Developing common protocols to measure tundra herbivory across spatial scales

Understanding and predicting large-scale ecological responses to global environmental change requires comparative studies across geographic scales with coordinated efforts and standardized methodologies. We designed, applied, and assessed standardized protocols to measure tundra herbivory at three spatial scales: plot, site (habitat), and study area (landscape). The plot- and site-level protocols were tested in the field during summers 2014–2015 at 11 sites, nine of them consisting of warming experimental plots included in the International Tundra Experiment (ITEX). The study area protocols were assessed during 2014–2018 at 24 study areas across the Arctic. Our protocols provide comparable and easy to implement methods for assessing the intensity of invertebrate herbivory within ITEX plots and for characterizing vertebrate herbivore communities at larger spatial scales. We discuss methodological constraints and make recommendations for how these protocols can be used and how sampling effort can be optimized to obtain comparable estimates of herbivory, both at ITEX sites and at large landscape scales. The application of these protocols across the tundra biome will allow characterizing and comparing herbivore communities across tundra sites and at ecologically relevant spatial scales, providing an important step towards a better understanding of tundra ecosystem responses to large-scale environmental change

Location of studies and evidence of effects of herbivory on Arctic vegetation: a systematic map

Herbivores modify the structure and function of tundra ecosystems. Understanding their impacts is necessary to assess the responses of these ecosystems to ongoing environmental changes. However, the effects of herbivores on plants and ecosystem structure and function vary across the Arctic. Strong spatial variation in herbivore effects implies that the results of individual studies on herbivory depend on local conditions, i.e., their ecological context. An important first step in assessing whether generalizable conclusions can be produced is to identify the existing studies and assess how well they cover the underlying environmental conditions across the Arctic. This systematic map aims to identify the ecological contexts in which herbivore impacts on vegetation have been studied in the Arctic. Specifically, the primary question of the systematic map was: “What evidence exists on the effects of herbivores on Arctic vegetation?”

Developing common protocols to measure tundra herbivory across spatial scales

Understanding and predicting large-scale ecological responses to global environmental change requires comparative studies across geographic scales with coordinated efforts and standardized methodologies. We designed, applied and assessed standardized protocols to measure tundra herbivory at three spatial scales: plot, site (habitat), and study area (landscape). The plot and site-level protocols were tested in the field during summers 2014-2015 at eleven sites, nine of them comprising warming experimental plots included in the International Tundra Experiment (ITEX). The study area protocols were assessed during 2014-2018 at 24 study areas across the Arctic. Our protocols provide comparable and easy-to-implement methods for assessing the intensity of invertebrate herbivory within ITEX plots and for characterizing vertebrate herbivore communities at larger spatial scales. We discuss methodological constraints and make recommendations for how these protocols can be used and how sampling effort can be optimized to obtain comparable estimates of herbivory, both at ITEX sites and at large landscape scales. The application of these protocols across the tundra biome will allow characterizing and comparing herbivore communities across tundra sites and at ecologically relevant spatial scales, providing an important step towards a better understanding of tundra ecosystem responses to large-scale environmental change.CGB was funded by the Estonian Research Council (grant IUT 20-28), and the European Regional Development Fund (Centre of Excellence EcolChange). JDMS was supported by the Research Council of Norway (262064). OG and LB were supported by the French Polar Institute (program “1036 Interactions”) and PRC CNRS Russie 396 (program “ICCVAT”). DSH, NL, MAG, JB and JDR were supported by the Natural Sciences and Engineering Research Council (Canada). NL, MAG, JB and JDR were supported by the Polar Continental Shelf Program. NL was supported by the Canada Research Chair program and the Canada Foundation for Innovation. NL and JB were supported by Environment Canada and Polar Knowledge Canada. NL and MAG were supported by the Government of Nunavut, the Igloolik Community, and Université de Moncton. NL, MAG and JB were supported by the Northern Scientific Training Program. JMA was funded by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum (QUEX-CAS-QP-RD-18_19). IHM-S was funded by the UK Natural Environmental Research Council Shrub Tundra (NE/M016323/1) grant. ISJ was funded by the University of Iceland Research Fund. Fieldwork in Yamal peninsula (Erkuta, Sabetta and Belyi) for DE, NS and AS was supported by the Russian Foundation for Basic Research (No: 18-05-60261 and No: 18-54-15013), Fram Centre project YaES (No: 362259), the Russian Center of Development of the Arctic, and the “Yamal-LNG” company. Fieldwork in Utqiaġvik was supported by the U.S. Fish and Wildlife Service. Fieldwork in Svalbard was supported by the Norwegian Research Council (AFG No: 246080/E10), the Norwegian Polar Institute, Climate-ecological Observatory for Arctic Tundra – COAT, the Svalbard Environmental protection fund (project number 15/20), and the University Centre in Svalbard (UNIS) and the AB-338/AB-838 students of 2018. Sampling at Billefjorden was supported by GACR 17- 20839S

Changes in fin whale (Balaenoptera physalus) song over a forty-four year period in New England waters

Although much marine mammal research has been conducted on whales in the waters surrounding Cape Cod, Massachusetts, little work has been done on the supposed resident fin whale (Balaenoptera physalus) population that resides in this area. Fin whales have three different sounds in their vocal repertoire but a relatively simple song that is composed of long trains of high intensity 20-Hz pulses. Performed only by males, the song may be a reproductive display with song characteristics evolutionarily fixed within populations. In this study, fin whale vocalizations recorded in the area of Cape Cod Bay during the years 1961, 1978, 2001, and 2005 were characterized and compared within years to detect possible divergence in the male song. Reliable comparisons with the earlier data proved difficult because of small sample sizes under the chosen song parameters. However, measurements of inter-pulse interval and song length did vary significantly between the short time span of 2001 to 2005. Mean inter-pulse intervals were shorter in 2001, and song duration was longer within the 2005 recordings. Negative correlations between the center frequency and the song duration, as well as the center frequency and inter-pulse interval were also found. Although we rejected the null hypothesis that song notes and features are fixed and do not change over time, we did discover valuable information regarding the relationships between different features of fin whale song. Based on comparison of the temporal and individual variation observed in our data with those collected on a population in Bermuda, it is unclear whether vocal characteristics are distinct and could be an identifying feature of separate breeding populations

The Changing Structure and Function of Arthropod Food Webs in a Warming Arctic

<p>Environmental changes, such as climate change, can have differential effects on species, with important consequences for community structure and ultimately, for ecosystem functioning. In the Arctic, where ecosystems are experiencing warming at twice the rate as elsewhere, these effects are expected to be particularly strong. A proper characterization of the link between warming and biotic interactions in these particular communities is of global importance because the tundra's permafrost stores a vast amount of carbon that could be released through decomposition as greenhouse gases and alter the global rate of climate change. In this dissertation, I examine how arthropod communities are responding to warming in the Arctic and how these responses might be affecting ecosystem functioning. </p><p>I first address the question of whether and how long-term changes in climate are affecting individual groups and overall community structure in a high-arctic arthropod food web. I find that increasingly warm springs and summers between 1996-2011 differentially affected some arthropod groups and that this led to major changes in the relative abundances of different trophic groups within the arthropod community. Specifically, spring and summer warming are associated with relatively more herbivores and parasitoids and fewer detritivores within the community. These changes are particularly pronounced in heath sites, suggesting that arthropod communities in dry habitats are more responsive to climate change than those in wet habitats. I also show that herbivores and parasitoids are sensitive to conditions at subzero temperatures, even during periods of diapause, and that all trophic groups benefit from a longer transition period between summer and winter. These results suggest that the projected winter and springtime warming in Greenland may have unexpected consequences for northern arthropod communities. Moreover, the relative increase in herbivores and loss of detritivores may be changing the influence of the arthropod community over key ecosystem processes such as decomposition, nutrient cycling, and primary productivity in the tundra. </p><p>Predator-induced trophic cascades have been shown to impact both community structure and ecosystem processes, yet it is unclear how climate change may exacerbate or dampen predator effects on ecosystems. In the second chapter of my dissertation, I investigate the role of one of the dominant tundra predators within the arctic ecosystem, wolf spiders, and how their impact might be changing with warming. Using results from a two-year-long field experiment, I test the influence of wolf spider density over the structure of soil microarthropod communities and decomposition rates under both ambient and artificially warmed temperatures. I find that predator effects on soil microarthropods change in response to warming and that these changes translate into context-specific indirect effects of predators on decomposition. Specifically, while high densities of wolf spiders lead to faster decomposition rates at ambient temperatures, they are associated with slower decomposition rates in experimentally warmed plots. My results suggest that if warming causes an increase in arctic wolf spider densities, these spiders may buffer the rate at which the massive pool of stored carbon is lost from the tundra. </p><p>Wolf spiders in the Arctic are expected to become larger with warming, but it is unclear how this change in body size will affect spider populations or the role of wolf spiders within arctic food webs. In the third chapter of my dissertation, I explore wolf spider population structure and juvenile recruitment at three sites of the Alaskan Arctic that naturally differ in mean spider body size. I find that there are fewer juveniles in sites where female body sizes are larger and that this pattern is likely driven by a size-related increase in the rate of intraspecific cannibalism. These findings suggest that across the tundra landscape, there is substantial variation in the population structure and trophic position of wolf spiders, which is driven by differences in female spider body sizes. </p><p>Overall, this dissertation demonstrates that arctic arthropod communities are changing as a result of warming. In the long-term, warming is causing a shift in arthropod community structure that is likely altering the functional role of these animals within the ecosystem. However even in the short-term, warming can alter species interactions and community structure, with important consequences for ecosystem function. Arthropods are not typically considered to be major players in arctic ecosystems, but I provide evidence that this assumption should be questioned. Considering that they are the largest source of animal biomass across much of the tundra, it is likely that their activities have important consequences for regional and global carbon dynamics.</p>Dissertatio

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

Author Posting. © The Author(s), 2017. This is the author's version of the work. It is posted here by permission of John Wiley & Sons for personal use, not for redistribution. The definitive version was published in Oikos 127 (2018): 460-471, doi:10.1111/oik.04398.A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.National Science Foundation Grants DEB 102683 to support the Arctic LTER to G. Shaver, OPP 0908502 to L. Gough, and DEB 1210704 to A. Koltz, as well as funding from CREOi and the National Geographic Committee for Research and Exploration to A. Koltz2018-10-0

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

Author Posting. © The Author(s), 2017. This is the author's version of the work. It is posted here by permission of John Wiley & Sons for personal use, not for redistribution. The definitive version was published in Oikos 127 (2018): 460-471, doi:10.1111/oik.04398.A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.National Science Foundation Grants DEB 102683 to support the Arctic LTER to G. Shaver, OPP 0908502 to L. Gough, and DEB 1210704 to A. Koltz, as well as funding from CREOi and the National Geographic Committee for Research and Exploration to A. Koltz2018-10-0

Imputation of average summer temperature at Zackenberg in 1995 from Differential arthropod responses to warming are altering the structure of Arctic communities

Summer temperature data from the year prior to the start of the study (1995) were unavailable from the Zackenberg climate station and were thus imputed from gridded temperature data for the Zackenberg area from the Climate Research Unit (CRU) TS3.23 Dataset

Changes in raw climate variables over study period at Zackenberg, Greenland from Differential arthropod responses to warming are altering the structure of Arctic communities

Variation in average temperatures, winter duration, and number of winter freeze-thaw events at Zackenberg, Greenland between 1996-2014