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Article

Response of Forest Bird Communities to Managed Landscapes in the Acadian Forest

1
Great Lakes Forestry Centre, Canadian Forest Service, Natural Resources Canada, 1219 Queen St. E., Sault Ste. Marie, ON P6A 2E5, Canada
2
Independent Researcher, Fredericton, NB E3B 6Y2, Canada
3
GWA Forestry and Applied Biosciences Consulting, 5 Rockridge Dr., Sussex Corner, Sussex, NB E4E 5R2, Canada
*
Author to whom correspondence should be addressed.
Retired.
Forests 2024, 15(1), 184; https://doi.org/10.3390/f15010184
Submission received: 21 November 2023 / Revised: 8 January 2024 / Accepted: 11 January 2024 / Published: 17 January 2024
(This article belongs to the Special Issue Forest Biodiversity Conservation)

Abstract

:
The loss of mature forests is a known stressor of forest management on biodiversity. Mature forests provide unique habitat for forest birds. Here, we examine the capacity of mature forest stands embedded in an intensively managed landscape to provide habitat for landbird species that are associated with mature, unfragmented habitats. We carry this out by comparing bird communities in forest stands in three landscapes with a gradient of management activity. We examined community-level indicators (richness, diversity, abundance and community structure), and trait-level indicators (species groups associated with cavity nesting, mature forests, interior forests and area sensitivity). We found no obvious negative effects on bird communities, species and trait groups in forest stands in the most intensively managed landscape relative to the less intensively managed landscapes. Our ability to draw inferences about the influence of management intensity is limited due to lack of replication; however, these results do provide evidence that mature forest stands within intensively managed landscapes can provide valuable habitat to mature forest associates. There are often trade-offs between generating wood products from the forest and the provision of mature forest habitats. Research on forest birds can provide some of the necessary information for assessing the size and shape of those trade-offs and help to inform the conversation about the desired structure, function and composition of forests.

1. Introduction

Forest management is often criticized for having negative impacts on forest health, forest biodiversity and forest landbirds in particular. Increasing the intensity of management is often associated with increasing impact, with implications for biodiversity decline worldwide [1,2]. There has been significant debate on the role of managed landscapes in supporting natural forest biodiversity [3,4,5]. Landscape-scale effects can be large drivers of community change at stand scales [6]. The impact of landscape on local scales in forested landscapes without land conversion is a matter of debate, however [7,8,9]. Much of the often-cited research on landscape-scale effects was conducted in deforested landscapes where forest patches are embedded in a sea of agriculture or non-forest [10,11,12,13]. In this study, we examine the evidence that in managed forested landscapes without significant land use change, local stands can continue to provide habitat for a range of forest birds, including those that are associated with older forests. We examine landbird communities in forested landscapes with limited land use change, but with intensive management, reflected by a high percentage of planted stands.
Sustainability is a widely held goal of forest management in Canada, including respecting the principles of conservation and biodiversity [14,15], and, in some cases, a legislated mandate [16]. Assessing sustainability is an extremely complex task [8], but one common approach is the use of indicators of sustainability, especially biodiversity indicators [17]. Birds have long been suggested as a good indicator of environmental sustainability [18,19] and forest sustainability in particular [18,20,21]. There are well-established standardized methods of measurement of the relative abundance and distribution of birds [22,23]. In particular, habitat specialists can be informative because they can be most sensitive to specific stressors to the system [24,25]. Here we focus on old forest associates.
Autonomous recording technology for collecting auditory data is also facilitating the collection of significantly more bird abundance data [26,27,28,29]. Auditory methods permit the collection of data for a large diversity of landbird species at the same time. This diversity provides increased potential to detect change in the bird community due to the large variety of habitats and resource use by distinct species. Forest birds also represent a fairly high trophic level, and, as such, they are integrators of the forest system and respond to multiple changes in the ecosystem. In addition, we know a lot about the life history of forest birds due to a long history of research and interest (Birds of the World online, https://birdsoftheworld.org/bow/home, accessed on 15 October 2023). This knowledge allows us to interpret changes in bird communities based on their functional traits that can link changes directly to potential stressors in the forest. Lastly, there is extensive bird data collection across provinces and territories through the Breeding Bird Atlas program (e.g., [30,31]) and the United States Geological Survey’s Breeding Bird Survey (BBS; [32]).
The objective of this study is to compare landbird communities in terms of richness, diversity, abundance, community structure and specific abundance in relation to functional traits in mature forest stands in three landscapes with different levels of management intensity. Bird communities in these landscapes have been studied for many years, and the landscape is known to have a high diversity of forest birds [33,34,35,36]. We were interested in understanding the ecological value of mature forests in the context of intensive forest management. This is a landscape-scale investigation in that birds were measured in matched forest stands in all three landscapes that were comparable in terms of ecosystem characteristics but differed in management intensity. We had no a priori expectation for the comparison of richness, diversity and abundance overall, but we hypothesized that if management intensity was impacting mature forest habitat at landscape scales, we would see community composition differences between landscapes. Specifically, we anticipated that cavity nesters, mature and overmature habitat associates, interior habitat associates and area-sensitive species would all be less common in the mature habitat of the most intensively managed landscape. We are aware that our capacity for statistical inference is limited here due to the lack of replication [37], although parsimony allows us to draw some reasonable conclusions about the value of older forest patches in intensively managed landscapes. Landscapes were chosen for their ecological similarity, all within the Acadian Forest. Landscape replication is often prohibitively difficult and expensive, and we must draw conclusions from more limited but still valuable data. An assessment of the ecological value of mature forests in management contexts will improve our ability to manage forests sustainably by allowing us to better understand and map elements of the ecological value of these mature forest stands.

2. Methods

2.1. Study Area

The three landscapes, Black Brook (BBRO), Quisibis (QUIS) and Debouille (DEBU), are all located within 100 km of each other in the province of New Brunswick, Canada, and the state of Maine, United States of America (Figure 1), all within the Acadian Temperate Forest [38], which covers an area including the Maritime provinces of Canada as well as the northeastern United States. Forest canopies are coniferous (e.g., balsam fir (Abies balsamea), white spruce (Picea glauca), red spruce (Picea rubens), black spruce (Picea mariana), eastern white cedar (Thuja occidentalis), deciduous (e.g., sugar maple (Acer saccharum), red maple (Acer rubrum), white birch (Betula papyrifera), yellow birch (Betula alleghaniensis), American beech (Fagus grandifolia) or a mixture). Anthropogenic disturbance is widespread in the Acadian Forest and includes forest harvesting, road building, agricultural conversion, rail and utility corridors and urban development. Windthrow, ice loading and insect infestations are the most widespread forms of natural disturbance. Wildfire is generally not a factor in the study areas [38]. In the study areas, the disturbances are primarily due to roads and harvest. Understory structure varies from dense to sparse and includes cold-deciduous broad-leaved shrubs, perennial herbs, tree regeneration and bryophytes.
BBRO (209,679 ha) is land owned and managed by J.D. Irving, Limited since the mid-1940s. The planting of stands began in the late 1950s. Twenty-five-year forest management plans are in place, which are revised on a five-year interval. These plans include multiple strategies and objectives related to sustainable wood supplies to support mills and communities; maintaining environmental quality, including water, site productivity and biological diversity; maintaining forest health; supporting non-timber use; and public accountability through third-party certification. The QUIS forest (37,294 ha) is provincially owned Crown land, also managed under a 25-year management plan according to strategies and objectives of the Province of New Brunswick (Crown Lands and Forests Act) and managed by an industrial licensee (part of Crown License 10, Twin Rivers Paper Company). The DEBU Township Forest (8489 ha) is managed for timber production and was acquired by the State of Maine in 1975. The area was first logged in the 1800s, heavily logged in the 1950s and 1960s and affected by the spruce budworm outbreak in the 1970s and 1980s and includes a nearly 364 ha ecological reserve. Timber management has been ongoing, with harvest levels at half of the calculated sustainable limit. The Deboullie Forest is 33% softwood, 25% hardwood and 42% mixedwood, with a general lack of stems under 10 m noted in the management plan.
GIS analysis was performed on forest resource inventories from each landowner along with forest management GIS updates to assess 5 landscape indicators of management activity, including area planted, area in softwood regeneration, area thinned in the last 10 years, area operated for hardwood and the amount of mature forest. BBRO had the highest area planted by a large margin (Table 1), the highest area thinned and the highest area of softwood regeneration and the least amount of mature forest and area of hardwood operated. QUIS and DEBU were more similar. Based on these values, we identified the BBRO landscape as the most intensively managed followed by QUIS and then DEBU. Primarily though, the large percentage of area planted in BBRO relative to the other forests is the most significant indicator of the amount of management activity.

2.2. Bird Sampling

All bird sampling was conducted in 2018. We identified 6 forest types to sample (Table 2) and attempted to sample 7 stands in each forest type for a total of 42 stands per landscape. We struggled to identify enough stands in the DEBU landscape that met size (at least 5 ha) and spatial (at least 100 m to all stand edges from the sampling location and at least 250 m from the next sampling location) requirements but were able to sample at least two stands in each forest type (Table 2). Stands ranged from 5 ha to approximately 100 ha. In each selected forest stand, we placed an autonomous recording unit (Song Meter 2, 3 or 4 from Wildlife Acoustics, Maynard, MA, USA) at a sampling location for a minimum of 20 days between June 1 and June 30 and recorded in stereo each day for 80 min, 10 min each hour from 30 min before sunrise to the 8th hour after sunrise. From the recordings, we chose the first 4 days where the recordings were free from noise (wind, rain and industrial) and interpreted two 5 min recordings from each day, one 20–30 min before sunrise and one within an hour after sunrise. Interpretation of recordings involves listening to the recording while examining a spectrogram and identifying all individuals heard by species. It is possible to identify multiple individuals of the same species on recording through the identification of counter singing with the use of stereo microphones. This gives a total of 8 recordings (40 min) per sampling location in the breeding season. Estimates of bird abundance in samples were generated using the maximum abundance for each species in the 8 recordings in each sampling location.

2.3. Analysis

We performed analyses in R version 4.2.1 (2022-06-23). We compared mean richness, diversity and abundance between landscapes and forest types using a one-way ANOVA and Tukey’s multiple comparisons (α = 0.05). Richness was estimated using the Margalef richness index [39], which controls for effort (number of samples). Diversity was measured using the Shannon diversity index [40]. We also conducted non-metric multidimensional scaling (metaMDS), PERMANOVA (Adonis II) and comparisons of Beta dispersion (Betadisp), all using the vegan package in R [41], to visually compare landscape types based on bird community structure and to compare bird community centroids and dispersion between landscapes (α = 0.05). Lastly, we grouped species by migration strategy, nesting strategy, habitat association, successional stage association, affinity for interior forest and area sensitivity (Table A1) and compared total abundance within groups between landscapes.

3. Results

3.1. Species Presence, Richness, Diversity and Abundance

Excluding rare species with fewer than five total observations, Scarlet Tanager was the only species not found in BBRO samples but present in QUIS (12 times) and DEBU (6 times). There were seven species absent from DEBU samples, excluding rare species (<5), American Bittern, American Crow, American Goldfinch, Bay-breasted Warbler, Evening Grosbeak, Fox Sparrow and White-winged Crossbill (Table A2). There were three species absent from QUIS samples: Alder Flycatcher, Mourning Warbler and Northern Flicker. The 10 most common species of each of the three landscapes had a high degree of overlap and included Ovenbird, Swainson’s Thrush, White-throated Sparrow, Ruffed Grouse, Hermit Thrush, Yellow-bellied Sapsucker and Black-throated Green Warbler (Table A2). We observed more individuals and more species in BBRO than QUIS and DEBU (Table 3).
A one-way ANOVA revealed that there was a statistically significant difference in the richness index, Shannon diversity and abundance between the landscapes (Table 4). Tukey’s HSD test for multiple comparisons supported that these differences are primarily between the BBRO and DEBU landscapes, with BBRO having higher richness, diversity and abundance (Table 3 and Table 4 and Figure 2). The data met the assumptions of normality, homoscedasticity of variance (assessed through visual examination of data) and independence (observations were spaced to maintain independence).

3.2. Community Composition

The non-metric multidimensional scaling (NMDS) results point to considerable overlap between landscape polygons, suggesting limited differences in bird community composition between the selected stands of landscapes (Figure 3). The Permanova test showed a significant difference at p < 0.05 for the comparison of BBRO with QUIS and DEBU but with very little variance explained. The beta dispersion paired comparisons showed no significant differences in community variance between BBRO and other landscapes (Table 5), suggesting that the variance between landscapes is homogeneous and, therefore, that the comparison of centroids is robust.

3.3. Bird Functional Trait Analysis

We saw no evidence of significant differences between the abundance of cavity nesters, mature forest associates, interior species and area-sensitive species in the three landscapes (Figure 4a–d). Overmature forest associates were too infrequent to assess (Figure 4b). In general, none of the classifications demonstrated obvious differences between landscapes and between functional classifications (Figure 4a–f), including habitat association and migration strategy (Figure 4e–f).

4. Discussion

We saw no obvious negative effects of greater management intensity of the BBRO landscapes at the stand scale. The effect of landscape-level influence on stand-level community composition and species abundance has been hypothesized for a long time. Landscape ecology theory predicts the existence of threshold levels of habitat in landscapes required to maintain species because of decreasing colonization rates and in increasing local extinction rates [43]. This effect has been clearly demonstrated in deforested landscapes, in particular in agricultural contexts [44,45,46]. The results for landscape-level effects have always been more equivocal in forests within forest management contexts [47,48,49]. Some studies have shown important landscape-scale effects on richness in one year only to demonstrate stronger stand-scale effects the next year [49]. Local patch size has also been demonstrated to be an important driver of species abundance, which can be both a local and landscape measure [3]. The low sensitivity of many boreal songbirds to harvesting at a landscape scale has been reported in several studies [50,51,52,53], and it has been suggested by others [7,19,54]. This may be because birds in boreal forests are adapted to frequent disturbances in natural boreal forest landscapes. In addition, though, it may also be because the context for the fragmentation is still forest, where there are gradients of habitat value, soft edges between patches and good connectivity, unlike agricultural areas with hard edges and habitat loss [55]. This is a more likely explanation for the Acadian Forest. Our results indicate a lack of landscape-scale effect that is consistent with other studies in forest management rather than agricultural contexts.
Significantly, there were no differences in the abundance of mature forest associates. We suggest that this should be the most sensitive indicator of landscape effect. The age class distribution of a forested region is one of the key characteristics that dictate the provision of habitat to wildlife [8]. All forest ages provide habitat, but currently in New Brunswick, forest management is reducing the distribution of the mature forest age classes, which could put stress on the species most associated with older age classes [56]. Mature forests provide unique habitat features for birds and other species [57,58,59,60], including larger trees and deadwood, which can support cavity nesters and saproxylics [54,61]. A reduction in the area of older forests can also be associated with shorter rotation planted forests.
There are a few studies on the Acadian Forest that have looked at landscape thresholds in species occurrence, predicting that there may be a threshold of habitat loss at the landscape scale below which ecological processes change abruptly [33,62]. This suggests that landscape-scale effects may not be evident until landscape-level habitat change reaches this threshold. There is some evidence for thresholds at which some species responded to landscape-level habitat change, ranging from 8.6 to 28.7% [33]. These results suggest that effects of landscape context have been identified, although at fairly low levels of landscape habitat. Our results suggest that if a threshold exists, it has not been reached on the BBRO landscape. This result suggests that the relatively high intensity of silviculture in the intensively managed forest landscape is not reducing the ability of mature forests to provide important habitat. This lends confidence to habitat estimates based on the prevalence and distribution of mature forests in ecosystems with forest continuity. Other studies have shown the ecological value of plantations themselves, where they contain diverse avian communities relative to native stands [63] or where they improve connectivity in landscapes [4,64].
The use of indicators that are known to respond to forest management stressors has long been a recommended approach to evaluating forest management practices [18]. This is supported by research that has demonstrated that generalist species are becoming more dominant while specialists are becoming rarer through a process of homogenization in response to human disturbance [24,25]. Our approach examined species with functional traits expected to be most impacted by common hypothesized stressors of intensive forest management. Cavity nesters [65] species that are area-sensitive or, alternatively, species that are associated with interior habitat [11,66] and species associated with mature or overmature habitat are all more likely to be impacted by a landscape-level reduction in older forest through intensive management. One of the key mechanisms that is used to explain this landscape-scale effect is meta-population dynamics [11,67], which suggests that patch occupancy will be higher in landscapes with more habitat or more connectivity [68]. Our results showed no evidence that these indicator groups were responding to the landscape-level management intensity. We speculate that this may be due to very high connectivity in these landscapes that are primarily forested even when mature patches of habitat are dispersed. The data suggest that all of these potentially sensitive specialists can be supported on these highly managed landscapes. This narrowing of the focus to forest birds at the highest risk of turnover or loss in abundance adds additional support to the ecological value of mature forest stands in a planted stand-rich context. We are not suggesting that there are no landscape-scale effects here but only that mature forest stands in BBRO appear to be providing similar habitat quality to mature forests in landscapes with a much smaller area of planted stands.
These results must be interpreted considering several caveats. First, our landscapes represent a gradient in planted stand management intensity (BBRO > QUIS > DEBU), but we do not have replication at the landscape scale, and so the results are confounded with landscape identity and all of the possible differences that might occur between landscapes we did not measure. However, these landscapes occur in a single forest system (the Acadian Forest) and have considerable overlap in bird communities based on the NMDS, suggesting that they represent a common forest system. In addition, the difference in the area of planted stands in these landscapes is so significant that parsimony would suggest that if management intensity was important, it would be visible here. Second, the sample size in the DEBU landscape was small relative to the other landscapes, and so this comparison should hold less weight but is consistent with the interpretation that high management intensity in BBRO is not compromising bird habitat quality. The third caveat is that perhaps the level of landscape management intensity in QUIS and DEBU is already sufficient to impact bird communities in the oldest forest stands. A comparison with an unmanaged landscape would be ideal but was not available in a comparable forest. It may be possible to use more generalized data from the Breeding Bird Survey and Breeding Bird Atlas to understand the pool of species that is expected as a point of comparison or reference condition, but, in the end, the vast majority of these data will come from managed landscapes. The reality that we have a limited selection of primary, or at least mature, unmanaged forest landscapes to act as reference conditions for sustainability assessments is significant and problematic.
An important implication of these results is that mature stands in these landscapes are able to provide mature forest habitat despite their context. The presence of species in these stands is not proof of the ability to breed here. There is a long-standing debate about the relationship between abundance and productivity [69], where some habitats contain sink populations that rely on immigration from outside of the stand to maintain populations [70]. Research in Ontario has demonstrated that many old forest associates do breed successfully in small patches of mature forest in a forest management context [53]. In contrast, there is evidence from areas fragmented by agriculture that reproductive success can be lowered in habitat fragments such that they become population sinks that cannot be maintained without immigration [71]. The BBRO landscape has been intensively managed for 70 years, and the area planted has been stable for the last 10 years, suggesting that these are persistent populations. Haché et al. [35] used demographic data collected from the BBRO forest to model ovenbird population dynamics under forestry as usual and climate change scenarios. They found that ovenbird populations act as demographic sinks in an intensively managed landscape under climate change but that without climate change (i.e., under current conditions), the number of territorial males would remain relatively constant. Given the inevitability of climate change, however, forest management planning and approaches may need to be altered to conserve old forest associates on these managed landscapes into the future.

5. Conclusions

Old forests provide unique habitat for forest birds, and the loss of that habitat in the Acadian Forest of New Brunswick has been linked directly to the long-term decline in old forest associates [56]. There is an inevitable trade-off between generating wood products from the forest and the provision of older forest habitat, although balance may be achievable through natural disturbance-based silvicultural systems that can better support mature forest birds [72]. Research on forest birds can provide some of the necessary information to assess the size and shape of those trade-offs and help to inform the conversation about the desired structure, function and composition of those forests. Specifically, improving our knowledge of the ecological value of mature forest stands in this managed context is an important step in our ability to assess trade-offs and find balance.

Author Contributions

Conceptualization, L.A.V.; Methodology, L.A.V. and K.M.; Investigation, L.A.V., K.P., G.A. and K.M.; Resources, L.A.V.; Data curation, L.A.V., K.P. and K.M.; Writing—original draft, L.A.V.; Writing—review & editing, L.A.V., K.P., G.A. and E.S.; Visualization, K.P. and E.S. All authors have read and agreed to the published version of the manuscript.

Funding

NSERC Collaborative Research and Development Grant no. CRDPJ 495007-16 with JD Irving Limited.

Data Availability Statement

Data generated or analyzed during this study are available from the corresponding author upon reasonable request.

Acknowledgments

We would like to thank Neil Thomson for his support in data collection and insights on the Debouille landscape.

Conflicts of Interest

Greg Adams has retired from his position as science director with JD Irving (owner of the Black Brook landscape). Kevin Porter is a consultant paid by JD Irving. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The JD Irving had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

Appendix A

Table A1. Functional classifications of bird species’ functional trait identities include (1) Migration Strategy: NEO = neotropical migrant, NOM = nomadic, RES = resident and SDM = short-distance migrant; (2) nesting strategy: C = cavity, G = ground, LTC = lower tree canopy, TC = tree canopy, TS = tall shrubs and W = wetland; (3) habitat association: CMI = conifer mixedwood, CON = conifer, HMI = hardwood mixedwood, HWD = hardwood, MIX = mixedwood and OHS = open herb and shrub; (4) successional association: G = generalist, HE = herbs, IM = immature, MA = mature, OM = overmature and TS = tall shrubs; (5) area sensitivity: Y = yes and N = no; (6) interior association: E = edge, I = Interior, I_E = both, S = shore and W = wetland.
Table A1. Functional classifications of bird species’ functional trait identities include (1) Migration Strategy: NEO = neotropical migrant, NOM = nomadic, RES = resident and SDM = short-distance migrant; (2) nesting strategy: C = cavity, G = ground, LTC = lower tree canopy, TC = tree canopy, TS = tall shrubs and W = wetland; (3) habitat association: CMI = conifer mixedwood, CON = conifer, HMI = hardwood mixedwood, HWD = hardwood, MIX = mixedwood and OHS = open herb and shrub; (4) successional association: G = generalist, HE = herbs, IM = immature, MA = mature, OM = overmature and TS = tall shrubs; (5) area sensitivity: Y = yes and N = no; (6) interior association: E = edge, I = Interior, I_E = both, S = shore and W = wetland.
Specie
Code
Common_NameMigration
Strategy
Nesting
Strategy
HabitatSuccessional
Association
Area
Sensitive
Interior
Association
ALFLAlder FlycatcherNEOTall
Shrubs
OHSTall ShrubsNE
AMBIAmerican BitternSDMGroundOHSHerbsNW
AMCRAmerican CrowSDMTree
Canopy
OHSGeneralistNE
AMGOAmerican GoldfinchSDMLower
Tree
Canopy
MIXTall ShrubsNE
AMREAmerican RedstartNEOTall
Shrubs
HMIImmatureYI_E
AMROAmerican RobinSDMLower
Tree
Canopy
MIXGeneralistNE
BAOWBarred OwlRESCavityHMIMatureYI
BAWWBlack-and-white
Warbler
NEOGroundCMIMatureYI
BBCUBlack-billed CuckooNEOLower
Tree
Canopy
HWDImmatureNI_E
BBWABay-breasted WarblerNEOLower
Tree
Canopy
CMIOvermatureNI_E
BBWOBlack-backed
Woodpecker
RESCavityCONMatureYI_E
BCCHBlack-capped
Chickadee
RESCavityMIXGeneralistNI_E
BHVIBlue-headed VireoSDMLower
Tree
Canopy
CMIMatureYI_E
BLBWBlackburnian WarblerNEOTree
Canopy
MIXMatureYI
BLJABlue JaySDMLower
Tree
Canopy
CMIGeneralistNI_E
BLPWBlackpoll WarblerNEOLower
Tree
Canopy
MIXOvermatureNI
BOCHBoreal ChickadeeRESCavityCONMatureNI
BRCRBrown CreeperSDMCavityMIXMatureYI
BTBWBlack-throated Blue
Warbler
NEOTall
Shrubs
HWDImmatureYI
BTNWBlack-throated Green
Warbler
NEOLower
Tree
Canopy
MIXMatureYI
CAWACanada WarblerNEOGroundMIXImmatureYI_E
CEDWCedar WaxwingSDMLower
Tree
Canopy
HMIImmatureNE
CHSPChipping SparrowSDMTall
Shrubs
MIXImmatureNE
CMWACape May WarblerNEOTree
Canopy
MIXMatureNI_E
CONICommon NighthawkNEOGroundOHSHerbsNE
CONWConnecticut WarblerNEOGroundCONOvermatureNI_E
CORACommon RavenRESTree
Canopy
MIXGeneralistNI_E
COYECommon YellowthroatNEOGroundOHSTall ShrubsNE
CSWAChestnut-sided
Warbler
NEOTall
Shrubs
HWDImmatureNE
DEJUDark-eyed JuncoSDMGroundCONImmatureNE
DOWODowny WoodpeckerRESCavityHWDImmatureNI_E
EAKIEastern KingbirdNEOLower
Tree
Canopy
OHSTall ShrubsNE
EAWPEastern Wood-PeweeNEOTree
Canopy
HWDMatureNI_E
EVGREvening GrosbeakNOMTree
Canopy
CONMatureNI_E
FOSPFox SparrowSDMGroundCONImmatureNE
GCFLGreatCrested
Flycatcher
NEOCavityHWDImmatureNI_E
GCKIGolden-crowned
Kinglet
SDMTree
Canopy
CONMatureNI
GHOWGreat Horned OwlRESCavityHMIMatureNI_E
GRAJCanada JayRESLower
Tree
Canopy
CMIOvermatureNI_E
GRHEGreen HeronNEOTall
Shrubs
OHSTall ShrubsNE
HAWOHairy WoodpeckerRESCavityMIXMatureYI
HETHHermit ThrushSDMGroundMIXMatureYI_E
HOWRHouse WrenSDMCavityHWDTall ShrubsNE
LEFLLeast FlycatcherNEOTree
Canopy
HWDImmatureYE
MAWAMagnolia WarblerNEOLower
Tree
Canopy
CONImmatureYI_E
MERLMerlinSDMTree
Canopy
CONGeneralistNI_E
MOWAMourning WarblerNEOGroundHMIImmatureNE
NAWANashville WarblerNEOGroundMIXImmatureNE
NOFLNorthern FlickerSDMCavityMIXImmatureNE
NOPANorthern ParulaNEOTree
Canopy
MIXMatureYI
NOWANorthern WaterthrushNEOGroundCMIMatureNI_E
NSOWNorthern Saw-whet
Owl
RESCavityCMIMatureNI_E
OCWAOrange-crowned
Warbler
SDMGroundCMITall ShrubsNE
OSFLOlive-sided FlycatcherNEOTree
Canopy
CONOvermatureNI_E
OVENOvenbirdNEOGroundHMIMatureYI
PAWAPalm WarblerSDMGroundCONTall ShrubsNE
PHVIPhiladelphia VireoNEOTree
Canopy
HWDImmatureNI_E
PIGRPine GrosbeakNOMLower
Tree
Canopy
MIXMatureNI_E
PIWAPine WarblerSDMTree
Canopy
CONMatureYI
PIWOPileated WoodpeckerRESCavityMIXMatureYI
PUFIPurple FinchSDMTree
Canopy
MIXMatureNI_E
RBGRRose-breasted
Grosbeak
NEOLower
Tree
Canopy
HMIImmatureNI_E
RBNURed-breasted
Nuthatch
RESCavityCMIMatureYI
RCKIRuby-crowned KingletSDMTree
Canopy
CONMatureNI_E
REVIRed-eyed VireoNEOLower
Tree
Canopy
HWDMatureNI_E
RTHURuby-throated
Hummingbird
NEOLower
Tree
Canopy
HWDGeneralistNE
RUBLRusty BlackbirdSDMLower
Tree
Canopy
MIXGeneralistNE
RUGRRuffed GrouseRESGroundHMIGeneralistNI_E
RWBLRed-winged BlackbirdSDMWetlandOHSTall ShrubsNW
SCTAScarlet TanagerNEOTree
Canopy
HWDMatureYI
SPSASpotted SandpiperNEOGroundOHSHerbsNS
SWTHSwainson’s ThrushNEOLower
Tree
Canopy
MIXMatureNI
TEWATennessee WarblerNEOGroundHMIImmatureNI_E
TRESTree SwallowSDMCavityOHSImmatureNE
VEERVeeryNEOTall
Shrubs
HMIImmatureYI
WAVIWarbling VireoNEOTree
Canopy
HWDImmatureNE
WBNUWhite-breasted
Nuthatch
RESCavityHWDMatureYI
WISNWilson’s SnipeSDMGroundOHSHerbsNW
WIWAWilson’s WarblerNEOGroundOHSTall ShrubsNE
WIWRWinter WrenSDMCavityCMIMatureYI
WOTHWood ThrushNEOLower
Tree
Canopy
MIXMatureNI_E
WTSPWhite-throated
Sparrow
SDMGroundMIXGeneralistNI_E
WWCRWhite-winged
Crossbill
NOMTree
Canopy
MIXMatureNI_E
YBFLYellow-bellied
Flycatcher
NEOGroundCONOvermatureNI_E
YBSAYellow-bellied
Sapsucker
SDMCavityMIXMatureYI_E
YEWAYellow WarblerNEOTall
Shrubs
HWDTall ShrubsNE
YRWAYellow-rumped
Warbler
SDMTree
Canopy
MIXMatureNI_E
Table A2. Bird species lists including common name, scientific name and abundance in each landscape.
Table A2. Bird species lists including common name, scientific name and abundance in each landscape.
Species CodeCommon NameScientific NameBBROQUIS DEBU
Abundance
ALFLAlder FlycatcherEmpidonax alnorum1105
AMBIAmerican BitternBotaurus lentiginosus430
AMCRAmerican CrowCorvus brachyrhynchos950
AMGOAmerican GoldfinchSpinus tristis9110
AMREAmerican RedstartSetophaga ruticilla2173
AMROAmerican RobinTurdus migratorius524114
BAOWBarred OwlStrix varia233
BAWWBlack-and-white WarblerMniotilta varia624
BBCUBlack-billed CuckooCoccyzus erythropthalmus200
BBWABay-breasted WarblerSetophaga castanea750
BBWOBlack-backed WoodpeckerPicoides arcticus131
BCCHBlack-capped ChickadeePoecile atricapillus46314
BHVIBlue-headed VireoVireo solitarius40338
BLBWBlackburnian WarblerSetophaga fusca30268
BLJABlue JayCyanocitta cristata29268
BLPWBlackpoll WarblerSetophaga striata200
BOCHBoreal ChickadeePoecile hudsonicus554
BRCRBrown CreeperCerthia americana222310
BTBWBlack-throated Blue WarblerSetophaga caerulescens424717
BTNWBlack-throated Green WarblerSetophaga virens565119
CAWACanada WarblerCardellina canadensis1252
CEDWCedar WaxwingBombycilla cedrorum010
CHSPChipping SparrowSpizella passerina352
CMWACape May WarblerSetophaga tigrina771
CONICommon NighthawkChordeiles minor310
CONWConnecticut WarblerGeothlypis agilis100
CORACommon RavenCorvus corax8233
COYECommon YellowthroatGeothlypis trichas302
CSWAChestnut-sided WarblerSetophaga pensylvanica426
DEJUDark-eyed JuncoJunco hyemalis275
DOWODowny WoodpeckerDryobates pubescens101
EAKIEastern KingbirdTyrannus tyrannus001
EAWPEastern Wood-PeweeContopus virens12115
EVGREvening GrosbeakCoccothraustes vespertinus630
FOSPFox SparrowPasserella iliaca810
GCFLGreat Crested FlycatcherMyiarchus crinitus001
GCKIGolden-crowned KingletRegulus satrapa373510
GHOWGreat Horned OwlBubo virginianus500
GRAJGray JayPerisoreus canadensis1061
GRHEGreen HeronButorides virescens110
HAWOHairy WoodpeckerDryobates villosus210
HETHHermit ThrushCatharus guttatus676518
HOWRHouse WrenTroglodytes aedon010
LEFLLeast FlycatcherEmpidonax minimus8129
MAWAMagnolia WarblerSetophaga magnolia412512
MERLMerlinFalco columbarius200
MOWAMourning WarblerGeothlypis philadelphia205
NAWANashville WarblerOreothlypis ruficapilla422110
NOFLNorthern FlickerColaptes auratus801
NOPANorthern ParulaSetophaga americana535014
NOWANorthern WaterthrushParkesia noveboracensis2423
NSOWNorthern Saw-whet OwlAegolius acadicus010
OCWAOrange-crowned WarblerOreothlypis celata100
OSFLOlive-sided FlycatcherContopus cooperi1334
OVENOvenbirdSeiurus aurocapilla728528
PAWAPalm WarblerSetophaga palmarum310
PHVIPhiladelphia VireoVireo philadelphicus8102
PIGRPine GrosbeakPinicola enucleator100
PIWAPine WarblerSetophaga pinus100
PIWOPileated WoodpeckerDryocopus pileatus142
PUFIPurple FinchHaemorhous purpureus9184
RBGRRose-breasted GrosbeakPheucticus ludovicianus20193
RBNURed-breasted NuthatchSitta canadensis555912
RCKIRuby-crowned KingletRegulus calendula35246
REVIRed-eyed VireoVireo olivaceus304619
RTHURuby-throated HummingbirdArchilochus colubris100
RUBLRusty BlackbirdEuphagus carolinus400
RUGRRuffed GrouseBonasa umbellus576132
RWBLRed-winged BlackbirdAgelaius phoeniceus100
SCTAScarlet TanagerPiranga olivacea0126
SPSASpotted SandpiperActitis macularius110
SWTHSwainson’s ThrushCatharus ustulatus827825
TEWATennessee WarblerOreothlypis peregrina591
TRESTree SwallowTachycineta bicolor010
VEERVeeryCatharus fuscescens21147
WAVIWarbling VireoVireo gilvus000
WBNUWhite-breasted NuthatchSitta carolinensis001
WISNWilson’s SnipeGallinago delicata411
WIWAWilson’s WarblerCardellina pusilla100
WIWRWinter WrenTroglodytes hiemalis494514
WOTHWood ThrushHylocichla mustelina050
WTSPWhite-throated SparrowZonotrichia albicollis946128
WWCRWhite-winged CrossbillLoxia leucoptera9120
YBFLYellow-bellied FlycatcherEmpidonax flaviventris37196
YBSAYellow-bellied SapsuckerSphyrapicus varius596028
YEWAYellow WarblerSetophaga petechia100
YRWAYellow-rumped WarblerSetophaga coronata132210

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Figure 1. Study areas. The three landscapes, Black Brook (BBRO), Quisibis (QUIS) and Debouille (DEBU), blue polygons, are all located within 100 km of each other in the province of New Brunswick, Canada, and the state of Maine, United States of America. QUIS polygon was drawn using a fixed-width buffer, creating an artificial overlap with BBRO. Bird survey sites located within the BBRO polygon contributed only to the BBRO data.
Figure 1. Study areas. The three landscapes, Black Brook (BBRO), Quisibis (QUIS) and Debouille (DEBU), blue polygons, are all located within 100 km of each other in the province of New Brunswick, Canada, and the state of Maine, United States of America. QUIS polygon was drawn using a fixed-width buffer, creating an artificial overlap with BBRO. Bird survey sites located within the BBRO polygon contributed only to the BBRO data.
Forests 15 00184 g001
Figure 2. Boxplot comparison of (a) Margalef richness index per sampled stand, (b) Shannon diversity index and (c) abundance for the 3 landscapes (BBRO = Black Brook, QUIS = Quisibis and DEBU = Debouille). Data points are represented by black dots.
Figure 2. Boxplot comparison of (a) Margalef richness index per sampled stand, (b) Shannon diversity index and (c) abundance for the 3 landscapes (BBRO = Black Brook, QUIS = Quisibis and DEBU = Debouille). Data points are represented by black dots.
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Figure 3. Non-metric multidimensional scaling (NMDS) plot of bird communities in the 3 landscapes. The overall stress value for the NMDS is 0.20. This level of stress suggests that too much reliance should not be placed on the details of the plot but that the overall plot can still correspond to a usable picture [42]. The non-metric R-squared is 0.96. Centroids of polygons are displayed as stars.
Figure 3. Non-metric multidimensional scaling (NMDS) plot of bird communities in the 3 landscapes. The overall stress value for the NMDS is 0.20. This level of stress suggests that too much reliance should not be placed on the details of the plot but that the overall plot can still correspond to a usable picture [42]. The non-metric R-squared is 0.96. Centroids of polygons are displayed as stars.
Forests 15 00184 g003
Figure 4. Comparison of landscapes according to functional traits for abundance (abun). Functional classifications include (a) nesting strategy: C = cavity, G = ground, LTC = lower tree canopy, TC = tree canopy, TS = tall shrubs and W = wetland; (b) successional associates: G = generalist, HE = herbs, IM = immature, MA = mature, OM = overmature and TS = tall shrubs; (c) interior associates: E = edge, I = interior, I_E = both, S = shore and W = wetland; (d) area sensitive: Y = yes and N = no; (e) habitat: CMI = conifer mixedwood, CON = conifer, HMI = hardwood mixedwood, HWD = hardwood, MIX = mixedwood and OHS = open herb and shrub; (f) migration strategy: NEO = neotropical migrant, NOM = nomadic, RES = resident and SDM = short-distance migrant. Box and whisker plots display quartiles and median values.
Figure 4. Comparison of landscapes according to functional traits for abundance (abun). Functional classifications include (a) nesting strategy: C = cavity, G = ground, LTC = lower tree canopy, TC = tree canopy, TS = tall shrubs and W = wetland; (b) successional associates: G = generalist, HE = herbs, IM = immature, MA = mature, OM = overmature and TS = tall shrubs; (c) interior associates: E = edge, I = interior, I_E = both, S = shore and W = wetland; (d) area sensitive: Y = yes and N = no; (e) habitat: CMI = conifer mixedwood, CON = conifer, HMI = hardwood mixedwood, HWD = hardwood, MIX = mixedwood and OHS = open herb and shrub; (f) migration strategy: NEO = neotropical migrant, NOM = nomadic, RES = resident and SDM = short-distance migrant. Box and whisker plots display quartiles and median values.
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Table 1. Landscape-scale measures of management activity taken from aerially interpreted forest resource inventory polygons.
Table 1. Landscape-scale measures of management activity taken from aerially interpreted forest resource inventory polygons.
ManagementBBROQUISDEBU
% Area planted to conifer (of total landbase)41.810.20
% Area softwood regeneration (of total landbase)4.41.03
% Area thinned in last 10 years (of total landbase)8.30.60
% Hardwood Area operated (of total hardwood area)21.121.832
% Mature forest (of total landbase)34.039.445.7
Table 2. The number of stands sampled in six forest types in this study (classified by stand attributes in GIS).
Table 2. The number of stands sampled in six forest types in this study (classified by stand attributes in GIS).
Forest TypeBBROQUISDEBU
Cedar Mature/Over Mature 1772
Mixedwood Mature/Over Mature 2773
Spruce Fir (16–45 years old) 3773
Spruce Fir Mature/Over Mature 4772
Tolerant Hardwood Mature–Over Mature–No Recent Harvest 5763
Tolerant Hardwood Mature–Over Mature–Selection Harvest 6772
Total424115
1 Cedar Mature/Over Mature: Eastern cedar-dominated stands, greater than 65 years old. Mature and overmature stands may have undergone selection harvest that favours cedar for retention and removes other softwood species unlikely to live another 25 years. 2 Mixedwood Mature/Over Mature: Natural stands with tolerant hardwoods and intolerant hardwoods mixed with softwood species that are older than 65 years and have had no silvicultural interventions. 3 Spruce Fir (16–45 years old): Natural spruce/fir forests that are 16 to 45 years old and have had no silvicultural interventions. 4 Spruce Fir Mature/Over Mature: Natural spruce/fir forests composed of white, red and/or black spruce and/or balsam fir. Stands are older than 45 years and have had no silvicultural interventions. 5 Tolerant Hardwood Mature–Over Mature–No Recent Harvest: Natural stands of tolerant hardwoods (sugar maple, yellow birch, American beech and red maple) that are older than 75 years and have had no silvicultural interventions. 6 Tolerant Hardwood Mature–Over Mature–Selection Harvest: Natural stands of tolerant hardwoods that are older than 75 years. Stands have typically undergone single tree selection or gap harvesting favouring healthy, vigorous stems conducive to selection harvest regimes (i.e., multiple entries in perpetuity).
Table 3. Bird richness, abundance and diversity summaries with standard deviation (SD) for 3 landscapes in the Acadian Forest in 2018.
Table 3. Bird richness, abundance and diversity summaries with standard deviation (SD) for 3 landscapes in the Acadian Forest in 2018.
Total Number of SpeciesNo. of Sampled StandsAbundanceMean Margalef Richness Per Sampled Stand (SD)Mean Abundance Per Sampled Stand (SD)Mean Shannon Diversity Per Sampled Stand
(SD)
All Landscapes 869931936.0131.16
BBRO774214566.28 (0.84)34.7 (7.1)3.03 (0.17)
QUIS654112785.95 (0.86)31.2 (7.4)2.96 (0.19)
DEBU56164595.48 (0.75)28.7 (5.8)2.83 (0.16)
Table 4. Results of one-way ANOVA and Tukey’s HSD test of multiple comparisons for the three landscapes. The F statistic is reported with degrees of freedom in brackets.
Table 4. Results of one-way ANOVA and Tukey’s HSD test of multiple comparisons for the three landscapes. The F statistic is reported with degrees of freedom in brackets.
Comparison RichnessShannon Diversity Abundance
Overall (one-way ANOVA)F(2, 96) = 5.634, p = 0.0049F(2, 96) = 6.345,
p = 0.0026
F(2, 96) = 5.017,
p = 0.0085
BBRO vs. QUIS (Tukey’s HSD test for multiple comparisons)p = 0.17p = 0.15p = 0.06
QUIS vs. DEBU (Tukey’s HSD test for multiple comparisons)p = 0.13 p = 0.096p = 0.45
BBRO vs. DEBU (Tukey’s HSD test for multiple comparisons)p = 0.004p = 0.002p = 0.013
Table 5. Pairwise comparisons of centroid and dispersion of the communities from the 3 landscapes.
Table 5. Pairwise comparisons of centroid and dispersion of the communities from the 3 landscapes.
ComparisonsDfPermanova FPermanova pR-SquaredBeta Dispersion FBeta Dispersion p
BBRO: QUIS13.530.0030.0410.8390.362
BBRO: DEBU13.460.0010.0582.970.090
QUIS: DEBU11.590.10.0284.500.038
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Venier, L.A.; Porter, K.; Adams, G.; McIlwrick, K.; Smenderovac, E. Response of Forest Bird Communities to Managed Landscapes in the Acadian Forest. Forests 2024, 15, 184. https://doi.org/10.3390/f15010184

AMA Style

Venier LA, Porter K, Adams G, McIlwrick K, Smenderovac E. Response of Forest Bird Communities to Managed Landscapes in the Acadian Forest. Forests. 2024; 15(1):184. https://doi.org/10.3390/f15010184

Chicago/Turabian Style

Venier, Lisa A., Kevin Porter, Gregory Adams, Kenneth McIlwrick, and Emily Smenderovac. 2024. "Response of Forest Bird Communities to Managed Landscapes in the Acadian Forest" Forests 15, no. 1: 184. https://doi.org/10.3390/f15010184

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