597:
53:
31:
512:
steppes, as well as varying altitudes from sea level to alpine regions. The highest species diversity are known from subtropical coastal areas with a
Mediterranean or dry climate. The species diversity of the Arthoniales is expected to be higher than currently reported from several areas worldwide like the tropical rain forests.
511:
The
Arthoniales is distributed in most habitats worldwide, as it ranges at latitudes from arctic to tropical regions. They grow on different types of substrates like bark, wood, rocks, bryophytes and living leaves. The order has adapted to live in both humid forests and dry habitats like savannas and
575:
The ascomata are usually apothecial and it produces bitunicate asci. The bitunicate asci are thick-walled, with an outer and inner layer. The outer and inner layer of the ascus wall are called exotunica and endotunica, and they separate during ascospore release. The asci usually contains eight
584:
The majority of the species in
Arthoniales are crustose lichens, with a thallus growing tight to the surface. Fruticose lichens have a bushy thallus and are known in Roccellaceae, where it has evolved and been lost multiple times. The thalli among crustose lichens can either grow within the
567:, chemistry, ascospore color and ascospore septation that can be used as uniting traits for delimiting families and genera in the Arthoniales. The reason is that the Arthoniales is an old order and taxa have evolved in parallel for a long time, which gives a high level of homoplasy.
532:
species are presented as well. The original state in the
Arthoniales is believed to be the lichenized state, and the non-lichenized and parasitic states have evolved independently several times. The lichenized fungi live in symbiosis with a
224:). Lecanographaceae, Roccellographaceae, Opegraphaceae and Roccellaceae are well-supported families within Arthoniales, and they were circumscribed in 2011. Andreiomycetaceae was described as a new family by Hodkinson and Lendemer in 2013.
1112:
Spatafora, Joseph W.; Sung, Gi-Ho; Johnson, Desiree; Hesse, Cedar; O'Rourke, Benjamin; Serdani, Maryna; Spotts, Robert; Lutzoni, François; Hofstetter, Valérie (November 2006). "A five-gene phylogeny of
Pezizomycotina".
1466:
1676:
1833:
1728:
1160:
1585:
1572:
1650:
1689:
1480:
Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil
Tawfeeq; Somayeh, Dolatabadi; Stadler, Marc; Haelewaters, Danny; et al. (2020).
1391:
1066:
Ertz, Damien; Tehler, Anders (2011-07-01). "The phylogeny of
Arthoniales (Pezizomycotina) inferred from nucLSU and RPB2 sequences".
1790:
1838:
978:"The Limitations of Ancestral State Reconstruction and the Evolution of the Ascus in the Lecanorales (Lichenized Ascomycota)"
1694:
1715:
1251:
Aptroot, André; Sparrius, Laurens B. (2008-12-01). "Crustose
Roccellaceae in The Galapagos Islands, With The New Species
169:
lichens are present as well. The order contains around 1500 species, while the largest order with lichenized fungi, the
596:
502:
as the outgroup. The cladogram shows the division of
Arthoniales into seven families, based on Hodkinson et al. 2013.
52:
1828:
1199:
1733:
1818:
39:
601:
949:
Frisch A, Thor G, Ertz D, Grube M (August 2014). "The
Arthonialean challenge: Restructuring Arthoniaceae".
1823:
1525:
529:
1460:
625:
have been tentatively classified in the Arthoniales but have not been placed with any certainty into a
1769:
1795:
1743:
1611:
525:
1642:
777:
585:
substrate, called endophloedal or it can grow at the surface of the substrate, called epiphloedal.
1298:
Grube, Martin (1998). "Classification and Phylogeny in the Arthoniales (Lichenized Ascomycetes)".
788:
563:
It is difficult to mention typical morphological characters like fruit body, exiple, hypothecium,
1448:
1315:
1280:
1183:
1156:
1091:
221:
145:
47:
1427:"Parallel evolution of lichen growth forms in the family Roccellaceae (Arthoniales, Ascomycota)"
1751:
1598:
1387:
1364:
1272:
1175:
1138:
1130:
1083:
1007:
999:
922:
521:
197:
121:
1563:
1756:
1681:
1503:
1493:
1438:
1354:
1346:
1307:
1264:
1233:
1191:
1122:
1075:
1043:
989:
958:
914:
799:
626:
546:
538:
209:
205:
166:
133:
129:
832:
722:
711:
542:
228:
186:
182:
159:
94:
1637:
1359:
1334:
1237:
645:
614:
1590:
1812:
1577:
1443:
1426:
766:
733:
213:
137:
1498:
1481:
1452:
1284:
1095:
1333:
Nelsen MP, Lücking R, Grube M, Mbatchou JS, Muggia L, Plata ER, Lumbsch HT (2009).
1048:
1031:
865:
217:
201:
141:
125:
1335:"Unravelling the phylogenetic relationships of lichenised fungi in Dothideomyceta"
1161:"Lichenicolous species of the Ascomycete genus Arthonia Ach. from Kangaroo Island"
576:
ascospores, while the shape and separation of the ascospores are more variating.
1720:
1663:
1603:
1557:
1268:
1126:
854:
821:
810:
678:
656:
170:
1079:
994:
977:
755:
700:
667:
534:
193:
84:
1548:
1276:
1179:
1134:
1087:
1003:
926:
1782:
1624:
843:
634:
64:
1368:
1350:
1142:
1011:
918:
1195:
30:
1777:
1702:
1542:
564:
551:
1187:
1508:
1319:
744:
689:
606:
1032:"Next-generation sequencing reveals sterile crustose lichen phylogeny"
1668:
190:
162:
74:
1707:
1655:
1519:
1311:
962:
1629:
901:
Sundin R, Tehler A (July 1998). "Phylogenetic Studies of the Genus
1764:
876:
622:
610:
595:
1523:
1616:
976:
Ekman, Stefan; Andersen, Heidi L.; Wedin, Mats (2008-02-01).
1386:(2nd ed.). London: The Natural History Museum.
490:
Figure 1. Cladogram of the Arthoniales, rooted with
1532:
109:Henssen ex D. Hawksw. & O.E. Erikss. (1986)
1382:Smith CW, Aptroot A, Coppins BJ, eds. (2009).
520:The majority of species in Arthoniales have a
944:
942:
940:
938:
936:
8:
1465:: CS1 maint: multiple names: authors list (
1520:
29:
20:
1507:
1497:
1442:
1425:Theler, Irestedt, Anders, Martin (2007).
1358:
1047:
993:
1412:The lichens of Great Britain and Ireland
1384:The Lichens of Great Britain and Ireland
1482:"Outline of Fungi and fungus-like taxa"
1168:Journal of the Adelaide Botanic Gardens
893:
227:The Arthoniales is the sister group to
1458:
617:familial placement in the Arthoniales.
158:is the second largest order of mainly
1834:Taxa named by David Leslie Hawksworth
1405:
1403:
196:. The order includes seven families (
7:
1770:8EECEB10-A206-ABE2-065E-A2D2AC720CAC
1744:cb7732d9-f2d7-4593-98ca-2b32b71098dd
1107:
1105:
1061:
1059:
1025:
1023:
1021:
173:, contains more than 14000 species.
537:that in most cases is a species of
319:
312:
305:
263:
256:
248:
241:
234:
1238:10.1111/j.1756-1051.1983.tb01473.x
14:
1444:10.1111/j.1096-0031.2007.00156.x
1414:. British Lichen society London.
51:
1216:Tehler, A (1983). "The genera
181:The Arthoniales is one of two
1:
639:Aptroot & Sparrius (2008)
804:R.C.Harris & Ladd (2007)
1269:10.1639/0007-2745-111.4.659
1127:10.3852/mycologia.98.6.1018
881:Aptroot & Tibell (1997)
672:Poelt & Döbbeler (1979)
1855:
694:Frisch & G.Thor (2014)
605:, shown here growing on a
1499:10.5943/mycosphere/11/1/8
1080:10.1007/s13225-010-0080-y
995:10.1080/10635150801910451
507:Distribution and habitats
473:
449:
425:
394:
381:
359:
337:
324:
317:
310:
303:
281:
268:
261:
254:
246:
239:
119:
114:
48:Scientific classification
46:
37:
28:
23:
1226:Nordic Journal of Botany
1049:10.5943/mycosphere/4/6/1
40:Cryptothecia rubrocincta
541:, but photobionts from
1839:Taxa described in 1986
1351:10.3114/sim.2009.64.07
1030:Hodkinson, BP (2013).
919:10.1006/lich.1998.0155
618:
609:stump, is one of many
602:Bactrospora brevispora
492:Curvularia brachyspora
428:Curvularia brachyspora
599:
549:and a few species in
1410:Smith, C. W (2009).
1253:Schismatomma spierii
1339:Studies in Mycology
1159:; M Wedin (2015).
982:Systematic Biology
619:
361:Rocellographaceae
270:Andreiomycetaceae
222:Roccellographaceae
146:Roccellographaceae
1829:Ascomycota orders
1806:
1805:
1752:Open Tree of Life
1526:Taxon identifiers
1393:978-0-9540418-8-5
1224:(Roccellaceae)".
907:The Lichenologist
571:Ascomatal anatomy
503:
500:Seynesia erumpens
496:Cudonia circinans
486:
485:
476:Seynesia erumpens
462:
461:
452:Cudonia circinans
438:
437:
414:
413:
405:
404:
383:Lecanographaceae
370:
369:
348:
347:
292:
291:
283:Chrysothricaceae
198:Andreiomycetaceae
152:
151:
122:Andreiomycetaceae
110:
1846:
1799:
1798:
1786:
1785:
1773:
1772:
1760:
1759:
1747:
1746:
1737:
1736:
1724:
1723:
1721:NHMSYS0001474981
1711:
1710:
1698:
1697:
1685:
1684:
1672:
1671:
1659:
1658:
1646:
1645:
1633:
1632:
1620:
1619:
1607:
1606:
1594:
1593:
1581:
1580:
1568:
1567:
1566:
1553:
1552:
1551:
1521:
1514:
1513:
1511:
1501:
1477:
1471:
1470:
1464:
1456:
1446:
1422:
1416:
1415:
1407:
1398:
1397:
1379:
1373:
1372:
1362:
1330:
1324:
1323:
1295:
1289:
1288:
1248:
1242:
1241:
1213:
1207:
1206:
1205:on 22 July 2020.
1204:
1198:. Archived from
1165:
1153:
1147:
1146:
1121:(6): 1018–1028.
1109:
1100:
1099:
1068:Fungal Diversity
1063:
1054:
1053:
1051:
1042:(6): 1028–1039.
1027:
1016:
1015:
997:
973:
967:
966:
946:
931:
930:
913:(4–5): 381–413.
898:
882:
871:
860:
849:
838:
827:
826:Müll.Arg. (1891)
816:
805:
794:
783:
782:Hafellner (1996)
772:
761:
750:
749:Müll.Arg. (1882)
739:
728:
717:
706:
695:
684:
683:A.Massal. (1860)
673:
662:
661:A.Massal. (1852)
651:
650:Hafellner (1998)
640:
547:Chrysotrichaceae
539:Trentepohliaceae
489:
320:
313:
306:
264:
257:
249:
242:
235:
210:Lecanographaceae
206:Chrysotrichaceae
134:Lecanographaceae
130:Chrysotrichaceae
108:
56:
55:
33:
21:
1854:
1853:
1849:
1848:
1847:
1845:
1844:
1843:
1819:Arthoniomycetes
1809:
1808:
1807:
1802:
1794:
1789:
1781:
1776:
1768:
1763:
1755:
1750:
1742:
1740:
1732:
1727:
1719:
1714:
1706:
1701:
1693:
1688:
1680:
1675:
1667:
1662:
1654:
1649:
1641:
1636:
1628:
1623:
1615:
1610:
1602:
1597:
1589:
1584:
1576:
1571:
1562:
1561:
1556:
1547:
1546:
1541:
1528:
1518:
1517:
1479:
1478:
1474:
1457:
1424:
1423:
1419:
1409:
1408:
1401:
1394:
1381:
1380:
1376:
1332:
1331:
1327:
1312:10.2307/3244176
1297:
1296:
1292:
1250:
1249:
1245:
1215:
1214:
1210:
1202:
1163:
1155:
1154:
1150:
1111:
1110:
1103:
1065:
1064:
1057:
1029:
1028:
1019:
975:
974:
970:
963:10.12705/634.20
948:
947:
934:
900:
899:
895:
890:
880:
869:
858:
847:
837:Sparrius (2009)
836:
833:Synarthothelium
825:
814:
803:
792:
781:
770:
759:
748:
737:
726:
723:Helminthocarpon
715:
712:Gossypiothallon
704:
693:
682:
671:
660:
649:
638:
594:
582:
573:
561:
545:are known from
543:Chlorococcaleae
524:lifestyle, but
518:
509:
487:
463:
439:
415:
406:
371:
349:
293:
229:Dothideomycetes
187:Arthoniomycetes
179:
144:
140:
136:
132:
128:
124:
107:
95:Arthoniomycetes
50:
17:
12:
11:
5:
1852:
1850:
1842:
1841:
1836:
1831:
1826:
1821:
1811:
1810:
1804:
1803:
1801:
1800:
1787:
1774:
1761:
1748:
1738:
1725:
1712:
1699:
1686:
1673:
1660:
1647:
1634:
1621:
1608:
1595:
1582:
1569:
1554:
1538:
1536:
1530:
1529:
1524:
1516:
1515:
1472:
1437:(5): 432–454.
1417:
1399:
1392:
1374:
1325:
1306:(3): 377–391.
1300:The Bryologist
1290:
1263:(4): 659–666.
1257:The Bryologist
1243:
1232:(5): 628–631.
1208:
1148:
1101:
1055:
1017:
988:(1): 141–156.
968:
957:(4): 727–744.
932:
892:
891:
889:
886:
885:
884:
873:
862:
851:
840:
829:
818:
807:
796:
793:Trevis. (1857)
785:
774:
763:
752:
741:
730:
719:
716:Aptroot (2014)
708:
705:Aptroot (2014)
697:
686:
675:
664:
653:
646:Arthophacopsis
642:
621:The following
593:
591:incertae sedis
587:
581:
578:
572:
569:
560:
557:
517:
514:
508:
505:
484:
483:
480:
479:
472:
469:
468:
465:
464:
460:
459:
456:
455:
448:
445:
444:
441:
440:
436:
435:
432:
431:
424:
421:
420:
417:
416:
412:
411:
408:
407:
403:
402:
399:
398:
393:
390:
389:
386:
385:
380:
377:
376:
373:
372:
368:
367:
364:
363:
358:
355:
354:
351:
350:
346:
345:
342:
341:
339:Opegraphaceae
336:
333:
332:
329:
328:
323:
318:
316:
311:
309:
304:
302:
299:
298:
295:
294:
290:
289:
286:
285:
280:
277:
276:
273:
272:
267:
262:
260:
255:
253:
247:
245:
240:
238:
233:
178:
177:Classification
175:
150:
149:
117:
116:
112:
111:
102:
98:
97:
92:
88:
87:
82:
78:
77:
72:
68:
67:
62:
58:
57:
44:
43:
35:
34:
26:
25:
16:Order of fungi
15:
13:
10:
9:
6:
4:
3:
2:
1851:
1840:
1837:
1835:
1832:
1830:
1827:
1825:
1824:Lichen orders
1822:
1820:
1817:
1816:
1814:
1797:
1792:
1788:
1784:
1779:
1775:
1771:
1766:
1762:
1758:
1753:
1749:
1745:
1739:
1735:
1730:
1726:
1722:
1717:
1713:
1709:
1704:
1700:
1696:
1691:
1687:
1683:
1678:
1674:
1670:
1665:
1661:
1657:
1652:
1648:
1644:
1639:
1635:
1631:
1626:
1622:
1618:
1613:
1609:
1605:
1600:
1596:
1592:
1587:
1583:
1579:
1574:
1570:
1565:
1559:
1555:
1550:
1544:
1540:
1539:
1537:
1535:
1531:
1527:
1522:
1510:
1505:
1500:
1495:
1492:: 1060–1456.
1491:
1487:
1483:
1476:
1473:
1468:
1462:
1454:
1450:
1445:
1440:
1436:
1432:
1428:
1421:
1418:
1413:
1406:
1404:
1400:
1395:
1389:
1385:
1378:
1375:
1370:
1366:
1361:
1356:
1352:
1348:
1345:: 135–144S4.
1344:
1340:
1336:
1329:
1326:
1321:
1317:
1313:
1309:
1305:
1301:
1294:
1291:
1286:
1282:
1278:
1274:
1270:
1266:
1262:
1258:
1254:
1247:
1244:
1239:
1235:
1231:
1227:
1223:
1219:
1212:
1209:
1201:
1197:
1193:
1189:
1185:
1181:
1177:
1173:
1169:
1162:
1158:
1152:
1149:
1144:
1140:
1136:
1132:
1128:
1124:
1120:
1116:
1108:
1106:
1102:
1097:
1093:
1089:
1085:
1081:
1077:
1073:
1069:
1062:
1060:
1056:
1050:
1045:
1041:
1037:
1033:
1026:
1024:
1022:
1018:
1013:
1009:
1005:
1001:
996:
991:
987:
983:
979:
972:
969:
964:
960:
956:
952:
945:
943:
941:
939:
937:
933:
928:
924:
920:
916:
912:
908:
904:
897:
894:
887:
879:
878:
874:
868:
867:
863:
857:
856:
852:
848:Dennis (1974)
846:
845:
841:
835:
834:
830:
824:
823:
819:
813:
812:
808:
802:
801:
797:
791:
790:
786:
780:
779:
775:
771:Matzer (1996)
769:
768:
767:Paradoxomyces
764:
758:
757:
753:
747:
746:
742:
736:
735:
734:Hormosphaeria
731:
725:
724:
720:
714:
713:
709:
703:
702:
698:
692:
691:
687:
681:
680:
676:
670:
669:
665:
659:
658:
654:
648:
647:
643:
637:
636:
632:
631:
630:
628:
624:
616:
612:
608:
604:
603:
598:
592:
588:
586:
579:
577:
570:
568:
566:
558:
556:
554:
553:
548:
544:
540:
536:
531:
527:
526:lichenicolous
523:
515:
513:
506:
504:
501:
497:
493:
482:
481:
478:
477:
471:
470:
467:
466:
458:
457:
454:
453:
447:
446:
443:
442:
434:
433:
430:
429:
423:
422:
419:
418:
410:
409:
401:
400:
397:
396:Arthoniaceae
392:
391:
388:
387:
384:
379:
378:
375:
374:
366:
365:
362:
357:
356:
353:
352:
344:
343:
340:
335:
334:
331:
330:
327:
322:
321:
315:
314:
308:
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185:of the class
184:
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138:Opegraphaceae
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1200:the original
1171:
1167:
1151:
1118:
1114:
1074:(1): 47–71.
1071:
1067:
1039:
1035:
985:
981:
971:
954:
950:
910:
906:
902:
896:
875:
870:Vain. (1890)
866:Tylophorella
864:
853:
842:
831:
820:
815:Grube (2001)
809:
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789:Phacothecium
787:
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395:
382:
360:
338:
326:Rocellaceae
325:
282:
269:
252:Arthoniales
226:
218:Roccellaceae
202:Arthoniaceae
180:
155:
153:
142:Roccellaceae
126:Arthoniaceae
120:
104:
38:
24:Arthoniales
18:
1664:iNaturalist
1564:Arthoniales
1558:Wikispecies
1534:Arthoniales
1509:10481/61998
1157:G Kantvilas
859:Rehm (1905)
855:Trichophyma
822:Synarthonia
811:Sporostigma
760:Syd. (1935)
738:Lév. (1863)
679:Catarraphia
657:Bactrospora
530:saprophytic
189:within the
171:Lecanorales
156:Arthoniales
105:Arthoniales
1813:Categories
1486:Mycosphere
1431:Cladistics
1222:Roccellina
1036:Mycosphere
888:References
778:Perigrapha
756:Nipholepis
727:Fée (1837)
701:Glyphopsis
668:Bryostigma
559:Characters
535:photobiont
522:lichenized
194:Ascomycota
85:Ascomycota
81:Division:
1783:100386959
1586:AusLichen
1277:0007-2745
1196:Q92902450
1180:0313-4083
1135:0027-5514
1115:Mycologia
1088:1560-2745
1004:1063-5157
927:1096-1135
844:Tarbertia
663:– 35 spp.
635:Angiactis
615:uncertain
167:fruticose
115:Families
71:Kingdom:
65:Eukaryota
1778:Tropicos
1703:MycoBank
1638:Fungorum
1591:30016188
1578:60013164
1573:AusFungi
1543:Wikidata
1453:84702559
1369:20169027
1285:84038943
1192:Wikidata
1188:44464886
1143:17486977
1096:19367958
1012:18300027
903:Arthonia
861:– 2 spp.
839:– 2 spp.
828:– 5 spp.
784:– 5 spp.
751:– 2 spp.
729:– 3 spp.
674:– 3 spp.
641:– 3 spp.
565:hymenium
552:Arthonia
160:crustose
61:Domain:
1549:Q148747
1360:2816970
1320:3244176
1174:: 1–6.
883:– 1 sp.
872:– 1 sp.
850:– 1 sp.
817:– 1 sp.
806:– 1 sp.
800:Phoebus
795:– 1 sp.
773:– 1 sp.
762:– 1 sp.
745:Minksia
740:– 1 sp.
718:– 1 sp.
707:– 1 sp.
696:– 1 sp.
690:Felipes
685:– 1 sp.
652:– 1 sp.
607:cypress
589:Genera
580:Thallus
516:Ecology
163:lichens
101:Order:
91:Class:
1796:147570
1757:124640
1741:NZOR:
1695:610642
1630:1ARTNO
1451:
1390:
1367:
1357:
1318:
1283:
1275:
1218:Dirina
1194:
1186:
1178:
1141:
1133:
1094:
1086:
1010:
1002:
925:
627:family
623:genera
191:phylum
183:orders
165:, but
1791:WoRMS
1765:Plazi
1734:39677
1708:90459
1682:12112
1677:IRMNG
1669:55452
1643:90459
1449:S2CID
1316:JSTOR
1281:S2CID
1203:(PDF)
1184:JSTOR
1164:(PDF)
1092:S2CID
951:Taxon
877:Wegea
75:Fungi
1729:NCBI
1690:ITIS
1656:1273
1651:GBIF
1625:EPPO
1617:5583
1467:link
1388:ISBN
1365:PMID
1273:ISSN
1220:and
1176:ISSN
1139:PMID
1131:ISSN
1084:ISSN
1008:PMID
1000:ISSN
923:ISSN
611:taxa
528:and
498:and
220:and
154:The
1716:NBN
1612:EoL
1599:CoL
1504:hdl
1494:doi
1439:doi
1355:PMC
1347:doi
1308:doi
1304:101
1265:doi
1261:111
1255:".
1234:doi
1123:doi
1076:doi
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613:of
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