497:
unlikely that algae would be preserved as coalified impressions. However, Edwards does note that the surface patterning could have been produced in a similar fashion to surface layers in green algae – that is, by the ends of tightly packed filaments causing indentation on the surface layer. (Just because they were formed in the same way doesn't mean they were formed by green algae, though.) The most likely affinity, for the aperture-bearing cuticles, is with the coralline red algae; the absence of biomineralization suggests a stem-group level affiliation. Whereas Smith & Butterfield contend that this material is separate from the charcoalified lichen-like material, Edwards insists that all the material comes from the same lichen-like organism.
458:
470:
122:
144:
104:
482:
337:, covered by a cuticle which preserved impressions of the underlying cells. He had found abundant disaggregated remains of all three features, none of which were connected to another, leaving his reconstruction of the phytodebris as parts of a single organism highly conjectural. Even so, it was picked up by Jonker (1973), who proposed that
430:
was originally recovered from the Red
Downtonian near the base of the Old Red Sandstone, between the clearly marine Ludlow deposits beneath it, and the unambiguously terrestrial Lower Old Red Sandstone. Further samples came from elsewhere in the Lower Old Red Sandstone (St Maughams Group, lowermost
364:
consisted of two layers of cuticle, although fossils giving this impression may in fact represent two layers which happened to overlap one another and become stuck. It is not readily established what the cuticle represents. Lang (1945) had it as an epidermal layer, similar to the waxy cuticle of
496:
closely resembles certain lichens, suggesting a lichenous habit – although as the lichens evolved at least six times independently, this does not fully establish the genetic affinity. The biochemistry of the organism is not inconsistent with an algal affinity, but
Edwards (1982) considers it
356:
should only apply to the tubes, until
Strother (1993) found more complete specimens, with tubes attached to the cuticle. He attempted to unite and formalise the genus, and extended it to include banded tubes, which are instead referred to as nematoclasts (Graham & Gray 2001).
418:
individuals, it is unlikely that they were involved in gas exchange. One suggestion based on fossil evidence for their purpose is the release of spores, although this may be the result of spores being trapped between two layers of cuticle.
348:
Further work failed to draw together all aspects of the organism: Edwards (1982) and
Edwards and Rose (1984) both provided thorough descriptions of the cuticular aspects of the plants, while Pratt
1168:
Burgess, N. D.; Edwards, D. (1991). "Classification of uppermost
Ordovician to Lower Devonian tubular and filamentous macerals from the Anglo-Welsh Basin".
1251:
McGregor, DC; Narbonne, GM (1978). "Upper
Silurian trilete spores and other microfossils from the Read Bay Formation, Cornwallis Island, Canadian Arctic".
1225:
Filipiak, P.; Zatoń, M. (2010). "Plant and animal cuticle remains from the Lower
Devonian of southern Poland and their palaeoenvironmental significance".
885:
Pratt, L.; Phillips, T. L.; Dennison, J. M. (1978). "Evidence of non-vascular land plants from the early
Silurian (Llandoverian) of Virginia, U.S.A".
1103:
1330:
1016:
1317:
1034:
352:(1978) and Niklas and Smocovitis (1983) focused on the anatomy of the tubes. Indeed, some workers suggested that the name
624:
1076:
Edwards, D.; Kerp, H.; Hass, H. (1998). "Stomata in early land plants: an anatomical and ecophysiological approach".
410:
Further work by
Edwards and Rose has identified oval-shaped growths in places on the cuticles of a limited number of
920:
Gensel, P. G.; Johnson, N. G.; Strother, P. K. (1990). "Early Land Plant Debris (Hooker's "Waifs and Strays"?)".
1018:
The Origin, Morphology, and
Ecophysiology of Early Embryophytes: Neontological and Paleontological Perspectives
324:
376:
The genus was later formalised by Strother, who discovered better preserved and more complete specimens in
143:
1062:
1363:
1358:
820:
Edwards, D. (1982). "Fragmentary non-vascular plant microfossils from the late Silurian of Wales".
538:, a similarly enigmatic organism with a similar cuticular patterning. Some material described as
414:
fragments, which develop into holes – whose purpose is unknown. Since they are not found in all
1285:
469:
1335:
457:
1097:
1260:
1126:
981:
929:
894:
721:
681:
582:
431:
Devonian), where they co-occur with plant spores. Additional, older material occurs with
8:
1264:
1130:
985:
933:
898:
725:
685:
586:
1181:
1150:
1142:
1050:
997:
945:
871:
833:
776:
737:
598:
138:
121:
1238:
1117:
Niklas, K. J. (1976). "Chemical Examinations of Some Non-Vascular Paleozoic Plants".
1030:
906:
712:
Lang, W. H. (1937). "On the Plant-Remains from the Downtonian of England and Wales".
694:
665:
1195:
1154:
1089:
1268:
1234:
1177:
1134:
1085:
1022:
989:
937:
902:
867:
829:
772:
729:
689:
643:
590:
327:
1308:
384:
is widespread from the late Silurian, but similar cuticle is reported from the
993:
594:
373:
comprised stringy tubes, with the cellular patterning produced by their ends.
1352:
625:"Contributions to the diversity in cryptogamic covers in the mid-Palaeozoic:
534:
61:
733:
526:
445:
377:
237:
103:
1026:
481:
380:, America – which appear to show tubes connected to the rims of cuticle.
194:
181:
168:
36:
369:
layer. Alternatively, Edwards (1982) proposed that the inner tissue of
1322:
1146:
1001:
949:
602:
433:
389:
366:
334:
330:
81:
46:
714:
Philosophical Transactions of the Royal Society B: Biological Sciences
647:
741:
439:
404:
394:
305:
86:
1279:
1272:
1138:
941:
1302:
76:
71:
56:
51:
41:
385:
91:
66:
323:
was first described by Lang in 1937, who envisioned it being an
309:
301:
297:
155:
972:
from the Silurian Bloomsburg Formation of Pennsylvania".
790:
Jonker, FP (1979). "Prototaxites in the lower Devonian".
465:
fringed with filaments and sporting further development
449:
in the Ludlow (upper Ludfordian) Whitcliffe formation.
761:
Prototaxites, Nematothallus, Parka, Foerstia, Orvillea
919:
884:
1014:
569:
Strother, P. K. (1993). "Clarification of the genus
670:: Coralline red algae from the Silurian of Gotland"
622:
1350:
1250:
663:
1075:
963:
961:
959:
1167:
564:
562:
560:
477:aperture fringed with multi-cellular filaments
1224:
849:
847:
845:
843:
815:
813:
811:
809:
807:
805:
618:
616:
614:
612:
399:). It is, however, difficult to distinguish
304:. Some of its constituents likely represent
1218:
1102:: CS1 maint: DOI inactive as of July 2024 (
956:
512:were both collapsed into a new form-species
16:A form genus comprising cuticle-like fossils
1161:
1069:
854:Edwards, D.; Rose, V. (1984). "Cuticles of
853:
659:
657:
623:Edwards, D.; Axe, L.; Honegger, R. (2013).
557:
1110:
1008:
913:
840:
802:
707:
705:
609:
120:
102:
783:
693:
664:Smith, M. R.; Butterfield, N. J. (2013).
1206:from the Lower Devonian of Poland].
1170:Botanical Journal of the Linnean Society
968:Strother, P. K. (1988). "New Species of
967:
878:
860:Botanical Journal of the Linnean Society
822:Botanical Journal of the Linnean Society
765:Botanical Journal of the Linnean Society
654:
636:Botanical Journal of the Linnean Society
568:
480:
468:
456:
819:
748:
702:
1351:
1116:
789:
345:, which he interpreted as a red alga.
315:
1284:
1283:
887:Review of Palaeobotany and Palynology
754:
711:
13:
1253:Canadian Journal of Earth Sciences
1182:10.1111/j.1095-8339.1991.tb02282.x
872:10.1111/j.1095-8339.1984.tb01563.x
834:10.1111/j.1095-8339.1982.tb00536.x
777:10.1111/j.1095-8339.1945.tb01351.x
485:Spores surrounding an aperture in
133:showing pseudocellular patterning
14:
1375:
759:and some anomalous early plants (
530:, a closely related sister taxon.
1239:10.1111/j.1502-3931.2010.00251.x
695:10.1111/j.1475-4983.2012.01203.x
142:
34:
1188:
1015:Graham, L.E.; Gray, J. (2001).
115:showing reproductive apertures
1078:Journal of Experimental Botany
1:
1090:10.1093/jexbot/49.suppl_1.255
1021:. Columbia University Press.
792:Palaeontographica Abteilung B
550:
245:Nematothallus pseudovasculosa
907:10.1016/0034-6667(78)90034-9
7:
519:
500:
452:
10:
1380:
492:Some material assigned to
422:
333:with tubular features and
308:, whereas others resemble
1292:
994:10.1017/S0022336000030237
595:10.1017/S0022336000025476
365:plants today, covering a
259:
254:
243:
236:
139:Scientific classification
137:
128:
119:
110:
101:
23:
1200:z dolnego dewonu Polski"
300:comprising cuticle-like
974:Journal of Paleontology
575:Journal of Paleontology
397: million years ago
1092:(inactive 2024-07-02).
734:10.1098/rstb.1937.0004
489:
478:
466:
341:represented leaves of
1027:10.7312/gens11160-009
858:: a further enigma".
755:Lang, W. H. (1945). "
484:
472:
460:
403:cuticle from that of
282:Edwards et al. (2013)
32:Ludfordian–Downtonian
360:It is possible that
1265:1978CaJES..15.1292M
1208:Acta Palaeobotanica
1131:1976Britt..28..113N
986:1988JPal...62..967S
934:1990Palai...5..520G
899:1978RPaPa..25..121P
726:1937RSPTB.227..245L
686:2013Palgy..56..345S
587:1993JPal...67.1090S
316:History of research
263:N. pseudo-vasculosa
514:Laevitubulus laxus
490:
479:
467:
1346:
1345:
1286:Taxon identifiers
1214:(1): 13–20. 1978.
1036:978-0-231-11161-4
648:10.1111/boj.12119
289:
288:
283:
275:
267:
232:
213:
1371:
1339:
1338:
1326:
1325:
1313:
1312:
1311:
1281:
1280:
1276:
1259:(8): 1292–1303.
1243:
1242:
1222:
1216:
1215:
1192:
1186:
1185:
1165:
1159:
1158:
1114:
1108:
1107:
1101:
1093:
1073:
1067:
1066:
1060:
1056:
1054:
1046:
1044:
1043:
1012:
1006:
1005:
965:
954:
953:
917:
911:
910:
882:
876:
875:
851:
838:
837:
817:
800:
799:
787:
781:
780:
771:(347): 535–552.
752:
746:
745:
720:(544): 245–291.
709:
700:
699:
697:
661:
652:
651:
633:
620:
607:
606:
581:(6): 1090–1094.
566:
487:Nematothallopsis
475:Nematothallopsis
463:Nematothallopsis
398:
281:
273:
265:
230:
223:
211:
206:
193:
180:
159:
147:
146:
131:Nematothallopsis
124:
113:Nematothallopsis
106:
96:
33:
29:Temporal range:
21:
20:
1379:
1378:
1374:
1373:
1372:
1370:
1369:
1368:
1364:Devonian plants
1359:Silurian plants
1349:
1348:
1347:
1342:
1334:
1329:
1321:
1316:
1307:
1306:
1301:
1288:
1273:10.1139/e78-136
1247:
1246:
1223:
1219:
1194:
1193:
1189:
1166:
1162:
1139:10.2307/2805564
1115:
1111:
1095:
1094:
1074:
1070:
1058:
1057:
1048:
1047:
1041:
1039:
1037:
1013:
1009:
966:
957:
942:10.2307/3514860
918:
914:
883:
879:
852:
841:
818:
803:
788:
784:
753:
749:
710:
703:
666:"A new view on
662:
655:
631:
621:
610:
567:
558:
553:
542:may in fact be
522:
503:
455:
425:
393:
318:
250:
247:
229:
221:
212:Strother (1993)
210:
208:Nematothalaceae
204:
191:
178:
157:
141:
97:
95:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
39:
31:
30:
27:
17:
12:
11:
5:
1377:
1367:
1366:
1361:
1344:
1343:
1341:
1340:
1327:
1314:
1298:
1296:
1290:
1289:
1278:
1277:
1245:
1244:
1233:(4): 397–409.
1217:
1187:
1160:
1125:(1): 113–137.
1109:
1084:(90001): 255.
1068:
1059:|journal=
1035:
1007:
980:(6): 967–982.
955:
928:(6): 520–547.
912:
893:(2): 121–149.
877:
866:(1–2): 35–54.
839:
828:(3): 223–256.
801:
782:
747:
701:
680:(2): 345–357.
653:
608:
555:
554:
552:
549:
548:
547:
531:
521:
518:
502:
499:
454:
451:
424:
421:
367:parenchymatous
317:
314:
287:
286:
285:
284:
276:
268:
257:
256:
252:
251:
248:
241:
240:
234:
233:
219:
215:
214:
202:
198:
197:
195:Nematophytales
189:
185:
184:
176:
172:
171:
166:
162:
161:
153:
149:
148:
135:
134:
126:
125:
117:
116:
108:
107:
99:
98:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
35:
28:
15:
9:
6:
4:
3:
2:
1376:
1365:
1362:
1360:
1357:
1356:
1354:
1337:
1332:
1328:
1324:
1319:
1315:
1310:
1304:
1300:
1299:
1297:
1295:
1294:Nematothallus
1291:
1287:
1282:
1274:
1270:
1266:
1262:
1258:
1254:
1249:
1248:
1240:
1236:
1232:
1228:
1221:
1213:
1210:(in Polish).
1209:
1205:
1204:Spongiophyton
1201:
1199:
1198:Spongiophyton
1191:
1183:
1179:
1175:
1171:
1164:
1156:
1152:
1148:
1144:
1140:
1136:
1132:
1128:
1124:
1120:
1113:
1105:
1099:
1091:
1087:
1083:
1079:
1072:
1064:
1052:
1038:
1032:
1028:
1024:
1020:
1019:
1011:
1003:
999:
995:
991:
987:
983:
979:
975:
971:
970:Nematothallus
964:
962:
960:
951:
947:
943:
939:
935:
931:
927:
923:
916:
908:
904:
900:
896:
892:
888:
881:
873:
869:
865:
861:
857:
856:Nematothallus
850:
848:
846:
844:
835:
831:
827:
823:
816:
814:
812:
810:
808:
806:
798:(1–3): 39–56.
797:
793:
786:
778:
774:
770:
766:
762:
758:
751:
743:
739:
735:
731:
727:
723:
719:
715:
708:
706:
696:
691:
687:
683:
679:
675:
674:Palaeontology
671:
669:
668:Nematothallus
660:
658:
649:
645:
641:
637:
630:
628:
627:Nematothallus
619:
617:
615:
613:
604:
600:
596:
592:
588:
584:
580:
576:
572:
571:Nematothallus
565:
563:
561:
556:
545:
544:Nematothallus
541:
540:Spongiophyton
537:
536:
535:Spongiophyton
532:
529:
528:
524:
523:
517:
515:
511:
507:
498:
495:
494:Nematothallus
488:
483:
476:
471:
464:
461:Apertures in
459:
450:
448:
447:
442:
441:
436:
435:
429:
428:Nematothallus
420:
417:
416:Nematothallus
413:
412:Nematothallus
408:
406:
402:
401:Nematothallus
396:
391:
387:
383:
382:Nematothallus
379:
374:
372:
371:Nematothallus
368:
363:
362:Nematothallus
358:
355:
354:Nematothallus
351:
346:
344:
340:
339:Nematothallus
336:
332:
329:
326:
322:
321:Nematothallus
313:
311:
307:
303:
299:
295:
294:
293:Nematothallus
280:
277:
272:
269:
264:
261:
260:
258:
253:
246:
242:
239:
235:
228:
227:
226:Nematothallus
220:
217:
216:
209:
203:
200:
199:
196:
190:
187:
186:
183:
182:Nematophytina
177:
174:
173:
170:
169:"Nematophyta"
167:
164:
163:
160:
154:
151:
150:
145:
140:
136:
132:
127:
123:
118:
114:
109:
105:
100:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
38:
26:
25:Nematothallus
22:
19:
1293:
1256:
1252:
1230:
1226:
1220:
1211:
1207:
1203:
1197:
1190:
1173:
1169:
1163:
1122:
1118:
1112:
1098:cite journal
1081:
1077:
1071:
1040:. Retrieved
1017:
1010:
977:
973:
969:
925:
921:
915:
890:
886:
880:
863:
859:
855:
825:
821:
795:
791:
785:
768:
764:
760:
756:
750:
717:
713:
677:
673:
667:
639:
635:
626:
578:
574:
570:
543:
539:
533:
527:Cosmochlaina
525:
513:
509:
505:
504:
493:
491:
486:
474:
462:
446:Steganotheca
444:
438:
432:
427:
426:
415:
411:
409:
400:
388:epoch (late
381:
378:Pennsylvania
375:
370:
361:
359:
353:
349:
347:
343:Prototaxites
342:
338:
320:
319:
292:
291:
290:
279:N. williamii
278:
270:
262:
244:
238:Type species
225:
224:
207:
130:
112:
24:
18:
335:sporophytes
274:Lang (1937)
266:Lang (1937)
249:Lang (1937)
231:Lang (1937)
129:Cuticle of
111:Cuticle of
1353:Categories
1042:2008-05-06
757:Pachytheca
642:(4): 505.
629:revisited"
551:References
434:Hostinella
405:arthropods
390:Ordovician
331:land plant
296:is a form
271:N. radiata
1176:: 41–66.
1119:Brittonia
1061:ignored (
1051:cite book
510:N. lobata
506:N. taenia
440:Cooksonia
306:red algae
158: (?)
152:Kingdom:
1309:Q1491485
1303:Wikidata
1202:[On
1155:21794174
763:n.g.)".
520:See also
501:Synonyms
453:Affinity
328:thallose
255:Species
201:Family:
165:Phylum:
1336:1084839
1323:4908157
1261:Bibcode
1227:Lethaia
1147:2805564
1127:Bibcode
1002:1305385
982:Bibcode
950:3514860
930:Bibcode
922:PALAIOS
895:Bibcode
722:Bibcode
682:Bibcode
603:1306131
583:Bibcode
573:Lang".
423:Habitat
386:Caradoc
310:lichens
302:fossils
218:Genus:
188:Order:
175:Class:
156:Plantae
1153:
1145:
1033:
1000:
948:
740:
601:
350:et al.
1331:IRMNG
1151:S2CID
1143:JSTOR
998:JSTOR
946:JSTOR
742:92244
738:JSTOR
632:(PDF)
599:JSTOR
325:early
298:genus
1318:GBIF
1104:link
1063:help
1031:ISBN
508:and
443:and
37:PreꞒ
1269:doi
1235:doi
1178:doi
1174:106
1135:doi
1086:doi
1023:doi
990:doi
938:doi
903:doi
868:doi
830:doi
796:171
773:doi
730:doi
718:227
690:doi
644:doi
640:173
591:doi
395:450
1355::
1333::
1320::
1305::
1267:.
1257:15
1255:.
1231:44
1229:.
1212:19
1172:.
1149:.
1141:.
1133:.
1123:28
1121:.
1100:}}
1096:{{
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