Zosterophyllopsida

The Zosterophylls are a group of extinct plants. The taxon was first
Zosterophyllopsida
established by Banks in 1968 as the subdivision Zosterophyllophytina; they
have since also been treated as the division Zosterophyllophyta and the class Temporal range: Ludlow to Devonian
Zosterophyllopsida. They were among the first vascular plants in the fossil Pre OS D C P T J K Pg N
record, and had a world-wide distribution. They were probably stem-group
lycophytes, forming a sister group to the ancestors of the living lycophytes.[1]
By the late Silurian (late Ludlovian, about 420 million years ago) a diverse
assemblage of species existed, examples of which have been found fossilised in
what is now Bathurst Island in Arctic Canada.[2]

Contents Zosterophyllum species fossils

Scientific classification
1 Morphology
2 Taxonomy and classification Kingdom: Plantae
3 Phylogeny and genera Division: Tracheophytes
4 See also
Subdivision: Lycophytina
5 References
6 External links
Class: Zosterophyllopsida

Morphology
The stems of zosterophylls were either smooth or covered with small spines known
as enations, branched dichotomously, and grew at the ends by unrolling, a process
known as circinate vernation. The stems had a central vascular column in which the
protoxylem was exarch, and the metaxylem developed centripetally. The sporangia
were kidney-shaped (reniform), with conspicuous lateral dehiscence and were borne
[3]
laterally in a fertile zone towards the tips of the branches.

The zosterophylls were named after the aquatic flowering plant Zostera from a
mistaken belief that the two groups were related. David P. Penhallow's generic
description of the type genus Zosterophyllum refers to "Aquatic plants with creeping
stems, from which arise narrow dichotomous branches and narrow linear leaves of
the aspect of Zostera."[4] Zosterophyllum rhenanum was reconstructed as aquatic,
the lack of stomata on the lower axes giving support to this interpretation.[3]
However, current opinion is that the Zosterophylls were terrestrial plants, and
Penhallow's "linear leaves" are interpreted as the aerial stems of the plant that had
become flattened during fossilization.[5]

Stomata were present, particularly on the upper axes. Their absence on the lower Reconstruction of the zosterophyll
Sawdonia ornata.
portions of the axes suggests that this part of the plants may have been submerged,
and that the plants dwelt in boggy ground or even shallow water.[3] In many fossils
these appear to consist of a slit-like opening in the middle of a single elongated guard cell, leading to comparison with the stomata of
some mosses.[6] However, this is now thought to result from the loss of the wall separating paired guard cells during
fossilisation.[7][8]

Taxonomy and classification

At first most of the fossilized early land plants other than bryophytes were placed in the class Psilophyta, established in 1917 by
Kidston and Lang.[9] As additional fossils were discovered and described, it became apparent that the Psilophyta were not a
homogeneous group of plants, and in 1975 Banks developed his earlier proposal to split it into three groups, which he put at the rank
of subdivision. One of these was the subdivision Zosterophyllophytina, named after the genus Zosterophyllum.[10][11] For Banks,
zosterophyllophytes or zosterophylls comprised plants with lateral sporangia which released their spores by splitting distally (i.e.
away from their attachment), and which hadexarch strands of xylem.[12] Bank's classification produces the hierarchy:

Division Tracheata
Subdivision Zosterophyllophytina = zosterophyllophytes, zosterophylls
Subdivision Lycophytina = lycopods
+ other subdivisions

Those who treat most of the extant groups of plants as divisions may raise both the zosterophylls and the Lycophytina sensu Banks to
the rank of division:[13]

Division Zosterophyllophyta = zosterophylls, zosterophyllophytes

Division Lycophyta = lycophytes

In their cladistic study published in 1997,[14] Kenrick and Crane provided support for a clade uniting both the zosterophylls and the
lycopsids, producing a classification which places the zosterophylls in a class Zosterophyllopsida of the subdivisionycophytina:
L [15]

Division Tracheata
Subdivision Lycophytina = lycophytes
Class Zosterophyllopsida = zosterophylls
Class Lycopodiopsida = lycopsids

This approach has been widely used alongside previous systems. A consequence is that "lycophyte" and corresponding formal names
such as "Lycophyta" and "Lycophytina" are used by different authors in at least two senses: either excluding zosterophylls in the
sense of Banks or including them in the sense of Kenrick and Crane.

A further complication is that the cladograms of Kenrick and Crane show that the zosterophylls, broadly defined, are paraphyletic,
but contain a 'core' clade of plants with marked bilateral symmetry and circinate tips. The class Zosterophyllopsida sensu Kenrick &
Crane may be restricted to this core clade,[16] leaving many genera (e.g. Hicklingia, Nothia) with no systematic placement other than
Lycophytina sensu Kenrick & Crane, but nevertheless still informally called "zosterophylls".

Under whatever name and rank, the zosterophylls have been divided into orders and families, e.g. the Zosterophyllales containing the
Zosterophyllaceae and the Sawdoniales containing the Sawdoniaceae. Since the publication of cladograms showing that the group is
paraphyletic[14][17] divisions of the class have been less used, being ignored, for example, in the 2009 paleobotany textbook by
Taylor et al.[13]

Phylogeny and genera

In 2004, Crane et al. published a unified cladogram for the polysporangiophytes (plants with branched stems bearing sporangia),
based on cladistic analyses of morphological features.[9] This suggests that the zosterophylls were a paraphyletic stem group, related
to the ancestors of modern lycophytes.

lycophytes Hicklingia
 basal groups (Adoketophyton, Discalis, Distichophytum (=Rebuchia), Gumuia, Huia,
Zosterophyllum myretonianum, Z. lianoveranum, Z. fertile)

'core' zosterophylls (Zosterophyllum divaricatum, Tarella, Oricilla, Gosslingia, Hsua,

Thrinkophyton, Protobarinophyton, Barinophyton obscurum, B. citrulliforme, Sawdonia,
Deheubarthia, Konioria, Anisophyton, Serrulacaulis, Crenaticaulis)

basal groups (Nothia, Zosterophyllum deciduum)

lycopsids (extant and extinct members)

Genera which are included at or around the zosterophyll position in the cladogram or have otherwise been included in the group by at
least one source, and hence may be considered zosterophylls in the broad sense, are listed below
.

Adoketophyton Konioria
Anisophyton Macivera
Barinophyton Nothia
Bathurstia (B) Oricilla
Crenaticaulis (B) Protobarinophyton
Danziella Rebuchia, see Distichophytum
Deheubarthia Sawdonia (B)
Demersatheca Serrulacaulis
Discalis Tarella
Distichophytum (B) Thrinkophyton
Gosslingia (B) Trichopherophyton
Gumuia Ventarura
Hicklingia Wenshania
Hsua Zosterophyllum (B)
Huia
B = included by Banks in his 1975 description of Zosterophyllophytina.

Genera may not be assigned to this group by other authors; for example, Adoketophyton was regarded by Hao et al., who named the
genus, as having evolved separately from the lycopsids, so that its taxonomic placement was uncertain.[18] Other genera, such as
Jugumella, may be members of this group.[19]

See also
Drepanophycales, another clade of early lycopods

References
1. Gensel, P.G. (1992), "Phylogenetic relationships of the zosterophylls and lycopsids: evidence from morphology
,
paleoecology, and cladistic methods of inference", Annals of the Missouri Botanical Garden, 79: 45073,
doi:10.2307/2399750 (https://doi.org/10.2307%2F2399750), JSTOR 2399750 (https://www.jstor.org/stable/2399750)
2. Kotyk, M.E.; Basinger, J.F.; Gensel, P.G. & de Freitas, T.A. (2002), "Morphologically complex plant macrofossils from
the Late Silurian of Arctic Canada",American Journal of Botany, 89: 10041013, doi:10.3732/ajb.89.6.1004 (https://d
oi.org/10.3732%2Fajb.89.6.1004), PMID 21665700 (https://www.ncbi.nlm.nih.gov/pubmed/21665700)
3. Stewart, W.N. & Rothwell, G.W. (1993), Paleobotany and the evolution of plants(2nd ed.), Cambridge, UK:
Cambridge University Press,ISBN 978-0-521-38294-6
4. Penhallow, D.P. (1892), "Additional notes onDevonian plants from Scotland",Can. Rec. Sci., 5: 113
 5. Zhu, W.-Q. & Kenrick, P. (1999), "A Zosterophyllum-like plant from the Lower Devonian of Y
unnan Province, China",
Review of Palaeobotany and Palynology, 105 (12): 111118, doi:10.1016/S0034-6667(98)00070-0(https://doi.org/
10.1016%2FS0034-6667%2898%2900070-0)
6. Paton, J.A. & Pearce, J.V. (1957), "The occurrence, structure and functions of the stomata in British bryophytes",
Transactions of the British Bryological Society, 3: 228259, doi:10.1179/006813857804829560(https://doi.org/10.11
79%2F006813857804829560)
7. Edwards, D.; Edwards, D.S. & Rayner, R. (1982), "The cuticle of early vascular plants and its evolutionary
significance", in Cutler, D.; Alvin, K.L. & Price, C.E., The Plant Cuticle, London: Academic Press,ISBN 978-0-12-
199920-9
8. Edwards, D.; Abbott, G.D. & Raven, J.A. (1996), "Cuticles of early land plants: a paleoecophysiological evaluation",
in Kerstiens, G., Plant Cuticles, an integrated functional approach
, Oxford: BIOS Scientific,ISBN 978-1-85996-130-8
9. Crane, P.R.; Herendeen, P. & Friis, E.M. (2004), "Fossils and plant phylogeny"(http://www.amjbot.org/cgi/content/full/
91/10/1683), American Journal of Botany, 91 (10): 168399, doi:10.3732/ajb.91.10.1683(https://doi.org/10.3732%2
Fajb.91.10.1683), PMID 21652317 (https://www.ncbi.nlm.nih.gov/pubmed/21652317)
10. Banks, H.P. (1968), "The early history of land plants", in Drake, E.T
., Evolution and Environment: A Symposium
Presented on the Occasion of the 100th Anniversary of the Foundation of Peabody Museum of Natural History at
Yale University, New Haven, Conn.: Yale University Press, pp. 73107, cited in Banks 1980
11. Banks, H.P. (1975), "Reclassification of Psilophyta",Taxon, 24: 401413, doi:10.2307/1219491 (https://doi.org/10.23
07%2F1219491)
12. Banks, H.P. (1980), "The role of Psilophyton in the evolution of vascular plants",Review of Palaeobotany and
Palynology, 29: 165176, doi:10.1016/0034-6667(80)90056-1(https://doi.org/10.1016%2F0034-6667%2880%29900
56-1)
13. Taylor, T.N.; Taylor, E.L. & Krings, M. (2009),Paleobotany, The Biology and Evolution of Fossil Plants (2nd ed.),
Amsterdam; Boston: Academic Press,ISBN 978-0-12-373972-8, p. 1028
14. Kenrick, Paul & Crane, Peter R. (1997a),The Origin and Early Diversification of Land Plants: A Cladistic Study
,
Washington, D.C.: Smithsonian Institution Press, ISBN 978-1-56098-730-7
15. See, e.g., Berry, C.M. & Fairon-Demaret, M. (2001), "The Middle Devonian Flora Revisited", in Gensel, P
.G. &
Edwards, D., Plants invade the Land : Evolutionary & Environmental Perspectives, New York: Columbia University
Press, ISBN 978-0-231-11161-4
16. Zhu, W.-Q. & Kenrick, P. (1999), "A Zosterophyllum-like plant from the Lower Devonian of Yunnan Province, China",
Review of Palaeobotany and Palynology, 105 (12): 111118, doi:10.1016/S0034-6667(98)00070-0(https://doi.org/
10.1016%2FS0034-6667%2898%2900070-0)
17. Kenrick, P. & Crane, P.R. (1997b), "The origin and early evolution of plants on land",Nature, 389 (6646): 3339,
Bibcode:1997Natur.389...33K (http://adsabs.harvard.edu/abs/1997Natur .389...33K), doi:10.1038/37918 (https://doi.o
rg/10.1038%2F37918)
18. Hao, Shougang; Wang, Deming & Beck, Charles B. (2003), "Observations on anatomy ofAdoketophyton
subverticillatum from the Posongchong Formation (Pragian, Lower Devonian) of unnan,
Y China", Review of
Palaeobotany and Palynology, 127 (34): 175186, doi:10.1016/S0034-6667(03)00119-2(https://doi.org/10.1016%2
FS0034-6667%2803%2900119-2)
19. Raymond, A.; Gensel, P. & Stein, W.E. (2006), "Phytogeography of Late Silurian macrofloras",Review of
Palaeobotany and Palynology, 142 (34): 165192, doi:10.1016/j.revpalbo.2006.02.005(https://doi.org/10.1016%2F
j.revpalbo.2006.02.005)

External links
Palaeos Plants : Lycopsida : Zosterophyllopsida
"Introduction to the Zosterophylls".

Retrieved from "https://en.wikipedia.org/w/index.php?title=Zosterophyllopsida&oldid=785853893

"

This page was last edited on 15 June 2017, at 19:53.

Text is available under theCreative Commons Attribution-ShareAlike License ; additional terms may apply. By using this
site, you agree to the Terms of Use and Privacy Policy. Wikipedia is a registered trademark of theWikimedia
Foundation, Inc., a non-profit organization.