Spatial scale of observation affects α, β and γ diversity of cavity-nesting bees and wasps across a tropical land-use gradient

Aim Anthropogenic changes in land use may have major consequences for global biodiversity. However, species diversity is determined by a suite of factors that may affect species differently at different spatial scales. We tested the combined effects of land use and spatial scale on α, β and γ diversity in the tropics using experimental communities of cavity‐nesting bees and waSPS (Hymenoptera: Aculeata). We aimed to determine whether: (1) land‐use intensity negatively affects species richness of cavity‐nesting Hymenoptera, (2) β diversity, both within and between plots, is higher in more natural systems, (3) species richness of flowering herbs correlates positively with species richness of Hymenoptera within and across habitats, (4) richness of cavity‐nesting Hymenoptera in highly modified habitats declines with increasing distance from natural or semi‐natural habitats, (5) the effects of land use, herb diversity and forest distance on Hymenoptera α and β diversity vary at different spatial scales, and (6) bees and waSPS respond to land use in a similar way. Location Manabi, south‐west Ecuador. Methods We examined diversity (species richness) within 48 plots of five habitat types that comprised a gradient of decreasing agricultural intensity from rice and pasture to coffee agroforests, unmanaged abandoned agroforests and forest fragments, using standardized nesting resources for reproducing communities of cavity‐nesting bees and waSPS. Results (1) Land use significantly affected α diversity of trap‐nesting bees and waSPS at the subplot (per trap) scale, but not subplot β diversity or plot‐scale species richness (γ diversity). (2) Beta diversity was surprisingly higher between plots within a land‐use type than between land‐use types. (3) Species richness of bees and waSPS increased with diversity of flowering herbs at the subplot (trap) scale only. (4) Forest distance correlated positively with bee species richness at the plot scale only. (5) Land use, herb diversity and forest distance each showed significant correlations with bee and wasp diversity at only one spatial scale. (6) Despite differences in life history, bees and waSPS responded to land‐use intensity in a similar way. Main conclusions The effects of land use on species richness were highly dependent on spatial scale. Subplot‐scale analyses showed that rice and pasture contained the highest species diversity, whereas plot‐scale analyses showed no significant difference in the diversity of different land‐use types. We emphasize caution in the estimation of biodiversity at only one spatial scale, and highlight the surprisingly large contribution of managed land to the regional biodiversity of these species.     

Factors controlling the spatial species richness pattern of four groups of terrestrial vertebrates in an area between two different biogeographic regions in northern Spain

Aim To examine the influence of environmental variables on species richness patterns of amphibians, reptiles, mammals and birds and to assess the general usefulness of regional atlases of fauna. Location Navarra (10,421 km²) is located in the north of the Iberian Peninsula, in a territory shared by Mediterranean and Eurosiberian biogeographic regions. Important ecological patterns, climate, topography and land‐cover vary significantly from north to south. Methods Maps of vertebrate distribution and climatological and environmental data bases were used in a geographic information systems framework. Generalized additive models and partial regression analysis were used as statistical tools to differentiate (A) the purely spatial fraction, (B) the spatially structured environmental fraction and (C) the purely environmental fraction. In this way, we can evaluate the explanatory capacity of each variable, avoiding false correlations and assessing true causality. Final models were obtained through a stepwise procedure. Results Energy‐related features of climate, aridity and land‐cover variables show significant correlation with the species richness of reptiles, mammals and birds. Mammals and birds exhibit a spatial pattern correlated with variables such as aridity index and vegetation land‐cover. However, the high values of the spatially structured environmental fraction B and the low values of the purely environmental fraction A suggest that these predictor variables have a limited causal relationship with species richness for these vertebrate groups. An increment in land‐cover diversity is correlated with an increment of specific richness in reptiles, mammals and birds. No variables were found to be statistically correlated with amphibian species richness. Main conclusions Although aridity and land‐cover are the best predictor variables, their causal relationship with species richness must be considered with caution. Historical factors exhibiting a similar spatial pattern may be considered equally important in explaining the patterns of species richness. Also, land‐cover diversity appears as an important factor for maintaining biological diversity. Partial regression analysis has proved a useful technique in dealing with spatial autocorrelation. These results highlight the usefulness of coarsely sampled data and cartography at regional scales to predict and explain species richness patterns for mammals and birds. The accuracy of models appears to be related to the range perception of each group and the scale of the information.     

From natural woodlands to cultivated land: diversity of fruit‐feeding butterflies and beetles in the mid‐Zambezi valley,northern Zimbabwe

We tested the effect of cultivation on butterfly (Nymphalidae: Charaxes) and beetle (Coleoptera: Scarabaeidae: Cetoniinae) species richness and abundance along a cultivation intensification gradient. Results showed significant differences in species richness and abundance between natural woodlands and cultivated landscapes with larger differences in areas of high cultivation intensity. The results indicate that natural woodland clearing for cultivation purposes has negative impacts on arthropod diversity, a situation more severe in highly intensified cultivated areas. Our results imply that mosaics of different land‐use units, each in a different phase of clearance‐cultivation‐abandonment‐recovery‐clearance cycle could counter the negative effects of cultivation intensity on arthropod diversity.     

The Impact of Anthropogenic Disturbance on Assembly Patterns of Termite Communities

Intensification of land‐use threatens biodiversity, especially in tropical ecosystems that harbor the planet's highest species richness. This negative impact of anthropogenic disturbance on species numbers is well established, but the mechanisms underlying the community assembly processes are less well understood. Termites are of fundamental importance in tropical ecosystems where they are critical for nutrient recycling and species diversity. We tested the impact of anthropogenic disturbance on termite species diversity and assembly processes in a West African savanna applying the newest techniques of phylogenetic community analyses. Species richness dropped in areas of intensive land‐use and compositional similarity between intensive land‐use areas was high. This contrasted with a protected National Park where communities were characterized by high species richness and intermediate species turnover between sites. Slightly disturbed areas in the buffer zone surrounding the park were intermediate, they still had high species richness but similarity between sites increased. Strikingly, the assembly pattern also changed with disturbance from more phylogenetic overdispersion to more clustering (coexisting species became phylogenetically more similar), but only when the fungus‐growing termite Macrotermes bellicosus was absent. Our data suggest that the major forces structuring termite communities depend: (1) on the presence of this dominant mound‐building termite; and (2) that they change to more environmental filtering with disturbance. Anthropogenic disturbance seems to function as a filter that allows only a specific subset of species to occur. Such an effect might be widespread in ecology but it is difficult to document quantitatively. Phylogenetic community analyses can help to contribute such evidence.     

Rapid redistribution of agricultural land alters avian richness,abundance, and functional diversity

The conversion of natural, or seminatural, habitats to agricultural land and changes in agricultural land use are significant drivers of biodiversity loss. Within the context of land‐sharing versus land‐sparing debates, large‐scale commercial agriculture is known to be detrimental to biodiversity, but the effects of small‐scale subsistence farming on biodiversity are disputed. This poses a problem for sustainable land‐use management in the Global South, where approximately 30% of farmland is small‐scale. Following a rapid land redistribution program in Zimbabwe, we evaluated changes in avian biodiversity by examining richness, abundance, and functional diversity. Rapid land redistribution has, in the near term, resulted in increased avian abundance in newly farmed areas containing miombo woodland and open habitat. Conversion of seminatural ranched land to small‐scale farms had a negative impact on larger‐bodied birds, but species richness increased, and birds in some feeding guilds maintained or increased abundance. We found evidence that land‐use change caused a shift in the functional traits of the communities present. However, functional analyses may not have adequately reflected the trait filtering effect of land redistribution on large species. Whether newly farmed landscapes in Zimbabwe can deliver multiple benefits in terms of food production and habitat for biodiversity in the longer term is an open question. When managing agricultural land transitions, relying on taxonomic measures of diversity, or abundance‐weighted measures of function diversity, may obscure important information. If the value of smallholder‐farmed land for birds is to be maintained or improved, it will be essential to ensure that a wide array of habitat types is retained alongside efforts to reduce hunting and persecution of large bird species.     

Habitat loss alters effects of intransitive higher-order competition on biodiversity: a new metapopulation framework

Recent studies have suggested that intransitive competition, as opposed to hierarchical competition, allows more species to coexist. Furthermore, it is recognized that the prevalent paradigm, which assumes that species interactions are exclusively pairwise, may be insufficient. More importantly, whether and how habitat loss, a key driver of biodiversity loss, can alter these complex competition structures (and therefore species coexistence) remain unclear. We thus present a new, simple yet comprehensive metapopulation framework that can account for any competition pattern and more complex higher-order interactions (HOIs) among species. We find that competitive intransitivity increases community diversity and that HOIs generally enhance this effect. Essentially, intransitivity promotes species richness by preventing the dominance of a few species, unlike the hierarchical competition, while HOIs facilitate species coexistence through stabilizing community fluctuations. However, variation in species’ vital rates and habitat loss can weaken or even reverse such higher-order effects, as their interaction can lead to a more rapid decline in competitive intransitivity under HOIs. Thus, it is essential to correctly identify the most appropriate interaction model for a given system before models are used to inform conservation efforts. Overall, our simple model framework provides a more parsimonious explanation for biodiversity maintenance than the existing theory.     

Preemption of space can lead to intransitive coexistence of competitors

Intransitive competition has the potential to be a powerful contributor to species coexistence, but there are few proposed biological mechanisms that could create intransitivities in natural communities. Using a three‐species model of competition for space, we demonstrate a mechanism for coexistence that combines a colonization–competition tradeoff between two species with the ability of a third species to preempt space from the other competitors. The combination of differential abilities to colonize, preempt, and overtake space creates a community where no single species can exclude both of its competitors. The dynamics of this kind of community are analogous to rock‐paper‐scissors competition, and the three‐species community can persist even though not all pairs of species can coexist in isolation. In distinction to prior results, this is a mechanism of intransitivity that does not require nonhierarchical local interference competition. We present parameter estimates from a subtidal marine community illustrating how documented competitive traits can lead to preemption‐based intransitivities in natural communities, and we describe methods for an empirical test of the occurrence of this mechanism.     

锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究

用样带法研究了草原群落植物多样性和初级生产力沿海拔和水分梯度的变化,结果表明;在样带梯度上,物种丰富度、多样性和群落初级生产力与海拔高度、年降水量和土壤有机C及全N含量呈正相关,而与年平均气温和干燥度呈负相关。随着海拔高度的降低,降水量的减少,热量和干燥度的增加,以及土壤有机C和全N含量的降低,草原群落的物种丰富度、多样性和初级生产力逐渐降低。典型相关分析的结果揭示出,土壤有机C含量和干燥度是对草原群落物种丰富度和初级生产力具有更大的影响。同时,草地的利用方式和强度对群落植物多样性和生产力也具有较强的影响。     

锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究

 用样带法研究了草原群落植物多样性和初级生产力沿海拔和水分梯度的变化,结果表明;在样带梯度上,物种丰富度、多样性和群落初级生产力与海拔高度、年降水量和土壤有机C及全N含量呈正相关,而与年平均气温和干燥度呈负相关。随着海拔高度的降低,降水量的减少,热量和干燥度的增加,以及土壤有机C和全N含量的降低,草原群落的物种丰富度、多样性和初级生产力逐渐降低。典型相关分析的结果揭示出,土壤有机C含量和干燥度是对草原群落物种丰富度和初级生产力具有更大的影响。同时,草地的利用方式和强度对群落植物多样性和生产力也具有较强的影响。     

锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究

 用样带法研究了草原群落植物多样性和初级生产力沿海拔和水分梯度的变化,结果表明;在样带梯度上,物种丰富度、多样性和群落初级生产力与海拔高度、年降水量和土壤有机C及全N含量呈正相关,而与年平均气温和干燥度呈负相关。随着海拔高度的降低,降水量的减少,热量和干燥度的增加,以及土壤有机C和全N含量的降低,草原群落的物种丰富度、多样性和初级生产力逐渐降低。典型相关分析的结果揭示出,土壤有机C含量和干燥度是对草原群落物种丰富度和初级生产力具有更大的影响。同时,草地的利用方式和强度对群落植物多样性和生产力也具有较强的影响。     

Land‐use change promotes avian diversity at the expense of species with unique traits

Land‐use change may alter both species diversity and species functional diversity patterns. To test the idea that species diversity and functional diversity changes respond in differing ways to land‐use changes, we characterize the form of the change in bird assemblages and species functional traits along an intensifying gradient of land use in the savanna biome in a historically homogeneous vegetation type in Phalaborwa, South Africa. A section of this vegetation type has been untransformed, and the remainder is now mainly characterized by urban and subsistence agricultural areas. Using morphometric, foraging and breeding functional traits of birds, we estimate functional diversity changes. Bird species richness and abundance are generally higher in urban and subsistence agricultural land uses, as well as in the habitat matrix connecting these regions, than in the untransformed area, a pattern mainly driven through species replacement. Functionally unique species, particularly ground nesters of large body size, were, however, less abundant in more utilized land uses. For a previously homogenous vegetation type, declines in the seasonality of energy availability under land‐use change have led to an increase in local avian diversity, promoting the turnover of species, but reduced the abundance of functionally unique species. Although there is no simple relationship between land‐use and diversity change, land‐use change may suit some species, but such change may also involve functional homogenization.     

Scale-dependent effects of land use on plant species richness of mountain grassland in the European Alps

Traditionally managed mountain grasslands in the Alps are species‐rich ecosystems that developed during centuries of livestock grazing. However, changes in land use including fertilisation of well accessible pastures and gradual abandonment of remote sites are increasingly threatening this diversity. In five regions of the Swiss and French Alps we assessed the relationship between land use, soil resource availability, cover of the unpalatable species Veratrum album, species richness and vegetation composition of mountain grasslands across four spatial scales ranging from 1 to 1000 m². Mean species richness and the increase in the number of species with increasing area were lower in intensively grazed, fertilised pastures than in traditional pastures or in abandoned pastures. Species composition of abandoned pastures differed from that of the other management types. Plant species richness was influenced by different factors at different spatial scales. At the 1 m² scale, plant species richness was negatively related to soil nitrate and influenced by the cover of V. album, depending on land use: species richness and cover of V. album were negatively correlated in abandoned pastures, but positively correlated in fertilised grasslands. At the 1000 m² scale, a negative effect of fertilization on richness was evident. These results indicate that at small scales species richness in mountain grasslands is determined by competition for light, which should be more important if nutrient availability is high, and by positive and negative interactions with unpalatable plants. In contrast, species richness at the large scale appears to be mainly influenced by land use. This result emphasizes the importance of studying such inter‐relationships at multiple scales. Our study further suggests that the maintenance of the traditional land use scheme is crucial for the conservation of plant species richness of mountain pastures as both intensification and abandonment changed species composition and reduced plant species diversity.     

Ecosystem engineering effects on species diversity across ecosystems: a meta‐analysis

Ecosystem engineering is increasingly recognized as a relevant ecological driver of diversity and community composition. Although engineering impacts on the biota can vary from negative to positive, and from trivial to enormous, patterns and causes of variation in the magnitude of engineering effects across ecosystems and engineer types remain largely unknown. To elucidate the above patterns, we conducted a meta‐analysis of 122 studies which explored effects of animal ecosystem engineers on species richness of other organisms in the community. The analysis revealed that the overall effect of ecosystem engineers on diversity is positive and corresponds to a 25% increase in species richness, indicating that ecosystem engineering is a facilitative process globally. Engineering effects were stronger in the tropics than at higher latitudes, likely because new or modified habitats provided by engineers in the tropics may help minimize competition and predation pressures on resident species. Within aquatic environments, engineering impacts were stronger in marine ecosystems (rocky shores) than in streams. In terrestrial ecosystems, engineers displayed stronger positive effects in arid environments (e.g. deserts). Ecosystem engineers that create new habitats or microhabitats had stronger effects than those that modify habitats or cause bioturbation. Invertebrate engineers and those with lower engineering persistence (<1 year) affected species richness more than vertebrate engineers which persisted for >1 year. Invertebrate species richness was particularly responsive to engineering impacts. This study is the first attempt to build an integrative framework of engineering effects on species diversity; it highlights the importance of considering latitude, habitat, engineering functional group, taxon and persistence of their effects in future theoretical and empirical studies.     

Hot,dry and different: Australian lizard richness is unlike that of mammals,amphibians and birds

Aims (1) To map the species richness of Australian lizards and describe patterns of range size and species turnover that underlie them. (2) To assess the congruence in the species richness of lizards and other vertebrate groups. (3) To search for commonalities in the drivers of species richness in Australian vertebrates. Location Australia. Methods We digitized lizard distribution data to generate gridded maps of species richness and β‐diversity. Using similar maps for amphibians, mammals and birds, we explored the relationship between species richness and temperature, actual evapotranspiration, elevation and local elevation range. We used spatial eigenvector filtering and geographically weighted regression to explore geographical patterns and take spatial autocorrelation into account. We explored congruence between the species richness of vertebrate groups whilst controlling for environmental effects. Results Lizard richness peaks in the central deserts (where β‐diversity is low) and tropical north‐east (where β‐diversity is high). The intervening lowlands have low species richness and β‐diversity. Generally, lizard richness is uncorrelated with that of other vertebrates but this low congruence is strongly spatially structured. Environmental models for all groups also show strong spatial heterogeneity. Lizard richness is predicted by different environmental factors from other vertebrates, being highest in dry and hot regions. Accounting for environmental drivers, lizard richness is weakly positively related to richness of other vertebrates, both at global and local scales. Main conclusions Lizard species richness differs from that of other vertebrates. This difference is probably caused by differential responses to environmental gradients and different centres of diversification; there is little evidence for inter‐taxon competition limiting lizard richness. Local variation in habitat diversity or evolutionary radiations may explain weak associations between taxa, after controlling for environmental variables. We strongly recommend that studies of variation in species richness examine and account for non‐stationarity.     

Species richness and evenness respond to diverging land‐use patterns – a cross‐border study of dry tropical woodlands in southern Africa

Cross‐border studies offer unique situations to study the impact of different land‐use regimes on ecosystems. Along the Angolan and Namibian border formed by the Okavango River, the environmental conditions and traditional land‐use practises are the same on either side of the river. However, decades of civil war in Angola led to a stagnant development while political stability in Namibia fostered a recent socio‐economic transformation. We investigated the impact of spatially diffuse land use on plant diversity of the dry tropical woodlands covering the vast, sandy hinterlands of the river. As accessibility is the major factor governing land use, we used distance to road as a proxy for land‐use intensity. Based on 58 vegetation plots sized 20 m × 50 m, we showed that species richness increased with distance to road in Angola while in Namibia it remained constant on a lower level. Evenness showed an inverse pattern to species richness and Shannon diversity index showed no response. Analysing diversity patterns according to life forms revealed that these patterns are primarily driven by woody species. The study showed that spatially diffuse land use has a measurable effect on plant diversity and illustrates that roads act as vectors of change.     

Effects of grazing intensity on plant richness and diversity: a meta‐analysis

Most of our knowledge of the effect of grazing on grassland structure is based on grazed–ungrazed contrasts. The effects of grazing in the most common scenario, where grazing intensity varies from low to high grazing intensity, are less known. The objectives of this paper were to 1) quantify the effect of stocking rates on species richness and diversity of grasslands world‐wide, and 2) evaluate the response under different environmental and experimental conditions. We conducted a meta‐analysis of experiments with at least two levels of controlled stocking rates and evaluated their effect on species richness and diversity. The results showed that the response of richness and diversity to either reducing or increasing stocking rate from a moderate level mostly fell within the range  25% or  5 species. Mean response of species richness and diversity to increasing stocking rate from moderate to high levels was negative. Mean response to lowering stocking rate from moderate levels was not different from zero. However, overall, species richness significantly decreased as stocking rate increased. The response of richness and diversity to stocking rate was not related to mean precipitation, productivity or aridity. However, the most negative responses of richness to stocking rate were larger in arid, low productivity systems than in subhumid and humid systems. The effects of grazing on richness and diversity found in this review were smaller than the effects on species composition shown by the literature. Thus, grazing drastically changes species composition, but the net change of species and diversity is much smaller.     

From competition to facilitation: how tree species respond to neighbourhood diversity

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large‐scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource‐use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

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.