Impact of marine protected areas on temporal stability of fish species diversity

Preserving biodiversity over time is a pressing challenge for conservation science. A key goal of marine protected areas (MPAs) is to maintain stability in species composition, via reduced turnover, to support ecosystem function. Yet, this stability is rarely measured directly under different levels of protection. Rather, evaluations of MPA efficacy generally consist of static measures of abundance, species richness, and biomass, and rare measures of turnover are limited to short-term studies involving pairwise (beta diversity) comparisons. Zeta diversity is a recently developed metric of turnover that allows for measurement of compositional similarity across multiple assemblages and thus provides more comprehensive estimates of turnover. We evaluated the effectiveness of MPAs at preserving fish zeta diversity across a network of marine reserves over 10 years in Batemans Marine Park, Australia. Snorkel transect surveys were conducted across multiple replicated and spatially interspersed sites to record fish species occurrence through time. Protection provided by MPAs conferred greater stability in fish species turnover. Marine protected areas had significantly shallower decline in zeta diversity compared with partially protected and unprotected areas. The retention of harvested species was four to six times greater in MPAs compared with partially protected and unprotected areas, and the stabilizing effects of protection were observable within 4 years of park implementation. Conversely, partial protection offered little to no improvement in stability, compared with unprotected areas. These findings support the efficacy of MPAs for preserving temporal fish diversity stability. The implementation of MPAs helps stabilize fish diversity and may, therefore, support biodiversity resilience under ongoing environmental change.     

Running to stand still: temperature effects on species richness, species turnover, and functional community dynamics

The information on temperature-mediated changes in biodiversity in local assemblages is scarce and mainly addresses the change in species richness. However, warming may have more consistent effects on species turnover than on the number of species. Moreover, very few studies extended the analysis of changes in biodiversity and species composition to questions of associated ecosystem functions such as primary production. Here, we synthesize 4 case studies employing microalgal microcosms within the Aquashift priority program to ask (1) do warming-related shifts in species richness correspond to changes in the rate of biomass production, (2) do similar relationships prevail for evenness, and (3) do warming-related shifts in species turnover stabilize or destabilize biomass production? Two of the four cases are previously unpublished, and for a third case, the link between diversity and functional consequences of temperature was not analyzed before. We found accelerated loss of species with warming in all cases. Biomass production was lower with lower species richness in most cases but increased with lower evenness. Most importantly, the relation between functional and compositional stability was different between cases: More rapid extinction resulted in more variable biomass in 2 cases conducted with a limited species pool, indicating that compositional destabilization relates to functional variability. By contrast, the only experiment with a large species pool (30 species) allowed previously rare species to become dominant in the community and showed more stable biomass at high turnover, indicating that compensatory dynamics (turnover) can promote functional stability. These 4 independent experiments highlight the need to consider both compositional and functional consequences of altered temperature regimes.     

Influence of environment, disturbance, and ownership on forest vegetation of coastal Oregon.

Information about how vegetation composition and structure vary quantitatively and spatially with physical environment, disturbance history, and land ownership is fundamental to regional conservation planning. However, current knowledge about patterns of vegetation variability across large regions that is spatially explicit (i.e., mapped) tends to be general and qualitative. We used spatial predictions from gradient models to examine the influence of environment, disturbance, and ownership on patterns of forest vegetation biodiversity across a large forested region, the 3-million-ha Oregon Coast Range (USA). Gradients in tree species composition were strongly associated with environment, especially climate, and insensitive to disturbance, probably because many dominant tree species are long-lived and persist throughout forest succession. In contrast, forest structure was strongly correlated with disturbance and only weakly with environmental gradients. Although forest structure differed among ownerships, differences were blurred by the presence of legacy trees that originated prior to current forest management regimes. Our multi-ownership perspective revealed biodiversity concerns and benefits not readily visible in single-ownership analyses, and all ownerships contributed to regional biodiversity values. Federal lands provided most of the late-successional and old-growth forest. State lands contained a range of forest ages and structures, including diverse young forest, abundant legacy dead wood, and much of the high-elevation true fir forest. Nonindustrial private lands provided diverse young forest and the greatest abundance of hardwood trees, including almost all of the foothill oak woodlands. Forest industry lands encompassed much early-successional forest, most of the mixed hardwood-conifer forest, and large amounts of legacy down wood. The detailed tree- and species-level data in the maps revealed regional trends that would be masked in traditional coarse-filter assessment. Although abundant, most early-successional forests originated after timber harvest and lacked legacy live and dead trees important as habitat and for other ecological functions. Many large-conifer forests that might be classified as old growth using a generalized forest cover map lacked structural features of old growth such as multilayered canopies or dead wood. Our findings suggest that regional conservation planning include all ownerships and land allocations, as well as fine-scale elements of vegetation composition and structure.     

Measuring the impact of the pet trade on Indonesian birds

The trade in wild animals involves one‐third of the world's bird species and thousands of other vertebrate species. Although a few species are imperiled as a result of the wildlife trade, the lack of field studies makes it difficult to gauge how serious a threat it is to biodiversity. We used data on changes in bird abundances across space and time and information from trapper interviews to evaluate the effects of trapping wild birds for the pet trade in Sumatra, Indonesia. To analyze changes in bird abundance over time, we used data gathered over 14 years of repeated bird surveys in a 900‐ha forest in southern Sumatra. In northern Sumatra, we surveyed birds along a gradient of trapping accessibility, from the edge of roads to 5 km into the forest interior. We interviewed 49 bird trappers in northern Sumatra to learn which species they targeted and how far they went into the forest to trap. We used prices from Sumatran bird markets as a proxy for demand and, therefore, trapping pressure. Market price was a significant predictor of species declines over time in southern Sumatra (e.g., given a market price increase of approximately $50, the log change in abundance per year decreased by 0.06 on average). This result indicates a link between the market‐based pet trade and community‐wide species declines. In northern Sumatra, price and change in abundance were not related to remoteness (distance from the nearest road). However, based on our field surveys, high‐value species were rare or absent across this region. The median maximum distance trappers went into the forest each day was 5.0 km. This suggests that trapping has depleted bird populations across our remoteness gradient. We found that less than half of Sumatra's remaining forests are >5 km from a major road. Our results suggest that trapping for the pet trade threatens birds in Sumatra. Given the popularity of pet birds across Southeast Asia, additional studies are urgently needed to determine the extent and magnitude of the threat posed by the pet trade.     

Structural gradients in an intertidal hard-bottom community: examining vertical,horizontal, and taxonomic clines in zoobenthic biodiversity

Measures of biodiversity along environmental gradients have long been the focus of marine ecological research. However, a general lack of comparability between studies and under-appreciation of co-occurring, less obvious clines has often undermined any general conclusions. Latitudinal, vertical, horizontal, and taxonomic gradients in intertidal biodiversity were assessed directly and indirectly using a large data set from one locality in southwest Ireland. A total of 153 epifaunal species, from 106 families and 13 phyla, was recorded from over 200,000 individuals in a sample area of 63 m². Multivariate analysis revealed that the vertical gradient of immersion was the primary structuring factor in the community but that a flow-driven horizontal gradient was also important in structuring low- and mid-shore zones. Crustaceans, annelids, and molluscs were most numerous over the entire site, but bryozoans and sponges may have played the most significant role in driving the horizontal (flow-related) trend in species richness at lower levels on the shore. Comparison of species richness with sites from other locations around the world proved inconclusive in assessing a global trend in richness from the poles to the tropics. However, interpolation of this and similar studies may prove useful in future latitudinal investigations of hard-substratum intertidal biota. Higher-taxon surrogacy for species trends proved significant and useful at the scale investigated here, whereby analysis of data from family or even order level may prove an efficient method of future monitoring. More data are required to assess if the top-down taxonomic approach will sufficiently mirror species trends at larger spatial scales. Overall, the generality of effects of environmental gradients on intertidal organisms may be more conclusive where entire communities are studied and consideration is given to other, less dominant clines.Communicated by J.P. Thorpe, Port Erin     

Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

Beyond Kyoto: Forest Management in a Time of Rapid Climate Change

Abstract: Policies to reduce global warming by offering credits for carbon sequestration have neglected the effects of forest management on biodiversity. I review properties of forest ecosystems and management options for enhancing the resistance and resilience of forests to climate change. Although forests, as a class, have proved resilient to past changes in climate, today's fragmented and degraded forests are more vulnerable. Adaptation of species to climate change can occur through phenotypic plasticity, evolution, or migration to suitable sites, with the latter probably the most common response in the past. Among the land-use and management practices likely to maintain forest biodiversity and ecological functions during climate change are (1) representing forest types across environmental gradients in reserves; (2) protecting climatic refugia at multiple scales; (3) protecting primary forests; (4) avoiding fragmentation and providing connectivity, especially parallel to climatic gradients; (5) providing buffer zones for adjustment of reserve boundaries; (6) practicing low-intensity forestry and preventing conversion of natural forests to plantations; ( 7) maintaining natural fire regimes; (8) maintaining diverse gene pools; and (9) identifying and protecting functional groups and keystone species. Good forest management in a time of rapidly changing climate differs little from good forest management under more static conditions, but there is increased emphasis on protecting climatic refugia and providing connectivity.     

Effects of productivity on biodiversity in forest ecosystems across the United States and China

In the global campaign against biodiversity loss in forest ecosystems, land managers need to know the status of forest biodiversity, but practical guidelines for conserving biodiversity in forest management are lacking. A major obstacle is the incomplete understanding of the relationship between site primary productivity and plant diversity, due to insufficient ecosystem‐wide data, especially for taxonomically and structurally diverse forest ecosystems. We investigated the effects of site productivity (the site's inherent capacity to grow timber) on tree species richness across 19 types of forest ecosystems in North America and China through 3 ground‐sourced forest inventory data sets (U.S. Forest Inventory and Analysis, Cooperative Alaska Forest Inventory, and Chinese Forest Management Planning Inventory). All forest types conformed to a consistent and highly significant (P < 0.001) hump‐shaped unimodal relationship, of which the generalized coefficients of determination averaged 20.5% over all the forest types. That is, tree species richness first increased as productivity increased at a progressively slower rate, and, after reaching a maximum, richness started to decline. Our consistent findings suggest that forests of high productivity would sustain few species because they consist mostly of flat homogeneous areas lacking an environmental gradient along which a diversity of species with different habitats can coexist. The consistency of the productivity–biodiversity relationship among the 3 data sets we examined makes it possible to quantify the expected tree species richness that a forest stand is capable of sustaining, and a comparison between the actual species richness and the sustainable values can be useful in prioritizing conservation efforts.     

Woody plant richness and NDVI response to drought events in Catalonian (northeastern Spain) forests

The role of species diversity on ecosystem resistance in the face of strong environmental fluctuations has been addressed from both theoretical and experimental viewpoints to reveal a variety of positive and negative relationships. Here we explore empirically the relationship between the richness of forest woody species and canopy resistance to extreme drought episodes. We compare richness data from an extensive forest inventory to a temporal series of satellite imagery that estimated drought impact on forest canopy as NDVI (normalized difference vegetation index) anomalies of the dry summer in 2003 in relation to records of previous years. We considered five different types of forests that are representative of the main climatic and altitudinal gradients of the region, ranging from lowland Mediterranean to mountain boreal-temperate climates. The observed relationship differed among forest types and interacted with the climate, summarised by the Thorntwaite index. In Mediterranean Pinus halepensis forests, NDVI decreased during the drought. This decrease was stronger in forests with lower richness. In Mediterranean evergreen forests of Quercus ilex, drought did not result in an overall NDVI loss, but lower NDVI values were observed in drier localities with lower richness, and in more moist localities with higher number of species. In mountain Pinus sylvestris forests NDVI decreased, mostly due to the drought impact on drier localities, while no relation to species richness was observed. In moist Fagus sylvatica forests, NDVI only decreased in plots with high richness. No effect of drought was observed in the high mountain Pinus uncinata forests. Our results show that a shift on the diversity-stability relationship appears across the regional, climatic gradient. A positive relationship appears in drier localities, supporting a null model where the probability of finding a species able to cope with drier conditions increases with the number of species. However, in more moist localities we hypothesize that the proportion of drought-sensitive species would increase in richer localities, due to a higher likelihood of co-occurrence of species that share moist climatic requirements. The study points to the convenience of considering the causes of disturbance in relation to current environmental gradients and historical environmental constraints on the community.     

Bird metacommunities in temperate South American forest: vegetation structure, area, and climate effects

Spatial structure in metacommunities and their relationships to environmental gradients have been linked to opposing theories of community assembly. In particular, while the species sorting hypothesis predicts strong environmental influences, the neutral theory, the mass effect, and the patch dynamics frameworks all predict differing degrees of spatial structure resulting from dispersal and competition limitations. Here we study the relative influence of environmental gradients and spatial structure in bird assemblages of the Chilean temperate forest. We carried out bird and vegetation surveys in South American temperate forests at 147 points located in nine different protected areas in central Chile, and collected meteorological and productivity data for these localities. Species composition dissimilarities between sites were calculated, as well as three indices of bird local diversity: observed species richness, Chao estimate of richness, and Shannon diversity. A stepwise multiple regression and partial regression analyses were used to select a small number of environmental factors that predicted bird species diversity. Although diversity indices were spatially autocorrelated, environmental factors were sufficient to account for this autocorrelation. Moreover, community dissimilarities were not significantly related to distance between sites. We then tested a multivariate hypothesis about climate, vegetation, and avian diversity interactions using a structural equation modeling (SEM) approach. The SEM showed that climate and area of fragments have important indirect effects on avian diversity, mediated through changes in vegetation structure. Given the scale of this study, the metacommunity framework provides useful insights into the mechanisms driving bird assemblages in this region. Taken together, the weak spatial structure of community composition and diversity, as well as the strong environmental effects on bird diversity, support the interpretation that species sorting has a predominant role in structuring avian assemblages in the region.     

The Silent Mass Extinction of Insect Herbivores in Biodiversity Hotspots

Abstract: Habitat loss is silently leading numerous insects to extinction. Conservation efforts, however, have not been designed specifically to protect these organisms, despite their ecological and evolutionary significance. On the basis of species–host area equations, parameterized with data from the literature and interviews with botanical experts, I estimated the number of specialized plant‐feeding insects (i.e., monophages) that live in 34 biodiversity hotspots and the number committed to extinction because of habitat loss. I estimated that 795,971–1,602,423 monophagous insect species live in biodiversity hotspots on 150,371 endemic plant species, which is 5.3–10.6 monophages per plant species. I calculated that 213,830–547,500 monophagous species are committed to extinction in biodiversity hotspots because of reduction of the geographic range size of their endemic hosts. I provided rankings of biodiversity hotspots on the basis of estimated richness of monophagous insects and on estimated number of extinctions of monophagous species. Extinction rates were predicted to be higher in biodiversity hotspots located along strong environmental gradients and on archipelagos, where high spatial turnover of monophagous species along the geographic distribution of their endemic plants is likely. The results strongly support the overall strategy of selecting priority conservation areas worldwide primarily on the basis of richness of endemic plants. To face the global decline of insect herbivores, one must expand the coverage of the network of protected areas and improve the richness of native plants on private lands.     

Effect of scale on trait predictors of species responses to agriculture

Species persistence in human‐altered landscapes can depend on factors operating at multiple spatial scales. To understand anthropogenic impacts on biodiversity, it is useful to examine relationships between species traits and their responses to land‐use change. A key knowledge gap concerns whether these relationships vary depending on the scale of response under consideration. We examined how local‐ and large‐scale habitat variables influence the occupancy dynamics of a bird community in cloud forest zones in the Colombian Chocó‐Andes. Using data collected across a continuum of forest and agriculture, we examined which traits best predict species responses to local variation in farmland and which traits best predict species responses to isolation from contiguous forest. Global range size was a strong predictor of species responses to agriculture at both scales; widespread species were less likely to decline as local habitat cover decreased and as distance from forest increased. Habitat specialization was a strong predictor of species responses only at the local scale. Open‐habitat species were particularly likely to increase as pasture increased, but they were relatively insensitive to variation in distance to forest. Foraging plasticity and flocking behavior were strong predictors of species responses to distance from forest, but not their responses to local habitat. Species with lower plasticity in foraging behaviors and obligate flock‐following species were more likely to decline as distance from contiguous forest increased. For species exhibiting these latter traits, persistence in tropical landscapes may depend on the protection of larger contiguous blocks of forest, rather than the integration of smaller‐scale woodland areas within farmland. Species listed as threatened or near threatened on the International Union for Conservation of Nature Red List were also more likely to decline in response to both local habitat quality and isolation from forest relative to least‐concern species, underlining the importance of contiguous forests for threatened taxa.     

Estimating the footprint of pollution on coral reefs with models of species turnover

Ecological communities typically change along gradients of human impact, although it is difficult to estimate the footprint of impacts for diffuse threats such as pollution. We developed a joint model (i.e., one that includes multiple species and their interactions with each other and environmental covariates) of benthic habitats on lagoonal coral reefs and used it to infer change in benthic composition along a gradient of distance from logging operations. The model estimated both changes in abundances of benthic groups and their compositional turnover, a type of beta diversity. We used the model to predict the footprint of turbidity impacts from past and recent logging. Benthic communities far from logging were dominated by branching corals, whereas communities close to logging had higher cover of dead coral, massive corals, and soft sediment. Recent impacts were predicted to be small relative to the extensive impacts of past logging because recent logging has occurred far from lagoonal reefs. Our model can be used more generally to estimate the footprint of human impacts on ecosystems and evaluate the benefits of conservation actions for ecosystems.     

Spatial Components of Bird Diversity in the Andes of Colombia: Implications for Designing a Regional Reserve System

Abstract:  Beta diversity, or the turnover in species composition among sampling sites in a region, is an important criterion for obtaining adequate representation of regional biodiversity in systems of protected areas. Recently, the additive model for partitioning regional (gamma) diversity (in opposition to the multiplicative model) has been proposed because it allows a direct measure of the contribution of beta diversity to gamma diversity. We determined avian beta diversity along latitudinal (among neighboring river drainages) and elevational axes in a 1347-km² region on the western slope of the Central Cordillera of the Colombian Andes, where a regional system of protected areas is being designed. We then compared avian beta diversity between sites based on rapid versus long-term (>1 year) inventories and between fragmented sites versus continuous forest. Overall, beta diversity represented 63.1% of gamma diversity among 16 sites. Elevational differences in species composition accounted for 43.3% of regional diversity, whereas differences among drainages accounted for 19.8%. A complementary cluster analysis showed that sites grouped by elevational zones. Rapid inventories overestimated beta diversity because of sampling effects, but the effect was biologically small. Estimators of species richness derived from species accumulation curves provided a useful alternative to compensate for undersampling in short-term surveys. Forest fragmentation increased beta diversity because of differential local extinction of populations. Nevertheless, in our region, forest fragments contributed to gamma diversity because they contained complementary sets of species. More importantly, they contained populations of special-interest species. Although the region is relatively small, our analyses indicate that spatial differentiation of the biota is an important factor for deciding number and location of protected areas in the Andean region.     

The role of time and species identities in spatial patterns of species richness and conservation

Many conservation actions are justified on the basis of managing biodiversity. Biodiversity, in terms of species richness, is largely the product of rare species. This is problematic because the intensity of sampling needed to characterize communities and patterns of rarity or to justify the use of surrogates has biased sampling in favor of space over time. However, environmental fluctuations interacting with community dynamics lead to temporal variations in where and when species occur, potentially affecting conservation planning by generating uncertainty about results of species distribution modeling (including range determinations), selection of surrogates for biodiversity, and the proportion of biodiversity composed of rare species. To have confidence in the evidence base for conservation actions, one must consider whether temporal replication is necessary to produce broad inferences. Using approximately 20 years of macrofaunal data from tidal flats in 2 harbors, we explored variation in the identity of rare, common, restricted range, and widespread species over time and space. Over time, rare taxa were more likely to increase in abundance or occurrence than to remain rare or disappear and to exhibit temporal patterns in their occurrence. Space–time congruency in ranges (i.e., spatially widespread taxa were also temporally widespread) was observed only where samples were collected across an environmental gradient. Fifteen percent of the taxa in both harbors changed over time from having spatially restricted ranges to having widespread ranges. Our findings suggest that rare species can provide stability against environmental change, because the majority of species were not random transients, but that selection of biodiversity surrogates requires temporal validation. Rarity needs to be considered both spatially and temporally, as species that occur randomly over time are likely to play a different role in ecosystem functioning than those exhibiting temporal structure (e.g., seasonality). Moreover, temporal structure offers the opportunity to place management and conservation activities within windows of maximum opportunity.     

Large-scale biodiversity patterns in freshwater phytoplankton

Our planet shows striking gradients in the species richness of plants and animals, from high biodiversity in the tropics to low biodiversity in polar and high-mountain regions. Recently, similar patterns have been described for some groups of microorganisms, but the large-scale biogeographical distribution of freshwater phytoplankton diversity is still largely unknown. We examined the species diversity of freshwater phytoplankton sampled from 540 lakes and reservoirs distributed across the continental United States and found strong latitudinal, longitudinal, and altitudinal gradients in phytoplankton biodiversity, demonstrating that microorganisms can show substantial geographic variation in biodiversity. Detailed analysis using structural equation models indicated that these large-scale biodiversity gradients in freshwater phytoplankton diversity were mainly driven by local environmental factors, although there were residual direct effects of latitude, longitude, and altitude as well. Specifically, we found that phytoplankton species richness was an increasing saturating function of lake chlorophyll a concentration, increased with lake surface area and possibly increased with water temperature, resembling effects of productivity, habitat area, and temperature on diversity patterns commonly observed for macroorganisms. In turn, these local environmental factors varied along latitudinal, longitudinal, and altitudinal gradients. These results imply that changes in land use or climate that affect these local environmental factors are likely to have major impacts on large-scale biodiversity patterns of freshwater phytoplankton.     

Carbohydrate storage and light requirements of tropical moist and dry forest tree species

In many plant communities, there is a negative interspecific correlation between relative growth rates and survival of juveniles. This negative correlation is most likely caused by a trade-off between carbon allocation to growth vs. allocation to defense and storage. Nonstructural carbohydrates (NSC) stored in stems allow plants to overcome periods of stress and should enhance survival. In order to assess how species differ in carbohydrate storage in relation to juvenile light requirements, growth, and survival, we quantified NSC concentrations and pool sizes in sapling stems of 85 woody species in moist semi-evergreen and dry deciduous tropical forests in the rainy season in Bolivia. Moist forest species averaged higher NSC concentrations than dry forest species. Carbohydrate concentrations and pool sizes decreased with the light requirements of juveniles of the species in the moist forest but not in the dry forest. Combined, these results suggest that storage is especially important for species that regenerate in persistently shady habitats, as in the understory of moist evergreen forests. For moist forest species, sapling survival rates increased with NSC concentrations and pool sizes while growth rates declined with the NSC concentrations and pool sizes. No relationships were found for dry forest species. Carbon allocation to storage contributes to the growth-survival trade-off through its positive effect on survival. And, a continuum in carbon storage strategies contributes to a continuum in light requirements among species. The link between storage and light requirements is especially strong in moist evergreen forest where species sort out along a light gradient, but disappears in dry deciduous forest where light is a less limiting resource and species sort out along drought and fire gradients.