Overcoming confusion and stigma in habitat fragmentation research.

Anthropogenic habitat loss is widely recognized as a primary environmental concern. By contrast, debates on the effects of habitat fragmentation persist. To facilitate overcoming these debates, here we: (i) review the state of the literature on habitat fragmentation, finding widespread confusion and stigma; (ii) identify consequences of this for biodiversity conservation and ecosystem management; and (iii) suggest ways in which research can move forward to resolve these problems. Confusion is evident from the 25 most-cited fragmentation articles published between 2017 and 2021. These articles use five distinct concepts of habitat fragmentation, only one of which clearly distinguishes habitat fragmentation from habitat area and other factors ('fragmentation per se'). Stigmatization is evident from our new findings that fragmentation papers are more charged with negative sentiments when compared to papers from other subfields in the environmental sciences, and that fragmentation papers with more negative sentiments are cited more. While most empirical studies of habitat fragmentation per se find neutral or positive effects on species and biodiversity outcomes, which implies that small habitat patches have a high cumulative value, confusion and stigma in reporting and discussing such results have led to suboptimal habitat protection policy. For example, government agencies, conservation organizations, and land trusts impose minimum habitat patch sizes on habitat protection. Given the high cumulative value of small patches, such policies mean that many opportunities for conservation are being missed. Our review highlights the importance of reducing confusion and stigma in habitat fragmentation research. To this end, we propose implementing study designs in which multiple sample landscapes are selected across independent gradients of habitat amount and fragmentation, measured as patch density. We show that such designs are possible for forest habitat across Earth's biomes. As such study designs are adopted, and as language becomes more precise, we expect that confusion and stigma in habitat fragmentation research will dissipate. We also expect important breakthroughs in understanding the situations where effects of habitat fragmentation per se are neutral, positive, or negative, and the reasons for these differences. Ultimately this will improve efficacy of area-based conservation policies, to the benefit of biodiversity and people.

Oil Infrastructure has Greater Impact than Noise on Stress and Habitat Selection in Three Grassland Songbirds.

Oil extraction may impact wildlife by altering habitat suitability and affecting stress levels and behavior of individuals, but it can be challenging to disentangle the impacts of infrastructure itself on wildlife from associated noise and human activity at well sites. We evaluated whether the demographic distribution and corticosterone levels of three grassland passerine species (Chestnut-collared Longspur, Calcarius ornatus; Baird's Sparrow, Centronyx bairdii; and Savannah Sparrow Passerculus sandwichensis) were impacted by oil development in southern Alberta, Canada. We used a landscape-scale oil well noise-playback experiment to evaluate whether impacts of wells were caused by noise. Surprisingly, higher-quality female Chestnut-collared Longspurs tended to nest closer to oil wells, while higher-quality Savannah Sparrows generally avoided nesting sites impacted by oil wells. Corticosterone levels in all species varied with the presence of oil development (oil wells, noise, or roads), but the magnitude and direction of the response was species and stimulus specific. While we detected numerous impacts of physical infrastructure on stress physiology and spatial demographic patterns, few of these resulted from noise. However, all three species in this study responded to at least one disturbance associated with oil development, so to conserve the grassland songbird community, both the presence of physical infrastructure and anthropogenic noise should be mitigated.

Avian behaviour changes in response to human activity during the COVID-19 lockdown in the United Kingdom.

Human activities may impact animal habitat and resource use, potentially influencing contemporary evolution in animals. In the United Kingdom, COVID-19 lockdown restrictions resulted in sudden, drastic alterations to human activity. We hypothesized that short-term daily and long-term seasonal changes in human mobility might result in changes in bird habitat use, depending on the mobility type (home, parks and grocery) and extent of change. Using Google human mobility data and 872 850 bird observations, we determined that during lockdown, human mobility changes resulted in altered habitat use in 80% (20/25) of our focal bird species. When humans spent more time at home, over half of affected species had lower counts, perhaps resulting from the disturbance of birds in garden habitats. Bird counts of some species (e.g. rooks and gulls) increased over the short term as humans spent more time at parks, possibly due to human-sourced food resources (e.g. picnic refuse), while counts of other species (e.g. tits and sparrows) decreased. All affected species increased counts when humans spent less time at grocery services. Avian species rapidly adjusted to the novel environmental conditions and demonstrated behavioural plasticity, but with diverse responses, reflecting the different interactions and pressures caused by human activity.

Reduced human activity during COVID-19 alters avian land use across North America.

The COVID-19 pandemic resulted in extraordinary declines in human mobility, which, in turn, may affect wildlife. Using records of more than 4.3 million birds observed by volunteers from March to May 2017­2020 across Canada and the United States, we found that counts of 66 (80%) of 82 focal bird species changed in pandemic-altered areas, usually increasing in comparison to prepandemic abundances in urban habitat, near major roads and airports, and in counties where lockdowns were more pronounced or occurred at the same time as peak bird migration. Our results indicate that human activity affects many of North America's birds and suggest that we could make urban spaces more attractive to birds by reducing traffic and mitigating the disturbance from human transportation after we emerge from the pandemic.

Diel patterns of predation and fledging at nests of four species of grassland songbirds.

Although it is common for nestlings to exhibit a strong bias for fledging in the morning, the mechanisms underlying this behavior are not well understood. Avoiding predation risk has been proposed as a likely mechanism by a number of researchers. We used video surveillance records from studies of grassland birds nesting in North Dakota, Minnesota, and Wisconsin to determine the diel pattern of nest predation and fledging patterns of four ground-nesting obligate grassland passerines (Grasshopper Sparrow (Ammodramus savannarum), Savannah Sparrow (Passerculus sandwichensis), Bobolink (Dolichonyx oryzivorus), and Eastern Meadowlark (Sturnella magna)). We used the nest predation pattern as a surrogate for predation activity to test whether nestlings minimized predation risk by avoiding fledging when predation activity was high and preferentially fledging when predation risk was low. Predation activity was significantly lower starting 3 hr before sunrise and ending 3 hr after sunrise, followed by a transition to a period of significantly higher activity lasting for 4 hr, before declining to an average activity level for the rest of the diel period. There was little evidence that the four grassland bird species avoided fledging during the high-risk period and Savannah Sparrow fledged at higher rates during that period. All four species had hours during the low-risk period where they fledged at higher rates, but only Grasshopper Sparrow fledged preferentially during that period. Bobolink and Eastern Meadowlark had multiple hours with high fledging rates throughout the daytime period, resulting in no relationship between probability of fledging and predation risk. Given the species variability in fledging pattern seen in our study, it is unlikely that there is a universal response to any driver that affects time of fledging. Further study is needed to understand the complex interplay between species ecology and drivers such as physiology, energetics, and predation in affecting grassland bird fledging behavior.

Effects of Natural Habitat Loss and Edge Effects on Wild Bees and Pollination Services in Remnant Prairies.

Habitat loss and edge effects resulting from habitat fragmentation are key processes implicated in the decline of bee populations globally. Their effects on wild bees and their pollination services in natural ecosystems are poorly understood, particularly in North American prairies. Our objectives were to determine whether natural habitat loss and edge effects affect bee abundance and pollination services in the Northern Great Plains. We sampled bee abundance and pollination services along transects beginning at road or tree edges in grasslands located in Manitoba, Canada. We measured bee abundance using pan traps, and pollination services using seed-set of Brassica rapa (L.) (Brassicales: Brassicaceae) phytometers. We collected local-scale habitat data by measuring occurrence of flowering species, vegetation type, and vegetation structure, and we measured habitat amount at 1-km radii using GIS analysis of landscape cover. Increasing amounts of habitat loss resulted in declines in bee abundance, and sometimes in pollination services. Results varied with bee life-history: proximity to road edges negatively affected social bees, and litter depth had negative effects on below- ground-nesting bees. Surprisingly, few effects on bees led to corresponding impacts on pollination services. This suggests that conservation of intact natural habitat across the northern Great Plains is important for maintaining resilient and diverse bee communities, but that efforts to conserve bee populations cannot be assumed to also maintain all associated pollination services.

Host dispersal and landscape conversion are associated with the composition of haemosporidian parasites of the golden-winged warbler.

Understanding factors that influence the spatial and temporal distributions of blood parasites is important to help predict how host species and their parasites may respond to global change. Factors that may influence parasite distributions are land cover and host dispersal patterns, which may result in exposure of a host to novel parasites, or escape from parasites of their origin. We screened golden-winged warblers from across the United States and Canada for blood parasites, and investigated whether land-use patterns or host dispersal affected the prevalence and composition of haemosporidian assemblages. Parasite prevalence varied strongly with study area, and areas with high agricultural cover had a significantly higher prevalence of Leucocytozoon and Parahaemoproteus parasites. Lineages of Parahaemoproteus and Leucocytozoon were genetically differentiated among study areas, and prevalence and composition of parasite assemblages indicated an increase in parasite prevalence and accumulation of unique parasite lineages from the southeast to the northwest. This matches the historical range expansion and natal dispersal patterns of golden-winged warblers, and suggests that golden-winged warblers may have been sensitive to novel parasites as they dispersed. The high prevalence and diversity of parasite lineages in the north-west extent of their breeding range (Manitoba) indicates that this population may face unique pressures.

Noisy anthropogenic infrastructure interferes with alarm responses in Savannah sparrows (Passerculus sandwichensis).

Many birds rely on anti-predator communication to protect their nests; however, anthropogenic noise from industrial activities such as oil and gas development may disrupt acoustic communication. Here, we conducted acoustic playback experiments to determine whether Savannah sparrows (Passerculus sandwichensis) responded to conspecific alarm calls by delaying feeding visits, and whether this response was impaired by noise-producing natural gas compressor stations, generator- or grid-powered screw pump oil wells, and noise amplitude. We played alarm calls, and, as a control, western meadowlark songs, to Savannah sparrows as they approached their nests to feed their nestlings, and measured feeding latency. The greatest impacts on behaviour were detected at the noisiest treatment, compressor stations; feeding latency was shortened here compared with control sites, which may expose nests to greater predation risk. As noise amplitudes increased, Savannah sparrows took longer to feed following meadowlark playbacks, perhaps because noise interfered with interpretation of acoustic cues. The effects of compressor stations on anti-predator behaviour may be best explained by the distracting effects of anthropogenic noise, while increases in feeding latency following meadowlark playbacks may be explained by a heightened response threshold caused by acoustic masking. Industrial infrastructure can influence the reproductive success of wildlife through its impact on perception and interpretation of conspecific signals, but these effects are complex.

Noise Source and Individual Physiology Mediate Effectiveness of Bird Songs Adjusted to Anthropogenic Noise.

Anthropogenic noise is a pervasive pollutant altering behaviour of wildlife that communicates acoustically. Some species adjust vocalisations to compensate for noise. However, we know little about whether signal adjustments improve communication in noise, the extent to which effectiveness of adjustments varies with noise source, or how individual variation in physiology varies with response capacity. We played noise-adjusted and unadjusted songs to wild Passerculus sandwichensis (Savannah Sparrows) after measurements of adrenocortical responsiveness of individuals. Playbacks using songs adjusted to noisy environments were effective in restoring appropriate conspecific territorial aggression behaviours in some altered acoustic environments. Surprisingly, however, levels of adrenocortical responsiveness that reduced communication errors at some types of infrastructure were correlated with increased errors at others. Song adjustments that were effective in communicating for individuals with lower adrenocortical responsiveness at pumpjacks were not effective at screwpumps and vice versa. Our results demonstrate that vocal adjustments can sometimes allow birds to compensate for disruptions in communication caused by anthropogenic noise, but that physiological variation among receivers may alter effectiveness of these adjustments. Thus mitigation strategies to minimize anthropogenic noise must account for both acoustic and physiological impacts of infrastructure.

Conventional oil and natural gas infrastructure increases brown-headed cowbird (Molothrus ater) relative abundance and parasitism in mixed-grass prairie.

The rapid expansion of oil and natural gas development across the Northern Great Plains has contributed to habitat fragmentation, which may facilitate brood parasitism of ground-nesting grassland songbird nests by brown-headed cowbirds (Molothrus ater), an obligate brood parasite, through the introduction of perches and anthropogenic edges. We tested this hypothesis by measuring brown-headed cowbird relative abundance and brood parasitism rates of Savannah sparrow (Passerculus sandwichensis) nests in relation to the presence of infrastructure features and proximity to potential perches and edge habitat. The presence of oil and natural gas infrastructure increased brown-headed cowbird relative abundance by a magnitude of four times, which resulted in four times greater brood parasitism rates at infrastructure sites. While the presence of infrastructure and the proximity to roads were influential in predicting brood parasitism rates, the proximity of perch sites was not. This suggests that brood parasitism associated with oil and natural gas infrastructure may result in additional pressures that reduce productivity of this declining grassland songbird.

Spatiotemporal variation in mechanisms driving regional-scale population dynamics of a Threatened grassland bird.

To achieve national population targets for migratory birds, landscape-level conservation approaches are increasingly encouraged. However, knowledge of the mechanisms that drive spatiotemporal patterns in population dynamics are needed to inform scale-variant policy development. Using hierarchical Bayesian models and variable selection, we determined by which mechanism(s), and to what extent, changes in quantity and quality of surrogate grassland habitats contributed to regional variation in population trends of an obligatory grassland bird, Bobolink (Dolichonyx oryzivorous). We used North American Breeding Bird Survey data to develop spatially explicit models of regional population trends over 25 years across 35 agricultural census divisions in Ontario, Canada. We measured the strength of evidence for effects of land-use change on population trends over the entire study period and over five subperiods. Over the entire study period, one region (Perth) displayed strong evidence of population decline (95% CI is entirely below 0); four regions displayed strong evidence of population increase (Bruce, Simcoe, Peterborough, and Northumberland). Population trends shifted spatially among subperiods, with more extreme declines later in time (1986-1990: 28% of 35 census divisions, 1991-1995: 46%, 1996-2000: 40%, 2001-2005: 66%, 2006-2010: 82%). Important predictors of spatial patterns in Bobolink population trends over the entire study period were human development and fragmentation. However, factors inferred to drive patterns in population trends were not consistent over space and time. This result underscores that effective threat identification (both spatially and temporally) and implementation of flexible, regionally tailored policies will be critical to realize efficient conservation of Bobolink and similar at-risk species.

Effects of shallow natural gas well structures and associated roads on grassland songbird reproductive success in Alberta, Canada.

Grassland songbird populations across North America have experienced dramatic population declines due to habitat loss and degradation. In Canada, energy development continues to fragment and disturb prairie habitat, but effects of oil and gas development on reproductive success of songbirds in North American mixed-grass prairies remains largely unknown. From 2010-2012, in southeastern Alberta, Canada, we monitored 257 nests of two ground-nesting grassland songbird species, Savannah sparrow (Passerculus sandwichensis) and chestnut-collared longspur (Calcarius ornatus). Nest locations varied with proximity to and density of conventional shallow gas well structures and associated roads in forty-two 258-ha mixed-grass prairie sites. We estimated the probabilities of nest success and clutch size relative to gas well structures and roads. There was little effect of distance to or density of gas well structure on nest success; however, Savannah sparrow experienced lower nest success near roads. Clutch sizes were lower near gas well structures and cattle water sources. Minimizing habitat disturbance surrounding gas well structures, and reducing abundance of roads and trails, would help minimize impacts on reproductive success for some grassland songbirds.

Effects of ambient noise on detectability and localization of avian songs and tones by observers in grasslands.

Probability of detection and accuracy of distance estimates in aural avian surveys may be affected by the presence of anthropogenic noise, and this may lead to inaccurate evaluations of the effects of noisy infrastructure on wildlife. We used arrays of speakers broadcasting recordings of grassland bird songs and pure tones to assess the probability of detection, and localization accuracy, by observers at sites with and without noisy oil and gas infrastructure in south-central Alberta from 2012 to 2014. Probability of detection varied with species and with speaker distance from transect line, but there were few effects of noisy infrastructure. Accuracy of distance estimates for songs and tones decreased as distance to observer increased, and distance estimation error was higher for tones at sites with infrastructure noise. Our results suggest that quiet to moderately loud anthropogenic noise may not mask detection of bird songs; however, errors in distance estimates during aural surveys may lead to inaccurate estimates of avian densities calculated using distance sampling. We recommend caution when applying distance sampling if most birds are unseen, and where ambient noise varies among treatments.

Urban Rights-of-Way as Reservoirs for Tall-Grass Prairie Plants and Butterflies.

Urban rights-of-way may be potential reservoirs of tall-grass prairie plants and butterflies. To determine if this is true, in 2007-2008, we conducted vegetation surveys of species richness and cover, and butterfly surveys of species richness and abundance, along 52 transmission lines and four remnant prairies in Winnipeg, Manitoba. We detected many prairie plants and butterflies within transmission lines. Some unmowed and infrequently managed transmission lines had native plant species richness and total percent cover of native plants comparable to that of similar-sized remnant tall-grass prairies in the region. Although we did not find significant differences in overall native butterfly numbers or species richness between rights-of-way and remnant prairies, we found lower numbers of some prairie butterflies along frequently mowed rights-of-way than within remnant tall-grass prairies. We also observed higher butterfly species richness along sites with more native plant species. By reducing mowing and spraying and reintroducing tall-grass prairie plants, urban rights-of-way could serve as extensive reservoirs for tall-grass prairie plants and butterflies in urban landscapes. Eventually, managing urban rights-of-way as reservoirs for tall-grass prairie plants and animals could contribute to the restoration of tall-grass prairie in the North American Midwest.

Direct effects of cattle on grassland birds in Canada.

Effects of grazing on grassland birds are generally thought to be indirect, through alteration of vegetation structure; however, livestock can also affect nest survival directly through trampling and other disturbances (e.g., livestock-induced abandonment). We extracted data on nest fates from 18 grazing studies conducted in Canada. We used these data to assess rates of nest destruction by cattle among 9 ecoregions and between seasonal and rotational grazing systems. Overall, few nests were destroyed by cattle (average 1.5% of 9132 nests). Nest destruction was positively correlated with grazing pressure (i.e., stocking rate or grazing intensity), but nest survival was higher in more heavily grazed areas for some species. Because rates of destruction of grassland bird nests by cattle are low in Canada, management efforts to reduce such destruction may not be of ecological or economic value in Canada.