Pinus elliottii
(slash pine)

Pictures

Picture	Title	Caption	Copyright
Title 25-year-old plantation Caption A 25-year-old plantation in southwestern Louisiana where every third row has been removed by thinning. Copyright James P. Barnett	25-year-old plantation	A 25-year-old plantation in southwestern Louisiana where every third row has been removed by thinning.	James P. Barnett
Title 13-year-old plantation Caption A 13-year-old plantation that was precommercially thinned to about 1200 trees/ha at age 5 years. Copyright James P. Barnett	13-year-old plantation	A 13-year-old plantation that was precommercially thinned to about 1200 trees/ha at age 5 years.	James P. Barnett
Title 19-year-old plantation Caption An unthinned plantation in central Louisiana at age 19 years. Copyright James P. Barnett	19-year-old plantation	An unthinned plantation in central Louisiana at age 19 years.	James P. Barnett
Title 5-year-old plantation Caption A 5-year-old plantation established on a site in southwestern Louisiana with a high water table. Copyright James P. Barnett	5-year-old plantation	A 5-year-old plantation established on a site in southwestern Louisiana with a high water table.	James P. Barnett
Title Plantation Caption Plantation, Tweefontein, South Africa. Copyright Environmentek, CSIR, South Africa	Plantation	Plantation, Tweefontein, South Africa.	Environmentek, CSIR, South Africa
Title Needles and cone Caption Copyright Gerrit van Wyk	Needles and cone		Gerrit van Wyk
Title Seeds Caption Mixed lot of P. elliottii seeds with the wings removed. Copyright James P. Barnett	Seeds	Mixed lot of P. elliottii seeds with the wings removed.	James P. Barnett
Title Cronartium fusiforme infestation. Caption Five-year-old P. elliottii with a severe infestation of Cronartium fusiforme. Such heavy infestation will cause mortality. Copyright James P. Barnett	Cronartium fusiforme infestation.	Five-year-old P. elliottii with a severe infestation of Cronartium fusiforme. Such heavy infestation will cause mortality.	James P. Barnett
Title Resin collection Caption Trees being worked for naval stores production. This use is declining because other sources of the resin products have become more readily available. Copyright James P. Barnett	Resin collection	Trees being worked for naval stores production. This use is declining because other sources of the resin products have become more readily available.	James P. Barnett

Identity

Preferred Scientific Name

• Pinus elliottii Engelm.

Preferred Common Name

• slash pine

Variety

• Pinus elliottii var. densa Little & Dorman
• Pinus elliottii var. elliottii

Other Scientific Names

• Pinus heterophylla

International Common Names

• English: swamp pine
• Spanish: pino amarillo; pino macho
• French: pin d'Elliott
• Chinese: shi di song

Local Common Names

• Brazil: pinheiro
• Germany: Elliotts Kiefer
• Italy: pino di Elliott
• South Africa: basden
• USA: Cuban pine; longleaf pitch pine; pitch pine; swamp pine; yellow slash pine

EPPO code

• PIUEL (Pinus elliottii)

Trade name

• American pitch pine
• slash pine
• yellow pine

Summary of Invasiveness

Pinus elliottii var. elliottii, commonly known as slash pine, is an important timber tree species native to the lower coastal plain within the southeastern USA. Trees grow up to 36 m in height and 0.9 m in diameter, producing a long, clear bole. Because of its rapid early growth and production of highly valuable wood products, it has been widely introduced into other countries. As an exotic, it is used in Africa, especially southern Africa, and in Australia and South America for various products, ranging from lumber to pulpwood. In Brazil, P. elliottii makes an important contribution to the resin production industry. Where it has been planted outside its native range it has escaped from cultivation and become naturalized, and in some areas become an aggressive invasive weed, threatening native species and ecosystems. Trees can form dense stands that shade out other plant species.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Gymnospermae
•                 Class: Pinopsida
•                     Family: Pinaceae
•                         Genus: Pinus
•                             Species: Pinus elliottii

Notes on Taxonomy and Nomenclature

The genus Pinus was first classified on evolutionary characteristics by Shaw (1914), and most taxonomists have since followed his general separation of the genus into two groups: section/subgenus Haploxylon or Strobus, and Diploxylon or Pinus, commonly called the "soft" (or "white") and "hard" pines, respectively. This division is based on the presence of one or two vascular bundles in the leaves, respectively. Shaw's original subdivision of these groups has been reworked by different authorities (e.g., Pilger, 1926; Duffield, 1952; Ferre, 1965; Landry, 1974, 1978), but botanists in recent years have generally recognized the classification described by Little and Critchfield (Little and Critchfield, 1969; Critchfield, 1986), who recognize three subgenera: (1) Ducampopinus (one species: P. krempfii), (2) Strobus and (3) Pinus. 

Pinus elliottii belongs to subgenus Pinus, section Australes. Leaves in this group occur in groups of 2-3 in fascicles, with biform hypodermis, endodermis with a thin outer cell wall, and with internal resin ducts (Little and Critchfield, 1969). There are two varieties of P. elliottii: the typical variety (var. elliottii) and the South Florida slash pine (var. densa).

Stephen Elliott in 1824 was the first botanist to distinguish slash pine, naming it P. taeda var. heterophylla. In a revision of the genus Pinus, George Engelmann (1880) published a detailed description of the new species, naming it P. elliottii in honour of Elliott, whose name for it was cited as a synonym. The name P. elliottii was, however, short-lived, with slash pine being named P. caribaea, P. palustris, P. cubensis and P. heterophylla at various times and by various authors. A study by Little and Dorman (1952) ended the confusion and showed that the hard pines with shiny brown cones generally classified as P. caribaea were in fact three geographically separated taxa, viz. P. caribaea with a tropical distribution in the West Indies and Central America, and P. elliottii vars. elliottii and densa from the southeastern USA and southern Florida, respectively.

The common name slash pine is derived from the local name ‘slash’ given to the overgrown swamps that were its original habitat.

Description

P. elliottii var. densa differs in a number of ways from var. elliottii (Little and Dorman, 1954); it is characterized by a grass-like, almost stemless stage that lasts for 2 to 6 years. It has a restricted range, a less desirable tree form than var. elliottii, and is more difficult to regenerate. As a result, almost all information available relates to var. elliottii.

P. elliottii var. elliottii trees range from 18 to 30 m in height and average about 0.6 m in diameter. Large trees are sometimes 36 m tall and 0.9 m in diameter. The trunk is straight and erect with a long clear bole. Self-pruning of the branches is common and results in a relatively short, narrow, ovoid crown with spreading and ascending limbs. P. elliottii develops an extensive lateral root system and a moderate taproot. Maximum length of the lateral roots may be double the tree height.

The needles are 18 to 25 cm long in fascicles of 2 and 3 on the same tree, dark glossy green, and tufted at the ends of tapering branches. They extend back some distance along the branch and are persistent until the end of the second season. Resin canals number 2 to 10 and are mostly internal. The fascicle sheath is usually 1.25 cm or less in length.

P. elliottii is monoecious and wind pollinated. The cones are 10 to 15 cm in length, ovoid-conic, and sessile. They usually persist on the tree until the following summer. Cones are reddish brown, lustrous, and are armed with a sharp spine. Seeds are about 0.6 cm long, dark brown mottled with black, with wings about 2.5 cm long that are thin and translucent. 

Plant Type

Distribution

P. elliottii is native to the southeastern USA, from southern South Carolina west to southeastern Louisiana and south to the Florida Keys. The range of P. elliottii var. elliottii extends over 8° latitude and 10° longitude. The latitude covered in the southeastern USA ranges from 27° to 35°N. P. elliottii var. densa has a more restricted native range, occurring only in the southern portion of Florida.

P. elliottii has been introduced into other US states, including Hawaii, as well as other countries for timber production. Large scale introductions have occurred in Brazil, Chile, Argentina, Venezuela, China, South Africa, New Zealand and Australia. In most of these situations, P. elliottii is an adequate seed producer for continued natural or artificial regeneration.

In Australia, P. elliottii is widely cultivated in forestry plantations and is also commonly grown as an ornamental and street tree. Widely naturalized in southeastern Queensland and less so in the coastal districts of northern New South Wales, it is classed as an environmental weed of roadsides, urban bushland, open woodlands, grasslands, disturbed sites and waste areas in subtropical regions (Queensland Government, 2011).

In South Africa, it is also widely cultivated for timber, and has invaded forest margins and grassland in Mpumalanga. It has also invaded lower-altitude, higher-rainfall areas in Zimbabwe (Coates Palgrave and Coates Palgrave, 2002).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The naturalized range of P. elliottii has been extended by planting into northern Alabama, Georgia, southern Tennessee and western Louisiana. It was introduced into Kentucky, Virginia and eastern Texas in the 1900s. It now regenerates naturally in these areas.

Of the Pinus species introduced and tested for timber production in the Hawaiian Islands in the 20th century, P. elliottii has been one of the few to thrive in the warm climate and exists in forest reserves on six of the islands, most significantly on Kauai and Molokai; in the latter case, 52,000 trees were planted there between 1954 and 1960 (Skolmen, 1980). Naturalized populations are known in the native forests near the original plantings on Kauai, Molokai and Oahu (Evenhuis and Eldredge, 2012).

In South Africa, P. elliottii has been grown as a commercial forest species in the Eastern Transvaal zone since the 1920s (Darrow, 1992).

In Australia, trials of conifers in infertile coastal lowlands of south-east Queensland were initiated in the 1920s (Nikles, 1996). P. elliottii was introduced to New South Wales in 1921 and to Queensland in 1925, with large scale plantings in Queensland north of Brisbane in the 1930s. Self-sown trees were observed soon after, but were first documented in the 1960s (Richardson and Higgins, 1998).

Seed of P. elliottii was first introduced to Brazil in 1936 by the Instituto Florestal in São Paulo, with large scale plantations being established and investigated for forestry potential in the south of the country in 1955. Due to its ease of management and fast growth, it was used in large reforestation programmes with P. taeda between 1955 and 1964 (Bracelpa, 2010). Subsequently, invasions of P. elliottii have occurred into grassland savannahs, wetlands and coastal dune systems.

Risk of Introduction

As an important timber tree, P. elliottii is likely to continue to be extensively planted in forest plantations throughout its native and introduced ranges.

Habitat

The common name for P. elliottii, slash pine, refers to the swampy overgrown ground known as ‘slashes’ that constitute its native habitat. It grows throughout the flatwood areas of north Florida and south Georgia, along streams and edges of swamps and bays where there is ample soil moisture. It colonizes abandoned fields and, with protection from fire, it has spread to drier sites, replacing P. palustris in mixed stands. P. elliottii var. densa also occurs on dry sites, either sandy flat lands or limestone outcrops.

Young P. elliottii trees are quite susceptible to injury by fire and mature stands can be damaged by fires in areas of heavy fuel accumulations. Trees may suffer severe damage from ice glaze (Derr and Enghardt, 1957) and are subject to windthrow on shallow soils and wherever a strong root system has not developed.

Habitat List

Hosts/Species Affected

P. elliottii can form dense stands that shade out other plant species. In Australia it is particularly common in eucalypt woodland areas. In Hawaii, Poa mannii, a federally listed endangered endemic grass growing only on the island of Kauai, is threatened by competition from P. elliottii and other alien invasive plant species (US Fish and Wildlife Service, 2010).

Biology and Ecology

Genetics

P. elliottii is a diploid species with a chromosome number of 2n = 24 (Doudrick et al., 1995).

Reproductive Biology

A 3-year period is required in P. elliottii from the time strobili are initiated until the time the seeds mature. The primordia of new strobili are detectable in late spring. These cone initials overwinter as buds the first year. Early in the second year the male strobili become purple and 5 cm long when pollen is shed in late January or February. The female strobili appear on stalks in the upper crown and are about 2.5 cm long and red to purple at the time of pollination. Soon after pollination, the pollen tube ceases to grow and appears to remain in a quiescent state for the summer and winter (Stanley, 1958). During the third growing season fertilization occurs some 12 to 14 months after pollination. The cones enlarge and seeds mature during the third summer.

Physiology and Phenology

Young P. elliottii trees are quite susceptible to injury by fire until they are 3.0 to 4.6 m tall and have bark thick enough to insulate the cambium from lethal temperatures (McCulley, 1950; Gruschow, 1952). Mature stands can be damaged by fires in areas of heavy fuel accumulations. Trees may suffer severe damage from ice glaze (Derr and Enghardt, 1957) and are subject to windthrow on shallow soils and wherever a strong root system has not developed.

Longevity

Although short-lived compared to other pines, slash pines may live as long as 200 years, growing to 18-30.5 m in height.

Associations

In its native range, P. elliottii is associated with coastal plant communities, coniferous and mixed forests, and wetlands. Plant associates include other canopy trees such as live oak (Quercus virginiana) and cabbage palm (Sabal palmetto), and understorey plants such as bluestems (Andropogon spp.), saw palmetto (Serenoa repens) and gallbery (Ilex glabra). Animals associated with slash pine include species that utilize it for protection and cover, as well as those that consume it. Seeds are eaten by birds and small mammals. Cattle and deer browse its seedlings, and some birds such as the bald eagle (Haliaeetus leucocephalus) and the endangered red-cockaded woodpecker (Leuconotopicus borealis) nest in slash pine trees (Hill, 2001).

Where naturalized in Australia it is particularly common in eucalypt woodlands.

Environmental Requirements

P. elliottii grows in a warm humid climate characterized by hot, wet summers, warm, dry autumns and springs, and mild, wet winters. In its native range, annual rainfall averages over 1270 mm, with approximately 915 mm falling during the 8 to 9 month growing season. The mean annual temperature is 18°C, with extremes from -18° to +41°C (Shoulders and Parham, 1983). Average monthly temperatures and temperature extremes are not important in limiting its range. However, the species' susceptibility to damage by ice storms may effectively limit its northern extension. Rainfall is also important in determining the range. 

Latitude/Altitude Ranges

Air Temperature

Rainfall

Rainfall Regime

Soil Tolerances

Soil drainage

• free
• impeded
• seasonally waterlogged

Soil reaction

• acid

Soil texture

• heavy
• light
• medium

Special soil tolerances

• infertile
• shallow

Natural enemies

Means of Movement and Dispersal

Natural Dispersal

Being winged, seeds are mostly spread short distances by wind (Queensland Government, 2011); over 90% fall within 45 m of the source tree (Cooper, 1957).

Vector Transmission

In Australia, Richardson and Higgins (1998) report that dispersal of slash pine cones by the glossy black cockatoo (Calyptorhynchus lathami) has aided the tree's spread into Eucalyptus forests.

Intentional Introduction

As an important forest tree, P. elliottii continues to be introduced to new areas for forestry purposes. 

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

Impact on Habitats

P. elliottii is classed as a Category 2 invasive species in South Africa, being a particular problem in Mpumalanga Province. Dense stands reduce water runoff and stream flow from mountain catchments, compete with native species, reduce grazing and pose a fire hazard (Invasive Species South Africa, 2015).

Studies have been carried out into P. elliottii invasions into grassland savannahs, wetlands and coastal dune systems in Brazil. Zenni and Ziller (2011) found it to be one of the most pervasive and aggressive invaders of subtropical ombrophilous forest and savannah. In coastal dunes in Florianópolis in Santa Catarina state, P. elliottii seed rain was observed to be continuous throughout the year, generating about 2 million viable seeds/ha annually which compensated for the lack of a soil seedbank. A 90% germination rate and a dense, permanent seedling bank contributed to the pine’s high level of invasiveness (Bechara et al., 2013, 2014). Abreu and Durigan (2011) observed that invasion by P. elliottii was one of the most serious threats to the remaining cerrado vegetation in São Paulo state. In one area, 22 years after the arrival of the first founding trees the savannah had become a dense pine forest, causing severe loss of native biodiversity.

In Argentina, P. elliottii has been observed invading overgrazed natural grasslands and other disturbed areas in Entre Rios, Corrientes and Misiones provinces, while in Queensland, Australia, the species has invaded heathland and open eucalypt forests (Richardson and Higgins, 1998).

Impact on Biodiversity

P. elliottii is a fast-growing tree that establishes dense stands that shade out native plants and prevent their regeneration. Invaded grasslands are transformed over time into species-poor shrublands and forests (Weber, 2003).

In Hawaii Poa mannii, a federally listed endangered endemic grass growing only on the island of Kauai, is threatened by loss and modification of habitat by alien invasive species such as P. elliottii. When it was listed in 1994, only four populations were known, containing a total of approximately 125 individuals (US Fish and Wildlife Service, 1994). From observations between 1991 and 2008, at least 13 populations are now known, with at least 100 individuals (US Fish and Wildlife Service, 2010).

P. mannii is one of three native Poa species in the Hawaiian Islands, all endemic to Kauai. It typically grows on moist vertical cliff faces or dripping, wet rock faces often on northern exposures in partial shade in lowland or montane diverse mixed mesic Metrosideros polymorpha or Acacia koa–M. polymorpha forests or shrubland at elevations of 327-1222 m (US Fish and Wildlife Service, 2003). Although described as a perennial, P. mannii appears to die back each year, with few rhizomes surviving to the following year. During this dieback period, several invasive introduced plant species have been known to overtake its cliff habitat (US Fish and Wildlife Service, 1994), competition from alien invasives, especially Erigeron karvinskianus and Lantana camara, but including P. elliottii, being a leading threat to the species’ survival. To reduce the threat to P. mannii, over 600 seeds were collected and stored, but no propagation had been carried out by 2010, and no plans for ecosystem-level restoration and management of habitats had been drawn up (US Fish and Wildlife Service, 2010).

Rolon et al. (2011) found that in the coastal plain of Rio Grande do Sul in southern Brazil, macrophyte species richness in ponds was reduced in areas invaded by P. elliottii.

Threatened Species

Risk and Impact Factors

• Proved invasive outside its native range
• Abundant in its native range
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Benefits from human association (i.e. it is a human commensal)
• Long lived
• Has high reproductive potential
• Has propagules that can remain viable for more than one year

• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Modification of fire regime
• Modification of hydrology
• Monoculture formation
• Reduced native biodiversity
• Threat to/ loss of endangered species
• Threat to/ loss of native species

• Competition - monopolizing resources
• Competition - shading

• Highly likely to be transported internationally deliberately

Uses

Economic Value

Slash pine is an important source of timber and wood pulp. Many of the products of P. elliottii forests are used in international trade. These range from solid wood products to kraft paper. The sapwood of P. elliottii is a yellowish white, while the heartwood is a reddish brown. The wood is very strong and heavy, very stiff, hard and moderately high in shock resistance. It has straight grain, is medium in texture and relatively difficult to work with hand tools (Alden, 1997). It is used for products such as sawlogs, veneer logs, poles, piling, posts, pulpwood, particleboard and chip-n-saw logs. The straightness of the bole makes P. elliottii particularly suited for poles, piling and solid wood products. Due to its rapid early growth it is also well suited for pulpwood rotations where paper is the final product. P. elliottii also yields the best quality and highest quantity of commercial turpentine of all American pitch pines (Streets, 1962).

Social Benefit

P. elliottii is also grown for its ornamental value and as a street tree. The species has been frequently used in agroforestry situations.

Environmental Services

Slash pines provide habitat and food for birds and small mammals, and are an important component of many natural communities (Hill, 2001). P. elliottii trees can be planted to stabilize the soil on eroding slopes and strip mine spoil banks.

Uses List

Environmental

• Agroforestry
• Erosion control or dune stabilization
• Revegetation

Fuels

• Fuelwood

General

• Ornamental

Materials

• Fibre
• Gum/resin
• Miscellaneous materials
• Wood/timber

Medicinal, pharmaceutical

• Traditional/folklore

Wood Products

Boats

Containers

• Boxes
• Crates

Pulp

• Long-fibre pulp

Roundwood

• Building poles
• Posts
• Transmission poles

Sawn or hewn building timbers

• Bridges
• Carpentry/joinery (exterior/interior)
• Engineering structures
• Flooring
• For heavy construction
• For light construction

Veneers

Wood-based materials

• Composite boards
• Laminated veneer lumber
• Laminated wood
• Plywood

Similarities to Other Species/Conditions

In the southeastern USA, P. elliottii can be readily distinguished from the other two southern yellow pines, P. palustris and P. taeda; from P. palustris by its thinner branchlet tips (1.75 vs. 2.5 cm diameter) and from P. taeda by having cones on a short stalk rather than no stalk (sessile) (Christman, 2011).

P. elliottii can be confused with several other naturalized pine species in Australia, but can be differentiated from them on the basis of cone and needle morphology. P. elliottii has relatively large symmetrical cones (7-20 cm long) borne on short stalks, and relatively long needles (15-30 cm) borne in groups (fascicles) of two or three (usually two). In P. radiata the cones are of similar size and borne on short curved stalks, and the needles are shorter (8-15 cm) and usually borne in groups of three. P. caribaea has relatively small symmetrical cones (5-12 cm) borne on short stalks, and relatively long needles (15-30 cm), usually in groups of three. In P. pinaster cones are large (9-20 cm long), slightly symmetrical or asymmetrical and sessile, while needles are long (10-30 cm) and usually borne in groups of two. P. halepensis has relatively small cones (5-14 cm) on curved stalks 10-30 mm long. Its needles are short (4-12 cm) and borne in groups of two (Queensland Government, 2011).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Where P. elliottii is growing as a weed, small individual plants can be removed by hand pulling, particularly in moist soil, taking care to remove the roots and applying mulch to discourage regrowth. Larger individuals may be dug out with a mattock or similar tool. Cutting and painting stumps with glyphosate can be done, but the herbicide mixture must be applied within 15 seconds of making the cut. In Australia, slash pines are sometimes used by native fauna as habitat trees (although they have little food value) so should be checked before felling. It is recommended that slash pines be replaced with native trees such as Eucalyptus to provide food and shelter for wildlife (Moreton Bay Regional Council, 2015).

References

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Alden HA, 1997. Softwoods of North America. General Technical Report - Forest Products Laboratory, USDA Forest Service, No. FPL-GTR-102:151 pp.; 216 ref.

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Barnett JP; Vozzo JA, 1985. Viability and vigor of slash and short-leaf pine seeds after 50 years of storage. Forest Science, 31(2):316-320; 11 ref.

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Bechara FC; Reis A; Bourscheid K; Vieira NK; Trentin BE, 2013. Reproductive biology and early establishment of Pinus elliottii var. elliottii in Brazilian sandy coastal plain vegetation: implications for biological invasion. Scientia Agricola, 70(2):88-92. http://www.scielo.br/sa

Bechara FC; Reis A; Trentin BE, 2014. Biological invasion of Pinus elliottii var. elliottii in Rio Vermelho State Park, Florianópolis, SC. (Invasão biológica de Pinus elliottii var. elliottii no Parque Estadual do Rio Vermelho, Florianópolis, SC.) Floresta, 44(1):63-72. http://ojs.c3sl.ufpr.br/ojs2/index.php/floresta/article/view/32013/22127

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Bennett FA, 1971. The role of thinning and some other problems in management of Slash Pine plantations. US For. Serv. Res. Pap. Stheast. For. Exp. Sta. No. SE-86, 1971. pp. 14. [5 ref.].

Berrios C; Menendez JM; Rodriguez M, 1987. Presence of Ips on new species of pines in the north of Matanzas province. Revista Forestal Baracoa, 17(2):113-115

Bethune JE, 1966. Performance of two Slash Pine [Pinus elliottii] varieties planted in south Florida. U.S. For. Serv. Res. Pap. Stheast. For. Exp. Sta. No. SE-24, 1966. pp. 9. [10 refs.].

Blakeslee GM; Dwinell LD; Anderson RL, 1980. Pitch canker of southern pines [Gibberella fujikuroi var. subglutinans]: identification and management considerations. Forestry Report, Southeastern Area, State and Private Forestry, USDA Forest Service, No. SA-FR 11, i + 15 pp.; 12 col. pl.; 19 ref.

Bracelpa, 2010. Pinus., Brazil: Bracelpa - Brazilian Pulp and Paper Association. http://bracelpa.org.br/bra2/?q=en/node/227

Byram TD; Lowe WJ, 1996. Forty-fourth progress report of the Cooperative Forest Tree Improvement Program. Circular Texas Forest Service, No. 296.

Carey JH, 1992. Pinus elliottii. Fire Effects Information System. Missoula, Montana, USA: USDA Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. http://www.feis-crs.org/beta/

Christman S, 2011. Pinus elliottii. Floridata Plant Encyclopedia. http://mobile.floridata.com/Plants/Pinaceae/Pinus%20elliottii/84

Ciesla WM, 2003. European woodwasp - a potential threat to North America's Conifer Forests. Journal of Forestry, 18-23.

Clarke SR; DeBarr GL; Berisford CW, 1990. Life history of Oracella acuta (Homoptera: Pseudococcidae) in loblolly pine seed orchards in Georgia. Environmental Entomology, 19(1):99-103

Clutter JL; Jones EP, 1980. Prediction of growth after thinning in old-field slash pine plantations [in southern USA]. Research Paper, Southeastern Forest Experiment Station, USDA Forest Service, No. SE-217, 19 pp.; 9 ref.

Coates Palgrave K; Coates Palgrave M, 2002. Palgrave's trees of southern Africa. New edition. Cape Town, South Africa: Struik Publishers, 1212 pp.

Cooper RW, 1957. Silvical characteristics of Slash Pine (Pinus elliottii Engelm. var elliottii). Station Paper. Southeastern Forest Experiment Station, No. 81:13.

Cremiere L; Ehrhart Y, 1990. Thirty years of introduced pine species in New Caledonia. [30 ans d'introduction d'espèces de pins en Nouvelle-Calédonie.] Bois et Forêts des Tropiques, No. 223, 3-23; 16 ref.

Critchfield WB, 1986. Hybridization and classification of the white pines (Pinus section Strobus). Taxon, 35(4):647-659; 44 ref.

Darrow WK, 1992. Heart shake in South African-grown Pinus elliottii. Wood and Fiber Science, 24(3):241-251.

Derr HJ; Enghardt H, 1957. Some forestry lessons from hurricane Audrey. Southern Lumberman, 195 (2441):142-144.

Derr HJ; Mann WF; Jr, 1971. Direct-seeding Pines in the South. Agric. Handbook No. 391. USA: USDA-Forest Service.

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Distribution References

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GBIF, 2015. Global Biodiversity Information Facility. http://www.gbif.org/species

Little E L Jr, Dorman K W, 1954. Stn. Pap. Steast. For. Exp. Sta, ii + 82.

Lohrey RE, Kossuth SV, 1990. (Pinus elliottii Engelm. Slash pine). In: Silvics of North American, Volume 1, Conifers. US Department of Agriculture Forest Service, Agriculture Handbook, 644 338-347.

Moore LM, Walker Wilson JD, 2006. USDA-NRCS Plant Guide: slash pine, Pinus elliottii Engelm., Baton Rouge, LA, USA: USDA-NRCS National Plant Data Center. 4 pp. http://plants.usda.gov/plantguide/pdf/pg_piel.pdf

Nikles D G, 1996. The first 50 years of the evolution of forest tree improvement in Queensland. In: Tree improvement for sustainable tropical forestry. QFRI-IUFRO Conference, Caloundra, Queensland, Australia, 27 October-1 November 1996. Volume 1. [Tree improvement for sustainable tropical forestry. QFRI-IUFRO Conference, Caloundra, Queensland, Australia, 27 October-1 November 1996. Volume 1.], [ed. by Dieters M J, Matheson A C, Nikles D G, Harwood C E, Walker S M]. Gympie, Australia: Queensland Forestry Research Institute. 51-64.

Ohnuki I, Ujiie T, Kajigaki J, Yamagaki K, Nagao K, 2003. The forests and forestry situation in the Eastern Region of Paraguay and an afforestation plan (2). Tropical Forestry. 17-24.

PROTA, 2015. PROTA4U web database., [ed. by Grubben GJH, Denton OA]. Wageningen, Netherlands: Plant Resources of Tropical Africa. http://www.prota4u.info

Queensland Government, 2011. Slash pine, Pinus elliottii. In: Weeds of Australia, Biosecurity Queensland Edition, Brisbane, QLD, Australia: Queensland Government. http://keyserver.lucidcentral.org/weeds/data/080c0106-040c-4508-8300-0b0a06060e01/media/Html/Pinus_elliottii.htm

Rolon A S, Rocha O, Maltchik L, 2011. Does pine occurrence influence the macrophyte assemblage in Southern Brazil ponds? Hydrobiologia. 157-165. http://springerlink.metapress.com/content/1573-5117/ DOI:10.1007/s10750-011-0813-2

Simberloff D, Nuñez M A, Ledgard N J, Pauchard A, Richardson D M, Sarasola M, Wilgen B W van, Zalba S M, Zenni R D, Bustamante R, Peña E, Ziller S R, 2010. Spread and impact of introduced conifers in South America: lessons from other southern hemisphere regions. Austral Ecology. 35 (5), 489-504. http://www.blackwell-synergy.com/loi/aec DOI:10.1111/j.1442-9993.2009.02058.x

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysimple.aspx

Yoshida M, Tsuzuki K, Satake K, 1981. On the growing process of loblolly pine and slash pine planted in the Shikoku region. Part 1. Height growth and preparation of the volume tables for standing trees. Bulletin, Forestry and Forest Products Research Institute, Japan. 1-35.

Zenni R D, Ziller S R, 2011. An overview of invasive plants in Brazil. Revista Brasileira de Botânica. 34 (3), 431-446. http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0100-84042011000300016&lng=en&nrm=iso&tlng=en DOI:10.1590/S0100-84042011000300016

Links to Websites

Contributors

13/08/15 Invasive Species Compendium sections added by:

Andrew Praciak, CABI, UK

Distribution Maps