{{Short description|Genus of lichens in the family Ramalinaceae}} {{Automatic taxobox | image = Biatora printzenii 123946.jpg | image_caption = Soralia of ''Biatora printzenii'', magnified 30X | taxon = Biatora | authority = Fr. (1817) | type_species = ''Biatora vernalis'' | type_species_authority = (L.) Fr. (1822) | synonyms_ref = <ref name="Species Fungorum synonymy"/> | synonyms = *''Ivanpisutia'' {{small|S.Y.Kondr., Lőkös & Hur (2015)}} }}
'''''Biatora''''' is a genus of lichens in the family Ramalinaceae. Originally circumscribed in 1817,<ref>Fries EM, Sandberg A. (1817). ''Lichenum dianome nova''. Lund.</ref> the genus consists of crustose and squamulose lichens with green algal photobionts, biatorine apothecia, colorless, simple to 3-septate ascospores, and bacilliform pycnospores.<ref name="Printzen 1999"/>
==Description==
''Biatora'' species are crustose lichens with a spreading ({{lichengloss|effuse}}) thallus that may appear thin and somewhat membranous in places. The surface is often cracked ({{lichengloss|rimose}}) and, in species that grow in association with mosses, may be {{lichengloss|granular}} or warted. The thallus is typically creamy white, dull green, glaucous green, or green-grey and lacks a distinct outer protective layer ({{lichengloss|cortex}}). Some species produce soredia, small reproductive {{lichengloss|granules}} that facilitate dispersal. A {{lichengloss|prothallus}}, the initial fungal layer that some lichens form before developing a full thallus, is absent. The photosynthetic partner ({{lichengloss|photobiont}}) is a {{lichengloss|chlorococcoid}} alga, a group characterized by spherical to broadly ellipsoidal cells.<ref name="Cannon et al. 2023"/>
The reproductive structures (apothecia) are {{lichengloss|biatorine}}, meaning they lack a {{lichengloss|thalline margin}} derived from the lichen thallus itself. They are {{lichengloss|sessile}} or closely {{lichengloss|appressed}} to the surface and range from weakly to strongly convex. In some species, the apothecia are initially flat with a shallow margin but later become {{lichengloss|immarginate}} (without a distinct border). Their colour varies widely, including light beige, dark reddish brown, green-grey, bluish green, or khaki. Black apothecia are rare but, when present, have a green or blue tint when wet. Most species lack {{lichengloss|pruina}}, the powdery surface coating found on some lichens.<ref name="Cannon et al. 2023"/>
A well-developed {{lichengloss|true exciple}} (the outer tissue of the apothecium) is present but becomes reflexed over time. It consists of tightly packed, radiating hyphae embedded in a gel matrix that remains stable in potassium hydroxide (K) solution and does not swell. The outer edge may be coated with a thin gel layer. The hymenium, where spores develop, is 30–100 μm tall and typically lacks a distinct {{lichengloss|epithecium}} (uppermost layer), though some species show pale pigmentation at the top. It does not contain granules or oil droplets and reacts with iodine (I+), staining red-brown when young and blue in older herbarium specimens. Below the hymenium, the {{lichengloss|subhymenium}} is distinct and slightly opaque due to the presence of ascogenous hyphae (spore-producing structures). The {{lichengloss|hypothecium}}, a supporting tissue beneath the hymenium, consists of interwoven hyphae embedded in a dense gel matrix.<ref name="Cannon et al. 2023"/>
The paraphyses, sterile filamentous structures within the hymenium, are coherent in KOH, have narrow {{lichengloss|lumina}} (0.5–2.5 μm wide), and are mostly unbranched, though occasional branching or connections (anastomoses) occur. The tips of the paraphyses are slightly swollen, sometimes reaching up to 5 μm in diameter, and rarely bear a distinct cap or hood. The asci, where spores develop, contain eight spores and have a ''Biatora''-type structure. They feature a blue-staining (K/I+) apical dome penetrated from below by a non-staining (K/I–) apical cushion, which is surrounded by a deeply blue-staining zone. The ascus walls themselves do not stain in K/I but are surrounded by an outer layer that reacts red-brown in iodine (I+) and blue in K/I. The ocular chamber, an internal structure within the ascus, is relatively small.<ref name="Cannon et al. 2023"/>
The {{lichengloss|ascospores}} are colourless, with a shape ranging from ellipsoidal to {{lichengloss|filiform}} (thread-like) or {{lichengloss|fusiform}} (spindle-shaped). They may be aseptate (lacking internal divisions) or have between one and seven septa. The spores are smooth and do not possess a distinct outer coating ({{lichengloss|perispore}}). Asexual reproduction occurs via conidia, which are produced in small, flask-shaped reproductive structures called pycnidia. These structures are {{lichengloss|immersed}} within the thallus and have an unpigmented or weakly pigmented wall, similar in colour to the hymenium. The conidia themselves are colourless, aseptate, and {{lichengloss|bacillar}} (rod-shaped).<ref name="Cannon et al. 2023"/>
Chemically, ''Biatora'' lichens can contain a variety of secondary metabolites, including gyrophoric acid and argopsin, and less commonly, other depsides, depsidones, xanthones, or usnic acid. Some species, however, lack detectable secondary metabolites when analysed using thin-layer chromatography.<ref name="Cannon et al. 2023"/>
==Species== *''Biatora alaskana'' {{small|Printzen & Tønsberg (2000)}} *''Biatora alnetorum'' {{small|S.Ekman & Tønsberg (2019)}} *''Biatora amylacea'' {{small|Palice, Malíček, Vondrák & Printzen (2023)}}<ref name="Palice et al. 2023"/> – Europe *''Biatora appalachensis'' {{small|Printzen & Tønsberg (2004)}}<ref name="Printzen & Tønsberg 2004"/> *''Biatora aureolepra'' {{small|T.Sprib. & Tønsberg (2009)}}<ref name="Spribille et al. 2009"/> *''Biatora australis'' {{small|Rodr.Flakus & Printzen (2016)}}<ref name="Printzen et al. 2016"/> *''Biatora bacidioides'' {{small|Printzen & Tønsberg (2003)}}<ref name="Printzen & Tønsberg 2003"/> *''Biatora britannica'' {{small|Printzen, Lumbsch & Orange (2001)}} *''Biatora carneoalbida'' {{small|(Müll. Arg.) Coppins (1992)}} *''Biatora chrysantha'' {{small|(Zahlbr.) Printzen (1994)}} {{Multiple image | image1 = Biatora chrysantha - Flickr - pellaea.jpg | caption1=''Biatora chrysantha'' | image2 =Biatora pontica - Flickr - pellaea.jpg | caption2 =''Biatora pontica'' }} *''Biatora chrysanthoides'' {{small|Printzen & Tønsberg (2003)}}<ref name="Printzen & Tønsberg 2003"/> *''Biatora cuprea'' {{small|(Sommerf.) Fr. (1831)}} *''Biatora cuyabensis'' {{small|(Malme) S.Y.Kondr. (2019)}} *''Biatora efflorescens'' {{small|(Hedl.) Räsänen (1935)}} *''Biatora ementiens'' {{small|(Nyl.) Printzen (2014)}} *''Biatora epirotica'' {{small|Printzen & T.Sprib. (2011)}}<ref name="Spribille et al. 2009"/> *''Biatora epixanthoides'' {{small|(Nyl.) Diederich (1989)}} *''Biatora globulosa'' {{small|(Flörke) Fr. (1845)}} *''Biatora hafellneri'' {{small|Rodr.Flakus & Printzen (2016)}}<ref name="Printzen et al. 2016"/> *''Biatora halei'' {{small|(Tuck.) S.Y.Kondr. (2019)}} *''Biatora hemipolia'' {{small|(Nyl.) S.Ekman & Printzen (2014)}} *''Biatora ivanpisutii'' {{small|S.Y.Kondr., Lőkös & Hur (2016)}} *''Biatora kalbii'' {{small|(Brako) S.Y.Kondr. (2019)}} *''Biatora kodiakensis'' {{small|Printzen & Tønsberg (2004)}}<ref name="Printzen & Tønsberg 2004"/> *''Biatora ligni-mollis'' {{small|T.Sprib. & Printzen (2009)}}<ref name="Spribille et al. 2009"/> *''Biatora loekoesiana'' {{small|S.Y.Kondr. & Hur (2018)}} *''Biatora longispora'' {{small|(Degel.) Lendemer & Printzen (2019)}} *''Biatora meiocarpa'' {{small|(Nyl.) Arnold (1887)}} *''Biatora nobilis'' {{small|Printzen & Tønsberg (2000)}} *''Biatora oligocarpa'' {{small|Printzen & Tønsberg (2004)}}<ref name="Printzen & Tønsberg 2004"/> *''Biatora oxneri'' {{small|(S.Y.Kondr., L.Lőkös & Hur) Printzen & Kistenich (2018)}} *''Biatora pacifica'' {{small|Printzen, Tønsberg & G.Thor (2016)}}<ref name="Printzen et al. 2016"/> *''Biatora pallens'' {{small|(Kullh.) Printzen (2005)}} *''Biatora pausiaca'' {{small|Printzen & Tønsberg (2003)}}<ref name="Printzen & Tønsberg 2003"/> *''Biatora pontica'' {{small|Printzen & Tønsberg (2003)}}<ref name="Printzen & Tønsberg 2003"/> *''Biatora printzenii'' {{small|Tønsberg (2002)}}<ref name="Tønsberg 2002"/> *''Biatora pseudosambuci'' {{small|(S.Y.Kondr., Lőkös & Hur) S.Y.Kondr., Lőkös & Hur (2016)}} *''Biatora pycnidiata'' {{small|Printzen & Tønsberg (2004)}}<ref name="Printzen & Tønsberg 2004"/> *''Biatora radicicola'' {{small|Printzen, Palice & J.P.Halda (2016)}}<ref name="Printzen et al. 2016"/> *''Biatora subduplex'' {{small|(Nyl.) Printzen (1995)}} *''Biatora subhispidula'' {{small|(Nyl.) S.Y.Kondr. (2019)}} *''Biatora terrae-novae'' {{small|Printzen & J.W.McCarthy (2016)}}<ref name="Printzen et al. 2016"/> *''Biatora toensbergii'' {{small|Holien & Printzen (1995)}}<ref name="Printzen 1995"/> *''Biatora vernalis'' {{small|(L.) Fr. (1882)}} *''Biatora veteranorum'' {{small|Coppins & Sérus. (2010)}}<ref name="Sérusiaux et al. 2010"/> *''Biatora vezdana'' {{small|S.Y.Kondr. (2019)}}
The taxon ''Biatora marmorea'', found in Alaska, was proposed as a new species in 2020;<ref name="Spribille et al. 2020"/> however, it is an illegitimate name as it had already been used for a species that is now known as ''Bagliettoa marmorea''.<ref name="Index Fungorum synonymy"/>
==References== {{Reflist|colwidth=30em|refs=
<ref name="Cannon et al. 2023">{{cite book |last1=Cannon |first1=P. |last2=Ekman |first2=S. |last3=Kistenich |first3=S. |last4=LaGreca |first4=S. |last5=Printzen |first5=C. |last6=Timdal |first6=E. |last7=Aptroot |first7=A. |last8=Coppins |first8=B. |last9=Fletcher |first9=A. |last10=Sanderson |first10=N. |last11=Simkin |first11=J. |year=2023 |title=Lecanorales: Ramalinaceae [revision 1], including the genera ''Bacidia'', ''Bacidina'', ''Bellicidia'', ''Biatora'', ''Bibbya'', ''Bilimbia'', ''Cliostomum'', ''Kiliasia'', ''Lecania'', ''Megalaria'', ''Mycobilimbia'', ''Phyllopsora'', ''Ramalina'', ''Scutula'', ''Thalloidima'', ''Toninia'', ''Toniniopsis'' and ''Tylothallia'' |series=Revisions of British and Irish Lichens |volume=35 |pages=23–24 |url=https://britishlichensociety.org.uk/sites/default/files/Ramalinaceae%20rev%201a_0.pdf}} {{OA}}</ref>
<ref name="Index Fungorum synonymy">{{cite web |title=Record Details: ''Biatora marmorea'' T. Sprib., in Spribille, Fryday, Pérez-Ortega, Svensson, Tønsberg, Ekman, Holien, Resl, Schneider, Stabentheiner, Thüs, Vondrák & Sharman, Lichenologist52(2): 89 (2020) |url=http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=830093 |publisher=Index Fungorum |access-date=28 February 2022}}</ref>
<ref name="Palice et al. 2023">{{cite journal |last1=Palice |first1=Zdeněk |last2=Malíček |first2=Jiří |last3=Vondrák |first3=Jan |last4=Printzen |first4=Christian |title=A distinctive new species of ''Biatora'' (Ramalinaceae, Lecanorales) described from native European forests |journal=The Lichenologist |volume=55 |issue=5 |year=2023 |doi=10.1017/S0024282923000464 |pages=325–334}}</ref>
<ref name="Printzen 1995">{{cite book |last=Printzen |first=Christian |year=1995 |title=Die Flechtengattung ''Biatora'' in Europa |trans-title=The lichen genus ''Biatora'' in Europe |series=Bibliotheca Lichenologica |volume=60 |page=137 |publisher=J. Cramer |location=Berlin/Stuttgart |isbn=978-3-443-58039-1}}</ref>
<ref name="Printzen 1999">{{cite journal |last1=Printzen |first1=C. |last2=Tønsberg |first2=T. |year=1999 |title=The lichen genus ''Biatora'' in northwestern North America |journal=The Bryologist |volume=102 |issue=4 |pages=692–713 |doi=10.2307/3244256 |jstor=3244256}}</ref>
<ref name="Printzen & Tønsberg 2003">{{cite journal |last1=Printzen |first1=C. |last2=Tønsberg |first2=T. |year=2003 |title=Four new species and three new apothecial pigments of ''Biatora'' |journal=Bibliotheca Lichenologica |volume=86 |pages=133–145}}</ref>
<ref name="Printzen & Tønsberg 2004">{{cite journal |last1=Printzen |first1=C. |last2=Tønsberg |first2=T. |year=2004 |title=New and interesting ''Biatora''-species, mainly from North America |journal=Symbolae Botanicae Upsalienses |volume=34 |issue=1 |pages=343–352}}</ref>
<ref name="Printzen et al. 2016">{{cite journal |last1=Printzen |first1=Christian |first2=Josef P. |last2=Halda |first3=John W. |last3=McCarthy |first4=Zdeněk |last4=Palice |first5=Pamela |last5=Rodriguez-Flakus |first6=Göran |last6=Thor |first7=Tor |last7=Tønsberg |first8=Jan |last8=Vondrák |year=2016 |title=Five new species of ''Biatora'' from four continents |journal=Herzogia |volume=29 |issue=2 |pages=566–585 |url=http://www.jjh.cz/jphbib/490.pdf}}</ref>
<ref name="Sérusiaux et al. 2010">{{cite journal |last1=Sérusiaux |first1=Emmanuël |last2=Brand |first2=A. Maarten |last3=Motiejunaite |first3=Jurga |last4=Orange |first4=Alan |last5=Coppins |first5=Brian J. |title=''Lecidea doliiformis'' belongs to ''Micarea'', ''Catillaria'' alba to ''Biatora'', and ''Biatora ligni-mollis'' occurs in Western Europe |journal=The Bryologist |volume=113 |issue=2 |year=2010 |doi=10.1639/0007-2745-113.2.333 |doi-access=free |pages=333–344 |url=https://orbi.uliege.be/bitstream/2268/175324/1/2010%20The%20Bryologist%20113%282%29.pdf }}</ref>
<ref name="Species Fungorum synonymy">{{cite web |title=Synonymy: ''Biatora'' Fr., Lichenum Dianome Nova: 7 (1817) |url=http://www.speciesfungorum.org/Names/SynSpecies.asp?RecordID=558 |publisher=Species Fungorum |access-date=10 September 2021}}</ref>
<ref name="Spribille et al. 2009">{{cite journal |last1=Spribille |first1=Toby |last2=Björk |first2=Curtis R.|last3=Ekman |first3=Stefan |last4=Elix |first4=John A. |last5=Goward |first5=Trevor |last6=Printzen |first6=Christian |last7=Tønsberg |first7=Tor |last8=Wheeler |first8=Tim |title=Contributions to an epiphytic lichen flora of northwest North America: I. Eight new species from British Columbia inland rain forests |journal=The Bryologist |volume=112 |issue=1 |year=2009 |pages=109–137 |doi=10.1639/0007-2745-112.1.109}}</ref>
<ref name="Spribille et al. 2020">{{cite journal |last1=Spribille |first1=Toby |last2=Fryday |first2=Alan M. |last3=Pérez-Ortega |first3=Sergio |last4=Svensson |first4=Måns |last5=Tønsberg |first5=Tor |last6=Ekman |first6=Stefan |last7=Holien |first7=Håkon |last8=Resl |first8=Philipp |last9=Schneider |first9=Kevin |last10=Stabentheiner |first10=Edith |last11=Thüs |first11=Holger |last12=Vondrák |first12=Jan |last13=Sharman |first13=Lewis |year=2020 |title=Lichens and associated fungi from Glacier Bay National Park, Alaska |journal=The Lichenologist |volume=52 |issue=2 |pages=61–181 |doi=10.1017/S0024282920000079 |pmid=32788812 |pmc=7398404 |doi-access=free |hdl=10261/232567 |hdl-access=free}}</ref>
<ref name="Tønsberg 2002">{{cite journal |last=Tønsberg |first=Tor |year=2002 |title=Additions to the Lichen Flora of North America XI |journal=The Bryologist |volume=105 |issue=1 |pages=122–125|doi=10.1639/0007-2745(2002)105[0122:ATTLFO]2.0.CO;2}}</ref>
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{{Taxonbar|from=Q4902679}}
Category:Biatora Category:Lecanorales genera Category:Lichen genera Category:Taxa named by Elias Magnus Fries Category:Taxa described in 1817