Macroneuropteris

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Macroneuropteris
Temporal range: Carboniferous
PB000058-large.jpg
Macroneuropteris scheuchzeri, specimen from Mazon Creek fossil beds in the collection at the Yale Peabody Museum of Natural History
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Tracheophytes
Division:† Pteridospermatophyta
Order:† Medullosales
Family:† Neurodontopteridaceae
Genus:† Macroneuropteris
Cleal et al., 1990
Species
  • M. scheuchzeri
  • M. macrophylla
  • M. britannica
  • M. subauriculata

Macroneuropteris is a genus of Carboniferous seed plants in the order Medullosales. The genus is best known for the species Macroneuropteris scheuchzeri, a medium-size tree that was common throughout the late Carboniferous Euramerica. Three similar species, M. macrophylla, M. britannica and M. subauriculata are also included in the genus.

Contents

Taxonomic history

Fossils on Plate 5 of Lythophylacii Britannici Ichnographia.jpg
This 1699 illustration is the first depiction of Macroneuropteris (lower right).
Early illustration of Macroneuropteris scheuchzeri.jpg
An illustration of Phyllites mineralis (now Macroneuropteris scheuchzeri) in Herbarium Diluvianum by Johann Jakob Scheuchzer, 1723.
Johann Jakob Scheuchzer.jpg
The most common species of the genus is named after Johann Jakob Scheuchzer.

The most abundant species of this genus, Macroneuropteris scheuchzeri, has had a long taxonomic history since it was first recognized in fossils found near Oxford, England by Edward Lhuyd in 1669. He referred to these leaves as Phyllites mineralis. It is illustrated and noted in Lhuyd's Lythophylacii Britannici Ichnographia, an early manuscript on English fossils published in 1699 with the financial help of Isaac Newton. [1] [2] The species was further described in the Herbarium Diluvianum written in 1723 by the botanist Johann Jakob Scheuchzer. [3] Nearly hundred years after Scheuchzer's death, the species was renamed Neuropteris scheuchzeri by professor Hoffmann in Christian Keferstein's 1826 atlas of German Geology, Deutschland, geognostisch-geologisch dargestellt. [4] In the 1800s, similar fossilized foliage was found in North America. The names Neuropteris cordata var. angustifolia, Neuropteris angustifolia, Neuropteris acutifolia, Neuropteris hirsuta, Neuropteris decipiens, and Odontopteris subcuneata were used for these but are now all considered to be forms of Macroneuropteris scheuchzeri. The genus was taxonomically refined in 1989 through epidermal research led by C. J. Cleal. Based on that work, the genus Neuropteris was divided into four genera, Neuropteris, Macroneuropteris, Neurocallipteris, and Laveineopteris. [5] [6] [7]

Description

Macroneuropteris and Reconstruction of Medullosales.jpg
Macroneuropterisfrond.jpg
Macroneuropteris scheuchzeri leaves and reconstruction of a generalized medullosalean tree. On right, an illustration showing the large leaf of Macroneuropteris scheuchzeri dividing at the base into two bipinnately compound halves.

The genus Macroneuropteris is used in some cases as a leaf organ taxon to refer to just the foliage of these trees. And in other cases, it is used to refer to the entire tree. The genus is associated with the stems and trunks of the wood organ taxon Medullosa noei. Together these fossils describe parts of a medullosalean seed fern tree that was likely about 8–10 meters tall with an upright trunk with large compound frond-like leaves.

Foliage

Macroneuropteris scheuchzeri specimen clearly showing the hair-like structures on the leaves. Macroneuropteris hair structure.jpg
Macroneuropteris scheuchzeri specimen clearly showing the hair-like structures on the leaves.

The foliage of the Macroneuropteris species consists of very large frond-like leaves that are bipartite (divided in two) near the base, forming two large bipinnately compound parts (see illustration). These compound fronds can be as large as several meters. [8]

In Macroneuropteris, each individual leaflet or pinnule of the compound frond were also notably large. In fact, the species M. scheuchzeri's pinnules were the largest of any seed plant of the Carboniferous Period. Individual pinnules are typically lanceolate with a round base. They have been found to be as long as 12 cm. These individual leaflets are often found fossilized by themselves separated from the frond. They have been inferred by some to be deciduous. In the famous Mazon Creek Fossil Beds of Illinois, these leaflets are one of the most commonly found plant fossils. The leaves have thick cuticles, sunken stomata, dense trichomes, and large hair-like structures. These foliar characteristics combined with the spiny stem structure where the leaflets drop, and the potential deciduous nature have led to many authors suggesting a xeromorphic tendency in the tree. Such adaptations may have allowed the genus to dominate the late Carboniferous landscape as other plants like Lepidodendrales steadily declined. [9] [10]

'Hairs' on the leaves

The presence of hair-like structures on the pinnules of Macroneuropteris has been noted since the mid 1800s. It has become an important taxonomic characteristic particularly for M. scheuchzeri, which has abundant epicuticular hair that can reach a maximum length of 1000 mm. It had been assumed that these were trichomes on the leaves and may have been used to help the plant conserve water. Recent molecular studies by Erwin L. Zodrow have discovered that although there are trichomes on the species of Macroneuropteris, the more noticeable dark 'hair-like' structures are likely not trichomes and may not be directly attached to the leaves. He suggests that these structures are material in the wax of the cuticle demonstrating a dynamic molecular Self-assembly. [11]

The hair-like structures of M. scheuchzeri (1) are not organically attached to the abaxial surface; (2) differ spectrochemically from the organic material of the lamina; (3) are composed, in contrast with the trichomes, of relatively long, unbranched aliphatic (polymythelinic) hydrocarbon chains [CH2]n, and (4) are acellular and black, unlike true trichomes of the species that are multicellular. Overall, the sum-total of these experimental results supports the postulate for dynamic molecular self-assembly. For this reason the term "extracuticular deposit" is proposed, reflecting the origin and emergent nature of such hair-like structures in the abaxial pinnule.

— Erwin L. Zodrow, "Molecular self-assembly: Hypothesized for 'hair' of Macroneuropteris scheuchzeri", International Journal of Coal Geology [12]

Reconstruction

Reconstructions of the entire Macroneuropteris trees have been based on various separate fossil parts. As noted above, the stems and trunks are usually referred to as Medullosa noei. Early attempts to reconstruct the entire tree were somewhat limited by this fragmentary material. One well-known reconstruction was illustrated for Stewart and Delevoryas paper in 1956. The illustration has been the basis for many reconstructions of the Medullosa noei tree. [13]

However, a complete Macroneuropteris tree was found in growth position in Nova Scotia that differs from the idealized reconstruction. This fossil tree was extensively studied by Howard Falcon-Lang who found many characteristics that differed from the previous reconstructions.

The fossil tree has a sharply tapering trunk surrounded in its lower part by a large number of downward-recurved senescent petioles, which form a skirt. Petioles borne in an upright or horizontal position, interpreted as fronds that were still photosynthetically active when buried, are confined to the uppermost preserved part of the tree. Adapted to growth in rapidly aggrading coastal wetlands, the skirt of Macroneuropteris scheuchzeri probably acted to prop up the trunk while additionally trapping large mounds of mud around the base of the tree and stabilizing coastal wetlands. The tree had a sprawling habit and a maximum height of about 2 m. Similar, but smaller, trees found in adjacent beds probably represent juvenile specimens of the same species.

— Howard Falcon-Lang, "A Macroneuropteris scheuchzeri tree preserved in growth position", Atlantic Geology [14]

Reproduction

Seed and pollen organs have not yet been found directly attached to the foliage of Macroneuropteris. For that reason, a variety of fossilized reproductive parts could be possible matches. However, a lot of evidence points toward some type of trigonocarpus as the seed/ovlule and Codonotheca caduca as the male pollen organ.

Ovules/seeds

Trigonocarpus, the seed/ovule of Medullosales such as Macroneuropteris Trigonocarpus.jpg
Trigonocarpus, the seed/ovule of Medullosales such as Macroneuropteris

In 1938, W. A. Bell studied the Sydney Coalfield in Nova Scotia, and suggested that the large fossilized seeds called Trigonocarpus noeggerati could be the ovules of Macroneuropteris scheuchzeri. [15] Erwin Zodrow in 2002 also noted that this ovule fossil was commonly in physical association with M. scheuchzeri foliage. [7] Specimens of Trigonocarpus can be quite large. The largest recorded was 10 cm and has been noted as the largest ovule produced by a non-angiosperm seed-plant. Some have noted that the large size of these seeds may have allowed them to float, like small coconuts, to be distributed in these coastal mangrove-like areas as well as inland wetland forests. [16] Their three-part symmetry gives them their name. A tube-like opening at the top brought pollen into the ovule. Pachytesta is a term that is also used for this type of seed/ovule. [17]

Male pollen organs

The male pollen organ called Codonotheca. This specimen was identified and named by E. H. Sellards in 1903. Sellards illustration of the specimen is on the right. Codonotheca.jpg
The male pollen organ called Codonotheca. This specimen was identified and named by E. H. Sellards in 1903. Sellards illustration of the specimen is on the right.

As early a 1903, the fossilized male pollen organs called Codonotheca caduca were speculated to be from plants bearing Macroneuropteris scheuchzeri foliage. [18] In 1907, E.H. Sellards further noted this connection. [19] In the late 1960s, both Laveine (1967) [20] and Darrah (1969) reinforced this association. [21] The pollen found in the fossilized Codonotheca caduca are monolete and exceptionally large (200–550 μm).

Possible insect pollination

The large size of the monolete pollen of Macroneuropteris and other seed ferns suggests that they may not have been well adapted to wind dispersal. This raises speculation about the possibility of insect pollination. [22] One of those possibilities is Arthropleura , a very large millipede of the Carboniferous. Scott and Taylor (1983) studied seed-fern pollen on the plates of Arthropleura and thought they might have a role in pollination. [23] W. A. Shear and others have noted that this is very unlikely due to the size of Arthropleura. [24] Other insects of the Carboniferous however may have been pollinators. The cycads, a modern seed plant with some similar affinities to seed ferns, were previously thought to only be pollinated by wind. New studies have confirmed the role of Thrips and other beetles in their pollination. This form of pollination is now known to be present as far back as the Cretaceous. A similar relationship may have occurred between these seed ferns and some Carboniferous insects. [25]

Distribution

Macroneuropteris scheuchzeri is a very recognizable species in the Late Carboniferous, and is found throughout what was known as Euramerica, a large supercontinent that included present-day North America, Europe, and northern Africa. M. macrophylla is found in many of the same locations. Because of their similarities, the two are easily misidentified. M. britannica and M. subauriculata are found mostly in Europe. In general, Macroneuropteris had a worldwide distribution over the tropical equatorial world of the late Carboniferous. [7]

The genus ranges from the Bashkirian stage of the Carboniferous to the early Asselian stage of the Permian. A range that is approximately 18 million years (approximately 298 to 316 million years ago). M. scheuchzeri became particularly common in the Moscovian stage. In two Moscovian-age (approximately 309 mya) fossil locations, Mazon Creek fossil beds in Illinois, U.S.A and Okmulgee in Oklahoma, U.S.A., Macroneuropteris is exceptionally abundant. Along with the leaves of Psaronius , it comprises nearly 60% of the flora in these fossil beds. [10] [26]

It is commonly found in the fossils above coal seams. It has been noted that the coal that formed during an evenly wet climate is dominated by lepidodendrales , and the layer above the coal formed during a transitional and more varied climate is dominated by Macroneuropteris and the tree fern Psaronius . The foliar adaptations described earlier in this article may have given Macroneuropteris an advantage during these transitional times.

As Lepidodendrons declined in the late carboniferous, Macroneuropteris continued to be common and even became a dominant element in these forests. An extinction event called the Carboniferous Rainforest Collapse occurred during the Kasimovian stage. This event decimated many of the Lepidodendrons . It also affected Macroneuropteris, however, the genus was able to recover quicker than other species after this event and became a dominant part of a new forest ecosystems alongside the tree fern Psaronius . [27]

Toward the end of the Carboniferous, the climate of Euramerica became increasingly dryer. Macroneuropteris disappeared from the fossil record for the most part. It was limited to isolated wet areas. It continued into the early Asselian stage of the Permian in these isolated locations. [10]

See also

Related Research Articles

The Carboniferous is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period 358.9 million years ago (Mya), to the beginning of the Permian Period, 298.9 million years ago. The name Carboniferous means "coal-bearing", from the Latin carbō ("coal") and ferō, and refers to the many coal beds formed globally during that time.

<span class="mw-page-title-main">Ginkgoales</span> Order of plants

Ginkgoales are a gymnosperm order containing only one extant species: Ginkgo biloba, the ginkgo tree. It is monotypic, within the class Ginkgoopsida, which itself is monotypic within the division Ginkgophyta. The order includes five families, of which only Ginkgoaceae remains extant.

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms are a group of seed-producing plants that includes conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, literally meaning 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or solitary as in yew, Torreya, Ginkgo. Gymnosperm lifecycles involve alternation of generations. They have a dominant diploid sporophyte phase and a reduced haploid gametophyte phase which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

<span class="mw-page-title-main">Paleobotany</span> Study of organic evolution of plants based on fossils

Paleobotany, which is also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeography), and the evolutionary history of plants, with a bearing upon the evolution of life in general. A synonym is paleophytology. It is a component of paleontology and paleobiology. The prefix palaeo- means "ancient, old", and is derived from the Greek adjective παλαιός, palaios. Paleobotany includes the study of terrestrial plant fossils, as well as the study of prehistoric marine photoautotrophs, such as photosynthetic algae, seaweeds or kelp. A closely related field is palynology, which is the study of fossilized and extant spores and pollen.

<i>Glossopteris</i> Genus of extinct seed ferns

Glossopteris [etymology: from Ancient Greek γλῶσσα + πτερίς ] is the largest and best-known genus of the extinct Permian order of seed ferns known as Glossopteridales. The genus Glossopteris refers only to leaves, within a framework of form genera used in paleobotany. Species of Glossopteris were the dominant trees of the middle- to high-latitude lowland vegetation across the supercontinent Gondwana during the Permian Period. Glossopteris fossils were critical in recognizing former connections between the various fragments of Gondwana: South America, Africa, India, Australia, New Zealand, and Antarctica.

<i>Archaeopteris</i> Extinct genus of Devonian vascular plants

Archaeopteris is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous, the oldest fossils being 385 million years old, and had global distribution.

<span class="mw-page-title-main">Pteridospermatophyta</span> Several distinct polyphyletic groups of extinct seed-bearing plants

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<span class="mw-page-title-main">Bennettitales</span> Extinct order of seed plants

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales are among the most common Mesozoic seed plants, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

The Mazon Creek fossil beds are a conservation lagerstätte found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately 309 million years ago in the mid-Pennsylvanian epoch of the Carboniferous period. These concretions frequently preserve both hard and soft tissues of animal and plant materials, as well as many soft-bodied organisms that do not normally fossilize. The quality, quantity and diversity of fossils in the area, known since the mid-nineteenth century, make the Mazon Creek lagerstätte important to paleontologists, in attempting to reconstruct the paleoecology of the sites. The locality was declared a National Historic Landmark in 1997.

<span class="mw-page-title-main">Coal forest</span> Land type during the late Carboniferous and Permian times

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<i>Psaronius</i> Genus of ferns

Psaronius was a Marattialean tree fern which grew to 10m in height, and is associated with leaves of the organ genus Pecopteris and other extinct tree ferns. Originally, Psaronius was a name for the petrified stems, but today the genus is used for the entire tree fern. Psaronius tree fern fossils are found from the Carboniferous through the Permian.

<i>Pecopteris</i> Extinct genus of ferns

Pecopteris is a very common form genus of leaves. Most Pecopteris leaves and fronds are associated with the marattialean tree fern Psaronius. However, Pecopteris-type foliage also is borne on several filicalean ferns, and at least one seed fern. Pecopteris first appeared in the Devonian period, but flourished in the Carboniferous, especially the Pennsylvanian. Plants bearing these leaves became extinct in the Permian period, due to swamps disappearing and temperatures on Earth dropping.

<span class="mw-page-title-main">Caytoniales</span> Extinct order of Gymnosperms

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<span class="mw-page-title-main">Medullosales</span> Extinct order of Late Carboniferous seed ferns

The Medullosales is an extinct order of pteridospermous seed plants characterised by large ovules with circular cross-section and a vascularised nucellus, complex pollen-organs, stems and rachides with a dissected stele, and frond-like leaves. Their nearest still-living relatives are the cycads.

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The Lyginopteridales were the archetypal pteridosperms: They were the first plant fossils to be described as pteridosperms and, thus, the group on which the concept of pteridosperms was first developed; they are the stratigraphically oldest-known pteridosperms, occurring first in late Devonian strata; and they have the most primitive features, most notably in the structure of their ovules. They probably evolved from a group of Late Devonian progymnosperms known as the Aneurophytales, which had large, compound frond-like leaves. The Lyginopteridales became the most abundant group of pteridosperms during Mississippian times, and included both trees and smaller plants. During early and most of middle Pennsylvanian times the Medullosales took over as the more important of the larger pteridosperms but the Lyginopteridales continued to flourish as climbing (lianescent) and scrambling plants. However, later in Middle Pennsylvanian times the Lyginopteridales went into serious decline, probably being out-competed by the Callistophytales that occupied similar ecological niches but had more sophisticated reproductive strategies. A few species continued into Late Pennsylvanian times, and in Cathaysia and east equatorial Gondwana they persisted into the Late Permian, but subsequently became extinct. Most evidence of the Lyginopteridales suggests that they grew in tropical latitudes of the time, in North America, Europe and China.

<i>Myriacantherpestes</i> Extinct genus of millipedes

Myriacantherpestes is an extinct genus of spiny millipedes from the Pennsylvanian subperiod of the Carboniferous period, known from fossils in Europe and North America.

<span class="mw-page-title-main">Callistophytales</span> Extinct order of plants

The Callistophytales was an order of mainly scrambling and lianescent plants found in the wetland "coal swamps" of Euramerica and Cathaysia. They were characterised by having bilaterally-symmetrical, non-cupulate ovules attached to the underside of pinnules that were morphologically similar to the "normal" vegetative pinnules; and small compound pollen-organs, also borne on the underside of unmodified pinnules, that produced saccate pollen. They were reproductively more sophisticated than most other Palaeozoic pteridosperms, some of which they seem to have out-competed and replaced in the "coal swamp" vegetation during Late Pennsylvanian and Permian times.

<span class="mw-page-title-main">Callistophytaceae</span> Extinct family of seed ferns

The Callistophytaceae was a family of seed ferns (pteridosperms) from the Carboniferous and Permian periods. They first appeared in late Middle Pennsylvanian (Moscovian) times, 306.5–311.7 million years ago (Ma) in the tropical coal forests of Euramerica, and became an important component of Late Pennsylvanian vegetation of clastic soils and some peat soils. The best known callistophyte was documented from Late Pennsylvanian coal ball petrifactions in North America.

<i>Asterotheca</i> Genus of plants

Asterotheca is a genus of seedless, spore-bearing, vascularized ferns dating from the Carboniferous of the Paleozoic to the Triassic of the Mesozoic.

<span class="mw-page-title-main">Paleontology in West Virginia</span>

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References

  1. ↑ Luyd, Edward (1699). Lithophylacii Britannici Ichonographica. London. pp. 144 plus pls. I–XVII.
  2. ↑ Westfall, Richard (1983-04-29). Never at rest" A biography of Isaac Newton . pp.  581. ISBN   978-0521274357.
  3. ↑ Scheuchzer, J. J. (1723). Herbarium diluvianum. Petrus Vander Aa.
  4. ↑ Keferstein, C. (1831). Deutschland geognostisch-geologisch Dargestellt und mit Charten und Durchschmittszeichnungen erlautert. Verlag des Landes-Industrie-Comptoirs.
  5. ↑ Cleal, C. J.; Zodrow, E. L. (1989). "Epidermal structure of some medullosan Neuropteris foliage from the Middle and Upper Carboniferous of Canada and Germany". Palaeontology. 32: 837–882.
  6. ↑ Cleal, C. J.; Zodrow, E. L. (1990). "A revised taxonomy for Palaeozoic neuropterid foliage". Taxon. 39 (3): 486–492. doi:10.2307/1223109. JSTOR   1223109.
  7. 1 2 3 Zodrow, Ervin (2003). "Foliar forms of Macroneuropteris scheuchzeri (Pennsylvanian, Sydney Coalfield, Nova Scotia, Canada)". Atlantic Geology. 39 (1). doi: 10.4138/1047 . ISSN   1718-7885.
  8. ↑ Beeler, H. E. (1983). "Anatomy and frond architecture of Neuropteris ovata and N. scheuchzeri from the Upper Pennsylvanian of the Appalachian Basin". Canadian Journal of Botany. 61 (9): 2352–2368. doi:10.1139/b83-259.
  9. ↑ José A. D'Angelo; Paul C. Lyons; Maria Mastalerz & Erwin L. Zodrow (2013). "Fossil cutin of Macroneuropteris scheuchzeri (Late Pennsylvanian seed fern, Canada)". International Journal of Coal Geology. 105: 137–140. doi:10.1016/j.coal.2012.10.012. ISSN   0166-5162.
  10. 1 2 3 Gregory W. Stull; William A. DiMichele; Howard J. Falcon-Lang; W. John Nelson & Scott Elrick (2012). "Palaeoecology of Macroneuropteris scheuchzeri, and its implications for resolving the paradox of 'xeromorphic' plants in Pennsylvanian wetlands". Palaeogeography, Palaeoclimatology, Palaeoecology. 331: 162–176. Bibcode:2012PPP...331..162S. doi:10.1016/j.palaeo.2012.03.019. ISSN   0031-0182.
  11. ↑ Zodrow, Erwin L.; D'Angelo, José (2014). "Hair-trichomes-files, and spectrochemistry of Macroneuropteris scheuchzeri (Basal Cantabrian, Sydney Coalfield, Canada)". Palaeontographica Abteilung B. Schweizerbart'sche Verlagsbuchhandlung. 290 (4–6): 141–153. doi:10.1127/palb/290/2014/141. S2CID   210635061.
  12. ↑ Zodrow, Erwin L. (2014). "Molecular self-assembly: Hypothesized for "hair" of Macroneuropteris scheuchzeri (Pennsylvanian-age seed-fern)". International Journal of Coal Geology Journal. 121: 14–18. doi:10.1016/j.coal.2013.11.002.
  13. ↑ Stewart, W. N.; Delevoryas, T. (1956). "The medullosan pteridosperms". Botanical Review. 22: 45–80. doi:10.1007/bf02872456. S2CID   31155702.
  14. ↑ Falcon-Lang, Howard (2009). "A Macroneuropteris scheuchzeri tree preserved in growth position in the Middle Pennsylvanian Sydney Mines Formation, Nova Scotia, Canada". Atlantic Geology. 45: 74–80. doi: 10.4138/atlgeol.2009.004 . ISSN   1718-7885.
  15. ↑ Bell, W. A. (1938). Fossil flora of Sydney Coal Field, Nova Scotia. Memoir (Geological Survey of Canada), 215. International Journal of Coal Geology. p. 334.
  16. ↑ DiMichele, William A. (2014). "Wetland-Dryland Vegetational Dynamics In The Pennsylvanian Ice Age Tropics". International Journal of Plant Sciences. Special Issue: Dynamics of Coal-Age Tropical Vegetation. 175 (2): 123–164. doi:10.1086/675235. S2CID   45084239.
  17. ↑ Gastaldo, R. A.; Matten, L. C. (1978). "Trigonocarpus leanus, a new species from the Middle Pennsylvanian of southern Illinois". American Journal of Botany. 65 (8): 882–890. doi:10.2307/2442184. JSTOR   2442184.
  18. ↑ Sellards, E. H. (1903). "Codonotheca, a new type of spore-bearing organ from the coal measures". American Journal of Science. 16 (91): 87–95. Bibcode:1903AmJS...16...87S. doi:10.2475/ajs.s4-16.91.87.
  19. ↑ Sellards, E. H. (1907). "Notes on the spore-bearing organ Codonotheca and its relationship with the Cycadofilices". New Phytologist. 6 (6–7): 175–178. doi: 10.1111/j.1469-8137.1907.tb06055.x .
  20. ↑ Laveine, J. P. (1967). Les Neuroptéridées du Nord de la France. Études Géologiques pour l'Atlas de Topographie Souterraine. Service Géologique des Houillères du Bassin du Nord et du Pas-de-Calais, 1(5):1–344, plus Atlas I to LXXXIV pls.
  21. ↑ Darrah, W. C. (1969). A Critical Review of the Upper Pennsylvanian Floras of Eastern United States with Notes on the Mazon Creek Flora of Illinois. Privately published, Gettysburg, Pennsylvania.
  22. ↑ Taylor, Thomas N; Taylor, Edith L; Krings, Michael (2009). Paleobotany: The biology and evolution of fossil plants. ISBN   978-0-12-373972-8.
  23. ↑ Scott, Andrew C.; Taylor, Thomas N. (1983). "Plant/animal interactions during the upper carboniferous". The Botanical Review. 49 (3): 259–307. doi:10.1007/bf02861089. ISSN   0006-8101. S2CID   34091045.
  24. ↑ Shear, W.A.; Kukalova-Peck, J. (1990). The Ecology of Paleozoic Terrestrial Arthropods: The Fossil Evidence. National Research Council of Canada.
  25. ↑ Peñalver, Enrique; Labandeira, Conrad C.; Barrón, Eduardo; Delclòs, Xavier; Nel, Patricia; Nel, André; Tafforeau, Paul; Soriano, Carmen (2012). "Thrips pollination of Mesozoic gymnosperms". PNAS. 109 (22): 8623–8. Bibcode:2012PNAS..109.8623P. doi: 10.1073/pnas.1120499109 . PMC   3365147 . PMID   22615414.
  26. ↑ Moore, Lillien C.; Wittry, Jack; DiMichele, William A. (2014). "The Okmulgee, Oklahoma fossil flora, a Mazon Creek equivalent: Spatial conservatism in the composition of Middle Pennsylvanian wetland vegetation over 1100 km". Review of Palaeobotany and Palynology. 200: 24–52. doi:10.1016/j.revpalbo.2013.08.002. hdl:10088/21841?show=full.
  27. ↑ Pfefferkorn, H. W.; Gastaldo, R. A.; DiMichele, William A.; Phillips, T.L. (2008). Pennsylvanian Tropical Floras of the United States as a Record of Changing Climate. Geological Society of America Special Paper. Vol. 441. pp. 305–316. doi:10.1130/2008.2441(21). ISBN   978-0-8137-2441-6.
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