identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03F587D3FFDAB947B5974D8CFEE9FBB9.text	03F587D3FFDAB947B5974D8CFEE9FBB9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aneuraceae	<div><p>a) Aneuraceae</p> <p>The Aneuraceae are the only thalloid liverworts found to date to contain basidiomycetous endophytes clearly identified as such by the presence of dolipore septa (Fig. 4D, F) (Bidartondo et al. 2003; Duckett &amp; Ligrone 2008a, b; Ligrone et al. 1993; Nebel et al. 2004; Preussing et al. 2010). The endophyte-host relationships recall those in orchids with the formation of hyphal coils in the colonised cells. These are then digested by the host to be followed by one or more reinfection cycles. As in the Marchantiophyta, the number of cell layers colonized by the fungus varies among taxa: from two to five in Chilean and New Zealand species of Aneura to approximately half the thallus in northern hemisphere A. pinguis (L.) Dumort. to virtually every cell in Cryptothallus.</p> <p>Whilst this kind of association is also found in Verdoornia R.M.Schust. (Duckett &amp; Ligrone 2008a) and in nearly all collections of Aneura Dumort. examined to date with the exception of A. pellioides (Horik.) and A. pinguis from California (Bidartondo &amp; Duckett 2010), the sister genus Riccardia Gray is almost fungus-free. In three Riccardia taxa, all New Zealand endemics, where an endophyte is present the associations appear distinctively different from those in Aneura. Two of these we describe here for the first time (Fig. 4A–D), but a third species R. intercellula A.E.Brown is known from but a single collection and has never been refound in nature (Brown &amp; Braggins 1989). In R. pennata A.E.Brown, prominent basidiomycetous hyphal coils, which show no signs of digestion by the host, fill the ventral epidermal cells (Fig. 4C). Even more remarkable is R. cochleata (Hook.f. &amp; Taylor) Kuntze. This species produces achlorophyllous radially symmetrical ‘rhizomes’ (Brown &amp; Braggins 1989) recalling the ‘roots’ of Haplomitrium but lacking the mucilage investiture of the latter (Fig. 4A). While the fleshy surface-growing thalli are fungus-free, the first investigation of the anatomy of the rhizomes revealed that the epidermal cells are packed with hyphal coils (Fig. 4B) that undergo digestion cycles.</p> <p>The fact that the Aneuraceae forms the crown group of an otherwise fungus-free lineage comprising Pleurozia Dumort., Phyllothallia E.A.Hodgs. and the Metzgeriaceae and the discovery from the initial molecular studies that the fungus from both Aneura and Cryptothallus belongs to the derived genus Tulasnella J.Schröt. (Bidartondo et al. 2003; Kottke et al. 2003) indicate that the aneuracean associations are of recent origin. Much more extensive sequencing investigations (Bidartondo &amp; Duckett 2010; Preussing et al. 2010), embracing over one hundred collections of the Aneuraceae, revealed that whilst in the vast majority the fungus is Tulasnella, two gatherings (from Chile and Switzerland) contained Sebacina Tul., the basidiomycete genus characteristic of leafy liverworts (Kottke et al. 2003). The molecular diversity discovered within the Tulasnella symbionts (Bidartondo &amp; Duckett 2010; Preussing et al. 2010), plus the rare occurrence of sebacinoids, in seemingly closely allied aneuracean taxa was at first sight most surprising. However recent molecular studies on the liverworts (Wackowiak et al. 2007; Baczkiewicz et al. 2008; Wickett &amp; Goffinet 2008) have revealed that conservative thallus morphology within the Aneuraceae conceals remarkable genetic diversity. In fact, the diversity in the endophytes closely mirrors that in the hosts and includes noteworthy congruence between the fungal and liverwort phylogenies. Indeed the nesting of Cryptothallus within a single clade of Aneura (Wickett &amp; Goffinet 2008), leading to the subsuming of the former into this genus is reinforced by the fungal data. Systematic considerations aside, with wholesale fungal digestion a standard feature of the aneuracean associations compared to those with basidiomycetes in leafy liverworts, it is perhaps not surprising that in this group we find the only achlorophyllose liverworts. Although on the one hand the fungus in the achlorophyllose rhizomes in Riccardia cochleata is most likely an independent acquisition, on the other the rhizomes perhaps provide an example as to how mycoheterotrophy may have evolved in the Aneuraceae.</p> </div>	http://treatment.plazi.org/id/03F587D3FFDAB947B5974D8CFEE9FBB9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pressel, Silvia;Bidartondo, Martin I.;Ligrone, Roberto;Duckett, Jeffrey G.	Pressel, Silvia, Bidartondo, Martin I., Ligrone, Roberto, Duckett, Jeffrey G. (2010): Fungal symbioses in bryophytes: New insights in the Twenty First Century. Phytotaxa 9: 238-253, DOI: 10.11646/phytotaxa.9.1.13
03F587D3FFDCB94DB5974E12FEFEFD69.text	03F587D3FFDCB94DB5974E12FEFEFD69.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Haplomitrium	<div><p>Haplomitrium and Treubia</p> <p>Until the recent and detailed electron microscope studies by Carafa et al. (2003) and Duckett et al. (2006a) our knowledge of the endophytes in these two genera was limited to early light microscope investigations (Goebel 1891; Grun 1914; Lilienfeld 1911). These suggested that fungal associations in Haplomitrium Dedecek and Treubia might be different from those found elsewhere in land plants and so this proved to be.</p> <p>Electron microscopy confirmed the presence of both extra and intracellullar hyphae in Treubia (Fig. 1). The former forms pseudoparenchyma-like structures within the mucilage-filled intercellular spaces in the thalli (Fig. 1D, E). In the older thalli these hyphae produce spores with highly distinctive multilamellate walls (Fig. 1F). The intracellular fungal zone lies in the lower part of the thalli and comprises coils of fine hyphae some of which terminate in short-lived fungal lumps (Fig. 1B, C). Exactly the same kind of intracellular cytology is found in the epidermal and sometimes the subepidermal cells in the subterranean mucilageinvested ‘roots’ of Haplomitrium. Spores with multilamellate walls like those in Treubia can also be seen in this mucilage. Unlike all other bryophyte-fungus associations where hyphal entry is via the rhizoids, in Treubia the rhizoids remain uninfected and entry is via mucilage-filled spaces between the epidermal cells. In Haplomitrium, which lacks rhizoids, hyphae penetrate directly through the epidermal cell walls.</p> <p>The presence of an intercellular phase for the infections in Treubia draws parallels with the same in Lycopodium L. (Duckett &amp; Ligrone 1992) and even more strikingly with the Devonian plant Nothia aphylla Lyon ex Høeg. (Krings et al. 2007a, b). Given this distinctive ultrastructure and the fact that Treubia and Haplomitrium are now considered to have had a very long evolutionary history in the land plant tree of life, it is extremely disappointing that the only published molecular study to date (Ligrone et al. 2007) revealed the presence of a Glomus in Haplomitrium chilensis R.M.Schust. closely allied to the Glomus Tul. &amp; C.Tul. Group A endophytes found in Conocephalum Hill, Fossombronia Hazsl. and Pellia Raddi. However, a much more extensive sequencing programme of the Treubia and Haplomitrium endophytes now in progress (Bidartondo &amp; Duckett 2010, unpublished data) is revealing that these two genera, plus several primitive taxa in the thalloid liverworts, lack Glomus but contain zygomycetous endophytes, a group of fungi recently resolved as ancestral to the glomeromycetes (James et al. 2006). Thus, we now have the first glimpses of what may well turn out to be a hitherto unsuspected spectrum of novel fungal symbioses very early in the ancestry of land plants.</p> </div>	http://treatment.plazi.org/id/03F587D3FFDCB94DB5974E12FEFEFD69	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pressel, Silvia;Bidartondo, Martin I.;Ligrone, Roberto;Duckett, Jeffrey G.	Pressel, Silvia, Bidartondo, Martin I., Ligrone, Roberto, Duckett, Jeffrey G. (2010): Fungal symbioses in bryophytes: New insights in the Twenty First Century. Phytotaxa 9: 238-253, DOI: 10.11646/phytotaxa.9.1.13
03F587D3FFD4B945B5974CB4FD26FCA8.text	03F587D3FFD4B945B5974CB4FD26FCA8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Jungermanniales	<div><p>b) Jungermanniales</p> <p>Electron microscopy and molecular studies (Read et al. 2000; Kottke et al. 2003; Kottke &amp; Nebel 2005; Nebel et al. 2004; Duckett et al. 2006b; Bidartondo &amp; Duckett 2010; Newsham &amp; Bridge 2010) have shown that eight or nine leafy liverwort genera; Barbilophozia Loeske, Diplophyllum (Dumort.) Dumort., Lophozia (Dumort.) Dumort., Scapania (Dumort.) Dumort. and Tritomaria Schiffn. ex Loeske in the Scapaniaceae, Nardia Gray in the Jungermanniaceae, Saccogyna Dumort. in the Geocalycaceae and Southbya Spruce, and almost certainly Gongylanthus Nees, in the Arnelliaceae, consistently and predominantly associate with members of the Sebacina vermifera Oberw. species complex, in striking contrast to Tulasnella as the predominant endophyte in the Aneuraceae. As in most liverwort-fungus associations, these endophytes enter the hosts via the rhizoids. However colonization by these basidiomycetes causes the rhizoids to become highly branched (Fig. 5A) rather than eliciting the terminal swellings typical of ascomycetous infections. Within the liverwort stems two kinds of association have been described. In the Arnelliaceae the endophytes colonize a cluster of cells in the centre of the stems (Fig. 5D) and there are no visible features in the host walls indicating why the fungus is so restricted. In the Scapaniaceae, Jungermanniaceae and Geocalycaceae, the central region of the stems contains a mosaic of infected and fungus-free cells. Growth of hyphae into the latter is prevented by overgrowths of host wall material (Fig. 5B). In both kinds of infection there is no evidence of fungus digestion by the host and healthy hyphae persist long after the death of the host cells in older stems (Fig. 5F). In fact, host cell death occurs very rapidly in the mosaic infections (Duckett et al. 2006b). One previously overlooked effect of fungal colonization in the Arnelliaceae is that oil bodies in infected cells rapidly break down (Fig. 5E) as hyphae proliferate suggesting that their contents may be a nutrient source for the endophytes.</p> <p>In striking contrast to the vast host range of the rhizoidal ascomycetes, cross-infection experiments on the leafy liverwort basidiomycetes strongly suggested these to be highly host specific (Duckett et al. 2006b). This situation has now been confirmed by a molecular study (Bidartondo &amp; Duckett 2010) revealing a high level of fungal specificity to the extent that where multiple liverworts co-occur they almost never share the same endophyte. This remarkable symbiotic conservatism differs fundamentally from the generalist pattern of mycorrhizas in seed plants that repeatedly evolved to form ectomycorrhizas simultaneously with a range of distantly related basidiomycete genera.</p> <p>The phylogenetic position of these basidiomycete-associated leafy liverworts (Davis 2004; Forrest et al. 2006; Heinrichs et al. 2005, 2007; Hentschel et al. 2006) and the fact that they are far more restricted than those with ascomycetes suggest that the basidiomycetes were a secondary acquisition following the loss of ascomycetes. However, the position of the Sebacinales in fungal phylogenies suggests that they may be considerably more ancient than the Tulasnella associations in the Aneuraceae (Kottke et al. 2003; Kottke &amp; Nebel 2005). The discovery that basidiomycetes are widespread in Scapania, Diplophyllum, Barbilophozia and Lophozia is congruent with their close evolutionary relationship (Schill et al. 2004; Yatsentyuk et al. 2004; He-Nygrén et al. 2006) whilst the placement of basidiomycete-associated liverworts (Lophozia sudetica (Nees &amp; Huebener) Grolle, Barbilophozia barbata (Schmieder ex Schreb.) Loeske, B. lycopodioides (Wallr.) Loeske and B. hatchery (A.Evans) Loeske) sister to other members of the Lophoziaceae (de Roo et al. 2007) suggests ancestral presence of fungi in this family.</p> </div>	http://treatment.plazi.org/id/03F587D3FFD4B945B5974CB4FD26FCA8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Pressel, Silvia;Bidartondo, Martin I.;Ligrone, Roberto;Duckett, Jeffrey G.	Pressel, Silvia, Bidartondo, Martin I., Ligrone, Roberto, Duckett, Jeffrey G. (2010): Fungal symbioses in bryophytes: New insights in the Twenty First Century. Phytotaxa 9: 238-253, DOI: 10.11646/phytotaxa.9.1.13
