102:
734:
131:
394:
593:
714:-shaped structures usually 3 to 8 mm in diameter with a hardened outer covering. This outer covering is tough and resistant to drying out, allowing the spores inside to survive unfavorable conditions such as winter frost or summer desiccation. Despite this toughness, the sporocarps will open readily in water if conditions are favorable, and specimens have been successfully
457:, which are also aquatic and heterosporous. However, both of these other fern families float freely on the surface of ponds or lakes instead of rooting in soil or mud. The close relationship of these groups to the Marsileaceae is supported by both morphologic and molecular analysis, as well as by the discovery of an intermediate
669:. Some of their leaves grow up to the surface of the water, and look just like leaves of species growing out of water. These plants also produce other leaves with shorter leaf stalks that are not long enough to reach the surface, and so the leaflets remain underwater. These leaves have different anatomical and
810:. The sorophore is a sorus-bearing structure unique to the Marsileaceae; it may extend to more than ten times the length of the sporocarp inside which it was coiled. This extension carries the numerous spore-producing sori attached along each side of the sorophore out into the water.
798:, it is very rare among other groups of plants. Also, most heterosporous plants produce their two kinds of sporangia in different places on the plant. Since the Marsileaceae grow both kinds together in a single cluster, they differ from other plants in this regard as well.
801:
The spores remain dormant inside the sporocarp through unfavorable conditions, but when conditions are suitable and wet, the sporocarp will germinate. It splits into halves, allowing the tissue coiled inside to become
445:
grows widely in temperate regions of both the northern and southern hemispheres. Its leaves do not subdivide into leaflets but are slender and tapered to a point, so that it is often overlooked and mistaken for a
624:
are not borne equally. Instead, they are borne in pairs with one pair of leaflets attached slightly higher than the other. Thus in the developing leaf, the leaflets are folded more like the wings of a
751:
The sporocarps are functionally and developmentally modified leaflets, although they have much shorter stalks than the vegetative leaflets. Inside the sporocarp, the modified leaflets bear several
718:
after being stored for more than 130 years. Each growing season, only one sporocarp typically develops per node along the rhizome near the base of the other leaf-stalks, though in some species of
551:
The
Marsileaceae share many of the basic structural characteristics common to most ferns, but the differences are more noticeable than the similarities. Species of this family have long, slender
342:, though at first sight they do not physically resemble other ferns. The group is commonly known as the "pepperwort family" or as the "water-clover family" because the leaves of the genus
688:-producing) leaflets at or near the base of the photosynthetic leaves. This reproductive portion looks and functions very differently from the vegetative portion of the leaves.
425:
can be distinguished from the other two genera by the presence of four leaflets on each leaf, although some species occasionally produce six leaflets per leaf. A second genus
765:). Each sorus includes a mix of two types of sporangium, each type producing only one of two kinds of spores. Toward the center of each sorus and developing first are the
502:
608:
ending in zero, two, or four (occasionally six) leaflets. The number of leaflets differs among the three genera and can therefore be used for identification. In
530:
262:
1638:
806:. As this internal tissue swells with water, it pushes the halves of the hard outer covering apart, and emerges as a long gelatinous worm-like
1677:
463:
1424:
386:
on the water surface. They grow in seasonally wet habitats, but survive the winter or dry season by losing their leaves and producing hard,
567:
along the rhizome, with wide spacing between leaf clusters. As a result, the plants appear to be more stem than leaf, unlike other ferns.
1586:
1651:
1599:
1011:
200:
407:
There are only three living genera in the family
Marsileaceae. The majority of species (about 45 to 70) belong to the genus
1656:
528:. The fossils include leaves with visible veins, as well as sporocarps. The currently oldest known member of the family is
573:
grow primarily from the same nodes as the leaves, but may also grow from other locations along the rhizome. The roots of
1197:
gen. et sp. nov., the Oldest
Macrofossil of Marsileaceae, from the Upper Jurassic to Lower Cretaceous of Western Japan".
467:. In general, the Salviniaceae and Azollaceae have a much better fossil record than the Marsileaceae. Until recently,
1369:
1341:
1313:
1135:
1068:
991:
894:
1777:
1682:
1625:
1731:
1772:
642:
which unroll as the leaf matures. At full maturity they are held erect with the leaflets unfolded, except in
1664:
1010:
Smith, Alan R.; Kathleen M. Pryer; Eric
Schuettpelz; Petra Korall; Harald Schneider; Paul G. Wolf (2006).
1495:
784:
Because the
Marsileaceae produce two kinds of spore (and thus two kinds of gametophyte), they are called
130:
1726:
1578:
1552:
1413:
882:
1604:
604:
The leaves are the most easily observed characteristic for the
Marsileaceae; they have a long slender
1692:
650:, losing their vegetative leaves in winter. Tropical species may also lose their leaves during the
1305:
1232:
Sun, Chunlin; Li, Tao; Na, Yuling; Wu, Wenhao; Li, Yunfeng; Wang, Lixia; Zhang, Lijun (July 2014).
921:"Phylogeny of Marsileaceous Ferns and Relationships of the Fossil Hydropteris pinnata Reconsidered"
1573:
1421:
1818:
1718:
1547:
1486:
1448:
831:
1234:"Flabellariopteris, a new aquatic fern leaf from the Late Triassic of western Liaoning, China"
983:
977:
1713:
1643:
739:
665:, especially those growing with their rhizome submerged, may have vegetative leaves that are
1754:
1705:
1565:
1297:
1813:
1508:
1245:
722:
there may be two or occasionally as many as twenty. The resemblance of the sporocarps to
8:
1397:
1298:
1539:
1249:
704:
when they reproduce. Unlike other ferns, the spores in this family are produced inside
1785:
1560:
1269:
1214:
1175:
1105:
1034:
951:
889:(volume 2 ed.). New York & Oxford: Oxford University Press. pp. 331–335.
125:
101:
1700:
1433:
1365:
1337:
1309:
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1261:
1233:
1152:
1131:
1064:
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706:
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335:
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1179:
955:
1790:
1253:
1206:
1167:
1097:
1026:
935:
803:
666:
605:
382:, or they may grow submerged in shallow water with some of the leaves extending to
347:
328:
280:
1009:
589:. These vessels have evolved independently of vessels in other groups of plants.
1500:
1428:
791:
774:
1669:
1764:
1471:
1391:
1153:"Marsileaceae Sporocarps and Spores from the Late Cretaceous of Georgia, U.S.A"
979:
A Field Manual of the Ferns & Fern-Allies of the United States & Canada
766:
670:
655:
582:
155:
1257:
1807:
1265:
786:
564:
536:
497:
367:
332:
231:
220:
59:
1744:
1088:
gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns".
947:
723:
454:
427:
238:
790:. While heterospory is the norm among all plants with seeds, such as the
773:. Surrounding them at the edge of the sorus and developing later are the
1612:
1480:
677:
512:
387:
246:
185:
34:
1151:
Lupia, R.; H. Schneider; G. M. Moeser; K. M. Pryer; P. R. Crane (2000).
1150:
1038:
1109:
849:
778:
758:
651:
612:, the leaves are narrowly cylindrical and taper to a point. Leaves of
521:
493:
79:
44:
1063:. Washington, D. C.: Smithsonian Institution Press. pp. 243–244.
1061:
1617:
1534:
1521:
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770:
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681:
647:
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441:
436:
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227:
110:
84:
1630:
1591:
1442:
1101:
393:
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1465:
1336:. New York and London: McGraw-Hill Book Company. pp. 197–223.
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1171:
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116:
74:
69:
54:
49:
39:
1526:
592:
795:
585:. Vessels have also been found in the rhizome of two species of
552:
506:, and pushed the known history of the Marsileaceae back into the
89:
64:
620:
bear four leaflets at the tip. The four leaflets on the leaf of
852:, so careful preparation methods must be used in order for the
823:
639:
458:
432:
379:
1304:(3rd ed.). New York: W. H. Freeman and Company. pp.
673:
characteristics better suited to their submerged environment.
319:
1513:
753:
697:
685:
556:
525:
489:
485:
478:
447:
431:
includes a single living species that grows only in southern
351:
142:
492:. In 2000, the discovery of fossilized sporocarps from the
286:
711:
701:
570:
560:
500:
was announced. These fossils were assigned to the species
375:
339:
313:
307:
301:
175:
165:
1396:(3rd ed.). New York: The Macmillan Company. pp.
1295:
1130:. Englewood Cliffs, NJ: Prentice Hall. pp. 434–435.
371:
295:
982:. Washington, D. C.: Smithsonian Institution. pp.
370:
or semi-aquatic. Plants often grow in dense clumps in
696:
473:
was the oldest fossil member known; it is a preserved
439:; it has only two leaflets per leaf. The third genus
1438:
article describing
Cretaceous fossils of Marsileaceae
1291:
1289:
1287:
1285:
1283:
1125:
1083:
1058:
535:
described in 2014 from isolated leaves dating to the
354:
and 50 to 80 species with most of those belonging to
310:
292:
289:
1084:
Rothwell, G. W.; R. A. Stockey (1994). "The role of
298:
1192:
1121:
1119:
848:. However, the sporocarps contain toxic levels of
769:, each of which will produce a single large female
646:whose leaves have no blade. Temperate species are
304:
283:
1280:
658:, and produce most of the food used by the plant.
453:The closest relatives of the Marsileaceae are the
1393:The Structure and Development of Mosses and Ferns
1805:
1385:
1383:
1381:
1116:
1052:
971:
969:
967:
965:
680:) leaflets, all species of Marsileaceae produce
1296:Gifford, Ernest M.; Adriance S. Foster (1988).
555:that creep along or beneath the ground. Their
876:
874:
872:
870:
868:
1378:
1355:
1353:
1327:
1325:
962:
777:, each of which will produce many small male
632:As with other ferns, the leaves develop in a
1364:. Portland: Timber Press. pp. 243–249.
1334:Morphology of Vascular Plants (Lower Groups)
1077:
1005:
1003:
914:
912:
910:
908:
906:
1300:Morphology and Evolution of Vascular Plants
1126:Taylor, Thomas N.; Edith L. Taylor (1993).
865:
450:. There are only about five species known.
316:
1350:
1322:
1193:Yamada, Toshihiro; Masahiro Kato (2002). "
1186:
1128:The Biology and Evolution of Fossil Plants
100:
1231:
1144:
1000:
975:
903:
616:bear two broad leaflets, while leaves of
1389:
732:
591:
392:
1199:International Journal of Plant Sciences
1160:International Journal of Plant Sciences
928:International Journal of Plant Sciences
880:
346:superficially resemble the leaves of a
1806:
1059:Kenrick, Paul; Peter R. Crane (1997).
887:Flora of North America north of Mexico
761:covered by a thin hood of tissue (the
1447:
1446:
1359:
1331:
918:
638:pattern. They begin as small, tight
1778:275e64d5-2202-44aa-b978-9c7c86060b66
1693:354625b8-9d4e-454c-aec9-5a18aaa9d971
757:, each of which consists of several
726:gives the family its common name of
629:than like the leaflets of a clover.
390:-resistant reproductive structures.
1390:Campbell, Douglas Houghton (1918).
1012:"A classification for extant ferns"
710:. These are hairy, short-stalked,
13:
1732:urn:lsid:ipni.org:names:77126791-1
361:
14:
1830:
1407:
676:In addition to their vegetative (
413:, which grows worldwide in warm-
366:Members of the Marsileaceae are
350:. In all, the family contains 3
279:
129:
32:
1414:Marsileacae description in the
1225:
596:Leaves of the Hawaiian species
563:) grow in distinct clusters at
581:are noteworthy for containing
1:
859:
813:
691:
546:
1422:Diversity map for US species
976:Lellinger, David B. (1985).
856:to be safe for consumption.
7:
919:Pryer, Kathleen M. (1999).
10:
1835:
1362:A Natural History of Ferns
1090:American Journal of Botany
881:Johnson, David M. (1993).
1455:
1332:Eames, Arthur J. (1936).
1258:10.1007/s11434-014-0359-6
435:and neighboring parts of
212:
207:
126:Scientific classification
124:
108:
99:
23:
1360:Moran, Robin C. (2004).
1238:Chinese Science Bulletin
661:Some aquatic species of
510:. Other remains include
503:Regnellidium upatoiensis
1416:Flora of North America
1195:Regnellites nagashimae
832:Indigenous Australians
748:
601:
513:Regnellites nagashimae
404:
397:An African species of
1714:Paleobiology Database
740:Pilularia globulifera
737:The European species
736:
595:
403:with floating leaves.
396:
374:along the shores of
30:Late Triassic–Recent
1250:2014ChSBu..59.2410S
1086:Hydropteris pinnata
654:. These leaves are
1427:2007-02-09 at the
842:sporocarps called
822:are cultivated in
749:
602:
470:Rodeites dakshinii
405:
223:(45 to 65 species)
1801:
1800:
1701:Open Tree of Life
1449:Taxon identifiers
1244:(20): 2410–2418.
531:Flabellariopteris
338:and semi-aquatic
272:
271:
263:Flabellariopteris
203:
1826:
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1041:. Archived from
1031:10.2307/25065646
1016:
1007:
998:
997:
973:
960:
959:
925:
916:
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818:Some species of
792:flowering plants
598:Marsilea villosa
348:four-leaf clover
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27:Temporal range:
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1429:Wayback Machine
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1408:External links
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1321:
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1279:
1224:
1211:10.1086/342036
1205:(5): 715–723.
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1172:10.1086/317567
1166:(6): 975–988.
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1136:
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1096:(4): 479–492.
1076:
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1048:on 2008-02-26.
1025:(3): 705–731.
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992:
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940:10.1086/314177
934:(5): 931–954.
902:
895:
883:"Marsileaceae"
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838:of pulverized
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1566:Marsileaceae
1487:Marsileaceae
1457:Marsileaceae
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275:Marsileaceae
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196:Marsileaceae
195:
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24:Marsileaceae
18:
1814:Salviniales
1613:iNaturalist
1481:Wikispecies
779:microspores
724:peppercorns
496:of eastern
481:, found in
477:containing
464:Hydropteris
388:desiccation
247:Regnellites
242:(1 species)
234:(6 species)
186:Salviniales
1808:Categories
1631:30024011-2
860:References
850:thiaminase
814:Human uses
745:sporocarps
728:pepperwort
716:germinated
707:sporocarps
700:, but not
692:Life cycle
652:dry season
606:leaf stalk
547:Morphology
522:Cretaceous
494:Cretaceous
421:regions.
162:Division:
114:(top) and
1535:FloraBase
1274:129560335
1266:1001-6538
826:pools or
808:sorophore
771:megaspore
759:sporangia
667:dimorphic
648:deciduous
644:Pilularia
635:circinate
627:butterfly
610:Pilularia
543:, China.
520:or Lower
475:sporocarp
442:Pilularia
437:Argentina
415:temperate
228:Pilularia
139:Kingdom:
120:(bottom)
111:Pilularia
1773:VicFlora
1762:VASCAN:
1745:42000403
1740:Tropicos
1472:Q1115420
1466:Wikidata
1425:Archived
1219:84631686
1180:16981770
1039:25065646
956:14073501
948:10506474
840:Marsilea
836:porridge
820:Marsilea
804:hydrated
796:conifers
763:indusium
743:bearing
720:Marsilea
671:cellular
663:Marsilea
622:Marsilea
618:Marsilea
587:Marsilea
575:Marsilea
553:rhizomes
541:Liaoning
518:Jurassic
508:Mesozoic
483:Tertiary
423:Marsilea
419:tropical
410:Marsilea
400:Marsilea
356:Marsilea
344:Marsilea
255:Rodeites
217:Marsilea
192:Family:
117:Marsilea
1246:Bibcode
1110:2445498
984:303–307
830:. The
828:aquaria
682:fertile
678:sterile
640:spirals
380:streams
368:aquatic
336:aquatic
258: †
250: †
208:Genera
182:Order:
172:Class:
143:Plantae
1755:447236
1752:uBio:
1706:164369
1690:NZOR:
1644:113879
1436:et al.
1434:Lupia
1368:
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1308:–313.
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854:nardoo
845:nardoo
824:garden
698:spores
561:leaves
557:fronds
479:spores
461:named
459:fossil
433:Brazil
352:genera
329:family
1719:54774
1683:13814
1657:17991
1639:IRMNG
1618:57886
1579:10539
1561:FoAO2
1553:10539
1540:22731
1527:1MASF
1501:54548
1400:–439.
1270:S2CID
1215:S2CID
1176:S2CID
1156:(PDF)
1106:JSTOR
1046:(PDF)
1035:JSTOR
1019:Taxon
1015:(PDF)
952:S2CID
924:(PDF)
702:seeds
686:spore
571:Roots
565:nodes
526:Japan
490:India
486:chert
448:grass
384:float
376:ponds
340:ferns
201:Mirb.
150:Clade
1727:POWO
1678:NCBI
1652:ITIS
1626:IPNI
1605:1257
1600:GRIN
1592:2375
1587:GBIF
1522:EPPO
1514:4498
1496:APNI
1366:ISBN
1338:ISBN
1310:ISBN
1262:ISSN
1132:ISBN
1065:ISBN
988:ISBN
944:PMID
891:ISBN
794:and
754:sori
712:bean
577:and
417:and
35:PreꞒ
1786:WFO
1765:164
1665:NBN
1574:FoC
1548:FNA
1509:EoL
1398:417
1306:305
1254:doi
1207:doi
1203:163
1168:doi
1164:161
1098:doi
1027:doi
936:doi
932:160
539:in
524:of
488:of
378:or
372:mud
331:of
287:ɑːr
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281:/
277:(
266:†
90:N
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75:J
70:T
65:P
60:C
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50:S
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