Ecological biogeography of southern ocean islands: species-area relationships, human impacts, and conservation

ABSTRACT Previous studies have concluded that southern ocean islands are anomalous because past glacial extent and current temperature apparently explain most variance in their species richness. Here, the relationships between physical variables and species richness of vascular plants, insects, land and seabirds, and mammals were reexamined for these islands. Indigenous and introduced species were distinguished, and relationships between the latter and human occupancy variables were investigated. Most variance in indigenous species richness was explained by combinations of area and temperature (56%)-vascular plants; distance (nearest continent) and vascular plant species richness (75%)-insects; area and chlorophyll concentration (65%)-seabirds; and indigenous insect species richness and age (73%)-land birds. Indigenous insects and plants, along with distance (closest continent), explained most variance (70%) in introduced land bird species richness. A combination of area and temperature explained most variance in species richness of introduced vascular plants (73%), insects (69%), and mammals (69%). However, there was a strong relationship between area and number of human occupants. This suggested that larger islands attract more human occupants, increasing the risk of propagule transfer, while temperature increases the chance of propagule establishment. Consequently, human activities on these islands should be regulated more tightly.     

Biogeography of the marine birds of a confined sea, the Mediterranean

Aim The Mediterranean sea is a winter productive oligotrophic basin where Atlantic water replaces water lost through evaporation, this influx being a major source of productivity and fertility. The long coastlines and the large number of islands cause high oceanographic heterogeneity. Moreover, during its geological history, it has dried out several times. So we describe the consequences of these particular features on species richness, distribution, and breeding ecology of marine birds. Location The Mediterranean sea (including the Black Sea and the Sea of Azov) communicates with the Atlantic Ocean only through a 14 km wide channel (Straits of Gibraltar), and since 1869, with the Red Sea through the Suez Canal. Methods The Mediterranean was subdivided into different areas, according to physical oceanographic entities and productivity, linked to numbers and distribution of both breeding and wintering marine birds (defined as species strongly dependent on marine resources, breeding only on islands and/or the coastline). Results The total marine bird biomass, and species diversity, are lower in the Mediterranean than in the near Atlantic. The eastern Mediterranean, with lowest primary productivity, contains fewer marine bird taxa than the more productive western part. Taxa which mainly occur in the western and southern parts of the Mediterranean migrate through the Straits of Gibraltar to winter in the southern Atlantic, while those inhabiting the northern and eastern parts are sedentary, as a result of differences in species composition. Northern coastal basin communities (i.e. the Tyrrhenian and the Balearic Seas), are composed of less pelagic, and earlier breeding species, that rear chicks during the productive season. These latter taxa are actually the most typical Mediterranean taxa, in terms of endemism. Main conclusions The Mediterranean marine bird community is not tropical, but rather, shows the highest affinity with the Atlantic temperate community. Its level of endemism is however high and comparable to other confined basins such as the Red Sea.     

Repopulation and colonization by birds in the Agmon Wetland,Israel

Lake Hula and its surrounding wetland in northern Israel were drained in the late 1950s and the dried wetlands were transformed into a diverse agricultural region with a 0.3 km2 nature reserve. A portion of the extinct Hula wetland was re-flooded in April 1994 by constructing a small lake, Agmon, and 90 km of canals. The purposes of this study were (a) to document the re-population and colonization of the new Agmon wetland by birds after its flooding, (b) to evaluate bird species richness in this new wetland in comparison to that in a nearby mature Hula Nature Reserve, and, (c) to investigate if the species originally present in the Hula Valley before the drainage had been re-established. The new lake has succeeded in attracting a large variety of water birds to the newly formed habitats, especially ducks, egrets and herons, plovers, waders and snipes. The distribution of bird species among the different habitats was not random. The northern section of the lake, which is shallow and has an open shore, had the largest number of species during 1995 and 1996 and was preferred by plovers, waders and snipes. A large roosting and breeding colony of thousands of egrets and herons was formed in the dense cattails ( Typha domingensis) in the southern section of the lake. Unlike the nearby Hula Nature Reserve, the new wetland lacks a large seasonally flooded area and is less attractive for feeding and breeding plovers, waders and stilts. The new wetland also has very little muddy area without vegetation and attracts few of the species that regularly feed in this habitat in the reserve. The new wetland enlarges the area of only a few of the habitats found in the nearby reserve and therefore attracts fewer species than it might otherwise. Recommendations for management of the new wetland include increased diversity of habitats, restriction of human activities and optimization of conflicts that have arisen between the needs of the wetland and those of nearby agriculture.     

Plants and birds on the shore of the Sea of Okhotsk: Balance, crisis, adaptation

Mutual relationships between sea colonial birds and plants on the northern shore of the Sea of Okhotsk are considered. The birds form huge flocks on small islands thus transforming the island landscape and providing specific vegetation at nesting sites. The intensity of ornithogenic load determines different relations between the productivity of plant communities and their species richness. As a rule, flora is depleted. The plants thriving at rookeries react to the intensive action of birds by forming specific ecobiomorphs. On the islands of the northern part of the Sea of Okhotsk tussocks of pine purple grass existing on the majority of nesting colonies are of the highest significance.     

Comparing alpha-diversity between plants and birds in a remnant wetland: evidence for a threshold and implication for management

In a Mediterranean patchy wetland of central Italy, we analyzed the relationship between the number of bird species, expressed in terms of bird alpha diversity, and plant alpha diversity, expressed as Hill number. This number (the exponential of the Shannon entropy) is considered one of the most strong and reliable indexes of alpha-diversity, synthesizing the information on evenness, richness and diversity in one single metric. We observed a progressive increase of the mean values of bird alpha diversity when plant alpha diversity increases along Hill number. Bird alpha diversity shows an abrupt increase between the first and the second of four categories of plant alpha diversity (0–1, >1–2, >2–3, >3), indicating a threshold response in all the groups considered (breeding, wintering and total bird assemblages). This marked decline of bird species richness at around 1 in the Hill index should represent an alarm for managers: wetland sites at or below this level of plant alpha diversity are likely to be experiencing a drastic decrease in bird species richness, both in spring (breeding birds) and in winter (wintering birds).     

Numbers of plant, insect and land bird species on nineteen remote islands in the Southern Hemisphere

The familiar species-area equation S = cA ^z may not account for as much of the variation in the numbers of species on islands as it has been held to. The undue emphasis on area as a factor probably results from early workers not considering as many factors as possible in their regression analyses.
In the case of 19 remote islands in the Southern Hemisphere, island area and elevation, latitude, mean temperature of coldest month and distance of island from mainland, nearest land and nearest land to west were used to consider variation in number of plant species with the technique of multiple regression. A nonlinear model explained 83% of the variation, with temperature and area accounting for 71% of the variation in number of plant species.
The above factors with number of plant species were then used to consider variation in number of insect species on the islands. The linear model was superior, accounting for 91% of the variation. Number of plant species accounted for 73% of the variation in number of insect species.
Variation in numbers of passerine bird species and total land bird avifauna was considered using all of the above nine factors. The linear model was superior in both cases. Number of plant species accounts for 60% and 71% of the variation respectively.
Various suggestions are made to account for the unexpected result of number of insect species making little contribution to the last two regressions.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Little evidence for sympatric speciation in island birds

It has been suggested that the presence of sister species in small circumscribed areas, such as isolated lakes or islands, might imply that these species originated sympatrically. To investigate this possibility in birds, we searched for endemic, congeneric species on isolated islands in the ocean. Among 46 islands and small archipelagos chosen because they contain at least one species of endemic land bird, we identified seven pairs of endemic congeners (excluding flightless rails). Of these seven, only four pairs are potentially sister species and thus possible candidates for sympatric speciation. However, three of these four pairs have always been considered the results of double invasion from a mainland source (in two of these cases, molecular-phylogenetic work has either confirmed a double invasion or is ambiguous). The one remaining pair may have speciated allopatrically on a small archipelago. Additional phylogenetic studies are required to understand these cases, and our results should also be considered in light of the large number of island-bird extinctions in historic time. We conclude that, at present, there is little evidence for sympatric speciation in island birds.     

Winter bird communities in the Åland archipelago: an island biogeographic point of view

The distribution of wintering land birds was examined in the archipelago of Åland (60°N, 20°E) along two ecological dimensions: (1) the island continuum from small skerries ((0.5 ha) to the mainland of Åland (97000 ha), and (2) the habitat range of the mainland of Åland. In the species-area relationship, both the exponential and power function models fitted well. The increase of the number of species with island size was interpreted as a consequence of increasing habitat diversity on larger islands. The smallest islands supported only few seed-eating species utilizing the islands in a fine-grained fashion; none of these birds belonged to the breeding fauna of the islands. With increasing island size and habitat diversity, insectivores and species tied to the human culture were added. On the mainland of Åland, more than half of the birds were observed near human habitations: bird communities wintering in pine forests were richer than in deciduous forests. Proportion of wintering species was greatest in the breeding communities of pine forests. The proportion of wintering species was presumably regulated by the degree of structural change in the habitat between summer and winter, this change being most drastic in deciduous habitats The species observed could be broadly classified into three feeding categories: opportunists ., omnivores and scavengers, dependent on the diverse food items provided by man; food specialists , species éating seeds and berries, ranging widely in the islands and habitats; and insectivorous habitat specialists , inhabiting coniferous habitats, and islands with coniferous (pine) forests. Opportunists, in particular, gel increasing wintering opportunities by the impact of man.     

A global review of island endemic birds

Although fewer than one-fifth of the world's bird species are restricted to islands, over 90% of bird extinctions during historic times have occurred on islands. The major identified cause has been the effects of exotic animal species introduced by man; the largest number of documented extinctions has occurred on islands of the Pacific Ocean.
Some 39% (402) of threatened bird species are restricted to islands and more than 907; of these are endemic to a single geopolitical unit. The largest numbers occurring in such units are in Indonesia (91) and the Philippines (34). As a region, the Pacific holds more threatened species (110) than any other, including almost half of those considered Endangered and over 40% of the Vulnerable species.
Most threatened island species are forest-dwelling. A high proportion of the Endangered species use seasonal/temperate forest. While habitat destruction now poses the greatest overall threat to island birds (affecting over half the species restricted to islands), the presence of introduced species threatens 30 of the 66 Endangered species.
Although immediate extinctions of island species can best be averted by mitigating the effects of introductions, the removal of native forests will be a more severe problem in the longer term. There is an urgent need for ecologists to provide detailed information on the habitats of both threatened and endemic species so that more appropriate and effective conservation programmes can be developed.     

THE BIRDS OF THE COCOS-KEELING ISLANDS (INDIAN OCEAN)

The birds of the Cocos-Keeling Islands, in the Indian Ocean, are discussed in the light of specimens and data collected during a stay of eleven months in 1941. Seventeen species were found to be breeding. One of these was an indigenous land bird, four were land birds known to have been introduced, and the remainder were sea birds mostly confined to North Keeling. During this period examples of seventeen species of strays or visitors were taken, of which ten probably occur regularly though in small numbers, while the other seven are true vagrants. Eight of these birds have not so far been recorded from Christmas Island.
Eleven birds listed from the islands by previous visitors were not present in 1941. Two of these were introduced but have failed to maintain themselves, and one, Egrctta garzetta nigripes , seems to be an occasional vagrant. The remaining eight records are all unsatisfactory.     

Does size really matter? Species–area relationships in human settlements

Aim To describe species–area relationships in human settlements and compare them with those from a non‐urban habitat. Location West‐central Mexico. Methods We surveyed breeding birds in 13 human settlements and five shrubland patches. We estimated bird species richness using an abundance‐based coverage estimator with equal sample sizes to eliminate biases related to sampling effort differences. To assess species–area relationships, we performed log–log linear regressions between the size of the studied patches and their estimated bird richness. We also used a logarithmic approach to determine how the species–area relationship asymptoted and made use of the Michaelis–Menten model to identify the size at which the studied patches reached their maximum species richness. We also investigated (1) possible relationships among the estimated bird richness and other variables known to affect urban‐dwelling birds (built cover, plant species richness, tree cover or human population density) and (2) changes in bird community composition related to the size of the studied human settlements. Results Species–area relationships exhibited different patterns among the studied habitats. The log–log regression slope was steeper in human settlements, while the intercept was higher in shrublands. The maximum number of species was more than twofold higher in shrublands. Human settlement patch size was the only variable significantly related to bird richness. Our community composition results show that two main bird groups are related to human settlement size, and that as the size of human settlements increases, bird community similarity in relation to the largest city increases. Main conclusions Human settlements act as ecological islands, with pronounced species–area relationships. Our results suggest that an important threshold for bird species richness and community composition is reached in human settlements > 10.2 km². This threshold is unlikely to be generalizable among bio‐regions, and thus should be quantified and considered when studying, managing and/or planning urban systems.     

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.     

Bird pollination syndrome is the plant's adaptation to ornithophily,but nectarivorous birds are not so selective

Many tropical plants are pollinated by birds and several bird phylogenetical lineages have specialised to a nectar diet. The long-assumed, intimate ecological and evolutionary relationship between ornithophilous plants and phenotypically specialised nectarivorous birds has nevertheless been questioned in recent decades, where such plant–pollinator interactions have been shown to be highly generalised. In our study, we analysed two extensive interaction datasets: bird–flower and insect–flower interactions, both collected on Mt Cameroon, west-central Africa. We tested if: 1) insects and birds interact with distinct groups of plants; 2) plants with a typical set of ornithophilous floral traits (i.e. bird pollination syndrome) interact mainly with birds; 3) birds favour plants with bird pollination syndrome and; 4) if and how the individual floral traits and plant level nectar production predict bird visitation. Bird-visited plants were typically also visited by insects, while approximately half of the plants were visited by insects only. We confirmed the validity of the bird pollination syndrome hypothesis, as plants with bird-pollination syndrome traits were visited by birds at a higher rate and mostly hosted a lower frequency of visiting insects. However, these ornithophilous plants were not more attractive than the other plants for nectar-feeding birds. Nectar production per plant individual was a better predictor of bird visitation than any other floral trait traditionally related to the bird pollination syndrome. Our study thus demonstrated the highly asymmetrical relationship between ornithophilous plants and nectarivorous birds.     

Parallel trends in the species diversity of west indian birds and bats

Summary In this paper I compare several biogeographic patterns of West Indian resident land birds and bats, including species-area and trophic diversity-area relationships, the number of islands inhabited per species and levels of endemism, trophic structure as compared with tropical mainland areas, and the degree of faunal simlarity between islands of similar sizes but different locations. In most respects, the bat and bird patterns are strikingly similar. Groups of birds that are conspicuously missing from the Antilles because of the absence of appropriate resources also have missing chiropteran counterparts. Plant-visiting bats and birds are better-represented in terms of relative number of species and, in birds, in biomass, on the Lesser Antilles than on the mainland (e.g. Panama). Small Antillean islands tend to share more species of birds and bats than do larger islands. Stochastic (sensu Simberloff 1978), deterministic, and interactive (e.g. competitive and trophic interactions) factors appear to underly these biogeographic trends. No evidence exists to suggest that Caribbean bats and birds have negatively affected each other's diversity.     

千岛湖陆桥岛屿繁殖鸟类的扩散能力差异 对群落动态的影响

岛屿生物地理学理论的核心过程是岛屿物种的周转, 包括迁入与灭绝。本研究旨在探讨扩散能力差异对岛屿繁殖鸟类群落动态的影响。2007年4月至2013年6月, 采用样线法调查了千岛湖36个陆桥岛屿的繁殖鸟类, 依据扩散能力强弱将其划分为两类, 结合陆桥岛屿参数, 并运用逻辑斯蒂回归模型和最大似然法, 来研究鸟类扩散能力的不同对其周转率的影响。结果表明, 千岛湖繁殖鸟类扩散能力强的物种具有较高周转率且受岛屿参数约束较小, 而扩散能力弱的物种周转率较低且对岛屿参数变化更敏感。因此, 千岛湖陆桥岛屿繁殖鸟类的扩散能力显著影响其群落动态。     

Flower visitation by birds in Europe

Most flowering plants depend on animal pollination. Several animal groups, including many birds, have specialized in exploiting floral nectar, while simultaneously pollinating the flowers they visit. These specialized pollinators are present in all continents except Europe and Antarctica, and thus, insects are often considered the only ecologically relevant pollinators in Europe. Nevertheless, generalist birds are also known to visit flowers, and several reports of flower visitation by birds in this continent prompted us to review available information in order to estimate its prevalence. We retrieved reports of flower–bird interactions from 62 publications. Forty‐six bird species visited the flowers of 95 plant species, 26 of these being exotic to Europe, yielding a total of 243 specific interactions. The ecological importance of bird–flower visitation in Europe is still unknown, particularly in terms of plant reproductive output, but effective pollination has been confirmed for several native and exotic plant species. We suggest nectar and pollen to be important food resources for several bird species, especially tits Cyanistes and Sylvia and Phylloscopus warblers during winter and spring. The prevalence of bird flower‐visitation, and thus potential bird pollination, is slightly more common in the Mediterranean basin, which is a stopover to many migrant bird species, which might actually increase their effectiveness as pollinators by promoting long‐distance pollen flow. We argue that research on bird pollination in Europe deserves further attention to explore its ecological and evolutionary relevance.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Effects of perennial wildflower strips and landscape structure on birds in intensively farmed agricultural landscapes

Farmland bird populations are in a deep crisis across Europe. Agri-environment schemes (AES) were implemented by the European Union to stop and reverse the general decline of biodiversity in agricultural landscapes. In Germany, flower strips are one of the most common AES. Establishing high-quality perennial wildflower strips (WFS) with species-rich native forb mixtures from regional seed propagation is a recent approach, for which the effectiveness for birds has not yet been sufficiently studied. We surveyed breeding birds and vegetation on 40 arable fields with WFS (20 with single and 20 with aggregated WFS) and 20 arable fields lacking WFS as controls across Saxony-Anhalt (Germany). Additionally, vegetation composition, WFS quantity and landscape structure (e.g. distance to nearest woody element) were considered in our analyses. All WFS were established with species-rich native seed mixtures (30 forbs) in agricultural practice as AES. Arable fields with WFS had a higher species richness and territory density of birds than controls, confirming the effectiveness of this AES. A forb-rich vegetation was the main driver promoting birds. Flower strip quantity at the landscape level had positive effects only on bird densities, but also single WFS achieved benefits. A short distance from WFS to woody elements increased total bird species richness. However, the density of farmland birds, which are target species of these AES, were negatively affected by the proximity and proportion of woody elements in the vicinity. The effect of the proportion of non-intensively used open habitats and overall habitat richness was unexpectedly low in the otherwise intensively farmed landscape. Species-rich perennial WFS significantly promoted breeding birds. Successful establishment of WFS, resulting in high-quality habitats, a high flower strip quantity as well as implementation in open landscapes were shown to maximise the effectiveness for restoring declining and AES target farmland birds.     

The effect of land type diversity and spatial heterogeneity on farmland birds in Norway

The decline in farmland birds observed throughout Europe during recent decades has attracted much attention. Agricultural intensification or land abandonment are commonly forwarded as key drivers. Several countries have established agri-environmental schemes (AES) to counter these negative trends among farmland birds. This paper reports a study of the relationship between land use and bird species in the agricultural landscape of Norway. The main objective was to investigate the effect of spatial heterogeneity and diversity of land use on total richness and abundance of farmland birds at a national level.Monitoring the distribution and abundance of birds is part of the Norwegian monitoring programme for agricultural landscapes. The monitoring programme is based on mapping of 1 × 1 km squares distributed across the entire agricultural landscape. Within these squares permanent observation points are established for bird monitoring. Detailed interpretation of aerial photographs provides the land classification. We tested the relationship between landscape metrics at different levels of land type detail and species richness and abundance of farmland and non-farmland birds.There was a positive relationship between species richness and abundance of farmland birds and agricultural area. For non-farmland birds the relationship was negative. Spatial heterogeneity of land use was a significant positive factor for both farmland and non-farmland species. High land type diversity was positive for farmland bird richness, but negative for abundance. Non-farmland bird richness was not affected by land type diversity, but abundance had a negative response.The results presented in this paper highlight the importance of a spatial heterogeneous landscape. However, we also found that land type diversity could negatively affect the abundance of both farmland and non-farmland birds. Our findings suggest a need for different management approaches depending on whether the aim is increased species richness or abundance. Achieving both aims with the same means might be difficult. We thus suggest a need for land use analyses before proper management strategies can be implemented.