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SignificanceIn species with internal fertilization, sperm spend an important part of their lives within the female. To examine the life history of the sperm during this time, we used semiquantitative proteomics and sex-specific isotopic labeling in fruit flies to determine the extent of molecular continuity between male and female reproductive tracts and provide a global catalog of sperm-associated proteins. Multiple seminal fluid proteins and female proteins associate with sperm immediately after mating. Few seminal fluid proteins remain after long-term sperm storage, whereas female-derived proteins constitute one-fifth of the postmating sperm proteome by then. Our data reveal a molecular "hand-off" from males to females, which we postulate to be an important component of sperm-female interactions.

Significance In species with internal fertilization, sperm spend an important part of their lives within the female. To examine the life history of the sperm during this time, we used semiquantitative proteomics and sex-specific isotopic labeling in fruit flies to determine the extent of molecular continuity between male and female reproductive tracts and provide a global catalog of sperm-associated proteins. Multiple seminal fluid proteins and female proteins associate with sperm immediately after mating. Few seminal fluid proteins remain after long-term sperm storage, whereas female-derived proteins constitute one-fifth of the postmating sperm proteome by then. Our data reveal a molecular “hand-off” from males to females, which we postulate to be an important component of sperm–female interactions.

Many sexually selected ornaments and weapons are elaborations of an animal’s outer body surface, including long feathers, colorful skin, and rigid outgrowths. The time and energy required to keep these traits clean, attractive, and in good condition for signaling may represent an important but understudied cost of bearing a sexually selected trait. Male fiddler crabs possess an enlarged and brightly colored claw that is used both as a weapon to fight with rival males and also as an ornament to court females. Here, we demonstrate that males benefit from grooming because females prefer males with clean claws over dirty claws but also that the time spent grooming detracts from the amount of time available for courting females. Males, therefore, face a temporal trade-off between attracting the attention of females and maintaining a clean claw. Our study provides rare evidence of the importance of grooming for mediating sexual interactions in an invertebrate, indicating that sexual selection has likely shaped the evolution of self-maintenance behaviors across a broad range of taxa.

Evolution explains both the unity and the diversity of all organisms, and developing students' ability to represent and communicate evolutionary relationships is an important component of a complete biology education. We present a series of student-centered, exploratory activities to help students develop their tree-thinking skills. In these activities, students use complementary phenotypic and molecular data to explore how to build phylogenetic trees and interpret the evolutionary relationships they represent. This learning module is designed to engage students in the process of science, provide them with active learning experiences using online bioinformatics tools, and foster their appreciation for the evolutionary connections across the tree of life.

In addition to increasing interest in the supply risk of minerals produced in China, there is also concern that China's efforts to mitigate its own mineral supply risk—through overseas foreign direct investment—may limit mineral availability for other countries in the short-term (due to production capacity constraints). However, little is publicly known about the quantity of global mineral production that is subject to Chinese ownership influence or how this influence may mitigate China's exposure to mineral commodity supply risk. In this analysis, we estimate China's ownership-share of foreign production for cobalt mine and intermediate materials (in the year 2016), as well as the net import reliance of China's cobalt refinery industry—which is then adjusted to reflect China's ownership share of foreign production. First, we find that China's foreign cobalt ownership is predominantly in the Democratic Republic of the Congo, the largest source of cobalt mine and intermediate imports for China's growing cobalt refinery industry. Second, overseas foreign direct investment provided China with ownership influence over roughly 29% of its cobalt mine and intermediate imports in 2016—which may have reduced the supply risk exposure of China's refinery industry from a net import reliance of 97% (on cobalt mine and intermediate materials) to an adjusted net import reliance of 68%. Third, China's global production share jumps from 2% to 14% (for cobalt mine material) and from 11% to 33% (for cobalt intermediate material) when China's ownership-share of foreign production is added to China's domestic production share. Fourth, over time China's foreign direct investment appears to have targeted facilities with progressively larger cobalt production capacities. Finally, China's global production share increases as cobalt material moves downstream (i.e. mine, 14%; intermediate, 33%; refined, 50% in 2016). If a primary motivation of China's Going Out Strategy is to secure natural resources, then China's global production share may be reserved for Chinese manufacturers. As a result, this analysis indicates that—for countries outside China—concerns related to critical mineral availability and supply risk may not be confined to minerals produced in China.

Mineral criticality is a subjective concept that has evolved throughout history. An abundance of literature on this topic has been published over the last decade, encompassing a variety of criteria and methodologies. To our knowledge, this work is the first large-scale effort to organize and analyze recent comprehensive criticality studies in order to determine if a consensus exists within the global community as to which elements are critical. Here, we set aside methodological differences and analyze the results of 32 comprehensive nonfuel mineral criticality studies that evaluate at least 10 elements. Of the 56 elements or elemental groups evaluated, the three most commonly identified as critical in these studies are the rare-earth elements (REE), the platinum-group metals (PGM), and indium. Most of the studies also identify tungsten, germanium, cobalt, niobium, tantalum, gallium, and antimony as critical. These results are consistent with the 11 most recent studies, published post-2014, which also identify bismuth as critical. Furthermore, the same elements identified in the complete dataset, except antimony, were designated as critical when normalized by geographic region. Magnesium was also deemed critical. Elements may be identified consistently as critical for several reasons; similarities in methodologies, which embody evolving perceptions of risk, or changing national and institutional priorities. This work compiles a large number of recent criticality studies in an effort to define a consensus of currently critical materials, essentially defining the modern criticality paradigm, which is valuable when interpreting an individual perspective in more global context.

Abstract Many animal signals are brightly colored and convey information about species identity as well as information about individual conspecifics. Colorful bird and lizard signals have received much attention, and many studies have related specific spectral properties of these signals to variation in mating success and territory defense. Far less attention has been given to invertebrates even though there are spectacularly colorful species. The enlarged claw of the male banana fiddler crab Austruca mjoebergi, for example, is bright yellow and contrasts vividly against the mudflat substrate. It is used in waving displays to attract females and in male territory defense and combat. Claw color varies among males in the degree of “yellowness,” ranging very pale yellow to orange. In this study, we examined female responses to claw color variation in two-choice tests using robotic crabs. We found that although females strongly discriminate against colors that fall outside the natural range of intensity, hue, and chroma, they show no consistent preferences for different claw colors within the natural range, and no single component of claw color (hue, chroma, or intensity) independently affected female choices. Using three-choice tests, we also showed that female preferences induce stabilizing selection on male claw color. We conclude that, although claw color is sufficient to facilitate species recognition, it is unlikely to be used in intraspecific mate choice to provide information about male quality. Significance statement Fiddler crabs are often brightly colored, are visually orientated animals, and have a highly complex social system. Despite this, there are few studies that have looked at the role of color in species recognition and mate choice in these animals. In this study, we use robotic crabs with painted claws to determine the role of claw color in species recognition and mate choice in the banana fiddler crab, Austruca mjoebergi. We found that color is important in conspecific mate recognition but the variation among males in claw “yellowness” is unlikely to be used by females in intraspecific mate choice decisions.

When females mate with more than one male, sexual selection acts both before and after mating. The interaction between pre- and postmating episodes of selection is expected to be context dependent, but few studies have investigated how total sexual selection changes under different ecological conditions. We examined how population density mediates the interaction between pre- and postmating sexual selection by establishing replicate populations of the horned dung beetle Onthophagus taurus at low, medium, and high densities, and then using microsatellite-based parentage analyses to measure male fitness. We found that mating success and fertilization success were positively correlated at all three densities, but the strength of the correlation decreased with increasing density. We also found a shift from negative to positive linear selection on testes mass as density increased, and opposing selection on weapons and testes at high densities. These patterns suggest that the importance of postmating processes increases with increasing population density, which reduces the selective advantage of weapons for premating contest competition, and increases the selective advantage of large ejaculates for postmating sperm competition. We expect that density-dependent selection on testes mass has contributed to the phenotypic variation observed between natural populations of O. taurus that differ in density.

Increasing reliance on non-renewable mineral resources reinforces the need for identifying potential supply constraints before they occur. The US National Science and Technology Council recently released a report that outlines a methodology for screening potentially critical minerals based on three indicators: supply risk (R), production growth (G), and market dynamics (M). This early-warning screening was initially applied to 78 minerals across the years 1996 to 2013 and identified a subset of minerals as “potentially critical” based on the geometric average of these indicators—designated as criticality potential (C). In this study, the screening methodology has been updated to include data for 2014, as well as to incorporate revisions and modifications to the data, where applicable. Overall, C declined in 2014 for the majority of minerals examined largely due to decreases in production concentration and price volatility. However, the results vary considerably across minerals, with some minerals, such as gallium, recording increases for all three indicators. In addition to assessing magnitudinal changes, this analysis also examines the significance of the change relative to historical variation for each mineral. For example, although mined nickel’s R declined modestly in 2014 in comparison to that of other minerals, it was by far the largest annual change recorded for mined nickel across all years examined and is attributable to Indonesia’s ban on the export of unprocessed minerals. Based on the 2014 results, 20 minerals with the highest C values have been identified for further study including the rare earths, gallium, germanium, rhodium, tantalum, and tungsten.

When females mate with multiple males, they set the stage for postcopulatory sexual selection via sperm competition and/or cryptic female choice. Surprisingly little is known about the rates of multiple mating by females in the wild, despite the importance of this information in understanding the potential for postcopulatory sexual selection to drive the evolution of reproductive behaviour, morphology and physiology. Dung beetles in the genus Onthophagus have become a laboratory model for studying pre- and postcopulatory sexual selection, yet we still lack information about the reproductive behaviour of female dung beetles in natural populations. Here, we develop microsatellite markers for Onthophagus taurus and use them to genotype the offspring of wild-caught females and to estimate natural rates of multiple mating and patterns of sperm utilization. We found that O. taurus females are highly polyandrous: 88% of females produced clutches sired by at least two males, and 5% produced clutches with as many as five sires. Several females (23%) produced clutches with significant paternity skew, indicating the potential for strong postcopulatory sexual selection in natural populations. There were also strong positive correlations between the number of offspring produced and both number of fathers and paternity skew, which suggests that females benefit from mating polyandrously by inciting postcopulatory mechanisms that bias paternity towards males that can sire more viable offspring. This study evaluates the fitness consequences of polyandry for an insect in the wild and provides strong evidence that female dung beetles benefit from multiple mating under natural conditions.

The U.S. Geological Survey National Minerals Information Center (NMIC) is the U.S. Government agency tasked with the collection, analysis, and dissemination of information on the production, consumption, import, export, and other measures of the flows of non-fuel mineral commodities of importance to the U.S. economy and national security. The NMIC and its agency predecessors have maintained a database of this information, collected and published annually, dating back to the beginning of the twentieth century. Time series analysis of annual information from the NMIC provides the opportunity to identify trends in the supply chains of the minerals and metals which are increasingly in demand for advanced technologies. The identification of trends in data for net import reliance, country concentration of production, global demand, price volatility, and other measures, when combined with world governance indicators, can be used to focus attention on individual mineral commodities where supply chain restrictions may develop. Specific examples for U.S. net import reliance, global tantalum primary mining, and mineral criticality screening are presented to illustrate the utility of time series analysis of trends in mineral commodity supply and demand, the types of data required, and the limitations of currently available information.

Understanding how organisms mitigate the impacts of climate warming is one of the biggest challenges facing modern-day biologists. For tropical ectotherms, staying cool is critical for avoiding thermal stress, so individuals that are able to maintain territories in cool microhabitats are likely to gain fitness advantages. This study evaluated the importance of shade availability in the tropical fiddler crab, Austruca mjoebergi, by investigating temperature variation and behavioral responses (distribution, activity level, and time budget) in sun-exposed and shaded microhabitats. We found that the daily temperatures in sun-exposed microhabitats were significantly higher than in shady microhabitats, and that more male crabs held territories in the shade. Shaded males were active on the sediment surface for longer periods each day, allowing more time for feeding and more opportunities to detect mate-searching females. The greater number of males resident in the shade, and shorter distances between male neighbors in the shade, suggested that there was less available space for individuals to move into the shaded microhabitats. The behavioral differences between sun- and shade-living residents are likely to have consequences for male fitness. We highlight the importance of shade in providing thermal refugia for tropical ectotherms. The impact of climate warming on tropical organisms could be less dire if individuals are able to access shade. Shade can offer thermal refugia for tropical organisms that may have important consequences for both behavior and fitness. Relatively low and stable temperatures in shaded areas allow male fiddler crabs to remain on the surface for longer compared to sun-exposed males, allowing for longer feeding periods and more mating opportunities.

Sexually selected weapons often function as honest signals of fighting ability. If poor-quality individuals produce high-quality weapons, then receivers should focus on other, more reliable signals. Cost is one way to maintain signal integrity. The costs of weapons tend to increase with relative weapon size, and thereby restrict large weapons to high-quality individuals who can produce and maintain them. Weapon cost, however, appears to be unpredictably variable both within and across taxa, and the mechanisms underlying this variation remain unclear. We suggest variation in weapon cost may result from variation in weapon composition—specifically, differences in the amount of muscle mass directly associated with the weapon. We test this idea by measuring the metabolic cost of sexually selected weapons in seven arthropod species and relating these measures to weapon muscle mass. We show that individuals with relatively large weapon muscles have disproportionately high resting metabolic rates and provide evidence that this trend is driven by weapon muscle mass. Overall, our results suggest that variation in weapon cost can be partially explained by variation in weapon morphology and that the integrity of weapon signals may be maintained by increased metabolic cost in species with relatively high weapon muscle mass.

Fertility depends on the progression of complex and coordinated postmating processes within the extracellular environment of the female reproductive tract (FRT). Molecular interactions between ejaculate and FRT proteins regulate many of these processes, including sperm motility, migration, storage, and modification, along with concurrent changes in the female. Although extensive progress has been made in the proteomic characterization of male-derived components of sperm and seminal fluid, investigations into the FRT have remained more limited. To achieve a comparable level of knowledge regarding female-derived proteins that comprise the reproductive environment, we utilized semiquantitative mass spectrometry-based proteomics to study the composition of the FRT tissue and, separately, the luminal fluid, before and after mating in Drosophila melanogaster. Our approach leveraged whole-fly isotopic labelling to delineate female proteins from those transferred male ejaculate proteins. Our results revealed several characteristics that distinguish the FRT fluid proteome from the FRT tissue proteome: (1) the fluid proteome is encoded by genes with higher overall levels of FRT gene expression and tissue specificity, including many genes with enriched expression in the fat body, (2) fluid-biased proteins are enriched for metabolic functions and (3) the fluid exhibits pronounced postmating compositional changes. The dynamic mating-induced proteomic changes in the FRT fluid informs our understanding of secretory mechanisms of the FRT, serves as a foundation for establishing female contributions to the ejaculate-female interactions that regulate fertility and highlights the importance of applying proteomic approaches to characterize the composition and dynamics of the FRT environment.

Male contest competition and female choice are key features of the mating systems of many species, and whole-organism performance may be targeted by both mechanisms given the dynamic body movements required during fighting and courtship displays. Using the dimorphic horned dung beetle Onthophagus taurus as a model system, we tested whether physical performance was important in determining a male’s victory in fights, and whether successful fighters were preferred by females as mates. We found that physical strength, horn length, and body mass were significant predictors of male fighting success, but males that won fights were not more attractive to females. Rather, females preferred males that delivered a high courtship rate, which was not correlated with strength, horn length, or body mass, but previously has been shown to be genetically correlated with body condition. The fact that there was no relationship between fighting success and mating success suggests that selection on traits favored by male–male competition and female choice can act relatively independently in this species, although both mechanisms appear to favor traits (strength and courtship, respectively) that are linked to a male’s ability to acquire and allocate resources for mass gain. Future work is needed to determine the relative contributions of these processes to the total strength of sexual selection acting on male phenotypes.

Emerald ash borer is expected to kill thousands of ash trees in the eastern U.S. This research develops tools to predict the effect of ash tree loss from the urban canopy on landslide susceptibility in Pittsburgh, PA. A spatial model was built using the SINMAP (Stability INdex MAPping) model coupled with spatially explicit scenarios of tree loss (0%, 25%, 50%, and 75% loss of ash trees from the canopy). Ash spatial distributions were estimated via Monte Carlo methods and available vegetation plot data. Ash trees are most prevalent on steeper slopes, likely due to urban development patterns. Therefore, ash loss disproportionately increases hillslope instability. A 75% loss of ash resulted in roughly 800 new potential landslide initiation locations. Sensitivity testing reveals that variations in rainfall rates, and friction angles produce minor changes to model results relative to the magnitude of parameter variation, but reveal high model sensitivity to soil density and root cohesion values. The model predictions demonstrate the importance of large canopy species to urban hillslope stability, particularly on steep slopes and in areas where soils tend to retain water. To improve instability predictions, better characterization of urban soils, particularly spatial patterns of compaction and species specific root cohesion is necessary. The modeling framework developed in this research will enhance assessment of changes in landslide risk due to tree mortality, improving our ability to design economically and ecologically sustainable urban systems.

Exaggerated horns are a characteristic feature of many male rhinoceros beetles. We surveyed and compared the scaling relationships of these sexually selected weapons for 31 Dynastinae species with different degrees of horn exaggeration. We found that nearly all rhinoceros beetle species were male dimorphic, that the allometric slope of major males was consistently shallower than the slope of minor males, and that the decrease in slope was greatest among species with the most exaggerated horns. These patterns are consistent with the curved allometries of stag beetle mandibles and giraffe weevil rostra, and suggest that the depletion of developmental resources is a general phenomenon limiting the continued exaggeration of insect weapons. The dimorphisms in horn morphology are expected to correspond to behavioural differences between major and minor males, but little is still known about the mating tactics of most rhinoceros beetle species. Future studies on the relative benefits and performance of horns during male–male combat are needed to fully understand the diversity of horn allometries and the evolution of exaggerated structures.

Infiltration-based green infrastructure has become a popular means of reducing stormwater hazards in urban areas. However, the long-term effects of green infrastructure on the geochemistry of roadside environments are poorly defined, particularly given the considerable roadside legacy metal contamination from historic industrial activity and vehicle emissions (e.g., Pb). Most current research on green infrastructure geochemistry is restricted to time periods of less than a year or limited sets of chemical species. This further limits our understanding of systems that evolve over time and are subject to seasonal variability. Between 2016 and 2018, two infiltration trenches in Pittsburgh, PA, were monitored to determine infiltration rates and dissolved nutrient and metal content. The trench water was analyzed to characterize seasonal patterns in both trench function and chemistry. Shifting patterns in infiltration rate and geochemical activity show trends corresponding with seasonal changes. Trench function is dependent on the local water table, with the highest infiltration rates occurring when evapotranspiration is active and groundwater elevation is low. Two seasonal chemical patterns were identified. The first is driven by road salt application in the winter and interaction of the salt pulse increase Pb and Cu concentrations. The second is driven by the formation of summer reducing environments that increase dissolved Fe and Mn. These findings suggest that chemical and hydrological activity in infiltration-based green infrastructure varies seasonally and may remobilize legacy contamination.

Sexual selection has equipped male rhinoceros beetles with large horns on their head and prothorax to aid in battle over access to females. Horns are used to pry and dislodge opponents from resource sites that attract females, so an optimal horn should be able both to withstand the high stresses imposed during fights, and to resist deflection in response to these loads. We examined the cross-sectional morphology of horns using microcomputed tomography scanning to determine how horn structure changes with horn length to withstand the different fighting loads. Specifically, we measured the second moment of area of horns within and among rhinoceros beetle species to assess whether changes in cross-sectional morphology accompany changes in body size in order to maintain high strength and stiffness during fights. We find that the second moment of area of horns increases with body size both intra-specifically and inter-specifically, and that these relationships closely fit those predicted if horns have been selected to be strong and stiff fighting structures. Our results therefore support the hypothesis that rhinoceros beetle horns are structurally adapted for combat. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 00: 000–000.