Bird community composition patterns in urban parks of Madrid: The role of age, size and isolation

This study intends to assess the influence of fragment age, size and isolation (from the regional species pool) on bird community composition patterns in urban parks in Madrid, and the role of local and regional factors on community structure. Park age was a good indicator of habitat complexity. Park age and area accounted for 62% of the variability in species richness, but two measures of isolation from the regional species pool were not included as significant factors. Species composition in urban parks showed a high degree of nestedness, which was associated with park age and area, but not with two measures of isolation from the regional species pool. The degree of nestedness increased with park age; the distribution of species varying from nested in old and mature parks to random in young parks. The incidence (% of species occurrence in parks) in young parks was correlated with regional densities, whereas in mature and old parks the incidence was correlated with local densities. In this urban landscape, species composition appears to be regulated by local factors (particularly in mature and old parks), such that species accumulate in an orderly (not random) fashion in relation to park age and area. Regional influences seem to be more pronounced only in young parks, which are mainly colonized by species from the regional species pool.     

Functional diversity of urban bird communities: effects of landscape composition,green space area and vegetation cover

In this study, we aim to gain a better insight on how habitat filtering due to urbanization shapes bird communities of Vienna city parks. This may help to derive implications for urban planning in order to promote and maintain high diversity and ecosystem function in an increasing urbanized environment. The structure of wintering bird communities of 36 Vienna city parks – surveyed once a month in January 2009, December 2009, December 2012, and January 2013 – was described by species richness and the functional diversity measurements FRic (functional richness), FEve (functional evenness), and FDiv (functional divergence). Environmental filtering was quantified by park size, canopy heterogeneity within the park, and the proportion of sealed area surrounding each park. Species richness, FRic, and FDiv increased with increasing park size. Sealed area had a strong negative effect on species richness and FDiv. Canopy heterogeneity played a minor role in explaining variance in FDiv data. FEve did not respond to any of these park parameters. Our results suggest a loss of species richness and functional diversity, hence most likely indicate a decline in ecosystem function, with decreasing park size and increasing sealed area of the surrounding urban landscape matrix.     

Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns

Summary Habitat subdivision by geography or human activity may be an important determinant of regional species richness. Cumulative species-area relationships for vertebrates, land plants, and insects on island archipelagoes show that collections of small islands generally harbor more species than comparable areas composed of one or a few large islands. The effect of the degree of habitat subdivision in increasing species richness appears to increase with the distance from potential sources of colonists. Mountaintop biotas show no clear differences between species richness on large alpine areas and collections of smaller peaks. National park faunas generally have more species in collections of small parks than in the larger parks. In all cases where a consistent effect of subdivision is observed, the more subdivided collection of islands or isolates contains more species. To the degree that these data provide guidance for establishing nature reserves, they suggest that increasing the numbers of reserves may be an important component of conservation strategies.     

Species richness and composition of amphibians and reptiles in a fragmented forest landscape in northeastern Bolivia

We quantified patterns of species richness and species composition of frogs and reptiles (lizards and snakes) among three habitats (continuous forest, forest islands, and a seasonally flooded savannah) and between forest island size and isolation classes in a floristic transition zone in northeastern Santa Cruz Department, Bolivia. Species richness was similar across macrohabitats, as was faunal composition of forested habitats, although savannah harbored a distinct herpetofauna. On forest islands, richness and composition of forest frogs was largely related to isolation, whereas reptiles were affected by both isolation and habitat. The observation that isolation rather than area was the primary driver of distribution patterns on forest islands stands in contrast to many studies, and may be a function of (1) the greater range in forest island isolation values compared to area or (2) the long history of isolation in this landscape.     

The adequacy of collecting techniques for estimating species richness of grassland invertebrates

1.  There is still considerable debate about the most effective methods of sampling invertebrates in monitoring and assessment programmes.
2.  The above-ground invertebrates of a limestone grassland in north-east England were compared between samples from pitfall traps and from a D-vac suction trap combined with a lightweight swish net (SW/DV).
3.  Over 14 000 individuals were captured, with similar numbers in the pitfall and SW/DV samples. A total of 480 species of Hemiptera, Coleoptera, Diptera and Araneae was identified and placed into 14 taxa for further analysis.
4.  The pitfall sample produced species/specimen curves from which it was possible to estimate species richness for all the Coleoptera and Araneae taxa and the calypterate Schizophora. The SW/DV sample was adequate to estimate the species richness of Hemiptera, most Diptera taxa, herbivorous Coleoptera and Linyphiidae.
5.  The proportion of Coleoptera and Araneae taxa that were method-unique was higher in the pitfall sample than the SW/DV sample and vice versa for the Hemiptera and Diptera taxa. Nevertheless, a relatively high proportion of method-unique species of most taxa was found in both sample types, indicating that they can each contribute to assessing species assemblages in grasslands.
6.  Both pitfall traps and SW/DV samples are needed to estimate species richness in grasslands for all taxa except Heteroptera, Homoptera and Lycosidae. Herbivorous Coleoptera and Linyphiidae were collected in numbers adequate for assessing richness in both sample types, but more specimens were required in one or other sample for the remaining taxa.     

Influence of settlement time, human population, park shape and age, visitation and roads on the number of alien plant species in protected areas in the USA

Abstract. I examined a data set of 77 protected areas in the USA (including national and state parks) to determine which of the following variables most strongly influence alien plant species richness: park area, climate (temperature and precipitation), native species richness, visitation rate, local human population size, total road length, park shape and duration of European settlement. Many of these predictor variables are intercorrelated, so I used multiple regression to help separate their effects. In support of previous studies, native species richness was the best single predictor of alien species richness, probably because it was a good estimator of both park area and habitat diversity available for establishment of alien species. Other significant predictors of alien species richness were years of occupation of the area by European settlers and the human population size of adjacent counties. Climate, visitation rate, road length and park shape did not influence alien species richness. The proportion of alien species (alien richness/native richness) is inversely related to park area, in agreement with a previous study. By identifying which variables are most important in determining alien species richness, such findings suggest ways to reduce alien species establishment.     

基于物种落差分析的公园鸟类多样性提升设计——以湖南常德螺湾湿地公园为例

随着城市化发展,许多公园鸟类栖息环境遭受破坏,导致当前的鸟类多样性与历史上的和周边的都存在较大差异,为此,我们从项目区域的当前-周边-历史三者间在鸟类物种及其栖息地上的落差分析入手,以此确定目标物种及其栖息地,并加以归类,从而有针对性地进行相关设计。2016年5月至2017年3月在湖南常德柳叶湖螺湾湿地公园开展的鸟类多样性提升设计中,通过实地调查并结合文献查阅,记录到项目区域16种,周边79种,历史上146种鸟类,运用三者间的物种落差分析法确定主要的21种可恢复目标物种和4种栖息地类型,结合项目区域及周边空间特点,将项目区域划分为4个区域,在各区域内设计并营建相应的栖息地,再配套采用多种鸟类招引措施。至2017年3月,项目区鸟类已可实地观察到39种,并于2017年12月吸引到超过2000只的野鸭群来此越冬。以上实际效果表明,设计有效地提升了湿地公园鸟类多样性且增加了景观要素,可充分发挥生物保护功能。     

Influence of urbanization on ant distribution in parks of Tokyo and Chiba City,Japan I. Analysis of ant species richness

To investigate how progressive urbanization influences the distribution of ant species in cities, I compared the ant species richness in urban parks of different areas and ages of Tokyo, the most intensively developed urban region in Japan, and its developing neighbor, Chiba City. A total of 43 ant species were found from 98 parks. Multiple regression analysis revealed that park area and age had a positive effect on the number of ant species in the parks and that the parks in Tokyo contained fewer species than did comparable parks in Chiba. Thus, the progression of urbanization reduces ant species richness in urban parks, most likely because it isolates the parks from the surrounding area. Next, in order to examine the relationships between the types of landform modification and the ant distribution in urban areas, I compared the ant species richness in urban parks of Chiba City among different landform types (upland + terraces, lowlands, cut and fill, banking, and reclamation). This study showed that the parks in the reclaimed lands and lowlands contained fewer ant species than did parks in other areas. The reason for this decline of ant fauna is discussed.     

Effects of seasonality on the species–area relationship: a case study with birds in urban parks

Aim To examine how parameters of the species–area curve and factors determining bird species richness are affected by seasonality. Location One hundred and thirty urban parks of the city of Valencia (Spain) ranging between 0.03 and 18.6 ha. Methods Bird censuses were conducted monthly during 1998 and 2004. For each park, 27 variables were recorded as measures of size, structure and isolation. Linear regression was used to test for patterns in species richness relative to independent variables. The results of each season were compared. Results Curves corresponding to the breeding period (spring–summer) had significantly higher intercepts, but slopes did not change across seasons. Turnover rates of resident species were dependent on patch size and isolation considering spring, but not autumn, from different years. Additionally, turnover rates in spring were lower in parks in close proximity to others than in isolated ones. In all seasons, the number of bird species showed a strong relationship with park area, which explained c. 60% of the variation in species numbers. Habitat variables and park isolation explained a small amount of the variation in species numbers. The data did not give support to the random placement hypothesis. Main conclusions The overall conclusions of this study give little support to the possible influence of seasonality on species–area relationship or on the three main hypotheses explaining it. This lack of influence of seasonality could be explained by the seasonal stability of the bird populations of the parks within the study area as a consequence of different factors, mainly the characteristics of the species pool, similarity in the features of the parks irrespective of their size, and stability of resources and conditions across seasons. The question of whether this lack of influence of seasonality on the species–area relationship occurs in those ecological systems that have a larger seasonal variation than the urban parks of the study area remains unresolved. Closer examination of the seasonal patterns reported in this study is likely to be useful in increasing our understanding of the species–area relationship.     

Determinants of regional species richness: an empirical analysis of the number of hawkmoth species (Lepidoptera: Sphingidae) on the Malesian archipelago

Aims Major patterns and determinants of the species richness of Sphingidae in the Malesian archipelago were investigated, including a distinction of richness patterns between subfamilies and range‐size classes. Location Southeast Asia, Malesia. Methods Using a compilation of specimen‐label data bases, geographic information system (GIS)‐supported estimates of distributional ranges for all Sphingidae species of Southeast Asia were used to assess the species richness of islands. Range maps for all species and checklists for 114 islands can be found at http://www.sphingidae‐sea.biozentrum.uni‐wuerzburg.de . Potential determinants of the species richness of islands were tested with general linear models. Results The estimated species richness of islands in the region is determined by biogeographical association, seasonality, availability of rain forest and island size. Species–area relationships are linear on a semi‐logarithmic representation, but not on a double‐logarithmic scale. Species richness of all sphingid subfamilies is influenced by biogeography. The presence of large rain‐forest areas affects mainly Smerinthinae, whereas distance from continental Asia is conspicuously irrelevant for this group. Widespread rather than geographically restricted species shape the overall distribution patterns of species richness. The altitudinal range of islands does not significantly affect species‐richness patterns, but its potential effects on geographically restricted species are discussed. Main conclusions As well as being affected by climatic and vegetation parameters, sphingid species richness is strongly influenced by a historical, directional dispersal process from continental Southeast Asia to the Pacific islands. This process did not apply equally to species of different taxonomic groups or range sizes. Widespread species decline in species richness towards the south‐east, whereas geographically restricted species exhibit an inverse pattern of species richness, probably because speciation becomes more important in this group within the more isolated island groups.     

Lizard predation alters the effect of habitat area on the species richness of insect assemblages on Bahamian isles

Aim Anolis lizard invasions are a serious threat world‐wide, and information about how this invasive predator affects the diversity of prey assemblages is important for many strategic conservation goals. It is hypothesized that these predators reduce the slope of species–area relationships (SARs) of their prey assemblages. The effects of island area and predation by anolis lizards on the species richness of insular insect assemblages were investigated. Location Twenty‐four isles around Staniel Cay, Exuma Cays, Bahamas. Methods Flying insects were sampled using half‐sized Malaise traps for three consecutive days on each island in May 2007. First, the effect of island area on the probability of lizard presence was evaluated. Then, the effects of the presence–absence of predatory lizards on SARs were analysed for the overall insect assemblage and for the assemblages of five dominant insect orders. Results Our results indicated that anolis lizards occurred primarily on larger islands. The species richness of the overall insect assemblage and five dominant insect orders significantly increased with island area. The interaction between island area and predator presence–absence significantly affected the overall insect assemblage and Diptera and Hymenoptera assemblages (but not Coleoptera, Hemiptera and Lepidoptera assemblages). The presence of predators caused decreases in the slope of the SARs. Main conclusions The presence of predatory lizards strongly affects species richness of insular insect assemblages with the island area being a crucial determinant of the species richness. Therefore, the slope of the SAR can serve as a measure of the consequence of invasive predatory species on native insect assemblages.     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.     

Terrestrial bird community patterns on the coralline islands of the Dahlak Archipelago, Red Sea, Eritrea

Aim This study aims to explain the patterns of species richness and nestedness of a terrestrial bird community in a poorly studied region. Location Twenty‐six islands in the Dahlak Archipelago, Southern Red Sea, Eritrea. Methods The islands and five mainland areas were censused in summer 1999 and winter 2001. To study the importance of island size, isolation from the mainland and inter‐island distance, I used constrained null models for the nestedness temperature calculator and a cluster analysis. Results Species richness depended on island area and isolation from the mainland. Nestedness was detected, even when passive sampling was accounted for. The nested rank of islands was correlated with area and species richness, but not with isolation. Idiosyncrasies appeared among species‐poor and species‐rich islands, and among common and rare species. Cluster analysis showed differences among species‐rich islands, close similarity among species‐poor and idiosyncratic islands, and that the compositional similarity among islands decreased with increasing inter‐island distance. Thus, faunas of species‐poor, smaller islands were more likely to be subsets of faunas of species‐rich, larger islands if the distance between the islands was short. Main conclusions Species richness and nestedness were related to island area, and nestedness also to inter‐island distances but not to isolation from the mainland. Thus, nestedness and species richness are not affected in the same way by area and distance. Moreover, idiosyncrasies may have been the outcome of species distributions among islands being influenced also by non‐nested distributions of habitats, inter–specific interactions, and differences in species distributions across the mainland. Idiosyncrasies in nested patterns may be as important as the nested pattern itself for conservation – and conservation strategies based on nestedness and strong area effects (e.g. protection of only larger islands) may fail to preserve idiosyncratic species/habitats.     

Biogeography of mammals on tropical Pacific islands

Aim We examine the influence of geography on species richness and endemism of mammals on tropical Pacific archipelagos to determine the importance of intra‐ and inter‐archipelago speciation in promoting local and regional species richness. Location Thirty tropical Pacific archipelagos. Methods A distributional list of mammals on 30 archipelagos was compiled, and values for 10 geographical variables were estimated for each archipelago. Mammal species were placed in three different categories (continental, Pacific and endemic) based on their distribution. The total number of species and numbers of species within each category were related to the geographical variables using Poisson regression analysis. Results Species richness was related positively to variables describing land area, numbers of large islands and elevation; and negatively to variables describing isolation. Levels of endemism did not differ between volant and non‐volant species, but differed between mega‐ and microchiropterans. Main conclusions Variation in species richness of mammals in the tropical Pacific region can be accounted for by a combination of intra‐archipelago speciation within archipelagos composed of large islands, and inter‐archipelago speciation, particularly among more isolated archipelagos. Mammals were less widely distributed throughout the study area than previously found for butterflies, skinks or birds. However, the level of endemism was similar to that of skinks and birds on the same archipelagos, and was higher than that of butterflies.     

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.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

Ranking the Azorean Natural Forest Reserves for Conservation Using their Endemic Arthropods

Endemic arthropods were used to evaluate the conservation value of the 16 Natural Forest Reserves (NFRs) of the Azores (Macaronesia). For each of the 280 known Azorean endemic species of arthropods, a rarity index was calculated, using distribution and abundance data obtained from the literature. In addition, several scoring indices were used to rank the 16 NFRs. Frequency distributions of the rarity index indicated that there was a tendency for a greater proportion of the commonest species being represented in the NFRs in contrast with a lower representation of the rarest species. About 60% of the endemic arthropod species that were recorded from the NFRs are single NFR endemics, that is, are known from only one of the 16 NFRs. Species richness was considered to be a very good surrogate measure of the conservation value of the 16 NFRs under study. The fact that the six highest ranked NFRs (using a composite multi-criteria index) are located in different islands has some important conservation management implications; to preserve a large proportion of the Azorean arthropod biodiversity there is a need to protect sites in all islands. If the five highest ranked NFRs are correctly managed in terms of conservation, then at least 80% of the endemic arthropods known from the NFRs could be protected. Most of the tested taxa (Acari-Oribatei; Lepidoptera; Diptera; Coleoptera) are good surrogates of the overall total set of species present in the 16 NFRs when using a species richness index.     

Vascular plant species richness in relation to habitat diversity and island area in the Finnish Archipelago

Aim The aim of this study is to explore the interrelationships between island area, species number and habitat diversity in two archipelago areas. Location The study areas, Brunskär and Getskär, are located in an archipelago in south‐western Finland. Methods The study areas, 82 islands in Brunskär and 78 in Getskär, were classified into nine habitat types based on land cover. In the Brunskär area, the flora (351 species) was surveyed separately for each individual habitat on the islands. In the Getskär area, the flora (302 species) was surveyed on a whole‐island basis. We used standard techniques to analyse the species–area relationship on a whole‐island and a habitat level. We also tested our data for the small island effect (SIE) using breakpoint and path analysis models. Results Species richness was significantly associated with both island area and habitat diversity. Vegetated area in particular, defined as island area with the rock habitat subtracted, proved to be a strong predictor of species richness. Species number had a greater association with island area multiplied by the number of habitats than with island area or habitat number separately. The tests for a SIE in the species–area relationship showed the existence of a SIE in one of the island groups. No SIE could be detected for the species–vegetated area relationship in either of the island groups. The strength of the species–area relationship differed considerably between the habitats. Main conclusions The general principles of island biogeography apply well to the 160 islands in this study. Vascular plant diversity for small islands is strongly influenced by physiographic factors. For the small islands with thin and varying soil cover, vegetated area was the most powerful predictor of species richness. The species–area curves of various habitats showed large variations, suggesting that the measurement of habitat areas and establishment of habitat‐based species lists are needed to better understand species richness on islands. We found some evidence of a SIE, but it is debatable whether this is a ‘true’ SIE or a soil cover/habitat characteristics feature.     

Sequential colonization and diversification of Galapágos endemic land snail genus Bulimulus (Gastropoda, Stylommatophora)

Species richness on island or islandlike systems is a function of colonization, within-island speciation, and extinction. Here we evaluate the relative importance of the first two of these processes as a function of the biogeographical and ecological attributes of islands using the Galápagos endemic land snails of the genus Bulimulus, the most species-rich radiation of these islands. Species in this clade have colonized almost all major islands and are found in five of the six described vegetation zones. We use molecular phylogenetics (based on COI and ITS 1 sequence data) to infer the diversification patterns of extant species of Bulimulus, and multiple regression to investigate the causes of variation among islands in species richness. Maximum-likelihood, Bayesian, and maximum-parsimony analyses yield well-resolved trees with similar topologies. The phylogeny obtained supports the progression rule hypothesis, with species found on older emerged islands connecting at deeper nodes. For all but two island species assemblages we find support for only one or two colonization events, indicating that within-island speciation has an important role in the formation of species on these islands. Even though speciation through colonization is not common, island insularity (distance to nearest major island) is a significant predictor of species richness resulting from interisland colonization alone. However, island insularity has no effect on the overall bulimulid species richness per island. Habitat diversity (measured as plant species diversity), island elevation, and island area, all of which are indirect measures of niche space, are strong predictors of overall bulimulid land snail species richness. Island age is also an important independent predictor of overall species richness, with older islands harboring more species than younger islands. Taken together, our results demonstrate that the diversification of Galápagos bulimulid land snails has been driven by a combination of geographic factors (island age, size, and location), which affect colonization patterns, and ecological factors, such as plant species diversity, that foster within-island speciation.