Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Prosopis glandulosa
(honey mesquite)

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Datasheet

Prosopis glandulosa (honey mesquite)

Summary

  • Last modified
  • 01 October 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Host Plant
  • Preferred Scientific Name
  • Prosopis glandulosa
  • Preferred Common Name
  • honey mesquite
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • P. glandulosa has been widely introduced and planted as a fuel and fodder tree. Seed are spread widely by grazing animals from established plantations or single trees around houses or water-holes, and will pers...

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Pictures

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PictureTitleCaptionCopyright
Typical shrubby form of invading P. glandulosa alongside a cultivated field in Texas, USA.
TitleShrubby form
CaptionTypical shrubby form of invading P. glandulosa alongside a cultivated field in Texas, USA.
CopyrightN.M. Pasiecznik
Typical shrubby form of invading P. glandulosa alongside a cultivated field in Texas, USA.
Shrubby formTypical shrubby form of invading P. glandulosa alongside a cultivated field in Texas, USA.N.M. Pasiecznik
P. glandulosa as an amenity tree in Texas, USA.
TitleAmenity tree
CaptionP. glandulosa as an amenity tree in Texas, USA.
CopyrightN.M. Pasiecznik
P. glandulosa as an amenity tree in Texas, USA.
Amenity treeP. glandulosa as an amenity tree in Texas, USA.N.M. Pasiecznik

Identity

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Preferred Scientific Name

  • Prosopis glandulosa Torr.

Preferred Common Name

  • honey mesquite

Other Scientific Names

  • Algarobia glandulosa Torr. (Cooper)
  • Neltuma constricta (Sarg.) Britton & Rose
  • Neltuma glandulosa (Torr.) Britton & Rose
  • Neltuma neomexicana Britton
  • Prosopis chilensis var. glandulosa Torr. (Standl.)
  • Prosopis glandulosa var. constricta Sarg.
  • Prosopis glandulosa var. glandulosa Torr. (Cockerell)
  • Prosopis glandulosa var. torreyana Benson
  • Prosopis juliflora var. glandulosa (Torr.) Cockerell
  • Prosopis juliflora var. torreyana L. D. Benson
  • Prosopis odorata Torr. & Frem.

International Common Names

  • English: algaroba; mesquite; western honey mesquite
  • Spanish: mezquite
  • Arabic: mesquite; mesquite wood

Local Common Names

  • Argentina: algarrobo
  • Chile: algarrobo
  • Germany: Mesquitebaum, Nektar-
  • Haiti: bayahonda
  • Mexico: algaroba; mesquite; mezquite
  • Peru: algarrobo
  • USA/Hawaii: kiawe

EPPO code

  • PRCJG (Prosopis juliflora var. glandulosa)
  • PRCJT (Prosopis juliflora var. torreyana)
  • PRSSGL (Prosopis glandulosa)

Summary of Invasiveness

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P. glandulosa has been widely introduced and planted as a fuel and fodder tree. Seed are spread widely by grazing animals from established plantations or single trees around houses or water-holes, and will persist for long periods in the seed bank. It has shown itself to be a very aggressive invader, especially in sub-tropical arid and semi-arid natural grasslands, both in its native range and where introduced. It is a nitrogen-fixing species and very drought and salt tolerant, rapidly out-competing other vegetation. Thorniness and a bushy habit enable it to quickly block paths and make whole areas impenetrable. Invasion in the native range generally involves an increase in plant density rather than an increase in its range. P. glandulosa is a declared noxious weed in Australia and South Africa, and the genus as a whole is regulated in several other countries. It is also reported as invasive in other southern African countries, notably Botswana and Namibia where it is known to hybridise with P. velutina, also in Australia, though P. glandulosa tends to dominate. In terms of ecology, uses, management and control, P. glandulosa and P. velutina can be effectively treated together, as a species complex.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Fabales
  •                         Family: Fabaceae
  •                             Subfamily: Mimosoideae
  •                                 Genus: Prosopis
  •                                     Species: Prosopis glandulosa

Notes on Taxonomy and Nomenclature

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P. glandulosa is used here in its original, restricted and certainly biological sense, re-established by Burkart (1940) and accepted by Benson (1941) and Johnston (1962). The all-embracing, collective P. juliflora concept of Bentham (1875) including P. glandulosa and several other species was maintained by other botanists for over a century and, although currently rejected by most taxonomists and researchers, it is still used occasionally to this day. Much of the literature from the USA has followed Bentham's classification of North American species as varieties of P. juliflora and great care must be taken when referring to old literature, with observation of authority and, more importantly, geographic origin required to ascertain the correct species being described. It can be said with certainty that all references to P. juliflora from the USA and northern Mexico prior to the publication of the accepted monograph (Burkart, 1976), and many from after this date, refer to either P. glandulosa or P. velutina. These two species are very closely related, and although it was hoped that the application of molecular techniques would clearly separate them, this has not been the case. Bessaga et al. (2000) separated them using isoezymes but not using RAPDs, and using only isoenzymes, Bessega et al. (2005) concluded that the two species originated from different ‘founder’ (speciation) events thus confirming their differences. However, using seed storage proteins, Freeman and Snyder (2005) were unable to differentiate between P. glandulosa var. glandulosa,P. glandulosa var. torreyana and P. velutina, amongst the 13 Prosopis species that were identifiable, confirming that they are very closely related indeed, which questions the conclusion of Bessega et al. (2005). P. glandulosa is certainly it is very closely related to P. velutina, which has a small native range and is largely sympatric in areas where both P. glandulosa var. glandulosa (to the east) and P. glandulosa var. torreyana (to the west) are present. All taxa hybridise readily and there appears to be some introgression in sympatric areas. Molecular analysis groups these three taxa together, confirming the close genetic relationships between them, supporting a reconsideration of them as a single species or species complex, Where introduced and invasive, the distinction is even less clear, adding to arguments that call for a reclassification.

This taxonomic confusion begun by Bentham (1875) has led to further problems when referring to literature on P. juliflora, particularly where introduced, caused by the simplification of binomials of introduced material by excluding the variety and authority. For example, when P. juliflora var. glandulosa (Torrey) Cockerell was introduced from the USA or Mexico before the 1970s, it was often called simply P. juliflora, when the seed was actually P. glandulosa Torrey. Initial misidentification without correction can be repeated continually leading many foresters and researchers to believe that a certain leaf and tree morphology is characteristic of one species when it can be seen by a trained eye to clearly belong to another. Several examples are widely quoted and have contributed to continued misidentification (Pasiecznik et al., 2004).

However, one study in Botswana by Muzila et al. (2011) and cited in Mosweu et al. (2013) should be ignored. Using morphological parameters, they proposed numerous hybrids which cannot possibly exist, including an inter-generic hybrid between Acacia and Prosopis species, and several between the known tetraploid P. juliflora, and diploid species. This does highlight, however, the very large morphological variation seen in Prosopis species, and that without clear reference material, can easily lead to misidentifications.

For accurate assessment, molecular techniques must be applied, and this will clarify the taxonomic confusion in coming years, and a further revision of the genus is to be expected.

The generic common name for Prosopis is 'mezquite' in Mexico and 'mesquite' in the USA, originating from the original Nahuatl Indian name from Mexico, 'misquitl' meaning 'bark that tans'. Mesquite is also the most common name for Prosopis in the English language, for example, in many areas where it has been introduced in Asia, Australia and Africa. The common English name for P. glandulosa is the more specific 'honey mesquite', used throughout the USA, with the closely related P. velutina known as ‘velvet mesquite’.

Description

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P. glandulosa exhibits high levels of variability in morphological characters. Variations are observed principally in native populations. In invading populations, clinal variations are obscured because of the rapid and widespread dispersal of diverse genetic material by humans and animals over a range of site and climatic conditions. The following description is adapted from Burkart (1976).

P. glandulosa is a tree up to 9 m tall, with a trunk diameter up to 1 m, though larger specimens are recorded. Spines are axillary, uninodal, 1-4.5 cm long, mostly solitary, sometimes very few, solitary or paired, sometimes with solitary and paired thorns on different nodes of the same twig. Leaves bipinnate, glabrous, 1-2 pairs of pinnae per leaf, up to 15 cm long. Pinnae 6-17 cm long, each with 6-17 pairs of leaflets. Leaflets distant on the rachis, 2-6.3 cm long by 1.5-4.5 mm broad, linear or oblong, 5-15 times as long as broad, obtuse, glabrous, subcoriaceous, prominently veined below. Flowers yellow, racemes 5-14 cm long, multiflorous, petals 2.5-3.5 mm long, ovary stalked, villous. Legume straight, 8-20 cm long by 0.7-1.3 cm broad, rarely subfalcate, compressed to subterete, submoniliform, glabrous, straw coloured or tinged with violet, short-stalked, with strong, varyingly acuminate. There are 5-18 seeds per pod, seeds 6-7 mm long, oblique to longitudinal. P. glandulosa var. torreyana has a similar habit to P. glandulosa var. glandulosa but with generally shorter pinnae and shorter leaflets that are less distant on the rachis. P. glandulosa var. prostrata is similar in foliar and floral morphology to var. glandulosa but differs in its habit, being generally a low-growing shrub.

Plant Type

Top of page Broadleaved
Perennial
Seed propagated
Shrub
Tree
Woody

Distribution

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P. glandulosa is native to North America, in northern Mexico and the southern USA from the Pacific coast to the Gulf of Mexico. However, the two varieties have mostly separate ranges. P. glandulosa var. torreyana is native to the western parts, common in California, Nevada and Arizona and neighbouring Mexico states, whereas P. glandulosa var. glandulosa is native to the eastern parts, very common in Texas and north-eastern Mexico, but also in New Mexico and occasionally in Arizona. A very prostrate form that rarely even becomes a shrub was given varietal rank by Burkart, and P. glandulosa var. prostrata is much less common, found only in parts of Texas, USA, and Tamaulipas, Mexico.

Although very widely introduced around the world, P. glandulosa is most common as an invasive weed in Australia and southern Africa. P. glandulosa is a generally subtropical species, as is P. velutina, in contrast to P. juliflora and P. pallida which are frost-sensitive tropical species. It is for this reason that P. glandulosa failed or performed relatively poorly in many species introduction trials in tropical climates.

Distribution Table

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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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasivePlantedReferenceNotes

Asia

BahrainPresent Planted CABI, 2005
IndiaPresentPresent based on regional distribution.
-HaryanaPresentIntroduced Not invasive Planted Tewari et al., 2000
-Indian PunjabPresentIntroduced Not invasive Planted Tewari et al., 2000
-MaharashtraPresent, few occurrencesIntroduced Not invasive Planted Tewari et al., 2000
-RajasthanPresent, few occurrencesIntroduced Not invasive Planted Tewari et al., 2000
-Uttar PradeshPresent, few occurrencesIntroduced Not invasive Planted Tewari et al., 2000
IsraelPresent, few occurrencesIntroduced Not invasive Planted Pasiecznik et al., 2001
JordanPresent, few occurrencesIntroduced Not invasive Planted Harris et al., 2003
KuwaitPresent Planted CABI, 2005
MyanmarPresent, few occurrencesIntroduced Not invasive Planted Burkart, 1976
PakistanPresent, few occurrencesIntroduced Not invasive Planted Burkart, 1976
QatarPresent Planted CABI, 2005
Saudi ArabiaPresent, few occurrencesIntroduced Not invasive Planted Burkart, 1976
United Arab EmiratesPresent Planted CABI, 2005

Africa

BotswanaPresentIntroduced Invasive Botswana Government, 2009
Cape VerdePresent, few occurrencesIntroduced Not invasive Planted Harris et al., 1996
NamibiaPresentIntroduced Invasive Planted Poynton, 1990
South AfricaPresentIntroducedpre 1880 Invasive Poynton, 1990; Zimmermann, 1991P. glandulosa var. torreyana
SudanPresentIntroduced1917Broun and Massey, 1929; El Fadl, 1997; CABI, 2005

North America

MexicoPresentNative Invasive Johnston, 1962; Burkart, 1976
USAPresentPresent based on regional distribution.
-ArizonaPresentNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-CaliforniaWidespreadNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-ColoradoPresentNative Invasive Natural USDA-NRCS, 2007
-KansasRestricted distributionNative Invasive Natural USDA-NRCS, 2007
-LouisianaRestricted distributionNative Invasive Natural USDA-NRCS, 2007
-NevadaPresentNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-New MexicoPresentNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-OklahomaRestricted distributionNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-TexasWidespreadNative Invasive Johnston, 1962; Burkart, 1976; USDA-NRCS, 2007
-UtahRestricted distributionNative Invasive Natural USDA-NRCS, 2007

Central America and Caribbean

CubaPresentIntroduced Planted Johnston, 1962

Europe

SpainPresent, few occurrencesIntroduced Not invasive Pasiecznik, 1989; Pasiecznik and Peñalvo, 2016

Oceania

AustraliaPresentIntroduced1900sPanetta and Carstairs, 1989
-Australian Northern TerritoryPresentIntroduced Invasive Natural Panetta and Carstairs, 1989
-New South WalesPresentIntroduced Invasive Natural Panetta and Carstairs, 1989
-QueenslandPresentIntroduced Invasive Burkhart, 1976; Panetta and Carstairs, 1989
-South AustraliaPresentIntroduced Invasive Planted Panetta and Carstairs, 1989
-Western AustraliaPresentIntroduced Invasive Planted Panetta and Carstairs, 1989

History of Introduction and Spread

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There are theories pertaining to the natural spread of woody legumes in arid ecosystems, which relate specifically to P. glandulosa in North America, and for which there is a large body of literature. The first is natural invasions proposed by ecologists (e.g. Archer, 1995), who suggest that the first trees that establish on arid land are Prosopis spp. which create 'islands of succession'. These 'islands' provide ameliorated conditions which aid the establishment of an increasing number of plants, and the ecosystem will stabilize over time. The second set of theories is proposed by weed scientists, who state that species of Prosopis are among the many weedy species that have a competitive advantage over native plants and require eradication or control (e.g. Jacoby and Ansley, 1991; Zimmerman, 1991). The third theory states that woody legumes have a competitive advantage only in ecosystems where soil nitrogen levels are very low (e.g. Geesing et al., 1999) and management interventions should be aimed at minimizing the nitrogen harvested as forage, and increasing soil fertility, which will reduce the competitive advantage of species of Prosopis over other species. In addition to these, Fredrickson et al. (2006) have recently presented a new hypothesis relating spread of P. glandulosa in North America to human history, the presence of animals and changing climate over millennia.

Intentional introductions of P. glandulosa within the native range must have occurred before Mexican independence, such as the introduction from mainland Mexico of P. glandulosa trees producing sweet pods to Baja California, Mexico. Ecological studies on the unintentional spread of P. glandulosa as a pioneer colonizer into savanna grassland in their native range over the past 100-200 years have analysed the ecological interactions and vegetation succession, and modelled future scenarios (Archer, 1995). No correlation was found between years of high rainfall and the spread of P. glandulosa, resulting in uneven-aged stands. Seed germination and seedling establishment of P. glandulosa has been studied in detail in relation to soil moisture, soil type, seed depth and plant cover (Scifres and Brock, 1970; Brown and Archer, 1990). This primary invasion is followed by the invasion of other woody perennials (e.g. Acacia farnesiana, Parkinsonia aculeata). The resulting vegetation community then facilitates the decline in Prosopis numbers, preventing further establishment, and is thought to lead to a stable woodland community in 400-500 years (Archer, 1995). A recent of study over 60 years showed that P. glandulosa shrub cover and patch numbers increased from 1936 to the 1970s, then stabilized until the present day at 43% cover and 83 patches ha-1 (Goslee et al., 2003).

Elsewhere in the Americas, Prosopis was introduced into north-east Brazil. Several papers in Habit and Saavedra (1990) state that the first introduction of Prosopis into Brazil was by J.B. Griffing, who in 1942 introduced seed from New Mexico, USA (presumably therefore P. glandulosa and/or P. velutina) to the Sierra Talhada area of Pernambuco state. This was followed by a second introduction to Rio Grande do Norte by S.C. Harland in 1947 of seed from Peru, and in 1948 of seed from Sudan. Only two trees of each of these later two introductions are believed to have survived and it is stated that these provided the basis for the entire population of Prosopis in the north-east of Brazil. There are no records of either P. glandulosa or P. velutina existing today as naturalized species, so it can be assumed that the introductions from the USA did not survive.

P. glandulosa was introduced into Australia around 1900. No records exist, but Prosopis species were first introduced to the country as a tree for shade, fodder and erosion control, with major planting and possibly further introductions in the 1920s and 1930s (Csurhes, 1996). Several introductions were clearly made, with four species now naturalized in Australia (Perry, 1998). Later introductions may have come from the Americas, or possibly from India or South Africa where Prosopis species had already become naturalized. P. glandulosa var. glandulosa, P. velutina, and a hybrid between the two, are widespread in southern areas and there is evidence of introgression with P. glandulosa var. torreyana (Perry, 1998). Another hybrid contains genes from an, as yet, uncertain species but it is thought to be a P. glandulosa x P. laevigata hybrid (Panetta and Carstairs, 1989), and van Klinken et al. (2006) notes the existence of a three-way hybrid P. velutina x P. glandulosa var. glandulosa x P. pallida, thus all combinations appear possible.

There appear to be several competing histories as to the introduction of Prosopis into the Indian sub-continent, with no doubt that it first occurred in the mid- to late 1800s. Luna (1996) uses the date of 1857 as that of first introduction of P. juliflora from Mexico, and Sindh province, now in Pakistan, is also often given as the region of first introduction and Mexico the origin. However, Reddy (1978) gives a compelling account of the request for Prosopis seed made by Lt. Col. R.H. Bedome, Conservator of Forests of Northern Circle (Madras) to the Secretary of the Revenue Department of Madras in 1876: "The Prosopis dulcis, the Prosopis pubescens and P. glandulosa - are stated to grow best on dry arid soil. They yield hard and valuable timber and also an abundance of sweet succulent pods which are used for cattle feeding and also ground into meal. It is very desirable to introduce these trees into the fuel plantations in our dry districts; and I have the honour to suggest that the British Consuls at Galveston and San Francisco should be applied to for the seed. The Prosopis juliflora is a species growing in Jamaica which I should be very glad to get seed of". This was sent to the Secretary of State and seeds arrived and were sown that same year and outplanted in 1878 (Reddy, 1978). Mohan (1940, in Raizada and Chatterji, 1954) reported the introduction of P. glandulosa var. glandulosa and other Prosopis species, but no records as to their performance are available (Raizada and Chatterji, 1954). Today, P. juliflora is the main species present, and records of the presence of naturalized P. glandulosa require confirmation. P. glandulosa has also been recorded in Myanmar (Burkart, 1976).

Much of the Prosopis in the Mediterranean zone of North Africa is likely to be P. glandulosa, P. velutina, P. chilensis (Burkart, 1976) or hybrid forms including these species. Trials showed that cold-sensitive P. juliflora was killed outright in Tunisia, whereas other species survived. Prosopis, possibly P. glandulosa, was introduced into Sudan by RE Massey from the Egyptian Department of Agriculture at Giza and from South Africa, both in 1917 (Broun and Massey, 1929; El Fadl, 1997). It is generally accepted that whatever the initial introductions may have been, subsequent introductions of P. juliflora from an unknown source are now the dominant 'common mesquite'. Records of introductions into South Africa are more complete. P. glandulosa var. torreyana was introduced into South Africa no later than 1880 and several times since (Poynton, 1990). Probably the source of much of the Prosopis to arrive in South Africa was the introduction of 23 seed lots from USA/Hawaii and Mexico from 1897 to 1916. Although they were all called P. juliflora, they almost certainly contained P. velutina and varieties of P. glandulosa. There are numerous other introductions of P. glandulosa around the world as a species included in research trials though what records exist show that most of these, especially in tropical regions, did not survive or performed very poorly.

Risk of Introduction

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P. glandulosa was widely introduced around the world intentionally, due to its value as a fuel/fodder species and also an ornamental in some regions. The seed is still available from mail order companies. However, its infamy as an invasive species has led to several governments banning the importation of seed, and the risk of further introduction is perceived as low. P. glandulosa is a declared noxious weed in Australia and South Africa, and the genus as a whole is regulated in several other countries.

Habitat

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P. glandulosa is principally a weed of grasslands both in the native ranges in Mexico and the USA, and where introduced and widespread in parts of Australia and South Africa. It is also known to invade watercourses, roadsides and disturbed areas. Harvard (1884, in Dahl, 1982) noted of P. glandulosa in the USA that "there is hardly any soil, if it is not habitually damp, in which mesquite cannot grow; no hill too rocky or broken, no flat too sandy or saline, no dune too shifting... to entirely exclude it". It has a broad ecological amplitude and is adapted to a very wide range of soils and site types from sand dunes to cracking clays. It is generally found in areas where water and soil fertility are the principal agents limiting plant growth, and is able to survive, and even thrive, on some of the poorest land, unsuitable for any other tree species. P. glandulosa dominates in dry, or seasonally dry, watercourses or depressions, and the presence and depth of the water table is a decisive factor in the distribution, size and growth of P. glandulosa. The height of P. glandulosa trees has been used successfully to estimate the depth of the water table in the USA (Simpson, 1977). It is also shown that local topography, related to soil moisture, affects where P. glandulosa becomes established (Wu and Archer, 2005).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Secondary/tolerated habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Productive/non-natural
Rail / roadsides Secondary/tolerated habitat Harmful (pest or invasive)
Rail / roadsides Secondary/tolerated habitat Productive/non-natural
Urban / peri-urban areas Secondary/tolerated habitat Productive/non-natural
Terrestrial ‑ Natural / Semi-naturalNatural grasslands Principal habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Natural
Riverbanks Principal habitat Harmful (pest or invasive)
Riverbanks Principal habitat Natural
Rocky areas / lava flows Principal habitat Harmful (pest or invasive)
Rocky areas / lava flows Principal habitat Natural
Scrub / shrublands Principal habitat Harmful (pest or invasive)
Scrub / shrublands Principal habitat Natural
Deserts Principal habitat Harmful (pest or invasive)
Deserts Principal habitat Natural
Arid regions Principal habitat Harmful (pest or invasive)
Arid regions Principal habitat Natural
Littoral
Coastal areas Secondary/tolerated habitat Harmful (pest or invasive)
Coastal areas Secondary/tolerated habitat Natural

Hosts/Species Affected

Top of page P. glandulosa is principally a common weed in natural grasslands in both its native range and where introduced.

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Poaceae (grasses)PoaceaeMain

Biology and Ecology

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Genetics

P. glandulosa, like most Prosopis taxa, has a chromosome number of n=14 (2n=28). It is closely related to other Prosopis species from North America on the basis of flavonoids (Solbrig et al., 1977), molecular markers (Ramirez et al., 1999) and seed storage proteins (Freeman and Snyder, 2005).

Physiology and Phenology

Seeds of P. glandulosa possess an inherently high level of dormancy. The hard seed coat must be broken or weakened to allow water absorption by the seed and for germination to occur. Hard seed coats will also degrade over time and older seed that is still viable tends to germinate without pre-treatment (Pasiecznik and Felker, 1992). Freshly harvested seeds gave germination rates of 90%, assumed to be because the seed coat has not yet hardened. Seeds in their endocarp shells exhibit decreased germination, thought to be due to the endocarp impeding water uptake by the seeds. The passage of seeds through different animals has varying effects on germination, through the removal of the mesocarp or endocarp, or other mechanical or chemical factors.

All Prosopis species are also able to survive in areas with exceptionally low annual rainfall or very lengthy dry periods but only if roots are able to tap ground water or another permanent water source within the first few years. Being adapted to arid and semi-arid climates, P. glandulosa generally germinates and establishes during the brief rainy season and seedlings must be sufficiently well established to survive the first dry season. The existence of two root systems, a deep tap root to reach ground water and a mat of surface lateral roots to make use of infrequent rainfall events, puts Prosopis species firmly in the category of phreatophytes, but they show a variety of mesophytic and xerophytic characteristics depending on water availability (Mooney et al., 1977). P. glandulosa exhibits marked growth flushes of new leaves during the year, with leaf production found to occur in two short, rapid bursts, considered to be a response to insect herbivory, as growth was mostly independent of climatic variables (Nilsen et al., 1987).

The main climatic variables involved in phenological variation are temperature and rainfall. These have been shown to affect the leaf morphology of P. glandulosa in Mexico and the USA (Graham, 1960). Temperature and rainfall also affected the timing of bud break and dormancy in populations of Prosopis species collected from a wide range of sites in the USA (Graham, 1960; Peacock and McMillan, 1965; Lee and Felker, 1992). Flowering time has evolved variously, with legume production generally coinciding with the beginning of the wet season for improved water dispersal and seedling establishment, or the beginning of the dry season, ensuring increased pod consumption and seed dispersal by wild animals. Flowering times vary between species and sites and are genetically controlled. In its native range, temperate races of P. glandulosa exhibit a delay in flowering compared with more subtropical races, even when planted at the same location, but each population had distinct, synchronized flowering (Graham 1960). Lee and Felker (1992) also noted synchronized flowering of P. glandulosa over a 120 km rainfall gradient, with increased nectar production and pod yields in drier years. The synchronized mass blooming of Prosopis species, irrespective of annual rainfall, was viewed as being due to their phreatophytic nature, with species able to produce flowers and seeds even during dry seasons (Simpson, 1977). Variation in the onset of flowering can be expected between populations of all species due to climatic variation within existing ranges. Flowering is also variable within and between trees of the same population.

Reproductive Biology

Anthesis is protogynous (Burkart, 1976). Styles emerge from most flowers prior to anthesis but they are probably not receptive at this stage and the flowers remain in this state for some days. Prosopis species are generally assumed to be self-incompatible (Solbrig and Cantino, 1975). Self-incompatibility has probably been positively selected in desert environments, with obligate outcrossing leading to high variability in the progeny produced, both within and between natural populations. Prosopis species are primarily insect pollinated, and this is facilitated by nutritional rewards. Flowers attract large numbers of potential pollinators with the production of copious amounts of pollen. For extensive lists of the numerous species of insect visitors, pollinators or seed-eating beetles of P. glandulosa, refer to Kingsolver et al. (1977), Ward et al. (1977) and Johnson (1983).

Although very large numbers of flowers are produced, not all are fertile (Solbrig and Cantino, 1975) and high rates of ovary abortion are found. Intra-populational dimorphism in nectar production was observed in P. glandulosa flowers in Mexico (López-Portillo et al., 1993). Very few legumes are produced compared with the large numbers of flowers produced per tree, and an average of 17% of the total seed produced by 15 American species was estimated to be immature (Solbrig and Cantino, 1975). Reports show that when collected and stored, seed of Prosopis species can remain viable for considerable periods of time. Seeds of P. glandulosa have been reported to maintain more than 50% viability over 10-15 years when stored in their pods (Pasiecznik and Felker, 1992). Seedlings are rarely observed under the canopy of a mature tree, possibly because of shading, allelopathic effects, or the presence of seed-eating insects.

Associations

Mares et al. (1977) summarised the ecological associations, describing American Prosopis tree species, in their native range, including P. glandulosa as "representative large desert trees which provide protection from grazing animals, shade, a moist microhabitat, a substrate for climbing or perching, and a reliable supply of nutrients for parasitic and semi-parasitic plants. In providing these habitat components, such desert scrub trees allow an increase in plant density and richness in the community as a whole. These species, which would be rare or absent without the presence of trees and shrubs such as Prosopis, in turn contribute to the support of other trophic levels by providing food sources (leaves, flowers and fruits) for desert scrub animals". Covering such a wide area, P. glandulosa forms part of many ecosystems and subsequently has a large variety of plant associations. In a majority of American arid and semi-arid ecosystems, it is often the dominant or co-dominant species.

Like many legume tree, P. glandulosa has evolved a symbiotic relationship with Rhizobium spp. and other nitrogen-fixing bacteria and also mycorrhizal associations to varying degrees. More Rhizobium strains are noted in native stands in the Americas than in areas where it has been introduced in Africa, possibly due to evolutionary adaptations favouring symbiotic relationships, with 98 strains of both fast-growing and slow-growing Bradyrhizobium types identified in association with P. glandulosa (Jenkins et al., 1987). Mycorrhizal fungi have been isolated from Prosopis roots, and their presence has positive benefits on plant survival and growth.

Mammals, in their native range, use P. glandulosa along with other desert trees for shade, protection and food; for an extensive list, see Mares et al. (1977). Physical protection is offered from the sun by a wide crown, and the thorns and shrubby form can offer excellent protection from large predators. Small mammals feed on the foliage as well as the pods. Birds use the trees as perches and nesting sites, and Prosopis species have been identified as important for native and migratory birds within their native range or where they have expanded their natural range into other habitat types. Some birds and mammals may prefer Prosopis species but most appear not to be restricted to them. The seeds and pods form an important part of the diet of many small desert mammals and, where P. glandulosa plants are common in the Americas, it is thought that their removal would cause a significant decrease in the populations of small wild mammals (Mares et al., 1977). One main association today is with domestic mammals, which have quickly developed a strong relationship with native P. glandulosa. Reptiles and amphibians seek the shade and shelter offered by P. glandulosa trees. Pollinating insects are very important, and other insects feed on the flowers or use them as a mating ground.

Environmental Requirements

P. glandulosa occupies soils overlying a variety of geological formations with no specific affinities, preferring clay soils but tolerates a wide variety of soil types. Soil nutrient status is rarely a limiting factor to distribution. Nitrogen is very rarely limiting, with nitrogen fixation and soil improvement leading to an increase in soil fertility as P. glandulosa trees mature (Geesing et al., 1999). Trees have been noted to fix nitrogen under conditions of high salinity and high water deficits (Felker et al., 1981). P. glandulosa is particularly able to tolerate saline and alkaline soils, but appears not to be well suited to acidic soils, and the possibility that low pH is a limiting factor to the distribution has been suggested (Peacock and McMillan, 1965). P. glandulosa is able to survive in areas with exceptionally low annual rainfall or very lengthy dry periods but only if roots are able to tap ground water or another permanent water source within the first few years, or if sufficient atmospheric moisture is available as in many coastal desert areas with persistent trade winds or seasonal fog. P. glandulosa is known to survive severe frosts to -20°C in the USA (Felker et al. 1982), but can also tolerate some of the highest temperatures in the world, being one of the few tree species growing in Death Valley, California, USA, with temperatures often exceeding 50°C.

Climate

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ClimateStatusDescriptionRemark
As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
B - Dry (arid and semi-arid) Preferred < 860mm precipitation annually
BS - Steppe climate Preferred > 430mm and < 860mm annual precipitation
BW - Desert climate Preferred < 430mm annual precipitation
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
38 20 50 1200

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) -20
Mean annual temperature (ºC) 20 30
Mean maximum temperature of hottest month (ºC) 20 40
Mean minimum temperature of coldest month (ºC) 5 15

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration612number of consecutive months with <40 mm rainfall
Mean annual rainfall501200mm; lower/upper limits

Rainfall Regime

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Summer
Winter

Soil Tolerances

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Soil drainage

  • free
  • impeded

Soil reaction

  • alkaline
  • neutral

Soil texture

  • heavy
  • light
  • medium

Special soil tolerances

  • infertile
  • saline
  • shallow
  • sodic

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Algarobius bottimeri Herbivore Seeds Zimmermann, 1991 South Africa Prosopis
Algarobius prosopis Herbivore Seeds Zimmermann, 1991 South Africa Prosopis
Anthonomus grandis
Evippe Herbivore Leaves Anderson et al., 2006 Australia Prosopis
Leveillula taurica
Mimosetes protractus Herbivore Seeds
Neltumius arizonensis Herbivore Seeds
Oncideres cingulata
Oncideres rhodosticta Herbivore Stems Polk and Ueckert, 1973
Prosopidopsylla flava Herbivore Growing point/Stems Van Klinken et al., 2009
Rotylenchulus reniformis

Notes on Natural Enemies

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Natural enemies of species of Prosopis native to the Americas are well documented with over 945 phytophagous insect species recorded (Van Klinken et al., 2009). A wide range of generalist natural enemies are known to occur on introduced trees. Beetles of the family Bruchidae are responsible for destroying a substantial percentage of seeds produced by P. glandulosa in its native range, and dispersal mechanisms may be an evolutionary response to destruction by such insects. Of the species of beetles found to feed on the pods of native American Prosopis, 93% were obligately restricted to Prosopis, showing a high degree of specialisation. Herbivory is an important factor, with an estimated 40% of immature P. glandulosa leaves removed by insects and 35% of shoots removed by rodents or insects (Nilsen et al., 1987). Defoliating insects vary in their severity of attack but have been found to be a major factor affecting the timing of bud break in North America. Locusts are not uncommon and twig girdlers (species of Oncideres) are damaging in some areas with adult beetles girdling small branches before ovipositing. Wood-boring beetles are also frequently found in the sapwood of fallen or cut trees but their effect on living trees is unknown. Nematodes are known to attack the roots of P. glandulosa (Freckman and Virginia, 1989), while occasionally infesting neighbouring crops.

Means of Movement and Dispersal

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Natural Dispersal

In arid and semi-arid zones it is particularly important that seeds are dispersed to sites with preferable water status. Water is an important dispersal agent in desert ecosystems. Water dispersal ensures widespread dissemination of seed during flooding or other high rainfall events when seedling establishment is favoured. P. glandulosa is often found colonizing ephemeral watercourses and dispersal is aided by water flow in the rainy season, particularly during very wet years (Solbrig and Cantino, 1975). Oceanic dispersal is important when trees are present along the coastal, and for crossing large bodies of water such as in the Caribbean. Pods and endocarps float and are impervious to water infiltration, protecting the seed from the harmful effects of extended periods in sea water. The spread of Prosopis species, where introduced, has followed similar patterns. In Australia and South Africa, the spread of P. glandulosa and related species followed periods of high rainfall (Zimmerman, 1991; Csurhes, 1996), possibly due to improved conditions for germination and establishment or increased water dispersion of the seed. This has led to the establishment of even-aged Prosopis stands. As Prosopis species have been naturalized in most countries for less than 100 years, this limits the possibility of detailed studies on long-term plant succession. However, studies on the ecology of spread in Australia (van Klinken et al., 2005) conclude that stands will continue to increase in area and density, and no self-thinning is expected to occur unlike that shown to happen in native range invasions, on the assumption that conditions are different.

Vector Transmission

Pods have a high sugar content, are low in anti-feedants, and are widely sought after by a variety of animals. Disjunct stands of trees near to old centres of population suggest that man has also been a dispersal agent in historic and prehistoric times (Fredrickson et al., 2006). Birds, bats, reptiles and ants also feed on Prosopis spp. fruits and are potential, if only minor, agents of dispersal. Livestock, except pigs and sheep, are now the primary dispersal agents, although the fruit are also avidly consumed by a wide variety of wild animals which play a major role in seed dispersal. The hard seeds tend to survive passage through the gut of most animals and, although there are conflicting reports on the effects of this passage on seed germination, it is generally accepted that the fruits and seeds are specialised for animal dispersion. Pods are eaten off the tree or off the ground and seeds are deposited in the faeces. However, removal of the endocarp is by no means ensured. Voided seed are given a positive advantage by being placed in faeces, with its improved water-holding capacity and high levels of nutrients. Livestock may tend to spend more time on better pasture or by water sources but voiding of seed in preferential locations is not guaranteed. However, different animals have very different effects on seed survival. Seed with the endocarp shell removed were common in the faeces of goats and cattle but rarely seen in the faeces of horses or foxes (Solbrig and Cantino, 1975). Passage through an animal is also thought to destroy seed-eating insects, but some survival of bruchid beetles has been noted (Kingsolver et al., 1977).

Agricultural Practices

Rapid invasions of Prosopis have accompanied the arrival and expansion of cattle ranching and its associated effects on the environment (Dahl, 1982). These include preventing grassland fires which otherwise killed young seedlings, over-stocking and so reducing herbaceous cover and plant competition which reduces seedling establishment, and increasing cattle numbers which act as effective disseminators of seed via their faeces. The soil erosion and soil degradation that follows over-grazing gives P. glandulosa seedlings a competitive advantage in poorer, nitrogen-deficient soils, and browsing and cutting result in the formation of multi-stemmed shrub forms. There have also been studies in North America showing a gradual northward expansion of P. glandulosa during the Holocene interglacial but at rates much lower than the recent explosion in P. glandulosa invasion following human interventions since 1800.

Intentional Introduction

All introductions of P. glandulosa around the world have been intentional, for their perceived benefits for fuel and fodder production and their tolerance to extreme drought and inhospitable environments.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Animal productionAs a fodder crop Yes Pasiecznik et al., 2001
Digestion and excretionLivestock and wild animals Yes Pasiecznik et al., 2001
DisturbanceAids establishment Yes Pasiecznik et al., 2001
Flooding and other natural disastersSpreading seeds Yes Solbrig and Cantino, 1975
ForestryAs a fuelwood tree Yes Pasiecznik et al., 2001
Landscape improvementAs a urban/street tree in native range Yes Pasiecznik et al., 2001
Ornamental purposesAs a urban/street tree in native range Yes Pasiecznik et al., 2001
ResearchSpecies introduction trials Yes Pasiecznik et al., 2001

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
WaterDuring rains and floods Yes Solbrig and Cantino, 1975

Impact Summary

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CategoryImpact
Animal/plant collections None
Animal/plant products Negative
Biodiversity (generally) Negative
Crop production None
Cultural/amenity Positive and negative
Economic/livelihood Positive and negative
Environment (generally) Positive and negative
Fisheries / aquaculture None
Forestry production None
Human health Negative
Livestock production Negative
Native fauna Positive and negative
Native flora Negative
Rare/protected species None
Tourism None
Trade/international relations None
Transport/travel Negative

Economic Impact

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The losses to the livestock industry solely from P. glandulosa invasion in the southern states of the USA, notably Texas, are counted in many millions of dollars due to the reduction in the availability of forage grasses (Jacoby and Ansley, 1991). Although the financial costs of eradication are also very high, it was seen for many decades to be rewarded by the increase in stocking densities afforded by control. The thorns also cause injury to livestock, and consumptions of the pods, when they may up the bulk of the animal's diet, can lead to ill health and even death. A similar situation to that of the south-western USA exists in Australia and South Africa, where P. glandulosa is called the number one scourge of livestock farmers.

Environmental Impact

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Impact on Habitats

The principal impacts of P. glandulosa invasion are believed to be on water and nutrient availability via direct competition. Where P. glandulosa has been removed in the USA, dry river beds have been observed to flow again, but the exact effects of this species on the level of water tables has yet to be fully elucidated. Dense stands in South Africa are estimated to use the equivalent of 1100 mm of rainfall per year (194 million m³), the equivalent to almost four times the average rainfall in the North Cape Province (Versfeld et al., 1998; Zimmermann et al., 2006). It also forms dense thickets and monocultures and crowds out other plant species. However, increased cover and shelter tends to have positive effects on animals, and a number of wild mammals, birds and even ants have been shown to increase in numbers following invasion by P. glandulosa in its native range.

Impact on Biodiversity

P. glandulosa var. torreyana and P. velutina have invaded large areas of arid southern Africa in the last century, and where are causing dieback and increased mortality of Acacia erioloba, a keystone species in the Kalahari Desert (Schachtschneider and February, 2013). This is attributed to competition for water with Prosopis as stable isotope results show that in riparian zones both A. erioloba and Prosopis are using the same water source, with a 50% increase in canopy dieback of A. erioloba in invaded river plots relative to the cleared river plots. This dieback cannot be related to changes in rainfall and temperature as there were no adverse fluctuations in the 10 years preceding this study. But, as A. erioloba is more water stressed in invaded river plots, this increase in mortality and dieback is due to plant moisture stress related to competition for water with Prosopis and not a clanging climate (Schachtschneider and February, 2013). Interestingly, a study on bush clump succession in South Africa concluded that the overall pattern especially of Acacia karroo invasion (encroachment) shows close parallels with that initiated by invasion of P. glandulosa into grassland in Texas, USA (O'Connor and Chamane, 2012).

A study of the impact of introduced P. glandulosa on an assemblage of dung beetles (Coleoptera: Scarabaeinae) in the northern Cape Province of South Africa showed that species richness and abundance were reduced in dense stands compared with a patch of open savanna (Steenkamp and Chown, 1996). Another study of bird and plant biodiversity in the southern Kalahari, South Africa, also showed reduced biodiversity in patches of introduced P. glandulosa and P. velutina compared with patches of native Acacia karroo (Dean et al., 2002). In Australia it was found that dense stands of hybrid P. glandulosa var. glandulosa almost totally excluded all herbaceaous cover (van Klinken et al., (2006). Whereas it may be expected that the presence of impenetrable stands of P. glandulosa decrease biodiversity, the opposite can occur. P. glandulosa invades mostly degraded grasslands where there is already a much reduced plant biodiversity, and although desirable forage grasses may be reduced, other native herbaceous species may flourish where cattle cannot browse. Also, birds and small mammals may benefit from protection from predators and hunters.

Impacts of increasing P. glandulosa cover from 0 to 100% in North America was assessed by Teague et al. (2014), and found it to be associated with significant declines in herbaceous biomass. Specifically, there were significant declines of C4 tall grasses, C4 mid-grasses, and forbs, small declines of C4 short grasses, and significant increases of C3 annual grasses, and both cool and warm season forb herbaceous functional groups declined with increasing P. glandulosa cover. Results indicate that the greatest benefits in species richness and herbage productivity could be achieved through reducing density of P. glandulosa especially on bottomland sites (Teague et al., 2014).

However, impacts on native grasses is not always clearly defined. For example, Ansley et al. (2013) found that quantifying woody cover/grass production relationships in North America savanna ecosystems where P. glandulosa dominates, is very complex. They found that maintaining productive stands of some grasses could be facilitated by maintaining woody cover below 30% threshold levels, and possibly by limiting grazing during episodic high rainfall events, i.e. that some tree cover is beneficial, and clearance should be accompanied by some form of rotational grazing.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Mentzelia leucophyllaNatureServe NatureServe; USA ESA listing as threatened species USA ESA listing as threatened speciesNevadaCompetition - monopolizing resourcesUS Fish and Wildlife Service, 1990

Social Impact

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In the southern states of the USA, Australia and South Africa, some livestock farmers have been forced to abandon their ranches due to P. glandulosa invasion with resulting social costs. The thorns also cause personal injury. When infestations are great, the dense thickets of P. glandulosa become impenetrable, blocking footpaths and restricting access.

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Highly mobile locally
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Has high genetic variability
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Increases vulnerability to invasions
  • Modification of fire regime
  • Modification of hydrology
  • Modification of nutrient regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts animal health
  • Negatively impacts livelihoods
  • Reduced amenity values
  • Reduced native biodiversity
  • Soil accretion
  • Threat to/ loss of native species
  • Transportation disruption
Impact mechanisms
  • Causes allergic responses
  • Competition - shading
  • Interaction with other invasive species
  • Poisoning
  • Rapid growth
  • Rooting
  • Produces spines, thorns or burrs
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult/costly to control

Uses

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P. glandulosa is a true multi-purpose species, and all parts of the tree have been, or continue to be, used. The timber is very hard with a desirable colour and good finish, comparing favourably with the world's best timbers, being used mainly for flooring and furniture in the USA and Mexico. Poles are used for fence posts and in rustic construction. The firewood is of excellent quality, and imparts a pleasant flavour to food cooked over it, and P. glandulosa charcoal ('mesquite coals') is traded commercially throughout the USA for the barbeque industry. The pods were once a staple food for numerous Native American tribes (e.g. Fredrickson et al., 2006) and are still a valuable fodder for livestock. The flowers are good bee forage giving a high quality honey, and the exudate gum is comparable to commercial gum Arabic from Acacia senegal. The bark is rich in tannins and fibres for leather treatment and rope making, and every part of the tree has been used for folk medicines.

Galactomannan seed gums from P. glandulosa have also been shown to be superior in quality to commonly used guar gum, with much potential for use in the food industry (Martínez-Ávila et al., 2014).

Harvested P. glandulosa has the potential for bioenergy under certain densities (Park et al., 2012), as the high harvesting and transport costs are offset by essentially no production costs. The costs of harvesting, storing and delivering biomass to a bioelectricity plant was assessed, under the assumption that the rights to harvest could be acquired in long-term leases. The advantage is that they grow on land that is not suitable for growing food or fibre and will not impact agricultural food markets as maize or sugarcane ethanol has done. In addition, there are no cultivation costs. Results indicated that tree density and harvesting costs are major factors affecting cost of delivered biomass, but that electricity production is a viable option for managing areas invaded by P. glandulosa (Park et al., 2012). A later study by Park et al. (2014) found that, compared to conventional coal, a lower economic return was found with P. glandulosa biomass used for electricity production, but a possible carbon emission tax would make this competitive with coal.

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed
  • Forage

Environmental

  • Agroforestry
  • Amenity
  • Erosion control or dune stabilization
  • Land reclamation
  • Landscape improvement
  • Revegetation
  • Shade and shelter
  • Soil conservation
  • Soil improvement
  • Wildlife habitat
  • Windbreak

Fuels

  • Biofuels
  • Charcoal
  • Fuelwood
  • Miscellaneous fuels

General

  • Ornamental

Human food and beverage

  • Beverage base
  • Emergency (famine) food
  • Flour/starch
  • Food additive
  • Gum/mucilage
  • Honey/honey flora
  • Seeds

Materials

  • Alcohol
  • Bark products
  • Dye/tanning
  • Wood/timber

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Wood Products

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Charcoal

Furniture

Other cellulose derivatives

Railway sleepers

Roundwood

  • Building poles
  • Posts
  • Roundwood structures
  • Stakes

Sawn or hewn building timbers

  • Carpentry/joinery (exterior/interior)
  • Exterior fittings
  • Fences
  • Flooring
  • For light construction
  • Gates
  • Wall panelling

Wood hydrolysates

Woodware

  • Industrial and domestic woodware
  • Tool handles
  • Toys
  • Turnery
  • Wood carvings

Similarities to Other Species/Conditions

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P. glandulosa has often been confused with other Prosopis species of section Algarobia due to the incorrect binomial being applied at or following introduction. Recent studies on the history of introduction (Pasiecznik et al., 2001) and guides to their identification (Pasiecznik et al., 2004) are resolving these causes of misidentification. The long and widely spaced leaflets of P. glandulosa var. glandulosa from Louisiana to New Mexico and neighbouring Mexican states clearly separate it from other Prosopis species, except some forms of P. chilensis from South America with which it is sometimes confused (where introduced), though the upright form, zig-zag branches and stouter thorns of P. chilensis are generally apparent. The closer-spaced leaflets of P. glandulosa var. torreyana from New Mexico to California, USA, and adjacent areas of Mexico are similar and not always separable from sympatric Prosopis species. Separation of the two varieties where introduced, is however, not always evident. However, P. glandulosa does hybridize with sympatric Prosopis species from section Algarobia, and zones of introgression with P. velutina in California, Arizona and New Mexico, USA, contain trees with intermediate foliar characteristics that are difficult to ascribe to one species or the other. Extensive populations of P. glandulosa x P. velutina hybrids have been identified in Australia and South Africa, where both species have been introduced.

Prevention and Control

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Prevention

Eradication

Major regional eradication and control programmes have been undertaken on Prosopis species in their native ranges, principally in the USA and Argentina, and also where they have been introduced, principally in Australia and South Africa.

The presence of P. glandulosa invading rangeland in southern USA, and threatening the livelihoods of ranchers for over 100 years, led to calls for its eradication. For over 60 years, researchers and ranchers in this region have worked together in an attempt to rid themselves of the menace of P. glandulosa invasion. However, no efficient and cost effective methods have been found. A wealth of literature exists on eradication programmes developed in the USA using mechanical and chemical techniques; these have been tested in the field against mainly P. glandulosa with limited success (see Cadoret et al., 2000). This information is well summarized in Jacoby and Ansley (1991) and forms the basis for evaluating the feasibility of eradication and control. Similar, but less intensive, eradication programmes have been implemented in South Africa and Australia. The term eradication has gradually been replaced with that of control, as it has become clear that total kill and exclusion of Prosopis from a site once invaded, if possible, cannot prevent further encroachment and reinvasion. Research still continues in the USA and Australia using fire, but it appears to have variable results and will not kill large trees anyway, so may be best used as part of an integrated control programme.

Control

Cultural Control

Hand clearance was the first method used to deal with Prosopis as a weed in the Americas. Work teams were sent into invaded pasture to fell all trees and seedlings and uproot stumps. Although very effective and obtaining a harvest of firewood and possibly fence posts and poles, the operation was too labour-intensive and expensive for the land owner to consider carrying it out on anything but a small scale. Hand clearing remains practical only for small land holdings of high value.

Young seedlings are sensitive to fire but older trees become increasingly protected by thick bark as they mature and will resprout rapidly after fire. However, fire can be used successfully as a management tool for preventing the re-establishment of young P. glandulosa seedlings and improving forage production, now seen as the most cost-effective control method for use in Texas, USA (Teague et al., 2001). Fire has been used in conjunction with other methods in the development of integrated eradication programmes in Australia. For example, spraying with herbicides produces dead wood that will ignite and support a sustained fire with more likelihood of killing the remaining trees. Marked differences were noted in the germination of ingested seed following passage through different animals (Mooney et al., 1977); germination was 82% with horses, 69% with cattle, but only 25% with sheep. Replacing free ranging cattle with other livestock, particularly sheep and pigs, possibly in conjunction with other control methods, could drastically reduce the spread of Prosopis species. Ponce-Guevara et al. (2016) also introduce the role of native mammals in controlling Prosopis, and that the reduction in their frequency is one of the reasons for the encroachment of P. glandulosa. Results from research demonstrated that black-tailed prairie dogs (Cynomys ludovicianus) and moderate grazing by cattle can suppress tree growth, and, when their populations are properly managed, they may interact synergistically to significantly limit P. glandulosa encroachment in desert grasslands (Ponce-Guevara et al., 2016).

Control by utilisation is increasingly being promoted as the best means to impact upon Prosopis invasions, and Fredrickson et al. (2006) concluded that one of the reasons for spread of P. glandulosa in North America was that it was no longer widely used as a resource, and concluded that “future control of mesquite may also arise from our appreciation of its attributes and the eventual use of mesquite to meet human needs”.

Mechanical Control

Mechanical site clearance involves tractor operations developed for removing trees, in which the roots are severed below ground level to ensure tree kill, and has been frequently used against P. glandulosa in the USA. These operations include root ploughing and chaining, which are often the most effective mechanical means, using a mouldboard plough pulled behind a Caterpillar tractor, or a heavy chain pulled between two machines.

With root ploughing, large trees must first be felled by hand, but this treatment has been used to remove stumps of up to 50 cm in diameter without difficulty and has a treatment life of 20 years or more (Jacoby and Ansley, 1991). Other advantages are that only a single pass is required, while site cultivation is effected leading to improved soil water conservation, and there is a chance to reseed with improved forage species. However, this method is one of the most expensive control treatments and is recommended only on deep soils that have a high potential for subsequent increased forage production (Jacoby and Ansley, 1991). The soil should be neither too wet nor too dry for effective root ploughing.

Chaining involves pulling a heavy chain between two slow moving Caterpillar tractors, with the effect of pulling over and uprooting larger trees. A second pass in the opposite direction ensures that roots on all sides are severed to ease tree removal (Jacoby and Ansley, 1991). Soil moisture is again important, with soil that is dry on the surface but moist below giving the optimal conditions. If the soil is too dry, the stem breaks leading to coppicing, if too wet, the soil and understorey is damaged (Jacoby and Ansley, 1991). Smaller, unbroken trees have to be removed by other means. Although this is an expensive treatment, it is effective where there are many mature trees. It is most widely used following herbicide application to remove dead standing trees.

Clearance with a biomass harvester produces wood chips that can be sold for energy production offsetting the operational costs (Felker et al., 1999).

Chemical Control

Chemical treatments involve the use of herbicides to kill trees, with the most effective being stem or aerial applications of systemic herbicides. Effectiveness is dependent upon chemical uptake, which in P. glandulosa is limited by the thick bark, woody stems and small leaves with a protective waxy outer layer. The formulation and application of chemicals for trees of mixed ages and sizes within a stand is difficult.

Many herbicides and herbicide mixtures have been tested on P. glandulosa. Until the banning of its use in the 1980s, 2,4,5-T was the herbicide of choice in the USA (Jacoby and Ansley, 1991) and Australia (Csurhes, 1996). Although 2,4-D provided excellent suppression of top growth, few trees were actually killed and such chemical treatments had to be applied periodically to ensure that forage yields were maintained. Infested sites often needed spraying every 5-7 years. The most effective chemical for high tree kill of P. glandulosa in the USA is clopyralid, but dicamba, picloram and triclopyr have also been successfully used, either alone or in combination (Jacoby and Ansley, 1991).

Biological Control

The high cost and poor success of mechanical and chemical eradication techniques have led to the investigation of alternative means of control. Several biological control programmes using species of seed-feeding bruchid beetles have been developed and implemented. In the native range, bruchid beetles can destroy substantial amounts of seed produced, thus severely limiting the potential for invasion. Species of bruchid beetles have been successfully introduced as part of control programmes in South Africa and Australia. The advantage with bruchids is their observed host specificity, with many species found to feed only on Prosopis, and some only on single species (Kingsolver et al., 1977). Other insect species known to have a deleterious effect on native and exotic Prosopis in the Americas, mainly twig girdlers and psyllids, have also been suggested as possible biological control agents. The twig girdler Oncideres rhodosticta is seen as a serious pest of P. glandulosa in the USA (Polk and Ueckert, 1973). Psyllids are known to severely affect the growth of Prosopis (Hodkinson, 1991) and have been suggested for use in controlling invasions.

Most work on biological control of Prosopis to date has been carried out in South Africa, where several programmes are underway. The seed-feeding insects Mimosetes protractus and Neltumius arizonensis were introduced to South Africa in conjunction with the bruchid beetles Algarobius prosopis and A. bottimeri for the control of invasive Prosopis species. N. arizonensis and A. prosopis were successful in establishing themselves in large numbers and having a significant effect on Prosopis, whereas the other species were only found in low numbers (Hoffmann et al., 1993). Maximum damage to seed was found where grazing was controlled, as the multiplication and progress is hampered by livestock devouring the pods before the insects destroy them.

In order to increase seed losses caused by existing agents, surveys were undertaken in Argentina by McKay et al. (2012), and a seed-feeding weevil Coelocephalapion gandolfoi Kissinger (Coleoptera: Brentidae: Apioninae) was identified. Aspects of the biology and the host range of this seed-feeding weevil were studied in Argentina and South Africa to evaluate its potential as a biocontrol agent, and field surveys found that the beetle was responsible for 51% of the seed damage on P. flexuosa with a host range restricted to Prosopis species in section Algarobia. Oviposition and feeding preference for Prosopis species native to Argentina and P. glandulosa from North America was very high, and it was considered a good candidate for biological control of invasive Prosopis species in South Africa.

In Australia, Prosopis infestations are at a relatively early stage and extreme care is being employed in the selection of suitable biological control agents, following the long history of problems caused there by plant and animal introductions. Insect species continue to be tested for their efficacy and host specificity as possible biological control agents of Prosopis species in Australia (van Klinken, 1999; van Klinken et al., 2009). Besides the two Algarobius species, the sap-sucking psyllid Prosopidopsylla flava and the leaf-tying moth, Evippe sp. have both been found to provide some control in Australia (Anderson et al., 2006).

However, in some countries, the current and potential future value of Prosopis as a resource means that biocontrol is not an option, and ‘control by utilization’ is being promoted as an alternative (Pasiecznik, 2006). The planned release of bruchid beetles in Kenya was aborted in 2007, when arguments that this could also destroy a fledging industrial use of pods for animal feed were accepted by the Forestry department (Nick Pasiecznik, personal communication, 2007).

Integrated Control

Mixed mechanical and chemical methods have proved more effective than either alone in several cases. Several integrated programmes that mix mechanical, chemical and fire have had reasonable success but are costly and require a high level of management input. For example, fire has been used in conjunction with other methods in the development of integrated eradication programmes in Australia. Spraying the tree with herbicides produces dead wood that will ignite and support a sustained fire with more likelihood of killing the remaining trees. However, methods of eradication that have been attempted for over half a century in the Americas have proved very expensive and largely unsuccessful in the long term. Total tree kill may be possible with some treatments, but adequate techniques for preventing the re-introduction of seeds and re-establishment of trees have yet to be developed. The potential environmental damage from the widespread use of herbicides must also be taken into consideration. It has been accepted that eradication is not possible using these techniques and, at best, only some form of control is feasible.

Integrated land management options are proposed as a sustainable solution to the problem of weedy invasions. The aim is to treat dense infestations to convert them to economically viable and sustainable agroforestry systems and this has already been evaluated with P. glandulosa in the USA (Felker et al., 1990). Following conversion, such systems involve management of the tree and understorey component to prevent further establishment of Prosopis seedlings. Systems have been devised that produce all of the above outputs on a single unit of land, showing internal rates of return far in excess of any alternative land use possible for such a site (Felker et al., 1990). Converting P. glandulosa stands in the USA from 356 trees/ha to 192 trees/ha produced 32.7 m³/ha of woody biomass, of which an estimated 55% was firewood, leaving 18m³/ha of solid wood for use as a timber, from the thinning operation (Cornejo-Oviedo et al., 1992). Felker et al. (1990) noted that because of small tree sizes, there was no timber produced from stands with densities above 3000 trees/ha, but as stand density decreased to 470 trees/ha, timber volume increased rapidly. Further periodic pruning maintains tree form and the value of saw logs as a system output.

Recruitment of Prosopis seedlings is prevented, or very much reduced, under the crown of mature Prosopis trees (Simpson, 1977). Correct management of understorey vegetation, maintaining ground cover and preventing over-grazing, will also restrict recruitment through competition. Maintenance and improvement of soil fertility is also thought to reduce the competitive advantage that woody legume seedlings have over other species (Geesing et al., 1999). Controlled burning, inter-row cultivation, collection of pods and grazing of livestock such as sheep and pigs, which kill ingested seeds, can be used to prevent further seedling recruitment (Harding, 1991). Livestock production can be significantly improved if conditions allow for understorey management.

Planting of productive forage species has proved to be economic in several countries, and inter-planting with agricultural crops may be possible on better soils in higher rainfall zones. Such an integrated agroforestry system, with a rotation length of 20-50 years, if established on open ground, has been shown to be profitable when using only the saw logs for timber as the system output (Felker et al., 1990). Further work is required on the economics of conversion of dense Prosopis infestations. Changes in land-use systems in the spread of Prosopis (D'Antoni and Solbrig, 1977) and the rapid expansion in areas invaded, are likely to be more important that climatic change. However, few cultural control programmes have been proposed or attempted (Jacoby and Ansley, 1991) even though some change in land-use management appears necessary for the effective control of P. glandulosa.

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05/12/16 Updated by:

Nick Pasiecznik, Consultant, France

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Nick Pasiecznik, Consultant, France

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Nick Pasiecznik, Consultant, France

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