The Magnitude of Local Host Specif icity for Phytophagous Insects and its Implications for Estimates of Global Species Richness

Abstract: Estimates of the global number of arthropod species range from 2 million to more than 30 million species. One of the most critical assumptions affecting the higher of these estimates is the assumed magnitude of host specificity of phytophagous insects, which varies considerably. Difficulties in estimating this value are caused by both lack of satisfactory data sets and lack of methods for its objective calculation. We provide a new method for predicting host specificity for phytophagous insects at a local scale based on the concept of effective specialization. When the insect-plant host associations of a restricted number of plant species are known, we can predict the magnitude of host specificity when more plant species are included. Based on 2561 host observations of 697 beetle species on 50 plant species in the canopies of a tropical dry forest in Panama, we predict that host specificity for adult phytophagous beetles in this forest range from 7% to 10%. These values suggest that the higher estimates of global species richness are not tenable.     

Effects of Growth Form and Functional Traits on Response of Woody Plants to Clearing and Fragmentation of Subtropical Rainforest

The conservation implications of large‐scale rainforest clearing and fragmentation on the persistence of functional and taxonomic diversity remain poorly understood. If traits represent adaptive strategies of plant species to particular circumstances, the expectation is that the effect of forest clearing and fragmentation will be affected by species functional traits, particularly those related to dispersal. We used species occurrence data for woody plants in 46 rainforest patches across 75,000 ha largely cleared of forest by the early 1900s to determine the combined effects of area reduction, fragmentation, and patch size on the taxonomic structure and functional diversity of subtropical rainforest. We compiled species trait values for leaf area, seed dry mass, wood density, and maximum height and calculated species niche breadths. Taxonomic structure, trait values (means, ranges), and the functional diversity of assemblages of climbing and free‐standing plants in remnant patches were quantified. Larger rainforest patches had higher species richness. Species in smaller patches were taxonomically less related than species in larger patches. Free‐standing plants had a high percentage of frugivore dispersed seeds; climbers had a high proportion of small wind‐dispersed seeds. Connections between the patchy spatial distribution of free‐standing species, larger seed sizes, and dispersal syndrome were weak. Assemblages of free‐standing plants in patches showed more taxonomic and spatial structuring than climbing plants. Smaller isolated patches retained relatively high functional diversity and similar taxonomic structure to larger tracts of forest despite lower species richness. The response of woody plants to clearing and fragmentation of subtropical rainforest differed between climbers and slow‐growing mature‐phase forest trees but not between climbers and pioneer trees. Quantifying taxonomic structure and functional diversity provides an improved basis for conservation planning and management by elucidating the effects of forest‐area reduction and fragmentation. Efectos de la Forma de Crecimiento y Atributos Funcionales en la Respuesta de Plantas Leñosas al Desmonte y Fragmentación de Bosque Lluvioso Subtropical     

Using Vascular Plants as a Surrogate Taxon to Maximize Fungal Species Richness in Reserve Design

Abstract:  Fungi are a hyperdiverse taxonomic group that may be disappearing at a very high rate. Identifying fungal species is difficult in the field, and the use of highly specialized taxonomists is required. Data and expertise on vascular plants are, on the other hand, much more common and easy to find. We tested the potential of using vascular plants as surrogates to select reserve sites that maximize the pooled number of fungal species. We used data from 25 forest plots in Tuscany, Italy, that were sampled for woody plants, all other plants, and fungi. Species richness of woody plants and all other plants did not correlate with species richness of fungi. The gradients in species composition were similar among the three considered groups, as indicated by a detrended correspondence analysis ordination and species complementarity between pairs of plots. Fungal communities of the 25 plots had a lower β diversity than plant communities, and there were no pairs of totally complementary sites. Site prioritization for conservation was obtained through integer linear programming to find for any given number of sites those combinations containing the maximum pooled species richness of woody plants or all plants. The combinations of sites obtained by optimizing vascular plant species did not maximize the pooled species richness of fungi, whereas those obtained by maximizing woody plant species provided better results for sets of four to eight plots, but not for all the possible combinations. These results indicated that, in general, vascular plants cannot be used to maximize fungal species richness.     

A Successful Predictive Model of Species Richness Based on Indicator Species

Abstract:  Because complete species inventories are expensive and time-consuming, scientists and land managers seek techniques to alleviate logistic constraints on measuring species richness, especially over large spatial scales. We developed a method to identify indicators of species richness that is applicable to any taxonomic group or ecosystem. In an initial case study, we found that a model based on the occurrence of five indicator species explained 88% of the deviance of species richness of 56 butterflies in a mountain range in western North America. We validated model predictions and spatial transferability of the model using independent, newly collected data from another, nearby mountain range. Predicted and observed values of butterfly species richness were highly correlated with 93% of the observed values falling within the 95% credible intervals of the predictions. We used a Bayesian approach to update the initial model with both the model-building and model-validation data sets. In the updated model, the effectiveness of three of the five indicator species was similar, whereas the effectiveness of two species was reduced. The latter species had more erratic distributions in the validation data set than in the original model-building data set. This objective method for identifying indicators of species richness could substantially enhance our ability to conduct large-scale ecological assessments of any group of animals or plants in any geographic region and to make effective conservation decisions.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

Effects of Land Use on Bird Species Richness in Sulawesi, Indonesia

Abstract:  There is still much debate over the potential value of land-use systems for the maintenance of tropical biodiversity. An increasing number of studies indicate that much forest biodiversity can also be found in the agricultural landscape matrix. Because there is little information on the potential value of land-use systems for tropical forest bird species, we conducted repeated point counts in near-primary forest, adjacent young secondary forest, modernized cacao agroforestry systems, and annual cultures at submontane elevations in central Sulawesi, Indonesia. Species richness decreased from natural forest and young secondary forest to agroforestry systems and annual cultures. Although species richness was similar between natural and young secondary forest, the number of endemic bird species was significantly lower in second-growth forest. Species composition gradually changed as the habitat changed from natural to secondary forest, agroforestry systems, and annual cultures. Despite close proximity to near-primary forest, the agroforestry systems studied supported only a few small frugivorous-nectarivorous species. Our results suggest that secondary forest could play an important role in the conservation of many Sulawesi bird species, but, although suitable for colonization, its potential to sustain populations over the long term is unknown. Improvement of the landscape matrix for biodiversity conservation through secondary habitats therefore seems desirable to enlarge the ranges of forest species, but the fight against land conversion within protected areas of the region should be of much higher importance, at least at present.     

Effects of Earthworm Invasion on Plant Species Richness in Northern Hardwood Forests

Abstract:  The invasion of non-native earthworms ( Lumbricus spp.) into a small number of intensively studied stands of northern hardwood forest has been linked to declines in plant diversity and the local extirpation of one threatened species. It is unknown, however, whether these changes have occurred across larger regions of hardwood forests, which plant species are most vulnerable, or with which earthworm species such changes are associated most closely. To address these issues we conducted a regional survey in the Chippewa and Chequamegon national forests in Minnesota and Wisconsin (U.S.A.), respectively. We sampled earthworms, soils, and vegetation, examined deer browse in 20 mature, sugar-maple-dominated forest stands in each national forest, and analyzed the relationship between invasive earthworms and vascular plant species richness and composition. Invasion by Lumbricus was a strong indicator of reduced plant richness in both national forests. The mass of Lumbricus juveniles was significantly and negatively related to plant-species richness in both forests. In addition, Lumbricus was a significant factor affecting plant richness in a full model that included multiple variables. In the Chequamegon National Forest earthworm mass was associated with higher sedge cover and lower cover of sugar maple seedlings and several forb species. The trends were similar but not as pronounced in Chippewa, perhaps due to lower deer densities and different earthworm species composition. Our results provide regional evidence that invasion by Lumbricus species may be an important mechanism in reduced plant-species richness and changes in plant communities in mature forests dominated by sugar maples.     

A Century of Plant Species Loss from an Isolated Fragment of Lowland Tropical Rain Forest

An isolated 4-ha fragment of lowland tropical rain forest has been preserved in the Singapore Botanic Gardens since their founding in 1859. The Botanic Gardens'Jungle has recently had enumerated all woody stems 5 cm diameter at breast height (dbh) and larger and the complete vascular plant flora inventoried. This inventory can be compared with the historic record of the flora of the Gardens'Jungle obtained from the extensive collection of herbarium specimens dating back to the 1890s. Of the 448 historically recorded native species, 220 are still present. Ninety-four native species for which there were no historic records and 80 introduced species were also recorded in the recent inventory. The 50.9% loss of plant species richness over approximately the last century has not been distributed uniformly across plant life-form groups. Tree species have been less likely to go extinct than shrubs, climbers, or epiphytes. But half of the tree species present in 1994 were represented by only one or two individuals ≥ 5 cm dbh and larger. Individual longevity may be the major correlate with persistence of plant species in isolated forest fragments. Shade-tolerant understory shrubs (mostly Rubiaceae ) and rattans ( Palmae ) have been particularly prone to extinction. Some species have probably proliferated during the period of isolation. The tree Calophyllum ferrugineum currently constitutes one quarter of all woody stems. A group of climbers has become very common and covers large areas, probably inhibiting tree regeneration. We conclude that tiny fragments will act as refuges for tropical rain-forest plant species for decades, possibly even centuries after isolation but on their own they will not provide a permanent guarantee of the conservation of tropical biodiversity.     

Cost-Efficiency of Decaying Wood as a Surrogate for Overall Species Richness in Boreal Forests

Abstract:  Decaying wood is one of the most important elements for species richness in boreal forests. We tested how well reserve selection based on the amount and quality of decaying wood results in a representation of four ecologically different taxa (beetles, birds, wood-inhabiting fungi, and vascular plants). We also compared the cost-efficiency of the use of dead-wood indicators with comprehensive species inventory. Our database included 32 seminatural old-forest stands located in northern Finland. Decaying wood was a relatively good indicator of saproxylic species but not overall species richness. Even though dead wood did not reflect accurately overall species richness, our results indicated that the use of decaying wood as an indicator in site selection was more cost-efficient than using information from large-scale species inventories. Thus, decaying wood is a valuable surrogate for species richness, but other cost-efficient indicators that reflect the requirements of those species which are not dependent on decaying wood should be identified.     

Beyond Species Richness: Community Similarity as a Measure of Cross-Taxon Congruence for Coarse-Filter Conservation

Abstract:  The use of a surrogate taxon in conservation planning has become questionable because recent evidence suggests that the correlation of species richness between pairs of taxa is highly variable both taxonomically and geographically. Species richness is only one measure of species diversity, however, and recent studies suggest that investigations of cross-taxon congruence should consider a broader range of assessment techniques. The cross-taxon congruence of community similarity between sites among taxa has rarely been examined and may be the most relevant measure of species diversity in the context of coarse-filter conservation strategies. We examined cross-taxon congruence patterns of species richness and community similarity (Bray-Curtis similarity) among birds, butterflies, and vascular plants in montane meadow habitats in the Greater Yellowstone Ecosystem. Although patterns of species richness (Spearman rank correlation) varied between taxa, we consistently found a positive correlation in community similarity (Mantel test) between all pair-wise comparisons of the three taxa (e.g., sites with similar bird communities also had similar butterfly communities). We suggest that the success of a surrogate taxon depends on the technique used to assess surrogacy and the specific approach to conservation planning. In the context of coarse-filter conservation, measures of community similarity may be more appropriate than measures of species richness. Furthermore, the cross-taxon congruency of community similarity in our study suggests that coarse-filter conservation may be tenable in montane meadow communities.     

Effects of Land-Use and Tsetse Fly Control on Bird Species Richness in Southwestern Ethiopia

Successful control of tsetse (Glossina spp.)-transmitted trypanosomiasis in the Ghibe Valley, Ethiopia, appears to have accelerated conversion of wooded grassland into cropland. Land conversion, in turn, may have fragmented wildlife habitat. Our objective was to assess the influence of the expansion of agricultural land-use, brought about by tsetse control, on ecological properties by using bird species richness and composition as indicators of environmental impacts. We sampled bird species richness and composition (using Timed-Species counts) and habitat structure (using field sampling and remote sensing) in four land cover/land-use types in areas subjected to tsetse fly control and adjacent areas without control. At the height of the growing season bird species numbers and vegetative complexity were greater in the small-holder, oxen-plowed fields and riparian woodlands than in wooded grasslands or in large-holder, tractor-plowed fields. Species composition was highly dissimilar (40–70% dissimilarity) comparing among land-use types, with many species found only in a single type. This implies that trypanosomiasis control that results in land conversion from wooded grasslands to small-holder farming in this region may have no adverse impacts on bird species numbers but will alter composition. These results also suggest that moderate land-use by humans (e.g., small-holder field mosaics) increases habitat heterogeneity and bird species richness relative to high levels of use (e.g., tractor-plowed fields). Tsetse control may be indirectly maintaining species richness in the valley by encouraging the differential spread of these small-scale, heterogeneous farms in place of large-scale, homogeneous farms. Nevertheless, if the extent of small-holder farms significantly exceeds that of present levels, negative impacts on bird species richness and large shifts in species composition may occur.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

Planting History and Propagule Pressure as Predictors of Invasion by Woody Species in a Temperate Region

Abstract:  We studied 28 alien tree species currently planted for forestry purposes in the Czech Republic to determine the probability of their escape from cultivation and naturalization. Indicators of propagule pressure (number of administrative units in which a species is planted and total planting area) and time of introduction into cultivation were used as explanatory variables in multiple regression models. Fourteen species escaped from cultivation, and 39% of the variance was explained by the number of planting units and the time of introduction, the latter being more important. Species introduced early had a higher probability of escape than those introduced later, with more than 95% probability of escape for those introduced before 1801 and <5% for those introduced after 1892. Probability of naturalization was more difficult to predict, and eight species were misclassified. A model omitting two species with the largest influence on the model yielded similar predictors of naturalization as did the probability of escape. Both phases of invasion therefore appear to be driven by planting and introduction history in a similar way. Our results demonstrate the importance of forestry for recruitment of invasive trees. Six alien forestry trees, classified as invasive in the Czech Republic, are currently reported in nature reserves. In addition, forestry authorities want to increase the diversity of alien species and planting area in the country.     

Reliability of a Higher-Taxon Approach to Richness, Rarity, and Composition Assessments at the Local Scale

Abstract:  A promising shortcut for quantifying species patterns is to use genera and families as surrogates of species. At large spatial scales, concurrence between patterns of richness, rarity, and composition of species and higher taxa is generally high. Only a few researchers, however, have examined this relationship at the local scale, which is frequently the relevant scale in land-use conflicts. We investigated the reliability of the higher-taxon approach in assessing patterns of species richness, rarity, and composition at the local scale. We studied diversity patterns of three commonly used surrogate taxa: vascular plants, ground-dwelling beetles, and moths. We conducted year-round field surveys for these taxa in the Jerusalem Mountains and the Judean foothills, Israel. Richness and composition of species were highly correlated with richness and composition of genera for all taxa. At the family level, correlations with richness and composition of species were much lower. Excluding monotypic genera and families did not affect these relations. Rarity representation based on higher taxa varied considerably depending on the taxon, and rarity scale and was weaker compared with richness and composition representation. Cumulative richness curves of species and genera showed similar patterns, leveling off at equivalent sampling efforts. Genus-level assessments were a reliable surrogate for local patterns of species richness, rarity, and composition, but family-level assessments performed poorly. The advantage of using coarse taxonomic scales in local diversity surveys is that it may decrease identification time and the need for experts, but it will not reduce sampling effort.