David C. Lightfoot

Diversity of Arthropod Responses to Host-plant Water Stress in a Desert Ecosystem in Southern New Mexico

Abstract Previous studies of insect-plant interactions have produced the contradictory Plant Stress Hypothesis (that stressed plants are more suitable hosts for phytophages) and Plant Vigor Hypothesis (that vigorous plants are more suitable hosts for phytophages). However, experimental studies of phytophage responses to host stress have involved only one, or a few, related phytophagous species, not whole communities of organisms associated with a particular plant species. We evaluated responses of various arthropods associated with creosotebush Larrea tridentata to manipulated water availability and plant stress in southern New Mexico during 1990–1991. Of 44 arthropod groups (taxa or functional groups) evaluated in our study only two taxa (including a lepidopteran folivore) showed significant negative response to water availability, thereby supporting the Plant Stress Hypothesis. Ten taxa (including eight phytophages) responded positively to water availability, supporting the Plant Vigor Hypothesis. One phytophage showed a nonlinear response, supporting neither hypothesis. Detrended Correspondence Analysis significantly distinguished the arthropod community on water-deprived shrubs from the communities on watered shrubs. The variation in responses among phytophagous insects on creosotebush indicated that the effect of plant water stress likely reflects the choice of phytophage, and perhaps the host plant, being studied. Therefore, neither the Plant Stress Hypothesis nor the Plant Vigor Hypothesis can explain responses of all phytophages on a particular plant species.

bottom-up regulation of plant community structure in an aridland ecosystem

We conducted a long-term rodent exclosure experiment in native grass- and shrub-dominated vegetation to evaluate the importance of top-down and bottom-up controls on plant community structure in a low-productivity aridland ecosystem. Using multiple regressions and analysis of covariance, we assessed how bottom-up precipitation pulses cascade through vegetation to affect rodent populations, how rodent populations affect plant community structure, and how rodents alter rates of plant community change over time. Our findings showed that bottom-up pulses cascade through the system, increasing the abundances of plants and rodents, and that rodents exerted no control on plant community structure and rate of change in grass-dominated vegetation, and only limited control in shrub-dominated vegetation. These results were discussed in the context of top-down effects on plant communities across broad gradients of primary productivity. We conclude that bottom-up regulation maintains this ecosystem in a state of low primary productivity that constrains the abundance of consumers such that they exert limited influence on plant community structure and dynamics.

Variation in Insect Densities on Desert Creosotebush: Is Nitrogen a Factor?

A field experiment was conducted to assess the effects of nitrogen and water on the densities and taxonomic and trophic compositions of arthropods on the foliage of desert creosotebush (Larrea tridentata). Nitrogen and water were manipulated through a series of fertilizer and irrigation treatments applied to naturally growing creosotebush shrubs at a site in the northern Chihuahuan Desert. Water with nitrogen, and nitrogen fertilization alone, significantly increased creosotebush foliage production and foliar nitrogen contents. Water alone had less effect on foliage production and no effect on foliar nitrogen contents. Foliage production, foliar nitrogen contents, and numbers of foliage arthropods were all higher in the late spring than in the mid or late summer. Numbers of foliage arthropods increased significantly on fertilized plants in the late spring. Water treatments alone had no effect on numbers of foliage arthropods, but a positive water-fertilizer interaction effect on numbers of foliage arthropods was evident in the late spring. Overall, there were sig? nificant positive correlations between foliage production, foliar nitrogen contents, and foliage arthropod densities. Phytophagous sap-sucking insects accounted for the majority of arthropods on creosotebush, and their densities varied most in relation to foliage pro? duction and foliar nitrogen contents. Results of our study demonstrated that sap-sucking phytophagous insects are more responsive than leaf-chewing insects to increased nitrogen content of creosotebush foliage, and that much ofthe between-plant variation in densities of phytophagous insects within a stand of creosotebush may be due to sap-sucking insects tracking variable foliar nitrogen.

Rapid Response of a Grassland Ecosystem to an Experimental Manipulation of a Keystone Rodent and Domestic Livestock

Megaherbivores and small burrowing mammals commonly coexist and play important functional roles in grassland ecosystems worldwide. The interactive effects of these two functional groups of herbivores in shaping the structure and function of grassland ecosystems are poorly understood. In North Americas central grasslands, domestic cattle (Bos taurus) have supplanted bison (Bison bison), and now coexist with prairie dogs (Cynomys spp.), a keystone burrowing rodent. Understanding the ecological relationships between cattle and prairie dogs and their independent and interactive effects is essential to understanding the ecology and important conservation issues affecting North American grassland ecosystems. To address these needs, we established a long-term manipulative experiment that separates the independent and interactive effects of prairie dogs and cattle using a 2 x 2 factorial design. Our study is located in the Janos-Casas Grandes region of northwestern Chihuahua, Mexico, which supports one of the largest remaining complexes of black-tailed prairie dogs (C. ludovicianus). Two years of posttreatment data show nearly twofold increases in prairie dog abundance on plots grazed by cattle compared to plots without cattle. This positive effect of cattle on prairie dogs resulted in synergistic impacts when they occurred together. Vegetation height was significantly lower on the plots where both species co-occurred compared to where either or both species was absent. The treatments also significantly affected abundance and composition of other grassland animal species, including grasshoppers and banner-tailed kangaroo rats (Dipodomys spectabilis). Our results demonstrate that two different functional groups of herbivorous mammals, burrowing mammals and domestic cattle, have distinctive and synergistic impacts in shaping the structure and function of grassland ecosystems.

Productivity of Creosotebush Foliage and Associated Canopy Arthropods Along a Desert Roadside

-A sampling study was designed to test the hypothesis that creosotebush (Larrea tridentata) shrubs along a roadside were larger, more vigorous and supported greater populations of foliage arthropods than shrubs growing 20 m away from the road margin. Structural and chemical attributes of shrubs and associated foliage arthropods were measured. Roadside shrubs were larger, had denser foliage, more flowers, higher foliar nitrogen contents and lower foliar resin contents than shrubs growing away from the road. Foliage arthropod densities were significantly higher on roadside shrubs. Sap-feeding herbivores dominated numerically and accounted for most of the differences in arthropod abundances between roadside and nonroadside shrubs. Numbers of foliage arthropods were positively correlated with shrub size, density and foliar nitrogen contents, and negatively correlated with foliar resin contents. These findings, in concordance with other studies, indicate that in arid environments productive, vigorous plants are preferred hosts for herbivorous insects.

Fruit Abortion in Yucca elata and Its Implications for the Mutualistic Association with Yucca Moths

The yucca moth Tegelicula yuccasella is the sole pollinator of Yucca elata, on yucca fruits to complete its life cycle. A high percentage of pollinated flowers aborts, killing the T. yuccasella eggs and larvae in them. We examined patterns of fruit production and abortion in Y. elata, and related these patterns to vegetative characteristics, moth abundance, and environmental conditions. We studied 38 inflorescences throughout their flowering period, during one season in southern New Mexico, USA. Each night we recorded the number of flowers opening, the number of fruit formed, the relative abundance of yucca moths, and climatic conditions. We monitored 11786 flowers, resulting in 699 mature fruit. Large inflorescences produced more, but proportionately fewer fruit than small inflorescences

Interplant variation in creosotebush foliage characteristics and canopy arthropods

SummaryWe conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.

Phytophagous insects enhance nitrogen flux in a desert creosotebush community

SummaryWe tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.

Pollination ecology of Yucca elata

The pollination biology of a population of 250 Yucca elata (Liliaceae) plants was studied in southern New Mexico. Yucca elata and the prodoxid yucca moth Tegeticula yuccasella have a mutualistic association that is essential for the successful sexual reproduction of both species. However, a wide range of other invertebrate species visit flowers during the day and at night. Our aim was to quantify the role of yucca moths and other invertebrate visitors in pollination and fruit set, using manipulative field experiments. Inflorescences were bagged during the day or night (N=12 inflorescences) to restrict flower visitors to either nocturnal or diurnal groups. Yucca moths were active exclusively nocturnally during the flowering period and thus did not visit inflorescences that were unbagged during the day. None of the 4022 flowers exposed only to diurnal visitors set fruit, whereas 4.6% of the 4974 flowers exposed only to nocturnal visitors (including yucca moths) produced mature fruit. The proportion of flowers producing fruit in the latter treatment was not significantly different from unbagged control inflorescences. In a series of experimental manipulations we also determined that: (1) flowers opened at dusk and were open for two days on average, but were only receptive to pollen on the first night of opening; (2) pollen must be pushed down the stigmatic tube to affect pollination; and (3) most plants require out-cross pollination to produce fruit. The combination of these results strongly suggests that yucca moths are the only species affecting pollination in Y. elata, and that if another species was to affect pollination, it would be a rare event.

Geographic Patterns of Ground-dwelling Arthropods Across an Ecoregional Transition in the North American Southwest

Abstract We examined the biogeographic patterns of ground-dwelling arthropod communities across a heterogeneous semiarid region of the Southern Rio Grande Rift Valley of New Mexico. Our 3 sites included portions of 5 ecoregions, with the middle site a transition area where all ecoregions converged. We addressed the following 3 questions: (1) Do the species assemblage patterns for ground arthropods across habitats and sites conform to recognized ecoregions? (2) Are arthropod assemblages in distinct vegetation-defined habitats within an ecoregion more similar to each other or to assemblages in similar vegetation-defined habitats in other ecoregions? (3) Is there a detectable edge effect with increased arthropod diversity in the area of converging ecoregions? We encountered 442 target arthropod species from pitfall traps operating continuously for 7 years over a series of different habitats at each of the 3 sites. We examined geographic distributions of spider and cricket/grasshopper species in detail, and they showed affinities for different ecoregions, respectively. Each habitat within a study site supported a unique overall arthropod assemblage; nevertheless, different habitats at the same site were more similar to each other than they were to similar habitats at other sites. Overall arthropod species richness was greatest in the area where all 5 ecoregions converged. Arthropod species and their geographic distributions are poorly known relative to vascular plants and vertebrate animals. Findings from this research indicate that ecoregional classification is a useful tool for understanding biogeographic patterns among arthropods.