Pest abundance on urban trees often increases with surrounding impervious surface. Gloomy scale (Melanaspis tenebricosa Comstock; Hemiptera: Diaspididae), a pest of red maples (Acer rubrum L.; Sapindales: Sapindaceae) in the southeast United States, reaches injurious levels in cities and reduces tree condition. Here, we use a chronosequence field study in Raleigh, NC, to investigate patterns in gloomy scale densities over time from the nursery to 13 yr after tree planting, with a goal of informing more efficient management of gloomy scale on urban trees. We examine how impervious surfaces affect the progression of infestations and how infestations affect tree condition. We find that gloomy scale densities remain low on trees until at least seven seasons after tree planting, providing a key timepoint for starting scouting efforts. Scouting should focus on tree branches, not tree trunks. Scale density on tree branches increases with impervious surface across the entire studied tree age range and increases faster on individual trees that are planted in areas with high impervious surface cover. There is a lag between the onset of pest infestations and a decline in tree condition, indicating that gloomy scale management should begin prior to a visible decline in tree condition. Our results inform management of gloomy scale in cities.
Urban trees often host greater insect pest abundance than trees in rural forests. This may be due, in part, to differences in tree diversity and canopy cover between these settings. Urban trees are often planted in isolation or monoculture, which favors pest accumulation. Gloomy scale, Melanaspis tenebricosa Comstock, is a pest of urban red maples (Acer rubrum L.) that is abundant where impervious surfaces dominate the local landscape. Increasing tree diversity and canopy cover around urban red maples may reduce gloomy scale abundance by supporting natural enemy communities. We investigated the effect that surrounding tree species richness and tree canopy cover had on gloomy scale abundance, natural enemy abundance, and biological control in red maple trees in Raleigh, NC, USA. We collected scales and natural enemies from red maples that spanned a gradient of tree species richness, canopy cover, and impervious surface values. We also measured gloomy scale parasitism and predation of sentinel prey in red maple canopies. Greater tree species richness and canopy cover were associated with lower gloomy scale density. Red maples in diverse settings also hosted fewer scales per natural enemy. Parasitoids were less common in maples in diverse settings, but generalist predator abundance was unaffected by tree diversity. Finally, tree species richness and canopy cover did not increase biological control of scales or sentinel prey. Our findings suggest that higher tree diversity and greater canopy cover may reduce gloomy scale density, but this is not entirely explained by the effects of natural enemies and biological control.
According to the Latitudinal Biotic Interaction Hypothesis (LBIH), the general increase in biodiversity towards lower latitudes can be partially explained by an increase in the intensity of biotic interactions. While LBIH received some support for plant-herbivores interactions, much less is known about how higher trophic levels may contribute to shape biotic interactions across latitudinal gradients. We hypothesized that the diversity of insectivorous birds increases towards lower latitude, leading to higher predation rates on insect herbivores.
We deployed plasticine caterpillars in 138 oak trees in 47 sites along a 19 degree latitudinal gradient in Europe to quantify bird insectivory through predation attempts. In addition, we used passive acoustic monitoring to (i) characterize the acoustic diversity of surrounding soundscapes; and (ii) infer both taxonomic and functional diversity of insectivorous birds from recordings.
The functional diversity of insectivorous birds increased towards lower latitude. Bird predation increased with latitude, forest cover and bird acoustic diversity but decreased with mean annual temperature and functional richness of insectivorous birds. Contrary to our predictions, latitudinal clines in bird predation attempts were not directly mediated by changes in insectivorous bird diversity or acoustic diversity, but latitude and habitat still had independent effects on predation attempts.
Our study does not fully support the predictions of the LBIH of more biotic interactions southwards and advocates for better accounting for activity and abundance of insectivorous birds when studying the large-scale variation in insect-tree interactions.
Cities are human dominated ecosystems providing novel conditions for organisms. Research on urban biodiversity is rapidly increasing, yet it is still hampered by the partial spatial coverage of cities and because of existing taxonomic biases. Predictive models have proved to be a key tool to solve this shortfall. However, predictive models have rarely been used in urban ecosystems due to either the lack of sufficient species records or high-quality predictors (e.g. meaningful ecological maps). Here, we assemble a large cross-taxa inventory of 1446 species from 12 taxonomic groups, including several understudied invertebrate groups, sampled in 251 sites in Zürich, Switzerland. We investigate the species diversity distributions and the structure of species assemblages along artificial urban ecological gradients by applying predictive models. We find that the general species diversity distribution law, where assemblages are dominated by a few very abundant and frequent species, applied consistently across all taxonomic groups (3% of the species accounting for approximately 50% of abundance). Furthermore, only species of intermediate abundance and frequency are spatially structured along urban intensity gradients, with rare species numbers keeping constant even in the most urbanised parts of the city. In addition, we show that green areas with low mowing regimes are associated with higher species diversity in the majority of taxonomic groups. Hence, this suggests management relaxation as a low-cost solution to promote species richness. Our study demonstrates the potential of predictive modelling for addressing ecological questions in urban environments and to inform management and planning.
Urbanization represents an unintentional global experiment that can provide insights into how species will respond and interact under future global change scenarios. Cities produce many conditions that are predicted to occur widely in the future, such as warmer temperatures, higher carbon dioxide (CO2) concentrations and exacerbated droughts. In using cities as surrogates for global change, it is challenging to disentangle climate variables-such as temperature-from co-occurring or confounding urban variables-such as impervious surface-and then to understand the interactive effects of multiple climate variables on both individual species and species interactions. However, such interactions are also difficult to replicate experimentally, and thus the challenges of cities are also their unique advantage. Here, we review insights gained from cities, with a focus on plants and arthropods, and how urban findings agree or disagree with experimental predictions and historical data. We discuss the types of hypotheses that can be best tested in cities, caveats to urban research and how to further validate cities as surrogates for global change. Lastly, we summarize how to achieve the goal of using urban species responses to predict broader regional- and ecosystem-level patterns in the future.
Urban habitats are characterized by impervious surfaces, which increase temperatures and reduce water availability to plants. The effects of these conditions on herbivorous insects are not well understood, but may provide insight into future conditions. Three primary hypotheses have been proposed to explain why multiple herbivorous arthropods are more abundant and damaging in cities, and support has been found for each. First, less complex vegetation may reduce biological control of pests. Second, plant stress can increase plant quality for pests. And third, urban warming can directly increase pest fitness and abundance. These hypotheses are not mutually exclusive, and the effects of temperature and plant stress are particularly related. Thus, we test the hypothesis that urban warming and drought stress combine to increase the fitness and abundance of the scale insect, Melanaspis tenebricosa, an urban tree pest that is more abundant in urban than rural areas of the southeastern U.S. We did this by manipulating drought stress across an existing mosaic of urban warming. We found support for the additive effect of temperature and drought stress such that female embryo production and body size increased with temperature and was greater on drought-stressed than watered trees. This study provides further evidence that drivers of pest insect outbreaks act in concert, rather than independently, and calls for more research that manipulates multiple abiotic factors related to urbanization and climate change to predict their effects on ecological interactions. As cities expand and the climate changes, warmer temperatures and drought conditions may become more widespread in the native range of this pest. These changes have direct physiological benefits for M. tenebricosa, and potentially other pests, that may increase their fitness and abundance in urban and natural forests.
A substantial amount of global carbon is stored in mature trees. However, no experiments to date test how warming affects mature tree carbon storage. Using a unique, citywide, factorial experiment, we investigated how warming and insect herbivory affected physiological function and carbon sequestration (carbon stored per year) of mature trees. Urban warming increased herbivorous arthropod abundance on trees, but these herbivores had negligible effects on tree carbon sequestration. Instead, urban warming was associated with an estimated 12% loss of carbon sequestration, in part because photosynthesis was reduced at hotter sites. Ecosystem service assessments that do not consider urban conditions may overestimate urban tree carbon storage. Because urban and global warming are becoming more intense, our results suggest that urban trees will sequester even less carbon in the future. © 2016 The Author(s) Published by the Royal Society. All rights reserved.
High-temperature tolerance in plants is important in a warming world, with extreme heat-waves predicted to increase in frequency and duration, potentially leading to lethal heating of leaves. Global patterns of high-temperature tolerance are documented in animals, but generally not plants, limiting our ability to assess risks associated with climate warming. To assess whether there are global patterns in high-temperature tolerance of leaf metabolism, we quantified Tcrit (high temperature where minimal chlorophyll a fluorescence rises rapidly, and thus where photosystem II is disrupted) and Tmax (temperature where leaf respiration in darkness is maximal, beyond which respiratory function rapidly declines) in upper-canopy leaves of 218 plant species spanning seven biomes. Mean site-based Tcrit values ranged from 41.5 °C in the Alaskan arctic to 50.8 °C in lowland tropical rainforests of Peruvian Amazon. For Tmax, the equivalent values were 51.0 and 60.6 °C in the Arctic and Amazon, respectively. Tcrit and Tmax followed similar biogeographic patterns, increasing linearly (˜8 °C) from polar to equatorial regions. Such increases in high temperature tolerance are much less than expected based on the 20 °C span in high temperature extremes across the globe. Moreover, with only modest high-temperature tolerance despite high summer temperature extremes, species in mid-latitude (~20°-50°) regions have the narrowest thermal safety margins in upper-canopy leaves; these regions are at the greatest risk of damage due to extreme heat-wave events, especially under conditions when leaf temperatures are further elevated by a lack of transpirational cooling. Using predicted heat-wave events for 2050 and accounting for possible thermal acclimation of Tcrit and Tmax, we also found that these safety margins could shrink in a warmer world, as rising temperatures are likely to exceed thermal tolerance limits. Thus, increasing numbers of species in many biomes may be at risk as heat-wave events become more severe with climate change. This article is protected by copyright. All rights reserved.
ContextSpatial heterogeneity has myriad influences on ecosystem processes, ecosystem services, and thus the sustainability of urban areas. It acts as a medium for urban design, planning, and management to determine how processes affecting sustainability can operate and interact. Therefore, how spatial heterogeneity is conceptualized and measured in cities is crucial for enhancing sustainability. Objectives
We show that the two most commonly used, but contrasting paradigms of urban ecology, ecology IN versus ecology OF the city, determine how spatial heterogeneity is thought of and used in different ways. We identify the key implications of these theoretical contrasts for the practice and assessment of sustainability in urban areas. Methods
We review and compare the different ways in which ecology IN versus ecology OF the city affect how to conceptualize, model and map urban spatial heterogeneity. We present a new framework to guide the comparison of spatial heterogeneity under the two paradigms. Results and conclusionThe integrative nature of this new framework becomes apparent under the ecology OF the city paradigm, because it recognizes the hybrid social and bioecological nature of heterogeneity in urban ecosystems. The hybrid approach to patchiness resonates with the three pillars of sustainability—environment, society, and economy. We exemplify how the more comprehensive and integrated framework of spatial heterogeneity under the ecology OF the city paradigm (1) supports more effective measurement and integration of the three components of sustainability, (2) improves management of heterogeneous urban ecosystems, and (3) satisfies calls for improved ecological tools to support urban ecosystem design.
Trees provide ecosystem services that benefit humans and the environment. Unfortunately, urban trees often do not provide maximum services due to abiotic stress and arthropod herbivores and borers. These problems often originate from trees being planted in unsuitable conditions. Cities are warmer than natural areas because impervious surfaces absorb and reradiate heat. Higher temperatures can increase pest insect abundance and water stress, and reduce street tree condition relative to natural forests. For example, the gloomy scale insect [Melanaspis tenebricosa Comstock (Hemiptera: Diaspididae)], a pest of red maple (Acer rubrum) street trees, is more abundant in warmer than cooler urban sites. Acer rubrum, at warmer urban sites with more M. tenebricosa, are typically in poor condition. Here, researchers demonstrate these relationships and illustrate how impervious surface cover can be used to predict the condition of A. rubrum street trees. Impervious surface thresholds were then developed to define suitable planting sites that can be used by individuals with access to GIS software. Researchers present the 'Pace-to-Plant' technique, which can be used by landscape professionals to quickly estimate impervious surface cover around a planting site. These thresholds predict future tree condition based on planting site impervious surface cover. The hope is that more informed planting will minimize pest infestations and maximize the future vigor and performance of street trees.
It has long been hypothesized that biotic interactions, including herbivory, are most intense at lower latitudes. However, this generalization has recently been challenged with studies showing that latitudinal gradients in damage may be rarer than previously believed. Additionally, most studies have focused on herbivory of native species, so it remains unknown whether natives and exotics follow similar patterns. This study compares rates of aboveground herbivory of multiple native and non-native Asteraceae across a latitudinal gradient, with a more detailed investigation of a focal exotic, Cirsium arvense. Herbivory of multiple tissue types was quantified for all species across an 815 km transect in Ontario, Canada. The native Asteraceae included in the survey typically experienced a decline in folivory with increasing latitude. Herbivory patterns for the exotic species were less clear; while most experienced high damage at the southernmost site, some also experienced high damage rates at mid-latitudes. For the focal species C. arvense, leaf and stem herbivory declined with increasing latitude, although seed damage showed strong regional variation across the invaded range. These results show that latitudinal variation in herbivory is highly dependent on the plant species being investigated, the tissue type being measured, and the type of herbivore(s) causing the damage. In some cases, populations in marginal areas might benefit from reduced damage by some groups of herbivores. In other cases, factors such as the availability of suitable habitat, the biology of specific enemies, and the origin of the host plant may override the influence of latitude on host performance.
1. In the past, it was widely accepted that herbivory decreased with latitude. However, several empirical studies have failed to support this hypothesis, leading to strong debate. In addition to providing a quantitative assessment of the latitudinal gradient in herbivory, our study aims to provide new information about the role of climatic factors in shaping the latitudinal gradient in herbivory.
2. Through the collection and analysis of published data on herbivory, we tested the latitudinal herbivory hypothesis (LH hypothesis) and the relationship of temperature and precipitation to herbivory.
3. Based on 1,890 data points distributed worldwide from more than 1,000 plant species, our results show that herbivory decreased with latitude and increased with temperature only in the Northern Hemisphere. In contrast, in the Southern Hemisphere, herbivory did not have a relationship with latitude and tended to decrease with temperature.
4. Synthesis. This study shows the LH hypothesis is supported only in the Northern Hemisphere, and highlights the importance of temperature in explaining the pattern of herbivory at the global scale. These possible hemispheric asymmetries in herbivory should not be overlooked in future studies.
Urban forest monitoring data are essential to assess the impacts of tree planting campaigns and management programs. Local practitioners have monitoring projects that have not been well documented in the urban forestry literature. To learn more about practitioner-driven monitoring efforts, the authors surveyed 32 local urban forestry organizations across the United States about the goals, challenges, methods, and uses of their monitoring programs, using an e-mailed questionnaire. Non-profit organizations, municipal agencies, state agencies, and utilities participated. One-half of the organizations had six or fewer urban forestry staff. Common goals for monitoring included evaluating the success of tree planting and management, taking a proactive approach towards tree care, and engaging communities. The most commonly recorded data were species, condition rating, mortality status, and diameter at breast height. Challenges included limited staff and funding, difficulties with data management and technology, and field crew training. Programs used monitoring results to inform tree planting and maintenance practices, provide feedback to individuals responsible for tree care, and manage tree risk. Participants emphasized the importance of planning ahead: carefully considering what data to collect, setting clear goals, developing an appropriate database, and planning for funding and staff time. To improve the quality and consistency of monitoring data across cities, researchers can develop standardized protocols and be responsive to practitioner needs and organizational capacities.
In light of catastrophic tree losses caused by Dutch elm disease, foresters recommended that the urban forest be diversified. The intent was to create a more sustainable urban forest that would not be decimated by a single pathogen or insect pest. However, recent introductions of deadly borers such as Asian longhorned beetle and emerald ash borer reveal that new introductions can have disastrous consequences for urban forests. An analysis of tree inventories from 12 cities in eastern North America reveals an overabundance of certain genera of trees such as Acer and Fraxinus. Introduced pests with broad host ranges such as the Asian longhorned beetle may be capable of killing or forcing intervention on large numbers of trees. Even cities that have diversified at a low taxonomic level (cultivar or species) may suffer greatly from the depredations of insect pests such as the emerald ash borer that specialize on plants at the generic level. Generalists capable of feeding on plants in several families will create problems for urban forests even when attempts have been made to diversify at higher taxonomic levels. Urban foresters should avoid planting susceptible taxa of trees, especially in cities that are overstocked in these taxa, and consider diversifying greatly the types of trees in cities.
Distinguishes the Urban Canopy Layer (UCL) from the ground to roof level, from the Urban Boundary Layer (UBL) which extends to the height where urban influences are no longer perceptible. Methods to establish the energy balance include micrometeorological approaches and numerical models. Data are presented under the heading of radiation budget, anthropogenic heat release, subsurface (storage) heat flux, turbulent heat transfer, advection, and urban-rural energy balance differences. -K.Clayton
The impact of anthropogenic climate change on terrestrial organisms is often predicted to increase with latitude, in parallel with the rate of warming. Yet the biological impact of rising temperatures also depends on the physiological sensitivity of organisms to temperature change. We integrate empirical fitness curves describing the thermal tolerance of terrestrial insects from around the world with the projected geographic distribution of climate change for the next century to estimate the direct impact of warming on insect fitness across latitude. The results show that warming in the tropics, although relatively small in magnitude, is likely to have the most deleterious consequences because tropical insects are relatively sensitive to temperature change and are currently living very close to their optimal temperature. In contrast, species at higher latitudes have broader thermal tolerance and are living in climates that are currently cooler than their physiological optima, so that warming may even enhance their fitness. Available thermal tolerance data for several vertebrate taxa exhibit similar patterns, suggesting that these results are general for terrestrial ectotherms. Our analyses imply that, in the absence of ameliorating factors such as migration and adaptation, the greatest extinction risks from global warming may be in the tropics, where biological diversity is also greatest.
• biodiversity
• fitness
• global warming
• physiology
• tropical
It is often claimed that evapotranspiration from urban trees can mitigate the urban heat island and adapt our cities to climate change; however, the relative effectiveness of different tree species has rarely been investigated. The current study addressed this shortcoming by comparing the growth and physiology of five commonly planted tree species: Sorbus arnoldiana, Crataegus laevigata, Malus ‘Rudolph’, Pyrus calleryana and Prunus ‘Umineko’. The study was conducted between March and November, 2011 in eight different streets of Manchester, UK where trees had been growing for 6 years in the same growing conditions. The study showed that evapotranspirational cooling is closely related to the growth and stress tolerance of tree species. Of the species tested, Prunus ‘Umineko’ and P. calleryana were the fastest growing and Malus ‘Rudolph’ was the slowest growing. In general faster growing species showed higher leaf area index (LAI) and higher stomatal conductivity and so provided more cooling. However, Prunus ‘Umineko’ had surprisingly low cooling and showed signs of drought stress. P. calleryana showed up to 100 % higher stomatal conductance than the other tree species. Combining the higher LAI and wider canopy, P. calleryana and C. laevigata provided cooling up to 2.2 kW tree−1, 3 to 4 times of cooling to that of Prunus ‘Umineko’ and S. arnoldiana and showed no signs of drought stress. Malus ‘Rudolph’ showed stress tolerance but provided low cooling. Prunus ‘Umineko’ and S. arnoldiana with their thin and sparse canopy provided low cooling and showed susceptibility to urban stress.
Trees are essential to urban habitats because they provide services that benefit the environment and improve human health. Unfortunately, urban trees often have more herbivorous insect pests than rural trees but the mechanisms and consequences of these infestations are not well documented. Here, we examine how temperature affects the abundance of a scale insect, Melanaspis tenebricosa (Comstock) (Hemiptera: Diaspididae), on one of the most commonly planted street trees in the eastern U.S. Next, we examine how both pest abundance and temperature are associated with water stress, growth, and condition of 26 urban street trees. Although trees in the warmest urban sites grew the most, they were more water stressed and in worse condition than trees in cooler sites. Our analyses indicate that visible declines in tree condition were best explained by scale-insect infestation rather than temperature. To test the broader relevance of these results, we extend our analysis to a database of more than 2700 Raleigh, US street trees. Plotting these trees on a Landsat thermal image of Raleigh, we found that warmer sites had over 70% more trees in poor condition than those in cooler sites. Our results support previous studies linking warmer urban habitats to greater pest abundance and extend this association to show its effect on street tree condition. Our results suggest that street tree condition and ecosystem services may decline as urban expansion and global warming exacerbate the urban heat island effect. Although our non-probability sampling method limits our scope of inference, our results present a gloomy outlook for urban forests and emphasize the need for management tools. Existing urban tree inventories and thermal maps could be used to identify species that would be most suitable for urban conditions.
The relationships among species' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals-a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species' physiology and the geography of climate change will advance assessments of species' vulnerability to climate change.
Maximum likelihood or restricted maximum likelihood (REML) estimates of the
parameters in linear mixed-effects models can be determined using the lmer
function in the lme4 package for R. As for most model-fitting functions in R,
the model is described in an lmer call by a formula, in this case including
both fixed- and random-effects terms. The formula and data together determine a
numerical representation of the model from which the profiled deviance or the
profiled REML criterion can be evaluated as a function of some of the model
parameters. The appropriate criterion is optimized, using one of the
constrained optimization functions in R, to provide the parameter estimates. We
describe the structure of the model, the steps in evaluating the profiled
deviance or REML criterion, and the structure of classes or types that
represents such a model. Sufficient detail is included to allow specialization
of these structures by users who wish to write functions to fit specialized
linear mixed models, such as models incorporating pedigrees or smoothing
splines, that are not easily expressible in the formula language used by lmer.
AimsUnderstanding how tree growth is influenced by climate is vital for predicting how forests will respond to climate change, yet there have been few studies of tree growth spanning macroclimatic gradients. The aim of this study is to correlate growth of a single lineage of broadleaf evergreen trees with continental-scale variability in climate.LocationAustralia's temperate mesic eucalypt forests, spanning latitudes from 23 to 43° S and longitudes from 115 to 153° E.Methods
We compiled and analysed a dataset containing around half a million measurements of growth in eucalypt tree diameter, collected from 2409 permanent forestry plots. These plots spanned a range of 558–2105 mm mean annual precipitation and 6–22 °C mean annual temperature. Generalized additive models were used to study the relationship between growth in tree diameter and several temperature and water availability variables.ResultsTree growth increased with precipitation, but with a diminishing response above a mean annual precipitation of 1400 mm. There was a peaked response to temperature, with maximum growth occurring at a mean annual temperature of 11 °C and maximum temperature of the warmest month of 25–27 °C. Lower temperatures directly constrain growth. High temperatures primarily reduced growth by reducing water availability, but they also appeared to exert a direct negative effect. Our best model, which included maximum temperature of the warmest month and the ratio of precipitation to evaporation, explained 28% of the deviance.Main conclusionsThe productivity of Australia's temperate eucalypt forests could decline substantially as the climate warms, given that 87% of these forests currently experience a mean annual temperature above 11 °C, where the highest growth rates were observed. This will reduce carbon sequestration and slow recovery after catastrophic disturbances such as wildfire.
Significance
We find that most terrestrial ectotherms are insufficiently tolerant of high temperatures to survive the warmest potential body temperatures in exposed habitats and must therefore thermoregulate by using shade, burrows, or evaporative cooling. Our results reveal that exposure to extreme heat can occur even at high elevations and latitudes and show why heat-tolerance limits are relatively invariant in comparison with cold limits. To survive climate warming, ectotherms in most areas may need to rely on behaviors—and have access to habitats—that provide a reprieve from extreme operative temperatures.
Favored for their rapid growth and brilliant fall color, maple (Acer spp.) trees are among the most commonly grown deciduous shade trees in urban landscapes and commercial production nurseries. Many maple species used as ornamental plants share a suite of important arthropod pests that have the potential to reduce the trees' economic and esthetic value. We review the biology, damage, and management for the most important pests of maples with emphasis on integrated pest management (IPM) tactics available for each pest. Unfortunately, the biology of some of these pests is not well studied. This knowledge gap, paired with the low esthetic threshold for damage on ornamental plants, has hindered development of IPM tactics for maple pests in nurseries and landscapes. Maples will likely remain a common landscape plant. Therefore, our challenge is to improve IPM of the diverse maple pest complex.
This paper presents and evaluates a method for the construction of long-range and wide-area temporal spatial datasets of daily precipitation and temperature (maximum and minimum). This method combines the interpolation of daily ratios/fractions derived from ground-based meteorological station records and respective fields of monthly estimates. Data sources for the described implementation over the conterminous United States (CONUS) are two independent and quality-controlled inputs: 1) an enhanced compilation of daily observations derived from the National Climatic Data Center digital archives and 2) the Parameter-Elevation Regressions on Independent Slopes Model (PRISM) maps. The results of this study show that this nonconventional interpolation preserves the spatial and temporal distribution of both the PRISM maps (monthly, topography-sensitive patterns) and the original daily observations. Statistics of a preliminary point comparison with the observed values at high-quality and independent reference sites show a reasonable agreement and a noticeable improvement over the nearest station method in orographically sensitive areas. The implemented datasets provide daily precipitation and temperature values at 2.5-min (around 4 km) resolution for 1960-2001. Combining seamless spatial and temporal coverage and topographic sensitivity characteristics, the datasets offer the potential for supporting current and future regional and historical hydrologic assessments over the CONUS.
Impervious surfaces are a ubiquitous urban feature that increase temperature and tree drought stress and are a demonstrated indicator of Acer rubrum L. tree condition and insect pest abundance. We examined the relationship between A. rubrum condition, impervious surface cover, and Melanaspis tenebricosa (Comstock) abundance, a primary herbivore of urban A. rubrum, in eight cities across the southern distribution of A. rubrum. We predicted that the effects of warming, due to impervious surface, would be greater in warmer southern cities than in cooler northern cities. We found that impervious surface was a robust predictor of tree condition, but this effect was not significantly affected by background temperature. Melanaspis tenebricosa abundance was a function of impervious surface and background temperature, with greatest abundances occurring at mid latitudes. Based on these relationships, we developed impervious surface thresholds to inform site selection for A. rubrum throughout the southeastern USA. Planting criteria based on habitat characteristics should maximize urban tree longevity and services provided.
Urban areas, a rapidly expanding land cover type, are composed of a mix of impervious surfaces, ornamental plants, and remnant habitat, which alters abiotic conditions and affects arthropod community assemblages and trophic interactions. Importantly, these effects often reduce arthropod diversity and may increase, reduce, or not change individual species or trophic interactions, which affects human and environmental health. Despite the pace of urbanization, drivers and consequences of change in urban arthropod communities remains poorly understood. Here, we review recent findings that shed light on the effects of urbanization on plants and abiotic conditions that drive arthropod community composition and trophic interactions, with discussion of how these effects conflict with human values and can be mitigated for future urbanization.
Urban forests provide important ecosystem services to city residents, including pollution removal and carbon storage. Climate change and urbanization pose multiple threats to these services. However, how these threats combine to affect urban trees, and thus how to mitigate their effects, remains largely untested because multi‐factorial experiments on mature trees are impractical.
We used a unique urban warming experiment paired with a laboratory chamber experiment to determine how three of the most potentially damaging factors associated with global change for urban and rural trees—warming, drought, and insect herbivory—affect growth of Quercus phellos (willow oak), the most commonly planted large shade tree in the southeastern US , which is known for its resilience to these potential stressors.
In a previous study, we found that the urban heat island effect was associated with reduced growth of Q. phellos and higher abundance of Parthenolecanium scale insects, key pests of oaks in cities. Here, we tested the hypothesis that tree water stress is the mechanism for these effects of warming. We found evidence that water stress is a major, interactive factor reducing urban tree growth, but found no evidence that water stress is associated with Parthenolecanium survival or abundance. Warming and Parthenolecanium only reduced growth in Q. phellos saplings that were simultaneously water stressed.
Synthesis and applications . Across many temperate cities worldwide, urban trees grow less than rural trees. Our results point to water stress as the most likely driver for this pattern. Importantly, we found that water stress both reduces tree growth on its own and exacerbates effects of warming and insect pests on tree growth. Therefore, management strategies targeted at increasing tree hydration in cities may reduce effects of these three key stressors that are expected to intensify with further urbanization and climate change.
The longstanding biotic interactions hypothesis predicts that herbivore pressure declines with latitude, but the evidence is mixed. To address gaps in previous studies, we measured herbivory and defence in the same system, quantified defence with bioassays, and considered effects of leaf age. We quantified herbivory and defence of young and mature leaves along a continental gradient in eastern North America in the native herb Phytolacca americana L. Herbivory in the field declined with latitude and was strongly correlated with lepidopteran abundance. Laboratory bioassays revealed that leaf palatability was positively correlated with latitude of origin. Young leaves were more damaged than mature leaves at lower latitudes in the field, but less palatable in bioassays. Both defence and palatability displayed non-linear latitudinal patterns, suggesting potential mechanisms based on biological or climatic thresholds. In sum, observational and experimental studies find patterns consistent with high herbivore pressure and stronger plant defences at lower latitudes.
Latitudinal patterns in herbivory are widely debated. A recent publication (Zhang et al ., Journal of Ecology , 104 , 2016 : 1089–1095) concluded that the absolute levels of herbivory (hypothesis 1), as well as latitudinal and climatic gradients in herbivory (hypothesis 2), differ between the hemispheres.
Zhang et al . (2016), among others, used a measure of plant damage from Moles & Westoby ( Oikos , 90 , 2000: 517–524) that greatly overestimates insect herbivory, and they did not include the main effect of hemisphere in their linear model that explored the relationship between herbivory and latitude. After correction of the values extracted from Moles & Westoby (2000), none of several statistical models that tested both hypotheses simultaneously (i.e. including hemisphere, latitude/climate and their interaction) confirmed the existence of statistically significant differences in the patterns of insect herbivory between the hemispheres.
Synthesis . The current level of knowledge does not provide grounds to conclude that hemispheric asymmetries exist either in the average levels of insect herbivory or in the relationships between herbivory and latitude or climate.
Kozlov & Klemola ( Journal of Ecology , 105 , 2017: 000‐000) argued that our conclusion about hemispheric asymmetries in herbivory is not supported after the replacement of data points extracted from Moles & Westoby ( Oikos , 90 , 2000: 517–524.).
To re‐evaluate the asymmetries in herbivory, we analysed the revised dataset according to Kozlov & Klemola's (2017) suggestion, as well as the dataset used in Kozlov et al . ( Global Ecology and Biogeography , 24 , 2015: 1126–1135.).
All the analyses showed that herbivory decreased with latitude only in the Northern hemisphere, but in the Southern hemisphere, herbivory showed no relation to latitude. Model selection showed that in both datasets, hemisphere is a key determinant of herbivory, with Southern hemisphere having higher levels of herbivory than the Northern hemisphere.
Synthesis . Based on currently available data collected from publications, we found evidence for hemispheric asymmetries in herbivory. This conclusion is unlikely to change even when the controversial data points are replaced.
Synopsis:
Few studies have quantified the relative importance of direct effects of climate change on communities versus indirect effects that are mediated thorough species interactions, and the limited evidence is conflicting. Trait-based approaches have been popular in studies of climate change, but can they be used to estimate direct versus indirect effects? At the species level, thermal tolerance is a trait that is often used to predict winners and losers under scenarios of climate change. But thermal tolerance might also inform when species interactions are likely to be important because only subsets of species will be able to exploit the available warmer climatic niche space, and competition may intensify in the remaining, compressed cooler climatic niche space. Here, we explore the relative roles of the direct effects of temperature change and indirect effects of species interactions on forest ant communities that were heated as part of a large-scale climate manipulation at high- and low-latitude sites in eastern North America. Overall, we found mixed support for the importance of negative species interactions (competition), but found that the magnitude of these interaction effects was predictable based on the heat tolerance of the focal species. Forager abundance and nest site occupancy of heat-intolerant species were more often influenced by negative interactions with other species than by direct effects of temperature. Our findings suggest that measures of species-specific heat tolerance may roughly predict when species interactions will influence responses to global climate change.
Despite the increasing rate of urbanisation, the consequences of this process on biotic interactions remain insufficiently studied. Our aims were to identify the general pattern of urbanisation impact on background insect herbivory, to explore variations in this impact related to characteristics of both urban areas and insect-plant systems, and to uncover the factors governing urbanisation impacts on insect herbivory. We compared the foliar damage inflicted on the most common trees by defoliating, leafmining and gall-forming insects in rural and urban habitats associated with 16 European cities. In two of these cities we explored quality of birch foliage for herbivorous insects, mortality of leafmining insects due to predators and parasitoids and bird predation on artificial plasticine larvae. On average, the foliage losses to insects were 16.5% lower in urban than in rural habitats. The magnitude of the overall adverse effect of urbanisation on herbivory was independent of the latitude of the locality and was similar in all 11 studied tree species, but increased with an increase in the size of the urban area: it was significant in large cities (city population 1–5 million) but not significant in medium-sized and small towns. Quality of birch foliage for herbivorous insects was slightly higher in urban habitats than in rural habitats. At the same time, leafminer mortality due to ants and birds and the bird attack intensity on dummy larvae were higher in large cities than in rural habitats, which at least partially explained the decline in insect herbivory observed in response to urbanisation. Our findings underscore the importance of top-down forces in mediating impacts of urbanisation on plant-feeding insects: factors favouring predators may override the positive effects of temperature elevation on insects and thus reduce plant damage. This article is protected by copyright. All rights reserved.
Biological effects of climate change are expected to vary geographically, with a strong signature of latitude. For ectothermic animals, there is systematic latitudinal variation in the relationship between climate and thermal performance curves, which describe the relationship between temperature and an organism's fitness. Here we ask whether these documented latitudinal patterns can be generalized to predict arthropod responses to warming across mid and high temperate latitudes, for taxa whose thermal physiology has not been measured. To address this question, we used a novel natural experiment consisting of a series of urban warming gradients at different latitudes. Specifically, we sampled arthropods from a single common street tree species across temperature gradients in 4 US cities, located from 35.8 to 42.4° latitude. We captured 6746 arthropods in 34 families from 111 sites that varied in summer average temperature by 1.7 to 3.4 °C within each city. Arthropod responses to warming within each city were characterized as Poisson regression coefficients describing change in abundance per °C for each family. Family responses in the two mid latitude cities were heterogeneous, including significantly negative and positive effects, while those in high latitude cities varied no more than expected by chance within each city. We expected high-latitude taxa to increase in abundance with warming, and they did so in 1 of the 2 high latitude cities; in the other, Queens (New York City), most taxa declined with warming, perhaps due to habitat loss that was correlated with warming in this city. With the exception of Queens, patterns of family responses to warming were consistent with predictions based on known latitudinal patterns in arthropod physiology relative to regional climate. Heterogeneous responses in mid latitudes may be ecologically disruptive if interacting taxa respond oppositely to warming. This article is protected by copyright. All rights reserved.
Both herbivory and plant defenses against herbivores have been predicted to increase toward tropical regions. Early tests of this latitudinal herbivory-defense hypothesis (LHDH) were supportive, but accumulating evidence has been mixed. We argue that the lack of clarity might be due to heterogeneity in methodology and problems with study design and interpretation. We suggest possible solutions. Latitudinal studies need to carefully consider spatial and phylogenetic scale, to link plant defense measurements to herbivore performance, and to incorporate additional concepts from plant defense theory such as tolerance and induced defense. In addition, we call for consistent measures of herbivory to standardize comparisons across biomes. Improving methodology in future studies of LHDH should resolve much of the current controversy.
Urbanization is one of the major environmental challenges facing the world today. One of its particularly pressing effects is alterations to local and regional climate through, for example, the Urban Heat Island. Such changes in conditions are likely to have an impact on the phenology of urban vegetation, which will have knock-on implications for the role that urban green infrastructure can play in delivering multiple ecosystem services. Here, in a human-dominated region, we undertake an explicit comparison of vegetation phenology between urban and rural zones. Using satellite-derived MODIS-EVI data from the first decade of the 20th century, we extract metrics of vegetation phenology (date of start of growing season, date of end of growing season, and length of season) for Britain's 15 largest cities and their rural surrounds. On average, urban areas experienced a growing season 8.8 days longer than surrounding rural zones. As would be expected, there was a significant decline in growing season length with latitude (by 3.4 and 2.4 days/degree latitude in rural and urban areas respectively). Although there is considerable variability in how phenology in urban and rural areas differs across our study cities, we found no evidence that built urban form influences the start, end, or length of the growing season. However, the difference in the length of the growing season between rural and urban areas was significantly negatively associated with the mean disposable household income for a city. Vegetation in urban areas deliver many ecosystem services such as temperature mitigation, pollution removal, carbon uptake and storage, the provision of amenity value for humans and habitat for biodiversity. Given the rapid pace of urbanization and ongoing climate change, understanding how vegetation phenology will alter in the future is important if we wish to be able to manage urban greenspaces effectively.
Plant–herbivore interactions occur in all ecosystems and provide a major avenue for energy flow to higher trophic levels. A long-standing hypothesis to explain the latitudinal gradient in species diversity proposes that the relatively stable and frost-free climate of the tropics should lead to more intense biotic interactions in tropical compared with temperate environments, giving rise to a greater diversity of plants and herbivores. Herbivory rates have been compared across latitudes to test this biotic interactions hypothesis, with herbivory typically being measured from observable leaf damage. However, we argue that a measure of percentage leaf damage alone does not straightforwardly reflect the cost of herbivory to the plant, and on its own does not constitute an appropriate test of the biotic interactions hypothesis. For a given amount of herbivory, the impact of herbivory is dependent upon many factors, such as the construction cost of the leaf, the growth and replacement rates and leaf life span. We investigate the latitudinal gradient in herbivory by analysing a large dataset of herbivory rates for 452 tree species and separating the species into those with short and long leaf life spans. We show that annual herbivory rates tend to be greater at lower latitudes for evergreen species (which have long-lived leaves), but no trend in herbivory rate with latitude was found for species with short leaf life spans. Phylogenetic least squares regression assuming Ornstein-Uhlenbeck processes also showed a negative effect of latitude on herbivory rate for evergreen trees, but we caution that viewing herbivory as a species trait is problematic. An integrative approach that incorporates leaf life span, as well as the costs of investment in growth and potential costs of losing leaf tissue, is needed to further our understanding of the ecological and evolutionary dynamics of herbivory.
This 'super-generalist' has low resource requirements and does many things reasonably well in a wide variety of ecological conditions.
Although a number of investigations have concluded that lower latitudes are associated with increases in herbivore abundance and plant damage, the generality of this pattern is still under debate. Multiple factors may explain the lack of consistency in latitude–herbivory relationships. For instance, latitudinal variation in herbivore pressure may be shaped entirely or not by climatic variables, or vary among herbivore guilds with differing life-history traits. Additionally, the strength of top–down effects from natural enemies on herbivores might also vary geographically and influence latitude–herbivory patterns. We carried out a field study where we investigated the effects of latitude and climate on herbivory by a seed-eating caterpillar and leaf chewers, as well as parasitism associated to the former across 30 populations of the perennial herb Ruellia nudiflora (Acanthaceae). These populations were distributed along a 5° latitudinal gradient from northern Yucatan (Mexico) to southern Belize, representing one-third of the species' latitudinal distribution and the entirety and one-third of the precipitation and temperature gradient of this species' distribution (respectively). We found opposing latitudinal gradients of seed herbivory and leaf herbivory, and this difference appeared to be mediated by contrasting effects of climate on each guild. Specifically, univariate regressions showed that seed herbivory increased at higher latitudes and with colder temperatures, while leaf herbivory increased toward the equator and with wetter conditions. Multiple regressions including temperature, precipitation and latitude only found significant effects of temperature for seed herbivory and latitude for leaf herbivory. Accordingly, that latitudinal variation in seed herbivory appears to be driven predominantly by variation in temperature whereas latitudinal variation in leaf herbivory was apparently driven by other unexplored correlates of latitude. Parasitism did not exhibit variation with latitude or climatic factors. Overall, these findings underscore that the factors driving latitudinal clines in herbivory might vary even among herbivore species coexisting on the same host plant.
Street trees are an integral element of urban life. They provide a vast range of benefits in residential and commercial precincts, and they support healthy communities by providing environmental, economic and social benefits. However, increasing areas of impermeable surface can increase the stresses placed upon urban ecosystems and urban forests. These stresses often lead tree roots to proliferate in sites that provide more-favourable conditions for growth, but where they cause infrastructure damage and pavement uplift. This damage is costly and a variety of preventative measures has been tested to sustain tree health and reduce pavement damage. This review explores a wide range of literature spanning 30 years that demonstrates the benefits provided by street trees, the perceptions of street trees conveyed by urban residents, the costs of pavement damage by tree roots, and some tried and tested measures for preventing pavement damage and improving tree growth.
Trees provide ecosystem services that counter negative effects of urban habitats on human and environmental health. Unfortunately, herbivorous arthropod pests are often more abundant on urban than rural trees, reducing tree growth, survival, and ecosystem services. Previous research where vegetation complexity was reduced has attributed elevated urban pest abundance to decreased regulation by natural enemies. However, reducing vegetation complexity, particularly the density of overstory trees, also makes cities hotter than natural habitats. We ask how urban habitat characteristics influence an abiotic factor, temperature, and a biotic factor, natural enemy abundance, in regulating the abundance of an urban forest pest, the gloomy scale, (Melanaspis tenebricosa). We used a map of surface temperature to select red maple trees (Acer rubrum) at warmer and cooler sites in Raleigh, North Carolina, USA. We quantified habitat complexity by measuring impervious surface cover, local vegetation structural complexity, and landscape scale vegetation cover around each tree. Using path analysis, we determined that impervious surface (the most important habitat variable) increased scale insect abundance by increasing tree canopy temperature, rather than by reducing natural enemy abundance or percent parasitism. As a mechanism for this response, we found that increasing temperature significantly increases scale insect fecundity and contributes to greater population increase. Specifically, adult female M. tenebricosa egg sets increased by approximately 14 eggs for every 1°C increase in temperature. Climate change models predict that the global climate will increase by 2–3°C in the next 50–100 years, which we found would increase scale insect abundance by three orders of magnitude. This result supports predictions that urban and natural forests will face greater herbivory in the future, and suggests that a primary cause could be direct, positive effects of warming on herbivore fitness rather than altered trophic interactions.
Cities experience elevated temperature, CO 2 , and nitrogen deposition decades ahead of the global average, such that biological response to urbanization may predict response to future climate change. This hypothesis remains untested due to a lack of complementary urban and long-term observations. Here, we examine the response of an herbivore, the scale insect Melanaspis tenebricosa, to temperature in the context of an urban heat island, a series of historical temperature fluctuations, and recent climate warming. We survey M. tenebricosa on 55 urban street trees in Raleigh, NC, 342 herbarium specimens collected in the rural southeastern United States from 1895 to 2011, and at 20 rural forest sites represented by both modern (2013) and historical samples. We relate scale insect abundance to August temperatures and find that M. tenebricosa is most common in the hottest parts of the city, on historical specimens collected during warm time periods, and in present-day rural forests compared to the same sites when they were cooler. Scale insects reached their highest densities in the city, but abundance peaked at similar temper-atures in urban and historical datasets and tracked temperature on a decadal scale. Although urban habitats are highly modified, species response to a key abiotic factor, temperature, was consistent across urban and rural-forest ecosystems. Cities may be an appropriate but underused system for developing and testing hypotheses about biological effects of climate change. Future work should test the applicability of this model to other groups of organisms.
Knowledge of the latitudinal patterns in biotic interactions, and especially in herbivory, is crucial for understanding the mechanisms that govern ecosystem functioning and for predicting their responses to climate change. We used sap-feeding insects as a model group to test the hypotheses that the strength of plant–herbivore interactions in boreal forests decreases with latitude and that this latitudinal pattern is driven primarily by midsummer temperatures. We used a replicated sampling design and quantitatively collected and identified all sap-feeding insects from four species of forest trees along five latitudinal gradients (750–1300 km in length, ten sites in each gradient) in northern Europe (59 to 70°N and 10 to 60°E) during 2008–2011. Similar decreases in diversity of sap-feeding insects with latitude were observed in all gradients during all study years. The sap-feeder load (i.e. insect biomass per unit of foliar biomass) decreased with latitude in typical summers, but increased in an exceptionally hot summer and was independent of latitude during a warm summer. Analysis of combined data from all sites and years revealed dome-shaped relationships between the loads of sap-feeders and midsummer temperatures, peaking at 17 °C in Picea abies, at 19.5 °C in Pinus sylvestris and Betula pubescens and at 22 °C in B. pendula. From these relationships, we predict that the losses of forest trees to sap-feeders will increase by 0–45% of the current level in southern boreal forests and by 65–210% in subarctic forests with a 1 °C increase in summer temperatures. The observed relationships between temperatures and the loads of sap-feeders differ between the coniferous and deciduous tree species. We conclude that climate warming will not only increase plant losses to sap-feeding insects, especially in subarctic forests, but can also alter plant-plant interactions, thereby affecting both the productivity and the structure of future forest ecosystems.
It is well known that trees can reduce the urban heat island and adapt our cities to climate change throughevapotranspiration. However, the effects of urbanization and anticipated climate change in the soil–rootrhizosphere have not been widely investigated. The current study studied the growth and physiology ofthe urban tree Pyrus calleryana grown in a factorial experiment with or without urbanization and simu-lated climate change between April 2010 and December 2012 in the Botanical Grounds of the Universityof Manchester, UK. The study indicated that urbanization and simulated climate change had small butcontrasting effects on tree growth and morphology. Urbanization increased tree growth by 20–30%, butdid not affect leaf area index (LAI) and showed reduced peak water loss and hence evapotranspirationalcooling. Although soil moisture content in the upper 20 cm was higher in the urbanized plots, urban-ization showed reduced sap flux density, reduced chlorophyll a:b and delayed recovery of chlorophyllfluorescence (Fv:Fm) throughout the experimental period. In contrast, simulated climate change had noeffect on growth but increased LAI by 10%. Despite being more water stressed, trees grown in simulatedclimate change plots lost more water both according to porometry and sap flow measurements. Simu-lated climate change increased peak energy and water loss by around 13%, with trees having an averagesap flux density of around 170 g cm−2d−1, 40% higher than trees grown in control plots. Our study sug-gested that transpirational cooling benefit might be enhanced with a longer growth season and higher soiltemperature in places such as Manchester, UK in future, but potentially at the expense of photosynthesisand carbon gain.
We used leaf gas exchange, sap flow, and eddy covariance measurements to investigate whether high temperature substantially limits CO2 uptake at the LBA-ECO (Large-scale Biosphere-Atmosphere) km-83 tropical forest site in Brazil. Leaf-level temperature-photosynthesis curves, and comparisons of whole-canopy net ecosystem CO2 exchange (NEE) with air temperature, showed that CO2 uptake declined sharply during warm periods. Observations of ambient leaf microclimate showed that leaves oscillate between two states: a cool, dimly lit stage and a hot, brightly illuminated stage where leaf temperatures are often greater than 35°C. The leaf-level rates of photosynthesis decreased when shaded leaves (~ambient air temperature and < 500 mumol m-2 s-1) were transferred into a prewarmed, brightly illuminated chamber (35° to 38°C and 1000 mumol m-2 s-1), coincident with increased leaf temperature, increased evaporative demand, and stomatal closure. The rates of whole-canopy CO2 uptake calculated at 5-min intervals increased initially at the onset of sunny periods that followed extended cloudy periods, but then decreased as the sunlight continued, leaf temperature and evaporative demand increased, and canopy conductance decreased. The forest at km-83 appears to be close to a high temperature threshold, above which CO2 uptake drops sharply. This sensitivity results in part from the covariance between leaf temperature and leaf illumination; the brightly illuminated leaves that contribute disproportionately to canopy photosynthesis are warmed to the point that leaf gas exchange is curtailed.
The incidence of alkaloid-bearing plants is dependent upon their habit and ecogeographical distribution. Among annual species the incidence of alkaloids is nearly twice that of perennials, among tropical floras nearly twice that of temperate floras, and a latitudinal cline is evident. In New Guinea, disparate communities differ in the incidence of alkaloid-bearing species and in the amount of alkaloid contained in their vegetative tissues. Families primarily distributed in the tropics have a higher percentage of alkaloid-bearing species than do those of temperate regions or those with cosmopolitan distributions. The most primitive orders, Magnoliales and Ranales, have a higher percentage of alkaloid-bearing species than the remainder of the dicots. These findings are discussed from the vantage point of coevolutionary theory. It is suggested that the ecogeographic patterns may be the result of differences in pest pressure, the alkaloids playing a defensive role in plants.
A primary focus of wildlife ecology is studying how the arrangement, quality, and distribution of habitat influence wildlife populations at multiple spatial scales. A practical limitation of conducting wildlife–habitat investigations in the field, however, is that sampling points tend to be close to one another, resulting in spatial clustering. Consequently, when ecologists seek to quantify the effects of environmental predictors surrounding their sampling points, they encounter the issue of using landscapes that are partially or completely overlapping. A presumed problem of overlapping landscapes is that data generated from these landscapes, when used as predictors in statistical modeling, might violate the assumption of independence. However, the independence of error is the critical assumption, not the independence of predictor variables. Nonetheless, many researchers strive to avoid such overlaps through sampling design or alternative analytical procedures and specialized software programs have been created to assist with this. We present theoretical arguments and empirical evidence showing that changing the amount of overlap does not alter the degree of spatial autocorrelation. Using data derived from 2 broad-scaled avian monitoring programs, we quantified the relationship between forest cover and bird abundance and occurrence at multiple landscapes ranging from 100 m to 24 km across. We found no clear evidence that increasing overlap of landscapes increased spatial autocorrelation in model residuals. Our results demonstrate that the concern of overlapping landscapes as a potential cause of violation of spatial independency among sampling units is misdirected and represents an oversimplification of the statistical and ecological issues surrounding spatial autocorrelation. Overlapping landscapes and spatial autocorrelation are separate issues in the modeling of wildlife populations and their habitats; non-overlapping landscapes do not ensure spatial independency and overlapping landscapes do not necessarily lead to greater spatial autocorrelation in model errors. © 2011 The Wildlife Society.