1. Introduction
Understanding the impacts of climate change on Arctic vegetation is a key requirement for prediction of how Arctic ecosystems will respond to future conditions [
1]. Passive warming manipulation methodologies, such as hexagonal open-top chambers (OTCs), that simulate future scenarios of warming on small plots (~1 m
2) have been extensively investigated in the Arctic [
1,
2,
3,
4,
5,
6,
7]. Previous warming studies using OTCs have been directed toward analyzing specific species responses [
8], radiation dynamics [
9], plant phenology [
10], snow regime shifts [
11], and trace gas exchange [
12,
13] to name a few. Studies have spanned multiple locations to include gradients; DeFrenne
et al. [
5,
6] examined transplants of forest understory species along a latitudinal gradient, and Oberbauer
et al. [
13] studied ecosystem CO
2 fluxes in response to latitudinal and moisture gradients. Meta-analyses of many sites combined have quantified the impacts of passive warming treatments on tundra vegetation and indicate a warming climate in high latitudes and high elevations is changing plant communities and ecosystem function [
14,
15]. A key variable in the interpretation of the treatment responses is the effectiveness of the OTC experimental warming within the different climate regimes, vegetation types, and soil microclimates of the tundra biome.
The increase in mean air temperatures within OTCs ranges from 1 to 3 °C and is dependent on size and design, site climate, ecosystem type, and soil properties [
3,
4,
16,
17,
18,
19,
20,
21,
22]. Soil temperatures at depth have been regularly tested in OTCs, but generally do not show a significant increase [
23,
24], although surface temperatures do [
22]. Most of the research has focused on air temperatures measured by shielded resistive sensors or thermocouples rather than tissue or surface temperatures, but the measurements have been limited to just a few locations per plot. Considerable emphasis has been placed on documenting gradients in air temperature within OTCs by measuring air temperature at specific positions. However, what determines metabolic rates are tissue and surface temperatures. Tissue surface temperatures are determined to a large degree by radiation loads, boundary layer, and evapotranspiraton rates, whereas air temperature is driven by sensible heat [
22]. Some measurements have been made with digital infrared thermometers in OTCs, but the resolution is limited. Given the small size of most OTCs, that they trap heat by increasing the boundary layer, and that they slightly reduce the light available to plants inside, detailed comparisons of the surface temperatures of vascular plants, moss, lichens, litter, and soil in and out of OTCs would be highly desirable.
Recent advances in thermal infrared photography (TIR) have made possible fine-scale, calibrated surface temperature measurement at resolutions previously unattainable with digital infrared thermometers. The objective of this study was to quantify the range and spatial variation in temperatures of tundra surfaces (vascular and non vascular plants, lichens, litter, and soil) induced by the presence of OTCs installed along moisture and latitudinal gradients in Arctic Alaska, USA.
2. Materials and Methods
The locations used for this research were situated along a gradient from High to Low Arctic (
Figure 1) including: Barrow (71°18′N, 156°40′W; 7 masl), Atqasuk (70°29′N, 157°25′W; 21 masl), and Toolik Lake, Alaska (68°37′N, 149°36′W; 736 masl). All of these locations are part of the International Tundra Experiment (ITEX) and the Arctic Observatory Network (AON-ITEX). Data for this project are archived at the ACADIS data repository [
25]. The OTCs used in this study for passive warming treatments were hexagonal in shape (~1 m in diameter, ~0.5 m in height) with an open top allowing for ventilation and precipitation inputs. They were constructed using 1.0 mm thick walls of Sun-Lite
® HP fiberglass glazing (Kalwall Corporation, Manchester, NH, USA) that have high solar transmittance in visible wavelengths (~86%) and low transmittance in longwave (thermal) wavelengths (~5%). The walls of the chambers are inwardly-inclined (60° with respect to the horizontal) to trap heat. Each location has had OTCs deployed in representative plant communities at two points on the local soil moisture spectrum (wet/moist and dry) since the mid 1990s. Occurrence of plant growth forms and surface features in the plots was recorded using point-intercept methods (0.75 m
2 area with 100 grid points [
17]). Percent cover of major surface feature types for each site and moisture regime are presented in
Figure 2. Detailed site information is presented below.
Figure 1.
Map of the three locations used in this study along with weather data during image acquisition. Soil moisture measurements recorded within three days of image acquisition are from plots along the ITEX transects immediately adjacent to the study sites as described in Healey
et al. [
26] that represent the wet/moist and dry ITEX OTC/Control plots.
Figure 1.
Map of the three locations used in this study along with weather data during image acquisition. Soil moisture measurements recorded within three days of image acquisition are from plots along the ITEX transects immediately adjacent to the study sites as described in Healey
et al. [
26] that represent the wet/moist and dry ITEX OTC/Control plots.
The Barrow wet site is located on the edge of a thaw lake basin and is dominated by graminoids, in particular
Carex aquatilis [
4]. Soils are organic (Histic Pergelic Cryaquepts) with shallow depth of thaw. As a result, standing water is frequently present at the wet site. The vegetation at the Barrow dry site is dominated by a dwarf deciduous shrub,
Salix rotundifolia, with evergreen shrubs (e.g.,
Cassiope tetragona) and graminoids (e.g.,
Luzula spp.,
Arctagrostis latifolia). Soils at the Barrow dry site are mostly mineral Pergelic Cryaquepts on silt, sand, and gravel on a raised beach ridge [
4].
Figure 2.
Aggregate percent cover of major plant growth forms and surface features at each site (BRW = Barrow; ATQ = Atqasuk, TLK = Toolik Lake) for dry (
a) and wet/moist (
b) moisture regimes (ctl = control, otc = open top chamber). Note: cover data were not available for all plots at Toolik Lake [
25].
Figure 2.
Aggregate percent cover of major plant growth forms and surface features at each site (BRW = Barrow; ATQ = Atqasuk, TLK = Toolik Lake) for dry (
a) and wet/moist (
b) moisture regimes (ctl = control, otc = open top chamber). Note: cover data were not available for all plots at Toolik Lake [
25].
The wet meadow at Atqasuk is located on the edge of a thaw lake basin and is dominated by
Carex aquatilis [
4]. Standing water is present throughout most of the summers in this site. Soils at the wet site are Histic Pergelic Cryaquepts. Soils at the dry site are Pergelic Cryopsamments on aolian sand of a stabilized sand dune. Vegetation at the dry site is dominated by the dwarf evergreen shrubs
Ledum palustre,
Cassiope tetragona, and
Vaccinium vitis-idaea and the graminoids
Hierochloe alpina and
Luzula confusa [
4].
At Toolik, the OTCs are placed on moist tussock tundra rather than wet sedge tundra because of the great abundance of tussock tundra in the region around Toolik Lake and the relative rarity of wet sedge vegetation. Tussock tundra at Toolik is dominated by the graminoids
Eriophorum vaginatum and
Carex bigelowii, the deciduous shrubs
Betula nana and
Salix pulchra, and the dwarf evergreen shrubs
Ledum palustre and
Vaccinium vitis-idaea [
11].
Eriophorum vaginatum, the dominant plant at the site, is a sedge that forms tussocks or raised mounds with the current year’s growth, persisting on top of the previous year’s growth. Soils are Pergelic Cryaquepts with thaw depths attaining 50–60 cm. Standing water is rarely present at the site except during snowmelt. The dry site vegetation is dry heath dominated by
Dryas octopetala,
Salix phlebophylla,
Arctous alpina, and fruticose lichens [
11,
21] located on well-drained rocky Pergelic Cryumbrepts. Thaw depth may attain 1 m or more.
The timing of this study was designed to analyze thermal properties of vegetation and uncovered soil/moss during peak growth in the Arctic growing season (late July–early August). Thermal infrared photographs were collected from a stable platform/tripod once at midday (~12:00–14:00 AST) from a height of 2.5–3.0 m between 4 and 6 August 2014 (Barrow: 4 August, Atqasuk and Toolik Lake: 6 August) at four randomly selected control plots and four coinciding OTCs within two different moisture regimes at each study location (wet/moist and dry: 24 plots overall). Imagery was collected at midday to minimize any shading effects from the OTC walls. Multiple images of each target were acquired to quantify measurement bias and ensure data integrity was upheld (
i.e., maintain nadir angle and ensure camera focus). At Barrow and Toolik Lake we used FLIR A655sc cameras (FLIR Systems, Wilsonville, OR, USA) which have an uncooled microbolometer detector equipped with a 13.1 mm lens exhibiting a 45° field of view, a spatial resolution of roughly 3 mm (640 × 480: over 300,000 pixels per image), and a spectral range of 7.5–14 μm. At Atqasuk, we used a FLIR A325sc camera which has an uncooled microbolometer detector equipped with a 10 mm lens exhibiting a 45° field of view, a spatial resolution of roughly 6 mm (320 × 240: over 76,000 pixels per image—not resampled to the A655sc resolution), and a spectral range of 7.5–13 μm. The thermal sensitivity of both camera models, defined by the Noise Equivalent Temperature Difference (NETD) is 0.03 °C at 30 °C. Accuracy of temperature measurements for the FLIR a655sc and a325sc cameras is ±2% of the reading with an average bias of ±0.104 °C. All images were atmospherically corrected via FLIR’s Research IR Software with inputs of atmospheric temperature and humidity acquired from weather stations onsite, optic temperature (assumed to be equal to air temperature), and surface emissivity was set to 0.98 following the average emissivity between 7.5 and 14 μm for tundra species reported in Wilber
et al. [
27]. Weather conditions for each day and location are presented in Map 1 and all sites experienced partly-mostly cloudy sky conditions on image dates. We calculated the average temperature of all pixels in each plot for each image, then consolidated the results for each treatment (wet/moist or dry) to examine differences within and among sites. To exclude non-uniform, microclimatic effects of the close proximity of the OTC to the vegetation around the inside perimeter of the structure, the OTC itself and the surface underlying the angled walls of the OTC were removed from image analysis to focus on the surface temperature of a circular area designated around the edge of the OTC opening that is visible from the nadir only (
Figure 3,
Figure 4,
Figure 5,
Figure 6,
Figure 7 and
Figure 8).
Figure 3.
Images depicting four Barrow dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the ring appearing in panels (d,h) is gas exchange collar not related to this study. Areas of image analysis are outlined with dotted white lines. Data within the transparent circular area has been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock.
Figure 3.
Images depicting four Barrow dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the ring appearing in panels (d,h) is gas exchange collar not related to this study. Areas of image analysis are outlined with dotted white lines. Data within the transparent circular area has been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock.
Figure 4.
Images depicting four Barrow wet site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the rings appearing in panels (b,f,i–p) are gas exchange collars not related to this study. Areas of image analysis are outlined with dotted white lines. Data in the transparent circular areas have been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. White grids symbolize chamber base points or plant identification tags.
Figure 4.
Images depicting four Barrow wet site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the rings appearing in panels (b,f,i–p) are gas exchange collars not related to this study. Areas of image analysis are outlined with dotted white lines. Data in the transparent circular areas have been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. White grids symbolize chamber base points or plant identification tags.
Figure 5.
Images depicting four Atqasuk dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted black lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock.
Figure 5.
Images depicting four Atqasuk dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted black lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock.
Figure 6.
Images depicting four Atqasuk wet site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the rings appearing in panels (i–p) are gas exchange collars not related to this study. Areas of image analysis are outlined with dotted white lines. Data in the transparent circular areas have been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. White grids symbolize chamber base points or plant identification tags.
Figure 6.
Images depicting four Atqasuk wet site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Note: the rings appearing in panels (i–p) are gas exchange collars not related to this study. Areas of image analysis are outlined with dotted white lines. Data in the transparent circular areas have been removed. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. White grids symbolize chamber base points or plant identification tags.
Figure 7.
Images depicting four Toolik Lake dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted white lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. Note: cover data were not available for all plots.
Figure 7.
Images depicting four Toolik Lake dry site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted white lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. Note: cover data were not available for all plots.
Figure 8.
Images depicting four Toolik Lake moist site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted white lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. Note: cover data were not available for all plots.
Figure 8.
Images depicting four Toolik Lake moist site plots (a–d) and the coinciding thermal images below (e–h); and images of four additional plots (i–l) with coinciding thermal images below (m–p). Areas of image analysis are outlined with dotted white lines. All temperatures are in degrees Celsius. Plant growth forms and surface features are depicted with the following abbreviations—BRYO: bryophyte/moss; DSHR: deciduous shrub; ESHR: evergreen shrub; FORB: forb; GRAM: graminoid; LICH: lichen; LITT: leaf litter; SOIL: bare soil/rock. Note: cover data were not available for all plots.
To quantify warming in this study, we subtracted the average surface temperature among all pixels within each control plot (CTL) from all pixels within the coinciding OTC immediately adjacent (
Figure 3,
Figure 4,
Figure 5,
Figure 6,
Figure 7 and
Figure 8) and as calculated percent difference as follows:
We present both absolute and percent differences because both approaches have value. However, careful consideration is required when examining percent differences between OTCs and controls due to the dependence on the base values involved in the calculation. Therefore, the absolute difference may be most appropriate for this analysis.
5. Conclusions
This study reports on the use of advanced thermal infrared imagery to analyze the peak growing season temperature patterns within OTCs, the passive warming devices commonly used for simulation of future temperature regimes in Arctic, Antarctic, and alpine communities. We selected long-term monitoring sites spanning latitudinal and moisture gradients where OTCs have been in place every growing season since the mid 1990s. The use of this technology enabled for the first time the quantification of the frequency distribution of surface temperatures within the OTCs and control plots at high spatial resolution. Major findings of our study are: (1) Similar to air temperature modification by OTCs, midday surface temperatures increased by 1–3 °C on average across all sites; (2) The greatest average surface warming (wet and dry sites combined) occurred at the highest latitude site (2.65 °C: Barrow, AK) and the lowest average surface warming occurred at our lowest latitude site at (1.27 °C: Toolik Lake); (3) Surface temperatures were higher and OTCs tended to increase temperatures more at dry sites than wet sites; (4) Surface temperatures within OTCs have much broader temperature distributions then those within control plots. These results imply that plant responses within OTCs should have greater variance than those in control plots.
This study serves as a foundation for future studies examining temperature-related phenomena (such as metabolism, heat stress, water stress,
etc.) at finer scales, potentially targeting thermal properties of specific species, rather than just a plot average. Ground-based Light Detection and Ranging (LIDAR), in combination with detailed species abundance, and spatial composition data (
i.e., proximity to neighboring plants and height differences among species present) would aid in quantifying the level to which surface structure plays a role in temperature modification to different tundra species at such a fine scale. Other variables to consider for future research are differences in temperature modification due to irradiance level, fraction direct/diffuse, sun angle, and duration of sunlight. Considering image processing techniques, when sites are geographically close, improvements are potentially attainable in similar studies if radiometric temperatures are analyzed which relieves the complications of atmospheric correction and emissivity requirements. Although, the instruments used in this study utilize on-the-fly radiometric calibration with an internal blackbody. Furthermore, the very small distance between our sensor and target reduces the impact of atmospheric effects and potential errors in emissivity assignment become negligible. Becker and Li found that if variation in emissivity and actual temperature within a pixel are small, then the radiometric temperature and the surface temperature are the same [
32]. Further research could quantify the magnitude of warming by each species, using radiometric temperatures for validation of results to potentially identify thresholds for survival or mortality throughout different time periods including diurnal, weekly, monthly, and seasonal timescales to investigate potential bias by midday image acquisition times at peak growing season. We foresee future studies expanding these findings to describe species-level thermodynamics that could prove highly valuable toward a better understanding of species-specific responses to changes in Arctic climate conditions.