Organisms Diversity & Evolution

Published by Springer Verlag
Online ISSN: 1439-6092
Publications
Article
Acoels are among the simplest worms and therefore have often been pivotal in discussions of the origin of the Bilateria. Initially thought primitive because of their "planula-like" morphology, including their lumenless digestive system, they were subsequently dismissed by many morphologists as a specialized clade of the Platyhelminthes. However, since molecular phylogenies placed them outside the Platyhelminthes and outside all other phyla at the base of the Bilateria, they became the focus of renewed debate and research. We review what is currently known of acoels, including information regarding their morphology, development, systematics, and phylogenetic relationships, and put some of these topics in a historical perspective to show how the application of new methods contributed to the progress in understanding these animals. Taking all available data into consideration, clear-cut conclusions cannot be made; however, in our view it becomes successively clearer that acoelomorphs are a "basal" but "divergent" branch of the Bilateria.
 
Maximum likelihood phylogram (unrooted), showing branch lengths (–ln likelihood=4275.79863, proportion of invariant sites=0.3491, gamma shape parameter=0.4952); support values above nodes are Bayesian posterior probabil- ities.  
Article
The phylogenetic relationships of microhylid frogs are poorly understood. The first molecular phylogeny for continental African microhylids is presented, including representatives of all subfamilies, six of the eight genera, and the enigmatic hemisotid Hemisus. Mitochondrial 12S and 16S rRNA sequence data were analysed using parsimony, likelihood and Bayesian methods. Analyses of the data are consistent with the monophyly of all sampled subfamilies and genera. Hemisus does not nest within either brevicipitines or non-brevicipitines. It is possibly the sister group to brevicipitines, in which case brevicipitines might not be microhylids. Phrynomantis and Hoplophryne potentially group with non-African, non-brevicipitine microhylids, in agreement with recent morphological and molecular data. Within brevicipitines, Breviceps is recovered as the sister group to a clade of Callulina+Spelaeophryne+Probreviceps. The relationships among the genera within this latter clade are unclear, being sensitive to the method of analysis. Optimal trees suggest the Probreviceps macrodactylus subspecies complex might be paraphyletic with respect to P. uluguruensis, corroborating preliminary morphological studies indicating that P. m. rungwensis may be a distinct species. P. m. loveridgei may be paraphyletic with respect to P. m. macrodactylus, though this is not strongly supported. Some biogeographic hypotheses are examined in light of these findings.
 
Article
The European marine fauna used to be considered to include 16 species of Discodoris sea slugs until a recent worldwide revision demonstrated that there is not a single Discodoris species in European waters. This exemplary case illustrates the fact that species checklists do not accurately represent biodiversity unless they are based on sound taxonomic work in which (1) the status of every available species name has been addressed, i.e. whether it is valid, synonymous, or of doubtful application, and (2) classification reflects phylogenetic relationships. It is argued that taxonomic revisions are critically needed, because the status of species names can only be addressed properly through revisions. It is discussed that fields which depend on taxonomic data, such as conservation biology and ecology, might be affected deeply if problematic species names (synonyms and nomina dubia) have not been recognized. Consequently, it is proposed that a taxon that has not been revised be red-flagged in checklists, so that non-taxonomists will know which species names should be applied with caution or not at all. KeywordsBiodiversity–Taxonomy–Nomenclature–Revision–Monograph
 
Article
Two new species of Ligia are described, L. persica sp. nov. from the Persian Gulf and L. yemenica sp. nov. from the Gulf of Aden. Ligia persica occurs along the northern coasts of the Persian Gulf and around some Iranian islands such as Qeshm and Kish. A comparison of SEM micrographs shows that the shape and ornamentation of distal parts of the appendix masculina are reliable characters for the identification of morphologically similar Ligia species. They are species-specific and of great importance in the taxonomy of the genus. KeywordsIsopoda- Ligia persica sp. nov.- Ligia yemenica sp. nov.-SEM-Persian Gulf-Gulf of Aden
 
Article
Sabellastarte Krøyer, 1856 (Sabellidae), a morphologically homogeneous group distributed in warm and temperate coasts of the Indo-Pacific and Caribbean Sea, is characterized by the presence of a unique combination of features. To date, the genus comprises eight species, but morphological characters traditionally used in diagnostics have shown intra-specific variability, making species boundaries and distributions unclear. The present study constitutes the first attempt to test the monophyly of Sabellastarte and its relationships to other sabellid genera by combining molecular (COI and 16S) and morphological data. Results include placement of a clade containing Stylomma, Sabella, Branchiomma and Bispira as the sister group to Sabellastarte. Phylogenetic analyses and genetic divergence among specimens from several localities around the world indicate the presence of at least six lineages within Sabellastarte. In the context of a discussion of species boundaries and diagnostic features, the distribution of some of those lineages can be explained by the presence of cryptic species and potential introductions. Keywords Sabellastarte -Sabellidae-Annelida-Integrative taxonomy-Morphology-Mitochondrial DNA
 
Box-and-whisker plot of intraspecific and intrageneric (x-axis) genetic distances measured in uncorrected p-distances (y-axis). Thick black lines indicate median values, boxes represent upper and lower quartiles, whiskers indicate value ranges, circles represent outliers
Tested primer sequences for 18S amplificates (M13 tails shown in italics; → =forward, ← =reverse)
Uncorrected p-distances given as average, minimum and maximum values; n = number of sequences (per strain) or number of individuals (per species/genus), respectively
(continued)
Article
Diatoms are present in all types of water bodies and their species diversity is influenced greatly by environmental conditions. This means that diatom occurrence and abundances are suitable indicators of water quality. Furthermore, continuous screening of algal biodiversity can provide information about diversity changes in ecosystems. Thus, diatoms represent a desirable group for which to develop an easy to use, quick, efficient, and standardised organism identification tool to serve routine water quality assessments. Because conventional morphological identification of diatoms demands specialised in-depth knowledge, we have established standard laboratory procedures for DNA barcoding in diatoms. We (1) identified a short segment (about 400bp) of the SSU (18S) rRNA gene which is applicable for the identification of diatom taxa, and (2) elaborated a routine protocol including standard primers for this group of microalgae. To test the universality of the primer binding sites and the discriminatory power of the proposed barcode region, 123 taxa, representing limnic diatom diversity, were included in the study and identified at species level. The effectiveness of the barcode was also scrutinised within a closely related species group, namely the Sellaphora pupula taxon complex and relatives. Keywords18S (SSU) rRNA gene–Bacillariophyta–DNA barcoding–Diatoms–Standard laboratory procedure
 
Article
AbstractThe deep-sea genusStorthyngura (family Munnopsididae) is revised. Three new genera (Rectisura, Sursumura and Vanhoeffenella) and four new species (Rectisura richardsoniae, Sursumura aberrata, Vanhoeffenella georgei, and V. moskalevi) are described from deep-sea basins and trenches of the South Atlantic Ocean. Thirty-three previously described species are placed in new combinations with one of the new genera. Diagnoses are presented for the subfamily Storthyngurinae Kussakin (2003), the genus Storthyngura VanhÏffen, and all new taxa. The accompanying Electronic Supplement offers a key to the six genera in the subfamily, and keys to species inStorthyngura and the three new genera.
 
Article
AbstractThe parabasalids include parasites (e.g. trichomonads) as well as many hypermastigid flagellates which live in termites and other wood-eating insects and contribute to the cellulose-digesting capacity of those animals. A hypermastigid, Joenina pulchella Grassi, is shown to have a “flagellar area” composed of 1300 flagella, including three privileged basal bodies which have homologues in the trichomonads. The cytoskeleton includes preaxostylar fibres, two parabasal fibres and two atractophores with the parabasal fibres subdividing to form many parabasals. The microtubular rows of the pelta-axostyle system surround the flagellar area and converge towards a multispiralled axostylar trunk. On the basis of similarities of ultrastructure, joeniids and devescovinids are argued to be members of the same clade. Projoenia Lavette is in the sister group to Devescovina Foa and gives rise to the series Placojoenia Radek, Joenia Grassi, Joenina Grassi. Projoenia has a “flagellar area” as in the joeniids, but also a recurrent flagellum with a paraxonemal fibre and a cresta as does Devescovina. Projoenia has a parabasal fibre twisted around the axostyle, as well as a multispiralled axostyle. In Placojoenia, Joenia and Joenina the recurrent flagellum is absent or reduced to the basal body as is the cresta; the parabasal apparatus becomes multibranched. The classical Hypermastigida is in need of major revision. Parabasalids such as Lophomonadidae, Joeniidae, Deltotrichonymphidae, and possibly Rhizonymphidae and Kofoidiidae, collectively the lophomonads, have conserved the trichomonad/devescovinid organization and have a trichomonad-like morphogenesis involving only the privileged basal bodies and attached fibres. They can be distinguished from the rest of hypermastigids and should be classified with the Devescovinidae and Calonymphidae in a large clade — the Cristamonadida (new order). The remaining hypermastigids (the Trichonymphina and Spirotrichonymphina) have a rostrum which separates in two hemi-rostra at division, form a sister group to all other parabasalids and are not closely related to the remainder of the hypermastigids.
 
Article
AbstractThe geographic variation of the circum-Mediterranean scorpion species Euscorpius carpathicus (L.) was traditionally analysed using morphological characters such as trichobothrial patterns, which resulted in the recognition of 23 subspecies; however, the biological reality of these subspecies remains unclear. Here, we focus on populations from the western Mediterranean and provide new molecular evidence that those from the island of Mallorca (Balearic Islands, Spain) represent a highly divergent lineage separate from E. carpathicusfrom the mainland of France (Vaucluse) and Italy (Liguria and Piemonte). This divergence is evidenced by morphological analysis. Moreover, allozyme and mtDNA divergences (about 10%) agree with our hypothesis that the Balearic island populations became isolated from the mainland about 5 Ma BP since the refilling of the Mediterranean Basin and have to be considered autochthonous. This hypothesis is additionally supported by the comparison of the genetic differentiation between artificially transplanted island populations and mainland populations in the congeneric species E. flavicaudis(de Geer). The phylogenetic species concept (PSC) is applied to elevate the subspecies E. carpathicus balearicus Caporiacco, 1950 to species rank. A lectotype is designated for this species.
 
Article
AbstractThe phylogenetic position of Corambidae has been subject to much speculation. Most of the confusion has arisen from insufficient anatomical knowledge and from interpretations which have not followed the rules of Hennigian phylogenetic systematics. In this study, as a model system, the poorly known Chilean species Corambe lucea Marcus, 1959 is redescribed in detail: digestive, reproductive, central nervous, circulatory, and excretory systems are examined anatomically and histologically. New biological data are also given. After critical comparison with congeners, C. lucea is confirmed to be a valid species. Major organ systems of C. lucea are discussed comparatively and used to resolve phylogenetic relationships. Gill structure and circulatory system of the Corambidae are homologous to that of the Anthobranchia bauplan but not to the secondary gills and circulatory system of the Phyllidiidae. The similar lateral position of respiratory organs and a posteroventral anus in some Corambidae and Phyllidiidae is clearly due to convergence; the two groups are true doridoidean nudibranchs but there is no indication for a sistergroup relationship. Corambids are shown to belong to the monophyletic group Suctoria which is characterized by the unique and complex autapomorphies “possession of a dorsal buccal pump” and “possession of a large first lateral tooth with a long, denticulated hook”.
 
Article
Rich samples of benthic Peracarida collected from abyssal plains of the Angola Basin (South Atlantic) contained several new and rare asellote isopods. In this contribution, Eurycope tumidicarpus n. sp. is described from well-preserved specimens. This species is remarkable because the second pereopod has an enlarged carpus, and pereopod 4 is shaped like the posterior swimming legs. New specimens of Acanthocope galathea Wolff, 1962 (Fig. 1a) are used for a redescription, hitherto unknown characters of the male are shown.
 
Article
In the course of the DIVA 1 expedition (RV “Meteor”, cruise M48/1) to the Angola Basin (South Atlantic) two myriotrochid holothurians were collected at abyssal depths – Neolepidotrochus parvidiscus angolensis subsp. nov. and Siniotrochus myriodontus Gage and Billett, 1986. The latter, until now, was known only from few specimens from the north-eastern Atlantic Ocean. This new record of S. myriodontus for the South Atlantic Ocean indicates a wide geographical distribution. Both species are described in detail, including parameters for the wheel deposits.
 
Hyalogyrina depressa, histological details of genital system (semithin sections). (a) Overview: longitudinal section of gonoducts and associated structures, anterior is at right. (b) Detail: longitudinal section of proximal part of gonoduct. (c) Cross section of receptaculum seminis. (d) Longitudinal section of vesicula seminalis with autosperm. (e) Longitudinal section of bursa copulatrix with degraded allosperm. (f) Cross section of ovotestis lobes, each with inner ovary and outer testis portion. (g) Detail of f: spermatogenesis in peripheral portion of ovotestis. Labels: b = bursa, bs1/2 = blind sac1/2, dg = digestive gland, fs = allosperm, gd1/2/3 = proximal/median/distal portion of gonoduct, hd = hermaphroditic duct, n/n' = nucleus of ripe/ juvenile egg, nl = nucleolus, re = rectum, rs = receptaculum seminis, s = autosperm, sc = spermatocyte, spg = spermatogonium, x = degraded allosperm, yg = yolk granules
Article
Whereas Hyalogyrina Marshall, 1988 was originally considered a skeneid vetigastropod, the family Hyalogyrinidae Warén & Bouchet, 1993 has later been classified as basal Heterobranchia despite their rhipidoglossate radula. In order to evaluate this placement and to shed more light on the origin of all higher Gastropoda, we investigated five representatives of all three nominal hyalogyrinid genera by means of semithin serial sectioning and computer-aided 3D reconstruction of the respective anatomy, which we present in an interactive way. In general the morphological features (shell, external morphology, anatomy) fully confirm the placement of Hyalogyrinidae in the Heterobranchia, but in particular the conditions of the genital system vary substantially within the family. The ectobranch gill of Hyalogyrinidae is shared with Valvatidae, Cornirostridae, and Xylodisculidae; consequently all these families are united in Ectobranchia Fischer, 1884. The rhipidoglossate hyalogyrinid radula suggests independent acquisition of taenioglossate radulae in the Caenogastropoda and other Ectobranchia. Therefore, the origin of the Heterobranchia—and thus of all higher gastropods—looks to have taken place already on the rhipidoglossate, i.e. the ‘archaeogastropod’, level of evolution. Ectobranchia are considered the first extant offshoot of the Heterobranchia; implications for the stem species of the latter are outlined. KeywordsGastropoda–Ectobranchia–Hyalogyrinidae–Interactive 3D anatomy–Systematics–Phylogeny–Heterobranchia
 
Article
The well-preserved histology of the geologically oldest sauropod dinosaur from the Late Triassic allows new insights into the timing and mechanism of the evolution of the gigantic body size of the sauropod dinosaurs. The oldest sauropods were already very large and show the same long-bone histology, laminar fibro-lamellar bone lacking growth marks, as the well-known Jurassic sauropods. This bone histology is unequivocal evidence for very fast growth. Our histologic study of growth series of the Norian Plateosaurus indicates that the sauropod sistergroup, the Late Triassic and early Jurassic Prosauropoda, reached a much more modest body size in a not much shorter ontogeny. Increase in growth rate compared to the ancestor (acceleration) is thus the underlying process in the phylogenetic size increase of sauropods. Compared to all other dinosaur lineages, sauropods were not only much larger but evolved very large body size much faster. The prerequisite for this increase in growth rate must have been a considerable increase in metabolic rate, and we speculate that a bird-like lung was important in this regard.
 
Linear morphometric traits obtained for Ephedrus persicae; see also Fig. 1
Article
We analysed linear measurements on various parts of the body and the configuration of 11 landmarks on the wing in a large sample of Ephedrus persicae that had emerged from 13 aphid host species, to assess whether static allometry (a measure of the scaling relationship between traits in a population of individuals at the same ontogenetic stage) accounts for variation in body shape. The analysed specimens came from several localities in Europe, Asia Minor, Japan and South America, and cover a large portion of the distribution area of E. persicae. We found that allometry accounts for variation in body shape among different biotypes within the E. persicae group. The allometric slopes for head size (HD), petiolus width (PETW), mesoscutum width (MSC), and ovipositor sheath length (OVPL) diverged significantly among biotypes, indicating biotype-specific allometries. The analysis of allometric variation in wing shape showed that the pattern and direction of allometric changes also differed among individuals that had emerged from different hosts. Our results (observed divergences in the directions of allometric slopes of particular morphometric traits and wing shape) suggest that allometric relations within E. persicae are not conserved, so that allometry itself changes, evolving differently in aphid parasitoids that emerge from different hosts. KeywordsAllometry-Morphometric variability-Geometric morphometrics- Ephedrus persicae
 
Article
There are few reliable diagnostic morphological characters for species of the asteracean genus Microseris, and quantitative differences in the shapes of the achenes and the paleaceous pappus parts play a decisive role in species recognition. The genetic basis of species and strain differences in various characters has been studied previously, but little is known about quantitative characters of the achenes.We performed a quantitative trait locus (QTL) analysis in the F2 of an interspecific cross between Microseris douglasii and M. bigelovii for achene length, achene diameter, achene shape, palea length, awn length, and achene pigmentation.Independent inheritance of the main heritable achene characters – achene length, palea length and achene pigmentation – was revealed by the detection of distinct and specific QTLs for these characters. For palea length five QTLs with about equal phenotypic effects were mapped on four different linkage groups. Achene length and achene shape (achene length / achene diameter) were determined by two different genetic systems with one major gene and two modifiers. The detection of QTLs with a polarity of the effects opposite to that in the parental strains for achene length and achene shape reveals genetic variation for a potential increase in species differences. For the highly heritable trait, achene pigmentation, the bimodal F2 distribution suggested single-factor inheritance for absence versus presence of spots, with dominance for the spotted condition. However, only relatively weak QTL effects on that trait could be detected. Additional molecular markers (RAPDs, AFLPs) have to be tested for cosegregation with that major gene. The results are discussed in the context of different theories for the evolution of morphological characters.
 
Courtship songs of Stenobothrus cotticus from Col d'Izoard in France (a-c) and from the Rila Mts. in Bulgaria (d). a Overview of a whole courtship sequence. b Section of 4 s duration. c Single syllable
Tree from neighbour-joining analysis of selected European Stenobothrus species based on Kimura-2-parameter distances among 1,000 bp sequences of the mitochondrial gene cytochrome oxidase, subunit 1 (co1). Chorthippus parallelus used as outgroup. Bootstrap support values≥50% among 1,000 bootstrap replicates given above branches
Current distribution of Stenobothrus cotticus ( white triangles ) and S. rubicundulus ( open circles ) in Europe; S. rubicundulus further ranges to western Ukraine (not shown). For localities and corresponding references, see “ Electronic Supplementary Material ” 
Article
A warming climate leads to shifts in distribution ranges to higher latitudes and altitudes. Consequently, cold-adapted alpine species can be trapped in interglacial Holocene refugia on high mountain summits if they fail to expand their ranges to the north. One example is the alpine grasshopper Stenobothrus cotticus. This species was assumed to be endemic to the southwestern Alps (France, Italy). However, we have found a second refugium in the Rila Mountains in southwestern Bulgaria. Analyses of the mitochondrial gene co1 and of phenotypic characters from morphology and behaviour did not reveal differences between the two geographically separated populations of S. cotticus studied. We suppose that S. cotticus had a wider distribution during colder periods, when its range was expanded to lower altitudes. This hypothesis is supported by the current distribution of the closely related montane S. rubicundulus. KeywordsDistribution-Glacial expansion-Interglacial refugia- Stenobothrus cotticus - Stenobothrus rubicundulus
 
Article
Gravel Bank Grasshopper (Chorthippus pullus) populations inhabit two contrasting environments, pebbly gravel banks with scarce vegetation cover in mountainous areas along the Alps and lowland grasslands dominated by Common Heather (Calluna vulgaris). Heath populations of C. pullus have been rediscovered only recently, and show a distribution scattered across Central Europe. The wings are reduced in this species; thus, it has low potential for long-distance dispersal. We used sequence data on a newly developed non-coding nuclear marker from three gravel-bank and four heath populations to test whether grasshoppers from the two environments represent distinct lineages. Gravel-bank populations were studied in southern Germany (Bavaria), heath populations in eastern Germany (Brandenburg and Saxony) and Ukraine. We compared those genetic data with an analysis of variation in a suite of morphometric traits. Finally, we combined genetic and morphometric data to reconstruct a plausible scenario for the ecological shift observed in C. pullus. Our newly developed marker did not sort populations from contrasting environments in two monophyletic lineages. Nevertheless, we found a general lack of gene flow between the gravel-bank and heath populations. There was pronounced variation among populations in morphometric traits. That variation was partially partitioned by habitat type, and populations from the same habitat tended to be more similar than those from different habitats. Our data suggest that heath populations originated through northward expansion from multiple southern European refugia, and that the gravel-bank populations represent one of these sources. Patterns of genetic and morphometric divergence suggest that gravel-bank and heath populations may be in the process of incipient speciation. Keywords Chorthippus pullus -Genetic divergence-Morphometrics-Nuclear DNA-Biogeography
 
Article
We analyzed the patterns of nucleotide sequence variation at three mitochondrial DNA loci, the noncoding mitochondrial control region and two genes (cytochrome b and cytochrome oxidase I) of Gavia immer in the largest collection of wintering individuals from Southern Europe to date. The sample consisted of 33 birds, oiled during the 2002/2003 Prestige tanker spill and washed ashore on the Galician coast (NW Iberian Peninsula). The aims of the study were to investigate the levels of standing genetic variation in the species, and to identify the geographic origin of these wintering birds. To do this, all available sequences of these loci, mostly from North American specimens collected from both the Atlantic and Pacific coasts, were retrieved from GenBank and included in the analysis. Overall, only 14 genetic variants were detected in the nearly 2Kb surveyed, which reflects very low levels of nucleotide site diversity in this species. Interestingly, all variants were found at very low frequencies, and there was no indication of any clear subdivision in the G. immer population. This genetic profile is consistent with G. immer being a single panmictic population of small effective population size as compared with other seabirds. These circumstances preclude identification of the breeding regions of these wintering birds relying solely on genetic data. In the light of these results, possible causes, and the genetic and ecological consequences, of this demographic scenario are discussed. Keywords Gavia immer –Mitochondrial DNA–Genetic diversity–Population genetics–Panmixia
 
Article
A skull of a new species of mousebird (Aves: Coliiformes) is described from the Middle Eocene of Messel in Germany. Chascacocolius cacicirostris n. sp. is the fifth coliiform species described from the Messel deposits, and a further example of the remarkable similarity between the early Eocene avifaunas of North America and Europe. As for its much smaller North American counterpart, C. oscitans Houde & Olson, 1992, the new species has greatly elongated retroarticular processes on the mandible, which are an adaptation to gaping, i.e. opening of the bill in a substrate. The cranium and upper beak of Chascacocolius are seen for the first time in the new specimen; the latter shows a striking resemblance to the upper beak of some modern New World Blackbirds (Passeriformes: Icteridae). C. cacicirostris n. sp. thus provides another example of the diversification of early Tertiary Coliiformes, some taxa of which apparently occupied feeding niches that today are filled by songbirds (Passeriformes).
 
Article
In addition to other potential causes, immigration into locally adapted populations has been suggested to maintain the genetic variance in fitness that is necessary for the good-genes hypothesis. Using population-genetic simulations, the present contribution shows that co-occurring local adaptation and migration can maintain genetic variance in fitness. In combination with an effect of local adaptation on condition and condition-dependent sexual signaling, such a scenario therefore enables the evolution and maintenance of female choice for locally adapted males. The simulations show that this mechanism can also work when choice is costly, and that the potential benefit is similar to that in other good-genes mechanisms. As a consequence of female choice in favor of locally adapted males, differentiation between populations can be expected to increase due to the decreased effective gene flow between populations. Based on such effects, choice of locally adapted males has the potential to play an important role in speciation and adaptive radiation.
 
Article
Coral reefs are renowned as complex ecosystems with an extremely large biodiversity. Parasite-host relationships contribute substantially to this, but are poorly known. We describe the results of a study in which approximately 60,000 corals were searched for parasitic Leptoconchus snails (Gastropoda: Coralliophilidae) in Indo-West Pacific waters of Egypt, the Maldives, Thailand, Palau and Indonesia. We discovered an adaptive radiation of 14 snail species, each of which lives in species-specific association with one or more of 24 mushroom coral species. The 14 snail species are described as new to science under the names Leptoconchus inactiniformis sp. nov., L. inalbechi sp. nov., L. incrassa sp. nov., L. incycloseris sp. nov., L. infungites sp. nov., L. ingrandifungi sp. nov., L. ingranulosa sp. nov., L. inlimax sp. nov., L. inpileus sp. nov., L. inpleuractis sp. nov., L. inscruposa sp. nov., L. inscutaria sp. nov., L. intalpina sp. nov., and L. massini sp. nov. Their separation is based on indisputable molecular differences, whereas the rudimentary shell characters or impoverished anatomical details do not allow identification. The coral hosts also serve to distinguish the snail species, as none of the former was found to contain more than one of the latter. The complexity of coral reefs is still underrated, as is shown here by the application of DNA taxonomy as an indispensable approach to unravel cryptic radiations, which must be known in order to understand the functioning of the ecosystem. KeywordsParasitic snails–Coral reefs–Coralliophilidae–Fungiidae–Indo-Pacific–DNA barcoding
 
Ventral nerve cord fusion in representatives of Syrphidae (A, B) and Bombyliidae (C, D). For each family, two species with widely divergent body shape have been chosen. In both cases, the familyspecific arrangement of ganglia is preserved (refer to Fig. 1). A, B are shown at the same scale, different from the scale for C, D. Body shape is drawn to scale. The nerves and ganglia are drawn schematically, although the positions of ganglia within the body are drawn to scale.
Article
Insect ganglia are often composed of fused segmental units or neuromeres. We estimated the evolution of the ventral nerve cord (VNC) in higher Diptera by comparing the patterns of neuromere fusion among 33 families of the Brachycera. Variation within families is uncommon, and VNC architecture does not appear to be influenced by body shape. The outgroup pattern, seen in lower Diptera, is fusion of neuromeres belonging to thoracic segments 1 and 2 (T1 and T2), and fusion of neuromeres derived from T3 and abdominal segment 1 (A1). In the abdomen, neuromeres A7–10 are fused into the terminal abdominal ganglion (TAG). Increased neuromere fusion is a feature of the Brachycera. No brachyceran shows less fusion than the outgroups. We established six pattern elements: (1) fusion of T1 and T2, (2) fusion of T3 and A1, (3) fusion of the T1/T2 and T3/A1 ganglia, (4) increase in the number of neuromeres comprising the TAG, (5) anteriorward fusion of abdominal neuromeres, and (6) the complete fusion of thoracic and abdominal neuromeres into a synganglion. States 1 and 2 are present in the outgroup lower Diptera, and state 3 in the Xylophagomorpha, Stratiomyomorpha, Tabanomorpha and Cyclorrhapha. State 4 is a feature of all Eremoneura. State 5 is present in Cyclorrhapha only, and state 6, fusion into a synganglion, has evolved at least 4 times in the Eremoneura. Synapomorphies are provided for the Cyclorrhapha and Muscoidea, and a grouping of three basal brachyceran infraorders Xylophagomorpha, Stratiomyomorpha and Tabanomorpha. The patterns of fusion suggest that VNC architecture has evolved irreversibly, in accordance with Dollo's law.
 
Article
A new genus is proposed for a new East African Phaneropterinae species, Lunidia viridis, occurring on Mt. Kilimanjaro, Tanzania. Based on 33 records, notes on distribution and habitat are given, as well as acoustical data provided. Climate and vegetation parameters obtained along several transects on Mt. Kilimanjaro were evaluated describing the ecological niche of the new species. This interdisciplinary approach allows not only a profound characterisation of the ecological demands of the new genus but also predictions of the potential distribution area, which is tested for the first time for an African bush cricket species. Lunidia viridis n. gen. n. sp. occurs within humid and perhumid forests and Chagga home gardens, avoiding subhumid conditions on the mountain. It is found from 1,330m upwards on the southern slopes, whereas the same ecological conditions are expressed from 1,930m upwards on the drier northern slopes. Lunidia viridis has an unusually complex and variable song, which is described from field and laboratory recordings. The FISH technique for characterizing chromosomes is applied for the first time for an African species; L. viridis exhibits a karyotype typical for most Tettigoniidae. KeywordsNew species-East Africa-Mt. Kilimanjaro-Eastern Arc-Ecological niche
 
Article
Data documenting skeletal development in rodents, the most species-rich ‘order’ of mammals, are at present restricted to a few model species, a shortcoming that hinders exploration of the morphological and ecological diversification of the group. In this study we provide the most comprehensive sampling of rodent ossification sequences to date, with the aim of exploring whether heterochrony is ubiquitous in rodent evolution at the onset of skeletal formation. The onset of ossification in 17 cranial elements and 24 postcranial elements was examined for eight muroid and caviomorph rodent species. New data are provided for two non-model species. For one of these, the African striped mouse, Rhabdomys pumilio, sampling was extended by studying 53 autopodial elements and examining intraspecific variation. The Parsimov method of studying sequence heterochrony was used to explore the role that changes in developmental timing play in early skeletal formation. Few heterochronies were found to diagnose the muroid and caviomorph clades, suggesting conserved patterning in skeletal development. Mechanisms leading to the generation of the wide range of morphological diversity encapsulated within Rodentia may be restricted to later periods in development than those studied in this work. Documentation of skeletogenesis in Rhabdomys indicates that intraspecifc variation in ossification sequence pattern is present, though not extensive. Our study suggests that sequence heterochrony is neither pivotal nor prevalent during early skeletal formation in rodents. KeywordsRodent-Heterochrony- Rhabdomys -Skeletogenesis-Development-Intraspecific variation
 
Article
For almost all groups of pathogens, unusual and rare host species have been reported. Often, such associations are based on single or few collections only, which are frequently hard to access. Many of them later prove to be due to misidentification of the host, the pathogen, or both. Therefore, such reports are often disregarded, or treated anecdotally in taxonomic and phylogenetic studies, regardless of their potential importance to unravelling the evolution of the entire group. Concerning oomycete biotrophs there are several reports of unusual and rare hosts for hardly known pathogens. In the order Fabales, for example, a single species of Albugo, A. mauginii, was described as parasitic to Onobrychis crista-galli about 80years ago, but not recorded again. All other confirmed members of Albugo s.str. are parasitic to representatives of the families Brassicaceae, Capparaceae, Cleomaceae, and Resedaceae in the order Brassicales. In the present study, molecular phylogenetic analysis of cox2 mtDNA sequences and morphological investigations on an original specimen confirmed the occurrence of a member of Albugo on Fabaceae hosts, with the characteristic thin wall of the secondary sporangia, which is almost uniform in thickness. In phylogenetic analyses the species results as embedded within Albugo s.str. Therefore, it is concluded that the natural host range of Albugo s.str. extends from Brassicales to Fabales via host jumping. Our results underscore that unrevised reports of pathogens from unusual hosts should be reconsidered carefully to obtain a more complete picture of pathogen diversity and evolution. KeywordsAncient DNA– cox2 mtDNA–Host jumping–Pathogen evolution–Unusual hosts
 
Article
It has been argued that adopting alternative nomenclatural procedures would jeopardize the importance of natural history collection by relegating reference specimens to a secondary role. Based on published statements and applications, the present contribution argues that reference specimens are an essential aspect of both the PhyloCode and the cladotypic procedures. Consequently, the latter procedures might actually result in a revival of interest in those collections, mostly by augmenting the number of reference specimens and, for the cladotypic approach, by necessitating appropriate curation and access to up-to-date research facilities. KeywordsHolotype-Specifier-Cladotype-PhyloCode-Nomenclature
 
Article
AbstractA long-term monitoring program has been established in Tasmania, Australia, as a Satellite Project for the International Biodiversity Observation Year (IBOY). This program aims to monitor distributional change in vegetation and fauna assemblages along an altitudinal gradient (70–1300 m) in response to climate change and other environmental events. Baseline data collected over a two-year period will be available for comparison with data collected in future decades.The vegetation varies with altitude and fire history. The rate of change in vegetation is not continuous along the altitudinal gradient, but is most rapid above 700 m and below the treeline at 1000–1100 m. Most vascular plant species reach the limit of their distribution within this zone.Despite their preliminary nature, the invertebrate data also display altitudinal and seasonal patterns. The treeline and the 700–1000 m zone again appear to be notable in terms of invertebrate distribution. While the composition of ground-based taxa may be closely related to the floristic composition of the vegetation (or its environmental drivers), the airborne invertebrate fauna appears to be more closely related to structural characteristics such as height and density. Of all taxa, the Coleoptera appear to be the best potential indicators across most altitudes and times.Although the current data provide a wealth of inventory and distributional information over altitude, their greatest potential value lies in long-term comparative information. Future sampling should focus not only on changes at and above the treeline, but also on the zone below this where many species are at their altitudinal limits and may be particularly sensitive to climate change.
 
Minimum-evolution phylogenetic reconstruction using partial nrLSU (a), cox2 (b), and ypt1 (c) sequences; where bootstrap support values in minimum-evolution or maximum-likelihood analyses were above 50%, these are shown on the branches in that order of analyses, while “–” labels denote lack of such support
Article
Plasmopara viticola is the causal agent of grapevine downy mildew and is among the most important diseases in viticulture. It originates from North America, where it coevolved with wild Vitis species. Beginning in the 1870s it turned into a global epidemic that has been causing severe yield losses. It is generally believed that a single species is causing downy mildew on a large variety of economically important cultivars. Here we report, based on one nuclear and two mitochondrial markers, that isolates from vineyards in the United States fall into three highly distinct phylogenetic lineages. One of these contains European strains and affects Vitis vinifera cultivars, while the other two lineages affect also other species of Vitis. The divergence between these lineages is high, and, judging from the genetic variation in other Plasmopara lineages, might reflect distinct species. Due to the potentially significant implications for quarantine regulations and resistance breeding, detailed studies will be necessary to clarify whether these genetically distinct lineages occur outside of North America or are still confined there. KeywordsCryptic species–Grape downy mildew–Herbarium specimens– cox2–nrLSU– ypt1
 
Maximum Likelihood tree for haplotypes of Chelonoidis carbonaria and C. denticulata, including sequences of C. chilensis and C. nigra. Support values above branches are ML bootstrap values; below branches, Bayesian posterior probabilities and MP bootstrap
Divergence time estimates for nodes in Fig. 4
Parsimony networks for mtDNA haplotypes of Chelonoidis carbonaria (left) and C. denticulata (right), based on alignments of 420 bp. Circle size approximately reflects haplotype frequency. Missing haplotypes are shown as small solid circles. Each line connecting haplotypes corresponds to one mutational step. Large upper-case letters indicate geographic origin of haplotypes within the range of C. carbonaria (N, north; NE, northeast; etc.). Haplotype frequencies for C. carbonaria are B1: n=30; (C1, C2, C3, C4, C5, C6, C9, C10): n=16; C11: n=6; (P1, P2): n=5; (C8, U1): n=4; (G1, G2):
Estimated divergence times for extant species of Chelonoidis and for clades of C. carbonaria (Bayesian tree). Grey lines correspond to credibility intervals, grey bars to one standard deviation above and
Chelonoidis carbonaria ; left: Brazil (Museum of Zoology Dresden MTD D 3620); right: Filadelfia, Chaco, Paraguay (Museum of Zoology Dresden MTD D 43485). Scale bars: 10 cm. Note distinct shell shapes and colorations 
Article
Using sequence data of the mitochondrial cytochrome b gene, we investigated phylogeographic differentiation of the Amazonian tortoise species Chelonoidis carbonaria and C. denticulata. While C. carbonaria is generally restricted to savannah habitats and adjacent forests, C. denticulata is associated with wet tropical and subtropical forests. Our study suggests a correlation between distinct habitat preferences and phylogeography of the two species. In Maximum Parsimony, Maximum Likelihood and Bayesian analyses, haplotypes of C. carbonaria cluster in several distinct clades reflecting the species’ patchy distribution in savannah habitats. By contrast, haplotypes of C. denticulata are only weakly differentiated; a finding also confirmed by parsimony network analysis. This suggests that the contiguous Amazonian rainforest allows gene flow between populations of the forest-dwelling C. denticulata throughout the range, but significantly impedes gene flow in C. carbonaria. The phylogeographic structure and extant distribution pattern of C. carbonaria is supportive of former Amazonian rainforest fragmentation, enabling the dispersal of savannah species. Based on fossil calibration, we dated divergence times for the C. carbonaria clades using a relaxed molecular clock, resulting in average estimates ranging from 4.0–2.2mya. This implies that the onset of rainforest fragmentation could predate the Pleistocene considerably. Furthermore, our findings call for further research on geographic and taxonomic variation in C. carbonaria and for a reassessment of the conservation status of the distinct genetic units. KeywordsPhylogeography-Cytochrome b gene-Testudinidae-Rainforest-Forest refugia hypothesis
 
Article
The dung beetles Amidorus obscurus and A. immaturus are nearly indistinguishable, being characterized by a marked constancy in external morphological traits and little sexual dimorphism in adults. We studied two syntopic populations from the Italian Alps by means of geometric morphometric analyses. To identify possible undetected shape differences between species, we focused on the head, pronotum and scutellum (three external traits) and the epipharynx. Results indicate that the external traits are rather similar in the two species, whereas the epipharynx is clearly different. Interspecific differences in the aedeagus were also taken into account; these are noteworthy because parameres of A. immaturus differ in shape and are at least three times longer than those of A. obscurus. If it is assumed that the diversification of the two species took place during the quaternary ice age, A. immaturus would have evolved these marked differences rather quickly, in keeping with the hypothesis of rapid genital evolution. In an ontogenetic trajectory framework, we also considered the morphology of larvae. Interspecific divergence in the shape of the epipharynx is already evident at the preimaginal stage, whereas that of the genital disc is not. Accordingly, we hypothesise that the feeding and reproductive traits of these two species diverged morphologically when they become functional. Finally, by considering recent advances in ecological and evolutionary knowledge of dung beetles, the pattern of relative constancy in external morphology exhibited by the tribe Aphodiini, and that of great morphological diversification displayed by Onthophagini, were compared, and hypotheses about the origins of these differences discussed.
 
Morphological characters included in the analysis. (a-c) Inner lobe of maxilliped, showing flattened spines; a Niphargus aggtelekiensis "f. lurensis", b N. tatrensis "f. schneebergensis", c N.. aggtelekiensis, topotype. (d, e) Dactylus with (d) and without (e) additional spiniform seta. (f-h) Niphargus tatrensis, telson narrowed and concave in apical part (f = "forma schneebergensis", g = topotype, h = "forma reyersdorfensis"). (i-l) Niphargus aggtelekiensis, telson broad convex (i = topotype, j = "forma oethscerensis", k = "forma lunzensis", l = "forma salzburgensis")
Distribution of character states in the Niphargus tatrensis aggregate
Article
Natural history museum collections harbour valuable information on species. The usefulness of such data critically depends on the accurate identification of species, which has been challenged by introduction of molecular techniques into taxonomy. However, most collections may suffer from DNA degradation, due to age and/or improper preservation; hence the identification of specimens depends solely on morphological features. This study explores how and to what extent morphological data can help to solve ambiguous taxonomic cases based on selected species concepts and with the use of operational criteria in a species-hypothesis testing procedure. The studied taxon, the Niphargus tatrensis species complex, comprises freshwater subterranean amphipods, distributed across Central Europe, the taxonomic status of which was debated extensively between 1930 and 1960. Using the general species concept, character- and tree-based operational criteria reveal northern and southern diagnosable and exclusive lineages identified here as N. tatrensis Wrześniowski, 1888 and N. scopicauda sp. n., respectively. The remaining populations represent the non-exclusive N. aggtelekiensis Dudich, 1932, which occurs from the eastern Alps to Hungary. In the entire complex, altitudinal distribution is largely limited to areas above 400m, where the mean annual temperature never exceeds 9°C. Seemingly well-defined distributional ranges of N. tatrensis and N. aggtelekiensis are fragmented in an ecological sense, which raises the question whether two of the three species recognised here actually consist of several unidentified taxa. Morphological similarity between the species, numerous polymorphic features, and the association with cool temperatures lead to a hypothesis in which fragmentation of the ancestral range occurred during post-Pleistocene climate warming, reducing gene flow across lowland populations due to niche conservatism of the ancestral species and/or to invasion of competitive species along the Danube and Drava rivers. The results are discussed regarding how old museum samples are conducive to more detailed molecular-taxonomic and conservation studies. KeywordsGeneral species concept-Population aggregation analysis-Tree based species delimitation-Mantel test-Museum collections-Polymorphic characters
 
Article
AbstractAmong genes coding for proteins with basic structural functions in all eukaryotes, the highly conserved and functionally essential gene for β-tubulin is receiving increasing attention in the reconstruction of phylogenies within a broad organismic range. We therefore constructed a set of twelve universally applicable primers that allow reliable amplification of β-tubulin genes among all major eukaryotic kingdoms including fungi (Fungi), animals (Animalia) and green plants (Planta). For primer design, the amino acid sequences of 35 β-tubulin genes from Ascomycota, Basidiomycota, Chytridiomycota, Zygomycota, Animalia, Oophyta and Planta were aligned and used for the definition of four well-conserved regions. These are suitable priming sites in PCR amplification experiments. Out of these amino acid regions twelve primers were designed which initiate especially the amplification of fungal β-tubulin genes. In four pairwise primer applications gene fragments of up to 1500 bp in size could be isolated, which comprise nearly complete β-tubulin genes from twelve species representative of the Fungi. The sequences of seven β-tubulin fragments were obtained from Allomyces moniliformis, A. neomoniliformis, Blastocladiella britannica, Chytridium confervae, Mortierella isabellina and Trametes versicolor, respectively. Reliable amplification of β-tubulin over a broad spectrum of organisms provides a strong basis for the establishment of both deep-level phylogenies and studies of complex species groups based on β-tubulin gene trees.
 
Article
The phylogeny of the genus Cynanchum s. str. is studied using cpDNA spacers and ITS. Morphological, anatomical and latex triterpenoid data are interpreted in light of the molecular results, and discrepancies are discussed. Vegetative characters are better indicators of relationship than floral characters, especially corona characters. The monophyly of all Malagasy species and, nested within the latter, of all stem-succulent taxa is ascertained and the genera Folotsia, Karimbolea, Platykeleba and Sarcostemma are subsumed under Cynanchum. One African species, C. galgalense, is excluded from Cynanchum.
 
Article
Four new species of Cyatheaceae from Ecuador are described: Alsophila conantiana Lehnert, Cyathea brucei Lehnert, C. moranii Lehnert, and C. sylvatica Lehnert. Range extensions are documented for Alsophila esmeraldensis R.C. Moran and Cyathea macrocarpa (C. Presl) Domin.For full article, see Electronic Supplement at: http://www.senckenberg.de/odes/06-13.htm
 
Article
AbstractFormaldehyde was first prepared in 1859, and since then has been in widespread use for fixing and preserving medical and biological specimens. The value of such archival material has increased considerably because several methods for extracting DNA from formaldehyde-fixed animal tissue have been developed. Most of these, however, either require large amounts of tissue (rarely available) or recover only short fragments of DNA. Here we summarize current knowledge of and experience with such published methods, look at some of the known problems, and develop an additional method based on embedding the tissue in agarose prior to treatment with proteinase-K and GeneReleaser. With this method we have obtained mitochondrial DNA useful for PCR reactions from as little as 3 mg tissue of more than 30 years old formaldehyde-fixed aplacophoran molluscs. We examine the conditions under which obtaining relatively high-quality DNA from formaldehyde-fixed material is possible, making previously collected samples accessible for molecular studies in genetics, systematics and related fields. The purpose of this short review is to acquaint molecular systematists with some of the methodological advances and considerations in using formaldehyde-preserved material.
 
Article
AbstractThe current revision provisionally lists 77 valid earthworm taxa in seven families from Japan, with approximately 80 further names (ca. 50% of the total) either in synonymy – including 40 new synonyms – or retained as species incertae sedis. Generic placement of the species yielded 17 new combinations. About 30 species are known introductions and another ten are possibly more widespread, thus the probable number of wholly endemic Japanese earthworms is around 40 species (ca. 50% of the total valid species). However, a definitive work on the systematics of Japan's earthworms is pending, and the current revision aims only to provide a status quo and to track changes from the last comprehensive revision by Easton (1981) that listed 74 taxa. Subsequently, 60 or so new pheretimoid names were added by Ishizuka in 1999–2001, but only a few are considered valid taxa, the remainder being synonyms or species incertae sedis. The substitute name ‘Pheretima’ palarva, nom. nov., is provided for the junior homonym Pheretima parvula Ishizuka et al., 2000.While much of Easton's synopsis is supported, Pontodrilus is now placed in Megascolecidae sensu Blakemore (2000) rather than Acanthodrilidae sensu Gates (1959); Amynthas carnosus (Goto & Hatai, 1899) is removed from synonymy with Amynthas gracilis; and an informal Amynthas corticis species-complex is established to accommodate the various morphs of this widely distributed species group. Pheretima (Parapheretima) koellikeri Michaelsen, 1928 is considered synonymous with Metaphire vesiculata (Goto & Hatai, 1899), thereby removing the genus Pheretima sensu stricto from the Japanese list. Polypheretima is also removed from Japanese indigeny, as the original description of Polypheretima iizukai (Goto & Hatai, 1899) failed to report intestinal caeca, and inspection of fresh material allows its placement in synonymy with Amynthas fuscatus (Goto & Hatai, 1898). Easton (1981) had listed this taxon as Metaphire fuscata, but further demonstration of superficial male pores qualifies it for Amynthas. Conversely, the discovery of copulatory pouches results in transfer from Amynthas to Metaphire for M. hilgendorfi (Michaelsen, 1892), comb. nov., Metaphire communissima (Goto & Hatai, 1898), comb. nov., and Metaphire megascolidioides (Goto & Hatai, 1899), comb. nov. The Metaphire hilgendorfi/Amynthas tokioensis species-complex (Amynthas hilgendorfi species-complex sensu Easton 1981) remains one of the most intractable and pressing problems for comprehension of the Japanese fauna, as most of the component taxa, e.g. Metaphire agrestis (Goto & Hatai, 1899), are parthenogenetically degraded morphs as yet unaffiliated with their ancestral and biparental populations. Resolution may be sought employing combinations of morphological and molecular (RNA, DNA) techniques to determine specific affinities while also complying with requirements of the International Code of Zoological Nomenclature (ICZN 1999).
 
Article
Annelida, traditionally divided into Polychaeta and Clitellata, are characterized by serial division of their body into numerous similar structures, the segments. In addition, there is a non-segmental part at the front end, the prostomium, and one at the back, the pygidium. New segments develop in a prepygidial proliferation zone. Each segment contains four groups of chaetae made up of β-chitin, a pair of coelomic cavities separated by mesenteries, and septa. The nervous system is a rope-ladder-like ventral nerve cord with a dorsal brain in the prostomium. For the last stem species a trochophore larva and a benthic adult are commonly postulated. There are two conflicting hypotheses describing the systematization of Annelida: the first postulates a sister-group relationship of Polychaeta and Clitellata, the second sees Clitellata as a highly derived taxon forming a subordinate taxon within the polychaetes which, consequently, are regarded as paraphyletic. Depending on the hypothesis, different characters have to be postulated for the stem species of Annelida. Besides segmentation other characters such as nuchal organs, palps and antennae, body wall musculature, cuticle, parapodia as well as structure of the central nervous system and the foregut play an important role in this discussion. Here, the different characters and character states are critically reviewed and analyzed with respect to morphology and function. The consequences for systematization of their phylogenetic interpretation as autapomorphies, synapomorphies or plesiomorphies are outlined. The resulting hypotheses are compared with those relying on molecular data sets.
 
Article
In shallow sublittoral sediments of the north-west coast of the Island of Elba, Italy, a new gutless marine oligochaete, Olavius ilvae n. sp., was found together with a congeneric but not closely related species, O. algarvensis Giere et al., 1998. In diagnostic features of the genital organs, the new species differs from other Olavius species in having bipartite atria and very long, often folded spermathecae, but lacking penial chaetae. The Elba form of O. algarvensis has some structural differences from the original type described from the Algarve coast (Portugal). The two species from Elba share characteristics not previously reported for gutless oligochaetes: the lumen of the body cavity is unusually constricted and often filled with chloragocytes, and the symbiotic bacteria are often enclosed in vacuoles of the epidermal cells. Regarding the bacterial ultrastructure, the species share three similar morphotypes as symbionts; additionally, in O. algarvensis a rare fourth type was found. The divergence of the symbioses in O. algarvensis, and the coincidence in structural and bacteria-symbiotic features between the two taxonomically different, but syntopic host species at Elba are discussed.
 
Article
The diversity, taxonomy and distribution of the Opheliidae (Annelida: Polychaeta) in Icelandic waters is reviewed based on material collected during the BIOICE project. Nine opheliid species are recorded from Iceland; of these, three were previously reported in the area (Ophelia limacina, Ophelina cylindricaudata and O. acuminata), four are new for Icelandic waters (Ammotrypanella cf. arctica, Ophelina abranchiata, O. helgolandica and Tachytrypane jeffreisii), and two are new to science. Ophelina basicirra sp. nov. is distinguished by having a narrow anal tube with one short proximal anal cirrus in ventral position. Ophelina bowitzi sp. nov. is characterized by the small size of its anterior branchiae, which become larger in the middle and posterior body regions, and by an upwardly bent anal tube that is wide at the base and narrowing distally. The genera Euzonus, Armandia and Polyophthalmus are not represented in the BIOICE samples. The distribution of each species off Iceland is presented; one species is restricted to shallow waters of northwestern fjords, three species are found south of the GIF Ridge, and five species are circumicelandic. Several body characters with taxonomic relevance in some species are reviewed based on SEM images. Furthermore, as a first step towards a future revision of the genus Ophelina Örsted 1843 in North Atlantic waters, the status of each species originally described or subsequently reported from the area is commented on, and a key to the currently valid species is presented. Ophelina longicephala Hartmann-Schröder, 1977, formerly a subspecies of O. delapidans (Kinberg, 1866), is raised to species status. Keywords Annelida–Ophelina–GIF Ridge–Deep sea–BIOICE project–SEM–Taxonomy–New species
 
Distances measured for morphometric characters ScBaC and ScBW  
Article
The Mediterranean harvester ant species Messor minor, M. cf. wasmanni, and M. capitatus can co-occur in the same habitat. In Italian populations, we encountered colonies that contained workers from more than one species as identified via standard morphology, as well as colonies with workers that appeared to be morphologically intermediate between species. This unusual finding required further analysis. We analysed such colonies using microsatellites, mitochondrial DNA and refined morphometrics, and a simple inference key for the colony-level interpretation of data from the three sources combined. We infer that Messor minor and M. cf. wasmanni engage in bidirectional interspecific gene flow. Hybrids between these two species are inferred to produce fertile offspring, which would indicate that barriers to hybridisation do not exist or can be completely overcome. This is unexpected, given that they are non-sister species and broadly sympatric in nature. Our findings also indicate the possible occurrence of hybrid-hybrid crosses, a phenomenon rarely observed in ants. We cautiously interpret the data at hand as in support of the interspecific gene flow considerably shaping the genetic makeup of populations, raising the question about a potential adaptive value of this hybridisation. Messor capitatus mixes with hybrids of the other two species, but we found no indication of hybridisation involving this species. We discuss various hypotheses on the causations of colony mixing and hybridisation in the three Messor species at the proximate and ultimate level. KeywordsHybridisation–Bidirectional interspecific gene flow–Non-sister species–Inference key
 
ANOVA
Article
Species delimitations in the predominantly Antarctic and South American group of neuropogonoid species of the lichen-forming fungal genus Usnea are poorly understood. Morphological variability has been interpreted as a result of harsh ecological conditions, but preliminary molecular data have led to doubts about the current species delimitations in these lichenized fungi. We examined species boundaries using a phylogenetic approach and a cohesion species recognition method generating haplotype networks and looking at associations of phenotypic characters with clades found in the networks. In addition, we estimated gene flow among detected clades and currently circumscribed species. We identified several clades that were significantly associated with phenotypic characters, but did not necessarily agree with current species circumscriptions. In one case (U. aurantiaco-atra/U. antarctica), network analysis and the estimation of gene flow provided no evidence of distinct species. The distinctness of another species pair (U. subantarctica/U. trachycarpa) remains dubious, showing evidence for gene flow among currently accepted species. KeywordsLichens– Usnea –Species delimitation–Cohesion species
 
Maximum likelihood tree obtained by heuristic searches (10 random-addition sequence replicates), based on 1084 base pairs of the 12S and 16S rRNA genes in selected representatives of the major dendrobatid lineages as identified by preliminary analysis (Fig. 1). Xenopus laevis was used as the outgroup. First numbers at nodes are results of maximum likelihood bootstrapping (in percent; 100 replicates), second numbers are results of maximum parsimony bootstrapping (in percent; 2,000 replicates), third numbers are Bayesian posterior probabilities (500,000 generations, every tenth generation sampled, burn-in set at 2%). Values are shown only for nodes that received support >70% in at least one analysis. Circles denote coloration (black = aposematic, grey = near to aposematic, white = non-aposematic), squares denote presence of skin alkaloids (black = present, grey = present in traces, white = not present, ? = not yet tested).
Examples of live coloration of aposematic and non-aposematic dendrobatid frogs from distinct clades (Table 1, Fig. 1) identified through our analysis, and geographic distribution of all species from the respective clusters for which genetic data were available (undescribed and unidentified species not considered). In the distribution maps, red color marks areas of aposematic species, grey those of cryptic species. A. Cryptophyllobates and Colostethus clade 4: A1 Cryptophyllobates azureiventris; A2 Colostethus bocagei. B. Epipedobates (from west of the Andes) and Colostethus clade 3: B1 Epipedobates tricolor; B2 Colostethus pratti. C. Allobates and Colostethus clade 1: C1 Allobates zaparo; C2 Colostethus humilis. D. Mannophryne, Nephelobates and Colostethus clade 2: D1 Mannophryne herminae; D2 Nephelobates molinarii. Distribution areas are given for the following species: 1 Colostethus subpunctatus; 2 Colostethus bocagei; 3 Cryptophyllobates azureiventris; 4 Colostethus pratti; 5 Epipedobates boulengeri; 6 E. tricolor and E. anthonyi; 7 Allobates femoralis; 8 A. zaparo; 9 Colostethus talamancae; 10 C. trilineatus; 11 C. humilis; 12 C. marchesianus; 13 Mannophryne and Nephelobates; 14 Colostethus palmatus; 15 C. stepheni; 16 C. baeobatrachus and C. degranvillei. Distributional information is based on Silverstone (1975, 1976), Schulte (1999), De la Riva et al. (2000), Lescure & Marty (2001), Lima & Caldwell (2001), Caldwell et al. (2002), Caldwell & Lima (2003), and the Amphibian Species of the World database ( gy/amphibia/) as of October 2002. Photographs by K.-H. Jungfer (A1, B1, B2, C1), M. Read (A2), and M. Vences (C2, D1, D2). 
Article
Poison frogs of the family Dendrobatidae contain cryptic as well as brightly colored, presumably aposematic species. The prevailing phylogenetic hypothesis assumes that the aposematic taxa form a monophyletic group while the cryptic species (Colostethus sensu lato) are basal and paraphyletic. Analysis of 86 dendrobatid sequences of a fragment of the 16S rRNA gene resulted in a much more complex scenario, with several clades that contained aposematic as well as cryptic taxa. Monophyly of the aposematic taxa was significantly rejected by SH-tests in an analysis with additional 12S and 16S rDNA fragments and reduced taxon sampling. The brightly colored Allobates femoralis and A. zaparo (Silverstone) comb. nov. (previously Epipedobates) belong in a clade with cryptic species of Colostethus. Additionally, Colostethus pratti was grouped with Epipedobates, and Colostethus bocagei with Cryptophyllobates. In several cases, the aposematic species have general distributions similar to those of their non-aposematic sister groups, indicating multiple instances of regional radiations in which some taxa independently acquired bright color. From a classificatory point of view, it is relevant that the type species of Minyobates, M. steyermarki, resulted as the sister group of the genus Dendrobates, and that species of Mannophryne and Nephelobates formed monophyletic clades, corroborating the validity of these genera. Leptodactylids of the genera Hylodes and Crossodactylus were not unambiguously identified as the sister group of the Dendrobatidae; these were monophyletic in all analyses and probably originated early in the radiation of Neotropical hyloid frogs.
 
Potential distribution ( gray shaded ) of Dorcas gazelles ( G. dorcas ) according to IUCN antelope survey reports (East 1988a, b; Mallon and Kingswood 2001) including sampling locations of wild specimens of Dorcas ( ♦ ) and Saudi gazelles ( ) used in this study. 
Article
Many species of gazelles (Gazella spp.) are nowadays threatened by hunting, poaching, habitat loss and habitat deterioration. Conservation efforts for this group not only face the problem of maintaining remnant populations, but often natural populations have been extirpated from the wild. In some cases, though, captive breeding programs exist that might provide a valuable source for future reintroductions. A major problem in this context is that phylogeographic relationships among different (potentially locally adapted) populations, and even basic phylogenetic relationships between species, are poorly understood, thus hampering the assignment of management units, breeding groups or stocks for reintroduction projects. Our present study focused on Dorcas gazelles (G. dorcas and G. saudiya) from the species’ entire distribution range, with samples originating from western Saharan Africa into Saudi Arabia. In stark contrast to previous studies reporting on pronounced genetic structure in taxa such as Mountain gazelles (G. gazella), we detected low genetic diversity and no evidence for major phylogenetic lineages when analyzing two mitochondrial genetic markers. Using a coalescent approach we infer a steep population decline that started approximately 25,000years before present and is still ongoing, which coincides with human activities in Saharan Africa. Our phylogenetic analyses, statistical parsimony network analysis and inferred colonization patterns shed doubt on the validity of various described subspecies of G. dorcas. KeywordsDesert ungulate– Gazella dorcas –Conservation unit–Phylogeography–Range expansion
 
Strict consensus tree of all eight most parsimonious trees, with Bremer support values (TreeBASE S2624) shown besides nodes; for tree section with 'distal erigonines', see Fig. 2. Topology congruent  
Strict consensus tree of the 'distal erigonines', with unambiguous character optimisation (squares) and clade numbers (in circles); see also Figs. 4, 5 and 6. Squares mark homoplasious (white) and non-homoplasious (black) character-state changes; character and character-state numbers given above and below squares, respectively  
Continuation of Fig. 3, showing clade 21  
Continuation of Fig. 4, showing clade 35  
Continuation of Fig. 5, showing clade 52  
Article
We present the most inclusive study on the higher-level phylogeny of erigonine spiders, including about 30% of all erigonine genera. By expanding the previously most comprehensive analysis (Miller and Hormiga Cladistics 20:385–442, 2004) we tested the robustness of its results to the addition of closely related taxa, and also the monophyly of the Savignia-group defined by Millidge (Bulletin of the British Arachnological Society 4:1–60, 1977). The character matrix was expanded by adding 18 newly scored species in 15 genera, and also includes all species scored by other authors. This adds up to 98 species in 91 erigonine genera plus 13 outgroup taxa. The parsimony analysis led to eight fully resolved most parsimonious trees (L=1084). The topology concerning the taxa basal to the ‘distal erigonines’ remained unchanged, and the latter clade still shares 67% of all nodes with the original analysis. The Savignia-group is not monophyletic at genus level with respect to Saloca diceros and Alioranus pastoralis, and the same applies at species level in Diplocephalus and Erigonella. From the Savignia-group, Glyphesis servulus, Diplocephalus cristatus, Savignia frontata, and two representatives each of Erigonella, Dicymbium and Araeoncus combine to form a monophyletic clade. KeywordsPhylogeny-Morphology-Complex genital organs-Dwarf spiders-Erigoninae
 
18SrRNA-gene sequences used in this study for the Neighbor-Joining tree construction.
(Continued).
Article
Analysis of the monosaccharide composition of purified cell walls of unicellular and filamentous ascomycetous fungi shows three patterns: (1) the mannose glucose type (for most hemiascomycetous yeasts) (2) the mannose glucose galactose type (for several members of all three main ascomycetous clades) and (3) the mannose glucose galactose rhamnose type (for members of the Euascomycetes and the Protomyces/ Schizosaccharomyces group).In order to estimate the usefulness of the carbohydrate patterns for phylogenetic analysis we compared them with a phylogenetic tree based on 18SrRNA-gene sequences using the Neighbor-Joining Method. In contrast with the situation for basidiomycetous fungi, the Ascomycota show no fixed cell wall type for the three classes. Based on cell wall carbohydrates, sequence data and molecular characters the Hemiascomycetes appear as the first branch within the Ascomycota. A second clade, comprising the genera Schizosaccharomyces, Pneumocystis, Taphrina, Protomyces, Neolecta and Saitoella, appears as a sister group of the Euascomycetes. We discuss the erection of a new class for this group of ascomycetous fungi for which we propose the name Protomycetes.
 
GenBank accession numbers for sequences used in this study, and corresponding voucher specimens (provided only for sequences not used in previous publications)
50% majority rule consensus tree from four Bayesian analyses of 1,000,000 generations each of the rbcL data set, excluding burn-in trees. Posterior probability support values≥0.90 from Bayesian analyses shown above branches, bootstrap support values>50% from Maximum Likelihood analyses below branches
50% majority rule consensus tree from Bayesian analyses of the three-gene ( rbc L, rps 4- trn S, and trn L- trn F) data set. Numbered solid circles indicate nodes receiving good support in both Bayesian (PP ≥ 0.90) and Maximum Likelihood (BS ≥ 70) analyses; open circles indicate nodes not receiving good support from either mea- sure; see also Figure 3. Single letters in dark and light grey squares on branches refer to synapomorphic indels in the rps 4- trn S and trn L- trn F alignments, respectively. “ P ” denotes polystichoid ferns; major groups also indicated at right 
Article
Cyrtomidictyum Ching and Cyrtogonellum Ching are two eastern Asian endemic genera whose taxonomic affinities and phylogenetic relationships have long been controversial. The main uncertainty surrounds the separation of the two genera from the species-rich genus Polystichum. Here we present a phylogenetic study focusing on the phylogenetic relationships of these polystichoid ferns. We reconstructed the relationships based on DNA sequence variation in four chloroplast genome regions, rbcL, atpB, and the intergenic spacers (IGS) rps4-trnS and trnL-trnF. Maximum likelihood and Bayesian inference analyses confirm earlier results that were based on less comprehensive taxon sampling and either only a single gene (rbcL) or two IGS (rps4-trnS and trnL-trnF). Cyrtomidictyum is the sister of the clade of polystichoid ferns that includes Cyrtogonellum, Cyrtomium subser. Balansana and three sections of Polystichum. Cyrtogonellum groups with several species of Polystichum, and constitutes the sister taxon to Polystichum sect. Sphaenopolystichum. We support the recognition of Cyrtomidictyum as circumscribed initially, rather than expansion of the genus to include either several Polystichum species or Cyrtogonellum, some Polystichum and Cyrtomium species. The monophyly of Cyrtomidictyum is supported by morphological characters such as once-pinnate leaves, free venation, prolongated leaf apices, and exindusiate sori. Two synapomorphic indels in the chloroplast genome, one 15-bp deletion in rps4-trnS, and one 3-bp insertion in trnL-trnF further differentiate Cyrtomidictyum from other polystichoid ferns. The close affinity of Cyrtogonellum to section Sphaenopolystichum of Polystichum s.s. is highly supported by molecular data. However, no shared morphological characters or molecular indels have been detected, although the distinctness of Cyrtogonellum is shown by a 13-bp insertion in the rps4-trnS alignment. KeywordsEast Asia-Molecular phylogeny- atpB- rbcL- rps4-trnS- trnL-trnF-IGS
 
External morphology of manus and pes in the trionychid Pelodiscus sinensis ( a , b ) and the carettochelyid Carettochelys insculpta ( c , d ). ( a ) Dorsal view; ( b – d ) ventral views. Both trionychids and carettochelyids are pentadactylous, but claws are present only on the first three digits (manus and pes) in trionychids, on the first two digits in C. insculpta 
Article
In order to examine the evolution of the phalangeal formula in a diverse clade of turtles, including hyperphalangy as a rare condition in this group, we studied 210 specimens representing all extant genera of Trionychidae and their sister taxon, Carettochelyidae. Both groups consist of highly aquatic species with elongated autopods that are either paddle-like (Trionychidae) or transformed to flippers (Carettochelyidae). Phalangeal formulae were obtained mostly by radiographs of alcohol-preserved or dry specimens, as well as by direct counts from skeletons. All trionychids and Carettochelys are pentadactylous, but their phalangeal formulae differ. Carettochelys exhibits the turtle-plesiomorphic state (manus and pes: 2-3-3-3-3), with no variation in adults. Trionychids exhibit intraspecific variation, ranging from 2-3-3-3-2 to 2-3-3-6-5 for the manus, and from 2-3-3-3-2 to 2-3-3-5-3 for the pes. The extant Carettochelys as well as the Middle Eocene Allaeochelys crassesculpta are characterized by an elongation of phalanges, whereas trionychids consistently have shorter phalanges. All trionychid genera exhibit some degree of hyperphalangy in digits IV and V, in both the manus and pes. Phalanges of the clawed digits I–III are very robust compared to phalanges of the non-clawed digits IV and V. The latter contribute significantly to the enlargement of the paddle by their additional phalanges. We hypothesize that this phalangeal pattern is coupled with prolongation of growth processes in the non-clawed digits. The differences in autopod morphology between carettochelyids and trionychids reflect different locomotor patterns related to different natural histories (elongated flippers for high-speed escape in the mainly herbivorous Carettochelys; broad paddles for rapid turns during hunting in the mainly carnivorous trionychids). The autopod of Pelodiscus sinensis is proposed as an experimental model to examine the developmental basis of adult autopod variation.
 
Article
Crinoid associates represent an abundant and diverse, but poorly explored, component of the hidden biodiversity of coral-reef ecosystems. We studied data from 5years of collecting in the Bay of Nhatrang (BN), Vietnam, to assess the diversity of crinoids and their symbionts, to compare it with other areas of the Indo-West Pacific, and to elucidate the extent to which the observed diversity of crinoids and their symbionts corresponds to their true diversity. In total, about 2,287 specimens of symbionts belonging to 70 species were found on 203 specimens of crinoids belonging to 33 species. Among the crinoids, the most numerous species were Himerometra robustipinna (36 specimens) and Cenometra bella (29 specimens), among the symbionts the polychaete Paradyte crinoidicola (c. 850 specimens) and the galatheid crustacean Allogalathea elegans (180 specimens). Species accumulation curves suggest that we have sampled most of the crinoid diversity in the BN, whereas the diversity of their symbionts remained undersampled. Estimated species richness of crinoids was higher than previously observed richness, and varied from 39 (estimated by bootstrap) to 46 (jackknife 2). Estimated species richness of symbionts was higher than observed richness, and varied from 71 (bootstrap) to 93 (jackknife 2). We suggest a slight increase in the number of crinoid species to result from more detailed studies of nocturnal species, and an increase in the number of symbiotic species when studies of nocturnal crinoid associates and sibling species among decapods are included. Our study revealed a rather rich crinoid fauna in the bay compared to other areas of the Indo-West Pacific, and the highest species richness of crinoid associates known from anywhere in the World Ocean. KeywordsBiodiversity–Species richness–Crinoids–Symbionts–Vietnam
 
Article
We analyzed the flowering phenodynamics of 43 Asteraceae species co-occurring in natural populations of Chaco Serrano forests in central Argentina. We explored the potential influence of factors such as photoperiod and climate (variations in temperature, rainfall, and frost), animal-plant interactions (richness of floral visitors, frequency of visits), some plant attributes (plant growth form, seed dispersal mechanism), and evolutionary relationships among species on flowering phenodynamics. Cluster Analysis (CA) and Principal Component Analysis (PCA) were the multivariate statistical methods used to analyze emerging patterns associated with these co-occurring species. Null-model analyses were used to evaluate whether flowering times are aggregated, segregated, or random. Results showed that flowering phenology was significantly correlated with the seasonal variation in temperature, photoperiod, rainfall, and frost. The multivariate statistical methods separated all the species in three groups: 1) species with short flowering time, large plant floral display, high frequency of visits by a large number of species of floral visitors, anemochorous fruits, and shrubby growth form, with a tendency to a segregated flowering pattern; 2) species with long flowering time, small plant floral display, low frequency of visits by few insect species, anemochorous fruits, and herbaceous growth form; and 3) species with long flowering time, small plant floral display, intermediate values for frequency of visits and number of species of floral visitors, seed dispersal mechanisms other than anemochory, and herbaceous growth form. In addition, all but one species belonging to early-branching tribes (tribes phylogenetically close to the root of the Asteraceae tree) were grouped together and clustered in the same region of the two-dimensional PCA ordination. All species belonging to the late-branching tribes (Asteroideae subfamily tribes) included in group 1 were separated from the other Asteroideae species in the PCA. In conclusion, it seems that climatic factors restrict the phenological period of most species, and that plant attributes and taxonomic membership are strongly related to flowering phenodynamics in this group of Asteraceae studied. KeywordsCompositae–Photoperiod and climate–Evolutionary relationships–Plant-animal interactions–Flowering phenology–Argentina
 
Top-cited authors
Daniele Silvestro
  • Université de Fribourg
Ingo Michalak
  • RANA - Büro für Ökologie und Naturschutz Frank Meyer
Charles Oliver Coleman
  • Museum für Naturkunde - Leibniz Institute for Research on Evolution and Biodiversity
Gonzalo Giribet
  • Harvard University
Uwe Fritz
  • Senckenberg Research Institute