Indicators to monitor the status of the tree of life.

Following the failure to fully achieve any of the 20 Aichi biodiversity targets, the future of biodiversity rests in the balance. The Convention on Biological Diversity's Kunming-Montreal Global Biodiversity Framework (GBF) presents the opportunity to preserve nature's contributions to people (NCPs) for current and future generations by conserving biodiversity and averting extinctions. There is a need to safeguard the tree of life-the unique and shared evolutionary history of life on Earth-to maintain the benefits it bestows into the future. Two indicators have been adopted within the GBF to monitor progress toward safeguarding the tree of life: the phylogenetic diversity (PD) indicator and the evolutionarily distinct and globally endangered (EDGE) index. We applied both to the world's mammals, birds, and cycads to show their utility at the global and national scale. The PD indicator can be used to monitor the overall conservation status of large parts of the evolutionary tree of life, a measure of biodiversity's capacity to maintain NCPs for future generations. The EDGE index is used to monitor the performance of efforts to conserve the most distinctive species. The risk to PD of birds, cycads, and mammals increased, and mammals exhibited the greatest relative increase in threatened PD over time. These trends appeared robust to the choice of extinction risk weighting. EDGE species had predominantly worsening extinction risk. A greater proportion of EDGE mammals (12%) had increased extinction risk compared with threatened mammals in general (7%). By strengthening commitments to safeguarding the tree of life, biodiversity loss can be reduced and thus nature's capacity to provide benefits to humanity now and in the future can be preserved.

Indicadores para monitorear el estado del árbol de la vida Resumen El futuro de la biodiversidad peligra tras no haberse logrado ninguno de los 20 Objetivos de Aichi. El Marco Global de Biodiversidad (GBF) de Kunming-Montreal del Convenio sobre la Diversidad Biológica (CDB) representa la oportunidad de preservar las contribuciones de la naturaleza a las personas (PNC) para las generaciones actuales y futuras mediante la conservación de la biodiversidad y la prevención de las extinciones. Es necesario salvaguardar el árbol de la vida -la historia evolutiva única y compartida de la vida en la Tierra- para mantener en el futuro los beneficios que aporta. En el GBF se han adoptado dos indicadores para supervisar los avances hacia el cuidado del árbol de la vida: el indicador de diversidad filogenética y el índice de especies evolutivamente distintas y globalmente amenazadas (EDGE). Aplicamos ambos a los mamíferos, las aves y las cícadas del mundo para demostrar su utilidad a escala mundial y nacional. El indicador de diversidad filogenética puede utilizarse para supervisar el estado de conservación general de grandes partes del árbol evolutivo de la vida, una medida de la capacidad de la biodiversidad para mantener los PNC para las generaciones futuras. El índice EDGE se utiliza para supervisar el rendimiento de los esfuerzos por conservar las especies más distintivas. El riesgo para la diversidad filogenética de aves, cícadas y mamíferos aumentó, y los mamíferos mostraron el mayor aumento relativo de la diversidad filogenética amenazada a lo largo del tiempo. Estas tendencias parecieron sólidas a la hora de elegir la valoración del riesgo de extinción. Las especies EDGE tuvieron un riesgo de extinción predominante cada vez peor. Una mayor proporción de mamíferos EDGE (12%) presentó un riesgo de extinción creciente en comparación con los mamíferos amenazados en general (7%). Si se refuerza el compromiso de salvaguardar el árbol de la vida, se puede reducir la pérdida de biodiversidad y preservar así la capacidad de la naturaleza para proporcionar beneficios a la humanidad ahora y en el futuro.

Mapping extrinsic traits such as extinction risks or modelled bioclimatic niches on phylogenies: does it make sense at all?

An increasing variety of extrinsic traits are used in comparative studies aimed at testing evolutionary hypotheses. After briefly reviewing the relevant literature, it appears that three different problems are implied by this trend. Some extrinsic traits are only surrogates for phenotypic traits, and should be redefined to better fit the requisites for phylogenetic analysis, such as selective regimes and extinction risks. Some others are already adequately defined and cannot be made less extrinsic, such as taxon age, geographical distribution, associates (parasites, symbionts, etc.), and bioclimatic modelled niches. Because they are not heritable, they should not be analysed by optimization onto a tree, but are better considered in sister-group comparisons or within a reconciliation procedure, as already done for areas of biogeography. © The Willi Hennig Society 2010.

Arthropod-Microbiota Integration: Its Importance for Ecosystem Conservation.

Recent reports indicate that the health of our planet is getting worse and that genuine transformative changes are pressing. So far, efforts to ameliorate Earth's ecosystem crises have been insufficient, as these often depart from current knowledge of the underlying ecological processes. Nowadays, biodiversity loss and the alterations in biogeochemical cycles are reaching thresholds that put the survival of our species at risk. Biological interactions are fundamental for achieving biological conservation and restoration of ecological processes, especially those that contribute to nutrient cycles. Microorganism are recognized as key players in ecological interactions and nutrient cycling, both free-living and in symbiotic associations with multicellular organisms. This latter assemblage work as a functional ecological unit called "holobiont." Here, we review the emergent ecosystem properties derived from holobionts, with special emphasis on detritivorous terrestrial arthropods and their symbiotic microorganisms. We revisit their relevance in the cycling of recalcitrant organic compounds (e.g., lignin and cellulose). Finally, based on the interconnection between biodiversity and nutrient cycling, we propose that a multicellular organism and its associates constitute an Ecosystem Holobiont (EH). This EH is the functional unit characterized by carrying out key ecosystem processes. We emphasize that in order to meet the challenge to restore the health of our planet it is critical to reduce anthropic pressures that may threaten not only individual entities (known as "bionts") but also the stability of the associations that give rise to EH and their ecological functions.

Revisiting species and areas of interest for conserving global mammalian phylogenetic diversity.

Various prioritisation strategies have been developed to cope with accelerating biodiversity loss and limited conservation resources. These strategies could become more engaging for decision-makers if they reflected the positive effects conservation can have on future projected biodiversity, by targeting net positive outcomes in future projected biodiversity, rather than reflecting the negative consequences of further biodiversity losses only. Hoping to inform the post-2020 biodiversity framework, we here apply this approach of targeting net positive outcomes in future projected biodiversity to phylogenetic diversity (PD) to re-identify species and areas of interest for conserving global mammalian PD. We identify priority species/areas as those whose protection would maximise gains in future projected PD. We also identify loss-significant species/areas as those whose/where extinction(s) would maximise losses in future projected PD. We show that our priority species/areas differ from loss-significant species/areas. While our priority species are mostly similar to those identified by the EDGE of Existence Programme, our priority areas generally differ from previously-identified ones for global mammal conservation. We further highlight that these newly-identified species/areas of interest currently lack protection and offer some guidance for their future management.

Distribution and relative age of endemism across islands worldwide.

Islands have remarkable levels of endemism and contribute greatly to global biodiversity. Establishing the age of island endemics is important to gain insights into the processes that have shaped the biodiversity patterns of island biota. We investigated the relative age of monocots across islands worldwide, using different measures of phylogenetic endemism tested against null models. We compiled a species occurrence dataset across 4,306 islands, and identified 142 sites with neo-, paleo-, mixed and super-endemism. These sites were distributed across the world, although they tended to be more common at low latitudes. The most frequent types of endemism were mixed and super-endemism, which suggests that present-day island biodiversity has frequently been shaped by processes that took place at different points in times. We also identified the environmental factors that contributed most to different types of endemism; we found that latitude, habitat availability and climate stability had a significant impact on the persistence of ancient taxa and on recent diversification events. The islands identified here are irreplaceable both for the uniqueness and the evolutionary history of their flora, and because they are a source of "option values" and evolutionary potential. Therefore, our findings will help guide biodiversity conservation on a global scale.

Vulnerability to climate change of islands worldwide and its impact on the tree of life.

Island systems are among the most vulnerable to climate change, which is predicted to induce shifts in temperature, rainfall and/or sea levels. Our aim was: (i) to map the relative vulnerability of islands to each of these threats from climate change on a worldwide scale; (ii) to estimate how island vulnerability would impact phylogenetic diversity. We focused on monocotyledons, a major group of flowering plants that includes taxa of important economic value such as palms, grasses, bananas, taro. Islands that were vulnerable to climate change were found at all latitudes, e.g. in Australia, Indonesia, the Caribbean, Pacific countries, the United States, although they were more common near the equator. The loss of highly vulnerable islands would lead to relatively low absolute loss of plant phylogenetic diversity. However, these losses tended to be higher than expected by chance alone even in some highly vulnerable insular systems. This suggests the possible collapse of deep and long branches in vulnerable islands. Measuring the vulnerability of each island is a first step towards a risk analysis to identify where the impacts of climate change are the most likely and what may be their consequences on biodiversity.

Taxonomic revision of the genus Monastria Saussure, 1864 (Blattodea: Blaberidae, Blaberinae) from the South American Atlantic forest, with the descriptions of five new species.

 The genus Monastria Saussure, 1864 includes medium to large sized (40-55 mm) dark brown or black cockroaches found in the understory of the Neotropical Atlantic Forest from the Northeast of Brazil to Paraguay and Argentina. The genus shows evident sexual dimorphism: males are elongated with fully developed wings extending beyond the apex of cerci and females are oval and brachypterous. This study is a revision of the genus with redescription of the three already known species, Monastria biguttata (Thunberg, 1826), Monastria similis (Serville, 1838) and Monastria angulata Saussure, 1864, and description of five new ones, Monastria itubera sp. n. and Monastria itabuna sp. n. from state of Bahia, Monastria cabocla sp. n. from state of Sergipe, Monastria kaingangue sp. n. from state of São Paulo and Monastria sagittata sp. n. from state of Minas Gerais. The morphology of the genus and all species is described in details, including male genitalia. Based on this revision, we proposed a new combination for Hiereoblatta papillosa (Thunberg, 1826) comb. n., excluding it from the genus Monastria. For the first time, the juvenile stages of Monastria are characterized and compared to other genera of Blaberinae of the Atlantic forest. Three determination keys are provided. The two first are aimed at identifying the adults and juvenile stages of the five genera of the Blaberinae radiation endemic to the Atlantic forest, respectively. The third concerns the identification of the species of the genus Monastria. A map indicating the localities where species were sampled is also provided.

The informative value of museum collections for ecology and conservation: A comparison with target sampling in the Brazilian Atlantic forest.

Since two decades the richness and potential of natural history collections (NHC) were rediscovered and emphasized, promoting a revolution in the access on data of species occurrence, and fostering the development of several disciplines. Nevertheless, due to their inherent erratic nature, NHC data are plagued by several biases. Understanding these biases is a major issue, particularly because ecological niche models (ENMs) are based on the assumption that data are not biased. Based on it, a recent body of research have focused on searching adequate methods for dealing with biased data and proposed the use of filters in geographical and environmental space. Although the strength of filtering in environmental space has been shown with virtual species, nothing has yet been tested with a real dataset including field validation. In order to contribute to this task, we explore this issue by comparing a dataset from NHC to a recent targeted sampling of the cockroach genus Monastria Saussure, 1864 in the Brazilian Atlantic forest. We showed that, despite strong similarities, the area modeled with NHC data was much smaller. These differences were due to strong climate biases, which increased model's specificity and reduced sensitivity. By applying two forms of rarefaction in the environmental space, we showed that deleting points at random in the most biased climate class is a powerful way for increasing model's sensitivity, so making predictions more suitable to the reality.

Parsimony analysis and ecological communities, phytosociology, nested subsets analysis, forest fragmentation: a reply to Giannini and Keller.

We suggested using parsimony analysis to study community evolution in terms of species composition and to apply these results in the context of forest fragmentation as a replacement for the so-called "nested subsets analysis" or other phenetic synecological or phytosociological methods (Pellens et al., 2005). Giannini and Keller (2007) took issue with this new application on the basis of three misunderstandings. We re-emphasize that communities themselves are analyzed, not landscape parts such as forest fragments. Therefore, it must be clear that communities are analogous to taxa and landscape parts such as fragments are analogous to distributions of taxa. Community evolution is the change in community composition by immigration, emigration and local extinction. Thus, gains and losses of species should not be confused with horizontal transfer. Parsimony analysis does not necessarily group communities based on shared absences of rare species. Rare species are not necessarily absent in the same communities and these absences are not necessarily inferred to be synapomorphies after rooting. This is the main advance expected when cladistics is used instead of the previously cited phenetic methods working with overall similarity.

The conservation-refugium value of small and disturbed Brazilian Atlantic forest fragments for the endemic ovoviviparous cockroach Monastria biguttata (Insecta: Dictyoptera, Blaberidae, Blaberinae).

The Brazilian Atlantic forest is a biodiversity hotspot and harbors many endemic species showing peculiar and unique traits. However, it has been reduced to less than 8% of its original surface and is distributed in scattered fragments, the great majority of which are smaller than 20 hectares and very disturbed, making it worth asking about their value for conservation. In this paper we assess the refugium value of small fragments to the conservation of one of the endemics of the Atlantic forest, the ovoviviparous cockroach Monastria biguttata. Our results showed that this species was ubiquitous in large and small forest fragments, but never present in plantations or pastures. The population age structure and sex ratio were balanced in every fragment, and total population size in the smallest fragments was at least several hundreds of individuals. Colony size, sex ratio, age structure, and density per piece of dead tree trunk indicated that populations from small fragments were not unbalanced or at risk of extinction. According to the analysis of resource availability, small fragments can provide suitable habitat for this species. In this situation, even very small forest fragments have a high refugium value for some endemic insect species. Considering their number in the landscape, these fragments should be considered with more attention in strategies of biodiversity conservation.

Outstanding micro-endemism in New Caledonia: More than one out of ten animal species have a very restricted distribution range.

New Caledonia is a biodiversity hotspot, with an extremely high number of endemic species with narrow distribution ranges that are at high risk of extinction due to open-cast nickel mining, invasive species and seasonal man-induced fires. Mentions of micro-endemism permeate the literature on the biota of this archipelago. However, so far there has been no research comparing distribution range in different animal groups. The aim of this study is to examine the implication of different sampling effort variables in order to distinguish micro-endemicity from data deficiency, and evaluate the distribution range, frequency, and extent to which micro-endemism is common to several groups of organisms. We compiled a dataset derived from publications in Zoologia Neocaledonica, comprising 1,149 species, of which 86% are endemic to New Caledonia. We found that the sampling effort variables that were best correlated with distribution range were the number of sampling dates and the number of collectors per species. The median value of sampling dates was used to establish a cut-off point for defining adequately sampled species. We showed that, although only 52% of species were sampled adequately enough to determine their distribution range, the number of species with a very narrow distribution range was still high. Among endemics from New Caledonia, 12% (116 species) have ranges ≤5.2km2 and 3.9% (38 species) have ranges between 23 and 100 km2. Surprisingly, a similar trend was observed in non-endemic species: 22% occurred in areas ≤ 5.2 km2, and 8% in areas 23-100 km2, suggesting that environmental dissimilarity may play an important role in the distribution of these species. Micro-endemic species were predominant in 18 out of 20 orders. These results will contribute to a re-assessment of the IUCN red list of species in this archipelago, indicating that at least 116 species are probably critically endangered.

Updating the Phylogenetic Dating of New Caledonian Biodiversity with a Meta-analysis of the Available Evidence.

For a long time, New Caledonia was considered a continental island, a fragment of Gondwana harbouring old clades that originated by vicariance and so were thought to be locally ancient. Recent molecular phylogenetic studies dating diversification and geological data indicating important events of submergence during the Paleocene and Eocene (until 37 Ma) brought evidence to dismiss this old hypothesis. In spite of this, some authors still insist on the idea of a local permanence of a Gondwanan biota, justifying this assumption through a complex scenario of survival by hopping to and from nearby and now-vanished islands. Based on a comprehensive review of the literature, we found 40 studies dating regional clades of diverse organisms and we used them to test the hypothesis that New Caledonian and inclusive Pacific island clades are older than 37 Ma. The results of this meta-analysis provide strong evidence for refuting the hypothesis of a Gondwanan refuge with a biota that originated by vicariance. Only a few inclusive Pacific clades (6 out of 40) were older than the oldest existing island. We suggest that these clades could have extinct members either on vanished islands or nearby continents, emphasizing the role of dispersal and extinction in shaping the present-day biota.

Phylogenetic algorithms and the evolution of species communities in forest fragments.

In forest fragmentation studies, low specific richness in small fragments and community nestedness are usually considered to result from species loss. However, except in the case of fragmentation experiments, these studies cannot distinguish between original low richness and secondary species loss, or between original high richness and secondary colonizations in fragments. To distinguish between these possibilities is a matter of historical inference for which phylogenetic algorithms are designed. The methods of phylogenetic analysis, and especially parsimony analysis, can be used to find a tree of relationships between communities from different forest fragments, taking the presence or absence of species among different communities as characters. Parsimony analysis searches if species subsets can be classified in a nested hierarchy, and also establishes how the communities evolved, polarizing species changes into either extinctions or colonizations. By re-analyzing two classical studies in this new and powerful way, we demonstrate that the differences between fragments and large continuous forests cannot be attributed to species loss in all cases, contrary to expectations from models.

Evolution on a shaky piece of Gondwana: is local endemism recent in New Caledonia?

New Caledonia is well known as a hot spot of biodiversity whose origin as a land mass can be traced back to the Gondwanan supercontinent. The local flora and fauna, in addition to being remarkably rich and endemic, comprise many supposedly relictual groups. Does the New Caledonian biota date back to Gondwanan times, building up its richness and endemism over 100 Myr or does it result from recent diversifications after Tertiary geological catastrophic events? Here we use a molecular phylogenetic approach to answer this question with the study of the Neocaledonian cockroach genus Angustonicus belonging to the subfamily Tryonicinae from Australia and New Caledonia. Both geological and molecular dating show that the diversification of this group is less than two million years old, whatever the date of its origin itself. This dating is not consistent with hypotheses of Gondwanan richness and endemism in New Caledonian biota. In other terms, local richness and endemism at the specific level are not necessarily related to an old Gondwanan origin of the Neocaledonian groups.

Catalogue of Dictyoptera from Syria and neighbouring countries (Lebanon, Turkey, Iraq and Jordan).

This study is a catalogue of Dictyoptera (Mantodea, Isoptera and Blattaria) from five Eastern Mediterranean countries (Syria, Lebanon, Turkey, Iraq and Jordan). There are 75 species of Dictyoptera known to occur in these countries. These species belong to 15 families (eight of Mantodea, four of Isoptera and three of Blattaria). Mantodea is by far the dictyopteran group with the highest richness with 43 species occurring in this region, followed by Blattaria, with 21, and Isoptera with 11. Turkey is the place with the highest number of Dictyoptera (34%), followed by Iraq (23%) then Syria (22%), Jordan (15%) and Lebanon (7%). An analysis of accumulated number of species along time shows that most of this biodiversity was described during the 20th century, and that Mantodea is the group with the highest number of species described more recently. If this curve is taken as an estimator of the increase of diversity with new prospections, this indicates that the number of Mantodea in this region would be much higher than presently known. Conversely, the local richness of Blattaria and Isoptera are likely to be close to the present numbers, as the curves remain steady for about 100 years. An accumulation curve of species described with occurrence restricted to these five countries shows that most of them were described at the beginning of the 20th century. An analysis of the number of references dealing with each of these species along time reveals that Mantodea is the dictyopteran group most studied in all periods except the second half of the 20th century, when Isoptera was more cited. The types of these species are distributed in 29 institutions, but are mainly concentrated in four major European collections.

Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence.

Understanding the origin and diversification of organisms requires a good phylogenetic estimate of their age and diversification rates. This estimate can be difficult to obtain when samples are limited and fossil records are disputed, as in Dictyoptera. To choose among competing hypotheses of origin for dictyopteran suborders, we root a phylogenetic analysis (~800 taxa, 10 kbp) within a large selection of outgroups and calibrate datings with fossils attributed to lineages with clear synapomorphies. We find the following topology: (mantises, (other cockroaches, (Cryptocercidae, termites)). Our datings suggest that crown-Dictyoptera-and stem-mantises-would date back to the Late Carboniferous (~ 300 Mya), a result compatible with the oldest putative fossil of stem-dictyoptera. Crown-mantises, however, would be much more recent (~ 200 Mya; Triassic/Jurassic boundary). This pattern (i.e., old origin and more recent diversification) suggests a scenario of replacement in carnivory among polyneopterous insects. The most recent common ancestor of (cockroaches + termites) would date back to the Permian (~275 Mya), which contradicts the hypothesis of a Devonian origin of cockroaches. Stem-termites would date back to the Triassic/Jurassic boundary, which refutes a Triassic origin. We suggest directions in extant and extinct species sampling to sharpen this chronological framework and dictyopteran evolutionary studies.

Biodiversity sampling using a global acoustic approach: contrasting sites with microendemics in New Caledonia.

New Caledonia is a Pacific island with a unique biodiversity showing an extreme microendemism. Many species distributions observed on this island are extremely restricted, localized to mountains or rivers making biodiversity evaluation and conservation a difficult task. A rapid biodiversity assessment method based on acoustics was recently proposed. This method could help to document the unique spatial structure observed in New Caledonia. Here, this method was applied in an attempt to reveal differences among three mountain sites (Mandjélia, Koghis and Aoupinié) with similar ecological features and species richness level, but with high beta diversity according to different microendemic assemblages. In each site, several local acoustic communities were sampled with audio recorders. An automatic acoustic sampling was run on these three sites for a period of 82 successive days. Acoustic properties of animal communities were analysed without any species identification. A frequency spectral complexity index (NP) was used as an estimate of the level of acoustic activity and a frequency spectral dissimilarity index (Df ) assessed acoustic differences between pairs of recordings. As expected, the index NP did not reveal significant differences in the acoustic activity level between the three sites. However, the acoustic variability estimated by the index Df , could first be explained by changes in the acoustic communities along the 24-hour cycle and second by acoustic dissimilarities between the three sites. The results support the hypothesis that global acoustic analyses can detect acoustic differences between sites with similar species richness and similar ecological context, but with different species assemblages. This study also demonstrates that global acoustic methods applied at broad spatial and temporal scales could help to assess local biodiversity in the challenging context of microendemism. The method could be deployed over large areas, and could help to compare different sites and determine conservation priorities.

Climate and soil type together explain the distribution of microendemic species in a biodiversity hotspot.

The grasshopper genus Caledonula, endemic to New Caledonia, was studied to understand the evolution of species distributions in relation to climate and soil types. Based on a comprehensive sampling of 80 locations throughout the island, the genus was represented by five species, four of which are new to science, of which three are described here. All the species have limited distributions in New Caledonia. Bioclimatic niche modelling shows that all the species were found in association with a wet climate and reduced seasonality, explaining their restriction to the southern half of the island. The results suggest that the genus was ancestrally constrained by seasonality. A molecular phylogeny was reconstructed using two mitochondrial and two nuclear markers. The partially resolved tree showed monophyly of the species found on metalliferous soils, and molecular dating indicated a rather recent origin for the genus. Adaptation to metalliferous soils is suggested by both morphological changes and radiation on these soils. The genus Caledonula is therefore a good model to understand the origin of microendemism in the context of recent and mixed influences of climate and soil type.

The evolutionary transition from subsocial to eusocial behaviour in Dictyoptera: phylogenetic evidence for modification of the "shift-in-dependent-care" hypothesis with a new subsocial cockroach.

Cockroaches have always been used to understand the first steps of social evolution in termites because they are close relatives with less complex and integrated social behaviour. Termites are all eusocial and ingroup comparative analysis would be useless to infer the origin of their social behaviour. The cockroach genus Cryptocercus was used as a so-called "prototermite" model because it shows key-attributes similar to the termites (except Termitidae): wood-feeding, intestinal flagellates and subsocial behaviour. In spite of these comparisons between this subsocial cockroach and eusocial termites, the early and remote origin of eusocial behaviour in termites is not well understood yet and the study of other relevant "prototermite" models is however needed. A molecular phylogenetic analysis was carried out to validate a new "prototermite" model, Parasphaeria boleiriana which shows a peculiar combination of these key-attributes. It shows that these attributes of Parasphaeria boleiriana have an independent origin from those of other wood-eating cockroaches and termites. The case of P. boleiriana suggests that a short brood care was selected for with life on an ephemeral wood resource, even with the need for transmission of flagellates. These new phylogenetic insights modify evolutionary hypotheses, contradicting the assumption made with Cryptocercus model that a long brood care is necessary for cooperation between broods in the "shift-in-dependent-care" hypothesis. An ephemeral wood resource is suggested to prompt generation overlap and the evolution of cooperation, even if brood care is shortened.