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''Pinus elliottii'', commonly known as slash pine,Family, P. P. (1990). Pinus elliottii Engelm. slash pine. ''Silvics of North America: Conifers'', (654), 338. is a
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
tree native to the
Southeastern United States The Southeastern United States, also referred to as the American Southeast or simply the Southeast, is a geographical region of the United States. It is located broadly on the eastern portion of the southern United States and the southern por ...
. Slash pine is named after the "slashes" – swampy ground overgrown with trees and bushes – that constitute its habitat. Other common names include swamp pine, yellow slash pine, and southern Florida pine. Slash pine has two different varieties: ''P. e.'' var. ''elliottii'' and ''P. e.'' var. ''densa''. Historically, slash pine has been an important economic timber for naval stores,
turpentine Turpentine (which is also called spirit of turpentine, oil of turpentine, terebenthene, terebinthine and (colloquially) turps) is a fluid obtained by the distillation of resin harvested from living trees, mainly pines. Mainly used as a special ...
, and
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
. The wood of slash pine is known for its unusually high strength, especially for a pine. It exceeds many hardwoods and is even comparable to very dense woods such as ironwood.


Description and taxonomy

This tree is fast-growing, but not very long-lived by pine standards (to 200 years). It reaches heights of with a trunk diameter of . The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are needle-like, very slender, in clusters of two or three, and long. The
cones A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
are glossy red-brown, in length, with a short (), thick prickle on each scale. It is known for its conical shape and unusually high strength, especially for a pine. Its wood has an average crush strength of 8,140 lb/in2 (56.1 MPa), which exceeds many hardwoods such as white ash (7,410 lb/in2) and black maple (6,680 lb/in2). It is not as strong as black ironwood (9,940 lb/in2), but because its average density is less than half that of ironwood, slash pine has a far greater strength-to-weight ratio. It may be distinguished from the related
loblolly pine ''Pinus taeda'', commonly known as loblolly pine, is one of several pines native to the Southeastern United States, from East Texas to Florida, and north to southern New Jersey. The wood industry classifies the species as a southern yellow pine. ...
(''P. taeda'') by the somewhat longer, glossier needles and larger red-brown cones, and from
longleaf pine The longleaf pine (''Pinus palustris'') is a pine species native to the Southeastern United States, found along the coastal plain from East Texas to southern Virginia, extending into northern and central Florida. In this area it is also known as ...
(''P. palustris'') by the shorter, more slender needles and smaller cones with less broad scales. Two varieties of ''P. elliotii'' are described, but recent genetic studies have indicated that the varieties may not be more closely related to each other than they are to other pines in the Southeast. If this is the case, reclassifying these varieties as separate species would be warranted. ''P. elliottii'' can
hybridize Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
with ''P. taeda'', sand pine (''
Pinus clausa ''Pinus clausa'' is a species of pine endemic to the Southeastern United States. Its common names include sand pine, Florida spruce pine, Alabama pine, and scrub pine. Distribution The tree is found in two separate locations, one across central ...
''), and ''P. palustris''.Carey, Jennifer H. 1992. Pinus elliottii. In: Fire Effects Information System, U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. https://www.fs.fed.us/database/feis/plants/tree/pinell/all.html The two commonly accepted varieties are the following: * ''P. e.'' var. ''elliottii'' (typical slash pine) ranges from
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
to
Louisiana Louisiana , group=pronunciation (French: ''La Louisiane'') is a state in the Deep South and South Central regions of the United States. It is the 20th-smallest by area and the 25th most populous of the 50 U.S. states. Louisiana is borde ...
, and south to central
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. Its leaves occur in bundles, fascicles of twos and threes, mostly threes, and the cones are larger, . * ''P. e.'' var. ''densa'' (South Florida slash pine, Dade County pine) is found in the
pine rocklands The South Florida rocklands ecoregion, in the tropical and subtropical moist broadleaf forests biome, occurs in southern Florida and the Florida Keys in the United States, where they would naturally cover an area of . These forests form on lim ...
of southern Florida and the
Florida Keys The Florida Keys are a coral cay archipelago located off the southern coast of Florida, forming the southernmost part of the continental United States. They begin at the southeastern coast of the Florida peninsula, about south of Miami, and e ...
, including the
Everglades The Everglades is a natural region of tropical climate, tropical wetlands in the southern portion of the U.S. state of Florida, comprising the southern half of a large drainage basin within the Neotropical realm. The system begins near Orland ...
. Leaves are nearly all in bundles of two, with longer needles. The cones are smaller, , the wood is denser, and the tree has a thicker taproot. Unlike the typical variety of slash pine, seedlings of ''P. e.'' var. ''densa'' has a "grass stage" similar to longleaf pine. ''P. e.'' var. ''densa'' is not frost tolerant, which limits its range to South Florida.


Range and habitat

Communities dominated by slash pine are termed "slash pine forests". Slash pine is predominately found in Florida and
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
, and extends from South Carolina west to southeastern
Louisiana Louisiana , group=pronunciation (French: ''La Louisiane'') is a state in the Deep South and South Central regions of the United States. It is the 20th-smallest by area and the 25th most populous of the 50 U.S. states. Louisiana is borde ...
, and south to the Florida Keys. It is common in
East Texas East Texas is a broadly defined cultural, geographic, and ecological region in the eastern part of the U.S. state of Texas that comprises most of 41 counties. It is primarily divided into Northeast and Southeast Texas. Most of the region consi ...
, where it was first planted at the E.O. Siecke State Forest in 1926. The natural habitat is sandy subtropical maritime forests and wet
flatwoods Flatwoods, pineywoods, pine savannas and longleaf pine-wiregrass ecosystem are terms that refer to an ecological community in the southeastern coastal plain of North America. Flatwoods are an ecosystem maintained by wildfire or prescribed fire an ...
. Slash pine generally grows better in warm, humid areas where the average annual temperature is above , with extreme ranges from . Factors such as competition, fire, and precipitation may limit the natural distribution of these trees. Slash pines are able to grow in an array of soils, but pine stands that are close to bodies of water such as swamps and ponds grow better because of higher soil moisture and seedling protection from wildfire. These forests have been managed through controlled fires since the beginning of the 20th century. Within the first year, ''P. elliottii'' is particularly susceptible to seedling mortality caused by fire. ''P. e.'' var. ''densa'' is more fire resistant than ''P. e.'' var.'' elliottii'' because it has thicker bark.


Fire ecology


History

Fire has long been an important element in Southeastern forests. Native Americans burned land to improve grass growth for grazing and visibility for hunting.Johnson, A. S., & Hale, P. E. (2000, September). The Historical Foundations of Prescribed Burning for Wildlife: a Southeastern Perspective. In ''The Role of Fire in Nongame Wildlife Management and Community Restoration: Traditional Uses and New Directions Proceedings of a Special Workshop''(p. 11). When European settlers arrived in the New World, they brought new diseases that severely diminished the Native American populations. Over time, with the lack of consistent burning, much of the open land of the South reverted to forest land. Logging began to increase in the Southeast, which created some tension between the loggers and local farmers. The loggers wanted to continue to burn the forest, but the local farmers were concerned about how burning would affect cattle grazing and
turpentine Turpentine (which is also called spirit of turpentine, oil of turpentine, terebenthene, terebinthine and (colloquially) turps) is a fluid obtained by the distillation of resin harvested from living trees, mainly pines. Mainly used as a special ...
production. Fire maintenance has long been a controversial issue. In the 1940s, the
Smokey Bear Smokey Bear is an American campaign and advertising icon of the U.S. Forest Service. In the Wildfire Prevention Campaign, which is the longest-running public service announcement campaign in United States history, the Ad Council, the United St ...
campaign to prevent wildfires promoted a shift toward fire suppression. Subsequently, many of these fire-dependent ecosystems became increasingly dominated by more shade-tolerant tree species (hardwoods). Despite many reports from the U.S. Forest Service about the benefits fire has on forage production, pine regeneration, control of tree pathogens, and reducing risks of wildfires, controlled burning did not begin to regain traction until the 1950s and 1960s.


Uses

Without regular fire intervals in slash pine forests, the ecosystem can change over time. For example, in the northern range for slash pine, forests can convert from mesic
flatwoods Flatwoods, pineywoods, pine savannas and longleaf pine-wiregrass ecosystem are terms that refer to an ecological community in the southeastern coastal plain of North America. Flatwoods are an ecosystem maintained by wildfire or prescribed fire an ...
to denser mixed-hardwood canopies with trees such as
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
s,
hickory Hickory is a common name for trees composing the genus ''Carya'', which includes around 18 species. Five or six species are native to China, Indochina, and India (Assam), as many as twelve are native to the United States, four are found in Mexi ...
, and southern magnolia.Snyder, J. R., Ross, M. S., Koptur, S., & Sah, J. (2005). Developing ecological criteria for prescribed fire in south Florida pine rockland ecosystems. In South Florida, the
pine rocklands The South Florida rocklands ecoregion, in the tropical and subtropical moist broadleaf forests biome, occurs in southern Florida and the Florida Keys in the United States, where they would naturally cover an area of . These forests form on lim ...
can convert to a rockland hammock dominated by woody shrubs and invasive plants. Invasive species are a major management issue in the South. Many pine trees and native plants are adapted to fire, meaning they require fire disturbance to open their pine cones, germinate seeds, and cue other metabolic processes. Fire can be a good management strategy for invasive species because many invasive plants are not adapted to fire. Therefore, fire can eliminate the parental plant or reduce seed viability. Controlled burning is also used to help reduce pathogen load in an ecosystem. For example, fire can eliminate pest populations or resting fungal spores that could infect new seedlings. Low-intensity burns can also clear space in the understory and provide nutrient pulses that benefit the understory vegetation. Fire is also used to prevent "fuel" buildup, the highly flammable plants such as grasses and scrub under the canopy that could burn easily in a wildfire. Most prescribed burn intervals are about every 2–5 years, which allows the ecosystem to regenerate after the burn. Much of the South Florida pine rockland ecosystem is highly fragmented and has not been burned because of the proximity to buildings. Risks such as smoke, air quality, and residual particulate matter in the environment pose safety issues for controlled burns near homes and businesses.


Diseases and pests


Fusiform rust

Starting in the late 1950s, the emergence of
fusiform rust Fusiform means having a spindle-like shape that is wide in the middle and tapers at both ends. It is similar to the lemon-shape, but often implies a focal broadening of a structure that continues from one or both ends, such as an aneurysm on a ...
on Southeastern pine trees including slash pine, loblolly pine, and longleaf pine, led to massive tree mortality within the pine industry. This obligate parasitic pathogen is notorious for infecting young trees in newly planted areas within the first few years of growing. The pine industry was still rather new at the time of this initial outbreak, so many newly planted forests had large-scale mortality because the trees were not yet old enough to be resilient to the disease or harvested. Florida’s pine industry in particular was booming with an increase in plantation acreage from in 1952 to upwards of in 1990. Because of the complicated lifecycle of ''Cronaritum quercuum'' f. sp. ''fusiforme'', the fungal causal agent of fusiform rust, the management strategies of pruning diseased stems, reducing fertilization, and discarding infected seed were not sufficient to prevent million-dollar annual loses.
Rust Rust is an iron oxide, a usually reddish-brown oxide formed by the reaction of iron and oxygen in the catalytic presence of water or air moisture. Rust consists of hydrous iron(III) oxides (Fe2O3·nH2O) and iron(III) oxide-hydroxide (FeO(OH ...
pathogens are difficult to manage because of their complicated reproductive lifecycles. ''C. querecuum'' f. sp. ''fusiforme'' is
heteroecious A heteroecious parasite is one that requires at least two hosts. The ''primary host'' is the host in which the parasite spends its adult life; the other is the ''secondary host''. Both hosts are required for the parasite to complete its life cycle ...
, requiring two different plant hosts for reproduction, and is macrocyclic, meaning it contains all five spore stages typical for rust infections:
basidiospore A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are ...
s,
teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the p ...
s,
urediniospore Urediniospores (or uredospores) are thin-walled spores produced by the uredium, a stage in the life-cycle of rusts. Development ''Urediniospores'' develop in the uredium, generally on a leaf's under surface. Morphology *Urediniospores usually h ...
s, aeciospores, and spermatia. Oak trees are the secondary host for this pathogen.Gilman, E. F., & Watson, D. G. (1994). Pinus elliottii: Slash Pine. ''USDA Forest Service Fact Sheet ST-463 Google Scholar''. The primary inocula on pine are
basidiospore A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are ...
s, which infect the pine needles between March and May. The basidiospores germinate and grow into the stems of the tree where the fungus can overwinter for 4–6 months in the wood. In the fall, the spermatia form and fertilize the aceiospores in the following spring. The aceiospores are released from the pine and are the primary inocula that infect the oak trees in the following growing season. Aceiospores grow through the oak leaves producing urediniospores on the underside of the oak leaves. These
urediniospore Urediniospores (or uredospores) are thin-walled spores produced by the uredium, a stage in the life-cycle of rusts. Development ''Urediniospores'' develop in the uredium, generally on a leaf's under surface. Morphology *Urediniospores usually h ...
s can reproduce clonally, asexually, and can continue to infect oak plants as a secondary inoculum. Within two weeks of the primary
urediniospore Urediniospores (or uredospores) are thin-walled spores produced by the uredium, a stage in the life-cycle of rusts. Development ''Urediniospores'' develop in the uredium, generally on a leaf's under surface. Morphology *Urediniospores usually h ...
inoculation on the
oak tree An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...
,
teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the p ...
s are formed which germinate into
basidiospore A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are ...
s that infect the pine trees and complete the life rust cycle. Symptoms on the pine include gall formation, stem swelling, cankers, bushiness, and dieback. The cankers in the stem allow secondary fungal infections or other pests to enter the trees easily. Understanding the climate conditions that can lead to rust outbreaks is an important component for management strategies, but this was not well understood in the early decades of this epidemic. More recent information has shown that certain weather patterns such as high humidity, wet pine needles, and temperatures around for about 18 days can increase the spread of basiodiospores, so increase disease severity. The secondary hosts, oaks, are another economically and ecologically important trees in the Southeast. Therefore, eradication of the secondary host is not only not possible, but also not effective because basidiospores can travel up to a half mile, easily infecting distant pine trees. A combination of management strategies, though, such as reducing fertilization treatments (which can benefit the pathogen), planting more rust disease-resistant trees in plantations,


Pitch canker

Pitch canker, a monocyclic disease caused by the fungus ''
Fusarium circinatum ''Fusarium circinatum'' is a fungal plant pathogen that causes the serious disease pitch canker on pine trees and Douglas firs (''Pseudotsuga menziesii''). The most common hosts of the pathogen include slash pine (''Pinus elliottii''), loblolly ...
'' (previously named ''Fusarium moniliforme'' var. ''subglutinans''), was first described in 1946 by Hepting and Roth. When it was first described, disease levels were low until the 1970s, when a massive epidemic of pitch canker caused mass tree mortality in Florida slash pine. Some hypotheses suggest that the pathogen may have originated in
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and was then introduced in Florida and later transmitted to
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
on diseased seed. The pathogen has been reported in Mexico, however, high fungal diversity and low tree mortality from the disease suggests that this pathogen may have co-evolved in Mexico before being introduced to other parts of the world.Gordon, T. R. 2006. Pitch canker disease of pines. Phytopathology 96:657-659. Many reports describe the pathogen as endemic to
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
,Correll, J. C., Gordon, T. R., McCain, A. H., Fox, J. W., Koehler, C. S., Wood, D. L., & Schultz, M. E. (1991). Pitch canker disease in California: pathogenicity, distribution, and canker development on Monterey pine (Pinus radiata). ''Plant Disease'', ''75''(7), 676-682. likely because the disease was introduced a long time therefore the population has become more diverse.Gordon, T. R., Storer, A. J., & Okamoto, D. (1996). Population structure of the pitch canker pathogen, Fusarium subglutinans f. sp. pini, in California. ''Mycological Research'', ''100''(7), 850-854. By 1974, over half of the slash pine population in Florida was infected with ''Fusarium circinatum'' disease.Dwinell, David L. Barrows-Broaddus, Jane B. Kuhlman, G. E. (1985). Pitch Canker: A Disease Complex. ''Plant Disease'', ''69''(3), 270–276. In areas where the pathogen is newly introduced, the fungal population is mostly clonal, because fewer mating types are present within the population, so sexual reproduction may be lower. Pitch canker infects nearly all pine species, including longleaf pine,
shortleaf pine ''Pinus echinata'', the shortleaf pine, is a species of pine native to the southeastern United States. Description The tree is variable in form, sometimes straight, sometimes crooked, with an irregular crown. The tree reaches heights of with a ...
, and
eastern white pine ''Pinus strobus'', commonly called the eastern white pine, northern white pine, white pine, Weymouth pine (British), and soft pine is a large pine native to eastern North America. It occurs from Newfoundland, Canada west through the Great Lakes ...
. This disease continues to be a problem in nurseries, and has been reported in other countries. A major problem in Florida is that artificial replanting of pines may be contributing to high disease incidences. The disease can be passed through seed and spores, but requires open wounds to infect the tree from insect damage, mechanical damage, hail/weather damage, etc. The predominant symptoms include needle
chlorosis In botany, chlorosis is a condition in which leaves produce insufficient chlorophyll. As chlorophyll is responsible for the green color of leaves, chlorotic leaves are pale, yellow, or yellow-white. The affected plant has little or no ability to ...
and reddening of shoots (called "flagging") that later die. Cankers or lesions that form on the trunks can turn the bark yellow or dark brown and cause resin to exude. Stems may die and get crystalized in
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
-soaked lesions. Resin is generally produced in plants to protect against pathogens. Sometimes, the tissue above the canker dies, causing girdling of the stem. The severity of the disease depends on weather conditions and may require moisture and insect wounds or hail to infect the trees. Some insects such as
bark beetle A bark beetle is the common name for the subfamily of beetles Scolytinae. Previously, this was considered a distinct family (Scolytidae), but is now understood to be a specialized clade of the "true weevil" family (Curculionidae). Although the ...
s, spittle bugs,
weevil Weevils are beetles belonging to the Taxonomic rank, superfamily Curculionoidea, known for their elongated snouts. They are usually small, less than in length, and Herbivore, herbivorous. Approximately 97,000 species of weevils are known. They b ...
s, pine tip moths, and needle midges may vector the disease into the tree. ''F. circinatum'' was used to inoculate ''P. e.'' var. ''densa'' trees to try to increase resin production for extraction, but this approach was ineffective.


Other fungi

Fungus species '' Thozetella pinicola'' was found on leaf litter of ''Pinus elliottii'' in Hong Kong in 2009.


Uses

This tree is widely grown in tree plantations. It is also used in
horticulture Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
.


See also

*
Southern yellow pine In ecology and forestry, yellow pine refers to a number of conifer species that tend to grow in similar plant communities and yield similar strong wood. In the Western United States, yellow pine refers to Jeffrey pine or ponderosa pine. In the ...


References


External links

* * {{Authority control elliottii Trees of the Southeastern United States Trees of the United States