Fern spike

In paleontology, a fern spike is the occurrence of unusually high spore abundance of ferns in the fossil record, usually immediately (in a geological sense) after an extinction event. The spikes are believed to represent a large, temporary increase in the number of ferns relative to other terrestrial plants after the extinction or thinning of the latter. Fern spikes are strongly associated with the Cretaceous–Paleogene extinction event,[1][2] although they have been found in other points of time and space such as at the Triassic-Jurassic boundary.[3][4] Outside the fossil record, fern spikes have been observed to occur in response to local extinction events, such as the 1980 Mount St. Helens eruption.[5]

Illustration depicting an ecosystem in successive temporal stages. It begins as a biodiverse system with many species, followed by a sudden loss of plant life when a disaster strikes (depicted as dust in the air and on the ground). Gradually, ferns germinate in and colonize the area. Once ferns are established, conifer trees begin growing.
Succession of an ecosystem following disturbance, in the form of a fern spike.

Causes

Extinction events have historically been caused by massive environmental disturbances, such as meteor strikes. Volcanic eruptions can also wipe out local ecosystems through pyroclastic flows and landslides, leaving the ground bare for new colonization.[6] For a population to recover and thrive after such an event, it must be able to tolerate the conditions of the disturbed environment. Ferns have multiple characteristics which predispose them to grow in those environments.

Spore characteristics

Plants generally reproduce with spores or seeds, meaning those will be what germinates in a disaster's aftermath. But spores have advantages over seeds in the environmental conditions produced by a disaster. They are generally produced in higher numbers than seeds, and are smaller, aiding wind dispersal.[6] While many wind-dispersed pollens of seed plants are smaller and farther dispersed than spores,[7] pollen cannot germinate into a plant and must land in a receptive flower. Some seed plants also require animals to disperse their seeds, which may not be present after a disaster. These characteristics allow ferns to rapidly colonize an area with their spores.

Fern spores require light to germinate.[8] Following major disturbances that clear or reduce plant life, the ground would receive ample sunlight that may promote spore germination. Some species' spores contain chlorophyll, which hastens germination and may aid rapid colonization of clear ground.[9]

Environmental tolerance

After the eruption of El Chichón, the fern Pityrogramma calomelanos was observed to regenerate from rhizomes buried by ash, even though the plants' leaves were destroyed.[6] The rhizomes tolerated exposure to heat and sulfur from the volcanic matter. Their survival suggests resilience of ferns to the harsh environmental conditions imposed by certain kinds of disasters, and rhizome regeneration may have been a factor in fern recovery after other environmental events.

Ecology

Fern spikes follow the pattern of ecological succession. In the past and in modern times, ferns have been observed to act as pioneer species.[5] Eventually, their abundance at a site decreases as other plants such as gymnosperms begin to grow.[2]

Spore availability

Fern spikes cannot occur without ferns already existing in the area, so spikes occur primarily in regions where ferns are already a prominent part of the ecosystem. At the Cretaceous-Paleogene extinction event, a fern spike occurred in the New Zealand area, where ferns made up 25% of plant abundance pre-extinction. After the event, fern abundance increased to 90%.[2]

Detection

Prehistoric fern spikes can be detected by sampling sediment. Sources include sediment that has been accumulating in a lake since the event of interest and sedimentary rocks such as sandstone.[5] Because sediment accumulates over time and thus shows superposition, layers can be assigned to certain times. Spore concentration in a layer can be compared to the concentration at different times, and concentration of other particles such a pollen grains. A fern spike is characterized by a suddenly higher abundance of fern spores following a disaster, generally accompanied by a decrease in other plant species as indicated by their pollen. Eventually fern abundance will decrease, hence the term "spike" describing the pattern.

Modern fern spikes can simply be directly observed, and allow for observation of factors contributing to the spike that may not be detectable otherwise, such as rhizomes persisting in ash.[6]

Significance

Because fern spikes generally coincide with certain disasters such as meteorite strikes and volcanic eruptions, their presence in the fossil record can indicate those events. A fern spike is believed to support a meteorite impact as cause of the Triassic-Jurassic extinction event, similar to the one later causing extinction at the end of the Cretaceous period.[3]

Known events

A fern spike followed a fungal spike after the Permian–Triassic extinction event (252 Ma). It has been observed in Australia.[10]

After the Triassic-Jurassic extinction event (201.3 Ma), ferns drastically increased in abundance while seed plants became scarce. The spike has been detected in eastern North America and Europe.[3][11]

A very widespread fern spike occurred after the Cretaceous–Paleogene extinction event (66 Ma).[2] The spike has been predominantly observed in North America, with just one observance outside the continent in Japan.[1]

Fern spikes today are often observed after volcanic eruptions. The areas affected by the eruptions of Mount St. Helens (May 18, 1980) and El Chichón (March—April 1982) exhibited such a pattern.[5][6]

See also

References
  1. Schultz, P. H.; D'Hondt, S. (1996). "Cretaceous-Tertiary (Chicxulub) impact angle and its consequences". Geology. 24 (11): 963–967. Bibcode:1996Geo....24..963S. doi:10.1130/0091-7613(1996)024<0963:CTCIAA>2.3.CO;2.
  2. Vajda, V.; Raine, J. I.; Hollis, C. J. (2001). "Indication of global deforestation at the Cretaceous-Tertiary boundary by New Zealand fern spike". Science. 294 (5547): 1700–1702. Bibcode:2001Sci...294.1700V. doi:10.1126/science.1064706. PMID 11721051.
  3. Fowell, S. J.; Olsen, P. E. (1993). "Time calibration of Triassic-Jurassic microfloral turnover, eastern North-America". Tectonophysics. 222 (3–4): 361–369. Bibcode:1993Tectp.222..361F. doi:10.1016/0040-1951(93)90359-R.
  4. Olsen, P. E.; Kent, D. V.; Sues, H. D.; et al. (2002). "Ascent of dinosaurs linked to an iridium anomaly at the Triassic-Jurassic boundary". Science. 296 (5571): 1305–1307. Bibcode:2002Sci...296.1305O. doi:10.1126/science.1065522. PMID 12016313.
  5. Adams, Jonathan (2009). Species Richness: Patterns in the Diversity of Life. Environmental Sciences. Springer-Verlag Berlin Heidelberg. p. 125. ISBN 9783540742784.
  6. Spicer, Robert A.; Burnham, Robyn J.; Grant, Paul; Glicken, Harry (1985). "Pityrogramma calomelanos, the Primary, Post-Eruption Colonizer of Volcán Chichonal, Chiapas, Mexico". American Fern Journal. 75 (1): 1–5. doi:10.2307/1546571. JSTOR 1546571.
  7. Raynor, Gilbert S.; Ogden, Eugene C.; Hayes, Janet V. (July 1976). "Dispersion of Fern Spores Into and Within a Forest". Rhodora. 78 (815): 473–487. JSTOR 23311224.
  8. Gantt, Elisabeth; Arnott, Howard J. (January 1965). "Spore Germination and Development of the Young Gametophyte of the Ostrich Fern (Matteuccia struthiopteris)". American Journal of Botany. 52 (1): 82. doi:10.1002/j.1537-2197.1965.tb06760.x. ISSN 0002-9122. JSTOR 2439978.
  9. Lloyd, Robert M.; Klekowski, Edward J. (1970). "Spore Germination and Viability in Pteridophyta: Evolutionary Significance of Chlorophyllous Spores". Biotropica. 2 (2): 129–137. doi:10.2307/2989770. JSTOR 2989770.
  10. Retallack, G. J. (January 1995). "Permian-Triassic Life Crisis on Land". Science. American Association for the Advancement of Science. 267 (5194): 77–80. Bibcode:1995Sci...267...77R. doi:10.1126/science.267.5194.77. JSTOR 2886044. PMID 17840061.
  11. Van de Schootbrugge, B.; Quan, T.M.; Lindstrom, S.L.; Püttmann, W.; Heunisch, C.; Pross, J.; Fiebig, J.; Petschick, R.; Rohling, H.G. (August 2009). "Floral changes across the Triassic/Jurassic boundary linked to flood basalt volcanism". Nature Geoscience. 2 (8): 589–594. CiteSeerX 10.1.1.730.9425. doi:10.1038/ngeo577. ISSN 1752-0908.
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