Effects of vegetation, corridor width and regional land use on early successional birds on powerline corridors

Powerline rights-of-way (ROWs) often provide habitat for early successional bird species that have suffered long-term population declines in eastern North America. To determine how the abundance of shrubland birds varies with habitat within ROW corridors and with land use patterns surrounding corridors, we ran Poisson regression models on data from 93 plots on ROWs and compared regression coefficients. We also determined nest success rates on a 1-km stretch of ROW. Seven species of shrubland birds were common in powerline corridors. However, the nest success rates for prairie warbler (Dendroica discolor) and field sparrow (Spizella pusilla) were <21%, which is too low to compensate for estimated annual mortality. Some shrubland bird species were more abundant on narrower ROWs or at sites with lower vegetation or particular types of vegetation, indicating that vegetation management could be refined to favor species of high conservation priority. Also, several species were more abundant in ROWs traversing unfragmented forest than those near residential areas or farmland, indicating that corridors in heavily forested regions may provide better habitat for these species. In the area where we monitored nests, brood parasitism by brown-headed cowbirds (Molothrus ater) occurred more frequently close to a residential area. Although ROWs support dense populations of shrubland birds, those in more heavily developed landscapes may constitute sink habitat. ROWs in extensive forests may contribute more to sustaining populations of early successional birds, and thus may be the best targets for habitat management.     

Legacy forest structure increases bird diversity and abundance in aging young forests

Many studies have demonstrated the importance of early‐successional forest habitat for breeding bird abundance, composition, and diversity. However, very few studies directly link measures of bird diversity, composition and abundance to measures of forest composition, and structure and their dynamic change over early succession. This study examines the relationships between breeding bird community composition and forest structure in regenerating broadleaf forests of southern New England, USA, separating the influences of ecological succession from retained stand structure. We conducted bird point counts and vegetation surveys across a chronosequence of forest stands that originated between 2 and 24 years previously in shelterwood timber harvests, a silvicultural method of regenerating oak‐mixed broadleaf forests. We distinguish between vegetation variables that relate to condition of forest regeneration and those that reflect legacy stand structure. Using principal components analyses, we confirmed the distinction between regeneration and legacy vegetation variables. We ran regression analysis to test for relationships between bird community variables, including nesting and foraging functional guild abundances, and vegetation variables. We confirmed these relationships with hierarchical partitioning. Our results demonstrate that regenerating and legacy vegetation correlate with bird community variables across stand phases and that the strength with which they drive bird community composition changes with forest succession. While measures of regeneration condition explain bird abundance and diversity variables during late initiation, legacy stand structure explains them during stem exclusion. Canopy cover, ground‐story diversity, and canopy structure diversity are the most powerful and consistent explanatory variables. Our results suggest that leaving varied legacy stand structure to promote habitat heterogeneity in shelterwood harvests contributes to greater bird community diversity. Interestingly, this is particularly important during the structurally depauperate phase of stem exclusion of young regenerating forests.     

Cropland edge,forest succession,and landscape affect shrubland bird nest predation

The effects of habitat edges on nest survival of shrubland birds, many of which have experienced significant declines in the eastern United States, have not been thoroughly studied. In 2007 and 2008, we collected data on nests of 5 shrubland passerine species in 12 early successional forest patches in North Carolina, USA. We used model selection methods to assess the effect of distance to cropland and mature forest edge on nest predation rates and additionally accounted for temporal trends, nest stage, vegetation structure, and landscape context. For nests of all species combined, nest predation decreased with increasing distance to cropland edge, by nearly 50% at 250 m from the cropland edge. Nest predation of all species combined also was higher in patches with taller saplings and less understory vegetation, especially in the second year of our study when trees were 4–6 m tall. Predation of field sparrow (Spizella pusilla) nests was lower in landscapes with higher agricultural landcover. Nest predation risk for shrubland birds appears to be greater near agricultural edges than mature forest edges, and natural forest succession may drive patterns of local extirpation of shrubland birds in early successional forest patches. Thus, we suggest that habitat patches managed for shrubland bird populations should be considerably large or wide (>250 m) when adjacent to crop fields and maintained in structurally diverse early seral stages. © 2011 The Wildlife Society.     

王朗自然保护区地栖脊椎动物群落结构和生境类型的关系

2006年8—11月,采用围栏陷阱法对四川省王朗自然保护区地栖脊椎动物的群落结构进行了研究,结合森林砍伐、旅游和植被恢复方式等干扰因子所推动的动物栖息生境变化,探讨了动物群落结构和生境类型的关系。监测历时75 d,8个样地共捕获3纲6目17科21种778只地栖脊椎动物,物种组成包括3种两栖类、1种鸟类和17种哺乳类。每百陷阱日的捕获率为32.4%;以普通鼩鼱、高山姬鼠、纹背鼩鼱、高原林蛙等4种为优势种,占全部捕获数的76.9%;其余17种的个体数均较少,物种多度和其分布区域呈显著线性相关(P0.001)。动物群落多样性指数,以干扰相对弱的原生针叶林物种多样性指数和均匀度指数为最高,而优势度指数最低;其次为旅游干扰的原生林灌丛和自然更新的次生林灌丛;人工种植的次生针叶林多样性指数和均匀度指数最低,而优势度指数最高。以动物群落相似性指数进行系统聚类,8个样地的动物群落聚为3类:即干扰相对弱的原生针叶林动物群落、严重干扰的人工针叶林动物群落、部分干扰的原生林灌丛和次生林灌丛动物群落。     

Regional Synthesis of Habitat Relationships in Shrubland Birds

Abstract: Shrubland birds are declining throughout the eastern United States. To manage scrub-shrub habitats for birds, managers need information on avian habitat relationships. Past studies have produced contradictory results in some cases and may be of limited generality because of site- and habitat-specific factors. We studied shrubland birds across 6 habitats in 3 New England states to provide more general information on habitat relationships than has been possible in past studies. Our study sites included all major scrub-shrub habitats in New England: wildlife openings, regenerating clear-cuts, beaver ponds, utility rights-of-way, pitch pine (Pinus rigida) woodlands, and scrub oak (Quercus ilicifolia) barrens and ranged from Connecticut to northern New Hampshire, with research conducted from 2002 to 2007. Using N-mixture models of repeated point counts, we found that 6 of 12 shrubland birds preferred areas with greater shrub cover. An additional 4 species appeared to prefer areas with lower-stature vegetation and greater forb cover. Eight of 10 bird species showed relationships with cover of individual plant species, with Spiraea spp., willows (Salix spp.), alders (Alnus spp.), and invasive exotics being the most important. We recommend that shrubland management for birds focus on providing 2 distinct habitats: 1) areas of tall (>1.5 m) vegetation with abundant shrub cover and 2) areas of lower (<1.5 m) vegetation with abundant forb cover but fewer shrubs.     

Patterns of bird abundance and habitat use in rice fields of the Kanto Plain, central Japan

Rice fields, the most widespread cropland in Japan, provide important habitat for birds, but few studies have examined bird communities that are influenced by agriculture in rice fields. I studied the seasonal pattern of bird abundance and their habitat preferences in terms of agricultural habitats in the rice fields of the central Kanto Plain, central Japan. In total, 19 waterbird species and 31 landbird species were recorded over the study year. Waterbird occurrence was largely restricted to the cultivation season (i.e. the flooded season, May–August), with maximum abundance in August. Rice fields dried up during winter and supported few waterbirds except for irregular occurrences of little egrets (Egretta garzetta) and common snipes (Gallinago gallinago), which selectively used ditches with shallow water. Landbird abundance increased after the start of cultivation (May) to August, but more birds were attracted to the rice fields in midwinter (around January). For both waterbirds and landbirds, most species preferred levees and fallow grass fields, but avoided fields with rice vegetation. Unplowed fields after harvest were preferred by several species including skylarks (Alauda arvensis), while plowed fields were less attractive to most species. These results suggest the need for advanced agricultural practices to include improvement in habitat quality for birds.     

Seasonality,habitat type and locality influenced bird assemblage structure in Nigeria

This is the first report of the avian assemblage in the study area of Dutse, Nigeria. In addition to recording bird species, the effects of season, dominant vegetation structure, locality and anthropogenic activities on bird abundance, species richness and diversity were investigated. Using the point transect method, 264 points on 48 km of transect were used to count birds between 06:30 and 11:00 from August 2015 to February 2016. A total of 122 bird species of 41 families were recorded. Highest bird species richness was recorded in Warwade, highest abundance in Model, and highest diversity in Malamawa. The dry season and woodland habitat showed higher bird species richness, abundance and diversity than the wet season and shrubland habitat. Tree density was more important in increasing bird abundance than shrub density. Small-scale anthropogenic activities and habitat modification, such as farming, grazing, wood removal and human interference did not appear to have impacted the birds; however, loss of high tree-density woodland habitats may pose a major threat to the bird community in Dutse. The presence of birds of concern in the area suggests the need for conservation efforts of avifauna and as well as the forested habitats in Dutse.     

Effects of Clearfell Harvesting on Lucanid Beetles (Coleoptera: Lucanidae) in Wet and Dry Sclerophyll Forests in Tasmania

The effects of commercial forestry harvest and regeneration practices (clearfelling and slash-burning) on the lucanid fauna of the wet sclerophyll forests of southern Tasmania and the dry sclerophyll forests of eastern Tasmania were examined using pitfall catches. Lucanids are saproxylic beetles, dependent on dead, moribund and decaying wood. Samples taken from old-growth forest and from a chronosequence of sites regenerating after logging, in each forest type, were used to compare the species richness and abundance of the lucanid assemblages. In both forest types, species richness and abundance was highest in the youngest regeneration sites (1–3 year), reflecting the species richness of the original and adjacent unlogged forest, lowest in the older (20–25 year) sites, and variable in the old-growth sites. TWINSPAN cluster analysis showed no clear distinction between regeneration and old-growth forest. The post-harvest slash and stump residue provided an important refugium and initial habitat, but our research indicates that some species may not maintain populations in the long term. Our results suggest that most species of lucanids will find a continuous supply of suitable habitat only in old-growth forests; and such species may become less common as clearfell harvesting leads to a replacement of heterogeneous old-growth forest with single-aged monospecific stands. Continuity of supply of wood in all decay stages, the maintenance of sufficient source areas, and biological connectivity between old-growth stands to enable dispersal, are all likely to be essential to maintain lucanid beetle community integrity. If similar principles apply to other saproxylic species of invertebrate, then clearfelling and slash-burning may cause a gradual extinction of an important element of the forest biota.     

Responses of two species of heathland rodents to habitat manipulation: Vegetation density thresholds and the habitat accommodation model

The abundance of two native rodent species, Rattus lutreolus and Pseudomys gracilicaudatus, has been shown to correlate with vegetation density in coastal wet heath. Fox's habitat accommodation model relates relative abundances of such small mammal species to heathland vegetation regeneration following disturbance. Implicit in the model is recognition that it is successional changes in vegetation, not time per se, that drives the responses of small mammal species along a regeneration axis. Using a brush‐cutter we deliberately removed approximately 85% of vegetation around trapping stations and recorded significant reductions in the abundance of both P. gracilicaudatus (an earlier‐stage colonizing species) and R. lutreolus (a late seral‐stage species). A significant decrease in the abundance of only the latter had been demonstrated previously when 60–70% of the vegetation had been removed. Following the brush‐cutting both species re‐entered the mammalian secondary succession at different times, first P. gracilicaudatus followed by R. lutreolus after the vegetation cover thresholds of each species had been reached. The impact of this habitat manipulation experiment was to produce a retrogression of the small mammal succession, experimentally demonstrating causality between changes in vegetation density and subsequent small mammal habitat use.     

浙江天童苦槠+白栎灌丛群落的生物量研究

采用直接收获法研究了浙江天童地区苦槠＋白栎次生灌丛群落的地上部分生物量。结果表明该区次生灌丛的地上部分生物量为16．77t/hm^2,还不到常绿阔叶林幼林生物量（地上部分）的1／3。次生灌丛生物量的分布,从群落层次上看,主要集中于藻木层;从垂直高度上看,主要分布在0－4m之间;从种类组成上看,主要分布于苦槠、白栎、山矾、茅栗等优势种中。根据生物量的研究,文中还对如何加速次生灌丛的进展演替进行了讨论。     

Recolonization of regenerating open forest by terrestrial lizards following sand mining

The terrestrial lizard fauna of a regenerating mining path situated in open forest on coastal sand dunes was surveyed by pitfall trapping. The regeneration age of the series of chronosequence plots used ranged from 3.9 to 15.9 years. No lizards were recorded on plots less than 5.9 years old. Only four species of lizard were regularly found on the mining path. The skink Ctenotus robustus was the first colonizer and attained peak abundance and biomass after 9 years’ regeneration. Amphibolurus muricatus and Ctenotus taeniolatus were less abundant than C. robustus and did not appear on the mining path until 9 to 11 years’ regeneration. The abundance of C. taeniolatus continued to increase up to the oldest age class (16 years). Linear multiple regression analysis showed that sparse patchy vegetation in the 0–1 m layer and the amount of live shrub cover jointly accounted for 72% of the variance in abundance and 68% of variance in biomass for C. robustus. Fifty percent of the variance in C. taeniolatus abundance was explained by the proportion of plant species endemic to forest, regeneration age and the patchiness in understorey vegetation height. However, 67% of C. taeniolatus biomass was accounted for by plant species richness, plant species diversity, and the proportion of endemic forest and heath plant species. Some lizard species recorded from the surrounding forest were not found on the mining path. Lizards appear to recolonize revegetated areas more slowly than some other animal groups; this ‘sensitivity’ implies that they can act as important ‘indicator species’ of successful regeneration in disturbed areas. The non-linear response with time of some habitat variables makes it difficult to predict the time required for the complete rehabilitation of the mining path; however, it seems likely that a period well in excess of 20 years may be necessary before the lizard fauna on the mining path reaches premined levels.     

Demographic Differences of Black-Capped Vireos in 2 Habitat Types in Central Texas

Abstract: To understand the effects of habitat selection, we analyzed differences in abundance, age structure, and nesting success of black-capped vireos (Vireo atricapilla) in 2 early successional habitat types found on Fort Hood, a 87,890-ha Military Reservation in central Texas, USA. These habitats were 1) large areas of continuously shrubby vegetation (both natural and mechanically made), referred to as shrubland habitat, and 2) anthropogenically created small patches of shrubby vegetation centered on one or several large trees, known locally as donut habitat. The objectives of our study were to determine whether there were differences in abundance, age structure, and daily nest survival in these 2 habitat types and to determine whether donut habitat is high- or low-quality habitat. Donut habitat had a lower abundance of vireos (half as many as shrubland/point count) and a higher percentage of second-year males, suggesting donut habitat was lower-quality habitat than shrubland. Analyses of daily nest survival indicated that habitat, nest height, and year were all important variables. Nests initiated in 2004, located in shrubland habitats, and higher from the ground were more likely to succeed. Our study provided evidence that habitat is a limiting factor for this federally endangered species. Because habitat is limiting, wildlife biologists at Fort Hood should focus on managing higher quality, contiguous shrubland habitat. Wildlife biologists should also continue to monitor areas of donut habitat to determine whether they represent potential population sinks.     

Bats are Not Birds – Different Responses to Human Land‐use on a Tropical Mountain

Land‐use intensification has consequences for biodiversity and ecosystem functioning, with various taxonomic groups differing widely in their sensitivity. As land‐use intensification alters habitat structure and resource availability, both factors may contribute to explaining differences in animal species diversity. Within the local animal assemblages the flying vertebrates, bats and birds, provide important and partly complementary ecosystem functions. We tested how bats and birds respond to land‐use intensification and compared abundance, species richness, and community composition across a land‐use gradient including forest, traditional agroforests (home garden), coffee plantations and grasslands on Mount Kilimanjaro, Tanzania. Furthermore, we asked how sensitive different habitat and feeding guilds of bats and birds react to land‐use intensification and the associated alterations in vegetation structure and food resource availability. In contrast to our expectations, land‐use intensification had no negative effect on species richness and abundance of all birds and bats. However, some habitat and feeding guilds, in particular forest specialist and frugivorous birds, were highly sensitive to land‐use intensification. Although the habitat guilds of both, birds and bats, depended on a certain degree of vegetation structure, total bat and bird abundance was mediated primarily by the availability of the respective food resources. Even though the highly structured southern slopes of Mount Kilimanjaro are able to maintain diverse bat and bird assemblages, the sensitivity of avian forest specialists against land‐use intensification and the dependence of the bat and bird habitat guilds on a certain vegetation structure demonstrate that conservation plans should place special emphasis on these guilds.     

Restoring habitat connectivity over the road: vegetation on a fauna land‐bridge in south‐east Queensland

The Compton Road overpass in southern Brisbane is known to be allowing passage by numerous birds, reptiles and amphibians; but what type of vegetation is providing habitat suitable for these movements? An evaluation of the reconstructed vegetation on the overpass shows that 45 species of plants were present after 4 years of growth. Most of these were in the original planting list but over 40% have almost certainly come from salvaged native topsoil used in the bypass construction. The resulting habitat, 4 years later, now appears similar to a typical coastal eucalypt forest secondary succession, although not yet identical to nearby forest. The current presence of a dense mid‐layer of small trees and shrubs may be an important factor in allowing even the edge‐avoiding species to cross the road. On‐going investigation of successional changes and the changing responses of the birds and other taxa will contribute to understanding the functionality of such crossing structures.     

Afromontane old-field vegetation: secondary succession and the return of indigenous species

South‐west Uganda primarily holds afromontane forests within three protected areas: Mgahinga Gorilla National Park, Echuya Forest Reserve and Bwindi Impenetrable National Park. All forests contain portions of old‐field vegetation. The central question of this study, then, is whether and how natural regeneration of afromontane vegetation would take place. A successional pathway consisting of nine plant communities was found for the first 2 years after cessation of agricultural use. A return of afromontane species despite of an initial dominance of neophytes, a diversification of life form spectra and a growing importance of endozoochory with time belonged to the conspicuous characteristics of this secondary succession. To obtain an insight into the role of birds and buffaloes as possible vectors for seed input, I examined their faeces with regard to germinable seeds. Both, bird faeces, as well as buffalo faeces contained germinable seeds originating from the forest flora. Thus, animal dispersal from the forest into the regenerating zones could be documented. The regeneration potential of the soil‐seed bank seems to be limited in its time scale, and only an initial regeneration capacity could be found. A further progressive succession into a secondary afromontane forest depends on an input of diaspores from undisturbed forest sites. Zoochory appears to be one of the most important dispersal mechanisms.     

Disturbance-induced bird diversity in early successional habitats in the humid temperate region of northern Japan

The positive role of moderate natural disturbance is less known for a mobile organism such as birds, compared to sessile organisms. In the face of recent declines of grassland birds, it is necessary to identify the mechanism to maintain avian diversity in early successional open habitats in different regions. In the humid temperate region, the predominant habitat type is woody vegetation. Therefore, bird communities were studied along vegetational succession using a chronosequence method. TWINSPAN identified three distinctive habitat types (barren, grass-fen and shrubland) and four habitat guilds of birds (pioneer, grassland, ubiquitous and shrub) in the study sites. Path analysis determined the direct and indirect effects of disturbance on the habitat guilds of birds. Short and intermediate intervals of the water disturbance are important to pioneer guilds and grass-fen habitat, respectively. The frequent occurrence of large animals had a negative impact on grassland guild. It suggests that grassland birds less tolerate cattle grazing or human activities in a region with intensive land use. The chronosequence study revealed the dynamic nature of the bird community. Birds occupy habitats following the successional sere of vegetation, responding to animal occurrence and human activities in various manners.     

Invertebrate availability and vegetation characteristics explain use of nonnesting cover types by mature‐forest songbirds during the postfledging period

ABSTRACT Some species of mature‐forest‐nesting songbirds use regenerating clearcuts and forested wetlands during the postfledging period (between nesting and migration). Relatively dense vegetation structure and abundant food resources in non‐mature‐forest cover types have been hypothesized to explain this phenomenon. We examined the relative importance of vegetation structure and invertebrate availability on use of nonnesting cover types by adult and hatch‐year Ovenbirds (Seiurus aurocapilla) and American Redstarts (Setophaga ruticilla) during the postfledging period of 2009 in northern Minnesota. We used mist nets to sample bird use of forested wetlands and regenerating clearcuts of three age groups: 1–6, 7–12, and 16–19 yr after harvest. We modeled captures of birds using vegetation characteristics and invertebrate availability sampled around nets as explanatory variables. For all birds studied, captures were best explained by food availability and secondarily by vegetation characteristics including litter depth and woody debris for Ovenbirds and canopy height for American Redstarts. Shrub‐level invertebrate availability received a cumulative weight of 0.74–0.99 in Akaike's information criterion corrected ranked models for adult and hatch‐year birds of both species. Vegetation density and variation in vegetation density explained almost no variation in captures of either species. We conclude that both invertebrate availability and some vegetation characteristics influence use of nonnesting cover types by Ovenbirds and American Redstarts during the postfledging period, but that invertebrate availability is generally the stronger predictor of that use.     

Understory Vegetation Dynamics of North American Boreal Forests

Understory vegetation is the most diverse and least understood component of North American boreal forests. Understory communities are important as they act as drivers of overstory succession and nutrient cycling. The objective of this review was to examine how understory vegetation abundance, composition, and diversity change with stand development after a major stand replacing disturbance. Understory vegetation abundance and diversity increase rapidly after fire, in response to abundant resources and an influx of disturbance adapted species. The highest diversity occurs within the first 40 years following fire, and declines indefinitely thereafter as a result of decreasing productivity and increased dominance of a small number of late successional feather mosses and woody plant species. Vascular plant and bryophyte/lichen communities undergo very different successional changes. Vascular plant communities are dynamic and change more dramatically with time after fire, whereas bryophyte and lichen communities are much slower to establish and change over time. Considerable variations in these processes exist depending on canopy composition, site condition, regional climate, and frequently occurring non-stand-replacing disturbances. Forest management practices represent a unique disturbance process and can result in different understory vegetation communities from those observed for natural processes, with potential implications for overstory succession and long-term productivity. Because of the importance of understory vegetation on nutrient cycling and overstory composition, post-harvest treatments emulating stand-replacing fire are required to maintain understory diversity, composition, and promote stand productivity in boreal forests.     

Effects of fire on the vegetation of a lowland heathland in North-western Italy

This study focuses on the effect of fire on lowland heathlands at the extreme southern edge of their European distribution (Vauda Nature Reserve, NW Italy). Forty-nine plots (50 m radius) were surveyed between 1999 and 2006. Each year, fire occurrences were recorded and per cent cover of four vegetation types (grassland, heath, low shrubland, and tall shrubland) was estimated in each plot. Vascular plant species richness was also recorded in 255, 1 m² quadrats. After a fire, grassland vegetation expanded, but then declined rapidly as heath and shrubland recovered: 7 years after a fire, tall shrubland encroached on to more than 40% of the plots, and grassland declined from 50% to 20% cover. Between 1999 and 2006, Betula pendula shrubland greatly expanded, while grassland decreased over most of the Reserve, even where fire frequency was high. Tall shrubland had low plant diversity and was dominated by widespread species of lower conservation value. By contrast, early successional vegetation (grassland and low shrubland) had higher richness and more narrowly distributed species, indication that the development of tall shrubland causes significant species loss in the heathland. Italian lowland heathlands are characterized by high rates of shrubland encroachment that threatens both habitat and species diversity. Burning frequencies of once in 3–6 years seem appropriate in this habitat, but burning alone might not suffice without actions to increase herbivore grazing.     

Does selective logging change ground‐dwelling beetle assemblages in a subtropical broad‐leafed forest of China?

Selective logging with natural regeneration is advocated as a near‐to‐nature strategy and has been implemented in many forested systems during the last decades. However, the efficiency of such practices for the maintenance of forest species are poorly understood. We compared the species richness, abundance and composition of ground‐dwelling beetles between selectively logged and unlogged forests to evaluate the possible effects of selective logging in a subtropical broad‐leafed forest in southeastern China. Using pitfall traps, beetles were sampled in two naturally regenerating stands after clearcuts (ca. 50 years old, stem‐exclusion stage: selectively logged 20 years ago) and two mature stands (> 80 years old, understory re‐initiation stage: selectively logged 50 years ago) during 2009 and 2010. Overall, selective logging had no significant effects on total beetle richness and abundance, but saproxylic species group and some abundant forest species significantly decreased in abundance in selectively logged plots compared with unlogged plots in mature stands. Beetle assemblages showed significant differences between selectively logged and unlogged plots in mature stands. Some environmental characteristics associated with selective logging (e.g., logging strategy, stand age, and cover of shrub and moss layers) were the most important variables explaining beetle assemblage structure. Our results conclude that selective logging has no significant impacts on overall richness and abundance of ground‐dwelling beetles. However, the negative effects of selective logging on saproxylic species group and some unlogged forest specialists highlight the need for large intact forested areas for sustaining the existence of forest specialist beetles.