Pinyon pine mortality alters communities of ground-dwelling arthropods

We documented the effect of drought-induced mortality of pinyon pine (Pinus edulis Engelm.) on communities of ground-dwelling arthropods. Tree mortality alters microhabitats utilized by ground-dwelling arthropods by increasing solar radiation, dead woody debris, and understory vegetation. Our major objectives were to determine (1) whether there were changes in community composition, species richness, and abundance of ground-dwelling arthropods associated with pinyon mortality and (2) whether specific habitat characteristics and microhabitats accounted for these changes. We predicted shifts in community composition and increases in arthropod diversity and abundance due to the presumed increased complexity of microhabitats from both standing dead and fallen dead trees. We found significant differences in arthropod community composition between high and low pinyon mortality environments, despite no differences in arthropod abundance or richness. Overall, 22% (51 taxa) of the arthropod community were identified as being indicators of either high or low mortality. Our study corroborates other research indicating that arthropods are responsive to even moderate disturbance events leading to changes in the environment. These arthropod responses can be explained in part due to the increase in woody debris and reduced canopy cover created by tree mortality

Ground-dwelling arthropod responses to succession in a pinyon-juniper woodland

Stand-replacing wildfire is an infrequent but important disturbance in southwestern pinyon-juniper woodlands. A typical successional cycle in these woodlands is approximately 300 years or more after a stand-replacing fire. Arthropods, especially ground-dwelling taxa, are one of the most abundant and diverse fauna in terrestrial ecosystems and are typically responsive to microhabitat change. Little is known regarding community responses of ground-dwelling arthropods to changes in woodland successional stages from early ecosystems dominated by grasses, herbaceous plants, and fire adapted shrubs to tree-dominated old-growth ecosystems. In 2007 and 2008, within Mesa Verde National Park, Colorado, we compared the community composition of ground-dwelling arthropods between old-growth pinyon-juniper stands that were 300–400 years old and early successional areas recovering from a stand-replacing fire in 2002. The 2002 fire eliminated the dominant woody vegetation, which was replaced by increased herbaceous vegetation and bare ground. The early successional arthropod community showed a significantly higher abundance in major arthropod taxonomic groups, except spiders, compared to old-growth woodland. Old-growth species richness was greater in late August–September, 2007 and greater in early successional habitats during April–July, 2008. Spatial variability of the habitat was much greater in the recently burned early successional plots than the old-growth late successional plots. The differences in habitat were strongly correlated with arthropod community composition, suggesting that ground-dwelling arthropods are very sensitive to habitat changes. Habitat affiliation was strong, with 83% (early succession ruderal) and 91% (old-growth woodland) of the species found primarily or exclusively in one habitat. Many habitat indicator species (defined as species found in significantly greater abundance in one habitat) were found in both burned and old-growth habitats. Several species were found to be strict specialists exclusive to only one of these habitats. Collectively, the results suggest that heightened concern over loss of old-growth woodlands is warranted, given the distinct nature of the ground-dwelling arthropod community in old-growth habitats

The Indestructible Insect: Velvet Ants From Across the United States Avoid Predation by Representatives From All Major Tetrapod Clades

Velvet ants are a group of parasitic wasps that are well known for a suite of defensive adaptations including bright coloration and a formidable sting. While these adaptations are presumed to function in antipredator defense, observations between potential predators and this group are lacking. We conducted a series of experiments to determine the risk of velvet ants to a host of potential predators including amphibians, reptiles, birds, and small mammals. Velvet ants from across the United States were tested with predator\u27s representative of the velvet ants native range. All interactions between lizards, free-ranging birds, and a mole resulted in the velvet ants survival, and ultimate avoidance by the predator. Two shrews did injure a velvet ant, but this occurred only after multiple failed attacks. The only predator to successfully consume a velvet ant was a single American toad (Anaxyrus americanus). These results indicate that the suite of defenses possessed by velvet ants, including aposematic coloration, stridulations, a chemical alarm signal, a hard exoskeleton, and powerful sting are effective defenses against potential predators. Female velvet ants appear to be nearly impervious to predation by many species whose diet is heavily derived of invertebrate prey

Herbivore-mediated negative frequency-dependent selection underlies a trichome dimorphism in nature

Authors are grateful for funding provided by an NSF GRFP to J.K.G. (2015195769) and DEB‐1353970 to L.F.D.Negative frequency‐dependent selection (NFDS) has been shown to maintain polymorphism in a diverse array of traits. The action of NFDS has been confirmed through modeling, experimental approaches, and genetic analyses. In this study, we investigated NFDS in the wild using morph‐frequency changes spanning a 20‐year period from over 30 dimorphic populations of Datura wrightii. In these populations, plants either possess glandular (sticky) or non‐glandular (velvety) trichomes, and the ratio of these morphs varies substantially among populations. Our method provided evidence that NFDS, rather than drift or migration, is the primary force maintaining this dimorphism. Most populations that were initially dimorphic remained dimorphic, and the overall mean and variance in morph frequency did not change over time. Furthermore, morph‐frequency differences were not related to geographic distances. Together, these results indicate that neither directional selection, drift, or migration played a substantial role in determining morph frequencies. However, as predicted by negative frequency‐dependent selection, we found that the rare morph tended to increase in frequency, leading to a negative relationship between the change in the frequency of the sticky morph and its initial frequency. In addition, we found that morph‐frequency change over time was significantly correlated with the damage inflicted by two herbivores: Lema daturaphila and Tupiochoris notatus. The latter is a specialist on the sticky morph and damage by this herbivore was greatest when the sticky morph was common. The reverse was true for L. daturaphila, such that damage increased with the frequency of the velvety morph. These findings suggest that these herbivores contribute to balancing selection on the observed trichome dimorphism.Publisher PDFPeer reviewe

Seismic imaging of the metamorphism of young sediment into new crystalline crust in the actively rifting Imperial Valley, California

Plate-boundary rifting between transform faults is opening the Imperial Valley of southern California and the rift is rapidly filling with sediment from the Colorado River. Three 65–90 km long seismic refraction profiles across and along the valley, acquired as part of the 2011 Salton Seismic Imaging Project, were analyzed to constrain upper crustal structure and the transition from sediment to underlying crystalline rock. Both first arrival travel-time tomography and frequency-domain full-waveform inversion were applied to provide P-wave velocity models down to ∼7 km depth. The valley margins are fault-bounded, beyond which thinner sediment has been deposited on preexisting crystalline rocks. Within the central basin, seismic velocity increases continuously from ∼1.8 km/s sediment at the surface to >6 km/s crystalline rock with no sharp discontinuity. Borehole data show young sediment is progressively metamorphosed into crystalline rock. The seismic velocity gradient with depth decreases approximately at the 4 km/s contour, which coincides with changes in the porosity and density gradient in borehole core samples. This change occurs at ∼3 km depth in most of the valley, but at only ∼1.5 km depth in the Salton Sea geothermal field. We interpret progressive metamorphism caused by high heat flow to be creating new crystalline crust throughout the valley at a rate comparable to the ≥2 km/Myr sedimentation rate. The newly formed crystalline crust extends to at least 7–8 km depth, and it is shallower and faster where heat flow is higher. Most of the active seismicity occurs within this new crust

A three-dimensional model for the probabilistic intergranular failure of polycrystalline arrays

A three-dimensional grain model, in which the grains are represented by regular truncated octahedra, has been developed to study probabilistic time-dependent intergranular failure in polycrystalline arrays. In this model, grain boundary facets are assumed to fail randomly in time, as a function of the facet normal stress. A simple approximate method for calculating the load shed by failed facets and a reasonable choice of failure criterion complete the model. This leads to a conceptually simple, but computationally complex, model capable of handling assemblages consisting of relatively large numbers (> 5000) of grains. The predicted scatter in the times-to-failure and the variation in number of failed facets with time are in quite reasonable agreement with available experimental data.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/49082/2/ms6302.pd

Sexual Dimorphism of Staminate- and Pistillate-Phase Flowers of Saponaria officinalis (Bouncing Bet) Affects Pollinator Behavior and Seed Set

The sequential separation of male and female function in flowers of dichogamous species allows for the evolution of differing morphologies that maximize fitness through seed siring and seed set. We examined staminate- and pistillate-phase flowers of protandrous Saponaria officinalis for dimorphism in floral traits and their effects on pollinator attraction and seed set. Pistillate-phase flowers have larger petals, greater mass, and are pinker in color, but due to a shape change, pistillate-phase flowers have smaller corolla diameters than staminate-phase flowers. There was no difference in nectar volume or sugar content one day after anthesis, and minimal evidence for UV nectar guide patterns in staminate- and pistillate-phase flowers. When presented with choice arrays, pollinators discriminated against pistillate-phase flowers based on their pink color. Finally, in an experimental garden, in 2012 there was a negative correlation between seed set of an open-pollinated, emasculated flower and pinkness (as measured by reflectance spectrometry) of a pistillate-phase flower on the same plant in plots covered with shade cloth. In 2013, clones of genotypes chosen from the 2012 plants that produced pinker flowers had lower seed set than those from genotypes with paler flowers. Lower seed set of pink genotypes was found in open-pollinated and hand-pollinated flowers, indicating the lower seed set might be due to other differences between pink and pale genotypes in addition to pollinator discrimination against pink flowers. In conclusion, staminate- and pistillate-phase flowers of S. officinalis are dimorphic in shape and color. Pollinators discriminate among flowers based on these differences, and individuals whose pistillate-phase flowers are most different in color from their staminate-phase flowers make fewer seeds. We suggest morphological studies of the two sex phases in dichogamous, hermaphroditic species can contribute to understanding the evolution of sexual dimorphism in plants without the confounding effects of genetic differences between separate male and female individuals

Two's Company, Three's a Crowd: Experimental Evaluation of the Evolutionary Maintenance of Trioecy in Mercurialis annua (Euphorbiaceae)

Trioecy is an uncommon sexual system in which males, females, and hermaphrodites co-occur as three clearly different gender classes. The evolutionary stability of trioecy is unclear, but would depend on factors such as hermaphroditic sex allocation and rates of outcrossing vs. selfing. Here, trioecious populations of Mercurialis annua are described for the first time. We examined the frequencies of females, males and hermaphrodites across ten natural populations and evaluated the association between the frequency of females and plant densities. Previous studies have shown that selfing rates in this species are density-dependent and are reduced in the presence of males, which produce substantially more pollen than hermaphrodites. Accordingly, we examined the evolutionary stability of trioecy using an experiment in which we (a) indirectly manipulated selfing rates by altering plant densities and the frequency of males in a fully factorial manner across 20 experimental plots and (b) examined the effect of these manipulations on the frequency of the three sex phenotypes in the next generation of plants. In the parental generation, we measured the seed and pollen allocations of hermaphrodites and compared them with allocations by unisexual plants. In natural populations, females occurred at higher frequencies in denser patches, a finding consistent with our expectations. Under our experimental conditions, however, no combination of plant densities and male frequencies was associated with increased frequencies of females. Our results suggest that the factors that regulate female frequencies in trioecious populations of M. annua are independent of those regulating male frequencies (density), and that the stable co-existence of all three sex phenotypes within populations is unlikely