Physiological integration impacts nutrient use and stoichiometry in three clonal plants under heterogeneous habitats

Physiological integration facilitates clonal plants to deal with heterogeneous resources. However, little is known about how nutrient patchiness affects its use and stoichiometry in clonal plants. We conducted an experiment with Cynodon dactylon, Glechoma longituba, and Potentilla reptans to address the effects of physiological integration on nutrient use efficiency and N:P ratios. For C. dactylon, the effects of nutrient patchiness on N use efficiency (NUE), P use efficiency (PUE), and N:P ratio were stronger in daughter ramets than in parent ramets; for G. longituba, nutrient patchiness affected PUE and N:P ratio of parent and daughter ramets, but not NUE; for P. reptans, nutrient patchiness decreased NUE, PUE, and N:P ratio, regardless of parent or daughter ramets. PUE was associated with N:P ratios in three clonal plants and this association of NUE with N:P ratios varied with species. Our findings suggest that physiological integration alters nutrient use efficiency and N:P ratios of clonal plants under patchy nutrients and that these effects are linked to clonal species identity.     

Quantitative nuclear DNA differences associated with genome evolution in Guizotia (Compositae)

The present communication deals with 2C nuclear genome size variation in a fairly small genus Guizotia. Twenty-four accessions belonging to six species, out of seven known, were analysed in order to elucidate the extent of DNA variation both at an intra—as well as interspecific level. At the intraspecific level none of the species exhibited significant differences in their genome size. Between the species, the 2C DNA amounts ranged from 3.61 pg in G. reptans to 11.37 pg in G. zavattarii; over three-fold DNA variation is evident. Apparently these interspecific DNA differences have been achieved independent of the numerical chromosomal change(s), as all the Guizotias share a common chromosome number 2n=2x=30. The cultivated oilseed crop, G. abyssinica (7.57 pg), has accommodated nearly 78% extra DNA in its chromosome complement during the evolutionary time scale of its origin and domestication from the wild progenitor G. schimperi (4.25 pg). The extent of genomic DNA difference(s) between the species has been discussed in the light of their interrelationships and diversity.     

Microcladia exserta sp. nov. (Ceramiaceae,Rhodophyta) from the East coast of South Africa

Microcladia exserta, a new species of the red algal genus Microcladia Greville (Ceramiaceae, Ceramiales), is described from the Natal coast of South Africa. This small, creeping alga, an epiphyte on the coralline alga Amphiroa anceps (Lamarck) Decaisne, is distinguished from other species in the genus by the following combination of characteristics: the prostrate habit, the exerted position of the tetrasporangia, and the presence in the cortex of numerous and conspicuous vesicle cells. Evidence is presented to demonstrate that the criterion of an erect habit in Microcladia vs. a prostrate habit in Herpochondria used to separate these genera is not sound.     

Habitat quality matters for the distribution of an endangered leaf beetle and its egg parasitoid in a fragmented landscape

Fragmentation, deterioration, and loss of habitat patches threaten the survival of many insect species. Depending on their trophic level, species may be differently affected by these factors. However, studies investigating more than one trophic level on a landscape scale are still rare. In the present study we analyzed the effects of habitat size, isolation, and quality for the occurrence and population density of the endangered leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) and its egg parasitoid, the hymenopteran wasp Foersterella reptans Nees (Hymenoptera: Tetracampidae). C. canaliculata is strictly monophagous on meadow sage (Salvia pratensis), while F. reptans can also parasitize other hosts. Both size and isolation of habitat patches strongly determined the occurrence of the beetle. However, population density increased to a much greater extent with increasing host plant density ( = habitat quality) than with habitat size. The occurrence probability of the egg parasitoid increased with increasing population density of C. canaliculata. In conclusion, although maintaining large, well-connected patches with high host plant density is surely the major conservation goal for the specialized herbivore C. canaliculata, also small patches with high host plant densities can support viable populations and should thus be conserved. The less specialized parasitoid F. reptans is more likely to be found on patches with high beetle density, while patch size and isolation seem to be less important.     

PHYLOGENETIC RELATIONSHIPS AMONG LINEAGES OF THE CERAMIACEAE (CERAMIALES,RHODOPHYTA) BASED ON NUCLEAR SMALL SUBUNIT rDNA SEQUENCE DATA1

Phylogenetic relationships among 69 species of the Ceramiales (51 Ceramiaceae, six Dasyaceae, seven Delesseriaceae, and five Rhodomelaceae) were determined based on nuclear SSU rDNA sequence data. We resolved five strongly supported but divergent lineages among the included Ceramiaceae: (i) the genus Inkyuleea, which weakly joins other orders of the Rhodymeniophycidae rather than the Ceramiales in our analyses; (ii) the tribe Spyridieae, which is sister to the remainder of the included ceramialean taxa; (iii) the subfamily Ceramioideae, weakly including the tribe Warrenieae; (iv) the subfamily Callithamnioideae; and (v) the subfamily Compsothamnioideae, which emerges as sister to the Dasyaceae/Delesseriaceae/Rhodomelaceae complex, thus rendering the Ceramiaceae sensu lato unequivocally paraphyletic, as has been argued separately on anatomical grounds by Kylin and Hommersand. Our data support a restricted concept of the Ceramiaceae that includes only one of the five lineages (Ceramioideae) that we have resolved. In addition to failing to ally with the Ceramiales in our molecular analyses, species of Inkyuleea differ substantially from other Ceramiaceae sensu lato in details of pre‐ and postfertilization development. The genus Inkyuleea is here assigned to the Inkyuleeaceae fam. nov., which we provisionally retain in the Ceramiales. Species of Spyridia also differ from the remaining Ceramiaceae in their postfertilization development, and, in light of our molecular data, the genus Spyridia is assigned to the Spyridiaceae. The Callithamnioideae is strongly monophyletic (100% in all analyses), which, in combination with key anatomical differences, supports elevation to family status for this lineage as the Callithamniaceae. Similarly, the Compsothamnioideae is solidly monophyletic in our molecular trees and has a unique suite of defining anatomical characters that supports family status for a complex that we consider to include the tribes Compsothamnieae, Dasyphileae, Griffithsieae, Monosporeae, Ptiloteae, Spermothamnieae, Sphondylothamnieae, Spongoclonieae, and Wrangelieae, for which the reinstated family name Wrangeliaceae is available.     

Studies on marine algae of the British Isles. 10. The genus Rhodymenia

Antarcticothamnion polysporum gen. et sp. nov. (Rhodophyceae: Ceramiaceae) is described from the South Shetland Is. and the Antarctic Peninsula. It differs from all previously described Ceramiaceae in vegetative structure: an indeterminate apex, which divides by alternating oblique septa, produces simple alternate-distichous determinate branches, while indeterminate branches are initiated on main axes in a verticillate arrangement. Reproductive structures are borne on modified indeterminate branch systems. Sporangia are polyhedrally divided. Spermatangia are formed in loose heads. Procarps are borne near the apex, but not always on the subapical cell, of a fertile branch. Two auxiliary cells may participate in the formation of a carposporophyte. A fusion cell is lacking and most cells of the gonimoblast develop into carposporangia, which are binucleate.

The suite of morphological characters exhibited by Antarcticothamnion sets this genus sufficiently apart from all previously described Ceramiaceae to warrant placement in its own tribe, Antarcticothamnieae trib. nov. It shares important features, however, with Callithamnieae, Compsothamnieae, and Ptiloteae.

Apical division is reviewed in the various tribes of Ceramiaceae. It is concluded that a pattern of alternating oblique septa is correlated with transitory or chronic structural imbalance resulting from a particular combination of timing and spatial sequence in the initiation and development of branches. The distribution of oblique apical division within Ceramiaceae is strongly correlated with features generally considered to be important at the level of tribe.

Phylogenetic relationships within Ptiloteae, a tribe characterized inter alia by oblique apical division, are suggested from an analysis of branching patterns. The validity of Falklandiella as a genus distinct from Dasyptilon is emphasized. Gymnothamnion and Tokidaea are removed from Ptiloteae, but without being assigned to another tribe. Tanakaella is removed from Sphondylothamnieae to Compsothamnieae, while Mazoyerella is removed from Compsothamnieae to Spermothamnieae.

Spongoclonium orthocladum A. et E. S. Gepp is discussed as a possible species of Antarcticothamnion, while an undescribed alga from the South Orkney Is. is definitely indicated as a second species of the genus.     

Changes in chloroplast ultrastructure in some high-alpine plants: adaptation to metabolic demands and climate?

Summary. The cytology of leaf cells from five different high-alpine plants was studied and compared with structures in chloroplasts from the typical high-alpine plant Ranunculus glacialis previously described as having frequent envelope plus stroma protrusions. The plants under investigation ranged from subalpine/alpine Geum montanum through alpine Geum reptans, Poa alpina var. vivipara, and Oxyria digyna to nival Cerastium uniflorum and R. glacialis. The general leaf structure (by light microscopy) and leaf mesophyll cell ultrastructure (by transmission electron microscopy [TEM]) did not show any specialized structures unique to these mountain species. However, chloroplast protrusion formation could be found in G. reptans and, to a greater extent, in O. digyna. The other species exhibited only a low percentage of such chloroplast structural changes. Occurrence of protrusions in samples of G. montanum and O. digyna growing in a mild climate at about 50 m above sea level was drastically reduced. Serial TEM sections of O. digyna cells showed that the protrusions can appear as rather broad and long appendices of plastids, often forming pocketlike structures where mitochondria and microbodies are in close vicinity to the plastid and to each other. It is suggested that some high-alpine plants may form such protrusions to facilitate fast exchange of molecules between cytoplasm and plastid as an adaptation to the short, often unfavorable vegetation period in the Alps, while other species may have developed different types of adaptation that are not expressed in ultrastructural changes of the plastids. Correspondence and reprints: Institute of Botany, University of Innsbruck, Sternwartestrasse 15, 6020 Innsbruck, Austria.     

Comparative cytogenetics of six Indo‐Pacific moray eels (Anguilliformes: Muraenidae) by chromosomal banding and fluorescence in situ hybridization

A comparative cytogenetic analysis, using both conventional staining techniques and fluorescence in situ hybridization, of six Indo‐Pacific moray eels from three different genera (Gymnothorax fimbriatus, Gymnothorax flavimarginatus, Gymnothorax javanicus, Gymnothorax undulatus, Echidna nebulosa and Gymnomuraena zebra), was carried out to investigate the chromosomal differentiation in the family Muraenidae. Four species displayed a diploid chromosome number 2n = 42, which is common among the Muraenidae. Two other species, G. javanicus and G. flavimarginatus, were characterized by different chromosome numbers (2n = 40 and 2n = 36). For most species, a large amount of constitutive heterochromatin was detected in the chromosomes, with species‐specific C‐banding patterns that enabled pairing of the homologous chromosomes. In all species, the major ribosomal genes were localized in the guanine‐cytosine‐rich region of one chromosome pair, but in different chromosomal locations. The (TTAGGG)_n telomeric sequences were mapped onto chromosomal ends in all muraenid species studied. The comparison of the results derived from this study with those available in the literature confirms a substantial conservation of the diploid chromosome number in the Muraenidae and supports the hypothesis that rearrangements have occurred that have diversified their karyotypes. Furthermore, the finding of two species with different diploid chromosome numbers suggests that additional chromosomal rearrangements, such as Robertsonian fusions, have occurred in the karyotype evolution of the Muraenidae.     

Nuclear crystals, lampbrush-chromosome-like structures, and perinuclear cytoskeletal elements associated with nuclear fragmentation in characean internodal cells

Summary This article gives a survey of nucleus-associated structures and inclusions in a diverse range of characean algae includingChara braunii Gm.,Chara corallina Klein ex Willd., em. R.D.W.,Nitella cristata A.Br., em. R.D.W.,Nitella flexilis (L.) Ag.,Nitella furcata (Roxb. ex Bruz.) Ag. em. R.D.W.,Nitella hyalina (DC.) Ag.,Nitella pseudoflabellata A.Br., em. R.D.W.,Nitella pseudoflabellata var.imperialis T.F.A.,Nitella translucens var.axillaris (A.Br.) R.D.W. andNitellopsis obtusa (Desv. in Lois.) J.Gr. Lampbrushchromosome-like structures were found in nuclei ofNitella flexilis andNitellopsis obtusa and seem to be involved in the distribution of genetic material during nuclear fragmentation. Intranuclear tubular crystals of unknown protein composition were present in all species, especially in young, elongating cells, and could be important for establishing the main axis of the nuclei. Spindle-shaped protein crystals that originate in the nucleus and are released into the cytoplasm upon nuclear degeneration were observed in branchlet internodal cells of one population ofNitella flexilis. Perinuclear microtubules were present in all species, but perinuclear actin fibrils were hitherto only found in mostNitella species and inNitellopsis obtusa. None of these nucleus-associated structures seems to be responsible for the formation of constrictions leading to nuclear fragmentation. These constrictions were perpendicular to the main axis of the nucleus and symmetrical in theNitella species but asymmetric inC. braunii, C. corallina, and inNitellopsis obtusa. Statistical analysis of nuclear size, number and constriction sites indicate that fragmentation is a nonsynchronous process independent of the light-dark cycle.Abbreviations CLSM confocal laser scanning microscopy - DAPI 4,6-diamidino-2-phenylindole - DIC Nomarski differential interference contrast - LCLS lampbrush chromosome-like structure(s) Dedicated to Professor Walter Gustav Url on the occasion of his 70th birthday     

Comparative studies on the circulatory system of the compound ascidians,Botryllus, Botrylloides and Symplegma

The circulatory systems of four polystyelids, Botryllus schlosseri, B. primigenus, Botrylloides violaceus and Symplegma reptans, were compared. The palleal buds are connected to the parent zooid by a peduncle and to the colonial vascular system by connecting vessels. The peduncle of S. reptans disappears at an earlier stage of bud development than in B. primigenus; it survives the dissolution of the parent zooid in B. schlosseri and B. violaceus. The connecting vessel is formed by anastomosis between an epidermal outgrowth from the bud and a neighboring colonial vessel, and is characterized by the presence of a sphincter. The number of connecting vessels formed in a palleal bud is three in S. reptans, two in B. primigenus and one each in B. schlosseri and B. violaceus. In each species, the larva has eight rudiments of ampullae. In B. primigenus, the original ampullae degenerate soon after metamorphosis and new ampullae extend from the ventral epidermis of the oozooid. In the other species, the colonial vascular system is derived from the original ampullae. The whole colonial vascular system contracts and expands periodically, with regionally different phases. During each expansion cycle, the sphincter contracts once in B. primigenus and twice in S. reptans. The correlation may be due to blood pressure and the propagation of excitation through the colonial vascular system.     

Selective herbivory by an invasive cyprinid,the rudd Scardinius erythrophthalmus

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Grillotia australis n. sp. and G. pristiophori n. sp. (Cestoda: Trypanorhyncha) from Australian elasmobranch and teleost fishes

Two new species of Grillotia are described from elasmobranch and teleost fishes from south-eastern Australia. G. australis n. sp., from the Australian angel shark Squatina australis. Regan, most closely resembles G. smarisgora (Wagener, 1854) and G. angeli Dollfus, 1969, differing from both species in the presence of smaller bulbs, two or occasionally three hooks in each intercalary row in the basal region, reduced to one in the metabasal region compared with four or five hooks in the metabasal region of G. smarisgora and a single hook in G. angeli, and in the limited extent of the band of hooklets on the external surface in the basal region of the tentacle, a region which is covered with hooks in G. smarisgora. Plerocerci of this species were found in the mackerel Trachurus declivis (Jenys) (site not known) from Tasmania. G. pristiophori n. sp., from the saw sharks Pristiophorus cirratus (Latham) and P. nudipinnis Günther, most closely resembles G. spinosissima Dollfus, 1969 in possessing a scolex covered with spiniform microtriches, but differs in having six rather than five hooks in each principal row, no intercalary hooks and by possessing a band of hooklets on the external surface of the tentacle which diminishes distally into a single file, rather than persisting as a band eight to nine files wide. G. pristiophori is the first trypanorhynch to be recorded from saw-sharks.     

Ostracod fauna of Duvensee,an ancient lake in Northern Germany

About 4000 ostracod valves have been analysed from a sediment core of a cultivated bog NW of Hamburg. The deepest deposits are sands free of subfossils, the uppermost layers consist of peat. In the sediment in between, there are three layers containing undestroyed valves. The ostracod assemblies of the older claygyttja (Late-glacial) and the two younger, fine detritus deposits (Post-glacial) are strikingly different. Cytherissa lacustris, Candona neglecta, Ilyocypris bradyi, Herpetocyrpis reptans, and two Limnocythere species were found in the Late-glacial layers. Changes in abundance of these species indicate alterations in climate, lake ground, water inflow and waterlevels. Postglacial layers are rich in Metacypris cordata valves associated with numerous Candona species, L. inopinata, Darwinula stevensi, and Cyclocypris laevis. These species are characteristic of a small lake with a large littoral zone. A similar development in ostracod assemblages is found in middle Europe by Absolon (1973), although the main Candona species is not C. neglecta but C. candida. Ostracod analysis cannot demonstrate an influence of anthropogenic impacts on ostracod successions.     

The genus Gymnoascus Baranetzky

Summary The genusGymnoascus Baranetzky is monographed and found to include five valid species including one previously recognized species,G. reessii, one new combination,G. johnstoni, two new species,G. longitrichus andG. corniculatus, and a species originally described as aGymnoascus but included inMyxotrichum since 1893,G. uncinatus. Twenty species previously described as belonging toGymnoascus are excluded from the genus or are considered as doubtful representatives.Supported in part by Botany Department Research Grant # 1344, University of California, Los Angeles, California.     

Taxonomy and distribution of Sargassum (Phaeophyceae) in the Gulf of Thailand

Ten species of Sargassum (Sargassaceae, Phaeophyceae) were found along the Gulf of Thailand. Morphological characteristics of Sargassum baccularia (Mertens) C.A. Agardh, S. binderi Sonder, S. cinereum J.G. Agardh, S.crassifolium J.G. Agardh, S. longifructum Tseng et Lu, S. oligocystum Montagne, S. polycystum C.A. Agardh, S. siliquosum J.G. Agardh, S. swartzii (Turner) C.A. Agardh and one unidentified species were examined and are described in detail. The most common species were S. polycystum distributed widely in almost all the study sites, S. crassifolium restricted to Prachuap Khirikhan Province, S. longifructum restricted to Chumphon Province, S. siliquosum restricted to Surat Thani Province and one unidentified species restricted to Songkhla Province. Three species (S. cinereum, S. longifructum and S. swartzii) are new records for the algal flora of Thailand. Five species (S. baccularia, S. cinereum, S. longifructum, S. polycystum and the unidentified species) belong to the section Zygocarpicae (J.G. Agardh) Setchell.     

New records of three marine red algae from Japan

Three species in the red algal order Ceramiales, Dasya longifila Masuda et Uwai (Dasyaceae), Endosiphonia horrida (C. Agardh) P. Silva (Rhodomelaceae) and Laurencia flexilis Setchell (Rhodomelaceae), are reported from Japan for the first time, and their morphological features are described along with taxonomic comments. Our findings point to the northernmost limit of geographic distribution of these species in the north‐western Pacific. Dasya longifila is characterized by small, sparsely corticated axes, long pseudolaterals in which intercalary cell divisions take place, and a small number of tetrasporangial stichidia and spermatangial branches per fertile pseudolateral. Endosiphonia horrida is characterized by frequently anastomosing branches that form a bush‐like tuft without a percurrent axis, inner cortical cells becoming the same length as the axial and periaxial cells, and luxuriously developed, unbranched trichoblasts. Laurencia flexilis is characterized by numerous cartilaginous rigid axes developing from a basal disc without creeping branches, the production of 4 periaxial cells per vegetative segment and the absence of longitudinally oriented secondary pit‐connections between contiguous superficial cortical cells.     

Taxonomy,biogeography and DNA barcodes of Geodia species (Porifera,Demospongiae, Tetractinellida) in the Atlantic boreo‐arctic region

Geodia species north of 60°N in the Atlantic appeared in the literature for the first time when Bowerbank described Geodia barretti and G. macandrewii in 1858 from western Norway. Since then, a number of species have been based on material from various parts of the region: G. simplex, Isops phlegraei, I. pallida, I. sphaeroides, Synops pyriformis, G. parva, G. normani, G. atlantica, Sidonops mesotriaena (now called G. hentscheli), and G. simplicissima. In addition to these 12 nominal species, four species described from elsewhere are claimed to have been identified in material from the northeast Atlantic, namely G. nodastrella and G. cydonium (and its synonyms Cydonium muelleri and Geodia gigas). In this paper, we revise the boreo‐arctic Geodia species using morphological, molecular, and biogeographical data. We notably compare northwest and northeast Atlantic specimens. Biological data (reproduction, biochemistry, microbiology, epibionts) for each species are also reviewed. Our results show that there are six valid species of boreo‐arctic Atlantic Geodia while other names are synonyms or mis‐identifications. Geodia barretti, G. atlantica, G. macandrewii, and G. hentscheli are well established and widely distributed. The same goes for Geodia phlegraei, but this species shows a striking geographical and bathymetric variation, which led us to recognize two species, G. phlegraei and G. parva (here resurrected). Some Geodia are arctic species (G. hentscheli, G. parva), while others are typically boreal (G. atlantica, G. barretti, G. phlegraei, G. macandrewii). No morphological differences were found between specimens from the northeast and northwest Atlantic, except for G. parva. The Folmer cytochrome oxidase subunit I (COI) fragment is unique for every species and invariable over their whole distribution range, except for G. barretti which had two haplotypes. 18S is unique for four species but cannot discriminate G. phlegraei and G. parva. Two keys to the boreo‐arctic Geodia are included, one based on external morphology, the other based on spicule morphology. © 2013 The Linnean Society of London     

MORPHOLOGY AND MOLECULAR ANALYSES OF THREE TOXIC SPECIES OF GAMBIERDISCUS (DINOPHYCEAE): G. PACIFICUS, SP. NOV., G. AUSTRALES, SP. NOV., AND G. POLYNESIENSIS, SP. NOV.

Three new dinoflagellate species, Gambierdiscus polynesiensis, sp. nov., Gambierdiscus australes, sp. nov., and Gambierdiscus pacificus, sp. nov., are described from scanning electron micrographs. The morphology of the three new Gambierdiscus species is compared with the type species Gambierdiscus toxicus Adachi et Fukuyo 1979, and two other species: Gambierdiscus belizeanus Faust 1995 and Gambierdiscus yasumotoi Holmes 1998. The plate formula is: Po, 3′, 7", 6C, 8S, 5‴, 1p, 2". Culture extracts of these three new species displayed both ciguatoxin- and maitotoxin-like toxicities. The following morphological characteristics differentiated each species. 1) Cells of G. polynesiensis are 68–85 μm long and 64–75 μm wide, and the cell’s surface is smooth. They are identified by a large triangular apical pore plate (Po), a narrow fish-hook opening surrounded by 38 round pores, and a large, broad posterior intercalary plate (1p) wedged between narrow postcingular plates 2‴ and 4‴. Plate 1p occupies 60% of the width of the hypotheca. 2) Cells of G. australes also have a smooth surface and are 76–93 μm long and 65–85 μm wide in dorsoventral depth. They are identified by the broad ellipsoid apical pore plate (Po) surrounded by 31 round pores and a long and narrow 1p plate wedged between postcingular plates 2‴ and 4‴. Plate 1p occupies 30% of the width of the hypotheca. 3) Cells of G. pacificus are 67–77 μm long and 60–76 μm wide in dorsoventral depth, and its surface is smooth. They are identified by the four-sided apical pore plate (Po) surrounded by 30 round pores. A short narrow 1p plate is wedged between the wide postcingular plates 2‴ and 4‴. Plate 1p occupies 20% of the width of the hypotheca. These three newly described species were also characterized by isozyme electrophoresis and DNA sequencing of the D8–D10 region of their large subunit (LSU) rRNA genes. The consistency between species designations based on SEM microscopy and classification inferred from biochemical and genetic heterogeneities was examined among seven isolates of Gambierdiscus. Their classification into four morphospecies was not consistent with groupings inferred from isozyme patterns. Three molecular types could be distinguished based on the comparison of their LSU rDNA sequences. Although G. toxicus TUR was found to be more closely related to G. pacificus, sp. nov. than to other G. toxicus strains, the molecular classification was able to discriminate G. polynesiensis, sp. nov. and G. australes, sp. nov. from G. toxicus. These results suggest the usefulness of the D8–D10 portion of the Gambierdiscus LSU rDNA as a valuable taxonomic marker.