Global diversity of craneflies (Insecta,Diptera: Tipulidea or Tipulidae <Emphasis Type="Italic">sensu lato</Emphasis>) in freshwater

The Tipulidae s.l.—craneflies—are one of the largest groups of the Diptera containing over 15,270 valid species and subspecies. The immatures of the majority of species live in aquatic or semiaquatic habitats. Some aquatic species live entirely submerged and lack functional spiracles, others come to the surface to take oxygen by using spiracles positioned at the end of the abdomen. Semiaquatic species occur in a wide range of habitats. The semiterrestrial and terrestrial larvae live in environments that are moist or at least humous. All adult craneflies are terrestrial. Conflicting hypotheses on the phylogenetic position of the Tipuloidea within the Diptera continue to exist: some authors consider them to represent one of the oldest lineages of the Diptera, others suppose a close relationship to the Brachycera, the true flies. Current systematic knowledge of the Tipuloidea indicates that the Palaearctic region contains the highest number of genus-group taxa, while the Neotropical region has the highest number of species and subspecies. The Afrotropical and Australasian regions are relatively poor respectively in genera and subgenera and in species and subspecies. The oldest fossils that represent the Tipuloidea date back to the Lower Triassic at about 240 million years. Present-day general distribution patterns of many higher taxa of Tipuloidea probably have a Pangean or Gondwanan origin. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment.     

Comparative food web structure of larval macrolepidoptera and their parasitoids on two riparian tree species

Single species or groups of species can be subjected to differing levels of parasitism on different plants. Previous studies have reported that parasitism of larval macrolepidoptera in an assemblage on box elder (Acer negundo L.) was significantly greater than on black willow [Salix nigra (Marsh)]. In this study, quantitative food webs, parasitoid overlap diagrams and other food web attributes were used to identify and describe direct and indirect interactions, and to compare assemblages on each tree species. These comparisons helped identify possible mechanisms explaining the differential parasitism observed. Although links among numerically dominant species in each assemblage were not strong, links between numerically dominant and subdominant species were strong. That is, numerically dominant and subdominant species interacted via shared parasitoids. The degree of parasitoid sharing by numerically dominant and subdominant species differed in each tree. There was less sharing of parasitoids on black willow than on box elder. Further, on box elder, the majority of parasitoids affecting numerically subdominant species originated from numerically dominant species, unlike in willow. These results lead to a working hypothesis—the source/nursery hypothesis—that proposes that community-wide levels of parasitism are highest in circumstances in which numerically subdominant species share parasitoid species in common with numerically dominant species, and most parasitoids attacking subdominant species originate from numerically dominant species. Thus, differences in degree of sharing and the types of herbivores sharing parasitoids may explain differential parasitism. Further, the source/nursery hypothesis may explain why the vast majority of species in most assemblages are numerically subdominant. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ontogeny of flight initiation in the fly <Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: implications for the giant fibre system

There are two modes of flight initiation in Drosophila melanogaster—escape and voluntary. Although the circuitry underlying escape is accounted for by the Giant fibre (GF) system, the system underlying voluntary flight initiation is unknown. The GF system is functionally complete before the adult fly ecloses, but immature adults initially fail to react to a stimulus known to reliably evoke escape in mature adults. This suggests that escape in early adulthood, ∼2-h post-eclosion, is not automatically triggered by the hard-wired GF system. Indeed, we reveal that escape behaviour displays a staged emergence during the first hour post-eclosion, suggesting that the GF system is subject to declining levels of suppression. Voluntary flight initiations are not observed at all during the period when the GF system is released from its suppression, nor indeed for some time after. We addressed the question whether voluntary flight initiation requires the GF system by observing take-off in Shak-B ² mutant flies, in which the GF system is defunct. While the escape response is severely impaired in these mutants, they displayed normal voluntary flight initiation. Thus, the escape mechanism is subject to developmental modulation following eclosion and the GF system does not underlie voluntary flight. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal niche dynamics, relative abundance and phylogenetic signal of coexisting species

The coexistence of similar species accounts for some 30% of diversity within communities, yet the coexistence and relative abundance of similar species is a continuing ecological conundrum. Using close phylogenetic relatedness as a measure of similarity, we previously demonstrated that neither classic niche theory nor neutral theory can explain the relative abundances of co-occurring pairs of similar tree species in a diverse tropical forest. Here, we show that the stable, focused competition of a temporal niche dynamic fits the distribution of observed fractional abundances (pairwise relative abundances). Previously published, independent evidence of temporal dynamics in this community supports our results; our model identifies additional criteria for field tests of differential sensitivity (DS) temporal dynamics. The success of temporal dynamics at explaining the observed distribution—and the failure of alternative hypotheses to do so—indicates that current diagnostics of community structure and assembly needs general re-examination. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Regulatory Evolution of a Duplicated Heterodimer Across Species and Tissues of Allopolyploid Clawed Frogs (<Emphasis Type="Italic">Xenopus</Emphasis>)

Changes in gene expression contribute to reproductive isolation of species, adaptation, and development and may impact the genetic fate of duplicated genes. African clawed frogs (genus Xenopus) offer a useful model for examining regulatory evolution, particularly after gene duplication, because species in this genus are polyploid. Additionally, these species can produce viable hybrids, and expression divergence between coexpressed species-specific alleles in hybrids can be attributed exclusively to cis-acting mechanisms. Here we have explored expression divergence of a duplicated heterodimer composed of the recombination activating genes 1 and 2 (RAG1 and RAG2). Previous work identified a phylogenetically biased pattern of pseudogenization of RAG1 wherein one duplicate—RAG1β—was more likely to become a pseudogene than the other one—RAG1α. In this study we show that ancestral expression divergence between these duplicates could account for this. Using comparative data we demonstrate that regulatory divergence between species and between duplicated genes varies significantly across tissue types. These results have implications for understanding of variables that influence pseudogenization of duplicated genes generated by polyploidization, and for interpretation of the relative contributions of cis versus trans mechanisms to expression divergence at the cellular level. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Patterns in the distribution of Arctic freshwater zooplankton related to glaciation history

We analysed circumpolar samples from 68 lakes within the 10°C-July isotherm from Arctic Canada, Nunavut, Greenland, Svalbard, Eastern Siberia, the Beringia region, and Alaska. In total, we found 3 species of Anostraca, 17 of Diplostraca, 1 species of cyclopoid and 14 species of calanoid copepods. Our study identifies a wider distribution for some copepods—e.g. Eurytemora pacifica, Leptodiaptomus sicilis, Arctodiaptomus novosibiricus, Cyclops abyssorum—than previously known. Moreover, one anostracan species, Artemiopsis bungei, was recorded in North America for the first time; and one chydoriid, Chydorus gibbus, is a new species for Greenland. We observed that species richness of crustaceans is lower in lakes that were glaciated during the Quaternary period, compared to those not glaciated (e.g. Chukotski Peninsula, Siberia; Point Barrow, Alaska; and Disko Island, Greenland). This confirms the findings of classic studies: glaciation has strongly affected the biogeography of freshwater crustaceans in circumpolar areas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Floral fragrances in two closely related fruit fly orchids,Bulbophyllum hortorum and B. macranthoides (Orchidaceae): assortments of phenylbutanoids to attract tephritid fruit fly males

Floral chemical components are important cues used by plants to attract pollinators. One outstanding case is “fruit fly orchids” in the genus of Bulbophyllum to attract their pollinators by releasing characteristic fragrances. Dacini fruit flies are main or exclusive pollinators which are strongly attracted to certain natural chemicals, either methyl eugenol (ME: a phenylpropanoid) or raspberry ketone (RK: a phenylbutanoid). Furthermore, zingerone (ZN: a phenylbutanoid) has been characterized as the attractant for both ME- and RK-sensitive fruit fly species. In the present study, we examined chemical profiles of two closely related Bulbophyllum orchids—B. hortorum, and B. macranthoides subsp. tollenoniferum—distributed in Papua New Guinea and the Southeast Asian countries, respectively. We first observed that RK-sensitive flies were attracted to these orchids by ex situ cultivation in Penang, Malaysia. These Bulbophyllum orchids contained RK and/or ZN as their main floral components. Other than these attractants, multiple phenylbutanoids including potential attractants for RK-sensitive species were identified from these orchids. Therefore, we examined attractiveness of potential phenylbutanoid attractants to an RK-sensitive melon fly, Zeugodacus cucurbitae, using laboratory-reared flies. Furthermore, we analyzed molecular phylogenetic relationships among phenylpropanoid- or phenylbutanoid-producing orchids to see a relation between chemical profiles and phylogenetic classification in the related species.     

Comparative Genomics of Phylogenetically Diverse Unicellular Eukaryotes Provide New Insights into the Genetic Basis for the Evolution of the Programmed Cell Death Machinery

Programmed cell death (PCD) represents a significant component of normal growth and development in multicellular organisms. Recently, PCD-like processes have been reported in single-celled eukaryotes, implying that some components of the PCD machinery existed early in eukaryotic evolution. This study provides a comparative analysis of PCD-related sequences across more than 50 unicellular genera from four eukaryotic supergroups: Unikonts, Excavata, Chromalveolata, and Plantae. A complex set of PCD-related sequences that correspond to domains or proteins associated with all main functional classes—from ligands and receptors to executors of PCD—was found in many unicellular lineages. Several PCD domains and proteins previously thought to be restricted to animals or land plants are also present in unicellular species. Noteworthy, the yeast, Saccharomyces cerevisiae—used as an experimental model system for PCD research, has a rather reduced set of PCD-related sequences relative to other unicellular species. The phylogenetic distribution of the PCD-related sequences identified in unicellular lineages suggests that the genetic basis for the evolution of the complex PCD machinery present in extant multicellular lineages has been established early in the evolution of eukaryotes. The shaping of the PCD machinery in multicellular lineages involved the duplication, co-option, recruitment, and shuffling of domains already present in their unicellular ancestors. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adaptive radiation of photosynthetic physiology in the Hawaiian lobeliads: dynamic photosynthetic responses

Hawaiian lobeliads have radiated into habitats from open alpine bogs to densely shaded rainforest interiors, and show corresponding adaptations in steady-state photosynthetic light responses and associated leaf traits. Shaded environments are not uniformly dark, however, but punctuated by sunflecks that carry most of the photosynthetically active light that strikes plants. We asked whether lobeliads have diversified in their dynamic photosynthetic light responses and how dynamic responses influence daily leaf carbon gain. We quantified gas exchange and dynamic light regimes under field conditions for ten species representing each major Hawaiian sublineage. Species in shadier habitats experienced shorter and less numerous sunflecks: average sunfleck length varied from 1.4 ± 1.7 min for Cyanea floribunda in shaded forest understories to 31.2 ± 2.1 min for Trematolobelia kauaiensis on open ridges. As expected, the rate of photosynthetic induction increased significantly toward shadier sites, with assimilation after 60 s rising from ca. 30% of fully induced rates in species from open environments to 60% in those from densely shaded habitats. Uninduced light use efficiency—actual photosynthesis versus that expected under steady-state conditions—increased from 10 to 70% across the same gradient. In silico transplants—modeling daily carbon gain using one species’ photosynthetic light response in its own and other species’ dynamic light regimes—demonstrated the potential adaptive nature of species differences: understory Cyanea pilosa in its light regimes outperformed gap-dwelling Clermontia parviflora, while Clermontia in its light regimes outperformed Cyanea. The apparent crossover in daily photosynthesis occurred at about the same photon flux density where dominance shifts from Cyanea to Clermontia in the field. Our results further support our hypothesis that the lobeliads have diversified physiologically across light environments in Hawaiian ecosystems and that those shifts appear to maximize the carbon gain of each species in its own environment. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Multiple subfamilies of mariner transposable elements are present in stalk-eyed flies (Diptera: Diopsidae)

The Diopsid stalk-eyed flies are an increasingly well-studied group. Presented here is evidence of the first known transposable elements discovered in these flies. The vertumnana mariner subfamily was identified in the Diopsini tribe, but could not be amplified in species of the Sphyracephalini tribe. PCR screening with degenerate primers revealed that multiple mariner subfamilies are present within the Diopsidae. Most of the sequenced elements appear to be pseudogenes; however two subfamilies are shown to be evolving under purifying selection, raising the possibility that mariner is active in some Diopsid species. Evidence is presented of a possible horizontal transfer event involving an unknown Teleopsis species and the Tephritid fly Bactrocera neohumeralis. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of climate-driven temperature changes on the diversity of freshwater macroinvertebrates

Increasing temperatures due to climate change were found to influence abundance and timing of species in numerous ways. Whereas many studies have investigated climate-induced effects on the phenology and abundance of single species, less is known about climate-driven shifts in the diversity and composition of entire communities. Analyses of long-term data sets provide the potential to reveal such relationships. We analysed time series of entire communities of macrozoobenthos in lakes and streams in Northern Europe. There were no direct linear effects of temperature and climate indices (North Atlantic Oscillation index) on species composition and diversity, but using multivariate statistics we were able to show that trends in average temperature have already had profound impacts on species composition in lakes. These significant temperature signals on species composition were evident even though we analysed comparatively short time periods of 10–15 years. Future climate shifts may thus induce strong variance in community composition. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users. Priority programme of the German Research Foundation—contribution 6.     

An interrupted beta-propeller and protein disorder: structural bioinformatics insights into the N-terminus of alsin

Defects in the human ALS2 gene, which encodes the 1,657-amino-acid residue protein alsin, are linked to several related motor neuron diseases. We created a structural model for the N-terminal 690-residue region of alsin through comparative modelling based on regulator of chromosome condensation 1 (RCC1). We propose that this alsin region contains seven RCC1-like repeats in a seven-bladed beta-propeller structure. The propeller is formed by a double clasp arrangement containing two segments (residues 1–218 and residues 525–690). The 306-residue insert region, predicted to lie within blade 5 and to be largely disordered, is poorly conserved across species. Surface patches of evolutionary conservation probably indicate locations of binding sites. Both disease-causing missense mutations—Cys157Tyr and Gly540Glu—are buried in the propeller and likely to be structurally disruptive. This study aids design of experimental studies by highlighting the importance of construct length, will enhance interpretation of protein–protein interactions, and enable rational site-directed mutagenesis. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Taking stock: inventory of alien species in the Mediterranean sea

573 alien marine metazoan species have been recorded in the Mediterranean Sea. The present checklist is the first to present the species’ native range, presumed mode of introduction, spatial extent, and the date of the first record in each country. The majority of aliens are thermophilic species originating from the Indo-Pacific or Indian Oceans, which have entered the Mediterranean through the Suez Canal. However, the means of introduction differ greatly among the phyla, and the basins of the Mediterranean. The temporal records of the alien species reflect political crises, economic development and scientific interest in studying the phenomenon—in the past two decades on average about 10 alien species new to the Mediterranean are recorded annually. Many have established durable populations and extended their range: 125 alien species have been recorded from four or more countries. The possible impacts of regulatory instruments and environmental management options are examined. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A study of the salt lakes and salt springs of Eyre Peninsula, South Australia

An 18-month-study of 40 saline wetlands, ranging from 6 to 336 g l⁻¹, on the west and southern coasts of Eyre Peninsula yielded 88 species of invertebrates, some aquatic plants and a fish. The invertebrates are taxonomically diverse and include 38 crustaceans, 28 insects, 12 molluscs and significantly an aquatic spider, a nemertean, two polychaetes, two sea anemones, a sponge and a bryzoan. Most were tolerant of wide fluctuations in salinity, there being 51 halobionts, 21 halophils and only 16 salt-tolerant freshwater species. Many invertebrates are restricted to the thalassic springs where marine molluscs dominated. Athalassic wetlands were dominated by crustaceans and were of two basic types—coastal and continental. There is evidence of the former evolving biologically into the later, and for some lakes to be still in transition. There is also evidence of increasing salinity in recent decades and already two lakes exhibit severe secondary salinity. Like other salt lakes in Australia the fauna is regionally distinctive. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest Editors: J. John & B. Timms Salt Lake Research: Biodiversity and Conservation—Selected papers from the 9th Conference of the International Society for Salt Lake Research     

Understory plant diversity is related to higher variability of vegetative mobility of coexisting species

Theoretical studies claim that if co-occurring species have very different mobilities this will result in greater small-scale species richness, but empirical evidence is still lacking. We measured horizontal vegetative mobility (VM) of 48 herbaceous understory species and estimated small-scale species richness in early and late successional boreonemoral herb-rich coniferous forests in central Estonia. VM of erosulate growth forms was significantly higher than that of hemi-rosette and rosette growth forms. Erosulate species exhibited higher mobility in young stands, but their relative and total cover was considerably higher in old stands. Local plant richness (in 1 × 1 m plots) correlated positively with the variability of VM of species in a plot—larger differences in VM resulted in a higher number of coexisting species. Our results thus suggest that species differences in VM can contribute to small-scale coexistence by providing different ways to colonise empty space. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Grasshopper (Orthoptera: Acrididae) communities respond to fire,bison grazing and weather in North American tallgrass prairie: a long-term study

Because both intrinsic and extrinsic factors influence insect population dynamics, operating at a range of temporal and spatial scales, it is difficult to assess their contributions. Long-term studies are ideal for assessing the relative contributions of multiple factors to abundance and community dynamics. Using data spanning 25 years, we investigate the contributions of weather at annual and decadal scales, fire return interval, and grazing by bison to understand the dynamics of abundance and community composition in grasshopper assemblages from North American continental grassland. Each of these three primary drivers of grassland ecosystem dynamics affects grasshopper population and community dynamics. Negative feedbacks in abundances, as expected for regulated populations, were observed for all feeding guilds of grasshoppers. Abundance of grasshoppers did not vary in response to frequency of prescribed burns at the site. Among watersheds that varied with respect to controlled spring burns and grazing by bison, species composition of grasshopper assemblages responded significantly to both after 25 years. However, after more than 20 years of fire and grazing treatments, the number of years since the last fire was more important than the managed long-term fire frequency per se. Yearly shifts in species composition (1983–2005), examined using non-metric multidimensional scaling and fourth-corner analysis, were best explained by local weather events occurring early in grasshopper life cycles. Large-scale patterns were represented by the Palmer Drought Severity Index and the North Atlantic Oscillation (NAO). The NAO was significantly correlated with annual mean frequencies of grasshoppers, especially for forb- and mixed-feeding species. Primary grassland drivers—fire, grazing and weather—contributing both intrinsic and extrinsic influences modulate long-term fluctuations in grasshopper abundances and community taxonomic composition. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ontogenetic development in architecture and biomass allocation of 13 rattan species in Indonesia

Rattans are climbing, nonbranching palms with diverse growth forms ranging from stems that remain at the forest floor to stems that reach the canopy. We analyzed changes in architecture and biomass allocation during ontogenetic development of 13 Indonesian species in the genera Calamus, Ceratolobus, Daemonorops, Korthalsia, and Plectocomiopsis. Species included both nonclimbers (<5 m tall maximum) and climbers (10 to 50 m tall). Nonclimbers retain a rosette form—that is, thick, short internodes with length/internode diameter ratio <10 but with long petioles—throughout their lives. Leaf and internode shapes of climbers at the early stage resemble those of nonclimbers, but internodes later become longer (internode length/diameter ratio ≥10) and petioles become shorter and developed one of two kinds of climbing organs at the adult stage. These developmental changes reduce self-shading within the crown. Some climbers have dwarf blades at the early stage and skip the rosette form. Principal component analysis of biomass allocation indicated that growth strategies to attain the adult stage are diverse. These results suggest that rattans reach maturity at different phases along a series of ontogenetic development stages and generate diverse growth forms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Eutrophication and fragmentation are related to species’ rate of decline but not to species rarity: results from a functional approach

Due to ubiquitous eutrophication and fragmentation, many plant species are actually threatened in Europe. Most ecosystems face an overall nutrient input leading to changes in species composition. Fragmentation is effectively influencing species survival. We investigate if two different measures of species performance of 91 calcareous grassland species–rate of decline and rarity—are related to comparable traits and hence processes. On the one hand we expected that species rate of decline is mainly determined by the processes of eutrophication and fragmentation. On the other hand we hypothesized that the importance of site characteristics may overwhelm the effect of eutrophication and fragmentation for species rarity. Hence, we compared persistence traits responding to eutrophication, dispersal traits being related to fragmentation and ecological site factors for decreasing and increasing species and for rare and common species. The results suggest that increasing species had better means of long-distance dispersal and were more competitive than decreasing species. In contrast, there were hardly any differences in traits between rare and common species, but site characteristics were related to species rarity. Rare species were in the main those with ecological preferences for warm, dry, light and nutrient poor conditions. This study may represent a basis for the assessment of plant species threat. Applying the deduced knowledge about the life history of decreasing versus increasing species to habitat-scale approaches it is possible to predict which species may become threatened in the future simply from the combination of their trait values. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Modularity and Complete Natural Homeoses in Cervical Vertebrae of Extant and Extinct Penguins (Aves: Sphenisciformes)

The cervical system of extant penguins (Aves: Sphenisciformes) is organised into morphological modules, each with its biomechanical function. Indeed, for these marine pelagic birds to acquire hydrodynamic morphology, the folding of the neck is essential. Despite a common general structure, the cervical vertebrae exhibit morphological differences depending on their positioning. These characteristics are identified as apparent cases of complete natural homeotic transformations—therefore, the composition of some modules varies. Two types of complete cervical homeoses are identified between species, but the second type can also occur within some species when the post hatching development is considered. The fossil material analysed here makes it apparent that the two modular configurations characterising the anterior part of the neck—a consequence of the first homeosis—existed 36 My and 25 My ago, for one, and circa 10 My ago, for the other. These comparisons also reveal a clear differentiation in vertebral features between the fossil species of the Oligocene–Miocene ages and the more recent and extant penguins. Ultimately, these observations make the proposal of a hypothesis in relation to the ontogenetic influence of Hox genes, and their regulators, based on the changes observed in the cervical segment of Sphenisciformes.