Choice of Species-Area Function Affects Identification of Hotspots

Abstract: I tested the reliability of species-area curves for use in identifying hotspots, political or geographical regions of high species richness. On a species-area plot, hotspots are points (regions) that appear above the curve to a greater extent than other points. Because several different curves can be fit to species-area data, identification of hotspots may differ depending on the curve-fitting function used. I tested this hypothesis by comparing hotspots identified by the power function, the extreme value function, a linear function, and the exponential function. I examined several species-area data sets varying in size and in the presence of endemics. I defined hotspots as the highest 25% for small data sets and highest 10% and 25% for large data sets of standardized residuals from each function fitted to each data set. For some data sets, the functions agreed in identification of hotspots in that they identified 75% or more of the same hotspots. The extreme value function tended to identify hotspots not identified by the other three functions. For most data sets, the functions did not agree completely in identifying hotspots. Therefore, species-area curves should not be used as the sole means of identifying hotspots of species richness, although they can be used to examine the effect hotspot area has on richness for hotspots identified by other methods.     

A unified model of avian species richness on islands and continents

How many species in a given taxon should be found in a delimited area in a specified place in the world? Some recent literature suggests that the answer to this question depends strongly on the geographical, evolutionary, and ecological context. For example, current theory suggests that species accumulate as a function of area differently on continents and islands. Species richness-climate relationships have been examined separately on continents and on islands. This study tests the hypotheses that (1) the functional relationship between richness and climate is the same on continents and islands; (2) the species-area slope depends on distance-based isolation; (3) species-area relationships differ among land bridge islands, oceanic islands, and continents; (4) richness differs among biogeographic regions independently of climate and isolation. We related bird species numbers in a worldwide sample of 240 continental parcels and 346 islands to several environmental variables. We found that breeding bird richness varies similarly on islands and on continents as a function of mean annual temperature, an area x precipitation interaction, and the distance separating insular samples from the nearest continent (R2 = 0.86). Most studies to date have postulated that the slope of the species-area relationship depends upon isolation. In contrast, we found no such interaction. A richness-environment relationship derived using Old World sites accurately predicts patterns of richness in the New World and vice versa (R2 = 0.85). Our results suggest that most of the global variation in richness is not strongly context-specific; rather, it reflects a small number of general environmental constraints operating on both continents and islands.     

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.     

Estimating landscape-scale species richness: reconciling frequency- and turnover-based approaches

One hypothesis for why estimators of species richness tend to underestimate total richness is that they do not explicitly account for increases in species richness due to spatial or environmental turnover in species composition (beta diversity). I analyze the similarity of a data set of native trees in Great Smoky Mountains National Park, USA, and assess the robustness of these estimators against recently developed ones that incorporate turnover explicitly: the total species accumulation method (T-S) and a method based on the distance decay of similarity. I show that the T-S estimator can give reliable estimates of species richness, given an appropriate grouping of sites. The estimator based on distance decay of similarity performed poorly. There are two main reasons for this: sample size effects and the assumption that distance decay of similarity exhibits a power law relationship. I show that estimators based on distance-decay relationships exhibit systematically lower rates of distance decay for samples with few individuals per site independent of environmental variation. Second, the data presented here and many other survey data sets exhibit exponential rather than power law distance-decay relationships. Richness estimators that explicitly incorporate beta diversity can be improved by beginning from an exponential distance-decay relationship and adjusting for the systematic errors introduced by small sample sizes.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.     

Deforestation Predicts the Number of Threatened Birds in Insular Southeast Asia

The world's tropical forests are being cleared rapidly, and ecologists claim this is causing a massive loss of species. This claim has its critics. Can we predict extinctions from the extent of deforestation? We mapped the percentage of deforestation on the islands of the Philippines and Indonesia and counted the number of bird species found only on these islands. We then used the species-area relationship to calculate the number of species predicted to become globally extinct following deforestation on these islands. Next, we counted the numbers of insular southeast Asian endemic bird species considered threatened—i.e., those having "a high probability of extinction in the wild in the medium-term future"—in the latest summary Red Data Book. The numbers of extinctions predicted from deforestation and the numbers of species actually threatened are strikingly similar. This suggests we can estimate the size of the extinction crisis in once-forested regions from the extent of deforestation. The numbers of extinctions will be large. Without rapid and effective conservation, many of the species endemic to insular southeast Asia will soon be lost.     

Global Patterns of Mammalian Diversity, Endemism, and Endangerment

To assess the conservation status of the world's land mammals, we compiled data on the number of total species, endemic species, recently extinct species, and currently endangered species for 155 countries. Total species richness was significantly correlated with territorial land area, whereas number of endemic species was only weakly correlated with both area and total number of species. The large amount of variation left unexplained by species-area regressions reflects the influence of other factors, such as latitude, topographic and habitat heterogeneity, and historical biogeography, on species richness and especially on patterns of endemism. Countries of particular conservation concern, because they have rich mammalian faunas containing many endemic species, are the large islands of Australia, Madagascar, Indonesia, and the Philippines, as well as continental Mexico. Patterns of recent extinctions and the current endangered status of species were difficult to interpret, largely because of inadequate and inconsistent data. The majority of officially listed endangered species are large, well known, and popular mammals, such as primates, ungulates, and carnivores, whereas the majority of species known to have gone recently extinct and likely to be currently threatened are small and inconspicuous, such as rodents and bats. Our work not only illustrates the role of ecological, evolutionary, and biogeographic processes in the origin and maintenance of land mammal diversity, it also presents the information at the level of biogeographic regions and political units where management and policy must be applied in order to slow the loss of this diversity.     

A hidden species-area curve

The species-area curve, which describes the relationship between the number of observed species in a geographical region and the area of the region, plays a central role in biogeography. Beyond its scientific importance, the species-area curve is commonly used to assess the loss of species due to habitat loss. When the species-area curve is estimated from spatial samples, the existence of species with low or highly spatially variable abundance exaggerates the true rate at which species accumulate with area. Here, a hidden species-area curve is defined that accounts for this sampling effect and its estimation by maximum likelihood is outlined. Both the species-area curve and the hidden species-area curve are conditioned on the observed species list; thus the analysis does not depend on the total number of observed and unobserved species, that is, species richness. The method is tested by sub-sampling some tropical forest data and found to work well. It is also applied to a classic data set from the deep sea. For these data, accounting for this sampling effect has a large impact.     

Plant Diversity Hotspots in the Atlantic Coastal Forests of Brazil

Abstract:  Plant-diversity hotspots on a global scale are well established, but smaller local hotspots within these must be identified for effective conservation of plants at the global and local scales. We used the distributions of endemic and endemic-threatened species of Myrtaceae to indicate areas of plant diversity and conservation importance within the Atlantic coastal forests ( Mata Atlântica ) of Brazil. We applied 3 simple, inexpensive geographic information system (GIS) techniques to a herbarium specimen database: predictive species-distribution modeling (Maxent); complementarity analysis (DIVA-GIS); and mapping of herbarium specimen collection locations. We also considered collecting intensity, which is an inherent limitation of use of natural history records for biodiversity studies. Two separate areas of endemism were evident: the Serra do Mar mountain range from Paraná to Rio de Janeiro and the coastal forests of northern Espírito Santo and southern Bahia. We identified 12 areas of approximately 35 km² each as priority areas for conservation. These areas had the highest species richness and were highly threatened by urban and agricultural expansion. Observed species occurrences, species occurrences predicted from the model, and results of our complementarity analysis were congruent in identifying those areas with the most endemic species. These areas were then prioritized for conservation importance by comparing ecological data for each.     

Benefits to poorly studied taxa of conservation of bird and mammal diversity on islands

Protected area delineation and conservation action are urgently needed on marine islands, but the potential biodiversity benefits of these activities can be difficult to assess due to lack of species diversity information for lesser known taxa. We used linear mixed effects modeling and simple spatial analyses to investigate whether conservation activities based on the diversity of well‐known insular taxa (birds and mammals) are likely to also capture the diversity of lesser known taxa (reptiles, amphibians, vascular land plants, ants, land snails, butterflies, and tenebrionid beetles). We assembled total, threatened, and endemic diversity data for both well‐known and lesser known taxa and combined these with physical island biogeography characteristics for 1190 islands from 109 archipelagos. Among physical island biogeography factors, island area was the best indicator of diversity of both well‐known and little‐known taxa. Among taxonomic factors, total mammal species richness was the best indicator of total diversity of lesser known taxa, and the combination of threatened mammal and threatened bird diversity was the best indicator of lesser known endemic richness. The results of other intertaxon diversity comparisons were highly variable, however. Based on our results, we suggest that protecting islands above a certain minimum threshold area may be the most efficient use of conservation resources. For example, using our island database, if the threshold were set at 10 km² and the smallest 10% of islands greater than this threshold were protected, 119 islands would be protected. The islands would range in size from 10 to 29 km² and would include 268 lesser known species endemic to a single island, along with 11 bird and mammal species endemic to a single island. Our results suggest that for islands of equivalent size, prioritization based on total or threatened bird and mammal diversity may also capture opportunities to protect lesser known species endemic to islands. Beneficios de los Taxa Poco Estudiados para la Conservación de la Diversidad de Aves y Mamíferos en Islas     

Area-Based Refinement for Selection of Reserve Sites with the Benefit-Function Approach

Abstract:  Optimization of resource use is necessary for efficient conservation planning. Many reserve-selection algorithms aim to identify representative but inexpensive networks, which may lead to selecting small sites due to their lower costs and collectively higher species richness. Nevertheless, larger sites would be preferable regarding species' long-term persistence. An area-based refinement can be used to overcome this problem. We used a reserve-planning framework in which continuous benefit functions valued representation (numbers of populations), and differential species weights were based on a species' local rarity and threatened status. We introduced a refinement based on the species-area relationship that provides relatively higher values for larger sites. We applied the proposed method to rich fen vegetation in southern Finland. The species-area refinement resulted in a network of significantly larger sites with minor trade-offs with representation (numbers of populations). Giving endangered species higher weights ensured that the trade-off occurred mostly between site size and representation of low-priority species. We recommend using a species-area refinement for practical, maximum-coverage conservation planning.     

Variation in species losses from islands: artifacts, extirpation rates, or pre-fragmentation diversity?

Species are being lost from isolated reserves as predicted by ecological theory, prompting calls for larger reserves with higher species immigration rates. However, some large islands have lost a large proportion of their species, whereas some small islands have not lost any. Conservation efforts would be more efficient if the cause of such variation in the relationships among number of species lost, island size, and immigration rate were known. Observed species losses could be affected by the time since islands were isolated, species immigration rates, species extirpation rates, the pre-fragmentation diversity of the region relative to steady state, or overestimation of the pre-fragmentation diversity of islands. To test the last three hypotheses, I compared the intersection points of the island, intraprovincial, and interprovincial species-area relationships of terrestrial mammals from nine archipelagos of land-bridge islands and terrestrial habitat isolates. Species losses from three archipelagos were greater than expected due to reduced immigration rates alone, although I could not resolve if this was due to increased extirpation rates or overestimation of the pre-fragmentation diversity of the islands. Analysis of six archipelagos indicates that the diversity of mammals in two regions of North America is currently below steady state, probably due to the extinction of mammals and glacial retreat during the late Pleistocene. These results have direct implications for reserve planning. When provincial diversity is below steady state, some combinations of reserve size and species immigration rate will allow reserves to maintain their pre-isolation diversity. However, the diversity of provinces relative to steady state is likely to vary, so conservation of a given proportion of a province may not always conserve the same proportion of its species. I present a new species-area relationship for islands formed by fragmentation that replaces the parameter c (fitted constant) with a rotation point. Estimation of this rotation point will allow reserve planners to separate the effects of extirpation and immigration rates on species losses from islands, identify provinces that are below steady-state diversity, and estimate the combinations of reserve size and immigration rate that will prevent loss of species from reserves.     

Predicting the effects of habitat homogenization on marine biodiversity.

Seafloor habitats throughout the world's oceans are being homogenized by physical disturbance. Even though seafloor sediments are commonly considered to be simple and unstructured ecosystems, the negative impacts of habitat homogenization are widespread because resident organisms create much of their habitat's structure. We combine the insight gained from remote sensing of seafloor habitats with recently developed analytical techniques to estimate species richness and assess the potential for change with habitat homogenization. Using habitat-dependent species-area relationships we show that realistic scenarios of habitat homogenization predict biodiversity losses when biogenic habitats in soft sediments are homogenized. We develop a simple model that highlights the degree to which the reductions in the number of species and functional diversity are related to the distribution across habitats of habitat-specific and generalist species. Our results suggest that, by using habitat-dependent species-area relationships, we can better predict variation in biodiversity across seafloor landscapes and contribute to improved management and conservation.     

Mitochondrial DNA Phylogeography and the Conservation of Endangered Lesser Antillean Icterus Orioles

Abstract: Recent natural and anthropogenic disturbances have endangered two of the three oriole species endemic to single islands in the Lesser Antilles. The ongoing eruption of the Soufriere Hills volcano may have doomed the Montserrat Oriole (   Icterus oberi ), whereas high levels of nest parasitism by a cowbird threaten the Martinique Oriole (   I. bonana ). These orioles and related Antillean and Central American forms have been considered part of the Icterus dominicensis superspecies complex, but the taxonomic status of the different Antillean island populations has been long debated. To investigate levels of evolutionary differentiation among threatened Lesser Antillean orioles, we analyzed 2507 nucleotides of protein-coding mitochondrial DNA (mtDNA) sequence from orioles on Martinique, Montserrat, St. Lucia (   I. laudabilis ), Puerto Rico (   I. dominicensis dominicensis ), Mexico (   I. d. prosthemelas ), and three Icterus outgroup species. Phylogenetic analyses of the mtDNA data supported the monophyly of Antillean members of the I. dominicensis complex and identified a star-like pattern of relationship among them. Mitochondrial distances between the Antillean populations were large (4.5–5.8% nucleotide divergence) and suggested that the Lesser Antillean orioles have been isolated evolutionarily from one another since the late Pliocene. The oriole taxa on Montserrat, Martinique, and St. Lucia meet species criteria under the phylogenetic species concept and represent evolutionarily significant units. The impending extinction of the phylogenetically unique Montserrat oriole highlights the vulnerability of island endemics to habitat degradation followed by rare and unpredictable natural catastrophes.     

Distribution and conservation status of littoral vascular plant species along the European coasts

A comprehensive list of 1068 typical littoral plant species and subspecies has been composed. They are considered endemic in a wide sense and are subdivided into widespread, transregional, regional and local endemics, the latter three categories being considered as endemics s.s. For each taxon the distribution, habitat preference, endemic status and conservation status are given. The list, which is available upon request, is summarized in a number of figures and tables, from which it appears that 61% of all species are endemics s.s., that ca. 30% of all species are dune and beach species and another nearly 30% are maritime rock species. Species of wet habitats are concentrated in northern and northwestern Europe, dune species in western and southwestern Europe, western Mediterranean and Black Sea. The conservation status of most species is indicated; 37% is considered threatened. It is concluded that the Bern Convention and the European Habitat Directive offer an entirely insufficient framework for effective conservation action. It is suggested to take the present list as a starting point for a geographical/taxonomical/ecological data base of European coastal endemics.     

Significance of Specimen Databases from Taxonomic Revisions for Estimating and Mapping the Global Species Diversity of Invertebrates and Repatriating Reliable Specimen Data

Abstract:  We argue that the millions of specimen-label records published over the past decades in thousands of taxonomic revisions are a cost-effective source of information of critical importance for incorporating invertebrates into biodiversity research and conservation decisions. More specifically, we demonstrate for a specimen database assembled during a revision of the robber-fly genus Euscelidia (Asilidae, Diptera) how nonparametric species richness estimators (Chao1, incidence-based coverage estimator, second-order jackknife) can be used to (1) estimate global species diversity, (2) direct future collecting to areas that are undersampled and/or likely to be rich in new species, and (3) assess whether the plant-based global biodiversity hotspots of Myers et al. (2000) contain a significant proportion of invertebrates. During the revision of Euscelidia , the number of known species more than doubled, but estimation of species richness revealed that the true diversity of the genus was likely twice as high. The same techniques applied to subsamples of the data indicated that much of the unknown diversity will be found in the Oriental region. Assessing the validity of biodiversity hotspots for invertebrates is a formidable challenge because it is difficult to decide whether species are hotspot endemics, and lists of observed species dramatically underestimate true diversity. Lastly, conservation biologists need a specimen database analogous to GenBank for collecting specimen records. Such a database has a three-fold advantage over information obtained from digitized museum collections: (1) it is shown for Euscelidia that a large proportion of unrevised museum specimens are misidentified; (2) only the specimen lists in revisionary studies cover a wide variety of private and public collections; and (3) obtaining specimen records from revisions is cost-effective.     

Guidelines for the Design of Conservation Strategies for the Animals of Venezuela

The geographic distribution of the animals of Venezuela was analyzed as a basis for a series of guidelines to help develop strategies for their conservation. A total of 313 animal taxa is distributed among 24 geographic units, corresponding to the political divisions of the country. Three different criteria were considered in analyzing these data: (1) the number and density of threatened taxa were used to define a ranking system for geographic units; (2) "hotspots" were identified using the distribution of threatened endemics, and (3) a "critical faunas analysis" was used to determine the minimum number of geographic units needed to maximize the number of sampled taxa. The first two criteria emphasize the importance of protecting habitat in the northern portion of the country, where most human intervention has already taken place. But the southern portion of the country is where most of the undisturbed habitat remains. We suggest that the conservation of the animals of Venezuela must follow a mixed strategy, based on two principles: one, aimed mainly at threatened endemics, should focus on the protection of critical habitat north of the Orinoco river, the second, aimed at all threatened animals— particularly high-risk taxa—should focus on assuring the long-term persistence of the pristine habitat in the south.     

Habitat fragmentation and effects of herbivore (howler monkey) abundances on bird species richness

Habitat fragmentation can alter herbivore abundances, potentially causing changes in the plant community that can propagate through the food web and eventually influence other important taxonomic groups such as birds. Here we test the relationship between the density of red howler monkeys (Alouatta seniculus) and bird species richness on a large set of recently isolated land-bridge islands in Lago Guri, Venezuela (n = 29 islands). Several of these islands host relict populations of howler monkeys at densities up to more than 30 times greater than those on the mainland. These "hyperabundant" herbivores previously have been shown to have a strong positive influence on aboveground plant productivity. We predicted that this should lead to a positive, indirect effect of howler monkey density on bird species richness. After accounting for passive sampling (the tendency for species richness to be positively associated with island area, regardless of differences in habitat quality) we found a significant positive correlation between howler monkey density and bird species richness. A path analysis incorporating data on tree growth rates from a subset of islands (n = 9) supported the hypothesis that the effect of howler monkeys on the resident bird communities is indirect and is mediated through changes in plant productivity and habitat quality. These results highlight the potential for disparate taxonomic groups to be related through indirect interactions and trophic cascades.     

Evaluating Biodiversity and Conserving Lake Malawi's Cichlid Fish Fauna

The cichlid fishes of the Great Lakes of Africa are extraordinary biological assemblages. Conservation strategies are urgently needed for these fish faunas, due to increased threats from over-fishing, habitat degradation, and species introductions. Here I investigate the patterns of biodiversity of the rock-dwelling cichlid fishes (known as mbuna) of Lake Malawi and present a series of recommendations to enhance their conservation. Specifically, I examine intralacustrine biogeograpbic distributions and diversity, and levels of endemism for both mbuna species and genera. The biogeographic patterns show a high degree of variation for both taxonomic designations. Significant relationships are found between diversity and distribution patterns. Speciose genera are more widespread and habitats close to other lands support higher biodiversity than in isolated islands. The results indicate that the genus level may be the appropriate taxonomic designation to consider for conservation purposes, and that areas of high generic endemism, such as the Muleri Islands, are significant for conservation. In order to account for geographic variation in biodiversity, consideration should be given to expanding Lake Malawi National Park boundaries to include areas in the northern and central parts of the lake, especially Likoma and Chisumulu Islands and Nkhata Bay. I discuss the establishment of an international park and the expansion of conservation efforts to include the non-mbuna cichlids. An ecosystem approach to biodiversity conservation and management is advocated.     

Predicting the Spatial Distribution of a Seabird Community to Identify Priority Conservation Areas in the Timor Sea

Understanding spatial and temporal variability in the distribution of species is fundamental to the conservation of marine and terrestrial ecosystems. To support strategic decision making aimed at sustainable management of the oceans, such as the establishment of protected areas for marine wildlife, we identified areas predicted to support multispecies seabird aggregations in the Timor Sea. We developed species distribution models for 21 seabird species based on at‐sea survey observations from 2000–2013 and oceanographic variables (e.g., bathymetry). We applied 4 statistical modeling techniques and combined the results into an ensemble model with robust performance. The ensemble model predicted the probability of seabird occurrence in areas where few or no surveys had been conducted and demonstrated 3 areas of high seabird richness that varied little between seasons. These were located within 150 km of Adele Island, Ashmore Reef, and the Lacepede Islands, 3 of the largest aggregations of breeding seabirds in Australia. Although these breeding islands were foci for high species richness, model performance was greatest for 3 nonbreeding migratory species that would have been overlooked had regional monitoring been restricted to islands. Our results indicate many seabird hotspots in the Timor Sea occur outside existing reserves (e.g., Ashmore Reef Marine Reserve), where shipping, fisheries, and offshore development likely pose a threat to resident and migratory populations. Our results highlight the need to expand marine spatial planning efforts to ensure biodiversity assets are appropriately represented in marine reserves. Correspondingly, our results support the designation of at least 4 new important bird areas, for example, surrounding Adele Island and Ashmore Reef. Pronostico de la Distribución Espacial de una Comunidad de Aves Marinas para Identificar Áreas Prioritarias de Conservación en el Mar de Timor