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Global climate change is causing more frequent and severe droughts, which could have serious repercussions for the maintenance of biodiversity. Here, we compare native bee assemblages collected via bowl traps before and after a severe drought event in 2014 in San Diego, California, and examine the relative magnitude of impacts from drought in fragmented habitat patches versus unfragmented natural reserves. Bee richness and diversity were higher in assemblages surveyed before the drought compared to those surveyed after the drought. However, bees belonging to the Lasioglossum subgenus Dialictus increased in abundance after the drought, driving increased representation by small-bodied, primitively eusocial, and generalist bees in post-drought assemblages. Conversely, among non-Dialictus bees, post-drought years were associated with decreased abundance and reduced representation by eusocial species. Drought effects were consistently greater in reserves, which supported more bee species, than in fragments, suggesting that fragmentation either had redundant impacts with drought, or ameliorated effects of drought by enhancing bees’ access to floral resources in irrigated urban environments. Shifts in assemblage composition associated with drought were three times greater compared to those associated with habitat fragmentation, highlighting the importance of understanding the impacts of large-scale climatic events relative to those associated with land use change.

Ecological impacts associated with ant introductions have received considerable attention, but most studies that report on these impacts contrast species assemblages between invaded and uninvaded sites. Given the low inferential power of this type of space-for-time comparison, alternative approaches are needed to evaluate claims that ant invasions drive native species loss. Here, we use long-term data sets from two different Argentine ant eradication programs on the California Channel Islands to examine how the richness and composition of native ant assemblages change before and after invasion (but prior to the initiation of treatments). At four different sites on two different islands, pre-invasion native ant assemblages closely resembled those at uninvaded (control) sites in terms of species richness, species composition, and the presence of multiple indicator species. Invader arrival coincided with large (> 75%) and rapid (within 1 year) declines in species richness, shifts in species composition, and the loss of indicator species. These impacts will hopefully be reversed by the recolonization of formerly invaded areas by native ant species following Argentine ant treatment, and long-term studies of native ant recovery at these sites are ongoing. Unchecked spread of the Argentine ant on other islands in this archipelago, however, poses a grave threat to native ants, which include a number of endemic taxa.

Most plant-pollinator mutualisms are generalized. As such, they are susceptible to perturbation by abundant, generalist, non-native pollinators such as the western honey bee (Apis mellifera), which can reach high abundances and visit flowers of many plant species in their expansive introduced range. Despite the prevalence of non-native honey bees, their effects on pollination mutualisms in natural ecosystems remain incompletely understood. Here we contrast community-level patterns of floral visitation by honey bees with that of the diverse native pollinator fauna of southern California, USA. We show that the number of honey bees visiting plant species increases much more rapidly with flower abundance than does that of non-honey bee insects, such that the percentage of all visitors represented by honey bees increases with flower abundance. Thus, honey bees could disproportionately impact the most abundantly blooming plant species and the large numbers of both specialised and generalised pollinator species that they sustain. Honey bees may preferentially exploit high-abundance floral resources because of their ability to recruit nestmates; these foraging patterns may cause native insect species to forage on lower-abundance resources to avoid competition. Our results illustrate the importance of understanding foraging patterns of introduced pollinators in order to reveal their ecological impacts.

The western honey bee (Apis mellifera) is the most frequent floral visitor of crops worldwide, but quantitative knowledge of its role as a pollinator outside of managed habitats is largely lacking. Here we use a global data set of 80 published plant-pollinator interaction networks as well as pollinator effectiveness measures from 34 plant species to assess the importance of A. mellifera in natural habitats. Apis mellifera is the most frequent floral visitor in natural habitats worldwide, averaging 13% of floral visits across all networks (range 0 – 85%), with 5% of plant species recorded as being exclusively visited by A. mellifera. For 33% of the networks and 49% of plant species, however, A. mellifera visitation was never observed, illustrating that many flowering plant taxa and assemblages remain dependent on non-A. mellifera visitors for pollination. Apis mellifera visitation was higher in warmer, less variable climates and on mainland rather than island sites, but did not differ between its native and introduced ranges. With respect to single-visit pollination effectiveness, A. mellifera did not differ from the average non-A. mellifera floral visitor, though it was generally less effective than the most effective non-A. mellifera visitor. Our results argue for a deeper understanding of how A. mellifera, and potential future changes in its range and abundance, shape the ecology, evolution, and conservation of plants, pollinators, and their interactions in natural habitats.

Island biotas provide unparalleled opportunities to examine evolutionary processes. Founder effects and bottlenecks, e.g. typically decrease genetic diversity in island populations, while selection for reduced dispersal can increase population structure. Given that support for these generalities mostly comes from single-species analyses, assemblage-level comparisons are needed to clarify how (i) colonization affects the gene pools of interacting insular organisms, and (ii) patterns of genetic differentiation vary within assemblages of organisms. Here, we use genome-wide sequence data from ultraconserved elements (UCEs) to compare the genetic diversity and population structure of mainland and island populations of nine ant species in coastal southern California. As expected, island populations (from Santa Cruz Island) had lower expected heterozygosity and Watterson's theta compared to mainland populations (from the Lompoc Valley). Island populations, however, exhibited smaller genetic distances among samples, indicating less population subdivision. Within the focal assemblage, pairwise F st values revealed pronounced interspecific variation in mainland-island differentiation, which increases with gyne body size. Our results reveal population differences across an assemblage of interacting species and illuminate general patterns of insularization in ants. Compared to single-species studies, our analysis of nine conspecific population pairs from the same island-mainland system offers a powerful approach to studying fundamental evolutionary processes. [ABSTRACT FROM AUTHOR]

1. Protection mutualisms between ants and honeydew-producing insects influence arthropod food webs by altering the behaviour, activity and local abundance of ants on plants. Ants often tend multiple species of honeydew-producing insects; however, studies that examine such effects typically consider only pairwisemutualisms. This study investigates how multi-species mutualisms between ants and honeydew-producing insects structure arthropod food webs in citrus.2. In an organic lemon orchard in San Diego County, California, U.S.A., ants or honeydew-producing insects (or neither) were experimentally removed from individual, mature lemon trees and then abundances of the following were estimated over a 2-year period: the Argentine ant, five species of honeydew-producing hemipterans, and California red scale. Red scale produces no honeydew but indirectly benefits from the presence of ants, which disrupt parasitism by Aphytis wasps.3. Mutualism between ants and honeydew-producing hemipterans indirectly andpositively affected red scale. Levels of parasitism decreased with increasing antrecruitment, and red scale density increased with both increasing ant recruitment and increasing honeydew-producing hemipteran abundance. Moreover, abundances of each of three honeydew-producing hemipteran species emerged as positive predictors of redscale density in different analyses; this finding suggests that individual hemipteran taxa exhibit distinct and spatially localised effects on ants.4. Evidence of positive, indirect effects between two focal species of honeydew-producing Hemiptera was also detected. Guilds of honeydew-producing hemipterans may provide ants with honeydew more consistently or for longer than that produced by any single species. These results argue for an appreciation of how mutualist diversity affects the ecological consequences of mutualisms.

1. In ant–hemipteran mutualisms, ants receive carbohydrates in the form of honeydew, while hemipterans receive protection from natural enemies. In the absence of natural enemies, however, the direct effects of tending are generally less well known. We hypothesised that with increasing tending intensity (ant to aphid ratio), aphid performance would increase initially, then decrease at high tending levels due to the metabolic cost of producing high quality honeydew.2. We tested our hypothesis in a greenhouse experiment by manipulating Argentine ant (Linepithema humile Mayr) colony size while holding constant the initial size of aphid (Chaitophorus populicola Thomas) aggregations. The two parameters associated with survival, aphid survivorship to maturity and longevity, declined with increasing tending intensity, whereas per capita birth rate and time to first reproduction showed no relationship to attendance. The intrinsic rate of increase declined only at relatively high tending levels, suggesting a nonlinearity in the effect of tending intensity.3. Tending intensity measured in the experiment was similar to that observed in free-living aggregations of C. populicola. Furthermore, the per capita recruitment rate of ants to free-living aphid aggregations was negatively density-dependent, indicating that small aggregations tend to experience the highest levels of tending intensity. This finding suggests that the aphid's intrinsic rate of increase may be positively density-dependent, mediated by the aphid's mutualistic interaction with the ant.4. In the Argentine ant–C. populicola interaction, experimental manipulation of colony size revealed a direct cost of ant attendance that was conditional upon tending intensity. Experiments that manipulate only ant presence or absence may yield an incomplete understanding of the mutualistic interaction if underlying nonlinearities exist.

Studies of food webs often employ stable isotopic approaches to infer trophic position and interaction strength without consideration of spatio-temporal variation in resource assimilation by constituent species. Using results from laboratory diet manipulations and monthly sampling of field populations, we illustrate how nitrogen isotopes may be used to quantify spatio-temporal variation in resource assimilation in ants. First, we determined nitrogen enrichment using a controlled laboratory experiment with the invasive Argentine ant (Linepithema humile). After 12 weeks, worker δ15N values from colonies fed an animal-based diet had δ15N values that were 5.51% greater compared to colonies fed a plant-based diet. The shift in δ15N values in response to the experimental diet occurred within 10 weeks. We next reared Argentine ant colonies with or without access to honeydew-producing aphids and found that after 8 weeks workers from colonies without access to aphids had δ15N values that were 6.31% larger compared to colonies with access to honeydew. Second, we sampled field populations over a 1-year period to quantify spatio-temporal variability in isotopic ratios of L. humile and those of a common native ant (Solenopsis xyloni). Samples from free-living colonies revealed that fluctuations in δ15N were 1.6–2.4‰ for L. humile and 1.8–2.9‰ for S. xyloni. Variation was also detected among L. humile castes: time averaged means of δ15N varied from 1.2 to 2.5‰ depending on the site, with δ15N values for queens ≥ workers > brood. The estimated trophic positions of L. humile and S. xyloni were similar within a site; however, trophic position for each species differed significantly at larger spatial scales. While stable isotopes are clearly useful for examining the trophic ecology of arthropod communities, our results suggest that caution is warranted when making ecological interpretations when stable isotope collections come from single time periods or life stages.

Plant invasions have tremendous potential to alter food webs by changing basal resources. Recent studies document how plant invasions may contribute to increased arthropod abundances in detritus-based food webs. An obvious mechanism for this phenomenon—a bottom-up effect resulting from elevated levels of detritus from the invasive plant litter—has not been explicitly studied. We examined the effects of an annual grass invasion on ground arthropod assemblages in the coastal sage scrub (CSS) of southern California. Bottom-up food web theory predicts that the addition of detritus would increase generalist-feeding arthropods at all trophic levels; accordingly, we expected increases in fungi, Collembola, and common predators such as mites and spiders. For the common ant taxa, habitat alteration may also be important for predicting responses. Thus we expected that Forelius mccooki and Pheidole vistana, the most common ant species, would decline because of changes in soil temperature (F. mccooki) and habitat structure (P. vistana) associated with litter. We studied trends observationally and conducted a 3-year experiment in which we manipulated litter quantity. In contrast to other published studies, most detritus-based arthropod taxa declined in areas of high grass invasion, and, within trophic levels, responses often varied idiosyncratically. For the two most common taxa, a native ant (F. mccooki), and predatory mites in the Anystidae, we experimentally linked declines in abundance to increased levels of invasive grass litter. Such declines, especially those exhibited by the most common ant taxa, could have cascading effects on the CSS ecosystem, where ants are numerically dominant and thus may have broad influences on food web and ecosystem properties. Our results highlight that accurately predicting arthropod responses to invasive plant litter requires careful consideration of the structural and food resources provided by detritus to each particular food web.

Climate change alters environmental conditions in ways that directly and indirectly affect plants. Flowering plants, for example, modify reproductive allocation in response to heat and drought stress, and such changes can in turn affect pollinator visitation and, ultimately, plant reproduction. Although the individual effects of warming and drought on plant reproductive allocation are well known, these factors may interact to influence reproduction. Here, we conducted a fully crossed temperature by irrigation manipulation in squash (Cucurbita pepo) to test how temperature and soil moisture variation affect pollinator‐mediated reproduction. To tease apart the direct and indirect effects of temperature and soil moisture, we compared hand‐pollinated plants to bee‐pollinated plants and restricted bee foraging (i.e., pollen transfer) to one experimental group per day. Temperature and soil‐moisture limitation acted independently of one another: warming decreased flower size and increased pollen production, whereas the effects of soil‐moisture limitation were uniformly inhibitory. While treatments did not change squash bee (Xenoglossa spp.) behavior, floral visitation by the honey bee (Apis mellifera) increased with temperature in male flowers and decreased with soil moisture in female flowers. Pollen deposition by bees was independent of plant soil moisture, yet reducing soil moisture increased pollen limitation. This result stemmed at least in part from the effects of soil‐moisture limitation on pollen viability; seed set declined with increasing deposition of fluorescent pigment (a proxy for pollen) from plants experiencing decreased soil moisture. These findings suggest that the transfer of lower‐quality pollen from plants experiencing soil‐moisture limitation led to drought‐induced pollen limitation. Similar effects may occur in a wide variety of flowering plant species as climate warming and drought increasingly impact animal‐pollinated systems. [ABSTRACT FROM AUTHOR]

The San Bernardino flying squirrel (SBFS) is an isolated subspecies of Humboldt's flying squirrel, occurring in montane sky islands in the San Bernardino and San Jacinto Mountains in Southern California, USA. Recent small mammal surveys in the San Jacinto Mountains suggest the squirrel is extirpated. Our objectives were (1) determine habitat features, including forest metrics and topographical factors, that influence SBFS presence, in the San Bernardino Mountains; (2) use information collected in the San Bernardino Mountains to confirm squirrel occurrence and habitat preference in the San Jacinto Mountains; and (3) assess habitat and climatic differences between the two mountain ranges that could explain species persistence in one mountain range but not the other. We surveyed for SBFS using camera traps at 54 sites in the San Bernardino Mountains and 34 sites in the San Jacinto Mountains using both camera traps and acoustics. In the San Bernardino Mountains, we detected squirrels in sites that were more mesic, had higher structural heterogeneity, and had greater amounts of downed woody material compared to non‐detection sites. Habitat parameters were similar between the two ranges; however, squirrels were not detected in the San Jacinto Mountains. Conditions in the San Jacinto Mountains were hotter and drier. Increased temperatures due to climate change could potentially explain the absence of flying squirrels in the San Jacinto Mountains. [ABSTRACT FROM AUTHOR]

Cold narcosis is commonly used for immobilizing bees and has been found to have no effect on mortality and fecundity in honeybees. We present the first study examining the effect of cold narcosis on recruitment and foraging in bumblebees. In a controlled laboratory setting, we observed that the number of foraging Bombus occidentalis, a New World bumblebee, increased after the focal forager returned to the nest with rich 2.5 M sucrose solution. However, cold narcosis (˜4° C, ˜5 min) significantly reduced the ability of B. occidentalis foragers to incite nest mates to forage in a flight arena: 22% fewer total foragers and 56% fewer new recruits were present in the flight arena 10 min after the return of a cold-treated bee compared to the numbers present after the return of an unmanipulated bee. These results demonstrate unanticipated effects of cold anaesthesia on bumblebee foraging behaviour, suggesting that the use of cold anaesthesia should be treated with caution in bumblebee foraging studies. © 2006 International Bee Research Association.

Partitioning the relative contributions of cuticular and respiratory water loss in a tracheate arthropod is relatively easy if it undergoes discontinuous gas exchange cycles or DGCs, leaving its rate of cuticular water loss in primary evidence while its spiracles are closed. Many arthropods are not so obliging and emit CO2 continuously, making cuticular and respiratory water losses difficult or impossible to partition. We report here that by switching ambient air from 21 to 100% O2, marked spiracular constriction takes place, causing a transient but substantial - up to 90% - reduction in CO2 output. A reduction in water loss rate occurs at the same time. Using this approach, we investigated respiratory water loss in Drosophila melanogaster and in two ant species, Forelius mccooki and Pogonomyrmex californicus. Our results - respiratory water loss estimates of 23%, 7.6% and 5.6% of total water loss rates, respectively - are reasonable in light of literature estimates, and suggest that the 'hyperoxic switch' may allow straightforward estimation of respiratory water loss rates in arthropods lacking discontinuous gas exchange. In P. californicus, which we were able to measure with and without a DGC, presence or absence of a DGC did not affect respiratory vs total water loss rates.

Drought is an increasingly important consequence of climate change. Drought often causes plants to alter patterns of resource allocation, which in turn can affect how plants interact with other species. How these altered interactions subsequently influence plant reproductive success remains incompletely understood and may depend on the degree of specialization exhibited by antagonists and mutualists. Specialist pollinators, for example, are dependent on floral resources from their obligate hosts and under drought conditions may thus indiscriminately visit these hosts (at least in certain circumstances). Generalist pollinators, in contrast, may only forage on host plants in good condition, given that they can forage on other plant species. We tested this hypothesis and its consequences for plant reproduction in squash (Cucurbita pepo) grown along an experimental moisture gradient ranging from dry (growth and flowering compromised) to wet conditions. Floral visitation increased with plant soil moisture for generalist honey bees but was independent of plant soil moisture for specialist squash bees. Pollen production increased with plant soil moisture, and fluorescent pigments placed on flowers revealed that pollinators primarily moved pollen from male flowers on well‐watered plants to the stigmas of female flowers on well‐watered plants. Seed set increased with increasing plant soil moisture but, notably, was higher in bee‐pollinated plants compared to plants pollinated by hand with an even mix of pollen from plants grown at either end of the experimental moisture gradient. These results suggest that superior pollen rewards, perhaps combined with selective foraging by generalists, enhanced reproductive success in C. pepo when plant soil moisture was high and more generally illustrate that pollinator behavior may contribute to how drought conditions affect plant reproduction. [ABSTRACT FROM AUTHOR]