The high biodiversity of benthic organisms in a coastal ecosystem revealed by an integrative approach / A alta biodiversidade de organismos bentônicos em um ecossistema costeiro revelada por uma abordagem integrativa

Abstract Increasing habitat modification and species loss demand consistent efforts to describe and understand biodiversity patterns. The BIOTA/FAPESP Program was created in this context and it has been a successful initiative to promote studies on biodiversity and conservation in Brazil. The BIOTA/Araçá is an interdisciplinary project that provided a detailed evaluation of the biodiversity of Araçá Bay, a coastal seascape located on the North coast of the state of São Paulo, Southeast Brazil. The bay encompasses multiple habitats, such as beaches, mangroves, rocky shores, and a tidal flat, and provides important ecosystem services. Unfortunately, the bay is the subject of complex social-environmental conflicts that oppose economic, social, and environmental demands (i.e., the expansion of neighboring harbor activities vs. small-scale artisanal fisheries and protection of biodiversity). The present study presents a survey of the benthic species occurring in the different habitats of Araçá Bay, including data obtained during the BIOTA/Araçá project and previous assessments of the area. The benthic species play an important role in marine environments and studying the diversity of these organisms that live associated with the bottom is indispensable for comprehending the environment's functioning. The macrofauna, meiofauna, and microorganisms associated with soft and hard bottom were listed, and additional information, such as the habitat and geographical distribution, were provided for each species. The checklist includes 826 species, almost 70% recorded during the BIOTA/Araçá project. The most speciose taxa were the annelids (225 spp.), mollusks (194 spp.), and crustaceans (177 spp.). Seven benthic species are endemic to Araçá Bay, 14 are considered threatened, and seven are economically exploited. Furthermore, the bay is the type locality of many taxa, and 11 new benthic species were described based on specimens sampled during the project. This project shows the importance of Araçá Bay as a unique biologically rich environment and highlights the need for conservation efforts in light of the current threats.

Resumo O aumento da modificação dos habitats e da perda de espécies demanda esforços consistentes para descrever e compreender os padrões de biodiversidade. O programa BIOTA/FAPESP foi criado nesse contexto e é uma iniciativa de sucesso para promover estudos em biodiversidade e conservação no Brasil. O BIOTA/Araçá é um projeto interdisciplinar que promoveu uma avaliação detalhada da biodiversidade da Baía do Araçá, um ecossistema costeiro localizado ao Norte do estado de São Paulo, Sudeste do Brasil. A baía engloba múltiplos habitats, tais como praias, manguezais, costões rochosos, e uma planície de maré, e também fornece importantes serviços ecossistêmicos. Infelizmente, a baía está sujeita à conflitos sócio-ambientais complexos que contrastam demandas econômicas, sociais e ambientais (i.e. a expansão das atividades do porto vizinho vs. a pesca artesanal de pequena escala e a proteção da biodiversidade). O presente estudo apresenta um levantamento das espécies bentônicas que ocorrem nos diferentes habitats da Baía do Araçá, incluindo dados obtidos durante o projeto BIOTA/Araçá e de investigações realizadas anteriormente na área. As espécies bentônicas desempenham um papel importante no ambiente marinho, e estudar a diversidade desses organismos que vivem associados ao fundo é indispensável para compreender o funcionamento do meio ambiente. A macrofauna, meiofauna, e microorganismos associados aos fundos consolidado e inconsolidado foram listados, e informações adicionais foram fornecidas para cada espécie, tais como a distribuição geográfica e nos habitats. O checklist inclui 826 espécies, quase 70% registradas durante o projeto BIOTA/Araçá. Os taxa mais especiosos foram os anelídeos (225 spp.), moluscos (194 spp.), e crustáceos (177 spp.). Entre as espécies bentônicas listadas, sete são endêmicas da Baía do Araçá, 14 são consideradas ameaçadas de extinção, e sete são exploradas economicamente. A baía é a localidade tipo de vários taxa, e 11 novas espécies bentônicas foram descritas com base em espécimes amostrados durante o projeto. Este projeto mostra a importância da Baía do Araçá como um ambiente de riqueza biológica única e demonstra a necessidade de esforços para a sua conservação considerando as atuais ameaças.

Georeferenced database and interactive online map of limnoterrestrial and freshwater Tardigrada from Central and South America / Banco de dados georreferenciado e mapa interativo online de Tardigrada terrestres e aquáticos da América Central e América do Sul

Abstract Like other meiofaunal organisms, tardigrades suffer from a significant knowledge gap concerning many aspects of their biodiversity. The lack of an up-to-date digital collection with all species and details of limnoterrestrial and freshwater tardigrades in South and Central America is one of the most critical gaps to be filled. Therefore, the present work aims to develop a database containing all valid species of limnoterrestrial and freshwater tardigrades from South and Central America found until 2023 and provide open access to the results. Data for each species were obtained directly from the literature using Google Scholar and the website tardigrada.net. This compiled data resulted in the creation of a database with the species name, author and year of species description, genus, family, class, type country, type location, coordinates (longitude and latitude), if it is aquatic and/or limnoterrestrial, substrate where it was found, the country and location of collection, and manuscript containing the species identification. Furthermore, the coordinates of each occurrence were plotted on maps with political-administrative boundaries and Neotropical and Andean biogeographic regions. In addition, statistical analysis was performed related to the geographic distribution of the sampling effort. From the literature, 2157 records of valid non-marine Tardigrada species, endemic or not, were computed. From these records, 271 species of tardigrades have been identified in the two regions combined, with 223 species in South America and 129 species in Central America. We were able to show that there are still many biases in the sampling of tardigrades in the Neotropical and Andean regions and that further studies are needed on the biogeography of these meiofaunal organisms in these biogeographic regions. We expect this database to help better understand the richness and distribution patterns of limnoterrestrial and aquatic tardigrade species in Central and South America.

Resumo Tardígrados, assim como outros organismos meiofaunais, possuem uma lacuna de conhecimento significativa acerca de muitos aspectos da sua biodiversidade. A inexistência de um acervo digital, e atualizado, com todas as espécies e detalhes de tardígrados limnoterrestres e aquáticos na América Central e Sul é uma das lacunas mais importantes a serem preenchidas. Dessa maneira, o presente trabalho tem como objetivo elaborar e disponibilizar, de maneira gratuita, um banco de dados contendo todas as espécies válidas de tardígrados limnoterrestres e aquáticos das América do Sul e América Central encontradas até 2023. Os dados de cada espécie foram obtidos diretamente na literatura, utilizando o Google Scholar e o site tardigrada.net. Com todos esses dados compilados, foi elaborado um banco de dados com nome da espécie, autor e ano de descrição da espécie, gênero, família, classe, país tipo, local tipo, coordenadas (longitude e latitude), se é aquático e/ou limnoterrestre, substrato onde foi encontrado, país coletado, local de coleta e manuscrito com a identificação da espécie. Ademais, as coordenadas obtidas de cada ocorrência foram plotadas em mapas das fronteiras político-administrativas e das regiões biogeográficas Neotropical e Andina. Além disso, uma análise estatística quanto à distribuição geográfica do esforço amostral foi feita. Da literatura, foram computados 2157 registros de espécies válidas de tardígrados limnoterrestre, endêmicas ou não. Desses registros, foram descobertas, até hoje, 271 espécies de tardígrados entre as duas regiões, com 223 espécies na América do Sul e 129 espécies na América Central. Foi possível demonstrar que ainda há muito viés na amostragem de tardígrados nas regiões Neotropical e Andina, e mais estudos acerca da biogeografia desses organismos meiofaunais nessas regiões biogeográficas são necessários. A partir desse banco de dados, espera-se contribuir para um maior entendimento da riqueza e dos padrões de distribuição de espécies de tardígrados limnoterrestres e aquáticos nas América Central e Sul.

New records, neotype designation and DNA sequences of three species of Chaetonotus (Gastrotricha: Chaetonotidae) from Brazil.

Gastrotrichs are aquatic microinvertebrates (less than 1 mm long) commonly found in freshwater and marine environments worldwide. The taxonomy of gastrotrichs is mainly based on external morphological characters, such as body shape and size of the different body regions, presence and distribution of adhesive tubules, or the microstructure of the external cuticle (e.g. scales, spines, cephalic plates). The diversity of Gastrotricha is still poorly known, especially in tropical regions. The aim of this study is to use integrative taxonomy to report new records of Chaetonotus dadayi, C. paucisquamatus and C. furcatus from the states of São Paulo and Minas Gerais, Brazil. In this study we present 1) new records of C. dadayi in Brazil with new morphological information and a first-time comprehensive diagnosis of the species; 2) the redescription of C. paucisquamatus based on material collected at the type locality and the neotype designation; and 3) the first record of C. furcatus outside its type locality. We also discuss the actual occurrence of C. paucisquamatus and C. furcatus in Poland, given that both species were found in artificial environments with flora imported from South America.

New data on Brazilian semiplanktonic gastrotrichs (Gastrotricha: Chaetonotida).

Gastrotricha are free-living aquatic microinvertebrates with a ubiquitous distribution. With almost two centuries of accumulated knowledge, currently there are more than 860 described species, but our knowledge on Brazilian Gastrotricha is still far from complete. This statement is even more precise when considering freshwater semiplanktonic gastrotrichs. We aim to contribute to biodiversity knowledge of Brazilian gastrotrichofauna with new records of Dasydytes lamellatus Kisielewski, 1991, Haltidytes pseudosquamosus Minowa & Garraffoni, 2017 and Neogossea acantholla Kisielewski, 1991. We provide morphometric, molecular and phylogenetic data for these species, and the first electron microscopy record of D. lamellatus revealing previously inaccessible characters.

Geographical sampling bias on the assessment of endemism areas for marine meiobenthic fauna.

Species distribution patterns are constrained by historical and ecological processes in space and time, but very often the species range sizes are geographical sampling biases resulting from unequal sampling effort. One of the most common definitions of endemism is based on the "congruence of distributional areas" criterion, when two or more species have the same distributional limits. By acknowledging that available data of marine meiobenthic species are prone to geographical sampling bias and that can affect the accuracy of the biogeographical signals, the present study combines analyses of inventory incompleteness and recognition of spatial congruence of Gastrotricha, Kinorhyncha, meiobenthic Annelida and Tardigrada in order to better understand the large-scale distribution of these organisms in coastal and shelf areas of the world. We used the marine bioregionalization framework for geographical operative units to quantify the inventory incompleteness effect (by modelling spatial predictions of species richness) and to recognize areas of endemism. Our models showed that the difference between observed and expected species richness in the Southern Hemisphere is much higher than in the Northern Hemisphere. Parsimony Analysis of Endemicity delimited 20 areas of endemism, most found in the Northern Hemisphere. Distribution patterns of meiobenthic species are shown to respond to events of geographical barriers and abiotic features, and their distribution is far from homogeneous throughout the world. Also, our data show that ecoregions with distinct biotas have at least some cohesion over evolutionary time. However, we found that inventory incompleteness may significantly affect the explanatory power of areas of endemism delimitation in both hemispheres. Yet, whereas future increases in sampling efforts are likely to change the spatial congruence ranges in the Southern Hemisphere, patterns for the Northern Hemisphere may prove to be relatively more resilient.

Assessing biodiversity shortfalls of freshwater meiofauna from the Atlantic Forest: New species, distribution patterns and the first total-evidence phylogeny of semiplanktonic Gastrotricha.

The Brazilian Atlantic forest is a tropical rainforest recognized as a hotspot of biodiversity, with high species richness and endemicity. This forest extends over a wide latitudinal range, bordering the entire Brazilian coastline, from sea level to high mountains over 2000 m.a.s.L., and presents a variety of environmental conditions and forest physiognomy. Despite many years of intense studies on animal biodiversity in the biome, there is a lack of information on meiofauna taxa causing several shortfalls in biodiversity knowledge of these tiny organisms. In this study, we address some of these shortfalls by describing a new species of Neogossea (Gastrotricha: Chaetonotida) from a lentic ecosystem in southeastern Brazil, surrounded by fragments of Atlantic Forest by using an integrative approach combining different morphological techniques and molecular data. We also point out new hypotheses of homologous structures due to scanning electron microscope observations of the new species. Additionally, we used two numerical methods to assess distribution patterns and historical regionalization of four freshwater meiofaunal taxa (Gastrotricha, Rotifera, Copepoda and Cladocera). For the first time, we accessed the areas of endemism in this biological hotspot based on aquatic fauna with a very peculiar life history. Due to sampling issues and meiofauna species being widespread, our results raise incongruences with previous endemism analyses on vertebrates and arthropods. Finally, we performed the first total-evidence phylogenetic analyses of benthic and semiplanktonic gastrotrichs based on 59 morphological characters and three molecular markers, employing a parsimony approach. The phylogenetic reconstruction supports the hypothesis of a single origin of semiplanktonic gastrotrichs, and both Dasydytidae and Neogosseidae families are monophyletic taxa as well as four non-monotypic genera.

A synopsis of knowledge, zoogeography and an online interactive map of Brazilian marine gastrotrichs.

Information regarding the records of Brazilian marine gastrotrichs is presented. We systematized and organized the existing information associated with approximately 23 species (belonging to the genera Aspidiophorus, Chaetonotus, Crasiella, Dactylopodola, Dendrodasys, Draculiciteria, Halichaetonotus, Heteroxenotrichula, Kryptodasys, Macrodasys, Pseudostomella, Ptychostomella, Urodasys and Xenotrichula) from the Brazilian coast (eight endemic) and their 1,581 records from 36 marine ecoregions of the world. A link is provided to an on-line interactive map where all occurrences for each species are shown, accompanied by geographic coordinates, oceans, countries, cities, granulometric characteristics and ecoregions. Furthermore, a critical analysis of the geographical distribution of Brazilian marine gastrotrichs, an estimate of the number of undescribed species, a summary of the existence and status of taxonomical collections are also presented.

Integrative taxonomy of a new Redudasys species (Gastrotricha: Macrodasyida) sheds light on the invasion of fresh water habitats by macrodasyids.

The order Macrodasyida (Gastrotricha) includes over 350 marine species, and only 3 freshwater species (Marinellina flagellata, Redudasys fornerise, R. neotemperatus). Herein we describe a new freshwater species of Macrodasyida, Redudasys brasiliensis sp. nov., from Brazil through an integrative taxonomic approach. The external morphology and internal anatomy were investigated using differential interference contrast microscopy, confocal microscopy, scanning and transmission electron microscopy. The systematization of the new taxon was inferred by nuclear (18S and 28S) and mitochondrial (COI) genes, and its intra-order relationships were assessed using data from most of available macrodasyids. Phylogenetic analyses yielded congruent trees, in which the new taxon is nested within the family Redudasyidae, but it was genetically distinct from the other species of the genus Redudasys. The new species shares the gross morphology and reproductive traits with other Redudasyidae and the presence of only 1 anterior adhesive tube per side with Redudasys neotemperatus, but it has a specific pattern of ventral ciliation and muscle organization. Results support the hypothesis that dispersion into fresh water habitats by Macrodasyida and Chaetonotida taxa occurred independently and that within Macrodasyida a single lineage invaded the freshwater environment only once. Furthermore, the Neotropical region seems to be peculiar for the evolution of the freshwater macrodasyid clade.

Redescription of the dasydytid gastrotrich Haltidytesooëides (Brunson, 1950) based on type material.

The semi-pelagic gastrotrich species Haltidytesooëides (Brunson, 1950) is redescribed based on original type material deposited at the Smithsonian National Museum of Natural History. Herein we present a new diagnosis and figures of the species, detailing the insertion position of the lateral spines, misinterpreted in the original description. Furthermore, we reassess the taxonomic key for the genus Haltidytes Remane, 1936 based on our new findings.

New species and new records of freshwater Heterolepidoderma (Gastrotricha: Chaetonotidae) from Brazil with an identification key to the genus.

A new species of freshwater Heterolepidoderma (Gastrotricha) was found in Brazil. Heterolepidoderma mariae sp. nov. is unique in possessing a three-lobed head, three types of dorsal keeled scales, a thin band of cilia on the head, connecting the two bands of ventral cilia, and an interciliary area with elliptical keeled scales with short spines. Heterolepidoderma famaillense Grosso & Drahg, 1991 is reported for the first time outside the type locality in Argentina, and we make some initial remarks on H. aff. majus Remane, 1927, a possible undescribed species. A dichotomous key for all freshwater species of Heterolepidoderma , with distributional data, is also provided.

New data on freshwater psammic Gastrotricha from Brazil.

Current knowledge of freshwater gastrotrich fauna from Brazil is underestimated as only two studies are available. The present communication is a taxonomic account of the first-ever survey of freshwater Gastrotricha in Minas Gerais State. Samplings were carried out yielding six species of three Chaetonotidae genera: Aspidiophorus cf. pleustonicus, Ichthydium cf. chaetiferum, Chaetonotus acanthocephalus, Chaetonotus heideri, Chaetonotus cf. succinctus, Chaetonotus sp., and also an undescribed species belonging to the genus Redudasys (incertae sedis): this is the first finding of specimens of Redudasys outside of original type locality. These preliminary observations suggest that the knowledge of the biodiversity of Gastrotricha in the Minas Gerais State, as well as in the whole Brazil, will certainly increase as further investigations are undertaken, and that freshwater Macrodasyida may be more common than previously thought.