Effects of bird species-level environmental preference on landscape-level richness-heterogeneity relationships

The niche-based argument that species are filtered from environments in which they cannot sustain viable populations is the basis of the Richness-Heterogeneity Relationship (RHR). However, the multi-dimensionality of niches suggests that the RHR may take different shapes along different environmental axes, with potential confounding effects if filtering along the axes is not equally strong. Here, we explore how different structural and landscape variables drive the RHR as the accumulative outcome of environmental preferences at the species-level while considering the intercorrelation between heterogeneity levels along three niche axes. We used occurrence data of avifauna from 226 sites situated along a grassland-to-woodland gradient in a Midwestern USA study area. In each site, we quantified horizontal (habitat cover type), vertical (vegetation height structure), and spatial (habitat configuration) heterogeneity and explored the shape of the observed RHR at the landscape scale, as well as the correlations among heterogeneity levels at different axes. We then fitted species distribution models to environmental variables from the three axes separately and compared the stacked probabilities of occurrences of all species to the observed species richness. We found that predictions of richness patterns improved when more than one heterogeneity axis was included in RHR models, and that habitat suitability along different axes is not equally strong. Furthermore, a unimodal RHR along the vegetation height axis, which the species distribution models revealed to be a weak predictor for most species, may arise through intercorrelation with heterogeneity along the two other axes, along which we recorded stronger signals of environmental preference at the species level. Our results emphasize the importance of selecting relevant niche axes in studies of species richness patterns because ultimately, these patterns reflect the various environmental preferences of individual species.

File: Gavish-et-al_2021_Effects-of-bird-species-level-environmental-preferences-on-landscape-level-richness-heterogeneity-relationships.-Basic-and-Applied-Ecology-56-1-13..pdf

Assessing the effectiveness of a forest Habitat Conservation Plan for a threatened seabird, the Marbled Murrelet

Habitat Conservation Plans (HCPs) commonly facilitate habitat conservation on private land in the United States, yet the effectiveness of individual HCPs is rarely evaluated. Here, we assess the effectiveness of a high-profile HCP created by a lumber company to protect old-growth forests used for breeding by Marbled Murrelets (Brachyramphus marmoratus) on private land. We used 17 years of HCP-monitoring data to compare trends in murrelet occupancy and inland counts between private HCP areas and public reference areas over time. Based on occupancy models applied to audio-visual survey data, average occupancy was higher in public reference areas (0.85; 85% confidence intervals [CI]: 0.79–0.90) than in private HCP areas (0.46; 85% CI: 0.38–0.54). Numerically, trends in occupancy were slightly positive in public areas ( = 1.01; 85% CI: 0.94–1.08) and slightly negative in private areas ( = 0.97; 85% CI: 0.87–1.06), but CI did not preclude stable occupancy on both ownerships. Based on generalized linear mixed models applied to inland radar survey data, murrelet counts in private HCP areas (least-squares [LS] mean = 8.7; 85% CI: 6.2–12.2) were lower than those in public reference areas (LS mean = 14.8; 85% CI: 10.1–21.7), but CI overlapped. Murrelet counts declined by 12–17% annually on both ownerships over the study period based on the top model, but a closely competing interactive model suggested more rapid declines in public reference (14–20%) than in private HCP (10–15%) areas. Both models indicated that murrelet counts were negatively related to sea surface temperature, suggesting that warm ocean conditions negatively affect murrelet breeding effort. Collectively, these results suggest that while HCP habitat may be lower quality than public reference areas, the HCP has likely not exacerbated ongoing declines of murrelets in the region. This work highlights the importance of including reference areas when evaluating conservation policies.

File: Brunk-et-al-2021_Effectiveness-of-HCP-for-Marbled-Murrelet.pdf

Patterns of bird species richness explained by annual variation in remotely sensed Dynamic Habitat Indices

Bird species richness is highly dependent on the amount of energy available in an ecosystem, with more available
energy supporting higher species richness. A good indicator for available energy is Gross Primary Productivity
(GPP), which can be estimated from satellite data.
Our question was how temporal dynamics in GPP affect bird species richness. Specifically, we evaluated the
potential of the Dynamic Habitat Indices (DHIs) derived from MODIS GPP data together with environmental and
climatic variables to explain annual patterns in bird richness across the conterminous United States. By focusing
on annual DHIs, we expand on previous applications of multi-year composite DHIs, and could evaluate lag-effects
between changes in GPP and species richness.
We used 8-day GPP data from 2003 to 2013 to calculate annual DHIs, which capture three aspects of vegetation
productivity: (1) annual cumulative productivity, (2) annual minimum productivity, and (3) annual
seasonality expressed as the coefficient of variation in productivity. For each year from 2003 to 2013, we
calculated total bird species richness and richness within six functional guilds, based on North American
Breeding Bird Survey data.
The DHIs alone explained up to 53% of the variation in annual bird richness within the different guilds
(adjusted deviance-squared D2adj = 0.20–0.52), and up to 75% of the variation (D2adj = 0.28–0.75) when
combined with other environmental and climatic variables. Annual DHIs had the highest explanatory power for
habitat-based guilds, such as grassland (D2adj = 0.67) and woodland breeding species (D2adj = 0.75). We found
some inter-annual variability in the explanatory power of annual DHIs, with a difference of 5–7 percentage
points in explained variation among years in DHI-only models, and 3–7 points for models combining DHI,
environmental and climatic variables. Our results using lagged year models did not deviate substantially from
same-year annual models.
We demonstrate the relevance of annual DHIs for biodiversity science, as effective predictors of temporal
variation in species richness patterns. We suggest that the use of annual DHIs can improve conservation planning,
by conveying the range of patterns of biodiversity response to global changes, over time.

File: Hobi-et-al-2021_BirdSpeciesRichness_DynamicHabIndices_EcolIndicators.pdf

Reducing anthropogenic subsidies can curb density of overabundant predators in protected areas

Protected areas safeguard biodiversity and provide opportunities for human recreation. However, abundant anthropogenic food subsidies associated with human activities in protected areas can lead to high densities of generalist predators, posing a threat to rare species at broad spatial scales. Reducing anthropogenic subsidies could curb populations of overabundant predators, yet the effectiveness of this strategy is unclear. We characterized changes in the foraging ecology, body condition, and demography of a generalist predator, the Steller’s jay, three years after implementation of a multi-faceted management program to reduce anthropogenic subsidies in a protected area in California. Stable isotope analysis revealed that the proportional contribution of anthropogenic foods to jay diets declined from 88% to 47% in response to management. Overlap between jay home ranges decreased after management began, while home range size, body condition, and individual fecundity remained stable. Adult density in subsidized areas decreased markedly from 4.33 (SE: ±0.91) to 0.65 (±0.20) jays/ha after the initiation of management, whereas density in unsubsidized areas that were not expected to be affected by management remained stable (0.70 ± 0.22 pre-management, 0.58 ± 0.38 post-management). Thus, the response of jays to management was density-dependent such that reduced densities facilitated the maintenance of individual body condition and fecundity. Importantly, though, jay population size and collective reproductive output declined substantially. Our study provides evidence that limiting anthropogenic subsidies can successfully reduce generalist predator populations and be part of a strategy to increase compatibility of species protection and human recreation within protected areas.

File: Brunk-et-al-2021_Reducing-anthropogenic-subsidies_Stellers-Jays_Biological-Conservation.pdf

Flying insect abundance in northern Wisconsin depends on lakes, vegetation, and time of the summer

Mayfly emergence resulted in an increase in flying insect abundance in late June and early July.

Considering their outsized importance as prey for so many species one would assume that patterns of insect abundance and their determinants have been well-studied. On the contrary, insect ecology is poorly understood and documented. Our study sought to gain an understanding of the subgroup of insects that fly, with a particular emphasis on groups that spend part of their life in lakes and streams.

Insect trapping took place in the northern highland lake district. The study area consists of northern mixed forest with one of the highest densities of lakes in the world.
Insect trapping took place in the northern highland lake district. The study area consists of northern mixed forest with one of the highest densities of lakes in the world.

We conducted insect trapping over three years in the forest landscape of northern Wisconsin, near UW-Madison’s Trout Lake Research Station. We trapped insects May-August around five different lakes and identified them in the lab.

Mean insect abundance varied over the course of the summer with peak average insect abundance occurring in late May and early June (Julian Day 145-155) in both years.
Mean insect abundance varied over the course of the summer with peak average insect abundance occurring in late May and early June (Julian Day 145-155) in both years.

There were several patterns that stood out. Flying insects tended to be many times more abundant in nearshore areas compared to interior forests. Different groups of insects showed different patterns. Diptera, including deerflies, midges, and gnats were the most abundant insects overall. As expected, emergent aquatic groups such as midges, mayflies, and dragonflies were more abundant in nearshore areas while beetles and thrips were more abundant in forest interiors. There were also multiple peaks of abundance through the season with large emergence events of midges and mayflies driving much of the pattern. In addition, local canopy cover was negatively correlated with insect abundance.

We observed birds, bats, and fish consuming flying insects. Abundance of these insect predators likely tracks the abundance of their insect prey. In addition, insects perform other ecosystems services such as pollination and nutrient cycling. Understanding the patterns and drivers of insect abundance can help us better understand northern Wisconsin forest ecosystems.

Contrasting seasonal patterns of relative temperature and thermal heterogeneity and their influence on breeding and winter bird richness patterns across the conterminous United States

Environmental heterogeneity enhances species richness by creating niches and providing refugia. Spatial variation in climate has a particularly strong positive correlation with richness, but is often indirectly inferred from proxy variables, such as elevation and related topographic heterogeneity indices, or derived from interpolated coarsegrain weather station data. Our aim was to develop new remotely sensed metrics of relative temperature and thermal heterogeneity, compare them with proxy measures, and evaluate their performance in predicting species richness patterns. We analyzed Landsat 8’s Thermal Infrared Sensor data, calculated two thermal metrics during summer and winter, and compared their seasonal spatial patterns with those of elevation and topographic heterogeneity. We fit generalized least squares models to evaluate each variable’s effect in predicting seasonal bird richness using data from the North American Breeding Bird Survey. Generally speaking, neither elevation nor topographic heterogeneity were good proxies for temperature or thermal heterogeneity, respectively. Relative temperature had a non-linear relationship with elevation that was negatively quadratic in summer, but slightly positively quadratic in winter. Topographic heterogeneity had a stronger positive relationship with thermal heterogeneity in winter than in summer. The magnitude and direction of elevation–temperature and topographic heterogeneity–thermal heterogeneity relationships in each season also varied substantially across ecoregions. Remotely sensed metrics of relative temperature and thermal heterogeneity improved the predictive performance of species richness models, and both thermal variables had significant effects on bird richness that were independent of elevation and topographic heterogeneity. Thermal heterogeneity was positively related to total breeding bird richness, migrant breeding bird richness and resident bird richness, whereas topographic heterogeneity was negatively related to total breeding richness and unrelated to migrant or resident bird richness. Because thermal and topographic heterogeneity had contrasting seasonal patterns and effects on richness, they must be carefully contextualized when guiding conservation priorities.

File: ecog.05520.pdf

Responses to land cover and grassland management vary across life history stages for a grassland specialist

Grassland birds have exhibited dramatic and widespread declines since the mid-20th century. Greater prairie chickens (Tympanuchus cupido pinnatus) are considered an umbrella species for grassland conservation and are frequent targets of management, but their responses to land use and management can be quite variable. We used data collected during 2007–2009 and 2014–2015 to investigate effects of land use and grassland management practices on habitat selection and survival rates of greater prairie chickens in central Wisconsin, USA. We examined habitat, nest-site, and brood-rearing site selection by hens and modeled effects of land cover and management on survival rates of hens, nests, and broods. Prairie chickens consistently selected grassland over other cover types, but selection or avoidance of management practices varied among life-history stages. Hen, nest, and brood survival rates were influenced by different land cover types and management practices. At the landscape scale, hens selected areas where brush and trees had been removed during the previous year, which increased hen survival. Hens selected nest sites in hay fields and brood-rearing sites in burned areas, but prescribed fire had a negative influence on hen survival. Brood survival rates were positively associated with grazing and were highest when home ranges contained ≈15%–20% shrub/tree cover. The effects of landscape composition on nest survival were ambiguous. Collectively, our results highlight the importance of evaluating responses to management efforts across a range of life-history stages and suggest that a variety of management practices are likely necessary to provide structurally heterogeneous, high-quality habitat for greater prairie chickens. Brush and tree removal, grazing, hay cultivation, and prescribed fire may be especially beneficial for prairie chickens in central Wisconsin, but trade-offs among life-history stages and the timing of management practices must be considered carefully.

File: ece3.6805.pdf

Winter Habitat Indices (WHIs) for the contiguous US and their relationship with winter bird diversity

The seasonal dynamics of snow cover strongly affect ecosystem processes and winter habitat, making them an important driver of terrestrial biodiversity patterns. Snow cover data from the Moderate Resolution Imaging Spectroradiometer (MODIS) Aqua and Terra satellites can capture these dynamics over large spatiotemporal scales, allowing for the development of indices with specific application in ecological research and predicting biodiversity. Here, our primary objective was to derive winter habitat indices (WHIs) from MODIS that quantify snow season length, snow cover variability, and the prevalence of frozen ground without snow as a proxy for subnivium conditions. We calculated the WHIs for the full snow year (Aug-Jul) and winter months (Dec-Feb) across the contiguous US from 2003/04 to 2017/18 and validated them with ground-based data from 797 meteorological stations. To demonstrate the potential of the WHIs for biodiversity assessments, we modeled their relationships with winter bird species richness derived from eBird observations. The WHIs had clear spatial patterns reflecting both altitudinal and latitudinal gradients in snow cover. Snow season length was generally longer at higher latitudes and elevations, while snow cover variability and frozen ground without snow were highest across low elevations of the mid latitudes. Variability in the WHIs was largely driven by elevation in the West and by latitude in the East. Snow season length and frozen ground without snow were most accurately mapped, and had correlations with station data across all years of 0.91 and 0.85, respectively. Snow cover variability was accurately mapped for winter (r = 0.79), but not for the full snow year (r = 􀀀 0.21). The model containing all three WHIs used to predict winter bird species richness patterns across the contiguous US was by far the best, demonstrating the individual value of each index. Regions with longer snow seasons generally supported fewer species. Species richness increased steadily up to moderate levels of snow cover variability and frozen ground without snow, after which it steeply declined. Our results show that the MODIS WHIs accurately characterized unique gradients of snow cover dynamics and provided important information on winter habitat conditions for birds, highlighting their potential for ecological research and conservation planning.

File: GudexCross_etal_2021_MODIS_WHIs_BirdDiversity.pdf

Satellite image texture captures vegetation heterogeneity and explains patterns of bird richness

Addressing global declines in biodiversity requires accurate assessments of key environmental attributes determining patterns of species diversity. Spatial heterogeneity of vegetation strongly affects species diversity patterns, and measures of vegetation structure derived from lidar and satellite image texture analysis correlate well with species richness. Our goal here was to gain a better understanding of why image texture explains bird richness, by linking field-based measures of vegetation structure directly with both image texture and bird richness. In addition, we asked how image texture compares with lidar-based canopy height variability, and how sensor resolution affects the explanatory power of image texture. We generated texture metrics from 30 m (Landsat 8) and 10 m (Sentinel-2) resolution Enhanced Vegetation Index (EVI) imagery from 2017 to 2019. We compared textures with vegetation metrics and bird richness data from 27 National Ecological Observatory Network (NEON) terrestrial field sites across the continental US. Both 30 and 10 m resolution texture metrics were strongly correlated with lidar-based canopy height variability (|r| = 0.64 and 0.80, respectively). Texture was moderately correlated with field-based metrics, including variability of vegetation height and tree stem diameter, and foliage height diversity (range |r| = 0.31–0.52). Generally, 10 m resolution texture had stronger correlations with lidar and field-based metrics than 30 m resolution texture. In univariate linear models of total bird richness, 10 m resolution texture metrics also had higher explanatory power (up to R2adj = 0.45), than 30 m texture metrics (up to R2adj = 0.31). Among all metrics evaluated, the 10 m homogeneity texture was the best univariate predictor of total bird richness. In multivariate bird richness models that combined texture with lidar-based canopy height variability and field-based metrics, both 30 m and 10 m resolution texture metrics were selected in top-ranked models and independently contributed explanatory power (up to R2adj = 46%). Lidar-based canopy height variability was also selected in a top-ranked model of total bird richness, but independently contributed only 15% of the variance explained. Our results show satellite image texture characterized multiple features of structural and compositional vegetation heterogeneity, complemented more commonly used metrics in models of bird richness and for some guilds outperformed both lidar-based canopy height variability and field-based vegetation measurements. Ours is the first study to directly link image texture both to specific components of vegetation heterogeneity and to bird richness across multiple ecoregions and spatial resolutions, thereby shedding light on habitat features underlying the strong correlation between image texture and biodiversity.

File: Farwell-et-al-2021_Sat-image-texture_veg_birds_RemSensEnv.pdf

Habitat heterogeneity captured by 30-m resolution satellite image texture predicts bird richness across the United States

Species loss is occurring globally at unprecedented rates, and effective conservation planning requires an understanding of landscape characteristics that determine biodiversity patterns. Habitat heterogeneity is an important determinant of species diversity, but is difficult to measure across large areas using field-based methods that are costly and logistically challenging. Satellite image texture analysis offers a cost-effective alternative for quantifying habitat heterogeneity across broad spatial scales. We tested the ability of texture measures derived from 30-m resolution Enhanced Vegetation Index (EVI) data to capture habitat heterogeneity and predict bird species richness across the conterminous United States. We used Landsat 8 satellite imagery from 2013–2017 to derive a suite of texture measures characterizing vegetation heterogeneity. Individual texture measures explained up to 21% of the variance in bird richness patterns in North American Breeding Bird Survey (BBS) data during the same time period. Texture measures were positively related to total breeding bird richness, but this relationship varied among forest, grassland, and shrubland habitat specialists. Multiple texture measures combined with mean EVI explained up to 41% of the variance in total bird richness, and models including EVI-based texture measures explained up to 10% more variance than those that included only EVI. Models that also incorporated topographic and land cover metrics further improved predictive performance, explaining up to 51% of the variance in total bird richness. A texture measure contributed predictive power and characterized landscape features that EVI and forest cover alone could not, even though the latter two were overall more important variables. Our results highlight the potential of texture measures for mapping habitat heterogeneity and species richness patterns across broad spatial extents, especially when used in conjunction with vegetation indices or land cover data. By generating 30-m resolution texture maps and modeling bird richness at a near-continental scale, we expand on previous applications of image texture measures for modeling biodiversity that were either limited in spatial extent or based on coarse-resolution imagery. Incorporating texture measures into broad-scale biodiversity models may advance our understanding of mechanisms underlying species richness patterns and improve predictions of species responses to rapid global change.

File: Farwell-eta-l-2020_Habitat-heterogeneity-30-m-and-birds_Ecological-Applications.pdf