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  1. Abstract

    The encroachment of woody plants into grasslands is a global phenomenon with implications for biodiversity and ecosystem function. Understanding and predicting the pace of expansion and the underlying processes that control it are key challenges in the study and management of woody encroachment. Theory from spatial population biology predicts that the occurrence and speed of expansion should depend sensitively on the nature of conspecific density dependence. If fitness is maximized at the low‐density encroachment edge, then shrub expansion should be “pulled” forward. However, encroaching shrubs have been shown to exhibit positive feedbacks, whereby shrub establishment modifies the environment in ways that facilitate further shrub recruitment and survival. In this case there may be a fitness cost to shrubs at low density causing expansion to be “pushed” from behind the leading edge. We studied the spatial dynamics of creosotebush (Larrea tridentata), which has a history of encroachment into Chihuahuan Desert grasslands over the past century. We used demographic data from observational censuses and seedling transplant experiments to test the strength and direction of density dependence in shrub fitness along a gradient of shrub density at the grass–shrub ecotone. We also used seed‐drop experiments and wind data to construct a mechanistic seed‐dispersal kernel, then connected demography and dispersal data within a spatial integral projection model (SIPM) to predict the dynamics of shrub expansion. Contrary to expectations based on potential for positive feedbacks, the shrub encroachment wave is “pulled” by maximum fitness at the low‐density front. However, the predicted pace of expansion was strikingly slow (ca. 8 cm/year), and this prediction was supported by independent resurveys of the ecotone showing little to no change in the spatial extent of shrub cover over 12 years. Encroachment speed was acutely sensitive to seedling recruitment, suggesting that this population may be primed for pulses of expansion under conditions that are favorable for recruitment. Our integration of observations, experiments, and modeling reveals not only that this ecotone is effectively stalled under current conditions but also why that is so and how that may change as the environment changes.

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  2. Abstract

    Understanding mechanisms that generate range limits is central to knowing why species are found where they are and how they will respond to environmental change. There is growing awareness that biotic interactions play an important role in generating range limits. However, current theory and data overwhelmingly focus on abiotic drivers and antagonistic interactions. Here we explore the effect that mutualists have on their partner's range limits: the geographic “footprint” of mutualism. This footprint arises from two general processes: modification of a partner's niche through environment‐dependent fitness effects and, for a subset of mutualisms, dispersal opportunities that lead suitable habitats to be filled. We developed a conceptual framework that organizes different footprints of mutualism and the underlying mechanisms that shape them, and evaluated supporting empirical evidence from the primary literature. In the available literature, we found that the fitness benefits and dispersal opportunities provided by mutualism can extend species' ranges; conversely, the absence of mutualism can constrain species from otherwise suitable regions of their range. Most studies found that the footprint of mutualism is driven by changes in the frequency of mutualist partners from range core to range edge, whereas fewer found changes in interaction outcomes, the diversity of partners, or varying sensitivities of fitness to the effects of mutualists. We discuss these findings with respect to specialization, dependence, and intimacy of mutualism. Much remains unknown about the geographic footprint of mutualisms, leaving fruitful areas for future work. A particularly important future direction is to explore the role of mutualism during range shifts under global change, including the promotion of shifts at leading edges and persistence at trailing edges.

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  3. Abstract

    Mutualism benefits partner species, and theory predicts these partnerships can affect the abundance, diversity, and composition of partner and non‐partner species. We used 16 years of monitoring data to determine the ant partner species of tree cholla cacti (Cylindropuntia imbricata), which reward ants with extrafloral nectar in exchange for anti‐herbivore defense. These long‐term data revealed one dominant ant partner (Liometopum apiculatum) and two less common partners (Crematogaster opuntiaeandForelius pruinosus). We then used short‐term characterization of the terrestrial ant community by pitfall trapping to sample partner and non‐partner ant species across ten plots of varying cactus density. We found that the dominant ant partner tended a higher proportion cacti in plots of higher cactus density, and was also found at higher occurrence within the pitfall traps in higher density plots, suggesting a strong positive feedback that promotes ant partner occurrence where plant partners are available. Despite the strong association and increased partner occurrence, ant community‐wide effects from this mutualism appear limited. Of the common ant species, the occurrence of a single non‐partner ant species was negatively associated with cactus density and with the increased presence ofL. apiculatum. Additionally, the composition and diversity of the ant community in our plots were insensitive to cactus density variation, indicating that positive effects of the mutualism on the dominant ant partner did not have cascading impacts on the ant community. This study provides novel evidence that exclusive mutualisms, even those with a strong positive feedback, may be limited in the scope of their community‐level effects.

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  4. Abstract Background and Aims The processes that maintain variation in the prevalence of symbioses within host populations are not well understood. While the fitness benefits of symbiosis have clearly been shown to drive changes in symbiont prevalence, the rate of transmission has been less well studied. Many grasses host symbiotic fungi (Epichloë spp.), which can be transmitted vertically to seeds or horizontally via spores. These symbionts may protect plants against herbivores by producing alkaloids or by increasing tolerance to damage. Therefore, herbivory may be a key ecological factor that alters symbiont prevalence within host populations by affecting either symbiont benefits to host fitness or the symbiont transmission rate. Here, we addressed the following questions: Does symbiont presence modulate plant tolerance to herbivory? Does folivory increase symbiont vertical transmission to seeds or hyphal density in seedlings? Do plants with symbiont horizontal transmission have lower rates of vertical transmission than plants lacking horizontal transmission? Methods We studied the grass Poa autumnalis and its symbiotic fungi in the genus Epichloë. We measured plant fitness (survival, growth, reproduction) and symbiont transmission to seeds following simulated folivory in a 3-year common garden experiment and surveyed natural populations that varied in mode of symbiont transmission. Key Results Poa autumnalis hosted two Epichloë taxa, an undescribed vertically transmitted Epichloë sp. PauTG-1 and E. typhina subsp. poae with both vertical and horizontal transmission. Simulated folivory reduced plant survival, but endophyte presence increased tolerance to damage and boosted fitness. Folivory increased vertical transmission and hyphal density within seedlings, suggesting induced protection for progeny of damaged plants. Across natural populations, the prevalence of vertical transmission did not correlate with symbiont prevalence or differ with mode of transmission. Conclusions Herbivory not only mediated the reproductive fitness benefits of symbiosis, but also promoted symbiosis prevalence by increasing vertical transmission of the fungus to the next generation. Our results reveal a new mechanism by which herbivores could influence the prevalence of microbial symbionts in host populations. 
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  5. Abstract

    The effects of climate change on population viability reflect the net influence of potentially diverse responses of individual‐level demographic processes (growth, survival, regeneration) to multiple components of climate. Articulating climate–demography connections can facilitate forecasts of responses to future climate change as well as back‐casts that may reveal how populations responded to historical climate change.

    We studied climate–demography relationships in the cactusCyclindriopuntia imbricata; previous work indicated that our focal population has high abundance but a negative population growth rate, where deaths exceed births, suggesting that it persists under extinction debt. We parameterized a climate‐dependent integral projection model with data from a 14‐year field study, then back‐casted expected population growth rates since 1900 to test the hypothesis that recent climate change has driven this population into extinction debt.

    We found clear patterns of climate change in our central New Mexico study region but, contrary to our hypothesis,C. imbricatahas most likely benefitted from recent climate change and is on track to reach replacement‐level population growth within 37 years, or sooner if climate change accelerates. Furthermore, the strongest feature of climate change (a trend towards years that are overall warmer and drier, captured by the first principal component of inter‐annual variation) was not the main driver of population responses. Instead, temporal trends in population growth were dominated by more subtle, seasonal climatic factors with relatively weak signals of recent change (wetter and milder cool seasons, captured by the second and third principal components).

    Synthesis. Our results highlight the challenges of back‐casting or forecasting population dynamics under climate change, since the most apparent features of climate change may not be the most important drivers of ecological responses. Environmentally explicit demographic models can help meet this challenge, but they must consider the magnitudes of different aspects of climate change alongside the magnitudes of demographic responses to those changes.

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  6. Abstract

    Although rarely experimentally tested, biotic interactions have long been hypothesised to limit low‐elevation range boundaries of species. We tested the effects of herbivory on three alpine‐restricted plant species by transplanting plants below (novel), at the edge (limit), or in the centre (core) of their current elevational range and factorially fencing‐out above‐ and belowground mammals. Herbivore damage was greater in range limit and novel habitats than in range cores. Exclosures increased plant biomass and reproduction more in novel habitats than in range cores, suggesting demographic costs of novel interactions with herbivores. We then used demographic models to project population growth rates, which increased 5–20% more under herbivore exclosure at range limit and novel sites than in core habitats. Our results identify mammalian herbivores as key drivers of the low‐elevation range limits of alpine plants and indicate that upward encroachment of herbivores could trigger local extinctions by depressing plant population growth.

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