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1.
Although the combination of different genes in phylogenetic analyses is a promising approach, the methodology is not well established and analyses often suffer from inadequate, noncongruent taxon sampling, long-branch attraction, or conflicting evolutionary models of the genes analyzed. Conflicts or congruence between multigene and single-gene phylogenies, as well as the assumed superiority of the multigene approach, are often difficult to assess solely because of incongruent taxon sampling. In the present study, a data set of 43 nuclear-encoded SSU rDNA and plastid-encoded rbcL gene sequences was generated from the same strains of conjugating green algae (Zygnematophyceae, Streptophyta). Phylogenetic analyses used the genes individually and in combination, either as concatenated sequences or with the log-likelihood summation method. Single-gene analyses, although mostly congruent, revealed some conflicting nodes and showed different patterns of statistical support. Combined analyses confidently resolved the conflicts between the single-gene analyses, enhanced phylogenetic resolution, and were better supported by morphological information. Long-branch taxa were not the same for the two genes analyzed, and, thus, their effect on phylogenetic resolution was minimized in the combined analyses. 相似文献
2.
Concatenated sequences of small- and large-subunit rRNA genes were used to infer the phylogeny of 29 species in six genera of Tintinnida. We confirmed previous results on the positions of major clusters and the grouping of various genera, including Stenosemella, the paraphyletic Tintinnopsis, the newly investigated Helicostomella, and some species of the polyphyletic Favella. Tintinnidium and Eutintinnus were found to be monophyletic. This study contributes to tintinnid phylogenetic reconstruction by increasing both the number of species and the range of genetic markers analyzed. 相似文献
3.
Christopher S. Drummond John Hall Kenneth G. Karol Charles F. Delwiche Richard M. McCourt 《Journal of phycology》2005,41(5):1055-1064
DNA sequence data were obtained for the gene encoding the large subunit of RUBISCO (rbcL) from 26 strains of Spirogyra and seven of Sirogonium, using as outgroups 10 genera in the Zygnematales and Desmidiales (Closterium, Cosmarium, Cylindrocystis, Gonatozygon, Mesotaenium, Netrium, Penium, Zygnema, Zygnemopsis, Zygogonium). Sequence data were analyzed using maximum parsimony (MP), maximum likelihood (ML), and Bayesian inference (BI), with bootstrap replication (MP, ML) and posterior probabilities (BI) as measures of support. MP, ML, and BI analyses of the rbcL data strongly support a single clade containing Spirogyra and Sirogonium. The Spirogyra taxa are monophyletic, with the exception of Spirogyra maxima (Hassall) Wittrock, which is nested within a clade with Sirogonium and shares with them the characters of loosely spiraled chloroplasts (<1 complete turn per cell) and anisogamy of gametangial cells; S. maxima differs from Sirogonium in displaying well‐defined conjugation tubes rather than a tubeless connection involving bending (genuflection) of filaments. The ML and BI analyses place this Sirogonium/Spirogyra maxima clade sister to the remaining Spirogyra. Morphological differences among strains of Spirogyra grouped together on the basis of rbcL data, including laboratory strains derived from clonal cultures (Spirogyra communis, S. pratensis), indicate that some characters (filament width, chloroplast number) used in the traditional taxonomy of this group are poor measures of species identity. However, some characters such as replicate end walls and loose spiraling of chloroplasts may be synapomorphies for Spirogyra clades. 相似文献
4.
We newly sequenced the nuclear-encoded small subunit (SSU) rDNA coding region for 21 taxa of the genus Closterium. The new sequences were integrated into an alignment with 13 known sequences of conjugating green algae representing six traditional families (i.e. Zygnemataceae, Mesotaeniaceae, Gonatozygaceae, Peniaceae, Closteriaceae, and Desmidiaceae) and five known charophycean sequences as outgroups. Both maximum likelihood and maximum parsimony analyses supported with high bootstrap values one large clade containing all placoderm desmids (Desmidiales). All the Closterium taxa formed one clade with 100% bootstrap support, indicating their monophyly, but not paraphyly, as suggested earlier. As to the taxa within the genus Closterium , we found two clades of morphologically closely related taxa in both maximum likelihood and maximum parsimony trees. They corresponded to the C. calosporum species complex and the C. moniliferum-ehrenbergii species complex. It is of particular interest that the homothallic entity of C. moniliferum v. moniliferum was distinguished from and ancestral to all other entities of the C. moniliferum-ehrenbergii species complex. Superimposing all 50 charophycean sequences on the higher order SSU rRNA structure model of Closterium , we investigated degrees of nucleotide conservation at a given position in the nucleotide sequence. A characteristic \"signature\" structure to the genus Closterium was found as an additional helix at the tip of V1 region. In addition, eight base deletions at the tip of helix 10 were found to be characteristic of the C. calosporum species complex, C. gracile , C. incurvum , C. pleurodermatum , and C. pusillum v. maius. These taxa formed one clade with an 82% bootstrap value in maximum parsimony analysis. 相似文献
5.
Alexis M. Bellorin Amelia Buriyo Jelveh Sohrabipour Mariana C. Oliveira Eurico C. Oliveira 《Journal of phycology》2008,44(4):1022-1032
Two new species of Gracilariopsis from the Indian Ocean are proposed—Gracilariopsis (Gp.) mclachlanii Buriyo, Bellorin et M. C. Oliveira sp. nov. from Tanzania and Gracilariopsis persica Bellorin, Sohrabipour et E. C. Oliveira sp. nov. from Iran—based on morphology and DNA sequence data (rbcL gene and SSU rDNA). Both species fit the typical features of Gracilariopsis: axes cylindrical throughout, freely and loosely ramified up to four orders, with an abrupt transition in cell size from medulla to cortex, cystocarps lacking tubular nutritive cells and superficial spermatangia. Nucleotide sequence comparisons of rbcL and SSU rDNA placed both species into the Gracilariopsis clade as distinct species from all the accepted species for this genus, forming a deeply divergent lineage together with some species from the Pacific. The new species are very difficult to distinguish on morphological grounds from other species of Gracilariopsis, stressing the importance of homologous molecular marker comparisons for the species recognition in this character‐poor genus. 相似文献
6.
Ionel Ciugulea María A. Nudelman Stacy Brosnan Richard E. Triemer 《Journal of phycology》2008,44(2):406-418
Previous studies using the nuclear SSU rDNA and partial LSU rDNA have demonstrated that the euglenoid loricate taxa form a monophyletic clade within the photosynthetic euglenoid lineage. It was unclear, however, whether the loricate genera Trachelomonas and Strombomonas were monophyletic. In order to determine the relationships among the loricate taxa, SSU and LSU nuclear rDNA sequences were obtained for eight Strombomonas and 25 Trachelomonas strains and combined in a multigene phylogenetic analysis. Conserved regions of the aligned data set were used to generate maximum‐likelihood (ML) and Bayesian phylogenies. Both methods recovered a strongly supported monophyletic loricate clade with Strombomonas and Trachelomonas species separated into two sister clades. Taxa in the genus Strombomonas sorted into three subclades. Within the genus Trachelomonas, five strongly supported subclades were recovered in all analyses. Key morphological features could be attributed to each of the subclades, with the major separation being that all of the spine‐bearing taxa were located in two sister subclades, while the more rounded, spineless taxa formed the remaining three subclades. The separation of genera and subclades was supported by 42 distinct molecular signatures (33 in Trachelomonas and nine in Strombomonas). The morphological and molecular data supported the retention of Trachelomonas and Strombomonas as separate loricate genera. 相似文献
7.
Nicolas Chomérat Daniel Y. Sellos Frédéric Zentz Elisabeth Nézan 《Journal of phycology》2010,46(1):183-194
A new marine benthic Prorocentrum species from sandy habitats of South Brittany (northwestern France), P. consutum sp. nov., is described using LM and SEM and molecular phylogenetic analyses. Cells have a subcircular to broadly ovoid shape and are plainly flattened. They are 57–61 μm long and 52–55 μm wide. A central pyrenoid is present, and the kidney‐shaped nucleus is positioned in the posterior region. In right valve view, the periflagellar area is deeply excavated, and the left valve forms a prominent apical ridge. The periflagellar area consists of nine platelets, and a small narrow collar is present around the flagellar pore. The ornamentation of this new species is very peculiar and is characterized by a ring of round areolae located at the periphery of the valves, each areola containing three or four pores. Apart from this ring of areolae, the cell surface is smooth and with scattered pores. Pores are not present in the center of the right or left valve. The intercalary band is generally narrow and faintly striated horizontally. The molecular phylogenetic position of P. consutum sp. nov. was inferred using SSU and LSU rDNA. In both analyses, this species branched with high support in the clade comprising species with a symmetric shape and appeared to be a sister group to that formed by P. lima and other tropical benthic species, such as P. arenarium, P. belizeanum, P. hoffmannianum, and P. maculosum. 相似文献
8.
Young plants of E. intestinalis have been grown in culture. Cutting off the thalli of attached plants just above the basal rhizoids leads to the regeneration of new branches. After a short time the point of wounding is indistinguishable. The process can be repeated many times; each regeneration resulting in a more branched thallus. Segments of unattached plants kept in culture may give rise to rhizoids from the basal cut ends and to papillae from the upper ends. They may also produce “bottle brush” forms similar to those found amongst ship-fouling algae. Such forms seem to arise when “swarmers” are retained in the parent cell and germinate in situ to give aggregations of juvenile plants. Their formation can be stimulated by a temperature shock. 相似文献
9.
Francine M.G. McCarthy Haifeng Gu Kenneth N. Mertens Consuelo Carbonell‐Moore Andrea M. Krueger Yoshihito Takano Kazumi Matsuoka 《Phycological Research》2018,66(2):137-148
Distinctive spindle‐shaped thecae first described by Samuel Eddy in 1930 and assigned to the genus Peridinium Ehrenberg are commonly reported from freshwater environments in eastern North America. We demonstrate that thecae incubated from cysts of Peridinium wisconsinense Eddy have six cingular plates and a protuberant apical pore complex characteristic of the family Thoracosphaeraceae Schiller 1930 emend. Tangen in Tangen et al . 1982. Small subunit ribosomal DNA (SSU rDNA) and internal transcribed spacer (ITS) sequences confirm the close genetic similarity with Chimonodinium lomnickii (Wo?oszyńska) Craveiro, Calado, Daugbjerg, Gert Hansen & Moestrup and with species recently reassigned to the genus Apocalathium Craveiro, Daugbjerg, Moestrup & Calado that was inferred from previously published LSU rDNA analysis of cysts of P. wisconsinense . Despite sharing identical tabulation with the thoracosphaeracean genera Chimonodinium Craveiro, Calado, Daugbjerg, Gert Hansen & Moestrup and Apocalathium , substantial morphological differences in the morphology of both the thecate and cyst stages of P. wisconsinense led us to reassign this species to the genus Fusiperidinium gen. nov. The phylogenetic position of Fusiperidinium wisconsinense comb. nov., inferred from concatenated data of SSU and LSU sequences, suggests that it evolved from the brackish Scrippsiella lineage, independently of the transition that produced the family Peridiniaceae. Cysts described as Geiselodinium tyonekensis Engelhardt from nonmarine strata from Alaska are apparently identical to the resistant cysts produced by F. wisconsinense . The palynologically‐constrained late Middle Miocene age for the Tyonek Formation provides a minimum age of 11.6 Ma for the evolution of this lineage, coinciding with a rapid glacioeustatic decline in sea level. Our findings also call into question the inclusion of the family Thoracosphaeraceae within the order Peridiniales Haeckel. 相似文献
10.
Bente Edvardsen Wenche Eikrem Kamran Shalchian‐Tabrizi Ingvild Riisberg Geir Johnsen Lars Naustvoll Jahn Throndsen 《Journal of phycology》2007,43(5):1054-1070
Since 1998, a heterokont flagellate initially named Chattonella aff. verruculosa has formed recurrent extensive blooms in the North Sea and the Skagerrak, causing fish mortalities. Cells were isolated from the 2001 bloom off the south coast of Norway, and monoalgal cultures were established and compared with the Chattonella verruculosa Y. Hara et Chihara reference strain NIES 670 from Japan. The cells in Norwegian cultured isolates were very variable in size and form, being large oblong (up to 34 μm long) to small rounded (5–9 μm in diameter) with two unequal flagella, numerous chloroplasts, and mucocysts. The SSU and partial LSU rDNA sequences of strains from Norway and Japan were compared and differed by 0.4% (SSU) and 1.3% (LSU), respectively. Five strains from Norway were identical in the LSU rDNA region. Phylogenetic analyses based on heterokont SSU and concatenated SSU + LSU rDNA sequences placed C. aff. verruculosa and the Japanese C. verruculosa within the clade of Dictyochophyceae, with the picoflagellate Florenciella parvula Eikrem as the closest relative. Ultrastructure, morphology, and pigment composition supported this affinity. We propose the name Verrucophora farcimen sp. et gen. nov. for this flagellate and systematically place it within the class Dictyochophyceae. Our studies also show that C. verruculosa from Japan is genetically and morphologically different but closely related to V. farcimen. The species is transferred from the class Raphidophyceae to the class Dictyochophyceae and renamed Verrucophora verruculosa. We propose a new order, Florenciellales, to accommodate V. farcimen, V. verruculosa, and F. parvula. 相似文献
11.
12.
Ryohei Ueno Nobutaka Hanagata Naoto Urano Motofumi Suzuki 《Journal of phycology》2005,41(6):1268-1280
Species of the heterotrophic green microalgal genus Prototheca and related taxa were phylogenetically analyzed based on the nuclear small subunit (SSU) and the 5′ end of the large subunit (LSU) rRNA gene (rDNA) sequences. We propose restricting the genus Prototheca to the four species: P. moriformis Krüger, P. stagnora (Cooke) Pore, P. ulmea Pore, and P. zopfii Krüger. The main diagnostic feature of these taxa is the absence of growth on trehalose.Of these, it was suggested that P. moriformis should be merged into P. zopfii; P. moriformis and three varieties of P. zopfii constituted a paraphyletic assemblage with estimated short evolutionary distances. The trehalose‐assimilating strains (Prototheca wickerhamii Tubaki et Soneda strains and Auxenochlorella protothecoides (Krüger) Kalina et Pun?ochá?ová SAG 211‐7a), together with an invertebrate pathogen Helicosporidium sp., diverged before the radiation of the four species of Prototheca in the SSU rDNA and composite (SSU rDNA plus LSU rDNA) analyses. Comparison between the results from physiological data in this work (fermentative pattern) and those described earlier (growth requirements) lead us to propose a hypothesis that the phenotypic variation, which did not represent diagnostic characters for species delimitation, may reflect the history of genetic diversification within the genus Prototheca as inferred from rDNA sequence characters. 相似文献
13.
Diana Sarno Wiebe H. C. F. Kooistra Sergio Balzano Paul E. Hargraves Adriana Zingone 《Journal of phycology》2007,43(1):156-170
Skeletonema costatum (Grev.) Cleve emend. Zingone et Sarno and S. grevillei Sarno et Zingone were known only from the type material collected from Hong Kong waters more than a century ago. Both species have now been collected as live material, and their morphology and phylogenetic position are investigated in this study. Eight Skeletonema strains isolated from Florida, USA; Uruguay; and Brazil are attributed to S. costatum, while one strain from Oman is ascribed to S. grevillei based on morphological similarity to the type material of these species. In addition, a new Skeletonema species, S. ardens Sarno et Zingone, is described for a strain from Singapore and two from northern Australian waters. Skeletonema ardens has terminal fultoportula processes ending in a tapered, undulate protrusion and long intercalary fultoportulae with 1:1 junctions. The rimoportula of terminal valves is located at the margin of the valve face. No major morphological variations were observed within S. grevillei and S. ardens along a salinity gradient, whereas in S. costatum, the processes shortened and the valves came into close contact at low salinities, as already described for S. subsalsum (Cleve) Bethge. Consistent with their morphology, Skeletonema costatum and Skeletonema subsalsum also had similar rDNA sequences. Skeletonema grevillei and S. ardens were distinct in the large subunit (LSU) phylogeny. Skeletonema ardens exhibited consistent intraspecific genetic differences in both the LSU and small subunit (SSU) rDNA. 相似文献
14.
用核糖体SSURdna全序列对蝗总科(Acridoidea)进行了分子系统学研究。依据测定的8种蝗虫的SSU Rdna全序列 (平均 1.844 bp),并从GenBank中选取了6种内群种类和2种外群种类的SSU Rdna同源序列,进行序列分析。利用Clustal、MEGA 和 PHYLIP 软件构建分子系统树(距离邻接法Neighbor-Joining,NJ;最小进化法 Minimum Evolution)。结果显示: (1) 蝗总科是一个单系类群;(2) 锥头蝗科(Chrotogonidae)和瘤锥蝗科(Pyrgomorphidea)亲缘关系较近,为蝗总科最原始的类群;(3) 网翅蝗科(Arcypteridae)和槌角蝗科(Gomphoceridae)有较近的亲缘关系; (4) 斑翅蝗科 (Oedipodidae)为最进化的类群; (5) SSU Rdna序列保守性强,转换transition)取代的速率大于或接近颠换(transversion)取代的速率;(6) 在系统树中,总科首先分离,大多数同科不同属的类群以高置信度聚合在一起,说明SSU Rdna序列适合用于蝗总科的系统发育关系分析。 相似文献
15.
Rafael Carballeira Rosa Trobajo Manel Leira Xavier Benito Shinya Sato David G. Mann 《欧洲藻类学杂志》2017,52(3):342-359
A previously unknown member of the Bacillariaceae was discovered almost simultaneously in four different brackish coastal wetlands on the Atlantic and Mediterranean coasts of the Iberian Peninsula. It appears to tolerate a wide range of salinities but was never common in samples where it occurred. The frustules were consistently hantzschioid (i.e. with the raphe systems always on the same side of the frustule) and the valve outline was asymmetrical about the apical plane, two features that have until recently been considered characteristic of Hantzschia. Molecular phylogenies based on rbcL and LSU rDNA indicated, however, that the new species does not belong in Hantzschia but among the several disparate lineages that comprise the paraphyletic genus Nitzschia. This finding, coupled with the recent discovery of other diatoms with constant hantzschioid symmetry but with a morphology very similar to the type species of Nitzschia, is discussed in relation to the status and characterization of Hantzschia as an independent genus. It is concluded that, while a core of hantzschioid species may exist that can be classified together, corresponding to the traditional understanding of the genus Hantzschia, there is no single morphological feature common to all of them that can be used to diagnose the group and differentiate it from the various hantzschioid lineages that are separate from true Hantzschia and currently placed in e.g. Nitzschia or Cymbellonitzschia. Testing whether a hantzschioid species does or does not belong to Hantzschia will in many cases require molecular evidence. Although the new coastal species does not belong to the same lineage as the type species of Nitzschia, N. sigmoidea, it is described for the moment as N. varelae Carballeira, D.G. Mann & Trobajo, sp. nov., until there is a better understanding of generic limits in the Bacillariaceae following a wider molecular and morphological survey of that family. 相似文献
16.
Ceramium inkyuii sp. nov. is newly described based on samples collected from the east coast of Korea and compared with similar species such as C. paniculatum and C. tenerrimum. The new species is characterized by pseudo‐dichotomously branched thalli with a twist in the upper part, a single row of spines on the abaxial side, strongly inrolled apices, and the presence of gland cells. In contrast, C. paniculatum has alternate branches and lacks gland cells, and C. tenerrimum is spineless and also lacks gland cells. Ceramium inkyuii was observed to be an annual species producing tetrasporangia in the spring to summer and cystocarps in the fall. Plastid‐encoded rbcL and nuclear small subunit (SSU) rDNA sequences were determined in four samples of C. inkyuii from different locations and six samples of four putative relatives. All four C. inkyuii replicates from three different locations had identical sequences of each gene, and the interspecific sequence divergences were enough to warrant its natural entity. The phylogenies of the rbcL and SSU rDNA sequences also indicate the monophyly of C. inkyuii. The spinous C. inkyuii was more closely related to the spineless C. tenerrimum than to the spinous C. paniculatum. 相似文献
17.
Orlando Necchi Jr. Auro Silva Garcia Fo Eric D. Salomaki John A. West Marina Aboal Morgan L. Vis 《欧洲藻类学杂志》2013,48(2):152-162
Twenty-five specimens of the freshwater red alga Compsopogon were collected from locations in North America, South America, Europe, Asia, Australasia and Oceania, and from an aquarium, with the goal of determining genetic diversity among specimens and ascertaining the number of phylogenetic species. Specimens were morphologically identified as having either the ‘caeruleus’ morphology, with regular polyhedral cortical cells, or the ‘leptoclados’ morphology, with irregular cortical cells with rhizoidal outgrowths. The ‘leptoclados’ morphology has been used by some researchers to distinguish the genus Compsopogonopsis from Compsopogon, or at least to distinguish C. leptoclados from other Compsopogon species. Sequence data for the rbcL gene and cox1 barcoding region were obtained for most specimens. In addition, SSU and partial LSU (barcode) rDNA were explored for a few specimens, but all sequences were identical. For the 25 newly generated and eight previously published rbcL gene data, there were seven unique haplotypes, but the sequence divergence was very low (≤7 bp, ≤ 0.7%). One haplotype was widespread, represented by 21 specimens from diverse locations in all regions sampled. Likewise, the 22 new and one previously published cox1 barcode region sequences yielded seven unique haplotypes with little sequence divergence (≤13 bp, ≤ 2.0%). One haplotype was widespread, being shared among 16 specimens from all regions. The combined molecular and morphological data showed no genetic differentiation between the ‘caeruleus’ and ‘leptoclados’ morphologies. The ubiquitous distribution of Compsopogon in tropical/subtropical regions and its low genetic variation are probably facilitated by the alga's ability to tolerate a wide range of stream conditions and its propagation via asexual spores. Given the findings of previous culture-based studies, morphometric research and field observations, coupled with the results of our study, we conclude there is only a single monospecific genus worldwide and that the species is correctly called C. caeruleus, since this is the oldest validly published name; all other previously described species of Compsopogon and Compsopogonopsis are synonyms. 相似文献
18.
Primer sequences are described for amplifying and sequencing a large fragment (approximately 2500 b.p.) of the nuclear-encoded large-subunit ribosomal RNA gene (LSU) from red algae. In comparison to RuBisCo large-subunit gene (rbcL) and nuclear-encoded small-subunit ribosomal RNA gene (SSU) sequence data, LSU sequence data was intermediate in the number of phylogenetically informative positions and sequence divergence. Parsimony analysis of LSU sequences for 16 Gelidiales species resolved some nodes unresolved in rbcL and SSU parsimony trees. An analysis of LSU sequences from 13 species of red algae classified in 11 orders suggests that this gene may be useful in studies of higher-level relationships of red algae. 相似文献
19.
The status of the green algal genera Haematococcus and Stephanosphaera has been a source of debate among algal systematists. A phylogenetic alliance between Haematococcus (sensu lato) and the colonial Stephanosphaera was affirmed by earlier molecular phylogenetic investigations. Although the data suggested that the genus Haematococcus may not be a monophyletic group, taxon sampling limited the scope of any potential taxonomic revision. Results from new molecular phylogenetic analyses of data from the 18S and 26S rRNA genes support the establishment of a separate genus, Balticola, as originally proposed by Droop in 1956. Haematococcus remains as a valid genus, with H. pluvialis as its only member. The monotypic status of H. pluvialis is supported both by molecular phylogenetic analyses of the ribosomal RNA genes and assessments of molecular evolution in the ITS2 sequences of H. pluvialis strains. The near-complete absence of compensatory base changes in a sequence-structure analysis of the highly variable ITS2 gene from more than 40 geographically diverse isolates of H. pluvialis corroborates the unity of the species inferred from molecular phylogenetic analyses of 18S and 26S rRNA gene sequence data. 相似文献
20.
During a survey of insect gut micro-organisms, we consistently isolated Pichia stipitis-like yeasts (Fungi: Ascomycota, Saccharomycetes) from the wood-ingesting beetles, Odontotaenius disjunctus and Verres sternbergianus (Coleoptera: Passalidae). The yeasts were isolated from passalid beetles over a wide area, including the eastern and midwestern USA and Panama. Phylogenetic analyses of the nuclear encoded small and large subunit rRNA gene (rDNA) sequences distinguished a well-supported clade consisting of the passalid yeasts and Pichia stipitis, P. segobiensis, Candida shehatae and C. ergatensis. Members of this clade have the ability to ferment and assimilate xylose or to hydrolyse xylan, major components of the polysaccharide, hemicellulose. Sexual reproduction was present in the passalid isolates but was rare among the gut yeasts of other beetles to which they were compared. Minor genetic and phenotypic variation among some of the passalid yeasts was detected using markers from the internal transcribed spacer region of the rDNA repeat unit, morphology, and in vitro metabolic tests. The consistent association of xylose-fermenting yeasts of almost identical genotypes with passalid beetles across a broad geographical distribution, suggests a significant symbiotic association. 相似文献