{{Short description|Extinct family of Paleozoic plants}} {{Automatic taxobox | fossil_range = {{Geological range|Early Carboniferous|latest=323}} | image = FossilFernLeavesPennsylvanianOhio.jpg | image_caption = Neuropteris ovata, from northeastern Ohio. | parent_authority = Němejc, 1963 | taxon = Calamopityaceae | authority = Solms, 1896 | subdivision_ranks = | subdivision = | synonyms = * '''Calamopityaea''' Scott, 1909 * '''Stenomyeleae''' Scott, 1923 }}
'''Calamopityaceae''' is the largest family of the division of extinct seed-bearing plants (spermatophytes) known as Pteridospermatophyta.<ref name=":0">{{Cite book|title=Botany for degree students : Gymnosperms.|last1=K.|first1=Sinha, A.|last2=Kumar.|first2=Anil|date=2006-01-01|publisher=S Chand|isbn=9788121926188|oclc=857708675}}</ref><ref name=":1">{{Cite book|url=https://books.google.com/books?id=F6B_xHO-IpoC&q=calamopityaceae&pg=PA118|title=Gymnosperm (naked seeds plant) : structure and development|last=Singh|first=V. P.|date=2006-01-01|publisher=Sarup & Sons|isbn=9788176256711|language=en}}</ref> It is the only family in the monotypic order '''Calamopityales'''.<ref name="Taylor 09">{{cite book |author1=Taylor, Edith L. |url=https://books.google.com/books?id=_29tNNeQKeMC&q=Protolepidodendrales |title=Paleobotany: The Biology and Evolution of Fossil Plants |author2=Taylor, Thomas N. |author3=Krings, Michael |date=2009 |publisher=Academic Press |isbn=9780080557830 |pages=271–74}}</ref> This family is characterized by its petioles and specific wood pattern, and it grew only in the Paleozoic era, specifically in North America and Europe.<ref name=":0" /><ref name=":2">{{Cite journal|jstor=2995557|title=An Ontogenetic Model for the Mississippian Seed Plant Family Calamopityaceae|last1=Hotton|first1=Carol|last2=Stein|first2=William|journal=International Journal of Plant Sciences|date=1994-03-01|volume=155|issue=2|pages=119–142|doi=10.1086/297152|s2cid=85151157}}</ref><ref name=":3">{{Cite book|title=Paleobotany: The Biology and Evolution of Fossil Plants (2)|last1=Taylor|first1=Edith|last2=Taylor|first2=Thomas|last3=Krings|first3=Michael|publisher=Academic Press|year=2009|isbn=9780123739728}}</ref> Three form genera within the family are diagnosed by their stem structure: ''Calamopitys'', ''Stenomyelon'', and ''Diichinia''.<ref name=":0" /> It was named by Solms-Laubach in 1896. Since then, its genera have been added to and grouped differently.<ref name=":0" />
== Morphology == Calamopityaceae is the largest family in Pteriodspermatophyta.<ref name=":1" /> This family is composed of gymnosperms, and because of their stem structure discovered through fossil rocks, they are considered to be in this division.<ref name=":5">{{Cite book|url=https://books.google.com/books?id=syR9CgAAQBAJ&q=calamopityaceae&pg=PT262|title=An Introduction to Paleobotany|last=Arnold|first=Chester A.|date=2013-04-16|publisher=Read Books Ltd|isbn=9781447495093|language=en}}</ref> However, nothing is known of their reproductive organs, but they are classified as seed plants based on their similarities to the Lyginopteridaceae and Medullasaceae families within Pteriodspermatophyta.<ref name=":2" /> Calamopityaceae resemble Lyginopteridaceae and Medullasaceae in the monoxylic wood structures in their stem; this structure suggests the stem (diameter less than 1.5 cm) was narrow during the Calamopityaceae plant lifetime.<ref name=":3" /><ref name=":5" /> Only some petiole tissue has been found; it is classified to be of the genus ''Kalymma'' and suggests the plant had large fronds.<ref name=":2" /><ref name=":3" /> To identify a genus within this family, this petiole structure and monoxylic wood must be present, as well as a much larger cortex than vascular cylinder.<ref name=":2" /> No fossil evidence has been found to describe on their seed and pollen (reproductive) organs, and therefore the species within this family show more variance than other families.<ref name=":0" /><ref name=":3" />
Gymnosperms, including those that are extinct, can be classified by their wood: monoxylic vs pycnoxylic.<ref name=":4">{{Cite book|url=https://books.google.com/books?id=4dvyNckni8wC&q=pycnoxylic&pg=PA4|title=Gymnosperms|last1=Bhatnagar|first1=S. P.|last2=Moitra|first2=Alok|date=1996-01-01|publisher=New Age International|isbn=9788122407921|language=en}}</ref> Monoxylic wood is soft and spongy and has a large pith and cortex.<ref name=":5" /> Pycnoxylic wood, which is more dense with less pith and cortex, is more commercially used.<ref name=":4" /> The three genera of Calamopityaceae, ''Calamopitys, Stenomyelon,'' and ''Diichnia'', show monoxylic wood stem patterns, and this is considered to be an essential classification of the family Calamopityaceae (hence why ''Bilignea, Eristophyton, Endoxylon,'' and ''Shenoxylon'' were removed from this family).<ref name=":0" />
== Origins == Calamopityaceae fossils have been found in North America and Europe, and they have been dated back to the Paleozoic era, specifically the Upper Devonian and Lower Carboniferous (Mississippian) periods.<ref name=":3" /><ref name=":5" /> Being from this early period, Calamopityaceae are significant as an example of some of the earliest seed plants and ancestors of angiosperms.<ref name=":2" /><ref name=":3" />
=== Examples of specific varieties and discoveries ===
* ''Stenomyelon tuedianum'': Calciferous Sandstone Series of Britain, 1912<ref name=":3" /> * ''Diichnia kentuckiensis'' and ''Diichnia readii'': New Albany Shale of Kentucky, 1937<ref name=":3" /> * ''Calamopitys embergeri'': Mid-Tournaisian of France, 1970<ref name=":3" /> * ''Calamopitys americana'': America, 1914<ref name=":3" />
== History == In 1856, the Austrian paleontologist, Franz Joseph Andreas Nicolaus Unger, was the first to find ''Calamopitys'', a genus of Calamopityaceae.<ref name=":0" /> This genus, which later was the root for the family name, was found in the Thuringian Forest.<ref name=":0" /> Forty years later, the family was named Calamopityaceae by Solms-Laubach.
Though the original genera, ''Calamopitys'', ''Stenomyelon'', and ''Diichnia'', still remain under this family classification, there have been historical additions to these groupings. Because the family is defined loosely on stem structure with nothing known about the foliage and reproductive structure, different genera have been added and removed from this family.<ref name=":3" /> Four more genera, ''Bilignea'', ''Eristophyton'', ''Endoxylon'', and ''Sphenoxylon'', were added to the family in 1936. These genera were classified by their pycnoxylic secondary wood pattern. In 1953, they were removed from the family with the intention of keeping the family composed of genera with monoxylic secondary wood.<ref name=":0" />
Like many "seed ferns", the affinities of the Calamopityales are unclear. Meyen (1984) placed them within the class Ginkgoopsida, which also includes ginkgos. Apart from ''Calamopitys'' and ''Stenomyelon'', he also placed the genera ''Lyrasperma'', ''Deltasperma'', ''Eosperma'', and ''Kalymma'' within Calamopityaceae.<ref>{{Cite journal |last=Meyen |first=Sergei V. |date=1984 |title=Basic features of gymnosperm systematics and phylogeny as evidenced by the fossil record |url=http://link.springer.com/10.1007/BF02874305 |journal=The Botanical Review |language=en |volume=50 |issue=1 |pages=1–111 |doi=10.1007/BF02874305 |issn=0006-8101|url-access=subscription }}</ref>
A classification scheme by Doweld (2001) considered Calamopityales to be an ''incertae sedis'' order within the phylum "Moresnetiophyta". Doweld also listed Buteoxylaceae <small>Barnard & Long 1973</small><ref>{{Cite journal |last1=Barnard |first1=P. D. W. |last2=Long |first2=A. G. |date=1973 |title=4.—On the Structure of a Petrified Stem and some Associated Seeds from the Lower Carboniferous Rocks of East Lothian, Scotland |url=https://www.cambridge.org/core/product/identifier/S008045680001499X/type/journal_article |journal=Transactions of the Royal Society of Edinburgh |language=en |volume=69 |issue=4 |pages=91–108 |doi=10.1017/S008045680001499X |issn=0080-4568|url-access=subscription }}</ref> as a synonym of Calamopityaceae, implying that its constituent fossil wood genera (''Buteoxylon'' and ''Triradioxylon'') should be referred to Calamopityaceae. This would additionally require treating the order Buteoxylonales as a synonym of Calamopityales.<ref>{{Cite book |last=Doweld |first=Alexander B. |url=https://books.google.com/books?id=MBAIAf3OAPcC |title=Prosyllabus Tracheophytorum: Tentamen systematis plantarum vascularium |publisher=GEOS |year=2001 |location=Moscow |language=EN, RU}}</ref> ''Stenomyelon'' has been compared favorably to ''Buteoxylon'' in more recent studies as well.<ref>{{Cite journal |last1=Conceição |first1=Domingas Maria da |last2=Esperança Júnior |first2=Mário G.F. |last3=Iannuzzi |first3=Roberto |last4=Recamonde-Mendoza |first4=Mariana |last5=Malta |first5=Guilherme B.B.O. |date=2023 |title=PaleoWood: A machine learning approach for determining the affinity of Paleozoic gymnosperm woods |url=https://linkinghub.elsevier.com/retrieve/pii/S0895981122004114 |journal=Journal of South American Earth Sciences |language=en |volume=121 |article-number=104125 |doi=10.1016/j.jsames.2022.104125|url-access=subscription }}</ref>
== Genera == Three genera of fossil wood are currently classified as belonging to the family Calamopityaceae, and their differences are distinguished by their decreasing primary xylem from ''Stenomyelon'', to ''Calamopitys'', to ''Diichnia''.<ref>{{Cite book|url=https://books.google.com/books?id=P0IRAAAAIAAJ&q=diarch+vs+medullation&pg=PA90|title=Geological Survey professional paper|last1=(U.S.)|first1=Geological Survey|last2=Management|first2=Geological Survey (U S. ) Geographic Names Information|date=1936-01-01|publisher=Govt. Print. Off.|language=en}}</ref> Additionally, ''Galtiera'', ''Triichnia'', and ''Bostonia'' are classified within the family by some authors,<ref>{{Cite journal |last1=Stein |first1=William E. |last2=Beck |first2=Charles B. |date=1992 |title=New information on Bostonia perplexa—an unusual member of the calamopityaceae from North America |url=https://linkinghub.elsevier.com/retrieve/pii/003466679290177I |journal=Review of Palaeobotany and Palynology |language=en |volume=72 |issue=1–2 |pages=73–102 |doi=10.1016/0034-6667(92)90177-I|hdl=2027.42/30042 |hdl-access=free }}</ref><ref>{{Cite journal |last1=Beck |first1=Charles B. |last2=Galtier |first2=Jean |last3=Stein |first3=William E. |date=1992 |title=A reinvestigation of Diichnia Read from the New Albany Shale of Kentucky |url= |journal=Review of Palaeobotany and Palynology |language=en |volume=75 |issue=1–2 |pages=1–32 |doi=10.1016/0034-6667(92)90147-9|hdl=2027.42/29689 |hdl-access=free }}</ref> and ''Faironia''<ref>{{Cite journal |last1=Decombeix |first1=Anne-Laure |last2=Galtier |first2=Jean |last3=Meyer-Berthaud |first3=Brigitte |date=2006 |title=Faironia difasciculata, a new gymnosperm from the Early Carboniferous (Mississippian) of Montagne Noire, France |url= https://hal.science/hal-00112100|journal=Review of Palaeobotany and Palynology |language=en |volume=142 |issue=3–4 |pages=79–92 |doi=10.1016/j.revpalbo.2006.03.020}}</ref> and ''Chapelia''<ref>{{Cite journal |last1=Beck |first1=Charles B. |last2=Bailey |first2=Robert E. |date=1967 |title=Plants of the New Albany Shale. III: ''Chapelia campbellii'' gen. n. |url=https://bsapubs.onlinelibrary.wiley.com/doi/10.1002/j.1537-2197.1967.tb10726.x |journal=American Journal of Botany |language=en |volume=54 |issue=8 |pages=998–1007 |doi=10.1002/j.1537-2197.1967.tb10726.x |issn=0002-9122|hdl=2027.42/141984 |hdl-access=free }}</ref> may also have affinities with the family.
=== ''Calamopitys'' ===
Type species - ''Calamopitys saturnii''<ref name=":0" />
There are six species within this genus, and it has the most species of any Calamopityaceae genera.<ref name=":5" /> Although ''Eristophyton'' is sometimes considered to be a subgenus under this genus, the distinction between pycnoxylic and monoxylic secondary wood maintains these genera as separate.<ref name=":5" /> In terms of structure, these plants under this genera have narrow stems with diameter 2–3 cm (or larger in ''C. embergeri'' and ''C. schweitzeri'').<ref name=":3" />
=== ''Stenomyelon'' ===
Type species - ''Stenomyelon tuedianum''<ref name=":0" />
Originally known as "Tweed Mill fossil", this genus consists of fewer species than ''Calamopitys'', but in addition to ''S. tuedianum'', species include ''S. primaevum'', ''S. heterangioides,'' and ''S. muratum.''<ref name=":0" />
=== ''Diichnia'' ===
Type species - ''Diichnia kentuckiensis''<ref name=":0" />
Species of this genus are classified based on the characteristics of the smallest primary xylem of Calamopityaceae and a five-angled pith, as seen in the stem cross-section.<ref name=":5" /> These characteristics separate this genus from the other genera.
=== Other form genera === Non-wood form genera referred to Calamopityaceae include:<ref>{{Cite journal |last1=Anderson |first1=John M. |last2=Anderson |first2=Heidi M. |last3=Cleal |first3=Chris J. |date=2007 |title=Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology |url=http://opus.sanbi.org/bitstream/20.500.12143/5611/1/Strelitzia_20_2007.pdf |journal=Strelitzia |volume=20 |pages=1–280}}</ref>
* ''Dolichosperma'' (ovulate organ) * ''Eurystoma'' (ovulate organ) * ''Lyrasperma'' (ovulate organ) * ''Kalymma'' (foliage) * ''Sphenopteridium'' (foliage)
==References== {{Reflist}}
{{Taxonbar|from1=Q25060013|from2=Q1026226}}
Category:Pteridospermatophyta Category:Carboniferous plants Category:Prehistoric plant families Category:Mississippian life Category:Mississippian first appearances Category:Mississippian extinctions