Serotiny

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Fire has caused minimal damage to this Banksia serrata (saw banksia) fruiting structure, but has triggered the opening of the follicles and the release of seed. Banksia serrata2.jpg
Fire has caused minimal damage to this Banksia serrata (saw banksia) fruiting structure, but has triggered the opening of the follicles and the release of seed.

Serotiny in botany simply means 'following' or 'later'.

Contents

In the case of serotinous flowers, it means flowers which grow following the growth of leaves, [1] or even more simply, flowering later in the season than is customary with allied species. Having serotinous leaves is also possible, these follow the flowering.

Serotiny is contrasted with coetany. Coetaneous flowers or leaves appear together with each other. [1]

In the case of serotinous fruit, the term is used in the more general sense of plants that release their seed over a long period of time, irrespective of whether release is spontaneous; in this sense the term is synonymous with bradyspory.

In the case of certain Australian, North American, South African or Californian plants which grow in areas subjected to regular wildfires, serotinous fruit can also mean an ecological adaptation exhibited by some seed plants, in which seed release occurs in response to an environmental trigger, rather than spontaneously at seed maturation. The most common and best studied trigger is fire, and the term serotiny is used to refer to this specific case.

Possible triggers include: [2]

Some plants may respond to more than one of these triggers. For example, Pinus halepensis exhibits primarily fire-mediated serotiny, [3] but responds weakly to drying atmospheric conditions. [4] Similarly, Sierras sequoias and some Banksia species are strongly serotinous with respect to fire, but also release some seed in response to plant or branch death.

Serotiny can occur in various degrees. Plants that retain all of their seed indefinitely in the absence of a trigger event are strongly serotinous. Plants that eventually release some of their seed spontaneously in the absence of a trigger are weakly serotinous. Finally, some plants release all of their seed spontaneously after a period of seed storage, but the occurrence of a trigger event curtails the seed storage period, causing all seed to be released immediately; such plants are essentially non-serotinous, but may be termed facultatively serotinous.

Fire-mediated serotiny

In the southern hemisphere, fire-mediated serotiny is found in angiosperms in fire-prone parts of Australia and South Africa. It is extremely common in the Proteaceae of these areas, and also occurs in other taxa, such as Eucalyptus (Myrtaceae) and even exceptionally in Erica sessiliflora (Ericaceae). In the northern hemisphere, it is found in a range of conifer taxa, including species of Pinus , [5] Cupressus , Sequoiadendron , and more rarely Picea .

Since even non-serotinous cones and woody fruits can provide protection from the heat of fire, [6] [7] the key adaptation of fire-induced serotiny is seed storage in a canopy seed bank, which can be released by fire. [8] The fire-release mechanism is commonly a resin that seals the fruit or cone scales shut, but which melts when heated. [9] [10] This mechanism is refined in some Banksia by the presence inside the follicle of a winged seed separator which blocks the opening, preventing the seed from falling out. Thus, the follicles open after fire, but seed release does not occur. As the cone dries, wetting by rain or humidity causes the cone scales to expand and reflex, promoting seed release. [11] The seed separator thus acts as a lever against the seeds, gradually prying them out of the follicle over the course of one or more wet-dry cycles. The effect of this adaptation is to ensure that seed release occurs not in response to fire, but in response to the onset of rains following fire.

The relative importance of serotiny can vary among populations of the same plant species. For example, North American populations of lodgepole pine ( Pinus contorta ) can vary from being highly serotinous to having no serotiny at all, opening annually to release seed. [12] Different levels of cone serotiny have been linked to variations in the local fire regime: areas that experience more frequent crown-fire tend to have high rates of serotiny, while areas with infrequent crown-fire have low levels of serotiny. [3] [13] Additionally, herbivory of lodgepole pines can make fire-mediated serotiny less advantageous in a population. Red squirrels ( Sciurus vulgaris ) and red crossbills ( Loxia curvirostra ) will eat seeds, and so serotinous cones, which last in the canopy longer, are more likely to be chosen. [14] [15] Serotiny occurs less frequently in areas where this seed predation is common.

Pyriscence can be understood as an adaptation to an environment in which fires are regular and in which post-fire environments offer the best germination and seedling survival rates. In Australia, for example, fire-mediated serotiny occurs in areas that not only are prone to regular fires but also possess oligotrophic soils and a seasonally dry climate. This results in intense competition for nutrients and moisture, leading to very low seedling survival rates. The passage of fire, however, reduces competition by clearing out undergrowth, and results in an ash bed that temporarily increases soil nutrition; thus the survival rates of post-fire seedlings is greatly increased. Furthermore, releasing a large number of seeds at once, rather than gradually, increases the possibility that some of those seeds will escape predation. [16] Similar pressures apply in Northern Hemisphere conifer forests, but in this case there is the further issue of allelopathic leaf litter, which suppresses seed germination. Fire clears out this litter, eliminating this obstacle to germination.

Evolution

Serotinous adaptations occur in at least 530 species in 40 genera, in multiple (paraphyletic) lineages. Serotiny likely evolved separately in these species, but may in some cases have been lost by the related non-serotinous species.

In the genus Pinus, serotiny likely evolved because of the atmospheric conditions during the Cretaceous period. [5] The atmosphere during the Cretaceous had higher oxygen and carbon dioxide levels than our atmosphere. Fire occurred more frequently than it does currently, and plant growth was high enough to create an abundance of flammable material. Many Pinus species adapted to this fire-prone environment with serotinous pine cones.

A set of conditions must be met in order for long-term seed storage to be evolutionarily viable for a plant:

Related Research Articles

<span class="mw-page-title-main">Pine</span> Genus of plants in the conifer family Pinaceae

A pine is any conifer tree or shrub in the genus Pinus of the family Pinaceae. Pinus is the sole genus in the subfamily Pinoideae.

<span class="mw-page-title-main">Conifer</span> Group of cone-bearing seed plants

Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta, also known as Coniferophyta or Coniferae. The division contains a single extant class, Pinopsida. All extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 2002, Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species.

Hygroscopy is the phenomenon of attracting and holding water molecules via either absorption or adsorption from the surrounding environment, which is usually at normal or room temperature. If water molecules become suspended among the substance's molecules, adsorbing substances can become physically changed, e.g. changing in volume, boiling point, viscosity or some other physical characteristic or property of the substance. For example, a finely dispersed hygroscopic powder, such as a salt, may become clumpy over time due to collection of moisture from the surrounding environment.

<span class="mw-page-title-main">Pinyon pine</span> Group of conifers

The pinyon or piñon pine group grows in southwestern North America, especially in New Mexico, Colorado, Arizona, and Utah. The trees yield edible nuts, which are a staple food of Native Americans, and widely eaten as a snack and as an ingredient in New Mexican cuisine. The name comes from the Spanish pino piñonero, a name used for both the American varieties and the stone pine common in Spain, which also produces edible nuts typical of Mediterranean cuisine. Harvesting techniques of the prehistoric American Indians are still used today to collect the pinyon seeds for personal use or for commercialization. The pinyon nut or seed is high in fats and calories. In the western United States, pinyon pines are often found in pinyon–juniper woodlands.

<i>Pinus flexilis</i> Pine tree found in North America

Pinus flexilis, the limber pine, is a species of pine tree-the family Pinaceae that occurs in the mountains of the Western United States, Mexico, and Canada. It is also called Rocky Mountain white pine.

<i>Pinus contorta</i> Species of plant

Pinus contorta, with the common names lodgepole pine and shore pine, and also known as twisted pine, and contorta pine, is a common tree in western North America. It is common near the ocean shore and in dry montane forests to the subalpine, but is rare in lowland rain forests. Like all pines, it is an evergreen conifer.

<i>Pinus serotina</i> Species of conifer

Pinus serotina, the pond pine, black bark pine, bay pine,marsh pine, or pocosin pine, is a pine tree found along the Southeastern portion of the Atlantic coastal plain of the United States, from southern New Jersey south to Florida and west to southern Alabama. Pond pine distribution may be starting to spread west towards Mississippi and Tennessee.

<i>Pinus virginiana</i> Species of conifer

Pinus virginiana, the Virginia pine, scrub pine, Jersey pine, possum pine, is a medium-sized tree, often found on poorer soils from Long Island in southern New York south through the Appalachian Mountains to western Tennessee and Alabama. The usual size range for this pine is 9–18 m, but can grow larger under optimum conditions. The trunk can be as large as 20 inches diameter. This tree prefers well-drained loam or clay, but will also grow on very poor, sandy soil, where it remains small and stunted. The typical life span is 65 to 90 years.

<span class="mw-page-title-main">Jack pine</span> Species of tree

Jack pine, also known as grey pine or scrub pine, is a North American pine.

<span class="mw-page-title-main">Fire ecology</span> Study of fire in ecosystems

Fire ecology is a scientific discipline concerned with the effects of fire on natural ecosystems. Many ecosystems, particularly prairie, savanna, chaparral and coniferous forests, have evolved with fire as an essential contributor to habitat vitality and renewal. Many plant species in fire-affected environments use fire to germinate, establish, or to reproduce. Wildfire suppression not only endangers these species, but also the animals that depend upon them.

<span class="mw-page-title-main">Disturbance (ecology)</span> Temporary change in environmental conditions that causes a pronounced change in an ecosystem

In ecology, a disturbance is a temporary change in environmental conditions that causes a pronounced change in an ecosystem. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and abiotic elements. A disturbance can also occur over a long period of time and can impact the biodiversity within an ecosystem.

Pyrophytes are plants which have adapted to tolerate fire.

<i>Banksia prionotes</i> Species of shrub or tree in the family Proteaceae native to the southwest of Western Australia

Banksia prionotes, commonly known as acorn banksia or orange banksia, is a species of shrub or tree of the genus Banksia in the family Proteaceae. It is native to the southwest of Western Australia and can reach up to 10 m (33 ft) in height. It can be much smaller in more exposed areas or in the north of its range. This species has serrated, dull green leaves and large, bright flower spikes, initially white before opening to a bright orange. Its common name arises from the partly opened inflorescence, which is shaped like an acorn. The tree is a popular garden plant and also of importance to the cut flower industry.

The ecology of Banksia is the relationships and interactions among the plant genus Banksia and its environment. Banksia has a number of adaptations that have so far enabled the genus to survive despite dry, nutrient-poor soil, low rates of seed set, high rates of seed predation and low rates of seedling survival. These adaptations include proteoid roots and lignotubers; specialised floral structures that attract nectariferous animals and ensure effective pollen transfer; and the release of seed in response to bushfire.

<span class="mw-page-title-main">Closed-cone conifer forest</span>

A Closed-cone conifer forest or woodland is a plant community occurring in coastal California and several offshore islands. The forests typically have a single-aged single-species conifer overstory with dense ladder fuels. Overstory species include coulter pine, monterey pine, bishop pine, shore pine, and several endemic cypresses, species which generally rely on fire to open their cones and release seeds. Closed-cone forests often grow in low nutrient and/or stressed soils, which can lead to slow growth.

<span class="mw-page-title-main">Sierra Nevada subalpine zone</span> Biotic zone in California, United States

The Sierra Nevada subalpine zone refers to a biotic zone below treeline in the Sierra Nevada mountain range of California, United States. This subalpine zone is positioned between the upper montane zone at its lower limit, and tree line at its upper limit.

A seed bank stores seeds from plants and is significant in preserving plant genetic diversity. Seed banks can be categorized according to their location and the ecological functions they serve. The primary types include soil seed banks, which are found in terrestrial environments; wetland seed banks, located in aquatic habitats; and canopy seed banks, present in the upper layers of forest ecosystems. Each type of seed contributes uniquely to plant biodiversity, ecosystem resilience and human well-being. Also, they are unique in the way they retain their seed. For example, in canopy seed banks or aerial seed banks, the seeds are stored in the canopies of trees and plants.

<span class="mw-page-title-main">Cassia crossbill</span> Species of bird

The Cassia crossbill is a passerine bird in the family Fringillidae. It is endemic to the South Hills and Albion Mountains in southern Idaho. Cassia crossbill rarely interbreeds with other call types that move into the South Hills of Idaho yearly, and can be considered to represent a distinct species via ecological speciation. The Cassia crossbill have specialized beaks to access the seeds of the lodgepole pine cones in this region, but are poorly adapted to other pine cones in surrounding regions.

<span class="mw-page-title-main">Fire adaptations</span> Traits of plants and animals

Fire adaptations are traits of plants and animals that help them survive wildfire or to use resources created by wildfire. These traits can help plants and animals increase their survival rates during a fire and/or reproduce offspring after a fire. Both plants and animals have multiple strategies for surviving and reproducing after fire. Plants in wildfire-prone ecosystems often survive through adaptations to their local fire regime. Such adaptations include physical protection against heat, increased growth after a fire event, and flammable materials that encourage fire and may eliminate competition.

References

  1. 1 2 Goodrich, Sherel (31 October 1983). "Utah flora: Salicacea". Great Basin Naturalist. 43 (4): 536. Retrieved 1 December 2020.
  2. 1 2 Lamont, B.; Lemaitre, D.; Cowling, R.; Enright, N. (1991). "Canopy seed storage in woody-plants". Botanical Review. 57 (4): 277–317. doi:10.1007/bf02858770. S2CID   37245625.
  3. 1 2 Hernández-Serrano, A; Verdú M.; González-Martínez S.C.; Pausas J.G. (2013). "Fire structures pine serotiny at different scales" (PDF). American Journal of Botany. 100 (12): 2349–2356. doi:10.3732/ajb.1300182. PMID   24222682.
  4. Nathan, R; Safriel, U.; Noy-Meir, I.; Schiller, G. (1999). "Seed release without fire in Pinus halepensis, a Mediterranean serotinous wind-dispersed tree". Journal of Ecology. 87 (4): 659–669. CiteSeerX   10.1.1.534.8609 . doi:10.1046/j.1365-2745.1999.00382.x. S2CID   54592020.
  5. 1 2 He, T; Pausas JG; Belcher CM; Schwilk DW; Lamont BB. (2012). "Fire-adapted traits of Pinus arose in the fiery Cretaceous" (PDF). New Phytologist. 194 (3): 751–759. doi:10.1111/j.1469-8137.2012.04079.x. hdl:10261/48120. PMID   22348443.
  6. Michaletz, ST; Johnson EA; Mell WE; Greene DF (2013). "Timing of fire relative to seed development may enable non-serotinous species to recolonize from the aerial seed banks of fire-killed trees". Biogeosciences. 10 (7): 5061–5078. Bibcode:2013BGeo...10.5061M. doi: 10.5194/bg-10-5061-2013 .
  7. Pounden, E; Greene DF; Michaletz ST (2014). "Non-serotinous woody plants behave as aerial seed bank species when a late-summer wildfire coincides with a mast year". Ecology and Evolution. 4 (19): 3830–3840. doi:10.1002/ece3.1247. PMC   4301049 . PMID   25614797.
  8. Lamont, BB; Enright NJ (2000). "Adaptive advantages of aerial seed banks". Plant Species Biology. 15 (2): 157–166. doi: 10.1046/j.1442-1984.2000.00036.x .
  9. Beaufait, WR (1960). "Some Effects of High Temperatures on the Cones and Seeds of Jack Pine". Forest Science. 6: 194–199.
  10. Johnson, EA; Gutsell SL (1993). "Heat budget and fire behaviour associated with the opening of serotinous cones in two Pinus species". Journal of Vegetation Science. 4 (6): 745–750. Bibcode:1993JVegS...4..745J. doi:10.2307/3235610. JSTOR   3235610.
  11. Dawson, C; Vincent JFV; Rocca A-M (1997). "How pine cones open". Nature. 390 (6661): 668. Bibcode:1997Natur.390..668D. doi:10.1038/37745. S2CID   4415713.
  12. Muir, P. S.; Lotan, J. E. (1985). "Disturbance history and serotiny of Pinus contorta in western Montana". Ecology. 66 (5): 1658–1668. Bibcode:1985Ecol...66.1658M. doi:10.2307/1938028. JSTOR   1938028.
  13. Schoennagel, T.; Turner, M. G.; Romme, W. H. (2003). "The influence of fire interval and serotiny on postfire lodgepole pine density in Yellowstone National Park". Ecology. 84 (11): 2967–2978. Bibcode:2003Ecol...84.2967S. doi:10.1890/02-0277.
  14. Benkman, C.W.; Holimon, W.C.; Smith, J.W. (2001). "The influence of a competitor on the geographic mosaic of coevolution between crossbills and lodgepole pine". Evolution. 55 (2): 282–294. doi:10.1554/0014-3820(2001)055[0282:TIOACO]2.0.CO;2. PMID   11308086.
  15. Talluto, M. V.; Benkman, C. W. (2014). "Conflicting selection from fire and seed predation drives fine-scaled phenotypic variation in a widespread North American conifer". PNAS. 111 (26): 9543–9548. Bibcode:2014PNAS..111.9543T. doi: 10.1073/pnas.1400944111 . PMC   4084486 . PMID   24979772.
  16. Bradshaw, S. Don; Kingsley W. Dixon; Stephen D. Hopper; Hans Lambers; Shane R. Turner (2011). "Little evidence for fire-adapted plant traits in Mediterranean climate regions". Trends in Plant Science. 16 (2): 69–76. doi:10.1016/j.tplants.2010.10.007. PMID   21095155.
  17. Hernández-Serrano, Ana (2014). "Heritability and quantitative genetic divergence of serotiny, a fire-persistence plant trait" (PDF). Annals of Botany. 114 (3): 571–577. doi:10.1093/aob/mcu142. PMC   4204669 . PMID   25008363.