Breeding occupancy and success of two hole-nesting passerines: the impact of fragmentation caused by forestry

We studied the distribution and the reproductive success of the pied flycatcher Ficedula hypoleuca and the redstart Phoenieurus phoenicurus in relation to forest patch size, edge type (clearcut vs natural), distance from the forest edge, and vegetation characteristics in a forest-dominated landscape. Breeding performances were recorded in up to 72 forest patches during 1992–1994 for birds breeding in nest-boxes. In the spring, breeding individuals of both species arrived earlier in large forest patches (> 1 ha) than in smaller ones. Pied flycatchers arrived earlier on clearcut edges than natural edges but in the redstart there was no preference for a particular edge type. The territory distance from the forest to open land edge did not affect the arrival dates of either species. In the case of the pied flycatcher, the proportion of unpaired males was highest in patches < 1 ha in size and in the case of the redstart this applied to patches < 5 ha in size. Pairing success was not related to the forest edge type or the nest's distance from the edge. Nest predation was not patch-size nor edge-related for either of the species, but in the combined data for both species nest predation was higher at clearcut edges than at natural edges. Clutch size, brood size and the survival of nestlings to the fiedgling stage (fledgling/egg. %) were independent of the patch size, edge type and nest's "distance from the forest edge.     

The breeding ecology and behaviour of Wrens Troglodytes troglodytes under primaeval and secondary conditions

The breeding habits of Wrens inhabiting the primaeval forest of Biatowieza National Park, Poland, are described and compared with those of Wrens living in man-transformed habitats of Western Europe.
Significant inter-habitat differences in numbers exist in Biatowieza. On average there are 4-3 territories per 10 ha in ash-alder, 1 -7 territories per 10 ha in oak–hornbeam and 0–4 territories per 10 ha in coniferous habitats. In ash–alder stands the males have smaller territories, build fewer nests and have better pairing success than in oak–hornbeam stands. However, these two habitats do not differ significantly in average clutch size, timing of breeding or nesting success. Over 80% of nests used for breeding in Biatowieza are situated in uprooted trees. Inter-habitat differences in bird numbers reflect differences in distribution and density of such trees.
Low numbers of Biatowieza Wrens cannot be accounted for solely by high mortality away from the breeding grounds. Available data strongly suggest that numbers of both sexes are proximately limited by behavioural interactions on the breeding ground.
In west European Wrens, densities are higher, territories smaller, polygyny more frequent, clutches smaller and production of young higher than in Biatowieza. Differences in production of young result from different rates of nest predation. Indirect evidence suggests that the breeding habits of Wrens once living in the primaeval forests of western Europe were similar to those of Biatowieza birds, and that the differences observed nowadays result to a large extent from man-made transformations of habitat. It is suggested that man's lowering of the predation pressure acting both on proximate (enabling birds to produce more young) and ultimate levels (decreasing relative advantages of dispersed breeding) was the decisive factor allowing an increase in numbers of Wrens in secondary habitats.     

Reproductive success of Tree Pipits Anthus trivialis in relation to habitat selection in conifer plantations

Aspects of the reproductive success of Tree Pipits Anthus trivialis were examined in relation to broad‐scale habitat and nest‐site selection in Thetford Forest, a coniferous plantation forest in eastern England. Three habitat classes were defined corresponding to previously reported densities of Tree Pipits: clearfell and recently planted stands (habitat class A: low density), stands 2–5 years old (B: high density) and stands 6 years or older (C: low density). The preference for 2–5‐year‐old stands indicated by higher densities was supported by the timing of territory settlement. Tree Pipits also showed distinct preferences for nest‐site characteristics that were relatively consistent across habitat classes and throughout the breeding season. At the ‘habitat scale’, results were consistent with the predictions of the ideal despotic distribution model. First clutches were laid significantly earlier in the preferred habitat class B. Overall nesting success (i.e. the proportion of nests producing fledglings), but not clutch size, also varied between habitats, being greater in habitat classes B and C than in habitat class A. The variation in overall nesting success between habitats was primarily driven by low nest survival rates during the laying/incubation period in clearfell and recently planted stands. Nest survival rates during the nestling period were lower in the preferred 2–5‐year‐old (and older) stands and declined over the course of the study. Preferences for nest‐site characteristics (at least for those that were measured) provided no apparent benefit to nest survival rates. Overall nesting success thus appeared to be determined at the habitat scale, perhaps because the broad differences in cover between habitats affected the likelihood of nest predation (the main cause of nest failure). It is suggested that the very low nesting success experienced by Tree Pipits in clearfell and new stands may be one factor in the species’ relative avoidance of this habitat and preference for 2–5‐year‐old stands.     

Mating success of the endemic Des Murs’ Wiretail (Sylviorthorhynchus desmursii,Furnariidae) in fragmented Chilean rainforests

Abstract: We studied the effects of fragment size, vegetation structure and presence of habitat corridors on the reproductive success of the Des Murs’ Wiretail (Sylviorthorhynchus desmursii Des Murs, Furnariidae), a small (10 g) understorey bird, endemic to South American forests. In a rural landscape of Chiloé Island, southern Chile (42°S; 70°W), we determined the mating and nesting success of wiretails in 28 territories distributed in seven small (1–20 ha) and two large (>300 ha) forest fragments during the 1997–1998 breeding season. Wiretails inhabited dense bamboo thickets in the understorey of forest patches, dense shrublands covering old fields, and dense early successional forest vegetation. Wiretails avoided open pastures. Reproductive success depended solely on the probability of finding mates, and the main factor affecting mating success was the presence of corridors. Mated individuals occupied 72% of the territories in forest patches <20 ha connected by corridors, 73% of the territories in large (>300 ha) fragments, but only 20% of territories in isolated fragments surrounded by pastures. Because of the rapid expansion of pastures in southern Chile, the conservation of wiretails and other understorey birds will depend on the maintenance of travel corridors with dense understorey vegetation between forest fragments.     

Evidence of a Minimum Patch Size Threshold of Reproductive Success in an Endangered Songbird

ABSTRACT That area-sensitive songbirds breed only in relatively large patches suggests that there may be a minimum patch size threshold in which they will breed, even when controlling for the total amount of habitat in the landscape. We searched for minimum patch size thresholds of presence, territory establishment by males, pairing success, and reproductive success for 2 migratory songbirds that differed in sensitivity to patch size: golden-cheeked warblers (Dendroica chrysoparia) and white-eyed vireos (Vireo griseus). We assessed 2 potential limiting factors: brown-headed cowbird (Molothrus ater) parasitism and arthropod biomass (food resource). We determined whether either factor was related to patch size and compared measurements of each above and below the observed thresholds. We monitored 24 golden-cheeked warbler and 47 white-eyed vireo territories in 12 patches. We found evidence of a minimum patch size threshold (between 15.0 ha and 20.1 ha) of reproductive success for golden-cheeked warblers, but not for white-eyed vireos. We found no minimum patch size thresholds for presence, territory establishment by males, or pair formation for either species. Conservation practices based on thresholds of presence, territory establishment, or pair formation might not address issues of reproduction for golden-cheeked warblers. We failed to find evidence that cowbird parasitism or arthropod biomass were limiting factors. The ability to identify patch size thresholds of reproductive success for target species could be useful in conservation and management for setting goals for retention and restoration of target species’ habitat patch size.     

Factors affecting the density of the middle spotted woodpecker Dendrocopos medius: a macrohabitat approach

The main goals of this study were to investigate the relationships between densities of the breeding population of the middle spotted woodpecker Dendrocopos medius obtained on 19 large-scale plots, and the share of different habitats and age of forest stands. An average crude density based on an area defined arbitrarily by observers was 0.66±0.43 territories/10 ha, while the ecological density based on an area including habitats potentially used by organisms reached 1.09±0.64 territories/10 ha. Stepwise multiple regression indicated that the crude density was negatively affected by the share of coniferous forest thereby explaining 50% of variation. The ecological density was negatively correlated with the percentage of young deciduous stands (0–40 years old). This variable explained 36% of the total variation. These results suggest that on the level of macrohabitat analysis, the share of unsuitable forest stands seem to play a major role in determining the density of the middle spotted woodpecker.     

Habitat fragmentation influences nestling growth in Mediterranean blue and great tits

In patchy forest areas, the size of the forest patch where birds breed has a strong influence on their breeding success. However, the proximate effects contributing to lowering the breeding success in small forest patches remain unclear; and a shortage of crucial resources in those forest patches has been suggested to account in some degree for this failure. With the aim to further investigate this issue, we have monitored the breeding cycle of blue and great tits in three ‘large’ forest patches (ranging between 26.5 and 29.6 ha) and twelve ‘small’ forest patches (ranging between 1.1 and 2.1 ha) in a Mediterranean area in central Spain, during three years (2011–2013). We also recorded the nestling diet inside the nest-boxes with the aid of handy-cams. Only males significantly differed between forest patch size categories; being on average younger and with better body condition in small patches for great and blue tits respectively. Reproductive traits did not vary between forest patch size categories, but the body condition of blue tit nestlings and the size of great tit nestlings did, being significantly better and larger respectively in large forest patches. The recruitment rate of blue tit nestlings was also higher in large patches. Regarding nestling diet, blue tits did not differ but great tits did, delivering a larger amount of caterpillars in large forest patches. Most variation in the reproductive traits occurred between years, probably due to annual differences in environmental conditions. This study suggests that food supply could be limiting the breeding success of birds above all in small patches, but also in large patches under particular environmental conditions.     

The reproductive success of Blackbirds Turdus merula in relation to habitat structure and choice of nest site

The reproductive success of a population of Blackbirds Turdus merula occupying farmland and woodland was studied over 3 years to investigate the effects of habitat on breeding success. Territory distribution was patchy in both farmland and woodland; some areas were unoccupied, while other areas were occupied at variable densities. Habitat structure appeared to influence occupation: the index of habitat complexity ("cover score") was higher in occupied areas than in unoccupied areas and high-density territories had higher cover scores than low-density territories. However, habitat structure had no significant effect on reproductive success because the cover scores of territories where pairs were successful did not differ significantly from those of territories where there were no successful breeding attempts. There was no evidence of differential mortality rates in adults according to habitat. The height, bulk and exposure of c. 430 nests were measured to determine the effect of nest and nest-site characteristics on reproductive success. Nest exposure was the only feature that differed between successful and failed nests, successful nests being less exposed than failed nests. The major cause of breeding failure was nest predation, but the effect of nest exposure operated only during the laying and incubation period and not during the nestling period. The significance of habitat structure for variation in population densities between habitats is discussed.     

导致西滨鹬集群繁殖的直接因子和最终因子(英文)

该文报道了西滨鹬 (Calidris mauri) 在美国阿拉斯加州育空-卡斯科奎姆河三角洲相对较小的空间尺度上所表现的集群繁殖行为。在开始产卵之前, 西滨鹬雄鸟聚集在一处 36 hm2的区域鸣唱飞行, 进行求偶炫耀。随后在该区域营巢时, 西滨鹬也保持着集群状态。检验了三个常用于解释具领域行为的物种的集群繁殖的假说:社会成员选择假说、捕食假说和物质资源限制假说。结果表明, 与斑块可利用性相比, 分散的个体常栖息于地势平坦的苔原栖息地斑块, 而集群个体常筑巢于地势起伏的苔原栖息地斑块, 因此, 该研究支持物质资源限制假说。在地势起伏的苔原栖息地斑块集群营巢的西滨鹬繁殖成功率较低, 这表明西滨鹬的这种栖息地占有模式符合理想的等级分布。在研究样地中, 年长且更具攻击性的雄鸟向雌鸟求偶的次数更多, 并且保卫位于地势平坦的苔原栖息地斑块上更大的领域; 相比之下, 年幼且攻击性较弱的雄鸟则聚集在地势起伏的苔原栖息地斑块。通常情况下, 繁殖期的集群多集中于某种关键资源的分布区内或分布区附近, 但西滨鹬繁殖集群的出现是由于年长和处于优势地位的个体将年幼和处于从属地位的个体排斥出高质量的栖息地。虽然, 繁殖领域的集群现象在很多非群居的鸟类中都曾有报道, 但该文是首次对非群居的单配制鸻鹬类物种在孵卵之前的集群繁殖行为进行了定量研究。     

Density-dependent habitat selection: Testing the theory with fitness data

Summary According to density-dependent habitat selection theory, reproductive success should decline with increased density. Fitness should be similar between habitats if habitat selection follows an ideal free distribution; fitness should be dissimilar between habitats if habitat selection is modified by territorial behavior. I tested these assumptions by examining a variety of fitness estimates obtained from white-footed mice living in nest boxes in forest, forest edge and fencerow habitats in southwestern Ontario. As expected, mean litter size declined with increased population density. Litter sizes, adult longevity and the proportion of adult animals in breeding condition were not significantly different among the three habitats. The success at recruiting at least one offspring to the adult population and the number of recruits per litter were much greater in the forest than in either of the other two habitats. Fitness was thus unequal among habitats and the results confirm both assumptions of density-dependent habitat selection theory for territorial white-footed mice.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Determinants of male mating success in a natural population of a stream goby of the genus Rhinogobius

The mating success of individually identified males of a stream goby, Rhinogobius sp. CB (cross‐band type), was recorded over one breeding season. One to 3 year‐old males were active in mating, but 3 year‐old males, which accounted for 36% of the population, amounted to c . 70% of the total males guarding eggs in the nest. Three quarters of the breeding males had only one brooding cycle, but the others had two or three. All the latter males changed their nest sites between cycles within a riffle. In 1 and 2 year‐old males, the number of brooding cycles contributed more to the mating success than egg mass size in one brooding cycle. For mating success of 3 year‐old males, the egg mass size in one brooding cycle, which can be enhanced by spawning with a large female or multiple females, was as important as the number of brooding cycles. These male reproductive tactics could be attributed to the age‐related ability of nest construction and mate acquisition.     

An ‘ecological trap’ for yellow warbler nest microhabitat selection

Contrary to assumptions of habitat selection theory, field studies frequently detect ‘ecological traps’, where animals prefer habitats conferring lower fitness than available alternatives. Evidence for traps includes cases where birds prefer breeding habitats associated with relatively high nest predation rates despite the importance of nest survival to avian fitness. Because birds select breeding habitat at multiple spatial scales, the processes underlying traps for birds are likely scale‐dependent. We studied a potential ecological trap for a population of yellow warblers Dendroica petechia while paying specific attention to spatial scale. We quantified nest microhabitat preference by comparing nest‐ versus random‐site microhabitat structure and related preferred microhabitat features with nest survival. Over a nine‐year study period and three study sites, we found a consistently negative relationship between preferred microhabitat patches and nest survival rates. Data from experimental nests described a similar relationship, corroborating the apparent positive relationship between preferred microhabitat and nest predation. As do other songbirds, yellow warblers select breeding habitat in at least two steps at two spatial scales; (1) they select territories at a coarser spatial scale and (2) nest microhabitats at a finer scale from within individual territories. By comparing nest versus random sites within territories, we showed that maladaptive nest microhabitat preferences arose during within‐territory nest site selection (step 2). Furthermore, nest predation rates varied at a fine enough scale to provide individual yellow warblers with lower‐predation alternatives to preferred microhabitats. Given these results, tradeoffs between nest survival and other fitness components are unlikely since fitness components other than nest survival are probably more relevant to territory‐scale habitat selection. Instead, exchanges of individuals among populations facing different predation regimes, the recent proliferation of the parasitic brown‐headed cowbird Molothrus ater, and/or anthropogenic changes to riparian vegetation structure are more likely explanations.     

Distribution of birds in natural landscape mosaics of old-growth forests in northern Sweden: relations to habitat area and landscape context

We censused breeding birds for three years in natural landscape mosaics of virgin old-growth spruce forest and mire in a large protected forest area in northern Sweden Twenty forest patches, ranging from 0 2 to 17 8 ha in size, were selected in two matrix types, dominated by forest and mire, respectively Patches were very similar with regards to habitat features There was a strong effect of patch area on species richness, but no effect of matrix type Standardization of species richness by rarefaction revealed that small patches (<5 ha) had fewer and large patches (>10 ha) more species than expected Overall distribution of species across patches showed a highly significant nested pattern, indicating that a few habitat generalists occupy all size classes, whereas more demanding species avoid small patches regardless of landscape composition Individual species tended to be distributed evenly across patch classes and no significant edge effect in terms of density of birds was found Our results have bearings on actions to preserve avian diversity in northern boreal forests small patches (<5 ha) provide habitat only for habitat generalists, and therefore larger (>10 ha) patches should be preserved     

Determining habitat quality for the cooperatively breeding Rufous Treecreeper,Climacteris rufa

Abstract The theory of habitat selection predicts that organisms should use habitat that maximizes their fitness. The cooperatively breeding Rufous Treecreeper, Climacteris rufa, exhibits non‐random habitat use at a number of spatial scales. By assessing correlative relationships between nest‐site use and nest success, and territory use and reproductive success and survival, it was determined whether non‐random use of habitat yields fitness benefits. It was also determined whether breeding group size contributed significantly to fitness once differences in territory quality had been considered. Structural characteristics of nest sites that were positively correlated with the probability of a site being used had no relationship with nest success. This result probably reflects the relatively unrestricted access to an abundance of suitable nest sites in the study area. Habitat traits that predicted territory use by treecreepers were positively correlated with a number of fitness measures. They were also positively correlated with breeding group size and provisioning rate to nestlings, which in turn were correlated with fitness. However, group size was not significantly related to any measure of fitness, except primary male survival, once territory quality had been considered. The quality of territories occupied by Rufous Treecreepers appeared to be a significant factor for breeding group fitness.     

Habitat selection by breeding Curlews Nwnenius arquata on mosaic farmland

Territory establishment and habitat use by breeding Curlews Numenius arquata were studied during 1987 and 1988 on mosaic farmland (dominated by dry tillage) at two sites in central Sweden. Curlews preferred to breed in areas with a high proportion of grassland, close to rivers, while dry tillage was avoided. Territories at my study site were larger (mean = 4 5.2 ha) than in areas consisting entirely of grassland. Territory size seemed to depend on the spatial distribution of grasslands, which suggests that habitat fragmentation forces Curlews to establish larger territories in modern farmland than in areas of grassland. The number of territories in patches of grassland was correlated with patch area, and unoccupied patches were more isolated than occupied patches. However, patch area was a more important factor than isolation, since large patches (> 3 5 ha) were always occupied. Sown grassland was used significantly more than expected for foraging early in the season, possibly indicating the strong influence of the nutritional requirements in the pre-breeding period on territory establishment. Habitat selection when foraging seemed to be less important late in the season, since there was no significant habitat preference then. During this period distance to the nest site seemed to be more important than habitat, sinced the preferred foraging fields (including fields of all habitats used more than expected by area) were situated closer to nests than the less preferred fields, probably an adaptation to the high nest predation risk. The same fields were mostly preferred in the pre-breeding period also, suggesting that nests were built close to good foraging areas.
My results indicate that the decline of the Swedish Curlew population since 1950 is caused by changes in land use, resulting in decreased grassland area and increased habitat fragmentation, which probably have affected both breeding and foraging possibilities negatively.     

Complex effects of habitat loss on Golden Eagles Aquila chrysaetos

The development of commercial forests presents potential threats to large raptors that rely on prey caught in open country. We examined the effect of afforestation of breeding habitat used by a population of Golden Eagles Aquila chrysaetos in Scotland where, over the last 50 years, extensive stands of exotic conifers have been planted. Using data for 31 years on territory occupancy and breeding success, together with spatiotemporally dynamic mapping of forest cover and predicted areas of territory‐use in a Geographical Information System, we examined relationships between forest cover and Eagle ecology at landscape and individual territory scales. Several territories were abandoned during the earliest phases of forest planting, but relatively few were apparently lost to later plantings. Territories with poorer breeding productivity appeared to be more vulnerable to abandonment than territories with better breeding productivity. At the landscape scale, temporal differences in breeding productivity were negatively related to the extent of forest cover, although productivity of individual territories showed no clear relationship with forest cover. Several territories with less than a 5% increase in forest cover experienced reduced productivity; however, territories least constrained by neighbouring pairs of Eagles showed an increase in productivity. Territories experiencing the greatest increases in forest cover showed a greater use of spatially separated nest‐sites by occupying pairs. Hence, pairs that were less constrained by neighbours appeared to compensate for loss of open habitat by shifting their territory‐use, whereas pairs that were more constrained could not compensate for open habitat loss and suffered reduced productivity (and, probably in some cases, abandoned the territory). We suggest that simple guidelines based on the extent and locations of habitat loss are inadequate when predicting effects on large territorial raptors such as Golden Eagles. Consideration should also be given to the ‘quality’ of a territory or occupying pair, as well as the extent to which territory‐use is constrained by neighbouring pairs or other ‘unsuitable habitat’ which may have been affected by previous episodes of open habitat loss.     

Spawning substrata are important for breeding habitat selection but do not determine premating reproductive isolation in three sympatric Hexagrammos species

Habitat use and spawning substrata were surveyed to characterize the contribution of habitat divergence to reproductive isolation in greenling Hexagrammos species. The spawning substrata and microhabitat in breeding territories differed amongst the three Hexagrammos species studied: H. octogrammus, H. agrammus and H. otakii used small red algae, surfgrass and bryozoans, respectively, as spawning substrata, and breeding territories were established in areas where those substrata were abundant. In contrast, non-territorial individuals were observed in a comparatively wider range of habitats than conspecific territories. Consequently, the distributions of non-territorial individuals of the three species partially overlapped. Since hybrids have been frequently collected, the difference in spawning substrata and the subsequent microhabitat divergence in breeding territories do not prevent females from encountering males of other species. Thus, in addition to habitat divergence, other factors such as behavioural differentiation may be needed to complete premating reproductive isolation amongst these three Hexagrammos species.     

Edge effects reduce the nesting success of Acadian Flycatchers in a moderately fragmented forest

ABSTRACT.   Forest fragmentation can create negative edge effects that reduce the reproductive success of birds nesting near the forest/nonforest interface, and threaten bird populations deeper in remnant forest habitats. Negative edge effects may be more pronounced in landscapes that are moderately fragmented, particularly where agriculture is the primary land-use fragmenting forests. Information about the extent and strength of edge effects at a site can help guide conservation actions, and determine their effectiveness. We examined edge effects for birds breeding in a nearly contiguous forest fragmented by relatively narrow agricultural corridors in Illinois (USA). We measured rates of nest predation and brood parasitism for Acadian Flycatchers ( Empidonax virescens ) over a continuum of distances from the edge of an agricultural inholding. Nest predation and brood parasitism were highest near the edge and decreased with increasing distance from the edge. Given the cumulative effects of nest predation and brood parasitism on reproductive success, we determined that forest within 600 m of the inholding was sink habitat. We found, however, that deeper forest interior areas currently serve as source habitat, and that conversion of the entire 205 ha agricultural corridor to forest would add 1350 ha of source habitat for Acadian Flycatchers. Such results provide support for a local conservation strategy of forest consolidation and establish baseline measures necessary to determine the relative effectiveness of any subsequent reforestation efforts.