Phylogenetic risk assessment is robust for forecasting the impact of European insects on North American conifers.

Some introduced species cause severe damage, although the majority have little impact. Robust predictions of which species are most likely to cause substantial impacts could focus efforts to mitigate those impacts or prevent certain invasions entirely. Introduced herbivorous insects can reduce crop yield, fundamentally alter natural and managed forest ecosystems, and are unique among invasive species in that they require certain host plants to succeed. Recent studies have demonstrated that understanding the evolutionary history of introduced herbivores and their host plants can provide robust predictions of impact. Specifically, divergence times between hosts in the native and introduced ranges of a nonnative insect can be used to predict the potential impact of the insect should it establish in a novel ecosystem. However, divergence time estimates vary among published phylogenetic datasets, making it crucial to understand if and how the choice of phylogeny affects prediction of impact. Here, we tested the robustness of impact prediction to variation in host phylogeny by using insects that feed on conifers and predicting the likelihood of high impact using four different published phylogenies. Our analyses ranked 62 insects that are not established in North America and 47 North American conifer species according to overall risk and vulnerability, respectively. We found that results were robust to the choice of phylogeny. Although published vascular plant phylogenies continue to be refined, our analysis indicates that those differences are not substantial enough to alter the predictions of invader impact. Our results can assist in focusing biosecurity programs for conifer pests and can be more generally applied to nonnative insects and their potential hosts by prioritizing surveillance for those insects most likely to be damaging invaders.

High species richness and turnover of vascular epiphytes is associated with water availability along the elevation gradient of Volcán Maderas, Nicaragua.

Research that has been conducted documenting species richness patterns on tropical mountains has resulted in conflicting observations: monotonic declines with increasing elevation, monotonic increases with increasing elevation, and a mid-elevation "bulge." Currently, it is unclear if these differences are due to environmental differences among study areas, the taxonomic groups or ecological groups (e.g., growth form) sampled, or the scale of study along elevation gradients. Because of the difficulty in sampling and identifying canopy-dwelling plants, the number of inventories quantifying tropical epiphytes is relatively limited and recent. In this study, we provide a detailed qualitative and quantitative assessment of the vascular epiphyte flora and its spatial distribution on Volcán Maderas, Isla de Ometepe, Nicaragua, including weather and environmental measurements along the entire elevation gradient of the volcano. We sampled epiphytes in five distinct forest types associated with increasing elevation as follows: dry forest, humid forest, wet forest, cloud forest, and elfin forest. Five weather stations were placed along the elevation gradient for us to relate observed patterns to environmental conditions. A mid-elevation peak in species richness was detected for all vascular epiphytes at approximately 1000 m in elevation (cloud forest), yet epiphyte abundance increased with increasing elevation. In total we identified 206 taxa of vascular epiphytes belonging to 26 families and 73 genera. The most species-rich family was the Orchidaceae with 55 species for the entire elevation gradient, followed by Bromeliaceae (29 species), Araceae (23), Polypodiaceae (25), Dryopteridaceae (16), and Piperaceae (11), with all other families represented by fewer than 10 species each. We found that richness patterns differ phylogenetically across epiphyte groups, possibly due to different adaptive strategies, and species for the most part appear to be narrowly distributed within specific habitat zones along the elevation gradient. Variables associated with moisture, precipitation, humidity, mist, or cloud cover are key to understanding the observed patterns.

Collections Education: The Extended Specimen and Data Acumen.

Biodiversity scientists must be fluent across disciplines; they must possess the quantitative, computational, and data skills necessary for working with large, complex data sets, and they must have foundational skills and content knowledge from ecology, evolution, taxonomy, and systematics. To effectively train the emerging workforce, we must teach science as we conduct science and embrace emerging concepts of data acumen alongside the knowledge, tools, and techniques foundational to organismal biology. We present an open education resource that updates the traditional plant collection exercise to incorporate best practices in twenty-first century collecting and to contextualize the activities that build data acumen. Students exposed to this resource gained skills and content knowledge in plant taxonomy and systematics, as well as a nuanced understanding of collections-based data resources. We discuss the importance of the extended specimen in fostering scientific discovery and reinforcing foundational concepts in biodiversity science, taxonomy, and systematics.

Small herbaria contribute unique biogeographic records to county, locality, and temporal scales.

PREMISE: With digitization and data sharing initiatives underway over the last 15 years, an important need has been prioritizing specimens to digitize. Because duplicate specimens are shared among herbaria in exchange and gift programs, we investigated the extent to which unique biogeographic data are held in small herbaria vs. these data being redundant with those held by larger institutions. We evaluated the unique specimen contributions that small herbaria make to biogeographic understanding at county, locality, and temporal scales. METHODS: We sampled herbarium specimens of 40 plant taxa from each of eight states of the United States of America in four broad status categories: extremely rare, very rare, common native, and introduced. We gathered geographic information from specimens held by large (≥100,000 specimens) and small (<100,000 specimens) herbaria. We built generalized linear mixed models to assess which features of the collections may best predict unique contributions of herbaria and used an Akaike information criterion-based information-theoretic approach for our model selection to choose the best model for each scale. RESULTS: Small herbaria contributed unique specimens at all scales in proportion with their contribution of specimens to our data set. The best models for all scales were the full models that included the factors of species status and herbarium size when accounting for state as a random variable. CONCLUSIONS: We demonstrated that small herbaria contribute unique information for research. It is clear that unique contributions cannot be predicted based on herbarium size alone. We must prioritize digitization and data sharing from herbaria of all sizes.

Seeds attached to refrigerated shipping containers represent a substantial risk of nonnative plant species introduction and establishment.

The initial processes for successful biological invasions are transport, introduction, and establishment. These can be directly influenced or completely avoided through activities that reduce the number and frequency of entering nonnative propagules. Economic and environmental benefits through preventative monitoring programs at early stages of invasion far outweigh the long-term costs associated with mitigating ecological and economic impacts once nonnative species establish and spread. In this study, we identified 30 taxa of hitchhiking plant propagules on the air-intake grilles of refrigerated shipping containers arriving into a United States seaport from a port on the Pacific coast of South America. The four monocotyledonous taxa with the highest number of seeds collected were analyzed; we estimated propagule pressure, germination, and survivorship of these taxa, and we used the estimates to determine likelihood of establishment. At the levels of propagule pressure estimated here, non-zero germination and survival rates resulted in high establishment probabilities even when escape rates from shipping containers were modelled to be exceedingly low. Our results suggest high invasion risk for nonnative taxa including Saccharum spontaneum L., a listed Federal Noxious Weed. Currently, not all shipping containers arriving at USA ports are thoroughly inspected due to limited personnel and funding for biological invasion prevention. Our results indicate that there is a significant risk from only a few propagules escaping into the environment from this source, and we propose possible solutions for reducing this risk.

An initial industrial flora: A framework for botanical research in cooperation with industry for biodiversity conservation.

Humans have created an accelerating, increasingly connected, globalized economy, resulting in a more globalized, shared flora. The prevention of new, establishing species is less costly, both economically and ecologically, and is more manageable than eradicating nonnative invasive species once they are widespread and negatively impactful. We ask if international trade hubs and points-of-entry with high-volume trade, constant disturbance, and propagule rain have a higher number of nonnative species compared to surrounding areas and if they may serve as initial establishment sites and refugia of nonnative, invasive populations. Therefore, we partnered with various federal, state, and private interests to evaluate the floristic composition at the Garden City Terminal of the Port of Savannah, Georgia, USA. We conducted the following study to demonstrate the collaborative relationship-building between researchers and industry and to develop a framework for biodiversity conservation. In our study, we collected all reproductive vascular plants in the secured areas of the Garden City Terminal during four major seasonal time points over two years. The percent of nonnative species and number of nonnative plant species per hectare at this industrial location exceeded all other comparison floras. The mean coefficient of conservatism was lowest among the comparison floras, indicating a highly disturbed habitat with nonnative, weedy native, and other native species tolerant of disturbance. Our study represents one of the first inventories of an Industrialized Flora and indicates that such areas are hot-spots of nonnative plant diversity and possible sources of emergent plant invasions. We posit that industrial sites and international points-of-entry should be considered laboratories for research on species transport and introduction, adaptability, and taxonomic delineation to better understand the mechanisms and consequences of biotic homogenization due to the volume and frequency of anthropogenic activities.

Evolutionary history predicts high-impact invasions by herbivorous insects.

A long-standing goal of invasion biology is to identify factors driving highly variable impacts of non-native species. Although hypotheses exist that emphasize the role of evolutionary history (e.g., enemy release hypothesis & defense-free space hypothesis), predicting the impact of non-native herbivorous insects has eluded scientists for over a century.Using a census of all 58 non-native conifer-specialist insects in North America, we quantified the contribution of over 25 factors that could affect the impact they have on their novel hosts, including insect traits (fecundity, voltinism, native range, etc.), host traits (shade tolerance, growth rate, wood density, etc.), and evolutionary relationships (between native and novel hosts and insects).We discovered that divergence times between native and novel hosts, the shade and drought tolerance of the novel host, and the presence of a coevolved congener on a shared host, were more predictive of impact than the traits of the invading insect. These factors built upon each other to strengthen our ability to predict the risk of a non-native insect becoming invasive. This research is the first to empirically support historically assumed hypotheses about the importance of evolutionary history as a major driver of impact of non-native herbivorous insects.Our novel, integrated model predicts whether a non-native insect not yet present in North America will have a one in 6.5 to a one in 2,858 chance of causing widespread mortality of a conifer species if established (R 2 = 0.91) Synthesis and applications. With this advancement, the risk to other conifer host species and regions can be assessed, and regulatory and pest management efforts can be more efficiently prioritized.

Studies of Jatrogossone A as a Reactive Oxygen Species Inducer in Cancer Cellular Models.

Natural products continue to provide a platform to study biological systems. A bioguided study of cancer cell models led us to a new member of the jatrophane natural products from Jatropha gossypiifolia, which was independently identified and characterized as jatrogossone A (1). Purification and structure elucidation was performed by column chromatography and high-performance liquid chromatography-mass spectrometry and NMR techniques, and the structure was confirmed via X-ray crystallography. The unique molecular scaffold of jatrogossone A prompted an evaluation of its mode of action. Cytotoxicity assays demonstrated that jatrogossone A displays selective antiproliferative activity against cancer cell models in the low micromolar range with a therapeutic window. Jatrogossone A (1) affects mitochondrial membrane potential (ΔΨm) in a time- and dose-dependent manner. This natural product induces radical oxygen species (ROS) selectively in cancer cellular models, with minimal ROS induction in noncancerous cells. Compound 1 induces ROS in the mitochondria, as determined by colocalization studies, and it induces mitophagy. It promotes also in vitro cell death by causing cell arrest at the G2/M stage, caspase (3/7) activation, and PARP-1 cleavage. The combined findings provide a potential mechanism by which 1 relies on upregulation of mitochondrial ROS to potentiate cytotoxic effects through intracellular signaling.

The influence of herbivory and weather on the vital rates of two closely related cactus species.

Herbivory has long been recognized as a significant driver of plant population dynamics, yet its effects along environmental gradients are unclear. Understanding how weather modulates plant-insect interactions can be particularly important for predicting the consequences of exotic insect invasions, and an explicit consideration of weather may help explain why the impact can vary greatly across space and time. We surveyed two native prickly pear cactus species (genus Opuntia) in the Florida panhandle, USA, and their specialist insect herbivores (the invasive South American cactus moth, Cactoblastis cactorum, and three native insect species) for five years across six sites. We used generalized linear mixed models to assess the impact of herbivory and weather on plant relative growth rate (RGR) and sexual reproduction, and we used Fisher's exact test to estimate the impact of herbivory on survival. Weather variables (precipitation and temperature) were consistently significant predictors of vital rate variation for both cactus species, in contrast to the limited and varied impacts of insect herbivory. Weather only significantly influenced the impact of herbivory on Opuntia humifusa fruit production. The relationships of RGR and fruit production with precipitation suggest that precipitation serves as a cue in determining the trade-off in the allocation of resources to growth or fruit production. The presence of the native bug explained vital rate variation for both cactus species, whereas the invasive moth explained variation only for O. stricta. Despite the inconsistent effect of herbivory across vital rates and cactus species, almost half of O. stricta plants declined in size, and the invasive insect negatively affected RGR and fruit production. Given that fruit production was strongly size-dependent, this suggests that O. stricta populations at the locations surveyed are transitioning to a size distribution of predominantly smaller sizes and with reduced sexual reproduction potential.

Digitizing specimens in a small herbarium: A viable workflow for collections working with limited resources.

PREMISE OF THE STUDY: Small herbaria represent a significant portion of herbaria in the United States, but many are not digitizing their collections. METHODS: At the Arkansas State University Herbarium (STAR), we have created a viable workflow to help small herbaria begin the digitization process, including suggestions for publishing data on the Internet. We calculated hourly rates of each phase of the digitization process. We also mapped accessions at the county level to determine geographic strengths in the collection. RESULTS: All 17,678 accessioned flowering plant specimens at STAR are imaged, databased in Specify, and available electronically on the herbarium's website. Students imaged the specimens at a mean rate of 145/h. We found differences in databasing rates between the graduate student leading the project (47/h) and undergraduate assistants (25/h). The majority of specimens at STAR were collected within the counties neighboring the institution. DISCUSSION: With this workflow, we estimate that one person can digitize a 20,000-specimen collection in less than 2.5 yr by working only 10 h/wk. Because STAR is a small herbarium with limited resources, the application of the workflow described should assist curators of similar-sized collections as they contemplate and undertake the digitization process.

Digitization workflows for flat sheets and packets of plants, algae, and fungi.

Effective workflows are essential components in the digitization of biodiversity specimen collections. To date, no comprehensive, community-vetted workflows have been published for digitizing flat sheets and packets of plants, algae, and fungi, even though latest estimates suggest that only 33% of herbarium specimens have been digitally transcribed, 54% of herbaria use a specimen database, and 24% are imaging specimens. In 2012, iDigBio, the U.S. National Science Foundation's (NSF) coordinating center and national resource for the digitization of public, nonfederal U.S. collections, launched several working groups to address this deficiency. Here, we report the development of 14 workflow modules with 7-36 tasks each. These workflows represent the combined work of approximately 35 curators, directors, and collections managers representing more than 30 herbaria, including 15 NSF-supported plant-related Thematic Collections Networks and collaboratives. The workflows are provided for download as Portable Document Format (PDF) and Microsoft Word files. Customization of these workflows for specific institutional implementation is encouraged.

Semiochemicals from ex situ abiotically stressed cactus tissue: a contributing role of fungal spores?

Semiochemicals play a central role in communication between plants and insects, such as signaling the location of a suitable host. Fungi on host plants can also play an influential role in communicating certain plant vulnerabilities to an insect. The spiroketal conophthorin is an important semiochemical produced by developing fungal spores. Spiroketals are also used as signals for scolytid communication. Plants and fungi are known to emit varying volatile profiles under biotic and abiotic stress. This paper reports distinctive temporal-volatile profiles from three abiotic treatments, room temperature (control), -15 °C (cold), and -15 °C to room temperature (shock), of cactus tissue plugs. Volatiles from the three treatments included monoterpenes from control plugs, compounds of varying classes and origin at later stages for cold plugs, and known semiochemicals, including spiroketals, at later stages for shock plugs. The results highlight several important findings: a unique tissue source of the spiroketals; abiotic cold-shock stress is indicated as the cause of spiroketal production; and, given previous findings of spirogenesis, fungal spore involvement is a probable biosynthetic origin of the spiroketals. These findings suggest an important role of fungal volatiles as signaling plant vulnerability to insects.

Host plant defense signaling in response to a coevolved herbivore combats introduced herbivore attack.

Defense-free space resulting from coevolutionarily naïve host plants recently has been implicated as a factor facilitating invasion success of some insect species. Host plants, however, may not be entirely defenseless against novel herbivore threats. Volatile chemical-mediated defense signaling, which allows plants to mount specific, rapid, and intense responses, may play a role in systems experiencing novel threats. Here we investigate defense responses of host plants to a native and exotic herbivore and show that (1) host plants defend more effectively against the coevolved herbivore, (2) plants can be induced to defend against a newly-associated herbivore when in proximity to plants actively defending against the coevolved species, and (3) these defenses affect larval performance. These findings highlight the importance of coevolved herbivore-specific defenses and suggest that naïveté or defense limitations can be overcome via defense signaling. Determining how these findings apply across various host-herbivore systems is critical to understand mechanisms of successful herbivore invasion.

Underutilized resources for studying the evolution of invasive species during their introduction, establishment, and lag phases.

The early phases of biological invasions are poorly understood. In particular, during the introduction, establishment, and possible lag phases, it is unclear to what extent evolution must take place for an introduced species to transition from established to expanding. In this study, we highlight three disparate data sources that can provide insights into evolutionary processes associated with invasion success: biological control organisms, horticultural introductions, and natural history collections. All three data sources potentially provide introduction dates, information about source populations, and genetic and morphological samples at different time points along the invasion trajectory that can be used to investigate preadaptation and evolution during the invasion process, including immediately after introduction and before invasive expansion. For all three data sources, we explore where the data are held, their quality, and their accessibility. We argue that these sources could find widespread use with a few additional pieces of data, such as voucher specimens collected at certain critical time points during biocontrol agent quarantine, rearing, and release and also for horticultural imports, neither of which are currently done consistently. In addition, public access to collected information must become available on centralized databases to increase its utility in ecological and evolutionary research.

Translocation experiments with butterflies reveal limits to enhancement of poleward populations under climate change.

There is a pressing need to predict how species will change their geographic ranges under climate change. Projections typically assume that temperature is a primary fitness determinant and that populations near the poleward (and upward) range boundary are preadapted to warming. Thus, poleward, peripheral populations will increase with warming, and these increases facilitate poleward range expansions. We tested the assumption that poleward, peripheral populations are enhanced by warming using 2 butterflies (Erynnis propertius and Papilio zelicaon) that co-occur and have contrasting degrees of host specialization and interpopulation genetic differentiation. We performed a reciprocal translocation experiment between central and poleward, peripheral populations in the field and simulated a translocation experiment that included alternate host plants. We found that the performance of both central and peripheral populations of E. propertius were enhanced during the summer months by temperatures characteristic of the range center but that local adaptation of peripheral populations to winter conditions near the range edge could counteract that enhancement. Further, poleward range expansion in this species is prevented by a lack of host plants. In P. zelicaon, the fitness of central and peripheral populations decreased under extreme summer temperatures that occurred in the field at the range center. Performance in this species also was affected by an interaction of temperature and host plant such that host species strongly mediated the fitness of peripheral individuals under differing simulated temperatures. Altogether we have evidence that facilitation of poleward range shifts through enhancement of peripheral populations is unlikely in either study species.