Stigmaria: Difference between revisions
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== Description and morphology == |
== Description and morphology == |
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[[File: |
[[File:Stigmaria fossil root in sandstone (Pottsville Group, Middle Pennsylvanian; Frazeysburg Pit, Muskingum County, Ohio, USA) 5 (28407351158).jpg|thumb|250px|right|A close-up of the circular scars where the root-like appendages were attached from the stigmarian rhizome. In sandstone. [[Pottsville Group]] (Middle Pennsylvanian), Frazeysburg Pit, Muskingum County, Ohio, U.S.]] |
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=== Overview === |
=== Overview === |
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=== Appendages ("rootlets") === |
=== Appendages ("rootlets") === |
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The stigmarian systems had rhizomorph axes that |
The stigmarian systems had rhizomorph axes that shows circular scars where the root-like appendages were formerly attached. These appendages were branched dichotomously, establishing the root [[abscission]] areas of the stigmarian system. Since the stigmarian systems were root-like, these lateral appendages indicate that these were modified leaves adapted to serve the function of abscission. ''Stigmaria'' had a complex branching structure; thus, it is comparable to the rhizomes of the extant (living) relative, the quillworts (genus [[Isoetes]]). ''Stigmaria'' comprises of four proximal axes connected to the trunk of lycophytes of its underground rooting structures.<ref name="Hetherington" /> |
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[[File:05July2008 022 SpringhillMinesFm branchingStigmaria.jpg|thumb| |
[[File:05July2008 022 SpringhillMinesFm branchingStigmaria.jpg|thumb|200px|left|''Stigmaria'' displaying branching pattern. [[Cumberland Group|Springhill Mines Formation]] (Pennsylvanian), Nova Scotia, Canada.]] |
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Along the rhizomorph axes, the appendages are |
The lateral appendages have a helical arrangement and a growth abscission mechanism. Along the rhizomorph axes, the appendages are connected to each axis in a circular pattern which would shed during the growth stage, forming the helical arrangements of ''Stigmaria'' root abscission areas. The four proximal axes dichotomize, creating a long underground system ranging up to {{cvt|15|m}} in radius, while being up to {{cvt|40|cm}} long and {{cvt|0.5-1|cm}} wide. The stigmarian rootlets consists of monarch vascular bundle enclosed by the inner and outer cortex. Evidently, the inner cortex and outer cortex is anchored by a hollow middle cortex, and a network of vascular branches extends between them.<ref>{{cite journal |last1=Weiss |first1=Frederick Ernest |year=1902 |title=The Vascular branches of Stigmarian rootlets |journal=Annals of Botany |publisher=JSTOR |volume=16 |pages=559–573 |jstor=43235190 |number=63}}</ref> |
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| ⚫ | [[Endarch]] is defined as the primary xylem of ''Stigmaria'', and organized in forked vascular strands encompassed by [[vascular cambium]]. [[Tracheid]]s of the secondary xylem are formed in spiral lines and consist of scalariform wall thickenings, while the fibrils are similar to those in the aerial branches. The scalariform tracheids along the stigmarian rhizomorph axes had lateral vascular and cork cambium as evidenced by its secondary xylem and [[meristem|meristematic tissues]].<ref>{{cite journal |last1=Rothwell |first1=Gar W. |last2=Pryor |first2=Janelle S. |year=1991 |title=Developmental dynamics of arborescent lycophytes—apical and lateral growth in Stigmaria ficoides |journal=American Journal of Botany |publisher=iley Online Library |volume=78 |pages=1740–1745 |doi=10.1002/j.1537-2197.1991.tb14538.x |number=12}}</ref> |
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[[File:Estonian Museum of Natural History Specimen No 193540 photo (g23 g23-25 1 jpg).jpg|thumb|150px|right|''Stigmaria'' fossil casts commonly have a distinctive cylindrical shape with circular scars all around the stigmarian rhizome. From the [[Estonian Museum of Natural History]], Tallinn, Estonia.]] |
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| ⚫ | [[Endarch]] is defined as the primary xylem of ''Stigmaria'', and organized in forked vascular strands encompassed by [[vascular cambium]]. [[Tracheid]]s of the secondary xylem are formed in spiral lines and consist of scalariform wall thickenings, while the |
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=== Development === |
=== Development === |
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| ⚫ | ''Stigmaria'' development is linked to the changes in aerial stems found in typical rhizomic structures seen in present plants. ''Stigmaria's'' features are unrelated when connecting to present plant functionality. Moreover, the spiral structure of the stigmarian rootlet attachment is separate from the asymmetrical changes of roots and rhizomes commonly seen in modern plants. While there were lateral appendages in ''Stigmaria'', none were found in the root systems of modern plants. However, fungi has [[mycorrhiza]]e, which are functioned from cortical [[parenchyma]] cells.<ref name="KarlJNiklas"/> |
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[[File:Stigmaria fossil root in sandstone (Pottsville Group, Middle Pennsylvanian; Frazeysburg Pit, Muskingum County, Ohio, USA) 5 (28407351158).jpg|thumb|250px|right|A close-up of the circular scars where the root-like appendages were attached from the stigmarian rhizome. In sandstone. [[Pottsville Group]] (Middle Pennsylvanian), Frazeysburg Pit, Muskingum County, Ohio, U.S.]] |
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| ⚫ | ''Stigmaria'' development is linked to the changes in aerial stems found in typical rhizomic structures seen in present plants. ''Stigmaria's'' features are unrelated |
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Though vascular bundles in leaves are bilaterally symmetrical including the appendages of ''Stigmaria'' and the monarch vascular bundle, present rhizomes have a radial point of symmetric vascular tissues. Furthermore, within a certain growth stage, foliar abscission (active shedding) of the appendages occurs from the stigmarian axis. Nonetheless, root abscission is relatively absent in modern plants. The stigmarian rootlets |
Though vascular bundles in leaves are bilaterally symmetrical including the appendages of ''Stigmaria'' and the monarch vascular bundle, present rhizomes have a radial point of symmetric vascular tissues. Furthermore, within a certain growth stage, foliar abscission (active shedding) of the appendages occurs from the stigmarian axis. Nonetheless, root abscission is relatively absent in modern plants. The stigmarian rootlets have a similarity to [[Lepidodendrales|arborescent lycophytes]], with functions related to absorbent organs, branching, and forking of proximal axes.<ref>{{cite journal |last1=Frankenberg |first1=Julian M. |last2=Eggert |first2=Donald A. |year=1969 |title=Petrified Stigmaria from North America: Part I. Stigmaria ficoides, the underground portions of Lepidodendraceae |journal=Palaeontographica Abteilung B |publisher=Schweizerbart'sche Verlagsbuchhandlung |pages=1–47}}</ref> |
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Since many lycopsids from the Paleozoic had a height of up to {{cvt|50|m}} meters, and grew in unsteady engulfed and saturated soil, the lycopods and their stigmarian system grew around the river systems.<ref name=Alekhin/><ref>{{cite journal|author1=Davies, N.S. |author2=Gibling, M. R.|year=2011|title=Evolution of fixed-channel alluvial plains in response to Carboniferous vegetation|journal=Nature Geoscience|volume=21|pages=629–633|doi=10.1038/ngeo1237|bibcode = 2011NatGe...4..629D|issue=9}}</ref> Therefore, it is debatable to how the underground system could handle the plants. Evidence to support their height was compared to the extensive stigmarian system. Thus, |
Since many lycopsids from the Paleozoic had a height of up to {{cvt|50|m}} meters, and grew in unsteady engulfed and saturated soil, the lycopods and their stigmarian system grew around the river systems.<ref name=Alekhin/><ref>{{cite journal|author1=Davies, N.S. |author2=Gibling, M. R.|year=2011|title=Evolution of fixed-channel alluvial plains in response to Carboniferous vegetation|journal=Nature Geoscience|volume=21|pages=629–633|doi=10.1038/ngeo1237|bibcode = 2011NatGe...4..629D|issue=9}}</ref> Therefore, it is debatable to how the underground system could handle the plants. Evidence to support their height was compared to the extensive stigmarian system. Thus, progression of the rhizomorph axes appeared to have [[secondary growth]] in their growth stages of the cortex. They may have been preferred to stand upright since arborescent lycophytes had bushy branches and only a few secondary xylem. The branches of neighboring lycopsids could interweave and deliver foundational support to the base stem.<ref name=KarlJNiklas>{{cite book |last1=Niklas |first1=Karl J. |url=https://books.google.com/books?id=l3bRJVMbNMcC&pg=PR9 |title=Plant biomechanics: an engineering approach to plant form and function |publisher=University of Chicago press |year=1992 |isbn=9780226586311}}</ref> On the other hand, the branch density and development of the wood in present trees can prevent [[tree uprooting|uprooting]].<ref name="Šamonil Král Hort 2010 pp. 65–79">{{cite journal | last=Šamonil | first=Pavel | last2=Král | first2=Kamil | last3=Hort | first3=Libor | title=The role of tree uprooting in soil formation: A critical literature review | journal=Geoderma | publisher=Elsevier BV | volume=157 | issue=3-4 | year=2010 | issn=0016-7061 | doi=10.1016/j.geoderma.2010.03.018 | pages=65–79}}</ref> |
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== Gallery == |
== Gallery == |
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Revision as of 05:20, 9 March 2024
| Stigmaria | |
|---|---|
| |
| Stigmaria, a fossil lycopsid rhizome | |
Scientific classification 
| |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Lycophytes |
| Class: | Lycopodiopsida |
| Order: | †Lepidodendrales |
| Genus: | †Stigmaria Brongn. |
Stigmaria is a form taxon for common fossils found in Carboniferous rocks.[1] They represent the underground rooting structures of arborescent lycophytes such as Sigillaria and Lepidodendron under the order Lepidodendrales.
Description and morphology
_5_(28407351158).jpg)
Overview
The Paleozoic swamps had tree-like lycopsids that grew up to 30 m (98 ft),[2] and even 50 m (160 ft) in height.[3] These lycopsid plants were anchored by an extensive network of branching underground structures with root-like appendages attached to them. The underground organs or structures of these lycopsids is referred to as Stigmaria.[4] Lycopsids first evolved during a rapid diversification of terrestrial land plants in the Devonian period[5] and became common plants within the Carboniferous coal forest flora. Lycopsids grew in low-level swampy wetland areas which they flourished during the Pennsylvanian age.[6] [7] Analysis of the morphology and anatomy of the stigmarian systems suggests that the axes around the structure were shoot-like, and so they are called rhizomes or rhizomorphs.[8] In general, common species of Stigmaria (Stigmaria ficoides sp.) have been analyzed extensively to provide an understanding of its morphology and histology.[4]
Appendages ("rootlets")
The stigmarian systems had rhizomorph axes that shows circular scars where the root-like appendages were formerly attached. These appendages were branched dichotomously, establishing the root abscission areas of the stigmarian system. Since the stigmarian systems were root-like, these lateral appendages indicate that these were modified leaves adapted to serve the function of abscission. Stigmaria had a complex branching structure; thus, it is comparable to the rhizomes of the extant (living) relative, the quillworts (genus Isoetes). Stigmaria comprises of four proximal axes connected to the trunk of lycophytes of its underground rooting structures.[8]
The lateral appendages have a helical arrangement and a growth abscission mechanism. Along the rhizomorph axes, the appendages are connected to each axis in a circular pattern which would shed during the growth stage, forming the helical arrangements of Stigmaria root abscission areas. The four proximal axes dichotomize, creating a long underground system ranging up to 15 m (49 ft) in radius, while being up to 40 cm (16 in) long and 0.5–1 cm (0.20–0.39 in) wide. The stigmarian rootlets consists of monarch vascular bundle enclosed by the inner and outer cortex. Evidently, the inner cortex and outer cortex is anchored by a hollow middle cortex, and a network of vascular branches extends between them.[9]
Endarch is defined as the primary xylem of Stigmaria, and organized in forked vascular strands encompassed by vascular cambium. Tracheids of the secondary xylem are formed in spiral lines and consist of scalariform wall thickenings, while the fibrils are similar to those in the aerial branches. The scalariform tracheids along the stigmarian rhizomorph axes had lateral vascular and cork cambium as evidenced by its secondary xylem and meristematic tissues.[10]
.jpg)
Development
Stigmaria development is linked to the changes in aerial stems found in typical rhizomic structures seen in present plants. Stigmaria's features are unrelated when connecting to present plant functionality. Moreover, the spiral structure of the stigmarian rootlet attachment is separate from the asymmetrical changes of roots and rhizomes commonly seen in modern plants. While there were lateral appendages in Stigmaria, none were found in the root systems of modern plants. However, fungi has mycorrhizae, which are functioned from cortical parenchyma cells.[11]
Though vascular bundles in leaves are bilaterally symmetrical including the appendages of Stigmaria and the monarch vascular bundle, present rhizomes have a radial point of symmetric vascular tissues. Furthermore, within a certain growth stage, foliar abscission (active shedding) of the appendages occurs from the stigmarian axis. Nonetheless, root abscission is relatively absent in modern plants. The stigmarian rootlets have a similarity to arborescent lycophytes, with functions related to absorbent organs, branching, and forking of proximal axes.[12]
Since many lycopsids from the Paleozoic had a height of up to 50 m (160 ft) meters, and grew in unsteady engulfed and saturated soil, the lycopods and their stigmarian system grew around the river systems.[3][13] Therefore, it is debatable to how the underground system could handle the plants. Evidence to support their height was compared to the extensive stigmarian system. Thus, progression of the rhizomorph axes appeared to have secondary growth in their growth stages of the cortex. They may have been preferred to stand upright since arborescent lycophytes had bushy branches and only a few secondary xylem. The branches of neighboring lycopsids could interweave and deliver foundational support to the base stem.[11] On the other hand, the branch density and development of the wood in present trees can prevent uprooting.[14]
Gallery
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Bedding plane view of a flattened Stigmaria preserved atop a shallow-water carbonaceous limestone. Joggins, Nova Scotia, Canada. -
In situ Stigmaria fossil from the Joggins Formation (Pennsylvanian), Cumberland Basin, Nova Scotia, Canada -
In situ lycopsid with attached stigmarian system from Joggins, Nova Scotia, Canada -
Stigmaria impression with visible rootlets connecting from the rhizome
References
- ^ Rothwell, GW; Erwin, DM (1985). "The rhizophore apex of Paurodendron: implications for homologies among the rooting organs of Lycopsida". American Journal of Botany. 72: 86–98. doi:10.2307/2443571. JSTOR 2443571.
- ^ Wang, Deming; Qin, Min; Lu, Le; Liu, Lu; Zhou, Yi; Zhang, Yingyang; Huang, Pu; Xue, Jinzhuang; Zhang, Shihui; Meng, Meicen (8 August 2019). "The Most Extensive Devonian Fossil Forest with Small Lycopsid Trees Bearing the Earliest Stigmarian Roots". Current Biology. 29: 2604–2615. doi:10.1016/j.cub.2019.06.053. Retrieved 13 January 2024.
- ^ a b V. V. Alekhin (1961). Geografiia rastenii s osnovani botaniki (Geography of plants and basics of botany). Gos. nauchno-pedagog. izd-vo. p. 167. Retrieved 2020-10-05.
- ^ a b Williamson, William Crawford (1887). A monograph on the morphology and histology of Stigmaria ficoides. Vol. 40. Palaeontographical society.
- ^ Xu, Hong-He; Yang, Ning; Bai, Jiao; Wang, Yao; Liu, Feng; Ouyang, Shu (1 February 2022). "Palynological assemblage of the Lower Devonian of Hezhang, Guizhou, southwestern China". Review of Palaeobotany and Palynology. 297: 104561. doi:10.1016/j.revpalbo.2021.104561. ISSN 0034-6667. S2CID 244048051. Retrieved 25 November 2023.
- ^ Cleal, C. J. & Thomas, B. A. (2005). "Palaeozoic tropical rainforests and their effect on global climates: is the past the key to the present?" Geobiology, 3, p. 13-31.
- ^ Christopher J. Cleal; Barry A.Thomas (1994). Plant Fossils of the British Coal Measures. The Palaeontological Association. ISBN 0-901702-53-6.
- ^ a b Hetherington, AJ; Berry, CM; Dolan, Liam (June 14, 2016). "Networks of highly branched stigmarian rootlets developed on the first giant trees" (PDF). PNAS. 113 (24): 6695–6700. doi:10.1073/pnas.1514427113. PMC 4914198. PMID 27226309. Retrieved 15 February 2017.
- ^ Weiss, Frederick Ernest (1902). "The Vascular branches of Stigmarian rootlets". Annals of Botany. 16 (63). JSTOR: 559–573. JSTOR 43235190.
- ^ Rothwell, Gar W.; Pryor, Janelle S. (1991). "Developmental dynamics of arborescent lycophytes—apical and lateral growth in Stigmaria ficoides". American Journal of Botany. 78 (12). iley Online Library: 1740–1745. doi:10.1002/j.1537-2197.1991.tb14538.x.
- ^ a b Niklas, Karl J. (1992). Plant biomechanics: an engineering approach to plant form and function. University of Chicago press. ISBN 9780226586311.
- ^ Frankenberg, Julian M.; Eggert, Donald A. (1969). "Petrified Stigmaria from North America: Part I. Stigmaria ficoides, the underground portions of Lepidodendraceae". Palaeontographica Abteilung B. Schweizerbart'sche Verlagsbuchhandlung: 1–47.
- ^ Davies, N.S.; Gibling, M. R. (2011). "Evolution of fixed-channel alluvial plains in response to Carboniferous vegetation". Nature Geoscience. 21 (9): 629–633. Bibcode:2011NatGe...4..629D. doi:10.1038/ngeo1237.
- ^ Šamonil, Pavel; Král, Kamil; Hort, Libor (2010). "The role of tree uprooting in soil formation: A critical literature review". Geoderma. 157 (3–4). Elsevier BV: 65–79. doi:10.1016/j.geoderma.2010.03.018. ISSN 0016-7061.