White-Headed Woodpecker Nesting Ecology After Wildfire

ABSTRACT Within forests susceptible to wildfire and insect infestations, land managers need to balance dead tree removal and habitat requirements for wildlife species associated with snags. We used Mahalanobis distance methods to develop predictive models of white-headed woodpecker (Picoides albolarvatus) nesting habitat in postfire ponderosa pine (Pinus ponderosa)-dominated landscapes on the Fremont-Winema National Forests in south central Oregon, USA. The 1-km radius (314 ha) surrounding 45 nest sites was open-canopied before fire and a mosaic of burn severities after wildfire. The 1-ha surrounding nests of white-headed woodpeckers had fewer live trees per hectare and more decayed and larger diameter snags than at non-nest sites. The leading cause of nest failure seemed to be predation. Habitat and abiotic features were not associated with nest survival. High daily survival rates and little variation within habitat features among nest locations suggest white-headed woodpeckers were consistently selecting high suitability habitats. Management activities that open the forest canopy and create conditions conducive to a mosaic burn pattern will probably provide suitable white-headed woodpecker nesting habitat after wildfire. When making postfire salvage logging decisions, we suggest that retention of larger, more decayed snags will provide nesting habitat in recently burned forests.     

Roosts of Allen's Lappet-Browed Bat in Northern Arizona

ABSTRACT In Arizona, USA, Allen's lappet-browed bat (Idionycteris phyllotis) forms maternity colonies in ponderosa pine (Pinus ponderosa) snags. There is little information on the roosting habitat of males. We used radiotelemetry to locate 16 maternity, 3 postlactating, and 2 bachelor roosts and combined data with unpublished data for maternity roosts (n = 11) located in 1993–1995. Most (96%) maternity roosts were in large-diameter (x̄ ± SE: 64 ± 2.7 cm) ponderosa pine snags under sloughing bark. Models that best predicted the probability of a snag's use as a maternity roost indicated bats selected taller snags closer to forest roads than comparison snags. Maternity roosts averaged 11 bats per roost (SE = 2, n = 15; from exit counts) and were an average distance of 1.6 km from capture sites (SE = 0.3, n = 17). Bachelor roosts were in vertical sandstone cliff faces in pinyon-juniper (Pinus edulis-Juniperus spp.) woodlands approximately 12 km from capture sites; these and other capture records in Arizona indicated sexual segregation may have occurred during the maternity season. Of 11 maternity snag roosts located in 1993–1995, only one continued to function as a roost. Resource managers should maintain patches of large-diameter ponderosa pine snags with peeling bark to provide maternity roosting habitat for Allen's lappet-browed bat.     

Cavity use and breeding biology of endangered Bahama Swallows (Tachycineta cyaneoviridis): implications for conservation

Bird populations, especially on islands, have declined or gone extinct due to overhunting, habitat loss and fragmentation, and adverse effects of the introduction of non-native species. Bahama Swallows (Tachycineta cyaneoviridis), endangered secondary cavity nesters that breed on only three islands in the northern Bahamas, are an island species with a declining population, but the causes of this decline are unknown. During four breeding seasons on Great Abaco Island (2014–2017), we identified cavity-nesting resources used by breeding swallows in native pine forest and other habitats, and estimated phenology and breeding parameters from a subset of nests. Bahama Swallows nest in cavities in a variety of structures, but rely most on woodpecker-excavated cavities in pine snags and utility poles. Swallows nesting in cavities in pine snags had higher fledging success (92%) than those nesting in cavities in utility poles (50–62%), which were concentrated in non-pine habitat that may expose swallows to predation and increased competition for nest cavities from other species. The high reproductive success of Bahama Swallows nesting in the pine forest indicates that the decline in population cannot be attributed to poor productivity on southern Great Abaco. However, our results suggest that the dependence of Bahama Swallows on cavities excavated by Hairy Woodpeckers (Dryobates villosus) for nesting sites may be a factor in their decline, and highlight the potential importance of the protection and management of pine forests in future efforts to ensure the survival of Bahama Swallows.     

Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Temporal dynamics of roost snags of long-legged myotis in the Pacific Northwest,USA

Snags are used as roosting sites by many bats living in coniferous forests of western North America. Thus, providing sufficient numbers of snags both spatially and temporally in forested landscapes is critical to sustaining populations of these species. One aspect that remains poorly understood is length of time that roost snags persist on the landscape in a form suitable for use by bats. This information is critical for forest-planning efforts in ensuring long-term availability of snag resources on forested landscapes. We monitored condition of 339 snags used as roosting sites by long-legged myotis (Myotis volans) 1–5 years post-discovery from 2001 to 2006 across 6 watersheds in Washington, Oregon, and Idaho, USA. Persistence rates (i.e., probability a snag remains standing from year x to x + 1) of roost snags declined with year post-discovery in all study areas. Fir snags (Abies spp.) exhibited lower persistence rates than other conifer species. Data for the Washington area indicated only 4.3% of roost snags likely remain standing 10 years post-discovery, with half-lives of all snag species <3 roost-years. Model ranking of habitat models predicting fall year of roost snags revealed that snag condition models were the most parsimonious in all geographic locations. Roost snags larger in diameter, shorter in height, and with fewer branches on the bole were likely to persist for more years. These data indicate that snags used as roosts by long-legged myotis are suitable as roosting sites for only a few years before falling. We recommend management policies for coniferous forests in the Pacific Northwest, USA, that promote sufficient leave-trees in set-aside areas to provide for future suitable, large-diameter snags for bats in managed, forested landscapes. © 2012 The Wildlife Society.     

Enhancing bark- and wood-boring beetle colonization and survival in vertical deadwood during thinning entries

Snags are used as habitat by several organisms, including bark- and wood-boring beetles, which contribute to snag decomposition and facilitate subsequent snag colonization by other organisms. However, snags seldom occur in young managed forests. This study examines how snag characteristics and spatial arrangement affect bark- and wood-boring beetle colonization and survival in spruce plantations subjected to commercial thinning, thinning with biomass removal and thinning with snag creation where a few clumps of trees were girdled. To this end, we documented the volume of snags, their characteristics and the number of beetle emergence holes in their basal section. Beetle colonization and survival to adulthood in snags was ~33 times greater when expressed per unit area in plantations supplemented with clumps of girdled trees than in other thinning regimes where low densities of beetle emergence holes per hectare were documented. Snag diameter at breast height, snag species and thinning treatments influenced the number of emergence holes per snag. Positive spatial autocorrelation between the abundance of emergence holes per snag was detected but no other spatial effect was noticeable. This work suggests that vertical deadwood is a limiting factor for bark- and wood-boring beetle colonization within plantations and challenges current thinking about the effect of thinning on beetle communities that was developed from window trap studies. These results underline the importance of large diameter snags for beetle conservation within managed forests and demonstrate that tree girdling during thinning entries is a viable method for creating snags to enhance beetle colonization and survival.     

Autumn use of woody snags by fishes in backwater and channel border habitats of a large river

Snags are important to fish communities in small rivers and streams, but their importance to fishes in large rivers has not been investigated. This study examined snag use by fishes during autumn in backwater and channel border habitats in the upper Mississippi River, and compared these to fish communities in reference sites without snags. Species assemblages differed significantly between backwater and channel border habitats, and between snag and reference sites within the channel border, likely responding to differences in substrate, depth, and current velocity. In both habitats, average fish biomass and abundance were higher (2 to 50 ×) at snag sites than at reference sites, but these differences were significant only for channel border biomass. Fish taxa richness differed between backwater and channel border habitats, but not between snag and reference sites. Most large piscivorous fishes (e.g., Micropterus spp., Stizostedion spp.), several insectivorous fishes (Lepomis macrochirus, Ambloplites rupestris, Minytrema melanops), and a few prey fishes (L. macrochirus, Notropis atherinoides) were significantly more abundant at snag sites than at reference sites, suggesting active selection of snags for foraging or protection. Snag quality, as assessed by a snag rating index, had a direct effect on attracting fish communities with greater biomass, especially within the channel border habitat. These results indicate that snags are important habitat for fish communities in both backwaters and channel border habitats of the upper Mississippi River.     

Habitat suitability and nest survival of white-headed woodpeckers in unburned forests of Oregon

We evaluated habitat suitability and nest survival of breeding white-headed woodpeckers (Picoides albolarvatus) in unburned forests of central Oregon, USA. Daily nest-survival rate was positively related to maximum daily temperature during the nest interval and to density of large-diameter trees surrounding the nest tree. We developed a niche-based habitat suitability model (partitioned Mahalanobis distance) for nesting white-headed woodpeckers using remotely sensed data. Along with low elevation, high density of large trees, and low slope, our habitat suitability model suggested that interspersion–juxtaposition of low- and high-canopy cover ponderosa pine (Pinus ponderosa) patches was important for nest-site suitability. Cross-validation suggested the model performed adequately for management planning at a scale >1 ha. Evaluation of mapped habitat suitability index (HSI) suggested that the maximum predictive gain (HSI = 0.36), where the number of nest locations are maximized in the smallest proportion of the modeled landscape, provided an objective initial threshold for identification of suitable habitat. However, managers can choose the threshold HSI most appropriate for their purposes (e.g., locating regions of low–moderate suitability that have potential for habitat restoration). Consequently, our habitat suitability model may be useful for managing dry coniferous forests for white-headed woodpeckers in central Oregon; however, model validation is necessary before our model could be applied to other locations. © 2011 The Wildlife Society.     

Snag Benchmarks and Treatment Options for Mixed‐Pine Forest Restoration in Eastern Upper Michigan

Snags (dead‐standing trees) are biological legacies that remain after disturbances in forests. We enhanced the ecological underpinnings of snag management within the context of mixed‐pine forest restoration in the northern Lake States by quantifying characteristics of live trees and snags within eighty-five 500‐m² plots at Seney National Wildlife Refuge (NWR) in eastern Upper Michigan. Study plots represented reference conditions (i.e. no past harvesting, relatively unaltered fire regime) and altered (i.e. harvested, altered fire regime) conditions. We also compared three treatments for creating snags from live trees. Snags were found in 87% of the reference plots and 85% of the altered plots. The only snag variables that differed between plot types were mean snag basal area, which was greater in altered plots (Student's t‐test, p = 0.04), and mean percent total basal area of snags (greater in reference plots, p = 0.06). The composition of snags differed only in the 10‐ and 25‐cm diameter classes (Multi‐Response Permutation Procedure, p < 0.10). The percentage of snags that developed into the most advanced decay class (DC) differed among treatments after 4 years (χ² = 16.49, p < 0.01), with 26% of girdled trees, 3% of prescribed fire trees, and zero topped trees reaching DC5. Logistic regression illustrated that the influence of predictor variables on DC development varied by species and treatment. The findings from this study, past studies, and ongoing projects at Seney NWR are directly applicable to innovative management of snags in mixed‐pine forests.     

Factors affecting Florida scrub-jay nest survival on Ocala National Forest,Florida

One of the main populations of the Florida scrub-jay (Aphelocoma coerulescens), a federally threatened species, occurs on Ocala National Forest, Florida. We determined the nest daily survival rate (DSR) of 474 nests of Florida scrub-jays in stands subject to sand pine reforestation management after timber harvesting or wildfire on Ocala National Forest. We used the information-theoretic approach with logistic-exposure modeling to determine the most likely models to account for DSR for the incubation and nestling stages separately. The models consisted of 4 components (temporal, management, habitat, and helpers) with each consisting of one to several specific variables. In the incubation stage all the best models included the temporal component alone or in concert with the habitat, helpers, or habitat and helpers components. Model averaging in the incubation stage indicated support for the year and day × year in the temporal component and stand age in the habitat component. In the nestling stage, top models all included the temporal component alone or with helpers or habitat, helpers, and the interaction of habitat and helpers. Model averaging in the nestling stage showed support for the year, nest age, and to a lesser extent day in the temporal component and helpers. The management component, which consisted of no site preparation, wildfire burn, post-harvest burn, chop and seed, or seed only, had little influence on nest survival for the incubation or nestling stages. However, we identified several other management factors that may increase Florida scrub-jay populations despite having no effect on DSR. First, the proportion of Florida scrub-jay nests was significantly higher than expected in burned habitat based on habitat availability, indicating a potential preferred nesting habitat conducive to population growth. Second, incubation stage DSR with respect to stand age (habitat component) declined to stand age 10 yr and then began to increase, which may be attributed to the higher bird population in the prime habitat in the middle stand ages. The denser population may result in more competition for resources, and possibly may attract more predators, resulting in a lower DSR. Thus, although DSR may be lower at the mid-stand ages, the overall population may actually be optimal. Therefore, to increase the Florida scrub-jay population on Ocala National Forest, we recommend maintaining a mosaic of stands ≤20 yr of age and emphasizing natural reseeding and site preparation via burning. © 2011 The Wildlife Society.     

Red-Headed Woodpecker Nest-Habitat Thresholds in Restored Savannas

Abstract: The red-headed woodpecker (Melanerpes erythrocephalus) is a widespread, adaptable species that continues to decline across North America. We examined stand, nest-tree, and cavity characteristics of red-headed woodpeckers in restored savannas within the Necedah National Wildlife Refuge, Wisconsin, USA, during 2002 and 2003. Based on availability, red-headed woodpeckers selected snags and trees with greater dead limb length. Red-headed woodpeckers nested in areas with greater basal area, cavity density, snag density, limb-tree density, and total dead limb length. Red-headed woodpeckers exhibited a decadent-tree threshold that was most accurately measured by the number of trees with dead limbs around (0.04 ha) nests. We found that the probability of a red-headed woodpecker nest being present greatly increased above the decadent-tree threshold. Woodland managers throughout the red-headed woodpecker's extensive breeding range can use our results and recommendations to guide decadent-tree retention for this species.     

Spatial and Temporal Considerations in Restoring Habitat for Wildlife

An accumulated body of theory and empirical evidence suggests that habitat selection by animals is a scale‐dependent, hierarchical process. Hierarchy theory predicts that habitat suitability is influenced by the interaction of factors at multiple spatial scales from the microsite to the landscape and that higher‐order factors impose constraints at lower levels. For instance, large‐scale factors such as landscape context may make a site unsuitable for a species even if the vegetation structure and composition are appropriate. In addition, the spatial arrangement of habitat elements at all scales must be considered when planning restoration efforts. For example, the presence of snags does not ensure that the site will be suitable for snag‐dependent species. The size, age, and spacing of snags and their juxtaposition to other habitat elements must also be considered. Finally, all habitats are dynamic, and therefore the ecological processes that contribute to those dynamics must be maintained or suitable substitutes included in the recovery plan. When considering restoring habitat for wildlife, we recommend that managers: (1) identify the wildlife species they want to target for restoration efforts, (2) consider the size and landscape context of the restoration site and whether it is appropriate for the target species, (3) identify the habitat elements that are necessary for the target species, (4) develop a strategy for restoring those elements and the ecological processes that maintain them, and (5) implement a long‐term monitoring program to gauge the success of the restoration efforts.     

Evening Bat Summer Roost-Site Selection on a Managed Pine Landscape

ABSTRACT Creation and maintenance of forested corridors to increase landscape heterogeneity has been practiced for decades but is a new concept in intensively managed southern pine (Pinus spp.) forests. Additionally, more information is needed on bat ecology within such forest systems. Therefore, we examined summer roost-site selection by evening bats (Nycticeius humeralis) in an intensively managed landscape with forested corridors in southeastern South Carolina, USA, 2003–2006. We radiotracked 53 (26 M, 27 F) adult evening bats to 75 (31 M, 44 F) diurnal roosts. We modeled landscape-level roost-site selection with logistic regression and evaluated models using Akaike's Information Criterion for small samples. Model selection results indicated that mature (≥40 yr) mixed pine-hardwood stands were important roost sites for male and lactating female evening bats. Upland forested corridors, comprised of mature pine or mixed pine-hardwoods, were important roosting habitats for males and, to a lesser extent, lactating females. Male roosts were farther from open stands and lactating female roosts were farther from mid-rotation stands than randomly selected structures. Our results suggest roost structures (i.e., large trees and snags) in mature forests are important habitat components for evening bats. We recommend maintaining older (>40 yr old) stand conditions in the form of forest stands or corridors across managed landscapes to provide roosting habitat. Furthermore, our results suggest that an understanding of sex-specific roost-site selection is critical for developing comprehensive guidelines for creating and maintaining habitat features beneficial to forest bats.     

High Density of Tree-Cavities and Snags in Tropical Dry Forest of Western Mexico Raises Questions for a Latitudinal Gradient

It has been suggested that a latitudinal gradient exists of a low density of snags and high density of naturally-formed tree-cavities in tropical vs. temperate forests, though few cavities may have characteristics suitable for nesting by birds. We determined snag and cavity density, characteristics, and suitability for birds in a tropical dry forest biome of western Mexico, and evaluated whether our data fits the trend of snag and cavity density typically found in tropical moist and wet forests. We established five 0.25-ha transects to survey and measure tree-cavities and snags in each of three vegetation types of deciduous, semi-deciduous, and mono-dominant Piranhea mexicana forest, comprising a total of 3.75 ha. We found a high density of 77 cavities/ha, with 37 cavities suitable for birds/ha, where density, and characteristics of cavities varied significantly among vegetation types. Lowest abundance of cavities occurred in deciduous forest, and these were in smaller trees, at a lower height, and with a narrower entrance diameter. Only 8.6% of cavities were excavated by woodpeckers, and only 11% of cavities were occupied, mainly by arthropods, though 52% of all cavities were unsuitable for birds. We also found a high density of 56 snags/ha, with greatest density in deciduous forest (70 snags/ha), though these were of significantly smaller diameter, and snags of larger diameter were more likely to contain cavities. The Chamela-Cuixmala tropical dry forest had the highest density of snags recorded for any tropical or temperate forest, and while snag density was significantly correlated with mean snag dbh, neither latitude nor mean dbh predicted snag density in ten forest sites. The high spatial aggregation of snag and cavity resources in tropical dry forest may limit their availability, particularly for large-bodied cavity adopters, and highlights the importance of habitat heterogeneity in providing resources for primary and secondary cavity-nesters.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Forest Stand Characteristics Altered by Restoration Affect Western Bluebird Habitat Quality

Forest managers are setting Ponderosa pine (Pinus ponderosa) forests in the southwestern United States on a trajectory toward a restored ecosystem by reducing tree densities and managing with prescribed fire. The process of restoration dramatically alters forest stands, and the effects of these changes on wildlife remain unclear. Our research evaluated which aspects of habitat alteration from restoration treatments may be affecting the habitat quality of Western Bluebird (Sialia mexicana), an insectivorous songbird whose populations are declining. Habitat loss resulting from fire‐suppression activities may be partially responsible for their population declines; thus, the bluebird is a good representative species for assessing how the reconstruction of presuppression forest conditions can affect wildlife. We measured habitat variables at 63 successful and 19 unsuccessful Western Bluebird nests in 1999–2001 and 2003. We compared habitat models that represented bluebird biology and habitat changes from restoration. Two models of nest success that included (1) an increased herbaceous and bare ground cover and (2) increased Gambel oak (Quercus gambelii) densities and reduced Ponderosa pine densities were most supported by the data. Increased herbaceous ground cover and Gambel oak density likely represent improved invertebrate assemblages and thus improved forage abundance for nesting bluebirds. Lower Ponderosa pine densities may provide bluebirds with open perches from which to hunt and thereby improve the availability of invertebrates as a food source. We also provide a landscape‐scale example of changes to bluebird habitat quality from treatments, which we recommend as a useful tool in restoration planning.     

Den Site Selection by the Florida Spotted Skunk

The eastern spotted skunk (Spilogale putorius) is a species of conservation concern in much of its range and has experienced a decline since the early to mid-1990s. But the subspecies that inhabits peninsular Florida, the Florida spotted skunk (S. p. ambarvalis), might still be abundant and is an important nest predator of the endangered Florida grasshopper sparrow (Ammodramus savannarum floridanus). To gain insight on this little-studied subspecies and inform potential management strategies, we conducted a resource selection study on the Florida spotted skunk. We examined 5 hypotheses for den site selection related to den type, vegetation, and landscape characteristics in a dry prairie ecosystem in central Florida. We tracked 36 individual skunks to 757 den sites. Using discrete choice analysis, we found that male and nonbreeding female skunks at our study site were 5 times more likely to select a mammal burrow over a gopher tortoise (Gopherus polyphemus) burrow, that selection of a den site increased 34% for each 1-burrow increase in the number of nearby burrows, and that selection of a den site increased 3% for every 10-cm increase in a visual obstruction index. Similarly, breeding female skunks were more likely to select mammal burrows and shallow depressions over gopher tortoise burrows by 16-fold and 13-fold, respectively, and selection of a den site increased by 75% for every 1-burrow increase in the number of nearby burrows. In contrast to previous studies that occurred in forested, mountainous environments elsewhere in the species' range, our findings suggest that den characteristics might be more important than landscape or vegetation characteristics to Florida spotted skunk den site selection in dry prairie. Additionally, the frequency of prescribed fires on the landscape did not appear to affect Florida spotted skunk den site selection. Thus, Florida spotted skunks in this ecosystem might be landscape generalists, thereby potentially limiting the ability of managers to control nest predation by this subspecies through habitat management. © 2019 The Wildlife Society.     

Poor effectiveness of Natura 2000 beech forests in protecting forest-dwelling bats

With the establishment of the Natura 2000 (N2000) network, the European Union intends to develop strategies to conserve Europe's threatened habitats and species, including bats. Forest-dwelling bats are highly reliant on forest structures, such as snags and hollow trees, which the bats need as roosts. The decrease in such forest microhabitats significantly affects the habitat use and, therefore, the activity in forests. To determine whether N2000 beech forests under active timber production offer better habitats for bats compared to commercially used non-N2000 forests, we measured the bat activity and assessed the potential roosts in trees and snags in eleven pairs of stands. All survey stands represented mesotrophic beech forests (Fagus sylvatica L.) of the N2000 habitat type 9130 (Asperulo-Fagetum) in three European Biogeographic Regions. The activity of all bat species, the activity of priority N2000 species, the species number, the number of trees with roosts and the snag volume did not differ significantly between the N2000 and non-N2000 stands. We conclude that the current management of the N2000 beech forests is almost identical to that of non-N2000 commercial forests, and thus, the N2000 status has not led to an increase of bat-relevant habitat variables yet. Consequently, additional efforts beyond the administrative assignment of N2000 areas are required to build and ensure an ecologically effective and sustainable network of beech forests in Europe, including increasing important forest requirements for bats, such as roosts and snags.