Patterns of species richness on very small islands: the plants of the Aegean archipelago

Aim To investigate the species–area relationship (SAR) of plants on very small islands, to examine the effect of other factors on species richness, and to check for a possible Small Island Effect (SIE). Location The study used data on the floral composition of 86 very small islands (all < 0.050 km²) of the Aegean archipelago (Greece). Methods We used standard techniques for linear and nonlinear regression in order to check several models of the SAR, and stepwise multiple regression to check for the effects of factors other than area on species richness (‘habitat diversity’, elevation, and distance from nearest large island), as well as the performance of the Choros model. We also checked for the SAR of certain taxonomic and ecological plant groups that are of special importance in eastern Mediterranean islands, such as halophytes, therophytes, Leguminosae and Gramineae. We used one‐way anova to check for differences in richness between grazed and non‐grazed islands, and we explored possible effects of nesting seabirds on the islands’ flora. Results Area explained a small percentage of total species richness variance in all cases. The linearized power model of the SAR provided the best fit for the total species list and several subgroups of species, while the semi‐log model provided better fits for grazed islands, grasses and therophytes. None of the nonlinear models explained more variance. The slope of the SAR was very high, mainly due to the contribution of non‐grazed islands. No significant SIE could be detected. The Choros model explained more variance than all SARs, although a large amount of variance of species richness still remained unexplained. Elevation was found to be the only important factor, other than area, to influence species richness. Habitat diversity did not seem important, although there were serious methodological problems in properly defining it, especially for plants. Grazing was an important factor influencing the flora of small islands. Grazed islands were richer than non‐grazed, but the response of their species richness to area was particularly low, indicating decreased floral heterogeneity among islands. We did not detect any important effects of the presence of nesting seabird colonies. Main conclusions Species richness on small islands may behave idiosyncratically, but this does not always lead to a typical SIE. Plants of Aegean islets conform to the classical Arrhenius model of the SAR, a result mainly due to the contribution of non‐grazed islands. At the same time, the factors examined explain a small portion of total variance in species richness, indicating the possible contribution of other, non‐standard factors, or even of stochastic effects. The proper definition of habitat diversity as pertaining to the taxon examined in each case is a recurrent problem in such studies. Nevertheless, the combined effect of area and a proxy for environmental heterogeneity is once again superior to area alone in explaining species richness.     

Production of planktonic crustaceans in the White Sea

This paper analyzes different methods for assessment of the production of marine organisms. The estimated values of annual production and ration of planktonic copepods of the White Sea are 5.0 × 10¹² and 2.2 × 10¹³ kcal, respectively. Comparison of the planktonic copepod ration with phytoplankton production shows that the energy demands of the copepod community are met by consumption of phytoplankton and, probably, also bacterioplankton and detritus.     

Vascular plant colonization and vegetation development on sea-born volcanic islands in the Aegean (Greece)

Inside the seawater-filled Santorini caldera (S Cyclades) the uninhabited islands Palea Kaimeni (PK) and Nea Kaimeni (NK) have been formed by submarine/subaerial activity. About 197 B.C. PK (0.54 km²) emerged, harbouring (in 1987) 178 vascular plant species, and covered by a mosaic of evergreen sclerophyllous and halophytic shrub communities and short-lived therophytic plant communities. NK, actually colonized by 156 vascular taxa, emerged ca 1570 A.D., its surface having enlarged to 3.44 km² owing to four subsequent eruptive phases of the island's volcano until 1950. At present, the most evolved vegetation type on NK is a steppe-like pioneer community rich in grasses and legumes apart from some Ficus carica trees which survived this century's volcanic eruptions. The beginnings of shrub vegetation on NK, originating from PK's consolidated woody canopy, are documented and evidence is shown of several ways of zoogenic and anthropogenic import of new vascular plants to the island. Comparative plant censuses demonstrate that permanent immigration of plants into NK overbalances by far the loss of species caused by the repeated destructions of the island's plant cover. Cases of species turnover (verified extinctions and invasions), of pseudoturnover (due to sampling errors), of long distance dispersal and of successful stabilization of founder-populations are exemplified.     

Factors affecting the distribution of vascular plants, springtails, butterflies and birds on small tropical islands

Aim

The primary objective of our study was to examine the factors affecting the distribution of vascular plants, springtails, butterflies and birds on small tropical islands to understand how different groups of organisms with distinct biological traits respond to biogeographical variables, such as island area.

Location

The Republic of Singapore (103°50′E, 1°20′N) located at the southern tip of Peninsular Malaysia.

Methods

Seventeen islands were surveyed for vascular plants, springtails, butterflies and birds. Correlation analysis, simple linear and multiple regression analyses and the nestedness index were used to test the hypotheses that (1) area is the best predictor of species/genus richness at both the community and specific/generic levels; (2) there is no correlation between population density and island area; and (3) species/genera are distributed as nested subsets.

Results

Area was the most significant factor in determining the island distribution of springtails, butterflies and birds at both the community and specific/generic levels, although there were disparate responses to the biogeographical variables between the three taxonomic groups, as well as between common species within each group. Individual species displayed disparate responses to biogeographical variables, suggesting that patterns of distribution at the community level may not be a good indicator of the population dynamics of individual species/genera. Plant species richness did not show any correlation with any of the tested variables. Population densities of springtails, butterflies and birds were positively correlated with area, contradicting the assumption of the equilibrium theory of island biogeography that population density of island species is independent of area. Population densities of plants showed no correlation with any of the tested biogeographical variables. Vascular plant, springtail, butterfly and bird communities on the islands showed significant patterns of nestedness, indicating there may be species/genus‐specific responses to biogeographical variables.

Main conclusions

We conclude that although area was the most important factor affecting the island distribution of springtails, butterflies and birds, conservation planning must take into consideration how target taxonomic groups respond to biogeographical variables, instead of relying on general principles (e.g. those derived from the equilibrium theory). On a local scale, in order to preserve the island biodiversity of Singapore, the highest priority should be given to preserving the larger islands (e.g. Pulau Ubin) which not only have higher numbers of species, but also species that are absent on smaller islands.

     

Assessment of primary production in the White Sea

The literature and original data on the primary production of phytoplankton in the White Sea are analyzed. By this parameter, the White Sea is significantly inferior only to the Chukchi Sea; it is similar to the Barents Sea, and exceeds other Russian Arctic seas by two to three times (the Kara Sea, Laptev Sea, and East Siberian Sea).     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

中国台湾种子植物区系的性质、特点及其与大陆植物区系的关系

台湾位于欧亚大陆东南缘的海洋中,地处热带的北部和亚热带的南部,约为21°45′~15°56′N,119°18′~124°34′E,是中国最大的岛屿。它是受季风气候强烈影响的地区之一,热量丰富,雨量充沛,干湿季明显。具有一个非常丰富的岛屿和山区植物区系。就其种子植物而言,约有186科,1201属,3656 种,包括热带属742属,温带属346属。根据台湾植物区系中各大科、主要植物群落优势种和中国特有种的地理分布以及热带属在整个植物区系中的主导地位,台湾地区的植物区系主体具有明显的亚热带性质。中国台湾本地特有种十分丰富,其比例远高于中国特有种的比例。这似乎表明台湾植物区系是一个古老区系在多次地质事件侵袭后又起活化的历史演变的结果。新老成分并存、共同发展是台湾植物区系的重要特点。通过台湾全部属和非特有种在周边地区地理分布的分析,中国台湾植物区系与中国大陆的关系最为密切,是东亚植物区系的重要组成部分,因此在植物分区上应属于泛北极植物区的东亚植物区系。     

Habitat islands and the equilibrium theory of island biogeography: testing some predictions

Summary Species-area data from a study of marsh birds are used to test five predictions generated by the equilibrium theory of island biogeography. Three predictions are supported: we found a significant species-area relationship, a non-zero level of turnover, and a variance-mean ratio of 0.5. One prediction is rejected: the extinction rates were not greater on small islands. The results of one test are equivocal: the number of species on each island was not always the same. As Gilbert (1980) suggests, a strong species-area relationship alone does not validate the theory. The avian communities we studied were on habitat islands, not true islands, and underwent complete extinction annually. Thus caution must be used before applying the theory to these and other habitat islands.     

渤海区无居民海岛植物物种丰富度分布格局

为了解渤海区小型无居民海岛植物物种丰富度的分布格局, 本研究以渤海区9个无居民海岛为研究对象, 基于种-面积关系和种-多度对数正态分布模型探究了无居民海岛植物的物种丰富度分布格局。结果表明: (1)位于温带地区的无居民海岛木本植物相对多度符合对数正态分布特征, 表明海岛木本植物群落中多度较大的物种数较少, 而多度较小的物种甚至是稀有种或罕见种较多; (2)渤海区多数无居民海岛物种数介于60-130之间, 表明总体丰富度较高; (3)通过分析海岛与周边保护区的空间关系发现, 物种丰富度高的海岛基本位于保护区内, 总体上渤海区海岛植物热点区得到了有效保护。综上所述, 种-多度分布模型和种-面积关系均是揭示海岛植物物种丰富度分布的重要手段, 同时可初步推断, 生态位理论对海岛植物物种多样性的维持显得更为重要。     

Genetic comparison of salmon from the White Sea and north-western Atlantic Ocean

Samples of salmon Salmo salar from the River Kachkovka and the River Nilma in northern Russia were analysed by starch gel electrophoresis and compared to three Norwegian stocks, the Neiden river in northern Norway and Øyreselv and Hopselv rivers on the west coast. The comparison included the following polymorphic loci: AAT-4 *, IDDH-2 *, IDHP-3 *, MDH- 3,4 *, MEP-2 *, ESTD * as well as the newly discovered polymorphic loci FBALD-3 * and TPI-3 *. Samples were run side by side on gels, and the alleles found in the Russian stocks were the same as those found in the Norwegian stocks, although the electrophoretic methods used lead to differences in designations of alleles. A polymorphism in ESTD * which involves a slow allele was commonly observed in the three northern populations of the Nilma, Kachkovka and Neiden rivers. This allele was absent in the other Norwegian stocks and in a major brood stock of farmed salmon in Norway. The IDHP-3 * 116 allele was found in unusually high frequencies in the northern populations. Thus, the variability observed at these two loci indicates a barrier to gene flow between the northern salmon stocks and the more southern stocks in the East Atlantic area.     

Salinity Adaptations and Bathymetric Distribution of Bivalve Mollusks Portlandia arcticaand Nuculana pernula in the White Sea

We examined changes in the oxygen consumption rate, the number of active animals, rate of losses of salts, and the level of cell hydration under conditions of decreased salinity in the mollusks Portlandia arctica(Gray) and Nuculana pernula(O.F. Müller), inhabiting the White Sea in the deepest depths under high salinity (about 30) and negative temperatures of –1.3 to –1.4°C throughout the year. It has been shown that the mollusks demonstrate a relatively great tolerance of desalination at the organism level and a capability to regulate cell volume under decreased salinity of the environment. We concluded that the depletion of fauna of the White Sea in the deepest depths is not connected with the tolerance level of deep-sea animals to salinity fluctuations, but rather is due to environmental conditions (liquid silts and negative temperatures).     

Life at the front: history, ecology and change on southern ocean islands

Terrestrial ecosystems of southern ocean islands have enjoyed renewed attention recently owing to the discovery that their climates are changing dramatically. This has led to an enhanced understanding of the biogeography of this region, and an increased awareness that these ecosystems provide unrivalled opportunities for investigating the impacts of environmental change on interactions between invasive and indigenous species. Recent studies have revealed increases in the abundance of established alien species and in the strength of their negative impacts on local biota, especially through indirect interactions. Also, increases in island temperature and human visitor frequency are likely to result in increasing numbers of successful alien colonization events.     

Turnover of passerine birds on islands in the Aegean Sea (Greece)

Aim We wish to determine the effect of migratory status on turnover rates in island birds. Because turnover is influenced by factors other than migratory status, we also considered the influence of body size and physical characteristics of the islands inhabited on the probabilities of extinction and immigration. Location The Mediterranean islands of Delos, Astypalea, Paros, Naxos and Lesvos in the Aegean Sea, Greece. Methods The passerine birds of these islands were surveyed between 1954 and 1961 by G.E. Watson, and were resurveyed between 1988 and 1992. The effects of migratory status and body size on the probabilities of extinction and immigration were examined by G‐tests of linear trend in proportion, and analysis of variance, respectively. A combined analysis of migratory status, body size and physical characteristics of the islands was carried out using logistic regressions of the probabilities of extinction and immigration on these factors. Results Species number on each island changed little between surveys, with no island's species number changing by more than one species. Twelve population extinctions and 11 immigrations were recorded. The smallest island, Delos (6 km²), had the highest annualized relative turnover rate (1.08), while the four larger islands (96–1614 km²) had lower and mutually similar rates (0.21–0.27). Populations on higher elevation islands were less likely to go extinct. There is no evidence for an effect of body size on the probabilities of extinction or immigration. Migratory status affected extinction and immigration probabilities differently: migratory species were more likely to immigrate, but less likely to go extinct. Main conclusions The position of the Aegean islands along a major north–south flyway may account for the observed effects of migratory status. The annual passage of large numbers of migrants may, via the rescue effect, decrease the chances of extinction, while at the same time increasing the chances of colonization of unoccupied islands. The likelihood of both extinction and immigration involves a complex interaction between life‐history traits and island characteristics. The effects of migratory status will depend not only on consideration of vagility, vulnerability and stochasticity identified by previous authors, but also upon the location of the islands in relationship to migratory pathways.     

Population dynamics of shrews on small islands accord with the equilibrium model

Three of the six species of shrew in Finland, Sorex araneus, S. caecutiens, and S. minutus , are common on the mainland and widespread on islands in lakes. The islands range from 0.01 to 500 ha in area, and from 10 to 3000 m in isolation (distance from the mainland). The species-area relationship, the lack of importance of habitat diversity, the increasing frequency of unoccupied small islands with isolation, and direct observations of small populations, all suggest that populations on small islands have a high extinction rate. Demographic stochasticity is the main cause of extinctions in the superior competitor, S. araneus , which occurs consistently on islands greater than 2 ha. The small species, S. caecutiens and S. minutus , are more sensitive to environmental stochasticity than is S. araneus , and are inferior to it in interspecific competition; these factors probably contribute to the absence of the small species from many islands tens of hectares in area. Frequent colonization of islands less than 500 m from the mainland is indicated by large numbers of shrews trapped from tiny islets where breeding is not possible, by increasing epigenetic divergence of island populations with isolation, and by observations of dispersal to and colonization of islands. Dispersal ability decreases with decreasing individual size, which may partly explain the absence of the small shrews from many relatively large islands. The shrew populations persist in a dynamic equilibrium on the islands. Epigenetic morphological variation is a useful tool in ecological studies of island populations.     

Lakes and rivers as islands: species-area relationships in the fish faunas of Ontario

Synopsis Fish species richness in 82 lakes in Ontario, Canada was significantly correlated with surface area. In this region, latitude explained only a small amount of the variation in fish species richness. Thus, our study provides a clear demonstration of the relation between fish species richness and lake area without the confounding effects of latitude and physiography inherent in analyses from broader geographic regions. By comparison with the species-area relationship obtained, we show that acidification clearly depressed the number of fish species in 66 acid-stressed lakes in Ontario. Fish species richness was also significantly correlated with both drainage and surface areas of 21 Ontario rivers. Slopes of species-area regressions of lakes and rivers did not differ significantly, suggesting that species are added to these habitats at similar rates. However, our regression analyses show that rivers support more species of fish per unit surface area of water. Although these results are consistent with some predictions of island biogeography theory, we suggest that fish species richness is more likely to be a simple function of habitat diversity, rather than an equilibrial balance between immigration and extinction.     

The effect of insularity on the diversity of land birds in the Fiji islands: implications for refuge design

Ecologists have argued over rules of thumb that could be used to set priorities in configuring systems of reserves for preserving biological diversity. To evaluate these simple strategies, I assembled a particularly large and comprehensive data set on the land birds of the Fiji archipelago. I analyzed the species distribution on 220 islands to compare the running total of species preserved by differènt sequences of adding nature reserves to a hypothetical reserve system, treating each island as if it were a potential reserve. A strategy of maximizing the number of islands contributing to any given reserve area (maximum fragmentation) is much more effective at including species than a strategy of maximizing the size of the island components of a reserve (minimum fragmentation). Nevertheless the maximum fragmentation strategy is not a very good one. It is less effective than many random strategies, especially when about 2–10% of total area is to be set aside as reserve, and when only rare species are considered. A computer program was used to determine an optimal strategy by maximizing the number of additional species added for each unit of area added. This strategy is always substantially more effective at encompassing species diversity than either the maximum or minimum fragmentation strategies. It is suggested that the poor performance of the minimum fragmentation strategy is due principally to the presence of many smaller-island endemics within the Fiji archipelago. More generally, it is argued that the effect of fragmentation on species diversity depends on the geographic scale and isolation of the region under consideration. In these respects the Fiji Archipelago may be a particularly good model for continental reserve systems.     

The Influence of Salinity on Early Ontogeny of the Mussel Mytilus edulisand the Starfish Asterias rubensfrom the White Sea

The influence of increased and decreased salinity on early larvae of the mussel Mytilus edulisL. and the starfish Asterias rubensfrom the White Sea is studied. Fertilized egg cells, embryos, larvae, and yearlings of the two species were compared. Salinity tolerance changes with the stage of development of mussels and starfish. Morphological changes occurring in eggs and embryos exposed to extremely high and low salinities are reported.     

Predicting biodiversity loss in island and countryside ecosystems through the lens of taxonomic and functional biogeography

We investigate how variation in patch area and forest cover quantified for three different spatial scales (buffer size of 500, 1500 and 3000 m radius) affects species richness and functional diversity of bat assemblages in two ecosystems differing in fragment–matrix contrast: a landbridge island system in Panama and a countryside ecosystem in the Brazilian Amazon. Bats were sampled on 11 islands and the adjacent mainland in Panama, and in eight forest fragments and nearby continuous forest in Brazil. Species–area relationships (SAR) were assessed based on Chao1 species richness estimates, and functional diversity–area relationships (FAR) were quantified using Chao1 functional diversity estimates measured as the total branch length of a trait dendrogram. FARs were calculated using three trait sets: considering five species functional traits (FAR_ALL), and trait subsets reflecting ‘diet breadth’ (FAR_DIET) and ‘dispersal ability’ (FAR_DISPERSAL). We found that in both study systems, FAR_ALL was less sensitive to habitat loss than SAR, in the sense that an equal reduction in habitat loss led to a disproportionately smaller loss of functional diversity compared to species richness. However, the inhospitable and static aquatic matrix in the island ecosystem resulted in more pronounced species loss with increasing loss of habitat compared to the countryside ecosystem. Moreover, while we found a significant FAR_DISPERSAL for the island ecosystem in relation to forest cover within 500 m landscape buffers, FAR_DIET and FAR_DISPERSAL were not significant for the countryside ecosystem. Our findings highlight that species richness and functional diversity in island and countryside ecosystems scale fundamentally differently with habitat loss, and suggest that key bat ecological functions, such as pollination, seed dispersal and arthropod suppression, may be maintained in fragments despite a reduction in species richness. Our study reinforces the importance of increasing habitat availability for decreasing the chances of losing species richness in smaller fragments.     

BIODIVERSITY RESEARCH: Nestedness for different reasons: the distributions of birds,lizards and small mammals on islands of an inundated lake

Aim We examined whether the community compositions of birds, lizards and small mammals were nested in a fragmented landscape in the Thousand Island Lake, China. We also assessed whether the mechanisms influencing nestedness differed among these taxonomic groups. Location Thousand Island Lake, China. Methods Presence/absence matrices were compiled for birds (42 islands) and lizards (42 islands) using line‐transect methods, and for small mammals (14 islands) using live‐trapping methods from 2006 to 2009. Nestedness was analysed using BINMATNEST, and statistical significance was assessed using the conservative null model 3. We used Spearman rank correlations and partial Spearman rank correlations to examine associations of nestedness and habitat variables (area, isolation, habitat diversity and plant richness) as well as life‐history traits (body size, habitat specificity, geographical range size and area requirement) related to species extinction and immigration tendencies. Results The community compositions of birds, lizards and small mammals were all significantly nested, but the causal factors underlying nestedness differed among taxonomic groups. For birds, island area, habitat specificity and area requirement were significantly correlated with nestedness after controlling for other independent variables. For lizards, habitat heterogeneity was the single best correlate of nestedness. For small mammals, island area, habitat heterogeneity and habitat specificity were significantly correlated with nestedness. The nested patterns of birds, lizards and small mammals were not attributable to passive sampling or selective colonization. Main conclusions The processes influencing nested patterns differed among taxonomic groups. Nestedness of bird assemblages was driven by selective extinction, and lizard assemblage was caused by habitat nestedness, while nestedness of small mammals resulted from both selective extinction and habitat nestedness. Therefore, we should take taxonomic differences into account when analysing nestedness to develop conservation guidelines and refrain from using single taxa as surrogates for others.