"gigantopterid" = an English noun describing large leaves with complex reticulate venation resembling the Cathaysian fossil seed plant genus Gigantopteris and North American genus Delnortea of the Permian Period, 260 million years ago"

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[ Links to Web Pages on Biogeography and Paleobiogeography ]

The Devils Golf Course on the floor of Death Valley is pictured to the right. On the east side of Death Valley near here, sedimentary terrace deposits occur, and water marks are visible on rock cliffs. Terraces and water marks provide clues to size and shoreline dynamics of Pleistocene Lake Manly.

The lowest point in North America known as Badwater, is flanked by the Panamint Mountains to the west and the Amargosa Mountains to the east. Sediments at this site are more than two miles deep according to geologists. The photograph of the Devil's Golf Course by Homer Hobi, captured the unusually high water following a wet spring in 2005.

Charles Smith's Web Page:
This helpful web site contains a bibliography and full-text archive of classical studies of biogeography and evolution, which was compiled by Dr. Smith ... LINK

Chris Scotese's Paleomap Project:
All students of paleobiogeography should immerse themselves in this award-winning site. Christopher Scotese has provided us with continental reconstructions of Pangaea, and animations of the break-up of this supercontinent into continental cratons and island arcs. The site provides information on how to download software, purchase software and other teaching materials, and offers links to other interesting web sites which deal with paleobiogeography ...

Deep Time Maps:
Need paleogeographic maps of western North America for your research? This is where you may access Professor Ron Blakey's most recent maps of the accreting western edge of North America, and view older continental configurations ... LINK

National Geographic Society:
This is a link to the official web site of the National Geographic Society ...

Professor Alistair Rees's University of Arizona Paleointegration Project Website:
Spanning the disciplines of paleoclimatology, paleogeography, paleontology, and geology, Professor Rees has put together this important library resource. Parts of the Paleointegration Project Web Site contain graphics and data from the former PGAP Project at the University of Chicago ...

The International Biogeography Society:
The official web site for The International Biogeography Society is complete with more than 100 links to databases, phylogeny programs, statistical software, teaching aids, and other scientific web pages on biogeography and paleobiogeography. The site is an excellent online resource for research biogeographers, botanists, ecologists, paleontologists, and systematists ... LINK

The Missouri Botanical Garden's Malagasy/Indo-Australo-Malesian Phytogeography Web Page:
Despite the continuous close proximity of Madagascar to Africa since its separation from the continent ca. 165 MYA, the Malagasy flora exhibits a remarkably high affinity with the Indo-Australo-Malesian floras far to the east. Such phytogeographic connections are especially prevalent among eastern humid forest taxa, and represent both ancient vicariance that has resulted in relictual (Cretaceous) Gondwanan disjunctions, as well as continuous dispersal events across the Indian Ocean ... LINK

Commercial and educational software is available to researchers and students including several projections and plate tectonic animations of the late Paleozoic assembly of Pangaea, uplift of the great central Hercynian massif ("Central Pangaean Mountains," Abstract, Heavens et al. 2015, see below), and subsequent split of the Triassic supercontinent into Gondwana and Laurasia (and so-forth). An exquisitely detailed reconstruction of early Permian land masses by Professor Ron Blakey, follows. This illustration, © Ron Blakey, Colorado Plateau Geosystems, was posted here with his written permission.

Student Exercise in ARC-INFO/GIS, Geo-referencing, and Paleobiogeography:

Several books, book chapters, and articles in classical and contemporary scientific literature contain detailed but speculative arguments on the paleobiogeography of stem group flowering plants and possible ancestors (some key references appear below). A late Paleozoic origin of angiosperms has been proposed by Axelrod (1952), among others. While others suggest a Jurassic or early Cretaceous origin of the clade (Friis et al. 2011, among others).

"... we hypothesize that proto-South-East Asia served both as the cradle and centre of diversification of early angiosperms" (page 332, Is the Most Recent Common Ancestor of Angiosperms Now Lost in Proto-southeast Asia Terranes That Are Underwater or Subducted?, Buerki et al. 2014).

Hypotheses suggested by Buerki et al. (2014) contradict classical ideas expressed by Axelrod (1952, 1959) and Krassilov (1997). Dispersal of early angiosperm floras from the tropics poleward was proposed by Axelrod in 1959. A statement quoted from this often overlooked paper is probably pertinent.

"... since angiosperms [Vojnovskyales?] may already have been in existence in upland areas by the Permian [Axelrod 1952], and because the record of the group is exceedingly fragmentary into the Early Cretaceous, the data obviously do not permit us to suggest any one area as the cradle of origin" (page 207, Discussion, Axelrod 1959, the words in [brackets] are mine).

Further, an important study of pollen samples recovered from isolated sedimentary layers in [at least one] continuous stratigraphic sequence in two deep well cores, reports monosulcate, columellate palynomorphs, and Afropollis, from the Middle Triassic (Anisian) about 240 MYA (Hochuli and Feist-Burkhardt 2013). Definitive paleontological evidence published by Peter Hochuli and Susanne Feist-Burkhardt should be read together with a review of Sanmiguelia paleobiology, also reporting three new localities from the Lower Triassic (Norian) Chinle Formation of southwestern North America (Ash and Hasiotis 2013).

Based on Axelrod and Krassilov's cogent arguments and unequivocal palynological evidence published by Hochuli and Feist-Burkhardt (2004, 2013), the Buerki team (2014) should do more homework and rethink their ideas, in my opinion.

Based on controversial research in connection with oil and gas exploration, the Hovey Channel is a southerly bay on the edge of the prominent gulf (left-center of Ron Blakey's paleomap, above) just to the north of the westernmost terminus of the Central Pangaean Mountains at the edge of the Panthalassa Ocean (C. A. Hill 1999).

The well-reasoned "Coastal Hypothesis," which was proposed by Retallack and Dilcher in 1981, could be extended more deeply in geologic time to putative angiosperm stem group seed plant populations of tropical maritime swamps, riparian scrub and woodlands, and semiarid hills bordering the Hovey Channel described above, and reconstructed by Ron Blakey, Ph.D.

When vojnovskyalean elements in Angaran, Cathaysian, and Euramerian floras are considered together with wide-ranging paleopopulations of delnorteas and evolsonias (Mamay et al. 1988, Mamay 1989), among other gigantopteroids (Schachat et al. 2015), potential sites for natural hybridization and intergeneric gene flow were spread across thousands of kilometers.

"Analysis of a wider set of proxies confirms that semi-arid and then arid conditions gradually expanded from west to east across equatorial Pangaea during the latter half of the LPIA" [Late Palaeozoic Ice Age, 305 to 270 MYA] (page 111, 2. The LPIA, 2.2 Glaciation and tropical precipitation during the LPIA: geological evidence and modeling, Heavens et al. 2015).

As the LPIA drew to a close with the demise of Central Pangaean Mountain glaciers, and aridity spread from west to east across early Pangaea, possible new habitats opened opportunities and venues for natural interspecific hybridization in sympatric seed plant populations. Colorful reproductive short- [spur-] shoots of certain species of these seed plants ("coevolutionary compartments") might have attracted pollen-eating and predatory insects with paleobiological consequences.

Were insect and shrub coevolutionary compartments of the late Paleozoic hypoxic icehouse and later hot house, venues of the first angiosperms? Possibly.

A paraphyletic (or polyphyletic) "origin" of stem group flowering plants potentially involving insect- or wind-pollination, natural interspecific hybridization, and paleopolyploidy was possible, but in ancient zones of sympatry from potentially widespread seed plant populations indigenous to coastal and extrabasinal, upland habitats of Pangaea (or pre-Pangaea). This premise eliminates any of the late Mesozoic southwest Pacific Ocean archipelagos, island arcs, spreading centers, or now submerged continental cratons (Buerki et al. 2014) from consideration as "The Cradle of the Flowering Plants" (page 137, Chapter 12, Armen Takhtajan [1969 translation by C. Jeffrey], Flowering Plants Origin and Dispersal, Washington, Smithsonian Institution Press, 310 pp.).

Computer-assisted exercises. As a student drill in ARC-INFO/GIS, geo-referencing, and paleobiogeography, plot the Angaran, Cathaysian, and Euramerian localities where fossils of Permian gigantopteroids and Vojnovskyales have been reported by Mamay et al. (1988) and Naugolnykh (2001), among others. Trace the assembly of Lower Permian continental cratons with "rafting" populations of gigantopteroids and vojnovskyaleans to potential locations on Triassic Pangaea.

Published work reveals that populations of Delnortea abbottiae, Evolsonia texana, Lonesomia mexicana, and Sandrewia texana were spread across a couple thousand kilometers of coastal regions bordering the Hovey Channel of the Panthalassa Sea at the western end of the Central Pangaean Mountains, and to the south of the western tip of the massif where the Artinskian Palmarito Formation was deposited (center left on Ron Blakey's paleomap, above). Plot the source rocks and geo-referenced localities where these fossils have been reported on one or more of the paleomaps obtained or purchased from the sources listed above.

Two other gigantopteroid species, Cathaysiopteris yochelsonii and Zeilleropteris wattii co-occur with so-called "Taeniopteris sp. nov." Taeniopteris sp. [aff. T. multinervis] is the "probable [problematical?] cycadophyte" on page 856, and in Table 1 on page 858 of Schachat et al. (2014) in Permian rock layers. The words [in brackets] are mine. Students should add these geo-referenced sites to the Permian paleomaps constructed as suggested above.

Now apply PAUP or other software of your choice to the sample datasets located on another page of this web site. How do your phylogenetic analyses compare with published work?

Scholars realize that there was never one "source" of an allopolyploid angiosperm "ancestor." Further, a single "origin" of an ancient hybrid flowering plant population was probably unlikely, based on some evidence that species of several possible Permo-triassic flowering plant-antecedents and progenitors including delnorteas, evolsonias, and Vojnovskyales, were spread across thousands of kilometers of Permo-carboniferous terrestrial landscapes.

In reality, scientists might never fully understand paleoecologies of insect- and fungal-plant mutualisms, ancient saltational speciation and coevolution, or past episodes of pollen- and gene-flow in once sympatric seed plant populations, long dead and gone. This begs several questions, among others.

Is the single paleopolyploid event discerned from study of the Amborella genome including an epsilon (ε)- whole genome duplication (WGD), which is depicted as the asterisk in the figured Structured Abstract of Amborella Genome Project (2013), part of the late Paleozoic alpha (α)- swarm of whole genome duplications (WGDs) modeled by Jiao et al. (2011)?

If angiosperms as broadly defined, are fundamentally paraphyletic (and/or polyphyletic), and WGDs (including the γ-triplication) are a result of classic allopolyploidy in paleopopulations of genetically unrelated evolutionary lines, how can a single ancestral Amborella genome be manifest "throughout angiosperm history" (Structured Abstract Discussion, Amborella Genome Project 2013) without genetic input from unrelated, extinct seed plant populations?

"Whereas some authors considered it [Sanmiguelia] as an angiosperm [Brown 1956; Cornet 1986, 1989a] others suggested an attribution to ginkgophytes and rejected a possible relation to angiosperms [Crane 1987, Doyle and Donoghue 1993]" (Discussion-Cretaceous and Pre-cretaceous Records, Hochuli and Feist-Burkhardt 2013).

Surprisingly, Peter Crane (page 779, 1985) once drew a connection between foliar material of Vojnovskya paradoxa and Sanmiguelia lewisii.

How does the paleogeographic distribution of Permo-carboniferous vojnovskyalean seed plant populations match-up with paleomaps based on georeferenced localities of sanmiguelias from the Triassic Chinle Formation of southwestern North America reported by Ash and Hasiotis (2013)?

Based on leaf morphologies and vicariance are Triassic paleopopulations of Sanmiguelia possible descendants of Permo-carboniferous Vojnovskyales?

"... angiosperms could have descended from highlands where they grew for millions of years, perhaps even since the Paleozoic, without leaving fossil traces. The highland hypothesis accords with the Lyellian-Darwinian doctrine of imperfection of the fossil record" (page 109-110, Krassilov 1997).

Do your animated paleomaps fit with Daniel Axelrod's classical ideas on the Permo-triassic origin and early distribution of angiosperms?

"... the absence of Permo-Triassic records of angiosperms is not surprising, but is fully consistent with the thesis that they may have evolved in upland regions of that period" (page 34, Axelrod 1952).

Does palynological evidence published by Hochuli and Feist-Burkhardt (2004, 2013) suggesting that populations of angiosperms occurred in arid, boreal, and tropical environments of the Triassic Period fit with opinions expressed by Axelrod (1959), J. A. Doyle (2012), Buerki et al. (2014), Chaboureau et al. (2014), and others (references appear below)?

"... Allopatric speciation and radiation, usually enhanced by orogeny processes and relief creation, rather involves here the fragmentation of the supercontinent into many small land masses ... " (page 14068, Introduction: Global Climate and Angiosperm Expansion and Diversification, Chaboureau et al. 2014).

Does APG IV have any bearing on the origin and early dispersal of flowering plants, including ANA grade angiosperms, as suggested by J. A. Doyle (2012) and Buerki et al. (2014)? No.

"... The congruence of the stratigraphic sequence of pollen types and their presumed evolutionary sequence would make no sense if angiosperms had already diversified in a hidden homeland area; there is no reason to expect that groups would migrate into better-known areas in the order in which they evolved much earlier ... " (page 303, Status of the Problem Prior to Molecular Systematics, J. A. Doyle 2012).

Taking into account a possible "evolutionary sequence" (chronocline) of leaf morphologies seen in Permo-carboniferous Vojnovskyales and Triassic sanmiguelias, and a "stratigraphic sequence of pollen types," which is evident from palynological evidence published by Hochuli and Feist-Burkhardt (2004, 2013), is the preceding statement accurate or precise?

Together with animations and/or graphics produced by application of commercial and/or educational software (see above), what can your team conclude on the phylogeography of paraphyletic clades of gymnosperms (including delnorteas, evolsonias, and sanmiguelias, among other seed plant lineages) discerned from computer-assisted analyses?

Tropical maritime swamps, riparian scrub and woodlands, and dry tropical shrublands and glades and rocky outcrops in walchian coniferous forests of the Central Pangaean Mountains bordering the Hovey Channel might offer convenient paleogeographic settings for J. A. Doyle's "hidden homeland area" where the ancestors of flowering plants including delnorteas, evolsonias, and sandrewias, "had already diversified." Daniel Axelrod proposed a similar idea in 1952.

Finally, C. S. P. Foster et al. (2017) conclude that, "using analyses of near-complete chloroplast genomes, we have estimated that crown group Angiospermae arose 221 Ma (251-192 Ma) in the mid-Triassic."

Background Reading:

Amborella Genome Project. 2013. The Amborella genome and the evolution of flowering plants. Science 342(6165): 1467.

APG IV. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20.

Ash, S. R. and S. T. Hasiotis. 2013. New occurrences of the controversial late Triassic plant fossil Sanmiguelia Brown and associated ichnofossils in the Chinle Formation of Arizona and Utah, USA. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 268(1): 65-82.

Axelrod, D. I. 1952. A theory of angiosperm evolution. Evolution 6(1): 29-60.

Axelrod, D. I. 1959. Poleward migration of early angiosperm flora - angiosperms only displaced the relict Jurassic-type flora at high latitudes in late Cretaceous time. Science 130(3369): 203-207.

Buerki, S., F. Forest, and N. Alvarez. 2014. Proto-southeast Asia as a trigger of early angiosperm diversification. Botanical Journal of the Linnean Society 174(3): 326333.

Buys, J., C. Spandler, R. J. Holm, and S. W. Richards. 2014. Remnants of ancient Australia in Vanuatu: Implications for crustal evolution in island arcs and tectonic development of the southwest Pacific. Geology 42: 939-942.

Chaboureau, A.-C., P. Sepulchre, Y. Donnadieu, and A. Franc. 2014. Tectonic-driven climate change and the diversification of angiosperms. Proceedings of the National Academy of Sciences 111(39): 1406614070.

Chaloner, W. G. and S. V. Meyen. 1973. Carboniferous and Permian floras of the northern continents. Pp. 169-186 In: A. Hallam (ed.), Atlas of Palaeobiogeography. Amsterdam: Elsevier Scientific Publishing Company, 531 pp.

Crane, P. R. 1985. Phylogenetic analysis of seed plants and the origin of angiosperms. Annals of the Missouri Botanic Garden 72: 716-793.

DiMichele, W. A. and R. W. Hook. 1992. 5. Paleozoic terrestrial ecosystems. Pp. 205-325 In: A. K. Behrensmeyer, J. D. Damuth, W. A. DiMichele, R. Potts, H.-D. Sues, and S. L. Wing, (eds.), Terrestrial Ecosystems Through Time, Evolutionary Paleoecology of Terrestrial Plants and Animals. Chicago: University of Chicago Press.

Doyle, J. A. 2012. Molecular and fossil evidence on the origin of angiosperms. Annual Review of Earth and Planetary Sciences 40: 301326.

Foster, C. S. P., H. Sauquet, M. van der Merwe, H. McPherson, M. Rossetto, and S. Y. W. Ho. 2017. Phylogenomic timescale for angiosperm evolution: evaluating the impact of genomic data and priors on Bayesian estimates of the angiosperm evolutionary timescale. Systematic Biology XX: XXX-XXX.

Friis, E. M., P. R. Crane, and K. R. Pedersen. 2011. Early Flowers and Angiosperm Evolution. Cambridge: Cambridge University Press, 596 pp.

Haugh, B. N. 1981. Discussion [of Melville 1981]. Pp. 275-286 In: G. Nelson and D. E. Rosen (eds.), Vicariance Biogeography: A Critique. New York: Columbia University Press, 593 pp.

Heavens, N. G., N. M. Mahowald, G. S. Soreghan, M. J. Soreghan, and C. A. Shields. 2015. A model-based evaluation of tropical climate in Pangaea during the late Palaeozoic icehouse. Palaeogeography, Palaeoclimatology, Palaeoecology 425: 109127.

Hill, C. A. 1999. Reevaluation of the Hovey Channel in the Delaware Basin, West Texas American Association of Petroleum Geologists Bulletin 83(2): 277-294.

Hochuli, P. A. and S. Feist-Burkhardt. 2004. A boreal early cradle of angiosperms? Angiosperm-like pollen from the Middle Triassic of the Barents Sea (Norway). Journal of Micropalaeontology 23: 97-104.

Hochuli, P. A. and S. Feist-Burkhardt. 2013. Angiosperm-like pollen and Afropollis from the Middle Triassic (Anisian) of the Germanic Basin (Northern Switzerland). Frontiers in Plant Science, Plant Evolution and Development 4: Article 344.

Jiao, Y., N. L. Wickett, S. Ayyampalayam, A. S. Chanderbali, L. Landherr, P. E. Ralph, L. P. Tomsho, Y. Hu, H. Liang, P. S. Soltis, D. E. Soltis, S. W. Clifton, S. E. Schlarbaum, S. C. Schuster, H. Ma, J. Leebens-Mack, and C. W. dePamphilis. 2011. Ancestral polyploidy in seed plants and angiosperms. Nature 473(7345): 97-100.

Krassilov, V. A. 1997. Angiosperm Origins: Morphological and Ecological Aspects. Sofia: Pensoft, 270 pp.

Kroenke, L. W. 1996. Plate tectonic development of the western and southwestern Pacific: Mesozoic to the present. Pp. 19-34, In: A. Keast and S. E. Miller (eds.), The Origin and Evolution of Pacific Islands Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. Amsterdam: SPB Academic Publishing.

Mamay, S. H. 1989. Evolsonia, a new genus of Gigantopteridaceae from the Lower Permian Vale Formation, North-central Texas.  American Journal of Botany 76(9): 1299-1311.

Mamay, S. H., J. M. Miller, D. M. Rohr, and W. E. Stein, Jr. 1988. Foliar morphology and anatomy of the gigantopterid plant Delnortea abbottiae from the Lower Permian of West Texas.  American Journal of Botany 75(9): 1409-1433.

Melville, R. 1981. Vicarious plant distributions and paleogeography of the Pacific Region. Pp. 238-274 In: G. Nelson and D. E. Rosen (eds.), Vicariance Biogeography: A Critique. New York: Columbia University Press, 593 pp.

Miller, J. M. 1989. The archaic flowering plant family Degeneriaceae: its bearing on an old enigma. National Geographic Research 5(2): 218-231.

Morat, Ph., J.-M. Veillon, and H. S. MacKee. 1986. Floristic relationships of New Caledonian rainforest phanerogams. Telopea 2(6): 631-679.

Morley, R. J. 2001. Why are there so many primitive angiosperms in the rain forests of Asia-Australasia. Pp. 185-200 In: I. Metcalfe, J. M. B. Smith, M. Morwood, and I. Davidson (eds.), Faunal and Floral Migrations and Evolution in SE Asia-Australasia. Lisse: Swets and Zeitlinger, 419 pp.

Naugolnykh, S. V. 2001. Morphology and systematics of representatives of Vojnovskyales. Paleontologicheskii Zhurnal 35(5): 545-556.

Raven, P. H. and D. I. Axelrod. 1974 (1981 republication). Angiosperm biogeography and past continental movements. Annals Missouri Botanical Garden 61: 539-673.

Recy, J. and J. Dupont. 1982. Le sud-quest du Pacifique: Données structurales, carte à 1:12,000,000 à l'équateur, Notice Explicative N° 97. Paris: ORSTOM.

Retallack, G. J. and D. L. Dilcher. 1981. 2. A coastal hypothesis for the dispersal and rise to dominance of flowering plants. Pp. 27-77 In: K. J. Niklas (ed.) Paleobotany, Paleoecology, and Evolution, Volume 2. New York: Praeger, 279 pp.

Ricardi, F., O. Rösler, and O. Odreman. 1999. Delnortea taphoflora (Gigantopteridaceae) of Loma de San Juan (Palmarito Formation, NW of Venezuela) and its palaeophytogeographical relationships in the Artinskian (Neopaleozoic).  Plantula 2(1-2): 73-86.

Ricardi-Branco, F. 2008. Venezuelan paleoflora of the Pennsylvanian-early Permian: paleobiogeographical relationships to central and western equatorial Pangaea. Gondwana Research 14(3): 297-305.

Rodda, P. and L. W. Kroenke. 1984. Chapter 7. Fiji: A Fragment Arc. Pp. 87-109 In: L. W. Kroenke, Cenozoic Tectonic Development of the Southwest Pacific. United Nations ESCAP, CCOP/SOPAC Technical Bulletin No. 6, 122 pp.

Schachat, S. R., C. C. Labandeira, and D. S. Chaney. 2015. Insect herbivory from early Permian Mitchell Creek Flats of north-central Texas: opportunism in a balanced component community. Palaeogeography, Palaeoclimatology, and Palaeoecology 440(3-4): 830-847.

Schachat, S. R., C. C. Labandeira, J. Gordon, D. Chaney, S. Levi, M. N. Halthore, and J. Alvarez. 2014. Plant-insect interactions from early Permian (Kungurian) Colwell Creek Pond, north-central Texas: the early spread of herbivory in riparian environments. International Journal of Plant Sciences 175(8): 855-890.


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