Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

旅游干扰下五台山不同植被景观区物种多样性特征

利用双向指示种分析方法(Two-way indicator species analysis,TWINSPAN)和6个物种多样性指数,研究了旅游干扰下五台山不同植被景观区物种多样性的特征.结果表明:1)TWINSPAN将所有样地划分为9类植被景观区,从Ⅰ区到Ⅸ区,随着旅游干扰程度的增加,植被景观大致由乔灌草区向灌草区、草本区和居民区方向变化.2)乔木层物种的丰富度和综合多样性随着旅游干扰的减小而趋于增加,但是其均匀度没有表现出明显的规律性.3)灌木层物种的丰富度和综合多样性也随着旅游干扰的减小而趋于增加,至于其均匀度,则呈现出在中度干扰下值最大,干扰很小的地方次之,在重度干扰下则最小.4)草本层物种的丰富度、均匀度和综合多样性指数均在旅游干扰适度的地方达到了最大值,在旅游干扰很小的地方,各种值则次之,在干扰严重的地方为最小.5)从整个植被层物种多样性的角度看,随着旅游干扰程度的减少,物种丰富度指数、均匀度指数和综合多样性指数均呈现趋于增加的趋势.表4参15     

Relationships between species richness, evenness, and abundance in a southwestern savanna

Species richness and evenness are components of biological diversity that may or may not be correlated with one another and with patterns of species abundance. We compared these attributes among flowering plants, grasshoppers, butterflies, lizards, summer birds, winter birds, and rodents across 48 plots in the grasslands and mesquite-oak savannas of southeastern Arizona. Species richness and evenness were uncorrelated or weakly negatively correlated for each taxonomic group, supporting the conclusion that richness alone is an incomplete measure of diversity. In each case, richness was positively correlated with one or more measures of abundance. By contrast, evenness usually was negatively correlated with the abundance variables, reflecting the fact that plots with high evenness generally were those where all species present were about equally uncommon. Therefore richness, but not evenness, usually was a positive predictor of places of conservation value, if these are defined as places where species of interest are especially abundant. Species diversity was more positively correlated with evenness than with richness among grasshoppers and flowering plants, in contrast to the other taxonomic groups, and the positive correlations between richness and abundance were comparatively weak for grasshoppers and plants as well. Both of these differences can be attributed to the fact that assemblages of plants and grasshoppers were numerically dominated by small subsets of common species (grasses and certain spur-throated grasshoppers) whose abundances differed greatly among plots in ways unrelated to species richness of the groups as a whole.     

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.     

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.     

Woody encroachment decreases diversity across North American grasslands and savannas

Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.     

围封6年对温带典型草原坡地物种多样性及其与地上生物量的关系的影响

以内蒙古太仆寺旗地区典型克氏针茅-羊草草原为研究对象,通过调查围封6年后围栏内外典型草原群落特征,分析围封、放牧处理下典型草原植物群落结构及地上生物量的动态变化,揭示围封放牧对植被群落结构、物种丰富度和地上生物量的影响。结果发现,在围封6年后,各功能群物种丰富度与盖度围栏内外的空间变化趋势基本一致。群落物种丰富度和地上生物量均随坡位下降而显著增加,初步显示了物种多样性与生态系统功能的空间变化的一致性。围栏内物种丰富度显著低于围栏外,而地上生物量却显著大于围栏外。围栏内多年生禾草、一年生植物物种丰富度显著低于围栏外,而半灌木盖度、生物量显著高于围栏外。围栏内外在禁牧和放牧不同处理下,表现出不同的草原退化方式：在禁牧条件下,尽管生物量有所恢复,但灌木和半灌木的增加却代表了草原的退化;放牧、刈割等人类活动增加了耐牧物种和不适口植物生长的可能,表现为草原退化指示物种增加,同样是草原退化的表现。围栏外一年生植物盖度和生物量显著高于围栏内,而一年生植物的频数也远远大于围栏内（围栏内外频数比13：71）,且独行菜、猪毛菜等仅出现于围栏外。同时,地上生物量随着物种丰富度的增加而增加,但只有围栏外的关系达到统计显著程度,说明围封调制了生物多样性与生态系统功能的关系。本研究表明,对于干旱半干旱典型草原的恢复,不能简单地采用长期完全禁牧措施,应当根据区域环境、植被以及社会经济情况,制定季节性放牧或者间歇性禁牧的恢复措施,从而保证草场的可持续性恢复利用。     

Forest age influences oak insect herbivore community structure, richness, and density.

Plant succession is one of many factors that may affect the composition and structure of herbivorous insect communities. However, few studies have examined the effect of forest age on the diversity and abundance of insect communities. If forest age influences insect diversity, then the schedule of timber harvest rotation may have consequent effects on biodiversity. The insect herbivore community on Quercus alba (white oak) in the Missouri Ozarks was sampled in a chronoseries, from recently harvested (2 yr) to old-growth (approximately 313 yr) forests. A total of nine sites and 39 stands within those sites were sampled in May and August 2003. Unique communities of plants and insects were found in the oldest forests (122-313 yr). Density and species richness of herbivores were positively correlated with increasing forest age in August but not in May. August insect density was negatively correlated with heat load index; in addition, insect density and richness increased over the chronoseries, but not on the sunniest slopes. Forest structural diversity (number of size classes) was positively correlated with forest age, but woody plant species richness was not. In sum, richness, density, and community structure of white oak insect herbivores are influenced by variation in forest age, forest structure, relative abundance of plant species, and abiotic conditions. These results suggest that time between harvests of large, long-lived, tree species such as white oak should be longer than current practice in order to maintain insect community diversity.     

Endangered Cacti in the Chihuahuan Desert: I. Distribution Patterns

We mapped the geographical distributions of 93 endangered species of cacti from the Chihuahuan Desert Region. We divided the region into grids of 30 minutes latitude by 30 minutes longitude and calculated species frequencies for each grid. The grids with the highest species richness values are aggregated in areas of moderate elevation, particularly towards the southeastern and, to a lesser extent, the eastern margins of the Chihuahuan Desert Region, in northern San Luis Potosí and the southern portions of Coahuila, Nuevo León, and Tamaulipas. This vast area constitutes the most important nucleus of cactus species concentration in the continent. Species richness decreases toward the western segment of the Chihuahuan Desert Region and from the Cuatro Ciénegas region to the north and northwest. Another important area is the Queretaroan-Hidalgoan Arid Zone, where several grids containing an important assemblage of endangered species occur. Climatic factors, such as minimum temperatures and mean annual precipitation, explain the current distribution patterns of these plants, and the recent Pleistocene climatic episodes appear to have played a determinant role in the existence of areas of high species concentration and in the proliferation of narrow endemics. Special actions are urgently needed to conserve the endangered Cactaceae of this region. We propose that a carefully selected network of small areas would be an appropriate approach for the conservation of these plants. But species richness cannot be taken as the sole criterion in the determination of protected areas. Additional criteria, such as degree of endemicity, degree of threat to species and areas, habitat diversity, and biogeographic congruence with other plant and animal groups, should be analyzed before these areas are selected. Propagation in botanical gardens using scientific criteria and commercial propagation would be additional methods of conservation.     

半干旱区人工封育草场植物群落物种多样性与复杂性研究──以宁夏盐池为例

围栏封育促进植被恢复效果研究方法很多,其中通过研究植被的群落特征及物种多样性来阐述恢复状况仍是比较常用和有效的手段,但多样性测度对生物群落的自组织及有序性方面的特征描述欠缺,鉴于此,多样性与复杂性测度结合起来研究群落结构与功能显得更有意义。研究人工封育对植被恢复过程中群落的多样性与复杂性的影响,必然有助于揭示生态恢复过程机理。目前,利用多样性与复杂性测度相结合来对群落结构进行量化研究不多,用于对草原生态系统植物群落结构研究更少。本文通过样地调查,利用多样性和复杂性指数相结合定量研究半干旱区人工封育草场植物群落物种多样性、复杂性随封育年限的变化规律,并对比分析复杂性与物种多样性之间的相关性。结果表明,（1）老封育区、新封育区丰富度指数整体高于对照区,说明封育有利于物种丰富度的增加。但随封育时间的延长,封育区的优势种占据主要的资源空间,丰富度指数降低。长时间封育,草场植被丰富度将发生规律性波动变化,低峰值和高峰值出现频率均约为4~5年/次。老封育区、新封育区的SW、SP多样性指数值大部分也高于对照区,说明封育措施增加草场植被的多样性。物种多样性指数受降水影响显著,当降水量增加或减少时,多样性指数也相应出现增加或减少,但降水影响效应具有一定的滞后性。老封育区、新封育区均匀度变化比较平稳,且指数值高于对照区,说明封育有利于均匀度的增加。（2）老封育区、新封育区总复杂性指数、无序结构复杂性指数值大部分比对照区指数值高,且指数波动相对平稳,说明封育区群落结构比对照区稳定,更适合荒漠草原植被的生长,封育增加了群落总复杂性、无序结构复杂性。封育也提高群落有序结构复杂性,但?     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

The impact of motorcycle traffic on soil and vegetation of stabilized coastal dunes, Israel

This study aimed to assess the response of soil and annual plant vegetation of stabilized coastal dunes in Israel, to varying intensities of off-road motorcycle (ORM) traffic, and to assess their resistance and resilience to such a disturbance. A standard experimental procedure that included 0, 20, 50, 100 and 200 ORM straight passes and 150 ORM turn passes was used. Plant ground cover, mean plant height, species richness, species diversity, soil penetrable depth, organic matter and moisture contents were measured on several dates within a period of 372 days after the experiment. Results have shown that: (1) ORM passes had a significant immediate impact on annual plants that increased with traffic intensity. The impact on the soil was detected only as an increase of penetrable depth. (2) The maximum impact on annual plants was observed in the wheel ruts and turn areas. The impact on the area between the wheel ruts and on the margins outside the wheel ruts was indirect and smaller. (3) Annual plant ground cover and mean plant height were less sensitive parameters than species richness and species diversity for determining the impact of ORM traffic on the area. (4) One year after the experiment, soil and annual plant vegetation in all passes were very similar to their pre-experimental condition. This indicates high resilience and recovery potential of the Mediterranean stabilized coastal dune ecosystem to ORM disturbance.     

Biodiversity Differences between Managed and Unmanaged Forests: Meta-Analysis of Species Richness in Europe

Abstract: Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest‐dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan‐European scale. We used meta‐analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management‐intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta‐analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.     

植物多样性对土壤微生物的影响

生物多样性强烈地影响生态系统的过程.生态系统过程的变化可导致生物多样性衰减并因此导致生态系统功能衰退.植物种丰度和植物功能多样性对土壤细菌群落的代谢活性和代谢多样性有成正比的影响.土壤细菌的代谢活性和代谢多样性随植物种数量的对数和植物功能组的数量而直线上升.其原因可能是由植被流入土壤的物质和能量的多样性和数量的增加,也可能是由土壤动物区系起作用的土壤微生境的多样性的增加造成的.由于植物多样性的丧失所引起的植物生物量的减少对分解者群落有强烈的影响微生物生物量将可能减少,因为在大多数陆地生态系统中,有机碳源限制着土壤微生物的活性.     

江苏省陆栖濒危脊椎动物物种丰富度及分布格局

采用实地调查与资料收集相结合的方法,以县(市)为基本地理单元建立江苏省陆栖濒危脊椎动物二元分布数据库;基于GIS技术,研究濒危动物物种丰富度格局;依据地理单元中物种组成的相似程度进行聚类分析,探讨江苏省濒危动物相似性分布格局。结果显示,江苏省陆栖濒危脊椎动物共有35科62属83种;西南部丘陵山区、盐城、连云港及启东地区濒危物种丰富度较高,江苏中部地区(主要为泰州和淮安)濒危物种丰富度较低,这与江苏省生物多样性分布热点基本一致,濒危物种丰富度与总物种丰富度之间呈极显著正相关关系(P<0.01,n=73),Pearson相关系数为0.869;依据物种组成相似性,当组间距离分别为23.8、20.6、18.8时,江苏省73个基本单元可聚为3、4、5组,该聚类结果表明江苏省陆栖濒危脊椎动物分布受自然地理亚区、地形地貌以及人为活动的共同影响。     

Conservation Prioritization Using GAP Data

Data collected by the Gap Analysis Program in the state of Idaho (U.S.A.) are used to prioritize the selection of locations for conservation action and research. Set coverage and integer programming algorithms provide a sequence of localities that maximize the number of species or vegetation classes represented at each step. Richness maps of vegetation cover class diversity, terrestrial vertebrate species diversity ("hot spot analysis"), endangered, threatened, and candidate species diversity, and unprotected vertebrate species diversity ("gap analysis"), when prioritized, show a rapid accumulation of species as more localities are chosen for terrestrial vertebrates and unprotected vertebrates. Gap analysis identifies four target areas ("gaps") that include 79 of the 83 vertebrate species not currently protected. Accumulation of vegetation cover classes and endangered, threatened, and candidate species is much slower. Sweep analysis is used to determine how well prioritizing on one component of diversity accumulates other components. Endangered, threatened, and candidate species do not sweep total vertebrates as well as unprotected vertebrates do, but are better than vegetation classes. Total vertebrates sweep endangered, threatened, and candidate species better than unprotected vertebrates do, which in turn are better than vegetation classes. We emphasize that prioritization must be part of conservation efforts at multiple scales and that prioritization points out important localities where more detailed work mast be undertaken.     

Terrestrial vertebrates alter seedling composition and richness but not diversity in an Australian tropical rain forest

Although birds and mammals play important roles in several mechanisms hypothesized to maintain plant diversity in species-rich habitats, there have been few long-term, community-level tests of their importance. We excluded terrestrial birds and mammals from fourteen 6 x 7.5 m plots in Australian primary tropical rain forest and compared recruitment and survival of tree seedlings annually over the subsequent seven years to that on nearby open plots. We re-censused a subset of the plots after 13 years of vertebrate exclusion to test for longer-term effects. After two years of exclusion, seedling abundance was significantly higher (74%) on exclosure plots and remained so at each subsequent census. Richness was significantly higher on exclosure plots from 1998 to 2003, but in 2009 richness no longer differed, and rarefied species richness was higher in the presence of vertebrates. Shannon's diversity and Pielou's evenness did not differ in any year. Vertebrates marginally increased density-dependent mortality and recruitment limitation, but neither effect was great enough to increase richness or diversity on open plots relative to exclosure plots. Terrestrial vertebrates significantly altered seedling community composition, having particularly strong impacts on members of the Lauraceae. Overall, our results highlight that interactions between terrestrial vertebrates and tropical tree recruitment may not translate into strong community-level effects on diversity, especially over the short-term, despite significant impacts on individual species that result in altered species composition.     

Limitations of Biodiversity Databases: Case Study on Seed-Plant Diversity in Tenerife, Canary Islands

Abstract:  Databases on the distribution of species can be used to describe the geographic patterns of biodiversity. Nevertheless, they have limitations. We studied three of these limitations: (1) inadequacy of raw data to describe richness patterns due to sampling bias, (2) lack of survey effort assessment (and lack of exhaustiveness in compiling data about survey effort), and (3) lack of coverage of the geographic and environmental variations that affect the distribution of organisms. We used a biodiversity database (BIOTA-Canarias) to analyze richness data from a well-known group (seed plants) in an intensively surveyed area (Tenerife Island). Observed richness and survey effort were highly correlated. Species accumulation curves could not be used to determine survey effort because data digitalization was not exhaustive, so we identified well-sampled sites based on observed richness to sampling effort ratios. We also developed a predictive model based on the data from well-sampled sites and analyzed the origin of the geographic errors in the obtained extrapolation by means of a geographically constrained cross-validation. The spatial patterns of seed-plant species richness obtained from BIOTA-Canarias data were incomplete and biased. Therefore, some improvements are needed to use this database (and many others) in biodiversity studies. We propose a protocol that includes controls on data quality, improvements on data digitalization and survey design to improve data quality, and some alternative data analysis strategies that will provide a reliable picture of biodiversity patterns.     

A global ecological signal of extinction risk in terrestrial vertebrates

To determine the distribution and causes of extinction threat across functional groups of terrestrial vertebrates, we assembled an ecological trait data set for 18,016 species of terrestrial vertebrates and utilized phylogenetic comparative methods to test which categories of habitat association, mode of locomotion, and feeding mode best predicted extinction risk. We also examined the individual categories of the International Union for Conservation of Nature Red List extinction drivers (e.g., agriculture and logging) threatening each species and determined the greatest threats for each of the four terrestrial vertebrate groups. We then quantified the sum of extinction drivers threatening each species to provide a multistressor perspective on threat. Cave dwelling amphibians (p < 0.01), arboreal quadrupedal mammals (all of which are primates) (p < 0.01), aerial and scavenging birds (p < 0.01), and pedal (i.e., walking) squamates (p < 0.01) were all disproportionately threatened with extinction in comparison with the other assessed ecological traits. Across all threatened vertebrate species in the study, the most common risk factors were agriculture, threatening 4491 species, followed by logging, threatening 3187 species, and then invasive species and disease, threatening 2053 species. Species at higher risk of extinction were simultaneously at risk from a greater number of threat types. If left unabated, the disproportionate loss of species with certain functional traits and increasing anthropogenic pressures are likely to disrupt ecosystem functions globally. A shift in focus from species- to trait-centric conservation practices will allow for protection of at-risk functional diversity from regional to global scales.     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.