Glossopteris

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Glossopteris
Temporal range: Permian
Pre

O
S
D
C
P
T
J
K
Pg
N

Glossopteris sp.

Scientific classification
Kingdo
Plantae
m:
Divisio Pteridospermat
n:
ophyta
Order: Arberiales
Glossopteridace
Family:
ae
Genus: Glossopteris
Species

G. angustifolia

G. brasiliensis

G. browniana

G. communis

G. indica

G. occidentalis

Fossils of the gymnosperm

Glossopteris (dark green)
found in all of the southern
continents provide strong
evidence that the continents
were once amalgamated into
a supercontinent Gondwana

Glossopteris (Ancient Greek: glossa, meaning "tongue", because the leaves were
tongue-shaped) is the largest and best-known genus of the extinct order of seed ferns known as
Glossopteridales (or in some cases as Arberiales or Dictyopteridiales).

Contents

1 History

2 Taxonomy

3 Biology

4 Outcrops in Brazil

5 References

6 Sources

7 External links

History
The Glossopteridales arose in the Southern Hemisphere around the beginning of the Permian
Period (298.9 million years ago).[1] Their distribution across several, now detached, landmasses
led Eduard Suess, amongst others, to propose that the southern continents were once
amalgamated into a single supercontinentPangea.[2] These plants went on to become the
dominant elements of the southern flora through the rest of the Permian but disappeared in
almost all places at the end of the Permian (252.17 million years ago).[3][4][5] The only convincing
Triassic records are very earliest Triassic leaves from Nidpur, India,[6] but even these records are
somewhat questionable owing to faulting and complex juxtapositioning of Permian and Triassic

strata at Nidpur. Although most modern palaeobotany textbooks cite the continuation of
glossopterids into later parts of the Triassic and, in some cases into the Jurassic, these ranges are
erroneous and are based on misidentification of morphologically similar leaves such as
Gontriglossa,[7] Sagenopteris, or Mexiglossa.[8] Glossopterids were, thus, one of the major
casualties of the end-Permian mass extinction event.[9]

Location of Glossopteris remains shown in green in the former Supercontinent Gondwana.

More than 70 fossil species of this genus have been recognized in India alone,[10] with additional
species from South America, Australia,[11][12] Africa, Madagascar[13] and Antarctica.[14] Essentially,
Glossopteris was restricted to the middle- and high-latitude parts of Gondwana during the
Permian [15] and was an important contributor to the vast Permian coal deposits of the Southern
Hemisphere continents.[16] Most northern parts of South America and Africa lack Glossopteris
and its associated organs. However, in recent years a few disparate localities in Morocco, Oman,
Anatolia, the western part of the island of New Guinea, Thailand and Laos have yielded fossils
that are of possible glossopterid affinity.[17] These peri-gondwanan records commonly occur
together with Cathaysian or Euramerican plant speciesthe assemblages representing a zone of
mixing between the strongly provincial floras of the Permian.[18] Apart from those in India and
the peri-gondwanan localities, a few other fossils from the Northern Hemisphere have been
assigned to this group, but these are not identified with great certainty. For example, specimens
assigned to Glossopteris from the far east of Russia in the 1960s are more likely to be
misdentifications of other gymnosperms such as Pursongia.[19] Confident assignment of fossil
leaves to Glossopteris normally requires their co-preservation with the distinctive segmented
roots of this group (called Vertebraria) or with the distinctive fertile organs.[20]

Taxonomy
Long considered a fern after its discovery in the 1820s,[21] it was later assigned to the
gymnosperms. The genus is placed in the division Pteridospermatophyta. In reality, many of the
plant groups included within this division are only distantly related to one another. Glossopterids'
relationships with other groups remain obscure. Most recent phylogenetic analyses favour
placement of glossopterids as sister to a large group including Corystospermales, Caytoniales,
Bennettitales, Pentoxylales, Gnetales (in some analyses), and angiosperms.[22] A few analyses
favour alternative links with Ginkgoales, Cordaitales and Pinales.

Glossopteris should strictly be used to refer to the distinctive spathulate fossil leaves with
reticulate venation, however, the term has also been used to refer to the parent plant as a whole.
[23]

Biology

Glossopteris browniana fossil in the Artis zoo, Amsterdam.

Glossopteris was a woody, seed-bearing shrub or tree, some apparently reaching 30 metres
(98 ft) tall. They had a softwood interior that resembles conifers of the family Araucariaceae.[24]
Seeds were borne on one side of variably branched or fused structures,[25][26][27][28][29][30] and
microsporangia containing pollen were borne in clusters at the tips of slender filaments.[31] Both
the seed- and pollen-bearing organs were partially fused (adnate) to the leaves, or, in some cases,
possibly positioned in the axils of leaves. The homologies of the flattened seed-bearing structures
have remained particularly controversial with some arguing that the fertile organs represent
megasporophylls (fertile leaves) whereas others have interpreted the structures as flattened, seedbearing, axillary axes (cladodes). It is unclear whether glossopterids were monoecious or
dioecious.
They are interpreted to have grown in very wet soil conditions,[32][33] similar to the modern Bald
Cypress. The leaves ranged from about 2 cm to over 30 cm in length.
The profile of glossopterid trees is largely speculative as complete trees have not been preserved.
However, based on analogies with modern high-latitude plants Glossopteris trees probably
tapered upwards like a Christmas tree and were relatively widely spaced to take advantage of the
low-angle sunlight at high latitudes. Instead of needles, they had large, broad lance- or tongueshaped leaves that fell to the ground at the end of summer. The fossil leaves are commonly found
as dense accumulations representing autumnal leaf banks.[34][35] The broad fossilized growth rings
in many Glossopteris woods reveal that the plants experienced strong growth spurts each springsummer but underwent abrupt cessation of growth before each following winter.[36]

Glossopteris leaves are morphologically simple so there are few characters that can be used to
differentiate species.[37] Consequently, many past researchers have considered the Permian
Glossopteris flora to be rather homogeneous with the same species distributed throughout the
Southern Hemisphere. However, more recent studies of the more morphologically diverse fertile
organs have shown that taxa had more restricted regional distributions and several intragondwanan floristic provinces are recognizable. Seeds, much too large to be wind-borne, could
not have blown across thousands of miles of open sea, nor is it likely they have floated across
vast oceans. Observations such as these led the Austrian geologist Eduard Suess to deduce that
there had once been a land bridge between these areas. He named this large land mass
Gondwanaland (named after the district in India where the plant Glossopteris was found). These
same observations would also lend support to Alfred Wegener's Continental drift theory.
The first Antarctic specimens of Glossopteris were discovered by members of Robert Scott's
doomed Terra Nova Expedition. The expedition members abandoned much of their gear in an
effort to reduce their load, but kept 35 pounds of Glossopteris fossils; these were found alongside
their bodies.[38]

Outcrops in Brazil
The first investigation of a Glossopteris flora associated with coal seams within a
paleogeographic and palaeoclimatic context, in the Paran Basin, southern Brazil, was that by
geologist Israel Charles White in 1908. This allowed correlation between Gondwanan coal
deposits in southern Brazil and those documented in South Africa, Australia, India and
Antarctica, and showed that this flora flourished in latitudes near the south pole.
In Rio Grande do Sul, Glossopteris leaves were found in paleorrota at Mina Faxinal, in Arroio
dos Ratos at Mina Morro do Papalo in Mariana Pimentel and Quitria in Pantano Grande.
Various species were recovered from the Rio Bonito Formation at these sites including G.
angustifolia, G. brasiliensis, G. browniana, G. communis, G. indica and G. occidentalis.[39]

References
1.
McLoughlin, S., 2012. Glossopteris insights into the architecture and relationships of an
iconic Permian Gondwanan plant. Journal of the Botanical Society of Bengal 65(2), 114.
McLoughlin, S. 2001. "The breakup history of Gondwana and its impact on pre-Cenozoic
floristic provincialism". Australian Journal of Botany, 49: 271-300.
McLoughlin, S., Lindstrm, S. & Drinnan, A.N. 1997 "Gondwanan floristic and
sedimentological trends during the Permian-Triassic transition: new evidence from the Amery
Group, northern Prince Charles Mountains, East Antarctica". Antarctic Science, 9: 281-298.
Vajda, V. & McLoughlin, S. 2007. "Extinction and recovery patterns of the vegetation
across the CretaceousPalaeogene boundary a tool for unravelling the causes of the endPermian mass-extinction". Review of Palaeobotany and Palynology 144: 99112.

 Lindstrm, S. & McLoughlin, S. 2007. "Synchronous palynofloristic extinction and

recovery after the end- Permian event in the Prince Charles Mountains, Antarctica: implications
for palynofloristic turnover across Gondwana". Review of Palaeobotany and Palynology 145:
89-122.
Pant, D.D. & Pant, R., 1987. Some Glossopteris leaves from Indian Triassic beds.
Palaeontographica 205B, 165-178.
Anderson, J. M. & Anderson, H. M., 1985. "Palaeoflora of southern Africa. Prodomus of
southern African megafloras Devonian to Lower Cretaceous". A.A. Balkema, Rotterdam. 423 pp.
Delevoryas, T. & Person, C.P. 1975. "Mexiglossa varia gen. et sp. nov., a new genus of
glossopteroid leaves from the Jurassic of Oaxaca, Mexico". Palaeontographica A 154, 114-120.
McLoughlin, S., Lindstrm, S. & Drinnan, A.N. 1997. "Gondwanan floristic and
sedimentological trends during the Permian-Triassic transition: new evidence from the Amery
Group, northern Prince Charles Mountains, East Antarctica". Antarctic Science, 9: 281-298.
Chandra, S. & Surange, K.R. 1979. "Revision of the Indian species of Glossopteris".
Monograph 2. Birbal Sahni Institute of Palaeobotany, Lucknow. 301 pp.
McLoughlin, S. 1994. "Late Permian plant megafossils from the Bowen Basin,
Queensland, Australia: Part 2". Palaeontographica 231B: 1-29.
McLoughlin, S. 1994. "Late Permian plant megafossils from the Bowen Basin,
Queensland, Australia: Part 3. Palaeontographica 231B: 31-62".
Appert, O., 1977. "Die Glossopterisflora der Sakoa in sdwest Madagaskar".
Palaeontographica 162B, 1 50.
Pigg, K. B., 1990. "Anatomically preserved Glossopteris foliage from the central
Transantarctic Mountains". Review of Palaeobotany and Palynology 66, 105-127.
McLoughlin, S. 2001. "The breakup history of Gondwana and its impact on pre-Cenozoic
floristic provincialism". Australian Journal of Botany, 49: 271-300
Holdgate G.R., McLoughlin, S., Drinnan A.N., Finkelman, R.B., Willett, J.C. &
Chiehowsky, L.A., 2005." Inorganic chemistry, petrography and palaeobotany of Permian coals
in the Prince Charles Mountains, East Antarctica". International Journal of Coal Geology 63:
156-177.
McLoughlin, S., 2012. "Glossopteris insights into the architecture and relationships of
an iconic Permian Gondwanan plant". Journal of the Botanical Society of Bengal 65(2), 114.
Meyen, S.V., 1987. Fundamentals of palaeobotany Chapman and Hall, London. 432 pp.
Zimina. V.G. 1967. "On Glossopteris and Gangamopteris in Permian deposits of the
Southern Maratime Territory". Paleontological Journal 2, 98-106.
McLoughlin, S., 2012." Glossopteris insights into the architecture and relationships of
an iconic Permian Gondwanan plant". Journal of the Botanical Society of Bengal 65(2), 114.
Brongniart, A., 1828a-38: Histoire des vgtaux fossiles on reserches botaniques et
gologiques sur les vgtaux renferms dans les diverses couches du globe. G. Dufour & Ed.
D'Ocagne, Paris. XII+488 pp. (Vol. I) / Crochard et Compagnie, Paris. 72 pp. (Vol. II).
Crane, P.R. 1985. Phylogenetic analysis of seed plants and the origin of angiosperms.
Annals of the Missouri Botanical Gardens 72, 716 793.
Gould, R.E., Delevoryas, T., 1977. The biology of Glossopteris: evidence from petrified
seed-bearing and pollen-bearing organs. Alcheringa 1, 87-399.
Weaver, L., McLoughlin, S. & Drinnan, A.N. 1997. Fossil woods from the Upper
Permian Bainmedart Coal Measures, northern Prince Charles Mountains, East Antarctica. AGSO
Journal of Australian Geology and Geophysics, 16: 655-676.

 McLoughlin, S. 1990. Some Permian glossopterid fructifications and leaves from the
Bowen Basin, Queensland, Australia. Review of Palaeobotany and Palynology, 62: 11-40.
McLoughlin, S. 1990. Late Permian glossopterid fructifications from the Bowen and
Sydney Basins, eastern Australia. Geobios, 23: 283-297.
McLoughlin, S. 1995 Bergiopteris and glossopterid fructifications from the Permian of
Western Australia and Queensland. Alcheringa, 19: 175-192.
Adendorff, R., McLoughlin, S. & Bamford, M.K. 2002. A new genus of ovuliferous
glossopterid fruits from South Africa. Palaeontologia africana, 38: 1-17.
Prevec, R., McLoughlin, S. & Bamford, M.K., 2008. Novel wing morphology revealed in
a South African ovuliferous glossopterid fructification. Review of Palaeobotany and Palynology
150: 22-36.
McLoughlin, S., 2012. Two new Senotheca (Glossopteridales) species from the Sydney
Basin, Australia, and a review of the genus. Review of Palaeobotany and Palynology 171, 140
151.
Lindstrm, S., McLoughlin, S. & Drinnan, A,N. 1997. Intraspecific variation of taeniate
bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains,
Antarctica. International Journal of Plant Science, 158: 673-684.
McLoughlin, S. 1993. Plant fossil distributions in some Australian Permian non-marine
sediments. Sedimentary Geology, 85: 601-619.
McLoughlin, S. & McNamara, K. 2001. Ancient Floras of Western Australia. Publication
of the Department of Earth and Planetary Sciences, Western Australian Museum. 42 pp.
McLoughlin, S. 1993. Plant fossil distributions in some Australian Permian non-marine
sediments. Sedimentary Geology, 85: 601-619.
Hill, R.S., Truswell, E.M., McLoughlin, S. & Dettmann, M.E. 1999. The evolution of the
Australian flora: fossil evidence. Flora of Australia, 2nd Edition, 1 (Introduction): 251-320.
Ryberg , P.E., & Taylor, E.L., 2007 . Silicified wood from the Permian and Triassic of
Antarctica: Tree rings from polar paleolatitudes. In Antarctica: A keystone in a changing world;
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Pigg, K.B. & McLoughlin, S. 1997. Anatomically preserved Glossopteris leaves from the
Bowen and Sydney basins, Australia. Review of Palaeobotany and Palynology, 97: 339-359.
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1.

Adami-Rodrigues, Karen; Alves de Souza, Paulo; Iannuzzi, Roberto; Pinto,

Iraj Damiani (2004). "Herbivoria em Floras Gondunicas do NeoPalezoico do Rio
Grande do Sul" (PDF). Revista Brasileira de Paleontologia 7 (2): 93102.

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gologiques sur les vgtaux renferms dans les diverses couches du globe. G. Dufour
and E. DOcagne, Paris 1: 265-288.

Anderson, H.M. & Anderson, J.M. 1985. The Palaeoflora of Southern Africa: Prodromus
of Southern African Megafloras, Devonian to Lower Cretaceous. A.A. Balkema,
Rotterdam. 416 pp.

Chandra, S. & Surange, K.R. 1979. Revision of the Indian species of Glossopteris.
Monograph 2. Birbal Sahni Institute of Palaeobotany, Lucknow. 301 pp.

Davis, Paul and Kenrick, Paul. 2004. Fossil Plants. Smithsonian Books (in association
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petrified seed-bearing and pollen-bearing organs. Alcheringa, 1: 387-399.

Pant DD 1977 The plant of Glossopteris. J Indian Bot Soc 56: 1-23.

Pant, D.D. & Gupta, K.L. 1971. Cuticular structure of some Indian Lower Gondwana
species of Glossopteris Brongniart. Part 2. - Palaeontographica, 132B: 130-152.

Pant, D.D. & Nautiyal, D.D. 1987. Diphyllopteris verticellata Srivastava, the probable
seedling of Glossopteris from the Paleozoic of India. - Rev. Palaeobot. Palynol., 51: 3136.

Pant, D.D. & Pant, R. 1987. Some Glossopteris leaves from Indian Triassic beds. Palaeontographica, 205B: 165-178.

Pant, D.D. & Singh, K.B. 1971. Cuticular structure of some Indian Lower Gondwana
species of Glossopteris Brongniart. Part 3. - Palaeontographica, 135B: 1-40.

Pigg, K. B. 1990. Anatomically preserved Glossopteris foliage from the central

Transantarctic Mountains. Rev. Palaeobot. Palynol. 66: 105-127.

Pigg, K.B. & McLoughlin, S. 1997. Anatomically preserved Glossopteris leaves from the
Bowen and Sydney basins, Australia. Review of Palaeobotany and Palynology, 97: 339359.

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Antarctica. - Internat. J. Coal Geol., 12: 657-679.

External links

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