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Algarroba, Mesquite, Kiawe) (Fabaceae)1 Biology and Impacts of Pacific Island Invasive Species. 6. Prosopis pallida and Prosopis juliflora (Algarroba, Mesquite, Kiawe) (Fabaceae)1 Timothy Gallaher2,3 and Mark Merlin2 Abstract: Prosopis pallida and P. juliflora (commonly referred to as algarroba, mesquite, or kiawe) were introduced from South America to areas in Oceania, Asia, and Africa during the early nineteenth century. In many cases, they natu- ralized and became widespread. In some places, alien Prosopis species are highly valued for the products and services that they can provide such as shade, cattle fodder, wood for fuel and fence posts, and nectar for honey production. In Aus- tralia, four Prosopis species including P. pallida, P. juliflora, P. glandulosa, P. velu- tina, and their hybrids are considered invasive and are subject to control efforts. After its introduction to Hawai‘i in 1828, P. pallida became a dominant tree in arid areas of the main Hawaiian Islands, replacing the native lowland dry forest species that had been decimated by human activity, particularly by the introduc- tions of goats and cattle. Prosopis pallida also has become an important economic species in Hawai‘i. Prosopis juliflora, a more recent introduction to Hawai‘i, is now spreading and is considered to be a noxious weed. Competition between Prosopis and native species as well as negative impacts of Prosopis on soil and local hydrology have been reported; however in some cases Prosopis species are char- acterized as midsuccessional species that rehabilitate degraded soils, eventually facilitating later-successional woodland. This provides a potential opportunity to use these species in reforestation efforts. Management decisions regarding these species should include a consideration of both their positive and negative ecological roles. If control or eradication is desired, a number of methods have been employed with various degrees of success. Prosopis L. is a genus of medium-sized trees 1986, Bessega et al. 2006, Burghardt and and shrubs in the family Fabaceae (Legumi- Espert 2007). nosae), subfamily Mimosoideae, with species Well-known species of Prosopis include occurring naturally in Africa, Asia, and in P. glandulosa Torr. (honey mesquite) and P. North and South America. The most recent velutina Woot. (velvet mesquite), which are authoritative work on the taxonomy of the both native to areas of the southwestern genus was conducted by Burkart (1976), who United States and Mexico, and P. pallida recognized 44 species in five sections. Phy- (Humboldt & Bonpland ex Willd.) Kunth logeny within the genus has been revisited and P. juliflora (Sw.) DC. from western South recently, and changes to subgeneric relation- America. Prosopis pallida and P. juliflora are ships have been proposed (Hunziker et al. among the most notable invasive species in several arid and semiarid areas of the world. Although these two species are morphologi- 1 Manuscript accepted 21 December 2009. cally similar and have in the past been treated 2 Botany Department, University of Hawai‘i at together as a complex, these apparent similar- Ma¯noa, 3190 Maile Way, Room 101, Honolulu, Hawai‘i ities may mask important ecological differ- 96822. 3 Corresponding author ([email protected]). ences (Burkart 1976, Pasiecznik et al. 2001, Burghardt and Espert 2007). Pacific Science (2010), vol. 64, no. 4:489–526 names doi: 10.2984/64.4.489 : 2010 by University of Hawai‘i Press Within their native ranges, numerous indige- All rights reserved nous names are applied to species of Prosopis. 489 490 PACIFIC SCIENCE . October 2010 Mesquite is a common name that originally cific and elsewhere as P. chilensis, P. juliflora, applied to the North American P. glandulosa and P. dulcis. In Hawai‘i, this species was and P. velutina, and the name algarroba is also sometimes referred to as algarrobo or generally used to refer to South American algaroba. species of Prosopis (D’Antoni and Solbrig 1977, Felger 1977). In many places where Prosopis juliflora they have been introduced, both names are commonly applied to all Prosopis species. In Synonyms of P. juliflora include Mimosa juli- Australia, all species of Prosopis are referred flora Swartz, Prosopis pallida forma annularis to as mesquite and also as prickle bushes. In Ferreyra, Prosopis inermis H.B.K., and Prosopis Hawai‘i, most people today know Prosopis horrida Kunth. For a complete list of syn- as kiawe, which according to one definition onyms for this and other Prosopis species see means, ‘‘to stream gracefully, as rain in the Pasiecznik et al. (2001). Prosopis juliflora is wind; to sway, as branches’’ (Pukui and Elbert commonly referred to as long-thorned kiawe 1986:146). The origins and context of the in Hawai‘i. Hawaiian name are not known (Yzendoorn 1911). description and account of variation Prosopis pallida Species Description Synonyms of P. pallida include Acacia pallida Prosopis pallida often grows as a tree with a Humboldt & Bonpland ex Willdenow, Mi- straight trunk with a diameter of up to 1 m mosa pallida (Willdenow) Poiret, and Prosopis and occasionally reaching heights over 20 m limensis Bentham (Pasiecznik et al. 2001). Pro- (Figure 1). Tree form may be erect, flat- sopis pallida has been misidentified in the Pa- topped, or decumbent with branches touch- Figure 1. Growth form of Prosopis pallida. This tree is growing at a park in leeward O‘ahu, Hawai‘i. Prosopis pallida and P. juliflora, Pacific Island Invasive Species . Gallaher and Merlin 491 Figure 2. Growth form of Prosopis juliflora. This shrub is growing at a park in leeward O‘ahu, Hawai‘i. A meter stick is present in the image for scale. ing the ground. Prosopis pallida may also grow species, although typically less than 40 mm as a shrub with numerous branches forking in P. pallida. In both species, thorns are oc- just above ground level; in such cases, other- casionally small or absent, particularly on wise tall or erect individuals may assume a upper-canopy branches of mature individuals. shrubby form due to moisture deficit, wind, Thorns on older branches are incorporated salt spray, frost damage, browsing, insect into the secondary growth. infestations, or cutting (Pasiecznik et al. Plants in the genus Prosopis are phreato- 2001). Prosopis juliflora tends to grow as a phytes (literally, groundwater-loving plants), shrub or small tree rarely reaching heights referring to their root system, which is capa- over 10 m (Figure 2). Both species reach ble of accessing relatively deep groundwater maximum height and productivity in riparian sources. Prosopis is described as having two zones with access to shallow groundwater complementary root systems: a taproot sys- (Schade et al. 2003), and both will also pro- tem and a lateral root system. The taproot duce extensive coppice growth when cut system is made up of one to three thickened above ground level. The outer bark of both taproots that grow quickly, rarely branch un- species is rough, fissured, ropelike, and light til a water table is reached, and often reach a gray to dark brown, and the heartwood is depth of 20–25 m (Havard 1884, Heit- dark brown to red and very hard due to the schmidt et al. 1988, Pasiecznik et al. 2001). abundance of resin. On older stems, short In Arizona, roots believed to be from a spe- shoots called brachyblasts develop at the cies of Prosopis were found at a depth of 175 nodes of both species, producing a cluster feet (53 m), among the deepest roots ever ob- of 1–10 leaves, inflorescences, and thorns. served (Phillips 1963, Solbrig and Cantino Thorns may be over 60 mm long on either 1975). The second root system comprises a 492 PACIFIC SCIENCE . October 2010 series of lateral roots that form a dense mat with P. juliflora, the only known tetraploid near the soil surface to take advantage of (2n ¼ 56) in the genus, is known to occur available near-surface water during infre- and appears to result in individuals that are quent rainfall events (Pasiecznik et al. 2001). intermediate in form between the parent Lateral roots of Prosopis can spread 18 m or species (Harris et al. 2003, Trenchard et al. more from the base of the tree (Mooney et 2008; T.G., pers. obs.). Hybridization is also al. 1977). frequent between other species of Prosopis, Leaves of Prosopis are bipinnately com- particularly among species of sect. Algarobia. pound and have an alternate arrangement In some cases, hybrids may be more suitably on the stem. The length of the compound adapted to particular environments than leaf of P. pallida and P. juliflora is between 5 parental types due to higher intraindividual and 20 cm. Each leaf is made up of one to variability (Vega and Hernandez 2005). five pairs of opposite pinnae, and each pinna has between 6 and 29 pairs of oblong, oppo- Distinguishing Features site leaflets 2.5–23 mm long and 1.5–5.5 mm wide (Pasiecznik et al. 2004). Prosopis may be distinguished from other Inflorescences are axillary, spikelike, cy- mimosoid legumes such as Acacia species by lindrical racemes up to 15 cm long. Each is the combination of cylindrical inflorescences, made up of several hundred yellow flowers indehiscent pods, and the absence of phyl- 0.5 cm long. Namba (1956) found 250–525 lodes (Figure 3). Within the genus Prosopis, flowers per raceme and approximately 100 considerable overlap in morphological char- florets per inch (2.54 cm) on P. pallida. acteristics between P. pallida and P. juliflora The flower is connected to the rachis by a has resulted in frequent misidentification. A short pedicel. The calyx and corolla both guide to help distinguish between these spe- have five lobes, and sepals tend to be fused cies is available (see Pasiecznik et al. 2004). and petals free; each flower has 10 stamens In that analysis, the number of interfoliar and a single carpel.
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