Skip to main content Accessibility help

The “seeds” on Padgettia readi are insect galls: reassignment of the plant to Odontopteris, the gall to Ovofoligallites n. gen., and the evolutionary implications thereof

  • Gregory W. Stull (a1), Conrad C. Labandeira (a2) (a3), William A. Dimichele (a2) and Dan S. Chaney (a2)


The Early Permian (Asselian) Euramerican plant Padgettia readi Mamay is reassigned to Odontopteris Brongniart, as O. readi (Mamay) Stull et al. n. comb. Distinctive elongate structures on neuropteroid pinnules of this plant, previously interpreted as fructifications, are herein reinterpreted as foliar histoid galls, structurally analogous to blister or vein galls, and probably induced by an early lineage of hemipterans or mites. These distinctive features are assigned to the new gall ichnogenus Ovofoligallites Labandeira, n. ichnogen. n. ichnosp, as O. padgetti Labandeira. The Early Permian association between an Odontopteris host and Ovofoligallites gallers probably originated during the Middle Pennsylvanian as a similar, antecedent association between Macroneuropteris scheuchzeri (Hoffmann) Cleal, Shute, and Zodrow and the maker of U-shaped surface features long known as a distinctive, unattributed damage type, but now recognizable as a likely gall. The persistence of this association between the galler and certain medullosan pteridosperms into the Permian adds to the morphological richness of the Permian galler insect fauna. The Permian ecological expansion of galling insects resulted in colonization of new host plants, primarily through a shift from the consumption of entire, mostly pteridophyte axial organs during the Pennsylvanian to the partitioning of seed plant tissues in leaves and small branches in the Permian. The Ovofoligallites galler was part of a diverse Permian galler guild involving a variety of plant taxa, organs and tissues that overwhelmingly targeted multiple lineages of seed plants.



Hide All
Aoki, S. and Moran, N. A. 1994. Pemphigus obesinymphae, a new American aphid species with defenders and swollen nymphs (Homoptera: Aphidoidea: Pemphigidae). Journal of the New York Entomological Society, 102:251260.
Ash, S. 1997. Evidence of arthropod–plant interactions in the Upper Triassic of the southwestern United States. Lethaia, 29:237248.
Barthel, M. and Amelang, A. 2011. Der Farnsamer Odontopteris schlotheimii Brongniart aus der Manebach-Formation des Thüringer Wald-Beckens. Semana, 26:1324.
Baxter, R. W. 1978. Sphenopteris sellardsi, a problematical pteridosperm from the Permian of Texas. University of Kansas Paleontological Contributions, 94:17.
Banerjee, J. 2004. Evidence of insect–plant interactions from the Upper Gondwana Sequence (Lower Cretaceous) in the Rajmahal Basin, India. Gondwana Research, 7:205210.
Banerjee, M. and Bera, S. 1998. Record of zoocecidia on leaves of Glossopteris browniana Brong from Mohuda Basin, Upper Permian, Indian Lower Gondwana. Indian Biologist, 30:5861.
Beck, A. L. and Labandeira, C. C. 1998. Early Permian insect folivory on a gigantopterid-dominated riparian flora from north-central Texas. Palaeogeography, Palaeoclimatology, Palaeoecology, 142:139173.
Berry, E. W. 1923. Pathological conditions among fossil plants, p. 99114. InMoodie, R. L.(ed.), Paleopathology: An Introduction to the Study of the Ancient Evidences of Disease. University of Illinois Press, Urbana.
Bertling, M., Braddy, S. J., Bromley, R. G., Demathieu, G. R., Genise, J., Mikuláš, R., Nielsen, J. K., Nielsen, K. S. S., Rindsberg, A. K., Schlirf, M., and Uchman, A. 2006. Names for trace fossils: A uniform approach. Lethaia, 39:265286.
Blagoderov, V. A. and Grimaldi, D. A. 2004. Fossil Sciaroidea in Cretaceous ambers, exclusive of Cecidomyiidae, Sciaridae, and Keroplatidae. American Museum Novitates, 3433:176.
Broutin, J., Doubinger, J., Farjanel, G., Freytet, P., Kerp, H., Langiaux, J., Lebreton, M.-L., Sebban, S., and Satta, S. 1990. Le renouvellement des flores au passage Carbonifère Permien: Approches stratigraphique, biologique, sédimentologique. Comptes Rendus de Academie des Sciences, Paris, 311:15631569.
Castro, M. P. 2005. La flora estefaniense B de La Magdalena (León, España), un referente europeo. Tomo II: Descripción sistemática de las Gimnospermas. Instituto Geológico Minero de España, Cuadernos del Museo Geominero, 4:1229.
Cleal, C. J., Shute, C. H., and Zodrow, E. L. 1990. A revised taxonomy for Paleozoic neuropterid foliage. Taxon, 39:486492.
Cridland, A. A. and Morris, J. E. 1960. Spermopteris, a new genus of pteridosperms from the Upper Pennsylvanian series of Kansas. American Journal of Botany, 47:855859.
Darrah, W. C. 1969. A critical review of upper Pennsylvanian floras of eastern United States with notes on the Mazon Creek flora of Illinois. Privately published, 220p.
Diéguez, C., Nieves-Aldrey, J. L., and Barrón, E. 1996. Fossil galls (zoocecids) from the upper Miocene of La Cerdaña (Lérida, Spain). Review of Palaeobotany and Palynology, 94:329342.
Dimichele, W. A. and Aronson, R. B. 1992. The Pennsylvanian–Permian vegetational transition: a terrestrial analogue to the onshore–offshore hypothesis. Evolution, 46:807824.
Dimichele, W. A., Montañez, I. P., Poulsen, C. J., and Tabor, N. J. 2009. Climate and vegetational regime shifts in the late Paleozoic ice age earth. Geobiology, 7:200226.
Dimichele, W. A., Cecil, C. B., Montañez, I. P., and Falcon-Lang, H. J. 2010. Cyclic changes in Pennsylvanian paleoclimate and effects on floristic dynamics in tropical Pangaea. International Journal of Coal Geology, 83:329344.
Doubinger, J., Vetter, P., Langiaux, J., Galtier, J., and Broutin, J. 1995. La flore fossile du bassin houiller de Saint-Étienne. Mémoires du Muséum National d'Histoire Naturelle, 164:1357.
Erwin, D. and Schick, K. N. 2007. New Miocene oak galls (Cynipini) and their bearing on the history of cynipid wasps in western North America. Journal of Paleontology, 81:561580.
Falcon-Lang, H. J. and Dimichele, W. A. 2010. What happened to the coal forests during Pennsylvanian glacial phases? Palaios, 25:611617.
Falcon-Lang, H. J., Nelson, J., Elrick, S., Looy, C., Ames, P., and Dimichele, W. A. 2010. Incised valley-fills containing conifers imply that seasonally-dry vegetation dominated Pennsylvanian tropic lowlands. Geology, 37:923926.
Fernandes, G. W. and Price, P. W. 1988. Biogeographical gradients in galling species richness: tests of hypotheses. Oecologia, 76:161167.
Fielding, C. R., Frank, T. D., Birgenheier, L. P., Rygel, M. C., Jones, A. T., and Roberts, J. 2008. Stratigraphic record and facies associations of the late Paleozoic ice age in eastern Australia (New South Wales and Queensland), p. 4157. InFielding, C. R., Frank, T. D., and Isbell, J. L.(eds.), Resolving the Late Paleozoic Ice Age in Time and Space. Geological Society of America Special Paper 441.
Florin, R. 1945. Die Koniferen des Obercarbons und des unteren Perms. Palaeontographica, Abt. B (Paläophytologie), 85:162, pls. 1–30.
Florin, R. 1950. Upper Carboniferous and lower Permian conifers. Botanical Review, 16:258282.
Gagné, R. J. 1994. The Gall Midges of the Neotropical Region. Cornell University Press, Ithaca, New York.
Grauvogel-Stamm, L. 1978. La flore du Grès-a-Voltzia (Buntsandstein Supérieur) des Vosges du Nord (France). Morphologie, anatomie, interprétations phylogénique et paléogéographique. Sciences et Géologie, 50:1225.
Grauvogel-Stamm, L. and Kelber, K.-P. 1996. Plant-insect interactions and coevolution during the Triassic in western Europe. Paleontologia Lombarda (N.S.), 5:523.
Guleva, J. S. and Raman, A. 2007. Age of plant galls. Current Science 93:598.
Harris, T. M. 1942. Wonnacottia, a new bennettitalean microsporophyll. Annals of Botany, 4:577592.
Harris, T. M. 1969. The Yorkshire Jurassic Flora. III. Bennettitales. British Museum (Natural History), London.
Hentz, T. F. 1988. Lithostratigraphy and paleoenvironments of upper Paleozoic continental red beds, North-Central Texas: Bowie (new) and Wichita (revised) Groups: The University of Texas at Austin, Bureau of Economic Geology Report of Investigations No. 170, 55p.
Johnson, W. T. and Lyon, H. H. 1991. Insects that Feed on Trees and Shrubs, second edition. Cornell University Press, Ithaca, New York.
Krantz, G. W. and Lindquist, E. E. 1979. Evolution of phytophagous mites (Acari). Annual Review of Entomology, 24:121158.
Krassilov, V. 2008. Mine and gall predation as top down regulation in the plant–insect systems from the Cretaceous of Negev, Israel. Palaeogeography, Palaeoclimatology, Palaeoecology, 261:261269.
Krassilov, V. and Karasev, E. 2005. Paleofloristic evidence of climate change near and beyond the Permian–Triassic boundary. Palaeogeogrpahy, Palaeoclimatology, Palaeogeography, 284:326336.
Krassilov, V. and Karasev, E. 2008. First evidence of plant–arthropod interaction at the Permian–Triassic boundary in the Volga Basin, European Russia. Alavesia, 2:247252.
Krassilov, V. and Rasnitsyn, A. 2008. Plant–Arthropod Interactions in the Early Angiosperm History: Evidence from the Cretaceous of Israel. Pensoft, Sofia.
Küster, E. 1911. Die Gallen der Pflanzen. Hirzel, Leipzig.
Labandeira, C. C. 1998. Early history of arthropod and vascular plant associations. Annual Review of Earth and Planetary Sciences, 26:329377.
Labandeira, C. C. 2002. The history of associations between plants and animals, p. 26–74, 248261. InHerrera, C. and Pellmyr, O., (eds.), Plant–Animal Interactions: An Evolutionary Approach. Blackwell Science, Oxford, U.K.
Labandeira, C. C. 2006a. The four phases of plant–arthropod associations in deep time. Geologica Acta, 4:409438.
Labandeira, C. C. 2006b. Silurian to Triassic plant and insect clades and their associations: new data, a review, and interpretations. Arthropod Systematics and Phylogeny, 64:5394.
Labandeira, C. C. 2011. Evidence for an earliest Late Carboniferous divergence time and the early larval ecology and diversification of major Holometabola lineages. Entomologica Americana, 117:921.
Labandeira, C. C. and Allen, E. M. 2007. Minimal insect herbivory for the Lower Permian Coprolite Bone Bed site of north-central Texas, U.S.A., and comparison to other late Paleozoic floras. Palaeogeography, Palaeoclimatology, Palaeoecology, 247:197219.
Labandeira, C. C. and Beall, B. S. 1990. Arthropod terrestriality, p. 214256. InMikulic, D.(ed.), Arthropods: Notes for a Short Course. Short Courses in Paleontology, Vol. 3. Paleontological Society, Knoxville, Tennessee.
Labandeira, C. C. and Phillips, T. L. 1996. A Carboniferous petiole gall: insight into early ecologic history of the Holometabola. Proceedings of the National Academy of Sciences U.S.A., 93:84708474.
Labandeira, C. C. and Phillips, T. L. 2002. Stem borings and petiole galls from Pennsylvanian tree ferns of Illinois, U.S.A.: implications for the origin of the borer and galling functional-feeding-groups and holometabolous insects. Palaeontographica (A), 264:184, pls. 1–16.
Labandeira, C. C., Wilf, P., Johnson, K. R., and Marsh, F. 2007. Guide to Insect (and Other) Damage Types on Compressed Plant Fossils (Version 3.0–Spring 2007). Smithsonian Institution, Washington, D.C.,
Larew, H. 1992. Fossil galls, p. 5059. InShorthouse, J. D. and Rohfritsch, O.(eds.), Biology of Insect-Induced Galls. Oxford University Press, Oxford, U.K.
Laveine, J. P. and Behlis, A. 2007. Frond architecture of the seed-fern Macroneuropteris scheuchzeri, based on Pennsylvanian specimens from the Northern France coal field. Palaeontographica, Abt. B (Paläophytologie), 277:141.
Laveine, J. P. and Delbeque, S. 2011. The bifurcate “outer” semi-pinnate frond of the seed-fern Barthelopteris germarii, based on Late Pennsylvanian specimens from the Blanzy-Montceau Basin, Massif Central, France. Palaeontographica, Abt. B (Paläophytologie), 287:155.
Laveine, J. P. and Legrand, L. 2008. The degree of foliar-laminate segmentation of the Pennsylvanian seed-fern Macroneuropteris scheuchzeri. Revue de Paléobiologie, 27:475510.
Lesquereux, L. 1892. The flora of the Dakota Group. United States Geological Survey Monograph, 17:19400.
Mädler, A. K. 1936. Eine Blattgall an einem vorweltlichen Pappel-Blatt. Natur und Volk, 66:271274.
Maia, V. C. and Fernandes, G. W. 2004. Insect galls from Serra de São José (Tiradentes, MG, Brasil). Brazilian Journal of Biology, 64:423445.
Mamay, S. H. 1960. Padgettia, a new genus based on fertile neuropteroid foliage from the Permian of Texas. The Palaeobotanist, 9:5357.
Mapes, G. and Gastaldo, R. A. 1989. Upper Paleozoic non-peat accumulating floras, p. 115127. InBroadhead, T. W.(ed.), Fossil Land Plants: Notes for a Short Course. Studies in Geology, vol. 15. University of Tennessee, Nashville.
Meyer, J. 1987. Plant Galls and Gall Inducers. Gebrüder Borntraeger, Stuttgart.
Müller, A. H. 1982. Über Hyponome fossiler und rezenter Insekten, erster Beitrag. Freiberger Forschungshefte C, 366:727.
Nyman, T., Widmer, A., and Roininen, H. 2000. Evolution of gall morphology and host-plant relationships in willow-feeding sawflies. Evolution, 54:526533.
Pant, D. D. and Srivastava, P. C. 1995. Lower Gondwana insect remains and evidences of insect–plant interaction, p. 317326. InSurange, K. R., Bose, M. D., and Khare, P. K., (eds.), Proceedings of the International Conference on Global Environment and Diversification through Geological Time. Society of Indian Plant Taxonomists, Allahabad, India.
Pfefferkorn, H. 1980. A note on the term “upland flora.” Review of Palaeobotany and Palynology, 30:157158.
Plotnick, R. E., Kenig, F., Scott, A. C., Glasspool, I. J., Eble, C. F., and Lang, W. J. 2009. Pennsylvanian paleokarst and fills from northern Illinois, U.S.A.: a window into Late Carboniferous environments and landscapes. Palaios, 24:627637.
Plumb, G. H. 1953. The formation and development of the Norway spruce gall caused by Aldeges abietis L. Connecticut Agricultural Experiment Station Bulletin, 566:177.
Potonié, H. 1893. Die Flora des Rothliegenden von Thüringen. Abhandlungen der Königlich Preussischen geologischen Landesansalt, N.F., 2:1298.
Prevec, R., Labandeira, C. C., Neveling, J., Gastaldo, R. A., Looy, C., and Bamford, M. A. 2009. A portrait of a Gondwanan ecosystem: a new Late Permian locality form KwaZulu–Natal, South Africa. Review of Palaeobotany and Palynology, 156:454493.
Rothwell, G., Grauvogel-Stamm, L., and Mapes, G. 2000. An herbaceous fossil conifer: gymnospermous ruderals in the evolution of Mesozoic vegetation. Palaeogeography, Palaeoclimatology, Palaeoecology, 156:139145.
Russo, R. A. 2006. Field Guide to Plant Galls of California and other Western States. University of California Press, Berkeley and Los Angeles.
Sellards, E. H. 1908. Fossil plants of the Upper Paleozoic of Kansas. University Geological Survey of Kansas, 9:386499.
Scott, A. C., Stephenson, J., and Collinson, M. E. 1994. The fossil record of leaves with galls, p. 447470. InWilliams, M. A. J.(ed.), Plant Galls. Systematics Association Special Volume, no. 49. Oxford University Press, Oxford, U.K.
Shcherbakov, D. E. 2000. Permian faunas of Homoptera (Hemiptera) in relation to phytogeography and the Permo–Triassic Crisis. Paleontological Journal, Supplement 3, 34:S251S262.
Shcherbakov, D. E. 2008. On Permian and Triassic insect faunas in relation to biogeography and the Permian–Triassic boundary in the Volga Basin, European Russia. Paleontological Journal, 42:1531.
Stireman, J. O. III, Devlin, H., Carr, T. G., and Abbot, P. 2009. Evolutionary diversification of the gall midge genus Asteromyia (Cecidomyiidae) in a multitrophic ecological context. Molecular Phylogenetics and Evolution, 54:194210.
Thomas, B. A. 1969. A new British Carboniferous calamite cone, Paracalamostachys spadiciformis. Palaeontology, 12:253261.
Tubbs, P. K. 2003. Comment on the proposal to emend the Code with respect to trace fossils. Bulletin of Zoological Nomenclature, 60:141142.
Van Amerom, H. W. J. 1973. Gibt es Cecidien in Karbon bei Calamitien und Asterophylliten?, p. 6383. InJosten, K.-H.(ed.), Compte Rendus Septième Congrès International de Stratigraphie et de Geologie du Carbonifère. Van Acken, Krefeld, Germany.
Vasilenko, D. V. 2005. Damages on Mesozoic plants from the Transbaikalian locality Chernovskie Kopi. Paleontological Journal, 39:628633.
Vasilenko, D. V. 2007. Feeding damage on Upper Permian plants from the Sukhona River. Paleontological Journal, 41:207211.
Waggoner, B. M. 1999. Fossil oak galls from the Stinking Water paleoflora of Oregon (early Miocene). PaleoBios, 19:814.
Waggoner, B. M. and Poteet, M. F. 1996. Unusual leaf galls from the middle Miocene of northwestern Nevada. Journal of Paleontology, 70:10801084.
Wagner, R. H. and Castro, M. P. 1998. Neuropteris obtuse, a rare but widespread Late Carboniferous pteridosperm. Palaeontology, 41:122.
Weiss, C. E. 1876. Steinkohlen-Calimarien, mit besonderer Berücksichtigung ihrer Fructificationen. Neumann'schen Kartenhandlung, Berlin.
Yukawa, J. 1978. New midge galls from Kyushu. Memoirs of the Faculty of Agriculture of Kagoshima University, 14:93101.


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed