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Alpine Trophic Interactions, Aphids, and Plant-Animal Networks

Investigates how predator-prey dynamics, ant-aphid mutualisms, and plant phenology interact across trophic levels in Rocky Mountain alpine and subalpine ecosystems, with particular attention to how snowmelt timing and climate shifts cascade through food webs.

K. A. MooneyA. S. NelsonE. H. Mooneymutualistic relationshipsfacilitative interactionstrophic cascadeData from: Progressive sensitivity of trophic leveData from: Elevational cline in herbivore abundancData from: Testing trade-offs and the dominance-imExperimental predator manipulation with colony establishmentpitfall trapping (Formicidae)Ant exclusion experiment on aphid coloniesSnowmelt affects aphid (<i>Aphis asclepladis</i>) Vegetation and environmental patterns in the CrestAdvanced phenology of higher trophic levels shifts

Knowledge Graph (350 nodes, 1318 connections)

Research Primer

Background

Mountain ecosystems in the Gunnison Basin are woven together by webs of interaction among plants, insects, and vertebrates. A central thread in this web is the set of mutualistic relationships between ants and sap-feeding insects such as aphids and treehoppers. In these associations, the herbivorous insects tap plant phloem and excrete sugar-rich honeydew that ants collect as food; in return, ants protect their partners from predators and parasitoid wasps. Because ants can also prey on the very insects they tend, the cost-benefit ratio of these interactions is fluid, shifting with ant species identity, ant nutritional state, aphid colony size, host plant condition, and weather. Understanding how these shifting partnerships scale up to shape plant reproduction, insect populations, and ultimately the structure of subalpine meadows is the core purpose of research in this area.

Several concepts recur throughout the findings below. A trophic cascade is an indirect effect that travels through a food chain, as when a predator changes the behavior or abundance of an intermediate consumer and thereby alters plants at the base. Trophic-level sensitivity is the idea that organisms higher in the food chain are often more vulnerable to environmental change, such as warming or drought, than the plants they depend on. Sexual dimorphism in plant-insect interactions refers to the fact that, in dioecious species like the subalpine herb Valeriana edulis (valerian), male and female plants can differ in their water use, chemistry, and attractiveness to insects, producing different arthropod communities on each sex. Associative learning describes how ants link chemical cues from plants with food rewards, allowing them to navigate the landscape as foragers. Finally, an aridity gradient, produced by combinations of temperature and precipitation across elevation, structures how strongly each of these interactions plays out.

These concepts matter for the Gunnison Basin because subalpine meadows around Gothic, Colorado are undergoing earlier snowmelt, warmer growing seasons, shifting ungulate and carnivore populations, and rising levels of nitrogen deposition. Each of these pressures can ripple through ant-aphid-plant networks in ways that alter wildflower reproduction and insect abundance, with consequences for land managers, pollinator conservation, and long-term ecosystem monitoring.

Foundational work

The research area was anchored in the late 1970s and 1980s by studies at and around the Rocky Mountain Biological Laboratory that used aphids, ants, and butterflies as tractable systems for testing mutualism theory. Addicott documented how multiple aphid species on fireweed partitioned their host plants and competed for the services of tending ants, showing that ants were a limited and limiting resource and that their effects on aphid populations were density-dependent and species-specific (Addicott, 1978) (Addicott, 1978) (Addicott, 1979) (Cushman & Addicott, 1989). In parallel, Pierce and colleagues demonstrated that ants attending larvae of the lycaenid butterfly Glaucopsyche lygdamus reduced attack by parasitoid wasps, providing one of the clearest early cases of protection mutualism driving the evolution of an insect-ant association (Pierce & Mead, 1981) (Pierce & Easteal, 1986).

The botanical context for these interactions was set by Langenheim's classic description of vegetation and environmental patterns in the Crested Butte area (Langenheim, 1962), and by Bierzychudek and Eckhart's theoretical treatment of how male and female plants of dioecious species can occupy different microhabitats (Bierzychudek & Eckhart, 1988). Later work on gynodioecious Geranium richardsonii showed that female plants trade reduced pollinator attraction for increased seed production (Williams et al., 2000), foreshadowing a broader research program on how plant sex shapes interactions with insects.

Key findings

A central, repeatedly confirmed result is that ant tending of sap-feeding insects is highly conditional. Ants can simultaneously protect and prey on aphids, producing net effects that swing from positive to negative depending on context (Billick et al., 2007). Temporal variability across years often matters more than spatial variability among sites in determining whether a given ant-treehopper partnership is mutualistic (Billick & Tonkel, 2003), and not all ant species are equivalent partners: in Glaucopsyche lygdamus, only Formica podzolica clearly reduced parasitism, while other ant species were neutral or even parasitic on the association (Fraser et al., 2001). A recent synthesis places these patterns in a deep evolutionary frame, showing that ant-hemipteran mutualisms arose independently multiple times and that benefits and costs are inherently context dependent (Nelson & Mooney, 2022).

A second body of work shows that plant sex and plant phenology structure these interactions. Female Valeriana edulis plants support several-fold higher densities of aphids, predators, and ants than males (Mooney lab work, 2013), and aphids preferentially colonize female plants in the field (Perry, 2009). Ant tending and aphid colonization peak sharply during host plant flowering stages on Ligusticum porteri and decline afterward (Mooney et al., 2023) (Kennamer, 2022). Ants themselves behave as botanists, using associative learning to link plant chemical blends with honeydew rewards (Sentner, 2017) (Zapata, 2018) (Nelson et al., 2020), and their willingness to tend aphids depends on their own nutritional state, with protein-fed colonies tending aphids far more than carbohydrate-fed colonies (Petry & Mooney, 2009) (Petry et al., 2012).

Climate and other global-change drivers run through this network in ways that often hit higher trophic levels hardest. Snowmelt date is the single best predictor of year-to-year variation in aphid and predator abundance, with earlier snowmelt reducing aphid numbers by creating water-stressed host plants (Robinson et al., 2017) (Mooney et al., 2020) (Theibault, 2014). Across elevation, ant-aphid mutualism strength declines as conditions cool and aridity decreases, with ants boosting aphid colony survival by 66% at low elevations but having no detectable effect at high elevations (Nelson et al., 2019) (Nelson et al., 2019). In Valeriana edulis, sex-specific water use has driven populations to become more male-biased with warming, shifting at roughly 175 meters per decade (Petry et al., 2016). Meanwhile, chronic low-level nitrogen deposition increases the abundance of ant-protected herbivores on sagebrush (Grinath, 2021) and can dampen trophic cascades from bears through ants to plants (Grinath, 2018). Vertebrates matter too: human activity near the Gothic field station shifts mule deer vigilance and flight behavior at scales of hundreds of meters (Price et al., 2014), and coyotes avoid human-dense areas in ways that feed back to deer and wildflowers (Waser et al., 2014).

Current frontier

Early work in the 1980s and 1990s built the conceptual scaffolding of protection mutualisms and plant sexual systems. Research in the 2000s and 2010s emphasized context dependence, host plant effects, and ant learning. Since 2020, the frontier has shifted toward climate-driven phenology, elevational gradients in interaction strength, and the integration of vertebrate consumers into these insect-plant networks. Recent studies show that host plant phenological stage interacts with temperature to determine aphid colony growth and ant recruitment (Mooney et al., 2023), and that bird predation on insects intensifies with elevation and aridity through behavioral rather than density-dependent mechanisms (Dean et al., 2024). New work on ant community ecology shows that different facets of dominance, behavioral, numerical, and ecological, are not equivalent and predict foraging trade-offs in different ways (Nelson & Mooney, 2025) (Sheard et al., 2020).

At the same time, long-term demographic records for Valeriana edulis are being paired with soil and climate data to ask how sessile plants will track shifting conditions. Soil grain size emerges as a surprisingly strong driver of survival, flowering, and seed production (Boxwell, 2025), density dependence in valerian does not vary systematically with elevation (Davis, 2023), and climate-driven population models can now nowcast species distributions from vital rates (Zeh, 2021). Vertebrate-focused work is extending the network upward: studies of black bear foraging on thatching ant nests (Hunter, 2023), deer browse on Ligusticum porteri (Rittler, 2024), and differential responses of deer, coyotes, and foxes to recreation intensity (Uetrecht et al., 2023) connect insect-plant mutualisms to landscape-scale human impacts.

Open questions

Several questions stand out for the next decade. How will the mismatch between aphid, host plant, and ant phenology develop as snowmelt continues to advance, and will earlier-emerging predators like lygus bugs increasingly dismantle ant-aphid mutualisms before they form? How do chronic nitrogen deposition, warming, and changing ungulate pressure combine, rather than act singly, to reshape trophic cascades in meadows and sagebrush steppe? Can the learning and foraging behavior of individual ant species be used to predict which mutualisms will persist under novel plant chemistries and community compositions? For dioecious plants such as Valeriana edulis, what are the long-term consequences of male-biased sex ratio shifts for seed production, pollinator communities, and the arthropod assemblages that track female plants? And how tightly coupled are vertebrate consumers like bears, deer, and insectivorous birds to the insect mutualisms below them, so that managing recreation, grazing, or predator populations can be connected to conservation of pollinators and wildflowers in the Gunnison Basin?

References

Addicott, J. F. (1978). Competition for mutualists: aphids and ants. Canadian Journal of Zoology.

Addicott, J. F. (1978). Niche relationships among species of aphids feeding on fireweed. Canadian Journal of Zoology.

Addicott, J. F. (1979). A multispecies aphid-ant association: density dependence and species-specific effects. Canadian Journal of Zoology.

Evans (1989). Seed protection by ants foraging on the extrafloral nectaries of the aspen sunflower, Helianthella quinquenervis.

Petry & Mooney (2009). A balanced diet: effects of ant nutritional state on the balance between mutualism and predation upon aphids.

Perry (2009). Variation in host plant sex mediates ant-aphid interactions.

Petry et al. (2012). Influence of macronutrient imbalance on native ant foraging and interspecific interactions in the field.

Petry et al. (2013). Mechanisms underlying plant sexual dimorphism in multi-trophic arthropod communities.

Waser et al. (2014). Coyotes, deer, and wildflowers: diverse evidence points to a trophic cascade.

Theibault (2014). Effects of early snowmelt and climate warming on Valeriana edulis and the insects that depend on it.

Robinson et al. (2017). Multitrophic interactions mediate the effects of climate change on herbivore abundance.

Sentner (2017). The ability of ants to associatively learn based on olfactory chemical cues produced by plants.

Zapata (2018). Are ants botanists?: ant associative learning of plant volatiles.

Grinath (2018). Short-term, low-level nitrogen deposition dampens a trophic cascade between bears and plants.

Nelson et al. (2019). Elevational cline in herbivore abundance driven by a monotonic increase in trophic level sensitivity to aridity.

Nelson et al. (2019). Progressive sensitivity of trophic levels to warming underlies an elevational gradient in ant-aphid mutualism strength.

Nelson et al. (2020). Are ants botanists? Ant associative learning of plant chemicals mediates foraging for carbohydrates.

Mooney et al. (2020). Early snowmelt reduces aphid abundance Aphis asclepiadis by creating water stressed host plants.

Sheard et al. (2020). Testing trade-offs and the dominance-impoverishment rule among ant communities.

Zeh (2021). Nowcasting the distribution of Valeriana edulis using climate driven population models.

Bierzychudek, P., & Eckhart, V. (1988). Spatial segregation of the sexes of dioecious plants. American Naturalist.

Billick, I., & Tonkel, K. (2003). The relative importance of spatial vs. temporal variability in generating a conditional mutualism. Ecology.

Billick, I., et al. (2007). Ant-aphid interactions: are ants friends, enemies, or both? Annals of the Entomological Society of America.

Boxwell, (2025). The role of soil in regulating plant performance in Valeriana edulis.

Cushman, J. H., & Addicott, J. F. (1989). Intra- and interspecific competition for mutualists: ants as a limited and limiting resource for aphids. Oecologia.

Davis, (2023). Elevation does not predict density dependent population dynamics in Valeriana edulis.

Dean, et al. (2024). Decomposing an elevational gradient in predation by insectivorous birds. Ecosphere.

Fraser, A. M., et al. (2001). Assessing the quality of different ant species as partners of a myrmecophilous butterfly. Oecologia.

Grinath, J. (2021). Chronic, low-level nitrogen deposition enhances abundances of ant-protected herbivores inhabiting an imperiled foundation species. Acta Oecologica.

Hunter, (2023). Breakfast of champions: spatiotemporal variation in the quality of ant nests for bear consumers.

Kennamer, (2022). Ant behavioral responses to aphids colonizing Ligusticum porteri.

Langenheim, J. H. (1962). Vegetation and environmental patterns in the Crested Butte area, Gunnison County, Colorado. Ecological Monographs.

Mooney, et al. (2023). Host plant phenology shapes aphid abundance and interactions with ants. Oikos.

Nelson, A. S., & Mooney, K. A. (2022). The evolution and ecology of interactions between ants and honeydew-producing hemipteran insects. Annual Review of Ecology, Evolution, and Systematics.

Nelson, A. S., & Mooney, K. A. (2025). Different aspects of dominance are not equivalent when testing for trade-offs in ant communities. Ecology and Evolution.

Petry, W. K., et al. (2016). Sex-specific responses to climate change in plants alter population sex ratios and performance. Science.

Pierce, N. E., & Easteal, S. (1986). The selective advantage of attendant ants for the larvae of a lycaenid butterfly, Glaucopsyche lygdamus. Journal of Animal Ecology.

Pierce, N. E., & Mead, P. S. (1981). Parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae and ants. Science.

Price, M. V., et al. (2014). Human activity affects the perception of risk by mule deer. Current Zoology.

Rittler, (2024). Associations between deer browse and aphid colonization in a long-term monitoring study of Ligusticum porteri.

Uetrecht, et al. (2023). Differential response of three large mammal species to human recreation in the Rocky Mountains of Colorado, USA. Frontiers in Conservation Science.

Williams, C. F., et al. (2000). Floral dimorphism, pollination, and self-fertilization in gynodioecious Geranium richardsonii. American Journal of Botany.

Concept (42) →

mutualistic relationships

Mutually beneficial herbivore-predator associations where sap-feeding insects provide honeydew food for ants in exchange for protection against predat...

processcommunity ecology135 papers

trophic cascade

Indirect effects of predators on lower trophic levels mediated through changes in intermediate consumer behavior or abundance

processcommunity ecology95 papers

facilitative interactions

Positive species interactions such as nurse plant effects by cushion-forming species that increase reproductive success for neighboring plants

processcommunity ecology95 papers

trophic synchrony

Temporal alignment between consumer energy requirements and food source availability across trophic levels

phenomenoncommunity ecology32 papers

recreational trail impacts

Effects of human recreational activities on wildlife communities through habitat alteration, disturbance, and behavioral changes

phenomenoncommunity ecology27 papers

honeydew composition

Sugar composition of plant phloem sap which honeydew is derived from, potentially altered by temperature affecting ant recruitment or tending behavior

measurementcommunity ecology16 papers

biomass allocation

Distribution of plant growth and resources between aboveground (shoots) and belowground (roots) structures

processgeneral ecology15 papers

tripartite interactions

Interactions involving three species or groups, such as soil microbes, plants, and pollinators

processcommunity ecology15 papers

herbivore preference

Selective feeding behavior of herbivores that favors certain plant traits or genotypes over others

processcommunity ecology14 papers

associative learning

The formation of predictive relationships between contingent stimuli in the environment

processgeneral ecology13 papers
Show 32 more concepts

trophic-level sensitivity

The concept that higher trophic levels are more sensitive to environmental change due to their smaller population sizes and greater environmental dema...

hypothesiscommunity ecology11 papers

extrafloral nectaries

Nectar-secreting organs located on leaf laminae, petioles, rachis, bracts, stipules, pedicels, or fruit that attract arthropods

phenomenoncommunity ecology11 papers

factorial experiment

Experimental design testing multiple factors simultaneously in all combinations

frameworkmethodological8 papers

sexual dimorphism in plant-insect interactions

How male and female plants of dioecious species differ in their interactions with insects

phenomenoncommunity ecology7 papers

dominance-discovery trade-off

The hypothesis that species that are competitively dominant have reduced ability to discover new resources quickly

hypothesiscommunity ecology7 papers

chemical cues

Chemical signals that organisms use to obtain information about their environment

phenomenoncommunity ecology7 papers

ecological dominance

Having the greatest foraging success relative to abundance in the environment

measurementcommunity ecology6 papers

predator exclusion

Experimental technique to prevent predator access while maintaining other ecological interactions

methodologymethodological5 papers

non-invasive sampling

Sampling methods that do not require capturing or handling animals, such as hair tube traps that collect hair samples

frameworkmethodological5 papers

honeydew production

The excretion of sugar-rich liquid waste by herbivorous hemipterans feeding on plant sap

processcommunity ecology5 papers

temperature buffering

When clustered plants experience facilitative interactions that cause leaves to be cooler relative to surrounding soils

processclimate5 papers

small mammal abundance

Relative number of small mammals detected through camera trap monitoring expressed as activity indices

measurementpopulation ecology4 papers

relative growth rate

Aphid colony growth rate calculated as ln(n1)-ln(n0)/t where n1 and n0 are final and initial colony sizes

metricpopulation ecology4 papers

variance partitioning

Method for quantifying the proportion of overall variance explained by different factors or pathways using R² calculations

measurementmethodological4 papers

cost-benefit ratio

The balance between costs and benefits that determines whether species interactions are mutualistic or antagonistic

frameworkcommunity ecology4 papers

morphotype classification

Grouping of individuals based on body shape rather than species-level identification due to field identification constraints

methodologymethodological4 papers

aridity gradient

Environmental gradient characterized by declining water availability, derived from temperature and precipitation variables

measurementclimate4 papers

abiotic factors

Non-living environmental factors such as climate, light, and temperature that influence organism performance

phenomenongeneral ecology4 papers

resource density

The abundance or availability of resources in a given area

measurementpopulation ecology3 papers

pitfall trapping

Sampling method using buried containers with killing solution to capture ground-dwelling invertebrates

measurementmethodological3 papers

multicollinearity

Statistical phenomenon where predictor variables are correlated, potentially masking individual effects in regression analysis

methodologicalmethodological3 papers

trophallaxis

The sharing of liquid food resources between nestmates through specialized anatomical and behavioral adaptations

processcommunity ecology3 papers

operational sex ratio

Proportion of flowering individuals that are primarily male-expressing

metricpopulation ecology3 papers

spatial clustering

When plants are found adjacent to one another with less than two finger-widths between them or when one plant surrounds another

phenomenoncommunity ecology3 papers

distance gradient

Spatial sampling design measuring wildlife responses at increasing distances from disturbance sources

frameworklandscape2 papers

ungulate-aspen interactions

Ecological interactions between hoofed mammals and aspen trees including herbivory effects on tree health and forest composition

processcommunity ecology2 papers

foundation species

Species that define ecosystems and provide critical habitat structure, where big sagebrush is a foundation species in steppe ecosystems of western Nor...

theorycommunity ecology2 papers

life cycle

The complete sequence of developmental stages from germination through survival, growth, flowering, and reproduction in sessile plants

frameworkpopulation ecology2 papers

F-score

Harmonic mean of precision and recall used to evaluate classification model performance

metricmethodological2 papers

detection probability

The probability of detecting a species given that it is present at a site

measurementpopulation ecology2 papers

dominance-generalism trade-off

Predicts that dominant species may be more specialized on particular resources than subordinates, allowing subordinate species to coexist by better ca...

hypothesiscommunity ecology2 papers

parasitoid behavior

Behavioral patterns of wasps that parasitize other arthropods, particularly aphids

processcommunity ecology2 papers

Protocol (30) →

Experimental predator manipulation with colony establishment

Controlled experiment manipulating predator presence by establishing artificial aphid colonies, protecting them from natural predators, then adding ly...

experimentalstandardized94 papers

pitfall trapping (Formicidae)

Standardized 2-hour baiting experiments using 5 different resource types arranged in pentagonal plots to sample ant communities and assess dominance p...

samplingstandardized72 papers

Ant exclusion experiment on aphid colonies

Experimental manipulation using physical barriers to exclude ants from aphid colonies while maintaining control colonies with ant access. Uses paired ...

experimentalstandardized34 papers

focal animal sampling

Standardized pellet count method for estimating deer density using systematic transect surveys with random placement. Identifies scat to species level...

sampling23 papers

Cariveau allometric conversion (Apidae)

Field capture of wild bumblebees followed by temporary immobilization through cold-induced chill coma to enable precise measurement of tongue length a...

measurementstandardized22 papers

Visual and camera trap survey of small mammals

Combined direct visual observation during peak activity periods with motion-activated camera bucket traps to determine species presence/absence in mea...

observational20 papers

focal animal sampling (Cervidae)

Controlled experiment using feeding stations paired with predator scent cues to test antipredator behavioral responses. Individual feeding times are r...

experimental19 papers

mark-recapture (Cricetidae)

Standard live-trapping protocol using Longworth traps in grid arrays for mark-recapture population estimation of small mammals with individual marking...

sampling17 papers

camera trap survey

Camera traps deployed along trails in a stratified random design to detect wildlife presence and estimate occupancy while accounting for detection pro...

sampling14 papers

jar method

Standardized wet sieving method using Yoder device to determine soil aggregate size distributions. Measures water-stable aggregates across multiple si...

measurementstandardized13 papers
Show 20 more protocols

Snow melt manipulation experiment (Aphididae)

Experimental advancement of snow melt timing using shade cloth application to test effects on host plant phenology and aphid-plant interactions. Combi...

experimentalstandardized11 papers

scat transect surveys (Cervidae)

Systematic collection and GPS mapping of predator scats along established trail networks to quantify spatial patterns of predator activity relative to...

sampling11 papers

Honeydew collection and chemical analysis

Collection of aphid honeydew on aluminum foil substrates followed by LC-MS/MS analysis to quantify sugar and amino acid composition as indicators of p...

analyticalstandardized10 papers

GIS-based meadow habitat mapping for ground squirrels

Used raster analysis in ArcGIS to identify suitable ground squirrel habitat by combining landcover and canopy height data, then calculated distance-fr...

computational9 papers

Ant chemical associative learning field assay (Formicidae)

A two-phase field protocol using training baits scented with plant chemicals followed by two-choice tests to assess whether ants form associative memo...

experimental9 papers

quadrat sampling (Asteraceae)

Plant cover measured using 1 m² quadrats divided into 1 dm² cells, counting filled cells to nearest 0.25 dm². Height measured to maximum plant extensi...

measurementstandardized9 papers

Plant biomass allocation analysis (Valerianaceae)

Calculation and statistical analysis of plant biomass allocation between vegetative and reproductive tissues using arcsin transformations and three-wa...

analytical7 papers

Colony fragmentation and diet manipulation of Formica podzolica (Formicidae)

Wild ant colonies were collected and split into paired fragments, then maintained on controlled diets (carbohydrate vs protein-rich) to test effects o...

experimental6 papers

stable isotope analysis

Carbon and nitrogen stable isotope analysis of ant tissues to determine trophic position and dietary composition. Involves specimen preparation, dryin...

analytical5 papers

Gravimetric soil moisture determination

Standardized method using funnel, filter paper and drainage to measure maximum water holding capacity of soil samples. Involves saturating soil, drain...

laboratory5 papers

Multi-year arthropod community surveys on dioecious plants

Exhaustive visual surveys of arthropod abundance on randomly selected male and female plants conducted across multiple years to quantify sex-biased co...

sampling5 papers

Ant identification and behavioral observation on sunflowers (Asteraceae)

Field-based study of ant interactions with pollinators on Helianthella quinquenervis, involving ant identification and behavioral observations. Protoc...

observational4 papers

Dominance scoring (Formicidae)

Three dominance scores calculated for each species at each site where it occurred to assess competitive hierarchies.

analytical3 papers

eBird citizen science protocol

Use existing citizen science bird observation data from eBird database to characterize bird abundance and diversity patterns across environmental grad...

observational3 papers

nonmetric multidimensional scaling (Plantae)

PERMANOVA testing of volatile composition differences across species and years, with NMDS visualization of chemical dissimilarity patterns.

analytical3 papers

Trophic interaction field observation

Systematic field observation of trophic interactions between organisms, primarily plants and insects, with rotating temporal sampling across multiple ...

observational3 papers

clay caterpillar predation bioassay

Artificial prey made from green modeling clay are deployed on tree branches to estimate bird attack rates over a standardized time period. Attack mark...

experimental3 papers

Climate wetness index calculation

Calculation of site-specific wetness indices using mean annual precipitation and temperature data to characterize local climate conditions for experim...

computational2 papers

AIC model selection (Plantae)

Multiple candidate models incorporating different combinations of climate predictors were fitted and ranked using Akaike Information Criterion to sele...

analytical2 papers

Sagebrush arthropod visual encounter surveys

Visual counting method for arthropods on sagebrush plants involving systematic counting of mobile species first (to avoid double-counting) followed by...

samplingstandardized2 papers

Publication (114) →

Show 104 more publications

Coyotes, deer, and wildflowers: diverse evidence points to a trophic cascade

2014Naturwissenschaftenarticle

Elevational cline in herbivore abundance driven by a monotonic increase in trophic level sensitivity to aridity

2019Journal of Animal Ecologyarticle

Mechanisms underlying plant sexual dimorphism in multi-trophic arthropod communities

2013Ecologyarticle

Influence of macronutrient imbalance on native ant foraging and interspecific interactions in the field

2012Ecological Entomologyarticle

Elevated temperatures alter an ant aphid mutualism

2019Entomologia Experimentalis et Applicataarticle

Host plant phenology shapes aphid abundance and interactions with ants

2023Oikosarticle

Are ants botanists? Ant associative learning of plant chemicals mediates foraging for carbohydrates

2020Ecological Entomologyarticle

Bottom-up mediation of an ant-membracid mutualism: effects from different host plants

2006Evolutionary Ecologyarticle

Effects of early snowmelt and climate warming on Valeriana edulis and the insects that depend on it.

2014student paper

The Effect of Climate Change on Plant Communities in the Rocky Mountains: How floral traits differ along an elevational gradient and in reciprocally transplanted communities

2018student paper

A balanced diet: Effects of ant (Hymenoptera: Formicidae) nutritional state on the balance between mutualism and predation upon aphids (Hemiptera: Aphididae)

2009student paper

The Evolution and Ecology of Interactions Between Ants and Honeydew-Producing Hemipteran Insects

2022The Annual Review of Ecology, Evolution, and Systematicsarticle

The effect of an introduced predator scent on mule deer (</i>Odocoileus hemionus</i>) browsing activities in meadow habitats in Gothic, Colorado

2010student paper

Nowcasting the distribution of <i> Valeriana edulis </i> using climate driven population models

2021student paper

Protection at a price? Ant interactions with pollinators on aspen sunflower (<i>Helianthella quinquenervis</i>)

2009student paper

Insect herbivore stoichiometry: the relative importance of host plants and ant mutualists

2008Ecological Entomologyarticle

Differential response of three large mammal species to human recreation in the Rocky Mountains of Colorado, USA

2023Frontiers in Conservation Sciencearticle

Decomposing an elevational gradient in predation by insectivorous birds

2024Ecospherearticle

Ant-aphid interactions: are ants friends, enemies, or both?

2007Annals of the Entomological Society of Americaarticle

Chronic, low-level nitrogen deposition enhances abundances of ant-protected herbivores inhabiting an imperiled foundation species

2021Acta Oecologicaarticle

Different aspects of dominance are not equivalent when testing for trade-offs in ant communities

2025Ecology and Evolutionarticle

Role of floral nectaries and plant sex in mediating ant-aphid interactions on <i>Valeriana edulis</i>

2011student paper

The role of soil in regulating plant performance in Valeriana edulis

2025student paper

Ant Behavioral Responses to Aphids Colonizing <i> Ligusticum porteri </i>

2022student paper

The Landscape of Fear and Trophic Cascades: Does Human Presence at RMBL Affect Deer Behavior?

2010student paper

Effects of phenological stage and temperature on <i>Ligusticum porteri’s</i> volatiles and trophic interactions

2016student paper

The effects of recreational trail use on small mammal species richness and abundance

2020student paper

Spatial distribution of mule deer (<i>Odocoileus hemionus</i>): Effects of plant palatability and habitat structure

2011student paper

Variation of wasp behavior patterns and pollination behavior on Ligusticum porteri

2024student paper

Species Interactions in Arthropod Communities: Density Dependence and Ant Interactions on Aphid Per Capita Population Growth

2010student paper

The Effect of Ant Colony Proximity on Membracid Survivorship

2005student paper

The relative importance of spatial vs. temporal variability in generating a conditional mutalism

2003Ecologyarticle

Testing trade-offs and the dominance–impoverishment rule among ant communities

2020Journal of Biogeographyarticle

The effect of hermaphrodite density and frequency at three spatial scales on the polen receipt and seed set of gynodioecious Geranium richardsonii (Geraniaceae)

2001student paper

Assessing the quality of different ant species as partners of a myrmecophilous butterfly

2001Oecologiaarticle

Variation in host plant sex mediates ant-aphid interactions

2009student paper

Short-term, low-level nitrogen deposition dampens a trophic cascade between bears and plants

2018Ecology and Evolutionarticle

Associations Between Deer Browse and Aphid Colonization in a Long-Term Monitoring Study of Liguisticum porteri

2024student paper

Niche relationships among species of aphids feeding on fireweed

1978Canadian Journal of Zoologyarticle

Plant sex and induced responses independently influence herbivore performance, natural enemies and aphid-tending ants

2012Arthropod-Plant Interactionsarticle

Genetic variation in plant functional traits as drivers in arthropod community structure

2013student paper

Impacts of Mule Deer Herbivory on Herbaceous Vegetation in the Gothic Area, with Focus on Aquilegia coerulea

2005student paper

What is the nature of the ant-aphid relationship?

2004student paper

Global Climate Change-Induced Mutualism Breakdown Among Legumes and their Soil Synbionts

2019student paper

A multispecies aphid-ant association: density dependence and species-specific effects

1979Canadian Journal of Zoologyarticle

The timing of the ant-effect on nymph size and survivorship in an ant-treehopper mutualism

2006student paper

Variation in interaction zone size and influence of sex on fitness components of Valeriana edulis

2024student paper

Abiotic and multitrophic determinants of geographic distribution in an herbivorous insect

2013student paper

Human activity affects the perception of risk by mule deer

2014Current Zoologyarticle

The effect of ants on membracid nymph size and instar

2007student paper

Indirect effect of black bears on sunflowers in nitrogen-polluted and pristine steppe

2024student paper

Elevation Does Not Predict Density Dependent Population Dynamics in Valeriana edulis

2023student paper

The Effect of Predation on Ant-aphid Mutualism in <i>Ligusticum porteri</i>

2019student paper

Plant chemical mediation of ant behavior

2019Current Opinion in Insect Sciencearticle

Effects of recreational trails on small mammal communities in north-central Gunnison County, Colorado

2019student paper

Chemotypic variation in oshá (<i>Ligusticum porteri</i>) in Colorado, USA

2018J Appl. Res. Med. Aromat. Plantsarticle

Expansion of herbaria data based on historically surveyed herbaceous plants in the Crested Butte area, Colorado.

2014student paper

Sex ratio and reproductive success along elevational gradients of gynodioecious populations of <i>Geranium richardsonii</i>

2016student paper

The influence of human recreational trail use has on rodent and predator activity using motion triggered cameras

2021student paper

Assessing the forecastability & forecast skill of models to predict sex ratios of Valeriana edulis

2020student paper

Are ants botanists?: Ant associative learning of plant volatiles

2018student paper

Tests for Elevational Gradients in Herbivore Abundance and Plant Resistance in the Rocky Mountain Ecosystem

2021student paper

Pollination Preferences of <i> Geranium richardsonii </i> Between Bee and Fly Species at High and Low Elevations

2021student paper

A test of sexual dimorphism in <i>Valeriana edulis</i> resistance and induced responses to herbivory

2010student paper

Antipredatory response of mule deer (<i>Odocoileus hemionus</i>) to predator urines

2005student paper

Effect of location on plant species richness and diversity in Aspen (<i>Populus tremuloides</i>) understory: edge vs. inner forest habitat

2013student paper

The effect of ant tending on the fitness of aphids during and after colony establishment

2011student paper

Plant resource allocation and herbivory for <i>Helianthella quinquenervis</i> (Asteraceae) over an elevational gradient

2001student paper

Impact of the Western Thatching Ant (Formica obscuripes) on

2004URBEEarticle

Development of Keras image classification model for use with a study on the effects of recreational trail use on small mammal species richness and activity

2021student paper

Seed protection by ants foraging on the extrafloral nectaries of the aspen sunflower, <i>Helianthella quinquenervis</i>

1989student paper

Using GIS techniques to test a model of the coexistence of the golden-mantled ground squirrel, Callospermopholis lateralis, and the least chipmunk, Tamias minimus

2023student paper

Reconstruction and spatial analysis of alpine treeline in the Elk Mountains, Colorado, USA

2016Physical Geographyarticle

Effects of avalanches on rodent populations

2019student paper

The ability of ants to associatively learn based on olfactory chemical cues produced by plants.

2017student paper

Self-Similarity in the Distribution of Plant Species Across a Successional Gradient

2002student paper

Does <i>Aphis asclepiadis</i> colony size mediate <i>Formica rufa</i> and <i>Tapinoma sessile</i> competition for mutualist aphids

2019student paper

The Combined Effects of Aphid-Ant Mutualism and Fertilizer on the Resource Allocation of Valerian edulis

2002student paper

What's for lunch: deciphering ant omnivory on lupine

2010student paper

Comparing predictive measures and model functions for estimating plant biomass: lessons from a sagebrush–rabbitbrush community

2019Plant Ecologyarticle

Floral dimorphism, pollination, and self-fertilization in gynodioecious <i>Geranium richardsonii</i> (Geraniaceae)

2000American Journal of Botanyarticle

Dynamics of male inconstancy in <i> Valeriana edulis </i> in the abiotic and mating environment

2022student paper

Comparing Longworth live traps to hair tubes for describing small mammal communities

2018student paper

Does ant usage vary between the plants <i>Ligusticum porteri</i> and <i>Helianthella quinquenervis</i>?

2013student paper

Examining top-down and bottom up effects on aphid abundance on Ligusticum porteri

2012student paper

Benefits of ant attendance for aphid colonies of varying density

2013student paper

Effects of quantity and distribution of pollen on fertilization in the gynodioecious species <i>Geranium richardsonii</i>

2009student paper

Impact of mound-building ants on ecosystem properties create islands of fertility in alpine meadows

2011student paper

Mule deer (<i>Odocoileus hemionus</i>) detect coyote (<i>Canis latrans</i>) scent

2011student paper

Intra- and interspecific competition for mutalists: ants as a limited and limiting resource for aphids

1989Oecologiaarticle

Chemical camouflage and the consequences of changing host plants in a treehopper-ant mutualism

2013student paper

Causes of predation intensity in an ant/aphid mutualist system

2011student paper

Aboveground productivity and floristic structure of a high subalpine herbaceous meadow

1979Arctic and Alpine Researcharticle

Effects of aggregation size and host plant on the survival of an ant-tended Membracid (Hemiptera: Membracidae): potential roles in selecting for generalized host plant use

2008Annals of the Entomological Society of Americaarticle

The effect of ants on the population dynamics of a protective symbiont of aphids, <i>Hamiltonella defensa</i>

2012Annals of Entomological Society of Americaarticle

The relationship between ant-tending and maternal care in the treehopper Publilia modesta

2001Behavioral Ecology and Sociobiologyarticle

Competition for mutualists: aphids and ants

1978Canadian Journal of Zoologyarticle

Breakfast of champions: Spatiotemporal variation in the quality of ant nests for bear consumers

2023student paper

Dichotomous key of the onagraceae family

2009student paper

The amino acids of extrafloral nectar from Helianthella quinquenervis (Asteraceae)

1980American Journal of Botanyarticle

Spatial segregation of the sexes of dioecious plants

1988American Naturalistarticle

Parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae and ants

1981Sciencearticle

The selective advantage of attendant ants for the larvae of a lycanaenid butterfly, Glaucopsyche lygdamus

1986Journal of Animal Ecologyarticle

A modified clip cage for use with aphids and other small insects

1981Canadian Entomologistarticle

Dataset (11) →

Data from: Progressive sensitivity of trophic levels to warming underlies an elevational gradient in ant-aphid mutualism strength

Although species interactions are often proposed to be stronger at lower latitudes and elevations, few studies have evaluated the mechanisms driving s...

other2018

Data from: Testing trade-offs and the dominance-impoverishment rule among ant communities

Aim: Ant communities are believed to be structured by competition, with dominant species competitively excluding subordinates (the dominance-impoveris...

other2021

Data from: Elevational cline in herbivore abundance driven by a monotonic increase in trophic level sensitivity to aridity

1. The abiotic environment drives species abundances and distributions both directly and indirectly through effects on multi-trophic species interacti...

other2019

Decomposing an elevational gradient in predation by insectivorous birds

Insectivorous birds have ecologically important effects on prey abundance, behavior, and evolution, and through top-down control birds indirectly redu...

other2024

Multi-year census of arthropod abundance on the plant Ligusticum porteri near Crested Butte, CO

The purpose of this study was to track year-to-year variation in aphid abundance on the host plant Ligusticum porteri (Apiaceae). We censused arthropo...

other2021

Plant phenology, aphid colony growth, and honeydew deposition data

Changing phenological cues can lead to trophic mismatch for plants and herbivores, and this often shifts herbivore feeding to plant stages of lower ...

other2023

Multi-year census of arthropod abundance on the plant Ligusticum porteri near Gothic, CO

The purpose of this study was to track year-to-year variation in aphid abundance on the host plant Ligusticum porteri (Apiaceae). We censused arthropo...

other2022

Data from: Short-term, low-level nitrogen deposition dampens a trophic cascade between bears and plants

Human activities have substantially increased atmospheric nitrogen (N) deposition in ecosystems worldwide, often leading to higher plant quality for h...

other2018

Data from: Different aspects of dominance are not equivalent when testing for trade-offs in ant communities

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2025

Arthropod abundance censused on the host plant Ligusticum porteri near Gothic, CO.

The objective of this study is to understand how climate cues affect the abundance and phenology of aphids and the arthropods with which they interact...

other2021
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