Living on the edge: The sensitivity of arthropods to development and climate along an urban-wildland interface in the Sonoran Desert of central Arizona.

Preservation of undeveloped land near urban areas is a common conservation practice. However, ecological processes may still be affected by adjacent anthropogenic activities. Ground-dwelling arthropods are a diverse group of organisms that are critical to ecological processes such as nutrient cycling, which are sensitive to anthropogenic activities. Here, we study arthropod dynamics in a preserve located in a heavily urbanized part of the Sonoran Desert, Arizona, U.S.. We compared arthropod biodiversity and community composition at ten locations, four paired sites representing the urban edge and one pair in the Preserve interior. In total, we captured and identified 25,477 arthropod individuals belonging to 287 lowest practical taxa (LPT) over eight years of sampling. This included 192 LPTs shared between interior and edge sites, with 44 LPTs occurring exclusively in interior sites and 48 LPTs occurring exclusively in edge sites. We found two site pairs had higher arthropod richness on the preserve interior, but results for evenness were mixed among site pairs. Compositionally, the interior and edge sites were more than 40% dissimilar, driven by species turnover. Importantly, we found that some differences were only apparent seasonally; for example edge sites had more fire ants than interior sites only during the summer. We also found that temperature and precipitation were strong predictors of arthropod composition. Our study highlights that climate can interact with urban edge effects on arthropod biodiversity.

Oystershell scale (Hemiptera: Diaspididae) population growth, spread, and phenology on aspen in Arizona, USA.

Oystershell scale (OSS; Lepidosaphes ulmi L.) is an invasive insect that threatens sustainability of aspen (Populus tremuloides Michx.) in the southwestern United States. OSS invasions have created challenges for land managers tasked with maintaining healthy aspen ecosystems for the ecological, economic, and aesthetic benefits they provide. Active management is required to suppress OSS populations and mitigate damage to aspen ecosystems, but before management strategies can be implemented, critical knowledge gaps about OSS biology and ecology must be filled. This study sought to fill these gaps by addressing 3 questions: (i) What is the short-term rate of aspen mortality in OSS-infested stands in northern Arizona, USA? (ii) What are the short-term rates of OSS population growth on trees and OSS spread among trees in aspen stands? (iii) What is the phenology of OSS on aspen and does climate influence phenology? We observed high levels of aspen mortality (annual mortality rate = 10.4%) and found that OSS spread rapidly within stands (annual spread rate = 10-12.3%). We found first, second, and young third instars throughout the year and observed 2 waves of first instars (i.e., crawlers), one throughout the summer and a second in mid-winter. The first wave appeared to be driven by warming seasonal temperatures, but the cause of the second wave is unknown and might represent a second generation. We provide recommendations for future OSS research, including suggestions for more precise quantification of OSS phenology, and discuss how our results can inform management of OSS and invaded aspen ecosystems.

Oviposition behavior of the quasi-gregarious parasitoid, Ooencyrtus kuvanae (Hymenoptera: Encyrtidae).

The parasitoid wasp, Ooencyrtus kuvanae (Howard) (Hymenoptera: Encyrtidae), is a natural enemy of the spongy moth, a significant forest pest in North America. We investigated the oviposition behavior of O. kuvanae females on spongy moth egg masses by (i) presenting female parasitoids with a single spongy moth egg mass that was replaced every day, 2nd day, 4th day, 8th day, or 16th day (which is the total length of the oviposition period) and (ii) presenting female parasitoids with 1, 2, 4, or 8 egg masses at a time. Offspring developmental length ranged from 18 to 24 days. On average, male offspring exhibited faster developmental times, emerging approximately 1 day ahead of females. The amount of time that adult females spent on an egg mass affected the number of parasitized eggs. Specifically, more offspring emerged in the 4-, 8-, and 16-day treatments than in scenarios involving daily or every second-day egg mass replacement. The percentage of male offspring decreased as the number of egg masses presented to females increased. Interestingly, the total number of female offspring remained constant, but the number of male offspring decreased with an increase in the number of egg masses and time spent by the parent within a patch. The observed sexual dimorphism in development time, the influence of resource availability on offspring sex ratios, and flexible oviposition patterns illustrate the adaptability of O. kuvanae in response to varying conditions. These insights have implications for our understanding of parasitoid-host interactions and their potential role in biological control strategies.

Increased cover of native and exotic plants on the rims of harvester ant (Hymenoptera: Formicidae) nests under grazing and drought.

Harvester ants create habitats along nest rims, which some plants use as refugia. These refugia can enhance ecosystem stability to disturbances like drought and grazing, but their potential role in invasion ecology is not yet tested. Here we examine the effects of drought and grazing on nest-rim refugia of 2 harvester ant species: Pogonomyrmex occidentals and P. rugosus. We selected 4 rangeland sites with high harvester ant nest densities in northern Arizona, USA, with pre-existing grazing exclosures adjacent to heavily grazed habitat. Our objective was to determine whether nest refugia were used by native or exotic plant species for each site and scenario of drought and grazing. We measured vegetation cover on nest surfaces, on nest rims, and at 3 distances (3, 5, and 10 m) from nests. At each site, we sampled 2 treatments (grazed/excluded) during 2 seasons (drought/monsoon). We found that nest rims increased vegetation cover compared with background levels at all sites and in almost all scenarios of treatment and season, indicating that nest rims provide important refugia for plants from drought and cattle grazing. In some cases, plants enhanced on nest rims were native grasses such as blue gramma (Bouteloua gracilis) or forbs such as sunflowers (Helianthus petiolaris). However, nest rims at all sites enhanced exotic species, particularly Russian thistle (Salsola tragus), purslane (Portulaca oleracea), and bull thistle (Cirsium vulgare). These results suggest that harvester ants play important roles in invasion ecology and restoration. We discuss potential mechanisms for why certain plant species use nest-rim refugia and how harvester ant nests contribute to plant community dynamics.

Habitat Type Affects Elevational Patterns in Ground-dwelling Arthropod Communities.

Understanding factors that drive biodiversity distributions is central in ecology and critical to conservation. Elevational gradients are useful for studying the effects of climate on biodiversity but it can be difficult to disentangle climate effects from resource differences among habitat types. Here we compare elevational patterns and influences of environmental variables on ground-dwelling arthropods in open- and forested-habitats. We examine these comparisons in three arthropod functional groups (detritivores, predators, and herbivores) and two taxonomic groups (beetles and arachnids). We sampled twelve sites spanning 1,132 m elevation and four life zones, collecting 4,834 individual ground arthropods identified to 123 taxa. Elevation was a strong predicator for arthropod composition, however, patterns differed among functional and taxonomic groups and individual species between open- and forested-habitats. Beetles, arachnids, and predators decreased with elevation in open habitats but increased in forests showing a significant interaction between habitat type and elevation. Detritivores and herbivores showed no elevational patterns. We found 11 arthropod taxa with linear elevational patterns, seven that peaked in abundance at high elevations, and four taxa at low elevations. We also found eight taxa with parabolic elevational patterns that peaked in abundance at mid-elevations. We found that vegetation composition and productivity had stronger explanatory power for arthropod composition in forested habitats, while ground cover was a stronger predictor in open habitats. Temperature and precipitation were important in both habitats. Our findings demonstrate that relationships between animal diversity and elevation can be mediated by habitat type, suggesting that physiological restraints and resource limitations work differently between habitat types.

Chromosome-level genome assembly and population genomic analyses provide insights into adaptive evolution of the red turpentine beetle, Dendroctonus valens.

BACKGROUND: Biological invasions are responsible for substantial environmental and economic losses. The red turpentine beetle (RTB), Dendroctonus valens LeConte, is an important invasive bark beetle from North America that has caused substantial tree mortality in China. The lack of a high-quality reference genome seriously limits deciphering the extent to which genetic adaptions resulted in a secondary pest becoming so destructive in its invaded area. RESULTS: Here, we present a 322.41 Mb chromosome-scale reference genome of RTB, of which 98% of assembled sequences are anchored onto fourteen linkage groups including the X chromosome with a N50 size of 24.36 Mb, which is significantly greater than other Coleoptera species. Repetitive sequences make up 45.22% of the genome, which is higher than four other Coleoptera species, i.e., Mountain pine beetle Dendroctonus ponderosae, red flour beetle Tribolium castaneum, blister beetle Hycleus cichorii, and Colorado potato beetle Leptinotarsa decemlineata. We identify rapidly expanded gene families and positively selected genes in RTB, which may be responsible for its rapid environmental adaptation. Population genetic structure of RTB was revealed by genome resequencing of geographic populations in native and invaded regions, suggesting substantial divergence of the North American population and illustrates the possible invasion and spread route in China. Selective sweep analysis highlighted the enhanced ability of Chinese populations in environmental adaptation. CONCLUSIONS: Overall, our high-quality reference genome represents an important resource for genomics study of invasive bark beetles, which will facilitate the functional study and decipher mechanism underlying invasion success of RTB by integrating the Pinus tabuliformis genome.

Pinyon Engraver Beetle Acoustics: Stridulation Apparatus, Sound Production and Behavioral Response to Vibroacoustic Treatments in Logs.

Bark beetles are among the most influential biotic agents in conifer forests, and forest management often focuses on bark beetle chemical communication for tree protection. Although acoustic communication occurs in many bark beetle species, we have yet to utilize acoustic communication for bark beetle control. Here, we describe the stridulatory organs and 'stress' chirps of the pinyon engraver, Ips confusus, a significant pest and mortality agent of pinyon pine in western North America. Only females possessed stridulatory organs and their stress chirps varied significantly in duration, pulses per chirp, and dominant frequency. We tested an array of acoustic-vibrational treatments into logs but were unable to disrupt male entry into logs or alter female-male interactions, female tunneling, and female oviposition. We found acoustic-vibrational treatments had little effect on I. confusus behavior and suggest further studies if acoustic methods are to be utilized for bark beetle control.

Field Translocation of Mountain Pine Beetles Suggests Phoretic Mite Communities Are Locally Adapted, and Mite Populations Respond Variably to Climate Warming.

Temperature is a key determining factor in the population dynamics of forest insects and their associated biota. Bark beetles, often considered key agents of change in forest ecosystems, are particularly affected by warming in their environment. Beetles associate with various phoretic mite species that have direct/indirect effects on beetle fitness and population dynamics, although there is limited knowledge of how temperature affects these communities. Here, we use a field reciprocal translocation experiment with the addition of a novel "warming" environment to represent future changes in local environment in two populations of a keystone bark beetle species (Dendroctonus ponderosae). We hypothesize that mite community abundances as carried by bark beetles are significantly altered when not in their native environments and when subjected to climate warming. We use multivariate generalized linear models based on species abundance data to show that mite community compositions significantly differ across different field climates; and that these patterns diverge between source populations, indicating local adaptation. Our study offers foundational information on the general effects of simulated climate-warming on the compositional shifts of common and abundant biotic associates of mountain pine beetles and may be used as a model system for other important insect-mite systems.

Evidence for Semiochemical Divergence Between Sibling Bark Beetle Species: Dendroctonus brevicomis and Dendroctonus barberi.

We investigated geographic variation in the semiochemistry of major disturbance agents of western North American pine forests, Dendroctonus brevicomis Le Conte and Dendroctonus barberi Hopkins (Coleoptera: Curculionidae: Scolytinae), species separated by the Great Basin in the USA that until recently were synonymous. At 15 sites in the western USA and northern Mexico, beetle populations were examined to determine (1) pheromone production by solitary, mining females, (2) male electroantennogram amplitudes in response to known semiochemicals for the genus, or (3) relative attractiveness of two female-produced pheromone components (endo- and exo-brevicomin) and two host odors (alpha-pinene and myrcene) to beetles in the field. Compared to female beetles collected east of the Great Basin (D. barberi), western females (D. brevicomis) produced a consistently higher proportion of, and male antenna were correspondingly more sensitive to, the exo- than the endo-isomer of brevicomin. With the exception of one sampling location (where no preference was observed), beetles west of the Great Basin were more attracted to exo- than endo- brevicomin trap lures, whereas eastern beetles displayed the reverse preference. In contrast, there was not a consistent difference between these populations regarding relative attraction or olfactory response to myrcene or alpha-pinene, although some geographic variability was evident. These data show that the semiochemical systems of D. brevicomis and D. barberi have diverged and corroborate genetic and morphological evidence that they are distinct, allopatric species.

Climate and vegetation structure shape ant communities along elevational gradients on the Colorado Plateau.

Terrestrial animal communities are largely shaped by vegetation and climate. With climate also shaping vegetation, can we attribute animal patterns solely to climate? Our study observes ant community changes along climatic gradients (i.e., elevational gradients) within different habitat types (i.e., open and forest) on the Colorado Plateau in the southwestern United States. We sampled ants and vegetation along two elevational gradients spanning 1,132 m with average annual temperature and precipitation differences of 5.7°C and 645mm, respectively. We used regression analyses and structural equation modeling to compare the explanatory powers and effect sizes of climate and vegetation variables on ants. Climate variables had the strongest correlations and the largest effect sizes on ant communities, while vegetation composition, richness, and primary productivity had relatively small effects. Precipitation was the strongest predictor for most ant community metrics. Ant richness and abundance had a negative relationship with precipitation in forested habitats, and positive in open habitats. Our results show strong direct climate effects on ants with little or no effects of vegetation composition or primary productivity, but contrasting patterns between vegetation type (i.e., forested vs. open) with precipitation. This indicates vegetation structure can modulate climate responses of ant communities. Our study demonstrates climate-animal relationships may vary among vegetation types which can impact both findings from elevational studies and how communities will react to changes in climate.

Bark Beetle Population Dynamics in the Anthropocene: Challenges and Solutions.

Tree-killing bark beetles are the most economically important insects in conifer forests worldwide. However, despite >200 years of research, the drivers of population eruptions and crashes are still not fully understood and the existing knowledge is thus insufficient to face the challenges posed by the Anthropocene. We critically analyze potential biotic and abiotic drivers of population dynamics of an exemplary species, the European spruce bark beetle (ESBB) (Ips typographus) and present a multivariate approach that integrates the many drivers governing this bark beetle system. We call for hypothesis-driven, large-scale collaborative research efforts to improve our understanding of the population dynamics of this and other bark beetle pests. Our approach can serve as a blueprint for tackling other eruptive forest insects.

First report of luminous stimuli eliciting sound production in weevils.

Light-based stimuli elicited acoustic responses in male Hylesinus aculeatus Say (Curculionidae: Scolytinae: Hylesinina) instantaneously, with 100% reliability. Stridulations were elicited with a white light beam in a dark environment and recorded with an ultrasonic microphone. Acoustic responses were consistent, and, when compared with sounds produced under stressful conditions (i.e. physical stimulation), no significant differences were found. Hylesinus aculeatus possess an elytro-tergal stridulatory organ and acoustic communication is only present in males. This is also the first report of acoustic communication for this species. Instantaneous light-elicited acoustic communication has potential applications in the development of electronic traps and real-time acoustic detection and identification of beetles, border biosecurity, and noise-reduction in acoustic data collection.

Phoretic mite communities associated with bark beetles in the maritime and stone pine forests of Setúbal, Portugal.

The phoretic mite communities of prominent bark beetle pests associated with pine stands of southern Portugal were sampled to determine whether they vary across bark beetle species and stand type. Bark beetles were sampled for mites from two primary (aggressive) bark beetle species (Ips sexdentatus and Orthotomicus erosus) and the most common secondary species (Hylurgus ligniperda) in maritime pine (Pinus pinaster) and stone pine (Pinus pinea) in the Setúbal province of Portugal. Twelve mite species, spanning diverse ecological roles, are found associated with these bark beetle systems. The relative abundances of the 12 species that make up the phoretic mite communities of maritime and stone pine varied significantly between host beetle species as well as between stand type, indicating that the phoretic host and dominant tree type are important drivers of mite community composition. The functional roles of these mites are outlined and their ecological significance in pine forest ecosystems is discussed.

Genomoviruses associated with mountain and western pine beetles.

Genomoviruses are circular single-stranded DNA viruses (∼2 kb in size) classified into nine genera, they are highly diverse and have been identified in a variety of samples ranging from fungi to animal sera. Here we identify five genomoviruses belonging to the Gemycircularvirus genus and one to the Gemykibivirus genus from mountain pine beetle and western pine beetle sampled in Arizona. Collectively these six viral genomes share <77% genome-wide pairwise identity and hence represent six new species of genomoviruses. Four of the gemycircularviruses from the mountain pine beetles are recombinant, with one having a recombinant region that spans the entire capsid protein. Pine beetles have a symbiotic relationship with certain tree pathogenic fungi. Therefore given that Sclerotinia sclerotiorum hypovirulence-associated DNA virus 1, a gemycircularvirus, induces hypovirulence in the plant pathogenic fungus Sclerotinia sclerotiorum and infects the mycophagous insect Lycoriella ingenua, it is possible that the six genomoviruses identified here may be directly associated with the pine beetle fungal symbionts and/or with the insects themselves.

The Role of Multimodal Signals in Species Recognition Between Tree-Killing Bark Beetles in a Narrow Sympatric Zone.

When related species coexist, selection pressure should favor evolution of species recognition mechanisms to prevent interspecific pairing and wasteful reproductive encounters. We investigated the potential role of pheromone and acoustic signals in species recognition between two species of tree-killing bark beetles, the southern pine beetle, Dendroctonus frontalis Zimmermann, and the western pine beetle, Dendroctonus brevicomis LeConte, in a narrow zone of sympatry, using reciprocal pairing experiments. Given the choice of adjacent con- or heterospecific female gallery entrance in a log, at least 85% of walking males chose the entrance of the conspecific, and half the males that initially entered heterospecific galleries re-emerged and entered the conspecific gallery within 15 min. Waveform analysis of female acoustic "chirps" indicated interspecific differences in chirp timing. Males may use information from female acoustic signals to decide whether to enter or remain in the gallery. Individuals in forced heterospecific pairings (produced by confinement of a heterospecific male within the female entrance) did not differ in pheromone production from individuals of conspecific pairs. However, due to the absence of the right species of male, galleries with heterospecific pairs released an abnormal pheromone blend that lacked at least one key component of the aggregation pheromone of either species. The complete aggregation pheromone (i.e., the pheromone blend from entrances with pairs) does not appear to deter interspecific encounters or confer premating reproductive isolation per se; however, it may confer selective pressure for the maintenance of other reproductive isolation mechanisms.

Western Pine Beetle Populations in Arizona and California Differ in the Composition of Their Aggregation Pheromones.

We compared pheromone production and response for populations of western pine beetle, Dendroctonus brevicomis LeConte, from sites in northern Arizona and northern California. Volatiles were collected from individuals of both sexes that had mined as a pair in a Pinus ponderosa log for 1 d, and they were subsequently analyzed by gas chromatography coupled to mass-spectrometry. Principal component analysis of quantities of Dendroctonus pheromone components indicated strong site-associated clustering of blend composition for females but not males. Much of the clustering in females evidently was due to differences in the production of endo- and exo-brevicomin, which occurred in average ratios of 0.1:1 and 19:1 for populations in the California and Arizona sites, respectively. In the California site, exo- was better than endo-brevicomin in enhancing trap catches of both sexes to lures containing the host-tree odor α-pinene and the male-produced aggregation pheromone component frontalin. In an identical test in the Arizona site, endo- was a better adjuvant than exo-brevicomin for male attraction, whereas females did not show a significant preference. At neither location were the isomers antagonistic to one another in activity. Thus, one aggregation pheromone has apparently diverged between these populations, concurrent with published evidence that D. brevicomis on either side of the Great Basin are genetically distinct and are possibly different species. Furthermore, production of and response to the isomers of brevicomin by flying Dendroctonus frontalis Zimmermann in the Arizona site were similar to those of sympatric D. brevicomis. This interspecific signal overlap is likely sustainable since joint species mass-attacks may assist both species in overcoming host defenses, thereby increasing host availability.

Structure of Phoretic Mite Assemblages Across Subcortical Beetle Species at a Regional Scale.

Mites associated with subcortical beetles feed and reproduce within habitats transformed by tree-killing herbivores. Mites lack the ability to independently disperse among these habitats, and thus have evolved characteristics that facilitate using insects as transport between resources. Studies on associations between mites and beetles have historically been beetle-centric, where an assemblage of mite species is characterized on a single beetle species. However, available evidence suggests there may be substantial overlap among mite species on various species of beetles utilizing similar host trees. We assessed the mite communities of multiple beetle species attracted to baited funnel traps in Pinus stands in southern Wisconsin, northern Arizona, and northern Georgia to better characterize mite dispersal and the formation of mite-beetle phoretic associations at multiple scales. We identified approximately 21 mite species totaling 10,575 individuals on 36 beetle species totaling 983 beetles. Of the mites collected, 97% were represented by eight species. Many species of mites were common across beetle species, likely owing to these beetles' common association with trees in the genus Pinus. Most mite species were found on at least three beetle species. Histiostoma spp., Iponemus confusus Lindquist, Histiogaster arborsignis Woodring and Trichouropoda australis Hirschmann were each found on at least seven species of beetles. While beetles had largely similar mite membership, the abundances of individual mite species were highly variable among beetle species within each sampling region. Phoretic mite communities also varied within beetle species between regions, notably for Ips pini (Say) and Ips grandicollis (Eichhoff).

Technique for studying arthropod and microbial communities within tree tissues.

Phloem tissues of pine are habitats for many thousands of organisms. Arthropods and microbes use phloem and cambium tissues to seek mates, lay eggs, rear young, feed, or hide from natural enemies or harsh environmental conditions outside of the tree. Organisms that persist within the phloem habitat are difficult to observe given their location under bark. We provide a technique to preserve intact phloem and prepare it for experimentation with invertebrates and microorganisms. The apparatus is called a 'phloem sandwich' and allows for the introduction and observation of arthropods, microbes, and other organisms. This technique has resulted in a better understanding of the feeding behaviors, life-history traits, reproduction, development, and interactions of organisms within tree phloem. The strengths of this technique include the use of inexpensive materials, variability in sandwich size, flexibility to re-open the sandwich or introduce multiple organisms through drilled holes, and the preservation and maintenance of phloem integrity. The phloem sandwich is an excellent educational tool for scientific discovery in both K-12 science courses and university research laboratories.

Use of acoustics to deter bark beetles from entering tree material.

BACKGROUND: Acoustic technology is a potential tool to protect wood materials and eventually live trees from colonization by bark beetles. Bark beetles such as the southern pine beetle Dendroctonus frontalis, western pine beetle D. brevicomis and pine engraver Ips pini (Coleoptera: Curculionidae) use chemical and acoustic cues to communicate and to locate potential mates and host trees. In this study, the efficacy of sound treatments on D. frontalis, D. brevicomis and I. pini entry into tree materials was tested. RESULTS: Acoustic treatments significantly influenced whether beetles entered pine logs in the laboratory. Playback of artificial sounds reduced D. brevicomis entry into logs, and playback of stress call sounds reduced D. frontalis entry into logs. Sound treatments had no effect on I. pini entry into logs. CONCLUSION: The reduction in bark beetle entry into logs using particular acoustic treatments indicates that sound could be used as a viable management tool.