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Phylogenetic position of Alwisia and its three species within Reticulariaceae. Tree based on partial SSU sequences (ca. 700 bp, 374 alignable positions retained) constructed with RaxML and rooted with the genus Ceratiomyxa. Seventeen new sequences (in boldface) were added to known sequences of the Reticulariaceae. The abbreviation ''gt'' denotes the genotype (see SUPPLEMENTARY INFORMATION 1), followed by the accession number of one representative sample and the number of specimens with the respective genotype in brackets. ML bootstrap replicates/Bayesian posterior probabilities are shown for each branch; a dash indicates a lack of resolution in the respective ML tree. Bar indicates the fraction of substitutions per site.
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Based on morphological investigations and a phylogeny constructed with partial sequences of the SSU rDNA gene, we revalidate the genus Alwisia and propose the combination Alwisia bombarda Berk. & Broome to be used against Tubifera bombarda (Berk. & Broome) G.W. Martin. Two new species, Alwisia morula and A. repens, are described based on material c...
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Context 1
... the Reticulariaceae, Alwisia bombarda is most similar to A. morula and A. repens. A phylogeny based on partial sequences of the SSU rRNA gene confirms this (FIG. 22). Alwisia bombarda has erect, long fibrous stalks and sporothecae united by a common stalk to small clusters resembling those of Metatrichia Ing. Because sporocarps remain free, these fructifications may be considered as peculiar pseudoaethalia or as clustered ...
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... whether every clearly separated SSU genotype in myxomycetes should be described as a species of its own. We think that only both morphological and genetic differences justify describing a new species in the myxomycetes. As seen from the molecular side, unusually large genetic distances, such as between the two genotypes of Tubifera dictyoderma (FIG. 22) warrants a closer examination of the respective ...
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... speciation in Reticulariaceae.-Our phylogeny, obtained with partial SSU sequences from 92 speci- mens of the genera Tubifera and Reticularia, revealed a cryptic speciation within these genera (FIG. 22). Without support by molecular investigations, two of these genotypes recently were described as separate species, the first of which was Tubifera applanata ( Leontyev andFefelov 2009, 2012) and the second Reticularia dudkae (Leontyev and Moreno 2011). Both turned out to be represented by well circum- scribed SSU genotypes. Tubifera ...
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... second Reticularia dudkae (Leontyev and Moreno 2011). Both turned out to be represented by well circum- scribed SSU genotypes. Tubifera applanata is repre- sented by 17 identical sequences and is related to the tropical species T. microsperma and T. dimorphotheca. Reticularia dudkae produced six identical sequences and clustered within Tubifera (FIG. 22), thus being a candidate for a new taxonomic combination. The specimens determined to be T. ferruginosa form several distinct genotypes (gt). Most common was T. ferruginosa gt1, where all 27 investigated specimens produce identical sequences that deviate in one position (base 4) from the published reference sequence EF513171 (Fiore- ...
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... dictyoderma also separates into two very different genotypes (genetic distance 0.531), both re- presented by two specimens with identical sequences. Moreover, this species forms a separate clade, distant not only from another species of Tubifera but even from Lycogala and Reticularia (FIG. 22). More material is needed to study this species and the morphologically close taxon T. casparyi (Rostaf.) T. ...
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Based on a morphological investigation of a series of specimens collected in New South Wales and Tasmania and a phylogeny constructed with partial 18S ribosomal RNA gene sequences, we describe a new species Alwisia lloydiae; the fourth species within the recently re-validated genus Alwisia. This new species is characterized by short ovate sporothec...
Citations
... These studies increased the number of species of the Reticulariaceae to 52 (Lado 2005), the number of genera to six and one more genus, Licaethalium, was re-erected within the Cribrariaceae for species, previously classified within Reticulariaceae (Leontyev et al. 2019b). Fifteen species of the genus Lycogala, nine species and one subspecies of the genus Tubifera, three species of the genus Alwisia and two species of the genus Siphoptychium were described (Leontyev et al. 2014a, Leontyev et al. 2014b,Leontyev et al. 2015, Leontyev et al. 2019a, Lloyd et al. 2019, Leontyev et al. 2022a, Leontyev et al. 2022b, Leontyev and Schnittler 2023b. The classical species L. epidendrum and T. ferruginosa have proven to represent species complexes, which contain at least 13 and 76 biological species, respectively (Leontyev and Schnittler 2023a). ...
... Rostaf., which exhibit distinct morphological characteristics, were described as early as the 19 century (Lado 2005, Leontyev et al. 2019b. By adopting an integrative approach, at the core of which was the quest for morphological features correlated with molecular barcoding data, we were able to delimit species previously unknown to science that are widespread in extensively studied regions like Europe and North America (Leontyev et al. 2014b, Leontyev et al. 2015). ...
As a result of the ten years (2012–2022) work under the critical revision of the genera of Reticulariaceae, a set of papers was published. Collection data of hundreds of specimens, used as a material for these studies, were provided as supplements of corresponding papers, but remained unpublished in biodiversity databases.
Here, we represent an occurrence dataset "Barcoded Reticulariaceae of the World", published in GBIF. It includes data on 523 myxomycete collections (including 36 types) gathered from five continents and spanning 24 countries. The dataset encompasses 43 distinct species and one subspecies of myxomycetes, including rare, endemic, and recently-described taxa. Species included to the database mainly belong to the genera Alwisia , Lycogala , Reticularia , Siphoptychium , Thecotubifera and Tubifera (Reticulariaceae), but as well Lindbladia and Licaethalium (Cribrariaceae). Nearly all of the research material, with the exception of several old collections, underwent molecular barcoding, primarily involving the 18S rDNA gene, but also the elongation factor 1α gene and mitochondrial cytochrome oxidase subunit I gene. For those sequences that are stored in the NCBI GenBank, accession numbers are provided in the dataset. Newly-described species make up a significant part of the studied herbarium collections; many of them can be characterised as common for their region. A particularly high level of taxonomic novelty is observed in Australia, which may be explained by the endemism of the local myxomycete biota.
... The first evidence of this was reported in the 19th century by Berkeley (1859), who described three species of myxomycetes from Tasmania (Lamproderma echinulatum, Oligonema verrucosum, Prototrichia metallica). Recent studies (Leontyev et al. 2014;Lloyd et al. 2019) added four more species (Alwisia lloydiae, Tubifera glareata, T. tomentosa, T. vanderheuliae). In this paper, we describe another, very peculiar species occurring in Tasmania, southeastern Australia, and New Zealand. ...
A new genus and species of myxomycete, Tasmaniomyxa umbilicata, is described based on numerous observations in Tasmania and additional records from southeastern Australia and New Zealand. The new taxon is characterized by an unusual combination of characters from two families: Lamprodermataceae and Didymiaceae. With Lamprodermataceae the species shares limeless sporocarps, a shining membranous peridium, an epihypothallic stalk, and a cylindrical columella. Like Didymiaceae, it has a soft, flaccid, sparsely branched capillitium, with rough tubular threads that contain fusiform nodes and are firmly connected to the peridium. Other characters of T. umbilicata that also occur in many Didymiaceae are the peridium dehiscing into petaloid lobes, the yellow, motile plasmodium, and the spores ornamented with larger, grouped and smaller, scattered warts. The transitional position of the new taxon is reflected by a three-gene phylogeny, which places T. umbilicata at the base of the branch of all lime-containing Physarales, thus justifying its description as a monotypic genus.
... Our study revealed systematic problems with molecular barcoding of myxomycetes of the order Reticulariales (Leontyev et al. 2019b), for which it remains impossible to find universal primers and, accordingly, to establish a routine sequencing of marker genes (Leontyev et al. 2014(Leontyev et al. , 2015(Leontyev et al. , 2019a. For Lycogala spp., a sequential trial with several primer pairs (mostly S1a/R_SIP and SF1/SR6Bright) allowed us to achieve a relatively high success rate. ...
Lycogala epidendrum is one of the most widely known myxomycete species and the first-ever discovered representative of this group. Using 687 original DNA sequences from 330 herbarium specimens from Europe, Asia, North and Central America, and Australia, we constructed the first detailed phylogenies of the genus Lycogala, based on two independently inherited genetic markers, the ribosome small subunit 18S rRNA nuclear gene (18S rDNA) and the mitochondrial cytochrome oxidase subunit I gene (COI). In both phylogenies, L. epidendrum appeared to be a polyphyletic group, represented by numerous clades. The four other recognized species of the genus (L. confusum, L. conicum, L. exiguum, and L. flavofuscum) are scattered between branches corresponding to L. epidendrum. A barcode gap analysis revealed 60 18S rDNA phylogroups of L. epidendrum, which are distant from each other not less than from other species of the genus Lycogala. For 18 of these phylogroups with both 18S rDNA and COI sequences available, recombination patterns were analyzed to test for reproductive isolation. In contrast to the results of a simulation assuming panmixis, no crossing between ribosomal and mitochondrial phylogroups was found, thus allowing the conclusion that all tested phylogroups represent biospecies. More than one third (39.6%) of the studied specimens share a single 18S rDNA phylogroup, which we consider to be L. epidendrum s. str. This group displays the broadest geographic distribution and the highest intraspecific genetic variability. Nearly all (93.3%) of the remaining non-singleton 18S rDNA phylogroups are restricted to certain continents or even regions. At the same time, various reproductively isolated phylogroups occur sympatric at a given location.
... Aethalia or pseudoaethalia characterize approximately 80% of the species, and the majority exhibit pseudocapillitium and reticulated spores. In the delimitation of the family proposed by Leontyev et al. (2014aLeontyev et al. ( , 2019, the genus Dictydiaethalium was excluded, and the presence of capillitium was recognized in Alwisia and Lycogala species based on analyses by McHugh & Reid (2008) and Leontyev et al. (2014b, c). ...
In the second decade of the 20th century, Lycogala flavofuscum aethalia were collected in Brazil by Rev. Johannes Rick. Despite being easily observed in the field because of the size of the aethalia, the species was only found again in 2019, on a live trunk of Sebastiania commersoniana (branquilho, Euphorbiaceae) in the municipality of São Gabriel, Rio Grande do Sul, Brazil. The description and illustration of recently collected material, and data on exsiccates collected by Rick from Brazil, deposited at Herbarium BPI, are presented herein. Information on substrates, microhabitats, and geographic distribution of the species in the Neotropics is provided based on bibliographical sources and herbarium catalogs. Considering the large time interval between records and the restricted area of occurrence in the country, its inclusion as near threatened in the Brazilian Red List is recommended.
... Therefore, herbarium collections of myxomycetes usually contain small quantity of biological material. Since the first attempt to isolate DNA from herbarium specimens of myxomycetes (Martín, Lado & Johansen, 2003), there is an increasing interest to barcode myxomycete species and examine phylogenetic relationships between the taxonomic groups using molecular data and based on field-collected specimens deposited in herbarium collections (e.g., Fiore-Donno et al., 2005;Fiore-Donno et al., 2012;Fiore-Donno et al., 2013;Novozhilov et al., 2013b;Leontyev et al., 2014;Leontyev, Schnittler & Stephenson, 2015;Schnittler et al., 2017)). Almost all studies using molecular methods in myxomycetes are based on a large quantity of biological material derived from several up to 30 sporocarps per collection (see Table 1 for examples and references). ...
Herbarium collections provide an essential basis for a wide array of biological research and, with development of DNA-based methods, they have become an invaluable material for genetic analyses. Yet, the use of such material is hindered by technical limitations related to DNA degradation and to quantity of biological material. The latter is inherent for some biological groups, as best exemplified by myxomycetes which form minute sporophores. It is estimated that ca. two-thirds of myxomycete taxa are represented by extremely scanty material. As DNA isolation methods applied so far in myxomycete studies require destructive sampling of many sporophores, a large part of described diversity of the group remains unavailable for phylogenetic studies or barcoding. Here, we tested several procedures of DNA isolation and amplification to seek for an efficient and possibly non-destructive method of sampling. Tests were based on herbarium specimens of 19 species representing different taxonomic orders. We assayed several variants of isolation based on silica gel membrane columns, and a newly designed procedure using highly reduced amount of biological material (small portion of spores), based on fine disruption of spores and direct PCR. While the most frequently used column-based method led to PCR success in 89.5% of samples when a large amount of material was used, its performance dropped to 52% when based on single sporophores. Single sporophores provided amplicons in 89.5% of samples when using a kit dedicated to low-amount DNA samples. Our new procedure appeared the most effective (94.7%) while it used only a small fraction of spores, being nearly non-destructive; it was also the most cost-effective. We thus demonstrate that combination of adequate handling of spore micro-disruption coupled with application of direct PCR can be an efficient way to circumvent technical limitations for genetic studies in myxomycetes and thus can substantially improve taxon sampling for phylogeny and barcoding. Additionally, this approach gives a unique possibility to apply both molecular and morphological assays to the same structure (sporophore), which then can be further stored as documentation.
... The capillitium is of obvious adaptive significance as it effectively prevents the spore mass against detaching from the stalk and allows spores to disperse separately as they dry out 5 . An interesting adaptive trend in myxomycetes is from solitary to compound fruiting bodies and from stalked forms to sessile forms, typically with corresponding change of spore dissemination mode from wind dispersal, often facilitated by a well-developed capillitium, to dispersal via rainwater and/ or insects 43,44 . ...
Myxomycetes constitute a group within the Amoebozoa well known for their motile plasmodia and morphologically complex fruiting bodies. One obstacle hindering studies of myxomycete evolution is that their fossils are exceedingly rare, so evolutionary analyses of this supposedly ancient lineage of amoebozoans are restricted to extant taxa. Molecular data have significantly advanced myxomycete systematics, but the evolutionary history of individual lineages and their ecological adaptations remain unknown. Here, we report exquisitely preserved myxomycete sporocarps in amber from Myanmar, ca. 100 million years old, one of the few fossil myxomycetes, and the only definitive Mesozoic one. Six densely-arranged stalked sporocarps were engulfed in tree resin while young, with almost the entire spore mass still inside the sporotheca. All morphological features are indistinguishable from those of the modern, cosmopolitan genus Stemonitis, demonstrating that sporocarp morphology has been static since at least the mid-Cretaceous. The ability of myxomycetes to develop into dormant stages, which can last years, may account for the phenotypic stasis between living Stemonitis species and this fossil one, similar to the situation found in other organisms that have cryptobiosis. We also interpret Stemonitis morphological stasis as evidence of strong environmental selection favouring the maintenance of adaptations that promote wind dispersal.
... Among these, nine unique 18S rDNA genotypes were recognized (within one genotype all sequences were identical). To reconstruct the 18S rDNA phylogeny, a total of 98 sequences were analyzed, including 9 unique sequences obtained in this study, 53 sequences of other members of the family Reticulariaceae, and 36 sequences of Lucisporomycetidae from other families, reported in previous studies (Fiore-Donno et al. 2013;Leontyev et al. 2014aLeontyev et al. , 2014cLeontyev et al. , 2015Leontyev et al. , 2019. Five species of the family Cribrariaceae were used as outgroups. ...
The genus Siphoptychium is resurrected on the basis of comparative morphology and phylogeny of partial nuc 18S rDNA (18S) and translation elongation factor 1-alpha (EF1A) nucleotide sequences. The genus is characterized by the firm upper surface of the pseudoaethalium, accreted but easily separable sporothecae, a tubular or fibrous columella, and spores with a reticulate ornamentation consisting of 7–9 meshes across the diameter. In addition to the currently known single species S. casparyi (= Tubifera casparyi), two new members of Siphoptychium are described: S. violaceum from coniferous forests of Europe, east Asia, and southeast Asia, and S. reticulatum from temperate and subarctic regions of North America and alpine regions of Europe. A second genus, Thecotubifera, is described to accommodate Tubifera dictyoderma. The fruiting body of this species is transitional between a pseudoaethalium and a true aethalium. It is covered by a contiguous membranous cortex formed by the fused tips of the sporothecae, a feature typical for aethalia. However, the inner portions of sporothecae remain discernible, a feature more typical for pseudoaethalia. Columellae of Th. dictyoderma are formed by perforated plates, and the spores have a reticulate ornamentation consisting of 2–5 meshes across the diameter. For Th. dictyoderma, we could confirm records only for tropical regions and Japan, whereas all studied European specimens, including those mentioned in current monographs, represent species of Siphoptychium.
... The new circumscription of the order (Leontyev et al. 2014a) includes the genera Alwisia, Lycogala, Tubifera and Reticularia. The first two of them possess a true capillitium (McHugh and Reid 2008, Leontyev et al. 2014b, c, Leontyev 2016, and this is reflected in the emended description of the group. ...
The traditional classification of the Myxomycetes (Myxogastrea) into five orders (Echinosteliales, Liceales, Trichiales, Stemonitidales and Physarales), used in all monographs published since 1945, does not properly reflect evolutionary relationships within the group. Reviewing all published phylogenies for myxomycete subgroups together with a 18S rDNA phylogeny of the entire group serving as an illustration, we suggest a revised hierarchical classification, in which taxa of higher ranks are formally named according to the International Code of Nomenclature for algae, fungi and plants. In addition, informal zoological names are provided. The exosporous genus Ceratiomyxa, together with some protosteloid amoebae, constitute the class Ceratiomyxomycetes. The class Myxomycetes is divided into a bright-and a dark-spored clade, now formally named as subclasses Lucisporomycetidae and Columellomycetidae, respectively. For bright-spored myxomycetes, four orders are proposed: Cribrariales (considered as a basal group), Reticulariales, a narrowly circumscribed Liceales and Trichiales. The dark-spored myxomycetes include five orders: Echinosteliales (considered as a basal group), Clastodermatales, Meridermatales, a more narrowly circumscribed Stemonitidales and Physarales (including as well most of the traditional Stemonitidales with durable peridia). Molecular data provide evidence that conspicuous morphological characters such as solitary versus compound fructifications or presence versus absence of a stalk are overestimated. Details of the capillitium and peridium, and especially how these structures are connected to each other, seem to reflect evolutionary relationships much better than many characters which have been used in the past.
... The present record represents the first one for the American continent and the entire Neotropical region and widely broadens the known distribution of the species. Interestingly, this specimen was found in the Cerro de la Muerte area, previously reported as a region where myxomycete assemblages include species traditionally associated with temperate areas (Leontyev et al. 2014, Rojas et al. 2015. The affinity of this species for bryophytes and colder wet environments is supported by the present study. ...
During field surveys of myxomycetes in Central America, 2 previously unrecorded genera and 4 species, viz. Craterium muscorum Ing, Dictydiaethalium dictyosporum Nann.-Bremek., Physarina echinocephala Höhn, and Stemonaria rufipes , were identified. Some of these are new for the Neotropics. These noteworthy range extensions of these species to the Mesoamerican biodiversity hotspot adds to our knowledge of rarely recorded myxomycetes worldwide. Images of the more relevant taxonomic characters are provided, and for some species, this is the first illustrations with macro and microphotographs and scanning electron microscopy images.
... The geographical criteria on small Caribbean islands provided by Lado & Wrigley de Basanta (2008) in their review of Neotropical myxomycetes was also followed in the present study. The latter publication and a series of subsequent reports published by different authors on the regional countries considered herein (see Estrada-Torres et al. 2009, Izarduy et al. 2009, Montes 2009, Moreno et al. 2009, 2010, b, c, d, 2012, 2013, b, c, 2017, Poulain et al. 2011, Agra et al. 2014, Leontyev et al. 2014, Rojas & Calvo 2014, Rojas & Doss 2014, Salazar-Márquez et al. 2014, Walker et al. 2014, b, Lizárraga et al. 2015a, b, c, Rojas & Valverde 2015, Dagamac et al. 2017) has been used to update the distribution of all species. Other annotations were provided for selected species whose vouchers were not found but for which reliable information was available, or for species not previously included in . ...
... Despite all the results showing the shortcomings for ecological analysis of the sampling carried out in Costa Rica, the results provided herein also showed a very interesting list of species. As could have been noted in the references provided with the comments to selected species, Costa Rica has been a territory where five new species of myxomycetes have been described in the last seven years (see Leontyev et al. 2014, Moreno et al. 2010, Walker et al. 2014). Also, with the recent reports of Craterium muscorum, Dictydiaethalium dictyosporum, Physarina echinocephala and Stemonaria rufipes ( Lado et al. 2018), these elusive species were fully documented, and their distributional range largely widened to encompass the Neotropical area. ...
An updated analysis of myxomycete diversity in Costa Rica is provided herein as a product of an extensive data mining-based study intended to normalize the information on myxomycetes from that country. Within the context of the Mesoamerican Biodiversity Hotspot, the 242 species of myxomycetes are reported herein demonstrated the potential of this area for diversity-based studies. For the construction of the updated species list, a thorough analysis that included examination of vouchers and a literature review has been carried out, and some species previously included in other works have been rejected due to inconsistencies or misidentification. The present contribution is essential to address the Wallacean shortfall within Mesoamerica and important for providing updated data on microorganisms in the context of regional biodiversity monitoring.
