The testate lobose amoebae (order Arcellinida Kent, 1880) finally find their home within Amoebozoa

Testate lobose amoebae (order Arcellinida Kent, 1880) are common in all aquatic and terrestrial habitats, yet they are one of the last higher taxa of unicellular eukaryotes that has not found its place in the tree of life. The morphological approach did not allow to ascertain the evolutionary origin of the group or to prove its monophyly. To solve these challenging problems, we analyzed partial small-subunit ribosomal RNA (SSU rRNA) genes of seven testate lobose amoebae from two out of the three suborders and seven out of the 13 families belonging to the Arcellinida. Our data support the monophyly of the order and clearly establish its position among Amoebozoa, as a sister-group to the clade comprising families Amoebidae and Hartmannellidae. Complete SSU rRNA gene sequences from two species and a partial actin sequence from one species confirm this position. Our phylogenetic analyses including representatives of all sequenced lineages of lobose amoebae suggest that a rigid test appeared only once during the evolution of the Amoebozoa, and allow reinterpretation of some morphological characters used in the systematics of Arcellinida.     

Diversity and biogeography of testate amoebae

Testate amoebae are amoeboid protists inhabiting a test (shell). They occur globally in soils, wetlands and freshwater, especially peats and mosses. They are of ancient origin, dating from at least the Mesozoic, with possible ancestors as old as the Neoproterozoic. Approximately 2,000 taxa have been described—a number which could easily rise to 4,000 with comprehensive recording. Whilst many protists appear to be cosmopolitan as morphospecies, some of the larger testate species (exceeding 100 μm) have long been considered, controversially, to be geographically restricted. Definitive conclusions have often been confounded by gaps in distributional data and misidentification. Recent increases in recording from previously little known regions, and the rise of molecular taxonomy, have started to resolve outstanding issues—processes still far from complete. Accordingly, biogeographical studies have concentrated on “flagship” species—those which can be identified with certainty and are sufficiently recorded to determine their ecological ranges. Apodera vas (Certes) has been proved to be largely restricted to the Gondwanaland continents and sub-Antarctic islands, but absent from the Holartic despite the availability of much suitable habitat. An early analysis postulated a Mesozoic origin of the species and a distribution influenced by continental drift. Recent molecular evidence could imply a later origin. Either way, its current distribution is clearly influenced by the pattern of global wind currents and lack of lowland tropical habitat. By contrast a “Gondwana-tropical” group of species appears to be restricted to latitudes unaffected by glaciation. Instances of local endemism, such as restriction to a single island, are also known, which await molecular evidence for substantiation. Special Issue: Protist diversity and geographic distribution. Guest editor: W. Foissner.     

COI barcoding of Nebelid testate amoebae (Amoebozoa: Arcellinida): extensive cryptic diversity and redefinition of the Hyalospheniidae Schultze

We used Cytochrome Oxidase Subunit 1 (COI) to assess the phylogenetic relationships and taxonomy of Nebela sensu stricto and similar taxa (Nebela group, Arcellinida) in order to clarify the taxonomic validity of morphological characters. The COI data not only successfully separated all studied morphospecies but also revealed the existence of several potential cryptic species. The taxonomic implications of the results are: (1) Genus Nebela is paraphyletic and will need to be split into at least two monophyletic assemblages when taxon sampling is further expanded. (2) Genus Quadrulella, one of the few arcellinid genera building its shell from self-secreted siliceous elements, and the mixotrophic Hyalosphenia papilio branch within the Nebela group in agreement with the general morphology of their shell and the presence of an organic rim around the aperture (synapomorphy for Hyalospheniidae). We thus synonymise Hyalospheniidae and Nebelidae. Hyalospheniidae takes precedence and now includes Hyalosphenia, Quadrulella (previously in the Lesquereusiidae) and all Nebelidae with the exception of Argynnia and Physochila. Leptochlamys is Arcellinida incertae sedis. We describe a new genus Padaungiella Lara et Todorov and a new species Nebela meisterfeldi n. sp. Heger et Mitchell and revise the taxonomic position (and rank) of several taxa. These results show that the traditional morphology-based taxonomy underestimates the diversity within the Nebela group, and that phylogenetic relationships are best inferred from shell shape rather than from the material used to build the shell.     

Leaf structure of the cerrado (Brazilian savanna) woody plants

With the aim of recognizing the commonest leaf pattern found in the woody flora of the cerrado (the Brazilian savanna) we analyzed the leaf anatomy of 30 representative species. The leaves are mostly dorsiventral and hypostomatic and covered by trichomes and a thick layer of wax and cuticle; the vascular bundles are surrounded by a sheath of fibers. The mesophyll has a developed palisade tissue, dispersed sclerified cells and idioblasts bearing crystals and phenolic compounds. We compared the results with those reported for other species (60 species) from the same biome and for the families that the studied species belong. The present study suggests that the xeromorphism observed for the cerrado leaves is related to the evolutionary history of this biome, since its first floristic elements must have faced deficient water conditions as well as the consequent soil acidity and toxicity. Therefore we may infer that the leaf anatomical pattern here observed was already present in the first elements of the cerrado and was selected to guarantee the survival of those species in the new environment. Furthermore, the xeromorphic features present in those leaves continue nowadays to help the plants protecting themselves from the different biotic and abiotic factors they are subjected to.     

Soil-vegetation relationships in cerrado (Brazilian savanna) and semideciduous forest, Southeastern Brazil

Several studies pointed out soil properties as the prime determinant ofcerrado (the Brazilian savanna) physiognomies, and a gradient from campocerrado (a shrub savanna) to cerradão (a tallwoodland) has been correlated with a soil fertility gradient. Based on thishypothesis, we investigated soil-vegetation relationships in thePé-de-Gigante Reserve (São Paulo State,SoutheasternBrazil). We randomly distributed 10 quadrats (10 × 10 m) oneach ofthe following physiognomies: campo cerrado, cerradosensu stricto, cerradão, andseasonal semideciduous forest, previously defined by the analysis of satelliteimages (LANDSAT-5). We sampled the woody individuals with stem diameter> 3 cm at soil level, identifying their species. In each quadrat, wecollectedsoil samples at the depths of 0–5, 5–25, 40–60, and80–100 cm, and determined pH, K, Ca, Mg, P, Al, H + Al, basesaturation, aluminium saturation, cation exchange capacity, and percentage ofsand, clay and loam. Obtained data were submitted to a canonical correspondenceanalysis (CCA) and to a detrended correspondence analysis (DCA). Our resultsshowed a clear distinction between semideciduous forest and the cerradophysiognomies, based in soil parameters. The former was related to higherconcentrations of cations and clay in the soil, while the latter was related tohigher concentrations of exchangeable aluminium in the soil surface. The threecerrado physiognomies – campo cerrado, cerradosensu stricto, and cerradão– could not be distinguished considering plant density and the analysedsoil features.     

Floristics and environmental factors determining the geographic distribution of epiphytic bromeliads in the Brazilian Atlantic Rain Forest

We investigated how epiphytic species and subfamilies of Bromeliaceae change along the extent of the Atlantic Rain Forest, to answer the questions: (i) How do the epiphytic genera and subfamilies of Bromeliaceae change along the domain? (ii) How similar are the different regions of the Atlantic Rain Forest in relation to the epiphytic species of bromeliads? (iii) Which environmental variables are the most important factors in determining species composition along the domain? We found 114 species of Bromelioideae and 73 of Tillandsioideae. The predominance of Bromelioideae was unexpected, because they are not wind-dispersed as would be expected for most epiphytes. The smaller number of species of Tillandsioideae, and the high frequency of species of Vriesea with limited geographic distributions indicated that epiphytes with rather limited geographic distributions predominate in this domain. Species similarity was divided into one block of south–southeastern localities, and a second block of northeastern–southeastern localities. These results suggest that the distribution of epiphytic bromeliad species resembles that of the phorophyte trees, more than a previous pattern suggested for all epiphytes in the domain. Latitude, temperature and altitude were important factors affecting the species composition along the domain. In general, our results differ from those of other studies in Latin America, and we suggest that historical and evolutionary events generated these differences.     

Dermamoeba algensis n. sp. (Amoebozoa, Dermamoebidae): an algivorous lobose amoeba with complex cell coat and unusual feeding mode

The genus Dermamoeba unifies oblong, flattened amoebae of lingulate morphotype, possessing a thick multilayered cell coat. It includes two species, D. granifera and D. minor. In this paper we describe a third species of this genus, D. algensis n. sp. This species is algivorous; engulfing a large algal cell, it destroys part of the cell coat liberating the plasma membrane, which forms the food vacuole. Thus the glycocalyx never appears inside the phagosome. This observation confirms that some of the thick-coated amoebae may use this way to avoid energetically costly digestion of their own glycocalyx. Studies of the physiology of this organism show that it feeds most actively at a temperature of 22-25 °C. Below and above this temperature the feeding intensity drastically decreases. The new species can survive NaCl concentrations up to 5%, which roughly corresponds to 50 ppt salinity. Accordingly, D. algensis has a wide range of salinity tolerance.     

Description and Phylogenetic Relationships of Spumochlamys perforata n. sp. and Spumochlamys bryora n. sp. (Amoebozoa, Arcellinida)

ABSTRACT. Spumochlamys perforata n. sp. and Spumochlamys bryora n. sp. were isolated and described from dry epiphytic moss. The morphology and ultrastructure of both species clearly demonstrate that they belong to the genus Spumochlamys (family Microchlamyiidae). They differ from its only described member, Spumochlamys iliensis (as well as from species of Microchlamys ), in the relief of the dorsal surface of the test, revealed by scanning electron microscopy, which can represent a good characteristic for species identification. They also differ in the structure of the dorsal part of the test wall (especially S. perforata ). Small subunit ribosomal DNA-based molecular phylogenetic analyses show that Spumochlamys is a deeply branching lineage of the Arcellinida, without any close affinities. Actin gene sequence analysis places this genus within the Tubulinea, close to two other arcellinid lineages but without forming a monophyletic group with them. These data together strongly suggest that the lack of resolution in the arcellinid molecular phylogenies is due to serious undersampling of taxa, a limited number of sequence data, and high divergence rates in most of the species.     

Ribosomal RNA genes challenge the monophyly of the Hyalospheniidae (Amoebozoa: Arcellinida)

To date only five partial and two complete SSU rRNA gene sequences are available for the lobose testate amoebae (Arcellinida). Consequently, the phylogenetic relationships among taxa and the definition of species are still largely dependant on morphological characters of uncertain value, which causes confusion in the phylogeny, taxonomy and the debate on cosmopolitanism of free-living protists. Here we present a SSU rRNA-based phylogeny of the Hyalospheniidae including the most common species. Similar to the filose testate amoebae of the order Euglyphida the most basal clades have a terminal aperture; the ventral position of the pseudostome appears to be a derived character. Family Hyalospheniidae appears paraphyletic and is separated into three clades: (1) Heleopera sphagni, (2) Heleopera rosea and Argynnia dentistoma and (3) the rest of the species from genera Apodera, Hyalosphenia, Porosia and Nebela. Our data support the validity of morphological characters used to define species among the Hyalospheniidae and even suggest that taxa described as varieties may deserve the rank of species (e.g. N. penardiana var. minor). Finally our results suggest that the genera Hyalosphenia and Nebela are paraphyletic, and that Porosia bigibbosa branches inside the main Nebela clade.     

Ordovician (Caradoc) Gastropoda of the Katkoyeh Formation, Kerman Province, Iran

Seven taxa of gastropods are described from the Ordovician (Caradoc) upper Katkoyeh Formation of the Kerman region, east-central Iran. Three are named species and four are under open nomenclature. The most abundant is the minute bellerophontiform Tritonophon peeli Horný, 1977, indicative of a shallow water plectonotid community. Shell material is not preserved in this species, but a small, simple bulbous protoconch is present. Two new species are Slehoferia pachyta and Nonorios kleistos, both with prominent thick-shelled conchs. Shell repair is documented in both of these forms, interpreted as a result of failed predation. Other micromorphic species occur in the samples, but only Tropidodiscus sp. and Nonorios? sp. are named. The Kerman assemblage is comparable directly with similar, closely coeval faunas in Bohemia, France, Portugal, Morocco, and Italy, corresponding with the Palaeozoic Mediterranean Province of northern peri-Gondwana.     

Global diversity of caddisflies (Trichoptera: Insecta) in freshwater

The not yet uploaded Trichoptera World Checklist (TWC) [], as at July 2006, recorded 12,627 species, 610 genera and 46 families of extant and in addition 488 species, 78 genera and 7 families of fossil Trichoptera. An analysis of the 2001 TWC list of present-day Trichoptera diversity at species, generic/subgeneric and family level along the selected Afrotropical, Neotropical, Australian, Oriental, Nearctic and Palaearctic (as a unit or assessed as Eastern and Western) regions reveals uneven distribution patterns. The Oriental and Neotropical are the two most species diverse with 47–77% of the species in widespread genera being recorded in these two regions. Five Trichoptera families comprise 55% of the world’s species and 19 families contain fewer than 30 species per family. Ten out of 620 genera contain 29% of the world’s known species. Considerable underestimates of Trichoptera diversity for certain regions are recognised. Historical processes in Trichoptera evolution dating back to the middle and late Triassic reveal that the major phylogenetic differentiation in Trichoptera had occurred during the Jurrasic and early Cretaceous. The breakup of Gondwana in the Cretaceous led to further isolation and diversification of Trichoptera. High species endemism is noted to be in tropical or mountainous regions correlated with humid or high rainfall conditions. Repetitive patterns of shared taxa between biogeographical regions suggest possible centres of origin, vicariant events or distribution routes. Related taxa associations between different regions suggest that an alternative biogeographical map reflecting Trichoptera distribution patterns different from the Wallace (The Geographical Distribution of Animals: With a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth’s Surface, Vol. 1, 503 pp., Vol. 2, 607 pp., Macmillan, London, 1876) proposed biogeography patterns should be considered. Anthropogenic development threatens biodiversity and the value of Trichoptera as important functional components of aquatic ecosystems, indicator species of deteriorating conditions and custodians of environmental protection are realised.     

Benthic hydroids (Cnidaria: Hydrozoa) from the Bransfield Strait area (Antarctica) collected by Brazilian expeditions,with the description of a new species

A total of 36 species of benthic hydroids, belonging to nine families and 16 genera, were found in the hydroid collection gathered during the Brazilian Antarctic expeditions PROANTAR III and IV. Seven of the species were identified only to generic level. There is a clear dominance of the subclass Leptothecatae with 33 species. By far the most diversified family was the Sertulariidae, with 16 species (44%). Symplectoscyphus with eight species, including Symplectoscyphus magnificus sp. nov., is the most diversified genus. Almost 70% of the species diversity is restricted to just six genera (38%). Sixty-eight percent of the species is Antarctic endemics and 86% is restricted to Antarctic or Antarctic/sub-Antarctic waters. Eudendrium antarcticum and Amphisbetia operculata are recorded for the first time from Antarctic waters.     

Comparative gas exchange performance during the wet season of three Brazilian Styrax species under habitat conditions of cerrado vegetation types differing in soil water availability and crown density

We studied the influence of environmental conditions in different vegetation types of the Brazilian savanna (cerrado s.l.) on CO₂ assimilation and water use efficiency of Styrax ferrugineus, S. camporum and S. pohlii, which are morphologically similar but have distinct distribution patterns. We measured leaf gas exchange and water relations in adult and young plants. Adult plants of S. ferrugineus were measured in the cerrado s. str.; plants of S. camporum at the edge of cerradão, a woodland vegetation; and in a riparian forest those of S. pohlii. Eight-month-old young plants were planted in the cerrado s. str., at the edge and in the understory of the cerradão, and in the understory of the riparian forest. For young plants, the high light availability in the cerrado s. str. resulted in a threefold greater CO₂ assimilation rate (A) compared to the other sites. A of adult plants under full irradiation (1800 μmol photons m⁻² s⁻¹) was significantly lower in S. pohlii compared to S. ferrugineus and S. camporum. Although sufficient soil water was available at every site at the end of the wet season, the extreme high water content in the riparian forest soil led to increased stomatal conductance (gs) and transpiration rate (E) for young plants of S. camporum and S. pohlii, but not of S. ferrugineus. However, the large gs in S. pohlii and S. camporum young plants did not result in higher A rates. The differences may indicate some specialization of S. ferrugineus to best growth in open savanna and of S. pohlii in riparian forest habitats. The high water use efficiency of S. camporum young plants in the understory of cerradão seems not yet sufficient to emphasize that this species benefits from this particular habitat. Because, while water supply during the wet season is quite sufficient, soil water deficits and frequent fires can occur during the dry season and might modify strongly the habitat conditions for the three Styrax species.     

Host specificity of a Brazilian mistletoe, Struthanthus aff. polyanthus (Loranthaceae), in cerrado tropical savanna

The aim of this study was to determine if the mistletoe Struthanthus aff. polyanthus shows host specificity, and if host abundance and twig architecture influence this specificity in Cerrado (senso strictu). An area of 1.3 ha of natural savanna was sampled for the occurrence of the mistletoe. The twigs of the hosts were classified as vertical or horizontal, and the bark as smooth or rough. We sampled a total of 666 trees in the study site and 118 individuals (17.7%) hosted S. aff. polyanthus. The abundance of mistletoe was significantly affected by the bark type, but not by the twigs inclination. In the cerrado, S. aff. polyanthus seems to be a generalist, with a tendency to be more common on some hosts (Kielmeyera coriacea, Pouteria ramiflora and Styrax ferrugineus), prefering that with rough bark type.     

SSU rRNA phylogeny of Arcellinida (Amoebozoa) reveals that the largest Arcellinid genus, Difflugia Leclerc 1815, is not monophyletic

The systematics of lobose testate amoebae (Arcellinida), a diverse group of shelled free-living unicellular eukaryotes, is still mostly based on morphological criteria such as shell shape and composition. Few molecular phylogenetic studies have been performed on these organisms to date, and their phylogeny suffers from typical under-sampling artefacts, resulting in a still mostly unresolved tree. In order to clarify the phylogenetic relationships among arcellinid testate amoebae at the inter-generic and inter-specific level, and to evaluate the validity of the criteria used for taxonomy, we amplified and sequenced the SSU rRNA gene of nine taxa - Difflugia bacillariarum, D. hiraethogii, D. acuminata, D. lanceolata, D. achlora, Bullinularia gracilis, Netzelia oviformis, Physochila griseola and Cryptodifflugia oviformis. Our results, combined with existing data demonstrate the following: 1) Most arcellinids are divided into two major clades, 2) the genus Difflugia is not monophyletic, and the genera Netzelia and Arcella are closely related, and 3) Cryptodifflugia branches at the base of the Arcellinida clade. These results contradict the traditional taxonomy based on shell composition, and emphasize the importance of general shell shape in the taxonomy of arcellinid testate amoebae.     

Global diversity of amphipods (Amphipoda; Crustacea) in freshwater

Amphipods are brooding peracaridan crustaceans whose young undergo direct development, with no independent larval dispersal stage. Most species are epibenthic, benthic, or subterranean. There are some 1,870 amphipod species and subspecies recognized from fresh or inland waters worldwide at the end of 2005. This accounts for 20% of the total known amphipod diversity. The actual diversity may still be several-fold. Amphipods are most abundant in cool and temperate environments; they are particularly diversified in subterranean environments and in running waters (fragmented habitats), and in temperate ancient lakes, but are notably rare in the tropics. Of the described freshwater taxa 70% are Palearctic, 13% Nearctic, 7% Neotropical, 6% Australasian and 3% Afrotropical. Approximately 45% of the taxa are subterranean; subterranean diversity is highest in the karst landscapes of Central and Southern Europe (e.g., Niphargidae), North America (Crangonyctidae), and Australia (Paramelitidae). The majority of Palearctic epigean amphipods are in the superfamily Gammaroidea, whereas talitroid amphipods (Hyalella) account for all Neotropic and much of the Nearctic epigean fauna. Major concentrations of endemic species diversity occur in Southern Europe, Lake Baikal, the Ponto-Caspian basin, Southern Australia (including Tasmania), and the south-eastern USA. Endemic family diversity is similarly centered in the Western Palearctic and Lake Baikal. Freshwater amphipods are greatly polyphyletic, continental invasions have taken place repeatedly in different time frames and regions of the world. In the recent decades, human mediated invasions of Ponto-Caspian amphipods have had great impacts on European fluvial ecosystems. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Species diversity,endemism and conservation of the family Caryophyllaceae in Greece

The family Caryophyllaceae includes world-wide 86 genera and approximately 2100 species. Greece is one of its most important centres of diversity and endemism. A total of 428 Caryophyllaceae taxa are distributed in Greece, 161 of them being endemic to the Greek political territory. The endemic element represents approximately 5% of the global diversity of the family at the species level. The aim of this paper is to discuss the distribution patterns of the Greek endemic Caryophyllaceae, as well as those with a limited distribution range to the neighbouring areas of the Balkans and Anatolia, on the basis of the phytogeographical regions of Greece in order to identify the important regions for their conservation. The majority of the Greek endemic Caryophyllaceae (64.6%) are distributed in only one single phytogeographical region, or even a smaller area indicating the extremely restricted distribution ranges of the endemic plants in Greece. Actually 83 Greek endemic Caryophyllaceae can be grouped on cytotaxonomic criteria. Most of them belong to the category of schizoendemics (91.6%), indicating that the endemism of Caryophyllaceae in Greece has mainly originated in an active way. Cluster analysis has been used to classify the phytogeographical regions according to their floristic similarities. Two iterative complementarity methods were used to evaluate the importance of each phytogeographical region in the conservation of the endemic Caryophyllaceae in Greece. Peloponnisos, Kriti-Karpathos and Sterea Ellas are the most important phytogeographical regions in this respect, followed by North Central and North East. When adding the Balkan-Aegean-Anatolian endemics to the analysis, Peloponnisos, North Central, Kriti-Karpathos, North East and Eastern Aegean result as the most important areas. In every case, an elevated number of sites are required for the conservation of Caryophyllaceae in Greece, reflecting the great dissimilarities in the floristic composition of the various phytogeographical regions. The results provided by the different methods are compared. A catalogue of the Greek endemic Caryophyllaceae is appended.     

Patterns of diversity,endemism and conservation: an example with Mexican species of Ternstroemiaceae Mirb. ex DC. (Tricolpates: Ericales)

The Ternstroemiaceae is a family of vascular plants with many taxonomic problems, basically in the delimitation of some of its species. Little is known about its world distribution and about the real state of conservation of its species. An analysis of the distribution patterns of the Mexican species was undertaken to provide information about the present status of species and possible conservation areas. Also, an analysis of the species richness, endemism centres, extension and location of the areas occupied by these species with a 1:50000 chart index was made. Finally, we propose some of the species of this family to be included in the IUCN Red List of Threatened Species and in the next Mexican Norma Oficial Mexicana 059.