{{Use dmy dates|date=May 2025}} {{Use Oxford spelling|date=May 2025}} {{Short description|Family of lichen-forming fungi}} {{Automatic taxobox | image = Strigula sp. (poss. Strigula elegans) (5824734683).jpg | image_caption = ''Strigula'' species on plant foliage in Hawaii (possibly ''Strigula elegans'') | taxon = Strigulaceae | authority = [[Alexander Zahlbruckner|Zahlbr.]], 1898 | type_genus = ''[[Strigula]]'' | type_genus_authority = [[Alexander Zahlbruckner|Zahlbr.]] (1898) | subdivisions_rank = | subdivision = | synonyms_ref = <ref name="2024 Outline"/> | synonyms = *Phyllobatheliaceae {{au|[[Friedrich August Georg Bitter|Bitter]] & F.Schill. (1927)}} }}
'''''Strigulaceae''''' is a [[Family (taxonomy)|family]] of mostly [[lichen]]-forming [[fungi]], one of two families in the order [[Strigulales]] (class [[Dothideomycetes]]).<ref name="Wijayawardene et al. 2020"/> A [[molecular phylogenetics|molecular analysis]] of the [[type genus]], ''[[Strigula]]'', has led to a reallocation of the [[foliicolous lichen|foliicolous]] species into six genera that correspond to well-delimited [[clade]]s with diagnostic [[phenotype]] features. These lichens live almost exclusively in [[tropical]] rainforests, where they grow as thin films on the surface of living leaves rather than on bark or rock like many other lichens. The family includes around 140 species distributed across multiple continents, making them one of the most widespread groups of leaf-dwelling organisms. DNA studies have revealed that what scientists once thought was a single large genus actually represents several distinct evolutionary [[lineage (evolution)|lineages]].
==Taxonomy==
The family Strigulaceae was erected by [[Alexander Zahlbruckner]] in 1898 for a mixed assortment of leaf-inhabiting {{lichengloss|pyrenocarpous}} lichens. Mid-twentieth-century revisions restricted the concept to three epiphyllous genera—''[[Strigula]]'', ''[[Phylloporis]]'' and ''[[Raciborskiella]]''—but subsequent authors merged the latter two into an expanded ''Strigula'' and went on to add [[corticolous lichen|corticolous]] (bark-dwelling) and [[saxicolous lichen|saxicolous]] (rock-dwelling) species as well as the genera ''[[Phyllobathelium]]'', ''[[Phyllocratera]]'' and ''[[Flavobathelium]]'' on [[morphology (biology)|morphological]] grounds. Although multigene surveys in the late 2000s showed that these satellite taxa belonged inside Strigulaceae, they also hinted that ''Strigula'' ''sensu lato'' (in the loose sense) was [[paraphyletic]] and that the delimitation of genera within the family required re-examination. Strigulaceae is placed in the order [[Strigulales]] (class [[Dothideomycetes]]) and forms the core lichenised [[lineage (evolution)|lineage]] of that order; its closest relative is the recently described family [[Tenuitholiascaceae]].<ref name="Jiang et al. 2020"/>
A broad multilocus [[phylogeny]] published in 2020 analysed 65 specimens representing 27 foliicolous species of ''Strigula sensu lato'' using various genetic markers. The study recovered six strongly supported [[clade]]s that correlate with clear [[morphology (biology)|morphological]] and ecological characters, and it treated each clade at generic rank. The [[type species]], ''[[Strigula smaragdula|S. smaragdula]]'', anchors ''Strigula'' ''sensu stricto'' (in the strict sense); the novel ''[[Serusiauxiella]]'' groups species with rapidly elongating macroconidial appendages and a ''Trentepohlia''-like photobiont; ''[[Raciborskiella]]'' comprises hypophyllous taxa with large ascospores; ''[[Puiggariella]]'' accommodates species with pale, non-carbonised perithecia and papillose thalli; ''[[Racoplaca]]'' covers the ''S. subtilissima'' complex characterised by olive-brown, finely lobed thalli edged by a black line; and ''[[Phylloporis]]'' was reinstated for supracuticular species allied to a ''[[Phycopeltis]]'' photobiont. Together these six genera constitute a monophyletic foliicolous lineage within Strigulaceae, and the authors transferred or recombined more than a dozen species names to reflect the revised framework.<ref name="Jiang et al. 2020"/>
The same analysis showed that the only sequenced non-foliicolous taxon, ''[[Strigula jamesii]]'', does not belong to this core lineage but instead forms a separate clade together with ''Flavobathelium'' and ''Phyllobathelium ''. This finding implies that additional saxicolous and corticolous species presently kept in ''Strigula'' will require reassignment once their DNA data become available. As of 2020, therefore, Strigulaceae included at least the nine lichenised genera ''Strigula'', ''Serusiauxiella'', ''Raciborskiella'', ''Puiggariella'', ''Racoplaca'', ''Phylloporis'', ''Flavobathelium'', ''Phyllobathelium'' and ''Phyllocratera'', while the status of non-lichenised ''[[Oletheriostrigula]]'' remained doubtful pending further study.<ref name="Jiang et al. 2020"/>
==Description== [[File:Racoplaca melanobapha.png|thumb|right|''[[Racoplaca melanobapha]]'']] Members of Strigulaceae are tiny, leaf-inhabiting lichens that usually form thin, crust-like films rather than the [[foliose lichen|leafy]] or [[fruticose lichen|shrubby]] growth seen in many [[temperate]] lichens. Most species live inside the outer [[plant cuticle|cuticle]] of living leaves (a [[habit (biology)|habit]] called subcuticular), so their thalli show up only as faint green to grey patches or delicate olive-brown lobes on the leaf surface; in a few genera the lichen sits on top of the cuticle (supracuticular) and can be peeled away with a needle . Four easily recognised thallus patterns match the main genetic lineages: thick bright-green crusts (''Strigula'' sensu stricto and ''Serusiauxiella''), paper-thin bluish films on the leaf underside (''Raciborskiella''), folded pale-green mats with tiny white pimples (''Puiggariella''), and olive-brown, lobe-edged patches outlined by a hair-line black border (''Racoplaca''). Because these lichens are so inconspicuous and confined to tropical forests, many casual observers overlook them entirely.<ref name="Jiang et al. 2020"/>
Microscopically the family is unified by a ''Strigula''-type [[ascus]]—a tiny, double-walled ({{lichengloss|bitunicate}}) spore sac whose tip contains a short clear plug (the ocular chamber); this character separates Strigulaceae from its closest relative, [[Tenuitholiascaceae]]. Each ascus releases eight colourless [[ascospore]]s, usually with a single cross-wall ([[septum]]) and measuring 7–70 [[micrometre]]s (μm) depending on the genus; in ''Raciborskiella'' the spores are at the larger end of this range and bear gelatinous "tails" that probably aid in sticking to new leaves . The flask-shaped fruit bodies ([[perithecia]]) that house the asci are minute black dots, their walls commonly {{lichengloss|carbonised}} (hardened and darkened) except in ''Puiggariella''. Asexual spores ([[conidium|macroconidia]]) are rod-shaped and tipped with mucilaginous appendages; in most genera these swell slowly after a day in water, but in ''Serusiauxiella'' they elongate to roughly 70 μm within an hour—a rapid-fire trick thought to help the spores glue themselves to the slippery leaf surface. Chemical screening shows no detectable [[lichen product]]s, so chemical [[spot test (lichen)|spot tests]] and [[thin-layer chromatography]] are uninformative for this family.<ref name="Jiang et al. 2020"/>
==Photobiont==
Strigulaceae lichens partner with just a few groups of [[green alga]]e. The majority harbour ''[[Cephaleuros]]'', a filamentous alga that threads through the leaf cuticle and makes it difficult to lift the lichen away; ''Serusiauxiella'' instead teams up with a ''[[Trentepohlia (alga)|Trentepohlia]]''-like alga, while the supracuticular genus ''Phylloporis'' associates with ''[[Phycopeltis]]'', whose flat, radiating plates let the lichen detach cleanly.<ref name="Jiang et al. 2020"/>
==Genera== [[File:Phylloporis palmae.png|thumb|right|''[[Phylloporis palmae]]'']] *''[[Dichoporis]]'' {{au|Clem. (1909)}}<ref name="Clements 1909"/> – 20 spp. *''[[Flagellostrigula]]'' {{au|Lücking, S.H.Jiang & Sérus. (2020)}}<ref name="Hongsanan et al. 2020"/> – 2 spp. *''[[Flavobathelium]]'' {{au|Lücking, Aptroot & G.Thor (1997)}}<ref name="Lücking et al. 1997"/> – 1 sp. *''[[Oletheriostrigula]]'' {{au|Huhndorf & R.C.Harris (1996)}}<ref name="Huhndorf & Harris 1996"/> – 1 sp. *''[[Phyllobathelium]]'' {{au|(Müll.Arg.) Müll.Arg. (1890)}}<ref name="Müller 1890"/> – 8 spp. *''[[Phyllocharis (lichen)|Phyllocharis]]'' {{au|Fée}} – 1 sp. *''[[Phyllocraterina]]'' {{au|Sérus. & Aptroot (2020)}}<ref name="Hongsanan et al. 2020"/> *''[[Phylloporis]]'' {{au|Clem. (1909)}} – 6 spp. *''[[Puiggariella]]'' {{au|Speg. (1881)}}<ref name="Spegazzini 1881"/> – 3 spp. *''[[Raciborskiella]]'' {{au|Höhn. (1909)}}<ref name="Höhnel 1909"/> – 2 spp. *''[[Racoplaca]]'' {{au|Fée (1825)}}<ref name="Fée 1825"/> – 5 spp. *''[[Serusiauxiella]]'' {{au|S.H.Jiang, Lücking & J.C.Wei (2020)}}<ref name="Jiang et al. 2020"/> – 3 spp. *''[[Strigula]]'' {{au|Fr. (1823)}}<ref name="Fries 1823"/> – ca. 60 spp. *''[[Swinscowia]]'' {{au|S.H.Jiang, Lücking & Sérus. (2020)}}<ref name="Hongsanan et al. 2020"/> – 34 spp.
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|first292=Y.Z. |last293=Lumbsch |first293=H.T. |last294=Luo |first294=L. |last295=Luo |first295=M. |last296=Luo |first296=Z.L. |last297=Ma |first297=J. |last298=Madagammana |first298=A.D. |last299=Madhushan |first299=A. |last300=Madrid |first300=H. |last301=Magurno |first301=F. |last302=Magyar |first302=D. |last303=Mahadevakumar |first303=S. |last304=Malosso |first304=E. |last305=Malysh |first305=J.M. |last306=Mamarabadi |first306=M. |last307=Manawasinghe |first307=I.S. |last308=Manfrino |first308=R.G. |last309=Manimohan |first309=P. |last310=Mao |first310=N. |last311=Mapook |first311=A. |last312=Marchese |first312=P. |last313=Marasinghe |first313=D.S. |last314=Mardones |first314=M. |last315=Marin-Felix |first315=Y. |last316=Masigol |first316=H. |last317=Mehrabi |first317=M. |last318=Mehrabi-Koushki |first318=M. |last319=Meiras-Ottoni |first319=A.de |last320=Melo |first320=R.F.R. |last321=Mendes-Alvarenga |first321=R.L. |last322=Mendieta |first322=S. |last323=Meng |first323=Q.F. 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|last386=Ramírez-Cruz |first386=V. |last387=Rambold |first387=G. |last388=Rathnayaka |first388=A.R. |last389=Raza |first389=M. |last390=Ren |first390=G.C. |last391=Rinaldi |first391=A.C. |last392=Rivas-Ferreiro |first392=M. |last393=Robledo |first393=G.L. |last394=Ronikier |first394=A. |last395=Rossi |first395=W. |last396=Rusevska |first396=K. |last397=Ryberg |first397=M. |last398=Safi |first398=A. |last399=Salimi |first399=F. |last400=Salvador-Montoya |first400=C.A. |last401=Samant |first401=B. |last402=Samaradiwakara |first402=N.P. |last403=Sánchez-Castro |first403=I. |last404=Sandoval-Denis |first404=M. |last405=Santiago |first405=A.L.C.M.A. |last406=Santos |first406=A.C.D.S. |last407=Santos |first407=L.A. |last408=Sarma |first408=V.V. |last409=Sarwar |first409=S. |last410=Savchenko |first410=A. |last411=Savchenko |first411=K. |last412=Saxena |first412=R.K. |last413=Schoutteten |first413=N. |last414=Selbmann |first414=L. |last415=Ševčíková |first415=H. |last416=Sharma |first416=A. |last417=Shen |first417=H.W. |last418=Shen |first418=Y.M. |last419=Shu |first419=Y.X. |last420=Silva |first420=H.F. |last421=Silva-Filho |first421=A.G.S. |last422=Silva |first422=V.S.H. |last423=Simmons |first423=D.R. |last424=Singh |first424=R. |last425=Sir |first425=E.B. |last426=Sohrabi |first426=M. |last427=Souza |first427=F.A. |last428=Souza-Motta |first428=C.M. |last429=Sri-indrasutdhi |first429=V. |last430=Sruthi |first430=O.P. |last431=Stadler |first431=M. |last432=Stemler |first432=J. |last433=Stephenson |first433=S.L. |last434=Stoyneva-Gaertner |first434=M.P. |last435=Strassert |first435=J.F.H. |last436=Stryjak-Bogacka |first436=M. |last437=Su |first437=H. |last438=Sun |first438=Y.R. |last439=Svantesson |first439=S. |last440=Sysouphanthong |first440=P. |last441=Takamatsu |first441=S. |last442=Tan |first442=T.H. |last443=Tanaka |first443=K. |last444=Tang |first444=C. |last445=Tang |first445=X. |last446=Taylor |first446=J.E. |last447=Taylor |first447=P.W.J. |last448=Tennakoon |first448=D.S. |last449=Thakshila |first449=S.A.D. |last450=Thambugala |first450=K.M. |last451=Thamodini |first451=G.K. |last452=Thilanga |first452=D. |last453=Thines |first453=M. |last454=Tiago |first454=P.V. |last455=Tian |first455=X.G. |last456=Tian |first456=W.H. |last457=Tibpromma |first457=S. |last458=Tkalčec |first458=Z. |last459=Tokarev |first459=Y.S. |last460=Tomšovský |first460=M. |last461=Torruella |first461=G. |last462=Tsurykau |first462=A. |last463=Udayanga |first463=D. |last464=Ulukapı |first464=M. |last465=Untereiner |first465=W.A. |last466=Usman |first466=M. |last467=Uzunov |first467=B.A. |last468=Vadthanarat |first468=S. |last469=Valenzuela |first469=R. |last470=Van den Wyngaert |first470=S. |last471=Van Vooren |first471=N. |last472=Velez |first472=P. |last473=Verma |first473=R.K. |last474=Vieira |first474=L.C. |last475=Vieira |first475=W.A.S. |last476=Vinzelj |first476=J.M. |last477=Tang |first477=A.M.C. |last478=Walker |first478=A. |last479=Walker |first479=A.K. |last480=Wang |first480=Q.M. |last481=Wang |first481=Y. |last482=Wang |first482=X.Y. |last483=Wang |first483=Z.Y. |last484=Wannathes |first484=N. |last485=Wartchow |first485=F. |last486=Weerakoon |first486=G. |last487=Wei |first487=D.P. |last488=Wei |first488=X. |last489=White |first489=J.F. |last490=Wijesundara |first490=D.S.A. |last491=Wisitrassameewong |first491=K. |last492=Worobiec |first492=G. |last493=Wu |first493=H.X. |last494=Wu |first494=N. |last495=Xiong |first495=Y.R. |last496=Xu |first496=B. |last497=Xu |first497=J.P. |last498=Xu |first498=R. |last499=Xu |first499=R.F. |last500=Xu |first500=R.J. |last501=Yadav |first501=S. |last502=Yakovchenko |first502=L.S. |last503=Yang |first503=H.D. |last504=Yang |first504=X. |last505=Yang |first505=Y.H. |last506=Yang |first506=Y. |last507=Yang |first507=Y.Y. |last508=Yoshioka |first508=R. |last509=Youssef |first509=N.H. |last510=Yu |first510=F.M. |last511=Yu |first511=Z.F. |last512=Yuan |first512=L.L. |last513=Yuan |first513=Q. |last514=Zabin |first514=D.A. |last515=Zamora |first515=J.C. |last516=Zapata |first516=C.V. |last517=Zare |first517=R. |last518=Zeng |first518=M. |last519=Zeng |first519=X.Y. |last520=Zhang |first520=J.F. |last521=Zhang |first521=J.Y. |last522=Zhang |first522=S. |last523=Zhang |first523=X.C. |last524=Zhao |first524=C.L. |last525=Zhao |first525=H. |last526=Zhao |first526=Q. |last527=Zhao |first527=H. |last528=Zhao |first528=H.J. |last529=Zhou |first529=H.M. |last530=Zhu |first530=X.Y. |last531=Zmitrovich |first531=I.V. |last532=Zucconi |first532=L. |last533=Zvyagina |first533=E. |title=The 2024 Outline of Fungi and fungus-like taxa |journal=Mycosphere |volume=15 |issue=1 |pages=5146–6239 [5225] |year=2024 |doi=10.5943/mycosphere/15/1/25 |url=https://www.researchgate.net/publication/385939154 |hdl=11584/429245 |hdl-access=free}}</ref>
}}
{{Taxonbar |from1=Q10681823 |from2=Q7188649}}
[[Category:Strigulales]] [[Category:Dothideomycetes families]] [[Category:Lichen families]] [[Category:Taxa described in 1898]] [[Category:Taxa named by Alexander Zahlbruckner]]