The epithelial sodium channel in the Australian lungfish,Neoceratodus forsteri (Osteichthyes: Dipnoi)

Epithelial sodium channel (ENaC) is a Na⁺-selective, aldosterone-stimulated ion channel involved in sodium transport homeostasis. ENaC is rate-limiting for Na⁺ absorption in the epithelia of osmoregulatory organs of tetrapods. Although the ENaC/degenerin gene family is proposed to be present in metazoans, no orthologues or paralogues for ENaC have been found in the genome databases of teleosts. We studied full-length cDNA cloning and tissue distributions of ENaCα, β and γ subunits in the Australian lungfish, Neoceratodus forsteri, which is the closest living relative of tetrapods. Neoceratodus ENaC (nENaC) comprised three subunits: nENaCα, β and γ proteins. The nENaCα, β and γ subunits are closely related to amphibian ENaCα, β and γ subunits, respectively. Three ENaC subunit mRNAs were highly expressed in the gills, kidney and rectum. Amiloride-sensitive sodium current was recorded from Xenopus oocytes injected with the nENaCαβγ subunit complementary RNAs under a two-electrode voltage clamp. nENaCα immunoreactivity was observed in the apical cell membrane of the gills, kidney and rectum. Thus, nENaC may play a role in regulating sodium transport of the lungfish, which has a renin–angiotensin–aldosterone system. This is interesting because there may have been an ENaC sodium absorption system controlled by aldosterone before the conquest of land by vertebrates.     

Latisemenia longshania,gen. et sp. nov., a new Late Devonian seed plant from China

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.     

Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China

Background and Aims With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns.Methods Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix.Key Results A new fern-like taxon, Shougangia bella gen. et sp. nov., is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia.Conclusions Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO₂ concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.     

Northward expansion of a marine parasite: Testing the role of temperature adaptation

The known range of the eastern oyster (Crassostrea virginica) parasite, Perkinsus marinus, expanded into the northeastern United States in the early 1990s. We used both in vitro and in vivo data to test the hypothesis that the northward expansion was associated with a low-temperature adapted strain of the parasite. In vitro proliferation of nine P. marinus isolates from three geographic sites, Massachusetts and New Jersey in the new range, and South Carolina in the historic southern range, was measured at seven temperatures (5 to 35 °C) using a tetrazolium blue dye assay. We wanted to determine if there were between- and within-geographic location differences in the P. marinus proliferation rate, and if so, whether they were associated with temperature. We found no evidence of low-temperature adaptation based on the fact that net proliferation rates for isolates from all three geographic locations were similar at temperatures from 5 to 20 °C. On the other hand, at temperatures of 25 to 35 °C, the South Carolina isolates exhibited higher proliferation rates than the northern isolates suggesting possible high-temperature adaptation of parasite strains that are routinely exposed to higher temperatures. Although there was significant within-location variation among isolates, the data tended to group together by geographic location supporting the hypothesis that there is an important regional component to the proliferation rate of P. marinus isolates. A survey of published data showed that the temperature at which in vivo proliferation was first observed in oysters at sites from the Gulf of Mexico to Massachusetts was typically between 20 and 23 °C with no evidence of a geographic cline. These results lend support to the hypothesis that the recent warming trend in the northeastern US is the most likely explanation for the P. marinus range extension.     

Discussion of the Early Devonian rhynchonellid brachiopod genus Borealirhynchia Su, 1976

The Early Devonian brachiopod genus Borealirhynchia was established by Su, 1976. Phylum Brachiopoda (Cambrian to Devonian). In: Paleontological Atlas of North China, Inner Mongolia Volume 1. Geological Publishing House Beijing, pp. 155-227 (in Chinese). It has not been previously described in detail, nor have transverse serial sections of the internal features of Borealirhynchia delerensis, the type species, been published. In this paper Borealirhynchia, along with a few species previously assigned to it, is analysed and discussed. Detailed transverse serial sections of the internal features of B.? lata Su, 1976, are provided, based on well-preserved specimens collected from the Lower Devonian strata of Dong Ujimqin Qi, northeastern Inner Mongolia. Borealirhynchia? gigantea Su, 1976 and Latonotoechia multicosta Su, 1976 are considered to be synonyms of B.? lata Su with the same external and internal characters present in all three species. Some Lower Devonian strata, in which Borealirhynchia was found and reported, from Dong Ujimqin Qi, northeastern Inner Mongolia, are fully described.     

Experimental evidence of bark beetle adaptation to a fungal symbiont

The importance of symbiotic microbes to insects cannot be overstated; however, we have a poor understanding of the evolutionary processes that shape most insect–microbe interactions. Many bark beetle (Coleoptera: Curculionidae, Scolytinae) species are involved in what have been described as obligate mutualisms with symbiotic fungi. Beetles benefit through supplementing their nutrient‐poor diet with fungi and the fungi benefit through gaining transportation to resources. However, only a few beetle–fungal symbioses have been experimentally manipulated to test whether the relationship is obligate. Furthermore, none have tested for adaptation of beetles to their specific symbionts, one of the requirements for coevolution. We experimentally manipulated the western pine beetle–fungus symbiosis to determine whether the beetle is obligately dependent upon fungi and to test for fine‐scale adaptation of the beetle to one of its symbiotic fungi, Entomocorticium sp. B. We reared beetles from a single population with either a natal isolate of E. sp. B (isolated from the same population from which the beetles originated), a non‐natal isolate (a genetically divergent isolate from a geographically distant beetle population), or with no fungi. We found that fungi were crucial for the successful development of western pine beetles. We also found no significant difference in the effects of the natal and non‐natal isolate on beetle fitness parameters. However, brood adult beetles failed to incorporate the non‐natal fungus into their fungal transport structure (mycangium) indicating adaption by the beetle to particular genotypes of symbiotic fungi. Our results suggest that beetle–fungus mutualisms and symbiont fidelity may be maintained via an undescribed recognition mechanism of the beetles for particular symbionts that may promote particular associations through time.     

Host-specific adaptation governs the interaction of the marine diatom,Pseudo-nitzschia and their microbiota

The association of phytoplankton with bacteria is ubiquitous in nature and the bacteria that associate with different phytoplankton species are very diverse. The influence of these bacteria in the physiology and ecology of the host and the evolutionary forces that shape the relationship are still not understood. In this study, we used the Pseudo-nitzschia–microbiota association to determine (1) if algal species with distinct domoic acid (DA) production are selection factors that structures the bacterial community, (2) if host-specificity and co-adaptation govern the association, (3) the functional roles of isolated member of microbiota on diatom–hosts fitness and (4) the influence of microbiota in changing the phenotype of the diatom hosts with regards to toxin production. Analysis of the pyrosequencing-derived 16S rDNA data suggests that the three tested species of Pseudo-nitzschia, which vary in toxin production, have phylogenetically distinct bacterial communities, and toxic Pseudo-nitzschia have lower microbial diversity than non-toxic Pseudo-nitzschia. Transplant experiments showed that isolated members of the microbiota are mutualistic to their native hosts but some are commensal or parasitic to foreign hosts, hinting at co-evolution between partners. Moreover, Pseudo-nitzschia host can gain protection from algalytic bacteria by maintaining association with its microbiota. Pseudo-nitzschia also exhibit different phenotypic expression with regards to DA production, and this depends on the bacterial species with which the host associates. Hence, the influences of the microbiota on diatom host physiology should be considered when studying the biology and ecology of marine diatoms.     

The problematic cemented Devonian brachiopod Schuchertellopsis durbutensis Maillieux, 1939

The Devonian cemented brachiopod Schuchertellopsis durbutensis has proved difficult to classify and its possible taxonomic relationships are unknown. Morphologically Schuchertellopsis resembles more closely members of the Orthotetidina than the Davidsoniidina. Examination of the shell structure, a key diagnostic feature of the Orthotetidina, shows that Schuchertellopsis has the cross laminar secondary shell typical of all orthotetidines. However, the presence of both pseudopunctate and incipient an extropunctate fabric within the ventral valve is unique amongst orthotetidine brachiopods and is thought to represent a phase of shell fabric experimentation. Schuchertellopsis probably fits most comfortably within the Schuchertellidae, and is the earliest representative of that family.     

A reconsideration of Dimeripteris cornuta SCHWEITZER and CAI, a diminutive fossil plant from the Middle Devonian of Yunnan, China

Small plant fragments previously attributed to Dimeripteris cornuta SCHWEITZER and CAI and new specimens are described from the Middle Devonian (Givetian) Xichong Formation of Wuding, Yunnan Province, southwest China. A predominantly trifurcate branching pattern is recognized in lower orders of branching. Erect fusiform sporangia are borne in pairs terminally on a short pedicel on up to three times dichotomously-branching fertile unit. The beaked tips of the paired sporangia point outwards. This plant is distinguished from known plants by the trifurcate branching pattern and sporangia morphology, but is of problematic affinity. The plant is named Tauritheca cornuta (SCHWEITZER and CAI) WANG and BERRY nov. gen. and nov. comb.     

An inducible morphological defence is a passive by-product of behaviour in a marine snail

Many organisms have evolved inducible defences in response to spatial and temporal variability in predation risk. These defences are assumed to incur large costs to prey; however, few studies have investigated the mechanisms and costs underlying these adaptive responses. I examined the proximate cause of predator-induced shell thickening in a marine snail (Nucella lamellosa) and tested whether induced thickening leads to an increase in structural strength. Results indicate that although predators (crabs) induce thicker shells, the response is a passive by-product of reduced feeding and somatic growth rather than an active physiological response to predation risk. Physical tests indicate that although the shells of predator-induced snails are significantly stronger, the increase in performance is no different than that of snails with limited access to food. Increased shell strength is attributable to an increase in the energetically inexpensive microstructural layer rather than to material property changes in the shell. This mechanism suggests that predator-induced shell defences may be neither energetically nor developmentally costly. Positive correlations between antipredator behaviour and morphological defences may explain commonly observed associations between growth reduction and defence production in other systems and could have implications for the evolutionary potential of these plastic traits.     

Plasticity predicts evolution in a marine alga

Under global change, populations have four possible responses: ‘migrate, acclimate, adapt or die’ (Gienapp et al. 2008 Climate change and evolution: disentangling environmental and genetic response. Mol. Ecol. 17, 167–178. (doi:10.1111/j.1365-294X.2007.03413.x)). The challenge is to predict how much migration, acclimatization or adaptation populations are capable of. We have previously shown that populations from more variable environments are more plastic (Schaum et al. 2013 Variation in plastic responses of a globally distributed picoplankton species to ocean acidification. Nature 3, 298–230. (doi:10.1038/nclimate1774)), and here we use experimental evolution with a marine microbe to learn that plastic responses predict the extent of adaptation in the face of elevated partial pressure of CO₂ (pCO₂). Specifically, plastic populations evolve more, and plastic responses in traits other than growth can predict changes in growth in a marine microbe. The relationship between plasticity and evolution is strongest when populations evolve in fluctuating environments, which favour the evolution and maintenance of plasticity. Strikingly, plasticity predicts the extent, but not direction of phenotypic evolution. The plastic response to elevated pCO₂ in green algae is to increase cell division rates, but the evolutionary response here is to decrease cell division rates over 400 generations until cells are dividing at the same rate their ancestors did in ambient CO₂. Slow-growing cells have higher mitochondrial potential and withstand further environmental change better than faster growing cells. Based on this, we hypothesize that slow growth is adaptive under CO₂ enrichment when associated with the production of higher quality daughter cells.     

A New Eurypterid (Chelicerata: Eurypterida) from the Upper Devonian Gogo Formation of Western Australia, With A Review of the Rhenopteridae

A new eurypterid, Rhenopterus waterstoni sp. nov., is described from the Gogo Formation (Frasnian, Upper Devonian) of Western Australia. This species is distinguished from related forms by the tuberculation of the anteriormost tergite and crenulated posterior margins of the carapace and opisthosomal segments. It is the only eurypterid specimen known from the Gogo Formation, the most complete eurypterid from Australia, and also the youngest representative of Rhenopterus in the fossil record. Structures retrieved from between the prosomal-opisthosomal juncture comprise polygonal tubes 30–40 μm in diameter, which are interpreted as sarcomeral sheaths of muscular tissue. Rhenopterus is reviewed: R. latus Størmer, 1936 is synonymized with R. diensti Størmer, 1936 as it is here recognized as a female sexual dimorph; R. maccarthyi (Kjellesvig-Waering, 1934) is an orthocone nautiloid.     

New evidence for the Early Devonian age of the Jauf Formation in northern Saudi Arabia

Well-preserved Early Devonian continental and sparse marine palynomorph assemblages recovered from an exploration borehole located in northern Saudi Arabia are reported here and add to the understanding of Arabian Devonian stratigraphy. The units examined are the conventionally cored lower Murayr, upper Hammamiyat and Sha’iba Members of the Jauf Formation. A comparison with data sets from elsewhere on the Arabian Plate suggests that the units examined are Emsian in age. The age of the Murayr and Hammamiyat boundary interval is revised, based on the occurrence of Dibolisporites eifeliensis in northern Saudi Arabia, and placed in the Rhabdosporites minutus (Min) Interval Zone of the Emphanisporites foveolatus-Verrucosisporites dubia (FD) Oppel Zone. A new miospore, Apiculiretusispora densa with distinctive interradial thickenings, is described.     

Flower colour adaptation in a mimetic orchid

Although the tremendous variability in floral colour among angiosperms is often attributed to divergent selection by pollinators, it is usually difficult to preclude the possibility that floral colour shifts were driven by non-pollinator processes. Here, we examine the adaptive significance of flower colour in Disa ferruginea, a non-rewarding orchid that is thought to attract its butterfly pollinator by mimicking the flowers of sympatric nectar-producing species. Disa ferruginea has red flowers in the western part of its range and orange flowers in the eastern part--a colour shift that we hypothesized to be the outcome of selection for resemblance to different local nectar-producing plants. Using reciprocal translocations of red and orange phenotypes as well as arrays of artificial flowers, we found that the butterfly Aeropetes tulbaghia, the only pollinator of the orchid, preferred both the red phenotype and red artificial flowers in the west where its main nectar plant also has red flowers, and both the orange phenotype and orange artificial flowers in the east, where its main nectar plant has orange flowers. This phenotype by environment interaction demonstrates that the flower colour shift in D. ferruginea is adaptive and driven by local colour preference in its pollinator.     

The chitin synthase involved in marine bivalve mollusk shell formation contains a myosin domain

Chitin is a key component in mollusk nacre formation. However, the enzyme complex responsible for chitin deposition in the mollusk shell remained unknown. We cloned and characterized the chitin synthase of the marine bivalve mollusk Atrina rigida. We present here the first chitin synthase sequence from invertebrates containing an unconventional myosin motor head domain. We further show that a homologous gene for chitin synthase is expressed in the shell forming tissue of larval Mytilus galloprovincialis even in early embryonic stages. The new data presented here are the first clear-cut indication for a functional role of cytoskeletal forces in the precisely controlled mineral deposition process of mollusk shell biogenesis.     

Vampires in the oceans: predatory cercozoan amoebae in marine habitats

Vampire amoebae (vampyrellids) are predators of algae, fungi, protozoa and small metazoans known primarily from soils and in freshwater habitats. They are among the very few heterotrophic naked, filose and reticulose protists that have received some attention from a morphological and ecological point of view over the last few decades, because of the peculiar mode of feeding of known species. Yet, the true extent of their biodiversity remains largely unknown. Here we use a complementary approach of culturing and sequence database mining to address this issue, focusing our efforts on marine environments, where vampyrellids are very poorly known. We present 10 new vampyrellid isolates, 8 from marine or brackish sediments, and 2 from soil or freshwater sediment. Two of the former correspond to the genera Thalassomyxa Grell and Penardia Cash for which sequence data were previously unavailable. Small-subunit ribosomal DNA analysis confirms they are all related to previously sequenced vampyrellids. An exhaustive screening of the NCBI GenBank database and of 454 sequence data generated by the European BioMarKs consortium revealed hundreds of distinct environmental vampyrellid sequences. We show that vampyrellids are much more diverse than previously thought, especially in marine habitats. Our new isolates, which cover almost the full phylogenetic range of vampyrellid sequences revealed in this study, offer a rare opportunity to integrate data from environmental DNA surveys with phenotypic information. However, the very large genetic diversity we highlight within vampyrellids (especially in marine sediments and soils) contrasts with the paradoxically low morphological distinctiveness we observed across our isolates.     

Local adaptation within a hybrid species

Ecological divergence among populations may be strongly influenced by their genetic background. For instance, genetic admixture through introgressive hybridization or hybrid speciation is likely to affect the genetic variation and evolvability of phenotypic traits. We studied geographic variation in two beak dimensions and three other phenotypic traits of the Italian sparrow (Passer italiae), a young hybrid species formed through interbreeding between house sparrows (P. domesticus) and Spanish sparrows (P. hispaniolensis). We found that beak morphology was strongly influenced by precipitation regimes and that it appeared to be the target of divergent selection within Italian sparrows. Interestingly, however, the degree of parental genetic contribution in the hybrid species had no effect on phenotypic beak variation. Moreover, beak height divergence may mediate genetic differentiation between populations, consistent with isolation-by-adaptation within this hybrid species. The study illustrates how hybrid species may be relatively unconstrained by their admixed genetic background, allowing them to adapt rapidly to environmental variation.