Content uploaded by Ludwig Beenken
Author content
All content in this area was uploaded by Ludwig Beenken on Mar 01, 2017
Content may be subject to copyright.
A preview of the PDF is not available
Russula densifolia Sécr. ex Gil. + Fagus sylvatica L
Abstract and Figures
Short description: Ectomycorrhizae regularly monopodial-pinnate, light brown, with velvety surface, all
mantle layers plectenchymatous, all septa simple, outer and middle mantle layers with
gelatinous matrix, mantle bearing cystidia forming 1-2 thick-walled knobs, content of
some cystidia becoming dark brown in sulfo-vanillin; rhizomorphs running along roots,
with a peripheral layer of thick-walled hyphae and cystidia similar to those of the mantle
and with an inner part containing vessel-like and ladder-like hyphae with arched septa
in close distances.
Figures - uploaded by Ludwig Beenken
Author content
All figure content in this area was uploaded by Ludwig Beenken
Content may be subject to copyright.
... Our studies revealed the typical features of ectomycorrhizas, a hyphal mantle and a Hartig net in all the mycorrhizal morphotypes from the three tree species. Plectenchymatous hyphal mantles are common for the ectomycorrhizas of Thelephoraceae (Agerer, 1995), but there are also pseudoparenchymatous mantles (Raidl & Müller, 1996;Jakucs & Agerer, 1999, 2001. A heterogenous assemblage of mantle and rhizomorph types exists in the Russulaceae; characteristic of the genus Lactarius are smooth mantles with lactifers (Agerer, 1995). ...
... Henkel et al. (2000) described a similar morphology of the ectomycorrhizas Lactarius panuoides-Dicymbe altsonii from Guyana's western border with Brazil. According to the anatomy of the ectomycorrhiza the species of Russula can be divided in two groups: a group with plectenchymatous outer mantle layers with cystidia and a group with pseudoparenchymatous outer mantle layers without cystidia (Beenken, 2001). The ectomycorrhizas of Russula puiggarii possess a plectenchymatous outer mantle layer. ...
* Three members of the Nyctaginaceae, two Neea species and one Guapira species, occurred scattered within a very species-rich neotropical mountain rain forest. The three species were found to form ectomycorrhizas of very distinctive characters, while all other tree species examined formed arbuscular mycorrhizas. * The ectomycorrhizas were structurally typified according to light and transmission electron microscope investigations. The internal transcribed spacer (ITS) rDNA and part of the nuclear large subunit (LSU, 28S) rDNA of the mycorrhiza forming fungi were amplified and sequenced. Phylogenetic analyses were carried out. * Neea species 1 was found to form typical ectomycorrhizas with five different fungal species, Russula puiggarii, Lactarius sp., two Tomentella or Thelephora species, and one ascomycete. Neea species 2 and the Guapira species were associated with only one fungus each, a Tomentella/Thelephora species clustering closely together in an ITS-neighbour-joining tree. The long and fine rootlets of the Guapira species showed proximally a hyphal mantle and a Hartig net, but distally intracellular fungal colonization of the epidermis and root hair development. The ectomycorrhizal segments of the long roots of Neea species 2 displayed a hyphal mantle and a Hartig net around alive root-hair-like outgrowths of the epidermal cells. * The distribution and the evolution of ectomycorrhizas in the predominantly neotropic Nyctaginaceae are discussed.
... Moreover, close examination of ECM features revealed differences amongst Lactifluus species, a diversity supported by molecular analyses. This confirms, once again, the informative value of mycorrhizal structure, when joined to evidence coming from other approaches, in resolving phylogenetic relationships in ECM fungi (Nuytinck et al. 2004, Mleczko & Ronikier 2007, Benkeen 2011).SV, surface view; ML, middle layer; IV, inner view; RHIZ, rhizomorphs; cys, cystidia; dpl, densely plectenchymatous; hd, highly differentiated; hh, extramatrical hyphae; hn, hyphal net; lact, lactifers; m-p, monopodial pinnate; m-pyr, monopodial pyramidal; pl, plectenchymatous; ps, pseudoparenchymatous; undif, undifferentiated. ...
We describe the morpho-anatomical features of the ectomycorrhizas (ECMs) formed
by Lactifluus rugatus on Cistus, a genus of about 20 species of woody shrubs typical of the
Mediterranean maquis. The description of L. rugatus mycorrhizas on Cistus is the first ECM
description of a species belonging to Lactifluus subgen. Pseudogymnocarpi. The ECM identity was
verified through molecular tools. Anatomically, the characteristic of L. rugatus mycorrhiza is the
presence of abundant, long “bottle-shaped” cystidia on mantle surface. Indeed, the overwhelming
majority of milkcap mycorrhizas are acystidiate. This is the third Lactarius/Lactifluus mycorrhiza
to have been described associated with Cistus, the others being Lactarius cistophilus and L.
tesquorum. The phylogenetic distance between all these taxa is reflected by the diversity of the
principal features of their ECMs, which share host-depending ECM features known for Cistus,
but are otherwise distinguishable on the host roots. Comparison of Lactifluus rugatus ECM with
those formed by L. vellereus and L. piperatus on Fagus reveals elevated intrageneric diversity
of mycorrhizal structures. Such a diversity is supported by analysis of ITS sequences of relevant
species within European Lactifluus species. Our study extends knowledge of Cistus mycorrhizal
biology and confirms the informative value of mycorrhizal structures in understanding phylogenetic
relationships in ECM fungi.
... The rhizomorph of L. megalopterus has a typical anatomy, which is otherwise known only from the rhizomorphs of Lactarius, Lactifluus and Russula (Russulaceae) (Beenken 2001a, b; russoloid rhizomorphs sensu Agerer 1999): it is divided into an outer layer and an inner part containing lactifers, vessel-like hyphae and densely septate hyphae with arched septa (ladder-like hyphae sensu Agerer 1999). Lactiferous hyphae are common in the rhizomorphs of Lactarius but are absent in Russula species and Lactifluus piperatus (Beenken 2001aBeenken , b, 2004). ...
A new sequestrate Lactarius species was found in a humid evergreen tropical rainforest dominated by Fabaceae of the subfamily Caesalpinioideae in Cameroon, Central Africa. It is described here as new to science and is named Lactarius megalopterus, referring to its spore ornamentation of extraordinarily high wings. Anatomical characters and molecular systematic analyses confirm its relationship to Lactarius subgenus Plinthogali. Phylogenetic analyses based on two nuclear DNA regions revealed its close relationship to Lactarius angiocarpus, which is also an angiocarpous species from Zambia in Africa. Molecular studies have shown that tuber-like, sequestrate sporocarps evolved independently in several lineages of Basidiomycota. The findings of sequestrate fungi in tropical rainforests raise questions regarding the evolutionary benefit of enclosing the spore-producing hymenium. The enclosure of spore-producing tissue has often been associated with the protection of the delicate hymenium against desiccation in arid habitats or against frost in cold habitats. However, these cannot be the selective factors in warm and humid areas like the tropics. This controversy is exemplarily studied and discussed in the family of Russulaceae, especially in the genus Lactarius. Characters shown by the angiocarpous sporocarp of the new Lactarius, such as thick-walled statismospores, an aromatic smell and mild taste, can be interpreted as adaptations to endozoochorous spore dispersal by mammals. Therefore, here we prefer the alternative hypothesis that sequestrate sporocarps are the result of adaptation to endozoochorous spore dispersal.
Die Fruchtkörper und Rhizomorphen von Gastrosporium simplex werden detailliert beschrieben. Die Rhizomorphen sind charakterisiert durch ampullenförmige Anschwellungen an den Septen innerer Hyphen, durch das Fehlen von gefäßartigen Hyphen mit sich auflösenden Septen, durch eine auffällige gelatinöse Matrix und einer dicken und dichten Lage von sternförmig zusammengesetzten Kristallen, sowie an Hand ballonförmiger, terminaler Hyphenzellen der Rhizomophenoberfläche. Besonders die ampullenförmig aufgetriebenen Hyphen sind von Bedeutung, da Rhizomorphen mit solchen Hyphen innerhalb der Hymenomycetes nicht weit verbreitet sind. Sie kommen offensichtlich generell in den nahe verwandten Ordnungen Gomphales, Geastrales und Gautieriales vor und sind wahrscheinlich auch für die Phallales charakteristisch sowie für die Gattungen Hydnum, Kavinia und für einige Arten der Gattungen Phlebiella und Trechispora. All diese Gruppen werden mit Gastrosporium simplex verglichen. Wir schließen, dass die Gastrosporiaceae wahrscheinlich einerseits mit dem Ordnungskomplex ‘Gomphales-Geastrales-Gautieriales’ und andererseits mit den Phallales verwandt sind. Aber wahrscheinlicher ist sogar eine nähere Verwandtschaft mit den Gattungen Ramaricium, Kavinia, Phlebiella und Trechispora. Denn Gastrosporium simplex bildet weder Acantho-Oleocystiden oder Acantho-Oleohyphen, die von sehr dünnwandigen, rundlichen, mit gelbem Inhalt gefüllten Zel-len begleitet werden – diese Merkmalskombination ist kennzeichnend für den Komplex ‘Gomphales-Geastrales-Gautieriales’ – noch formen sie in blasenförmigen Zellen die für Phallales bekannten, zusammengesetzten Kristalle. Die Unterschiede in der Glebaontogenie im Vergleich zu den Phallales rechtfertigt eine separate Ordnung Gastrosporiales. Eine Beziehung zu den Lycoperdales kann schon hinsichtlich der vollkommen abweichenden Glebaontogenie ausgeschlossen werden, aber auch wegen der grundsätzlich andersartigen Ontogenie der Rhizomorphen. Im Gegensatz zu den Gastrosporiales sind die Lycoperdales nämlich durch so genannte agaricoide Rhizomorphen gekennzeichnet, die dicke, gefäßartige Hyphen aufweisen, wie sie auch für die ihnen verwandten Gattungen Agaricus, Macrolepiota und Bovista kennzeichnend sind.The rhizomorphs of Gastrosporium simplex and some ideas to the systematic position of the Gastrosporiaceae (Hymenomycetes, Basidiomycota)Fruitbodies and rhizomorphs of Gastrosporium simplex are described in detail. The rhizomorphs are characterized by ampullate inflations at the septa of internal hyphae, by the lack of vessel-like hyphae with dissolving septa, by a prominent gelatinous matrix, and a dense and thick layer of star-like arranged crystals on their surface and by balloon-shaped to globose terminal cells. Particularly the hyphae with ampullate inflations are important, as rhizomorphs with such hyphae are not widely distributed within Hymenomycetes. They apparently consistently occur in the closely related orders Gomphales, Geastrales and Gautieriales and are probably also characteristic of Phallales and of the genera Hydnum, Kavinia,and some Phlebiella and Trechispora species. All these groups are compared with Gastrosporium simplex and it is concluded that Gastrosporiaceae are likely related on the one hand to the complex ‘Gomphales-Geastrales-Gautieriales’ and, on the other, to the Phallales. But possibly, there is even a closer relation to the resupinate genera Ramaricium, Kavinia, Phlebiella and Trechispora, as Gastrosporium simplex neither forms acantho-oleocystidia or acantho-oleohyphae accompanied by very thin-walled, roundish cells with yellowish contents – characteristic of the complex ‘Gomphales-Geastrales-Gautieriales’ – nor do they form composed crystals in globular cells as they are found in Phallales. The differences in gleba ontogeny as compared to that of Phallales justifies a separate order for Gastrosporiaceae. A relation to Lycoperdales can be excluded with respect to the gleba formation and with respect to the completely different ontogeny of the rhizomorphs as well. In contrast to Gastrosporiales, the Lycoperdales are characterized by so-called agaricoid rhizomorphs with vessel-like thick hyphae as they are known to occur also in their relatives Agaricus, Macrolepiota and Bovista.
Aproximately 5,000–6,000 fungal species form ectomyorrhizae (ECM), the symbiotic organs with roots of predominantly trees. The contributing fungi are not evenly distributed over the system of fungi. Within Basidiomycota exclusively Hymenomycetes and within Ascomycota exclusively Ascomycetes contribute to the symbiosis. Hymenomycetes play a big part, Ascomycetes a minor role; Zygomycetes only form exceptionally ECM. Responsible for ascomycetous ECM are mostly Pezizales with their hypogeous derivatives, whereas Boletales, Gomphales, Thelephorales, Amanitaceae, Cantharellaceae, Cortinariaceae, Russulaceae, and Tricholomataceae are the most important ectomycorrhizal relationships within Hymenomycetes. ECM, as transmitting organs between soil and roots, are transporting carbohydrates for growth of mycelium and fruitbodies from roots and have to satisfy the tree’s demand for water and nutrients. The latter task particularly influences the structure of ECM as nutrients are patchily distributed in the soil and saprotrophic as well as ectomycorrhizal fungi can act as strong competitors for nutrients. In focusing these requirements, ECM developed variously structured hyphal sheaths around the roots, the so-called mantles, and differently organized mycelium that emanates from the mantle, the so-called extramatrical mycelium. The mantles can be plectenchymatous consisting of loosely woven, differently arranged hyphae or they are densely packed, forming a pseudoparenchyma similar to the epidermis of leaves. The extramatrical mycelium grows either as simple scattered hyphae from the mantle into the soil or it can be united to undifferentiated rhizomorphs with a small reach or to highly organized root-like organs with vessel-like hyphae for efficient water and nutrient transport from distances of decimeters. Cystidia, sterile and variously shaped hyphal ends, possibly appropriate for preventing animal attack, in addition, can cover mantles and rhizomorphs. Although only a limited number of species could be considered, some general conclusions are possible.The genus Tuber forms needle-shaped cystidia and lacks rhizomorphs and clamps. Gomphales ECM are identified by rhizomorphs with ampullate inflations at septa of some hyphae and by oleoacanthocystidia or/and oleoacanthohyphae. Thelephoraceae reveal a great diversity of mantle structures and of extramatrical mycelium, with some additional optional characters, i.e., dark brown color, cystidia, blue granules, amyloid hyphae, or amyloid septa. Bankeraceae are mostly characterized by plectenchymatous mantles with star-like pattern and chlamydospores. Russulaceae possess smooth and hydrophilic ECM. Russula forms plectenchymatous mantles with knob-bearing cystidia, so-called russuloid cystidia, or pseudoparenchymatous mantles without cystidia. Lactarius lacks cystidia and shows laticifers within plectenchymatous or within pseudoparenchymatous mantles. The Boletales families Boletaceae, Gyroporaceae, Melanogastraceae, Paxillaceae, Rhizopogonaceae, Sclerodermataceae, and Suillaceae have the most advanced rhizomorph type, the so-called boletoid rhizomorphs, and reveal generally plectenchymatous mantles, frequently with ring-like patterns. Gomphidiaceae and Albatrellaceae provide cystidia, plectenchymatous mantles, and amyloidy; Gomphidiaceae are generally growing in ECM of Suillaceae and Rhizopogonaceae. Cortinariaceae reveal plectenchymatous mantles and undifferentiated or differentiated rhizomorphs or lack rhizomorphs at all. Cortinarius and Dermocybe are distinct by irregularly shaped, bent to tortuous ECM with many rhizomorphs, some growing over the mycorrhizal tip into the soil. Inocybe lacks rhizomorphs and its emanating hyphae are furnished by many secondary septa and prominent clamps with a hole. Rozites lacks rhizomorphs, too, and reveals a distinctly amyloid gelatinous mantle matrix. Descolea and Descomyces are covered by bolbitioid cystidia. Lastly, the genus Tricholoma forms plectenchymatous mantles and a high diversity of rhizomorphs. Some of the ectomycorrhizal features are used to hypothesize relationships at different taxonomic levels. These conclusions are compared with recently developed molecular hypotheses. Correspondence between the two types of hypotheses are evident, while some conflicts wait for a settlement.
A data set of LSU DNA sequences of mainly European Russula and Lactarius species was subjected to molecular phylogenetic analysis. Species could be allocated to six clades, with an unresolved phylogeny.
One of these clades represents the genus Lactarius. The only analysed species of the section Archaeinae (Russula) was placed basal to both genera. Thus Lactarius appears to be derived from Russula. Russula was divided into four clusters, corresponding to the sections Plorantinae and Nigricantinae, subgenus Heterophyllidia including the section Foetentinae, and a cluster representing the remaining subgenera of the “Genuinae”. Even though the resulting groups can be considered
as valid classificatory groups, species associations resulting from molecular analyses neither support the division of Russula into the subgenus Compacta (including the sections Nigricantinae, Plorantinae, and Archaeinae) and the “Genuinae” (including all remaining taxa), nor do they support previously proposed evolutionary lineages within
the “Genuinae”. Ribosomal ITS DNA sequences of Russula species were analysed to achieve better infrageneric resolution. The results are discussed in relation to current classification
systems and to what is known about the mycorrhizae formed by Russula species. While the systematic value attached to many macroscopic and microscopic sporocarp features was not supported by
sequence data, mycorrhizal anatomy is in good correspondence with many of the results from the phylogenetic analysis.
Species in the large mushroom genus Russula are important ecologically as ectomycorrhizal fungi and economically as comestibles. Most infrageneric classification schemes of this genus have originated in Europe, but because of nomenclatural history and an evolving suite of characters these systems remain largely incongruent. Using ribosomal DNA sequences for 87 species representing all infrageneric taxa described from Europe, the phylogenetic position and relationships among these species were examined. Cladistic analysis of the ITS1, 5.8S, and ITS2 regions showed a cluster of five to six small to large clades basal in the topology and one large apical clade arising from the deeper nodes, none of which has been previously recognized in toto at the subgeneric level. Two of these groups, the Compactae and Pallidosporinae, which have been previously recognized as subsections of section Compacta, did not appear to be closely related. Bootstrap support and Bremer decay values indicated that collapse of the tree into monophyly at the deeper nodes would result in two large groups which are consistent with the classical subgeneric concept of the Eurussulae and a narrowed Compactae. The topology confirmed some previously described infrageneric taxa at the section level including the Tenellae and Heterophyllae and at the subsection level including the Cupreinae, Laricinae, Lilaceinae, Integroidinae, Violaceinae, Sphagnophilinae, Viridantinae, Emeticinae, Subvelatae, Pallidosporinae, and portions of the Polychromae and Sardoninae. The molecular analysis also indicated many interesting new combinations or relationships not previously conceived. Mapping of characters such as spore print colour, taste, and presence of acid resistant incrustations, which have been used to define infrageneric taxa in Russula, onto the phylogeny identified interesting patterns consistent with hypotheses regarding plesiomorphic and apomorphic characters. However, because of potential loss or reversal of character states, this analysis did not support their unequivocal use in infrageneric classification.
Thousands of ectomycorrhizal (ECM) fungal species exist, but estimates of global species richness of ECM fungi differ widely. Many genera have been proposed as being ECM, but in a number of studies evidence for the hypothesized ECM habit is lacking. Progress in estimating ECM species richness is therefore slow. In this paper we have retrieved studies providing evidence for the ECM habit of fungal species and for the identification of the mycobiont(s) in specific ECM associations, using published and web-based mycorrhiza literature. The identification methods considered are morphoanatomical characterization of naturally occurring ECMs, pure culture synthesis, molecular identification, and isotopic evidence. In addition, phylogenetic information is also considered as a relevant criterion to assess ECM habit. Of 343 fungal genera for which an ECM status has been alleged, about two thirds have supportive published evidence or ECM status can be at least hypothesized. For the remaining taxa, currently no indication exists as for their ECM nutritional habit, besides field observations of associations with putative hosts. Our survey clearly indicates that current knowledge of ECM fungal diversity, as supported by experimental evidence, is only partly complete, and that inclusion of many fungal genera in this trophic and ecological category is not verified at this stage. Care must thus be used when compiling lists of ECM and saprotrophic fungi in community-level studies on the basis of published information only. On the basis of our literature search we conservatively estimate ECM species richness around 7750 species. However, on the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate of ECM species richness would likely be between 20000 and 25000.
All available publications providing descriptions of ectomycorrhizas (ECM) were reviewed in order to build a database containing details on fungus forming the ECM, host tree, country where the material for description was collected, and habitat of the ECM. Other secondary data were also recorded. In all 1244 descriptions of ECM published since 1961 in 479 papers were reviewed. The number of different ECM morphotypes described was 814. Most ECM described were collected in Europe and North America. Gymnosperms were the most common tree associates, and boreal and temperate forests the most studied ecosystems. Fungal symbionts were mostly Basidiomycota, epigeous, and with mushroom-like morphology. The paper also addresses the gaps in ECM knowledge that mycorrhizologists should address in future studies.
Status et Noms nouveaux pour les taxa infrageneriques dans le Genere Russula. Doc. mycol
- Gnesi H
GNESI H ( 1987) Status et Noms nouveaux pour les taxa infrageneriques dans le Genere Russula. Doc. mycol. Tome XVlll, Fascicule 69, Novitates 3: 39-40. -SINGER R ( 1986) The Agaricales in Modern Taxonomy. 4. edition, Koeltz Scientific Books, Koenigstein. -YAMADA A (1998) Russula nigricans (Bull.: Fr.)