Secondary structure analysis of 34 internal transcribed spacer 2 (ITS-2) sequences showed that the current model for the green algae Scenedesmus and Desmodesmus is not accurate. In particular, helix I of the currently used model showed considerable deviations from our new model. The newly proposed model is supported by many two-sided compensated base pair changes and fully compensated insertions in all four helices. Phylogenetic analysis by maximum parsimony based on the new alignment confirmed the recent division of the old genus Scenedesmus into the new genera Scenedesmus and Desmodesmus. However, the analysis was not able to show phylogenetic relationships within these two genera. Hence, the ITS-2 region alone is not suitable for clarifying the phylogeny of Scenedesmus and Desmodesmus and new regions have to be found for future sequence analyses.

New holococcolith–heterococcolith life-cycle associations are documented based on observations of combination coccospheres. Daktylethra pirus is shown to be a life-cycle phase of Syracosphaera pulchra and Syracolithus quadriperforatus a life-cycle phase of Calcidiscus leptoporus. In addition, new observations from cultures confirm the life-cycle associations of Crystallolithus braarudii with Coccolithus pelagicus and of Zygosphaera hellenica with Coronosphaera mediterranea. In all four cases previous work has shown that the heterococcolithophorid species is associated with another holococcolithophorid. Two other examples of a heterococcolithophorid being associated with two holococcolithophorids have previously been identified, so this seems to be a common phenomenon. The six examples are reviewed to determine whether a single underlying mechanism is likely to be responsible for all cases. It is concluded that there is no single mechanism but rather that the six examples fall into three categories: (a) in two cases the holococcolith types are probably simply ecophenotypic morphotypes; (b) in two other cases the holococcolith types are discrete and are paralleled by morphometric differences in the heterococcolith types; (c) in the final two cases the holococcolith types are discrete but are not paralleled by any obvious morphological variation in the heterococcolith morphology. We infer that cryptic speciation may be widespread in heterococcolithophorid phases and that study of holococcolithophorid phases can provide key data to elucidate this phenomenon.

Grateloupia doryphora (Montagne) Howe, originally described from Peru, has repeatedly been reported as an invasive species in Atlantic and Mediterranean waters. Various attempts to explain this species’ route of introduction have been unsatisfactory. New evidence from comparative rbcL sequence analysis and morphology suggests that this adventive species in the NE and NW Atlantic corresponds with G. turuturu Yamada, originally described from Japan. This provenance follows a well-recognized trend of invasive marine organisms that have colonized the Atlantic Ocean and Mediterranean Sea from Pacific NE Asia.

The temperature dependence of UV effects was studied for Arctic and temperate isolates of the red macrophytes Palmaria palmata, Coccotylus truncatus and Phycodrys rubens. The effects of daily repeated artificial ultraviolet B and A radiation (UVBR: 280–320 nm, UVAR: 320–400 nm) treatments were examined for all isolates at 6, 12 and 18 °C by measuring growth, optimal quantum yield of PSII (Fv/Fm) and cyclobutane–pyrimidine dimer (CPD) accumulation. Furthermore, possible ecotypic differences in UV sensitivity between Arctic and temperate isolates were evaluated. Large species-specific differences in UV sensitivity were observed for all parameters: the lower subtidal species C. truncatus and P. rubens were highly sensitive to the UV treatments, whereas P. palmata, which predominantly occurs in the upper subtidal zone, was not affected by these treatments. Only minor differences were found between Arctic and temperate isolates, suggesting that no differences in UV sensitivity have evolved in these species. Relative growth rates were temperature-dependent, whereas species-specific UV effects on growth rates were relatively independent of temperature. In contrast, the species-specific decrease in Fv/Fm and its subsequent recovery were temperature-dependent in all species. UV effects on Fv/Fm were lower at 12 and 18 °C compared with 6 °C. In addition, UV effects on Fv/Fm decreased in the course of the experiment at all temperatures, indicating acclimation to the UV treatments. CPDs accumulated during the experiment in both isolates of P. rubens, whereas CPD concentrations remained low for the other two species. CPD accumulation appeared to be independent of temperature. The results suggest that summer temperatures occurring in temperate regions facilitate repair of UV-induced damage and acclimation to UV radiation in these algae compared with Arctic temperatures. Because the differences in UV effects on Fv/Fm, growth and CPD accumulation were relatively small over a broad range of temperatures, it was concluded that the influence of temperature on UV effects is small in these species.

Uncertainties about the identity of type specimens of red algae have frequently led to taxonomic and nomenclatural confusion. A procedure for extracting PCR-amplifiable DNA from formalin-fixed material and herbarium specimens was used to investigate the taxonomic status of several South African Gigartinaceae. We compared nucleotide sequences in the internal transcribed spacer 1 (ITS1) region in type specimens and other historically important collections presently referred to Gigartina bracteata, G. radula and G. stiriata. The following opinions are supported: (1) Fucus bracteatus S.G. Gmelin, Chondrodictyon capense Kützing and Iridaea clathrata Decaisne represent a single species of South African Gigartina (G. bracteata (S.G. Gmelin) Setchell & Gardner) in which the disintegration of tetrasporangial sori results in a reticulate thallus. (2) Mastocarpus polycarpus Kützing, M. incrassatus Kützing and Iridaea lapathifolia Kützing represent a single species of South African Gigartina (G. polycarpa (Kützing) Setchell & Gardner) that has often, but erroneously, been called G. radula (Esper) J. Agardh. (3) Mastocarpus verrucosus Kützing is a later heterotypic synonym of Iridaea papillosa Bory (Sarcothalia papillosa (Bory) Leister) and was based on material that probably came from southern South America rather than from South Africa, the provenance given by Kützing. (4) Fucus stiriatus Turner and Sphaerococcus burmannii C. Agardh represent a single species of South African Sarcothalia (S. stiriata (Turner) Leister).

The aim of the current investigation was to test the general convention that the Dasyaceae, Delesseriaceae and Rhodomelaceae are all monophyletic families of the red algal order Ceramiales. Phylogenetic relationships among 45 ceramialean taxa were determined, including eight ceramiacean, 18 dasyacean, nine delesseriacean and eight rhodomelacean species, plus two of uncertain ceramialean affinities, based on 34 anatomical characters and nuclear small subunit (SSU) rDNA sequences. Results from our ‘total-evidence’ approach were consistent with the notion that the Dasyaceae, Delesseriaceae and Rhodomelaceae have evolved from a common ancestor within a paraphyletic Ceramiaceae. Our data indicate, however, that the Rhodomelaceae alone is monophyletic at the family level, with both the Dasyaceae and Delesseriaceae polyphyletic. In particular we resolved two independent and well-supported lineages for the included Dasyaceae, viz., a Dasya group and a Heterosiphonia group, which were as distinct from one another as they were from the Rhodomelaceae and the various lineages of Delesseriaceae. The molecular data alone were equivocal on the issue of monophyly of the Dasyaceae. We therefore advocate conservative taxonomic revisions as an interim step towards eventual resolution of familial-level taxonomy in the Ceramiales. Based on our results, the Heterosiphonioideae H.-G. Choi, Kraft, I.K. Lee et G.W. Saunders subfam. nov. is proposed for Heterosiphonia and five closely related genera, and the Dasyoideae Schmitz et Falkenberg is emended for the remaining taxa. Although the Dasyoideae is a natural group, it is in need of a thorough systematic reinvestigation at the generic level. Our analyses indicate that the genus Dasya is polyphyletic or paraphyletic in excluding Dasysiphonia, Eupogodon and Rhodoptilum and that Heterosiphonia japonica also has affinities to this group, taxonomic issues that will be addressed in detail elsewhere.

Macroalgal assemblages of Posidonia oceanica rhizomes of the western Mediterranean have been studied, in order to describe the structure of this community and to evaluate changes due to recent macroalgal invasions. To achieve these objectives, P. oceanica rhizomes were sampled from 21 beds distributed throughout the western Mediterranean Sea and macroalgal assemblages were studied using both species and functional groups as descriptors. Moreover, modifications linked to temporal fluctuations and depth were evaluated in an insular and in a continental bed. A total of 74 macroalgal species were found, among them 7 Chlorophyta, 8 Fucophyceae and 59 Rhodophyta. The species number varied from 5±0 at Vada to 18±0·58 at Panarea Island. The total percentage cover ranged from 19·5±1·8 at Cabo de Creus to 77·2±13·3 at La Maddalena Island. Filamentous species were dominant in the localities colonized by Acrothamnion preissii or Womersleyella setacea, while a high functional diversity occurred where the two introduced species were not present or they did not constitute dense turfs; the importance of crustose and foliose species was remarkable, while low cover of articulate and corticated-terete species was found at all localities. Localities where turfs were not developed were grouped together, even if they appeared more dispersed than those dominated by filamentous algae. The two beds studied over a 1 year period showed higher values of macroalgal cover in March and December, although significant differences were not detected with ANOVA in relation to either depth or season. The development of turfs, which occurs widely in western Mediterranean seagrass beds, seems to modify strongly the structure of macroalgal assemblages of rhizomes, mostly via the decrease in species and functional diversity.

In the course of the cell cycles of synchronous cultures of the chlorococcal alga Scenedesmus quadricauda, the following were monitored: total protein and RNA accumulation as a measure of growth processes, the timing of the commitment points at which the cells trigger the sequence of reproductive processes (DNA replication, nuclear and cellular division) and the course of the reproductive processes. The synchronous cultures were grown either under various lighting regimes, or in the temporary presence of specific inhibitors of either proteosynthesis (cycloheximide) or DNA replication (5-fluorodeoxyuridine). By adjusting the length of the light period, the cell cycle could be manipulated. Cell cycle patterns could be altered to give different numbers of sequences of reproductive processes. The extent of their mutual overlap could be influenced and the number of daughter cells produced could be altered. Schematic illustrations of various cell cycle patterns and comparisons with those of higher plants and other algal species are presented.

Phylogenetic analyses, based upon nuclear small-subunit ribosomal RNA gene sequences, of four ‘chlorosarcinoid’ species referred to Planophila Gerneck show that the genus is polyphyletic. The type species, P. laetevirens Gerneck, is closely related to species in the Ulotrichales, Ulvophyceae. The monotypic sarcinoid genus Pseudendocloniopsis is the closest relative of Planophila; the two genera represent the addition of a new morphological type to the Ulotrichales. Planophila microcystis (Dangeard) Kornmann & Sahling forms a clade at the base of the Ulvophyceae with Oltmannsiellopsis, and thus belongs to the Oltmannsiellopsidales. This result is also supported by the Oltmannsiellopsis-like ultrastructure of P. microcystis zoospores. Planophila sp. B from Antarctica, which has Trebouxia-like pyrenoid structure, is a trebouxiophyte closely related to Chlorella-like unicellular coccoids, Stichococcus bacillaris and Prasiola species. This is the first robustly supported molecular phylogenetic analysis that places Prasiola in the Trebouxiophyceae. As shown previously, P. terrestris Groover & Hofstetter belongs to the Chaetopeltidales, Chlorophyceae. Dangemannia gen. nov. (type species: D. microcystis (Dangeard) comb. nov.), Floydiella gen. nov. (type species: F. terrestris (Groover & Hofstetter) comb. nov.) and Pabia gen. nov. (type: P. signiensis sp. nov.) are proposed.

Changes in the ultrastructure and chemistry of the cell wall of the unicellular volvocalean green alga Haematococcus pluvialis were investigated during the transformation of flagellates into aplanospores. The motile biflagellated state exhibited a distinct gelatinous extracellular matrix. Its ultrastructure resembled the typical volvocalean multilayered architecture with a median tripartite crystalline layer. The transformation into the non-motile cell state was characterized by formation of a new layer, a primary wall, within the extracellular matrix. During this process, the initial extracellular matrix remained intact except for the outer layers of the tripartite crystalline layer, which decomposed. Further morphogenesis of the aplanospore resulted in the formation of a voluminous multilayered cell wall. A trilaminar sheath was formed inside the primary wall and the innermost and thickest part was an amorphous secondary wall, consisting mostly of a mannan. Results obtained by staining with the fluorescent dye primuline as well as by acetolysis suggest the occurrence of sporopollenin-like material (algaenan) within the trilaminar sheath of the aplanospore cell wall. The primary wall and the outer remnants of the extracellular matrix disintegrated as the aplanospores aged, and were completely absent in the resting cell state.

Sequence variation among 22 isolates representing a global distribution of the prymnesiophyte genus Phaeocystis has been compared using nuclear-encoded 18S rRNA genes and two non-coding regions: the ribosomal DNA internal transcribed spacer 1 (ITS1) separating the 18S rRNA and 5·8S rRNA genes and the plastid ribulose-1,5-bisphosphate carboxylase/oxygenase (RUBISCO) spacer flanked by short stretches of the adjacent large and small subunits (rbcL and rbcS). 18S rRNA can only resolve major species complexes. The analysis suggests that an undescribed unicellular Phaeocystis sp. (isolate PLY 559) is a sister taxon to the Mediterranean unicellular Phaeocystis jahnii; this clade branched prior to the divergence of all other Phaeocystis species, including the colonial ones. Little divergence was seen among the multiple isolates sequenced from each colonial species complex. RUBISCO spacer regions are even more highly conserved among closely related colonial Phaeocystis species and are identical in Phaeocystis antarctica, Phaeocystis pouchetii and two warm-temperate strains of Phaeocystis globosa, with a single base substitution in two cold-temperate strains of P. globosa. The RUBISCO spacer sequences from two predominantly unicellular Phaeocystis isolates from the Mediterranean Sea and PLY 559 were clearly different from other Phaeocystis strains. In contrast, ITS1 exhibited substantial inter- and intraspecific sequence divergence and showed more resolution among the taxa. Distinctly different copies of the ITS1 region were found in P. globosa, even among cloned DNA from a single strain, suggesting that it is a species complex and making this region unsuitable for phylogenetic analysis in this species. However, among nine P. antarctica strains, four ITS1 haplotypes could be separated. Using the branching order in the ITS1 tree we have attempted to trace the biogeographic history of the dispersal of strains in Antarctic coastal waters.

Examination of material obtained from the Urdaibai estuary (North Spain) during a 2 year phytoplankton sampling programme revealed a rare species of the genus Chaetoceros, C. salsugineus Takano. The morphology of this small (4.0×5.5 μm) member of the subgenus Hyalochaetae has been described using both light and electron microscopy. Regarding the original material, two new features have been observed: longer chains up to 24 cells and a new type of aperture, bisected by the fusion of central protuberances of adjacent valves. This bloom-forming diatom reached high densities in the poly-euhaline zone of the estuary closely related with high temperatures, denoting its neritic and summer preferences. Maximal cell densities (106–107 cells l−1) were reached in August and September when the water column was completely mixed and the salinity and temperature were about 31 psu and 20 °C, respectively. This is the first report of C. salsugineus in Europe and contributes to the knowledge of the morphological and ecological features of this species. Relationships with other small species of the subgenus Hyalochaetae are also discussed.

A new member of Delesseriaceae (Ceramiales, Rhodophyta) is described from Southern Taiwan and the Philippines. On the basis of comparative vegetative and reproductive morphology, and phylogenetic analysis inferred from nuclear-encoded large-subunit ribosomal DNA sequences (LSU rDNA), we conclude that it belongs in the genus Drachiella, tribe Schizoserideae, subfamily Phycodryoideae. The new taxon shares with other Drachiella species the absence of macro- and microscopic veins; diffuse growth by marginal and intercalary meristematic cells; a polystromatic, lobed thallus; abundance of rhizoidal marginal proliferations used for attachment; convoluted plastids in surface cells; abundant secondary pit connections among adjacent vegetative cells; large intercellular spaces between surface cells; procarps confined to the upper side of the thallus, circular in outline, consisting of a supporting cell bearing a strongly curved carpogonial branch and two sterile groups that remain undivided; vertical division of gonimoblast initial from auxiliary cell, and unilateral, monopodial branching of gonimoblasts; and mature cystocarps with a massive candelabrum-like fusion cell of fused gonimoblasts bearing carposporangia in branched chains. It is distinguished from the other members of the genus by thalli that consist of extensive tangled mats of prostrate and overlapping decumbent blades, procarps confined to the upper side of the thallus, and the lack of basal stalks or stipes. Whereas the Schizoserideae is predominantly a Southern Ocean tribe, one of the tribe's four genera, Drachiella, was known only from the eastern Atlantic and Mediterranean. We herein report the first record of the genus for the Indo-Pacific Ocean, and describe Drachiella liaoii, sp. nov., as a fourth species in the genus.

We examined three species of diminutive Porphyra,Porphyra suborbiculata Kjellman from the North Pacific, Porphyra lilliputiana W. A. Nelson, G. A. Knight et M. W. Hawkes from the South Pacific, and Porphyra carolinensis Coll et J. Cox from the western North Atlantic. These taxa were compared in terms of morphology, habitat data and sequence haplotypes of nuclear small subunit rDNA (SSU) and internal transcribed spacers of the nuclear rDNA cistron (ITS). These three species have similar morphologies and growth habits, and share very similar type descriptions and habitat records. Haplotype variation was found within the 11 samples of P. lilliputiana we examined and within P. suborbiculata samples from two locations, but the single P. carolinensis haplotype (from collections from two separate locations) was identical to one found in several widespread P. lilliputiana samples. Unrooted phylogenetic trees based on sequence data do not support any of the three species as being a monophyletic group. We conclude that these three taxa represent a single species with the oldest name P. suborbiculata having nomenclatural priority. It is likely that P. suborbiculata has recently been introduced to the western Atlantic from the Pacific region.

Quantification of the iron nutritional status of phytoplankton is of great interest not only for the study of the oceans but also for fresh waters. Flavodoxin is a small flavoprotein proposed as a molecular marker for iron stress, since it is induced as a consequence of iron deprivation, replacing the iron-sulphur protein ferredoxin. Flavodoxin and ferredoxin from Scenedesmus vacuolatus have been immunoquantified in cells grown under different iron nutritional conditions. Flavodoxin and ferredoxin levels correlate with the iron availability, and the calculated flavodoxin index can be used as an iron-stress marker. Other physiological parameters such as copper deficiency, heterotrophic or mixotrophic growth, nitrogen source and salt stress were also tested as potential factors influencing flavodoxin expression. Salt stress and heterotrophic growth conditions alter flavodoxin and ferredoxin expression. Once flavodoxin expression is repressed by iron (and severe deficiency alleviated), S.vacuolatus still increases its ferredoxin from 0.5 to 1.6 mol of ferredoxin per mole of ferredoxin-NADP+ reductase, and this ratio can be used for the evaluation of mild deficiency.

Phylogenetic relationships within the Sphacelariales sensu stricto were investigated using both a molecular and a morphological approach. Twenty species were included, representing all three families (i.e. Cladostephaceae, Sphacelariaceae and Stypocaulaceae) and six of the eight genera. The outgroup consisted of six species representing the Syringodermatales, Dictyotales, Choristocarpaceae and Onslowiaceae. DNA sequences of partial rbcL [1255–1375 nucleotides (nt)] and the adjacent RUBISCO spacer (40–842 nt) were determined in order to assess the molecular phylogeny. Only the 3′-end of the spacer (112 nt positions) was alignable for the ingroup taxa. Partition homogeneity testing showed that rbcL and RUBISCO spacer sequences could be combined. However, analysis of rbcL alone or in combination with the RUBISCO spacer gave the same results with only slight differences in support (bootstrap, jackknife, decay). Support was low at the base of the ingroup. Four basal clades could be discerned: (1) Stypocaulaceae, (2) Sphacelaria radicans, (3) Sphacelaria caespitula and (4) all other included taxa: (Sphacella subtilissima+the Sphacelaria subgenus Propagulifera) and (Cladostephus spongiosus+Sphacelaria nana+the Sphacelaria subgenera Battersia+Pseudochaetopteris). The independent morphological analysis (using 23 unordered morphological characters) revealed high homoplasy and an almost completely unresolved tree in which only the subgenus Propagulifera was supported. The morphological characters were subsequently mapped onto the rbcL tree in order to identify diagnostic or phylogenetically informative characters. Blackening in response to bleach and the presence of secondary segments were found to be basal synapomorphies for the Sphacelariales; presence of propagules with a lenticular central apical cell defines the Propagulifera; and strict acroblastic branching mode and axillary zoidangia define the Stypocaulaceae. The remaining characters have been gained or lost multiple times. This study highlights the problem of extreme morphological convergence and/or plasticity. Four options for a new circumscription including nomenclatural changes are discussed. It is concluded that none of the options will lead to greater clarity with respect to either identification or classification.

An undescribed marine Pyramimonas, P. australis Andreoli et Moro, sp. nov., forming a bloom in a hole of Terra Nova Bay (Ross Sea, Antarctica) sea ice, was collected, but could not be cultured. Consequently, the description of this new species is based on light and electron microscope observations on samples that were fixed or stored at −80 °C, and its phylogenetic position inferred from nuclear-encoded small-subunit ribosomal DNA (SSU rDNA) and chloroplast-encoded rbcL gene sequences. This is the third Antarctic species described for this genus. The ultrastructure of the cell is consistent with species of the subgenus Trichocystis McFadden, but differs in that it has unique body and cyst scales, and a different encystment procedure. The outermost layer of body scales is formed by flat box scales with peripheral perforations oriented parallel to the four edges and with a further eight central perforations oriented perpendicular to the peripheral ones. Crown scales, which in many other species of the genus form the outermost layer over the entire cell body, were observed in this species in the flagellar pit over the box scales. The flagella are covered by a pentagonal underlayer of scales and by limuloid scales with two subsidiary spines, in addition to the central one. Encystment begins in the flagellate form resulting in a cyst with an irregular wall bearing spine scales. Ultrastructural and molecular data confirm that P. australis belongs to the subgenus Trichocystis.

The effects on photosynthesis of CO2 and desiccation in Porphyra haitanensis were investigated to establish the effects of increased atmospheric CO2 on this alga during emersion at low tides. With enhanced desiccation, net photosynthesis, dark respiration, photosynthetic efficiency, apparent carboxylating efficiency and light saturation point decreased, while the light compensation point and CO2 compensation point increased. Emersed net photosynthesis was not saturated by the present atmospheric CO2 level (about 350 ml m−3), and doubling the CO2 concentration (700 ml m−3) increased photosynthesis by between 31% and 89% at moderate levels of desiccation. The relative enhancement of emersed net photosynthesis at 700 ml m−3 CO2 was greater at higher temperatures and higher levels of desiccation. The photosynthetic production of Porphyra haitanensis may benefit from increasing atmospheric CO2 concentration during emersion.

Molecular and breeding studies on two pan-tropical marine red algae reveal vastly different levels of genetic variation and reproductive isolation. Sequenced DNA regions from the nuclear, mitochondrial and plastid genomes (partial LSU rRNA, cox2-3 spacer, RuBisCo spacer, respectively) revealed high genetic variation among individuals of Spyridia filamentosa. An rbcL analysis shows that Spyridia is a monophyletic genus distinct from other Ceramiaceae sampled, suggesting that the Ceramiaceae could be paraphyletic. There is complete congruence between all data sets of S. filamentosa, showing a well-supported phylogeographic pattern with samples from the Pacific distinct from Atlantic and Indian Ocean samples. One western Mediterranean sample is associated with Atlantic specimens, while an eastern Mediterranean sample has closer affinities to Pacific samples, possibly indicating a recent cryptic introduction into the eastern Mediterranean. Limited breeding studies imply that these samples are mostly reproductively isolated, whereas a successful cross demonstrated maternal inheritance of organellar DNA. These data indicate that S. filamentosa exists as several cryptic species. Murrayella periclados exhibits low levels of genetic variation and no phylogeographic structure, and almost complete reproductive compatibility between isolates. This suggests that all M. periclados samples share a recent common ancestor that may have dispersed relatively rapidly, or that rates of base pair substitution between these two species vary greatly. Rapid long-distance dispersal of M. periclados is not indicated by what is known of the biology of M. periclados, especially in comparison with S. filamentosa, which appears to be a much better candidate for long-distance dispersal. These data demonstrate that red algal morphospecies are not equivalent units of diversity, with implications for our view of red algal biodiversity and evolution.

An elachistacean epiphyte, Neoleptonema yongpilii E.-Y. Lee & I.K. Lee, gen. et sp. nov., is reported from Korean coasts. The plants are distinguished by having unbranched assimilatory filaments with intercalary plurilocular sporangia as well as lateral plurilocular sporangia from the cortex. The new genus differs from the genera Leptonematella P. Silva and Halothrix Reinke by having pod-shaped plurilocular sporangia on the medulla, and from Elachista Duby and Proselachista Y.P. Lee & Garbary by having intercalary plurilocular sporangia and a poorly developed medulla. The phylogenetic relationships of Neoleptonema yongpilii were inferred from the spacer sequences between the genes coding for the large and small subunits of the RuBisCO gene. The new genus is a member of a poorly resolved clade consisting of several genera within the Elachistaceae, within which it is more closely related to Halothrix and Elachista nipponica than to Leptonematella.