RAPD and microsatellite transferability studies in selected species of <Emphasis Type="Italic">Prosopis</Emphasis> (section Algarobia) with emphasis on <Emphasis Type="Italic">Prosopis juliflora</Emphasis> and <Emphasis Type="Italic">P. pallida</Emphasis>

The genus Prosopis (Leguminosae, Mimosoideae), comprises 44 species widely distributed in arid and semi-arid zones. Prosopis pallida (Humb. & Bonpl. ex Willd.) Kunth and P. juliflora (Sw.) DC. are the two species that are truly tropical apart from P. africana, which is native to tropical Africa (Pasiecznik et al. 2004), and they have been introduced widely beyond their native ranges. However, taxonomic confusion within the genus has hampered exploitation and better management of the species. The present study focusses primarily on evaluating the genetic relationship between Prosopis species from the section Algarobia, containing most species of economic importance, though P. tamarugo from section Strombocarpa is also included for comparison. In total, 12 Prosopis species and a putative P. pallida × P. chilensis hybrid were assessed for their genetic relationships based on RAPD markers and microsatellite transferability. The results show that P. pallida and P. juliflora are not closely related despite some morphological similarity. Evidence also agrees with previous studies which suggest that the grouping of series in section Algarobia is artificial.     

Development and characterization of microsatellite markers for Prosopis chilensis and Prosopis flexuosa and cross‐species amplification

Prosopis chilensis and Prosopis flexuosa (Fabaceae) are closely related hardwood arboreal species that are widely distributed in the arid regions of Argentina. The development of highly polymorphic markers, such as microsatellites, is desirable for genetic studies of these species. Here, we present the development and characterization of six polymorphic microsatellite markers in P. chilensis and P. flexuosa. These markers showed a polymorphism information content between 0.14 and 0.85 and the number of alleles varied from two to 13 considering both species. All markers revealed a broad cross‐species affinity when tested in seven other Prosopis species. All primers amplified in at least five species.     

Community Impacts of Prosopis juliflora Invasion: Biogeographic and Congeneric Comparisons

We coordinated biogeographical comparisons of the impacts of an exotic invasive tree in its native and non-native ranges with a congeneric comparison in the non-native range. Prosopis juliflora is taxonomically complicated and with P. pallida forms the P. juliflora complex. Thus we sampled P. juliflora in its native Venezuela, and also located two field sites in Peru, the native range of Prosopis pallida. Canopies of Prosopis juliflora, a native of the New World but an invader in many other regions, had facilitative effects on the diversity of other species in its native Venezuela, and P. pallida had both negative and positive effects depending on the year, (overall neutral effects) in its native Peru. However, in India and Hawaii, USA, where P. juliflora is an aggressive invader, canopy effects were consistently and strongly negative on species richness. Prosopis cineraria, a native to India, had much weaker effects on species richness in India than P. juliflora. We carried out multiple congeneric comparisons between P. juliflora and P. cineraria, and found that soil from the rhizosphere of P. juliflora had higher extractable phosphorus, soluble salts and total phenolics than P. cineraria rhizosphere soils. Experimentally applied P. juliflora litter caused far greater mortality of native Indian species than litter from P. cineraria. Prosopis juliflora leaf leachate had neutral to negative effects on root growth of three common crop species of north-west India whereas P. cineraria leaf leachate had positive effects. Prosopis juliflora leaf leachate also had higher concentrations of total phenolics and L-tryptophan than P. cineraria, suggesting a potential allelopathic mechanism for the congeneric differences. Our results also suggest the possibility of regional evolutionary trajectories among competitors and that recent mixing of species from different trajectories has the potential to disrupt evolved interactions among native species.     

Climatic signals in growth and its relation to ENSO events of two Prosopis species following a latitudinal gradient in South America

Semiarid environments throughout the world have lost a major part of their woody vegetation and biodiversity due to the effects of wood cutting, cattle grazing and subsistence agriculture. The resulting state is typically used for cattle production, but the productivity of these systems is often very low, and erosion of the unprotected soil is a common problem. Such dry‐land degradation is of great international concern, not only because the resulting state is hardly productive but also because it paves the way to desertification. The natural distribution of the genus Prosopis includes arid and semiarid zones of the Americas, Africa and Asia, but the majority of the Prosopis species are, however, native to the Americas. In order to assess a likely gradient in the response of tree species to precipitation, temperature and their connection to El Niño southern oscillation (ENSO) events, two Prosopis species were chosen along a latitudinal gradient in Latin America, from northern Peru to central Chile: Prosopis pallida from a semi‐arid land in northern and southern Peru and P. chilensis from a semiarid land in central Chile. Growth rings of each species were crossdated at each sampling site using classical dendrochronological techniques. Chronologies were related with instrumental climatic records in each site, as well as with SOI and N34 series. Cross‐correlation, spectral and wavelet analysis techniques were used to assess the relation of growth with precipitation and temperature. Despite the long distance among sites, the two Prosopis species presented similar responses. Thus, the two species' growth is positively correlated to precipitation, while with temperature it is not. In northern Peru, precipitation and growth of P. pallida present a similar cyclic pattern, with a period of around 3 years. On the other hand, P. pallida in southern Peru, and P. chilensis also present this cyclic pattern, but also another one with lower frequency, coinciding with the pattern of precipitation. Both cycles are within the range of the ENSO band.     

Symbiotic properties of sinorhizobia isolated from Acacia and Prosopis nodules in Sudan and Senegal

The host specificity, infection process and effectiveness of nodules produced by several African sinorhizobial strains on different Acacia and Prosopis species (Leguminosae, Mimosoideae) were studied. Sinorhizobium arboris strain HAMBI 1552^T, S. kostiense strains HAMBI 1489^T and HAMBI 1493, S. saheli strain HAMBI 1496 and S. terangae bv. acaciae strain ORS 1058 induced nitrogen fixing nodules on seedlings of the following African or Latin American species (marked with ^*): A. angustissima ^* , A. mellifera, A. nilotica, A. oerfota (synonym A. nubica), A. senegal, A. seyal, A. sieberiana, A. tortilis subsp. raddiana, P. chilensis ^* , P. cineraria, P. juliflora and P. pallida ^* . All strains increased plant yield significantly compared with uninoculated seedlings watered with nitrogen-free medium, but none appeared to be superior. The sinorhizobial strains were unable to effectively nodulate Sesbania rostrata (Papilionoideae).All roots had hairs, but particularly in the case of Acacia spp. they were often sparse. After inoculation root hairs were deformed and, in general, infection in Acacia spp. occurred through short root hairs and in Prosopis spp. through longer ones. After entry, the rhizobia filled infection pockets in the root hair, which later expanded into sac-like structures. When infection threads occurred, they usually started from sac-like structures. Elongation and ramification of the nodules indicated that Acacia spp. and Prosopis spp. have indeterminate nodules. A persistent apical meristem, which is the characteristic feature of the indeterminate nodule type, was much clearer in Prosopis spp. than in Acacia spp. Sinorhizobial strains formed tumour-like structures with undifferentiated cell tissue on the Australian acacia A. holosericea and ineffective, nodule-like structures on the African P. africana.     

Inter- and intra-genetic variation of four wild populations of Prosopis using rapd-pcr fingerprints

The species of Prosopis (Leguminosae) are welladapted to grow in the arid regions of the world. In Mexico, nearly 70%of its territory accounts for arid lands and nine species of this genus arefound in the country which are currently utilized for multiple purposes. Giventhat a sustainable exploitation of this important natural resource is necessary,knowledge of the genetic variability of natural populations is required. Thegenetic variation among and within four wild populations of Prosopisglandulosa, P. juliflora, and two ofP. laevigata, located in the Mexican States of Chihuahua(1), Sinaloa (2), Guanajuato (3) and Hidalgo (4), respectively, was determinedusing random amplified polymorphic DNA (RAPDs). There was a genetic variation of72.48% among these four populations, and within them the estimated geneticvariation was 27.52%. Between the populations of P.laevigata the genetic variation obtained was 7.15%, and within them92.85% (AMOVA, P < 0.001). Based on the assumption ofWright's island model, the genetic variation found among populations correspondsto a gene flow (N_em) of 0.09, while the gene flow detected betweenthe populations of P. laevigata was 3.2. A total of 27 RAPDphenotypes were identified in the four populations; among them, 11 correspondedto P. laevigata. The phylogenetic analyses showed that thethree species are differentiated in three clades and that probably P.juliflora was the origin of the other two species. The protogynoustrait of their flowers and the outcrossing are the main factors that may explainthe genetic variation observed in these populations.     

Bioaccumulation and effect of cadmium in the photosynthetic apparatus of Prosopis juliflora

In the present study Prosopis juliflora plants grown in hydroponics solution were exposed to 50,100 and 1000 μM CdCl₂. The cadmium uptake, transport and toxicity on the photosynthetic activities in the plants were measured at 48 h after starting cadmium treatments. The results showed that the concentration of Cd²⁺ in P. juliflora tended to increase with addition of Cd²⁺ to hydroponics solution. However, the increase of Cd²⁺ in roots and leaves varied largely. In this sense, the accumulation of Cd²⁺ in P. juliflora roots increased significantly in proportion with the addition of this metal. In contrast a relatively low level of Cd²⁺ transportation index, and bioaccumulation factor were found in P. juliflora at 48 h after of treatments. On the other hand the maximum photochemical efficiency of photosystem II (Fv/Fm) and the activity of photosystem II (Fv/Fo) ratios in P. juliflora leaf treated with Cd²⁺ not showed significantly changes during the experiment. These results suggested that the photosynthetic apparatus of P. juliflora was not the primary target of the Cd²⁺ action. Further studies will be focused in understanding the participation of the root system in Prosopis plants with the rhizosphere activation and root adsorption to soil Cd²⁺ under natural conditions.     

Comparative study (AFLP and morphology) of three species of Prosopis of the Section Algarobia: P. juliflora, P. pallida, and P. limensis. Evidence for resolution of the “P. pallida–P. juliflora complex”

The problems of delimitation of species of Prosopis originate from the few morphological discontinuities which exist among some of them; some, however, originated as a result of wide distribution of germplasm without proper knowledge of the species, in particular, much material catalogued as P. juliflora, but being of other species, was distributed for reforestation projects worldwide. This work tests the morphological results obtained for P. pallida and P. limensis of the Peruvian–Ecuadorian coast and for P. juliflora of the Caribbean Basin of Colombia and Venezuela utilizing a study of AFLPs and a study of the morphology of plantlets developed in a conventional garden study. The phenogram obtained for the AFLPs demonstrates each of the three species to be a well differentiated cluster and the molecular variance between them is significantly greater than the variance within each species. Study of the plantlets also indicates statistically significant differences for four morphological characters between P. juliflora and the other two species (P. pallida and P. limensis). These results, in addition to the morphological differentiation evident between adult plants of P. pallida and P. limensis and the clear separation of these two species from P. juliflora, corroborate the genetic identity of the three taxa analyzed.     

Discovery of a life history shift: precocious flowering in an introduced population of Prosopis

The genus Prosopis contains many valuable, long-lived, multi-purpose legume trees, some of which are also invasive species. The time of first flowering is important for increasing production of sweet, protein-rich pods in plantations, especially under short rotations, and affects the rate of spread as a weed. Trees generally begin flowering at 3–5 years of age. However, seedlings from a small seedlot collected from an introduced population in southern Mauritania were observed to begin flowering at 3–4 months after germination. This is unknown in the native range of any Prosopis species, and appears to represent a major evolutionary event triggered after naturalization. This paper reports a detailed investigation with seed collected from Aleg, Brakna region, Mauritania, in 1998 (Prosopis sp. ‘Aleg’). Two experiments were established, in glasshouses at Cirad, Montpellier, France, and Coventry University, UK. Flowering began at the two sites 104 and 169 days after sowing, respectively, and 97.5% of plants had flowered at Coventry after 306 days. Flowers produced abundant pollen with 50–60% viability indicated by FDA staining and 40–50% germination on an artificial medium. No pods were formed. All plants analysed were diploid. Morphologically, these juvenile plants were similar to American species of section Algarobia, and exhibited leaf characters typical of P. pallida. However, due to the large variation in morphology within this section of the genus, further studies are required to confirm the actual species. This is the first record of precocious flowering in Prosopis, and has been found only in a single, introduced population. Implications for genetic improvement of tropical Prosopis, and particularly the spread of these species as invasive weeds, are discussed. Such a life history shift in the increased reproductive ability of a species following introduction, with potentially significant environmental effects, may not be restricted to Prosopis and merits further detailed investigation.     

The host range and biology of the mesquite psyllid Heteropsylla texana

Prosopis spp. areintroduced rangeland weeds in Australia. In asearch for biological control agents, thepsyllid Heteropsylla texana, which causessevere distortion to growing leaf and floralshoots, was imported from Texas, USA forhost range testing in quarantine facilities.No-choice tests were conducted on 60 plantspecies including P. pallida, P. velutinaand P. juliflora. In these trials, adultssurvived on 45 non-target plants but ovipositedonly on the Prosopis spp., Dichrostachys spicata and Acaciabidwillii. Nymphs developed to ovipositingadults on all Prosopis spp. and both D. spicata and A. bidwillii. Developmentof small numbers of adults on D. spicataand A. bidwillii occurred for only onegeneration. Damage was noticed only on Prosopis spp. In multiple-choice trials usingthree plant species, oviposition and subsequentdevelopment of eggs and nymphs to adults tookplace on P. pallida and D. spicatabut not on A. bidwillii. The low numbersdeveloping, the longer development times toadults when reared on D. spicata and A.bidwillii, and the failure of these plantsto sustain populations beyond one generationindicate that these plants are not hosts ofH. texana. While adult survival on manytest plants may imply that adult feedingoccurred, the risk to populations of theseplants in the field is negligible. It wasconcluded that H. texana is specific toProsopis spp. and could be released inAustralia for control of Prosopis spp.Observations of the biology of this speciesmade during the course of rearing andexperimentation indicated that nymphs developedthrough five instars to adults in 7–8 days sothat total development from egg to adult takes13–17 days. Females produce up to 100 eggs.     

Genetic structure of two Prosopis species in Chaco areas: A lack of allelic diversity diagnosis and insights into the allelic conservation of the affected species

The Gran Chaco is the largest continuous region of the South American dry forest, spanning Argentina, Paraguay, Bolivia, and Brazil. Prosopis rubriflora and Prosopis ruscifolia are typical tree species of chaquenian area forests, which have been subjected to continuous fragmentation caused by cattle raising. This study evaluated P. rubriflora and P. ruscifolia in areas with varying levels of disturbance. We investigated the contemporary genetic diversities of both species in areas with distinct anthropogenic disturbances. Even with a lower heterozygote frequency, disturbed areas can provide important storage for alleles, allowing the maintenance of diversity. The genetic diversity of P. rubriflora was surprisingly similar to that of P. ruscifolia (H_e = 0.59 and H_e = 0.60, respectively) even with very different distribution ranges of both species. However, P. ruscifolia exhibited a higher intrapopulation fixation index than P. rubriflora. P. rubriflora showed evidence of bottlenecking in 64% of the sampled areas, while P. ruscifolia showed such evidence in 36% of the sampled areas. Additionally, P. rubriflora had two distinct populations due to its disjunctive geographic distribution, whereas P. ruscifolia had a single population that exhibited few signs of population structure in some areas, possibly due to the main pollinators presenting a short range of dispersion. Our results suggest that 42 Chaco areas should be conserved to retain the minimum of 500 individuals necessary to maintain genetic diversity for 100–1,000 generations. This study improves our understanding of these two Prosopis species and provides information for the conservation of their genetic diversities.     

Biofuels and invasive species from an African perspective – a review

A large number of proposed biofuel crops share the same traits as known invasive plant species, many of which are already present in Africa and include species such as Prosopis glandulosa Torrey (Mimosaceae), P. juliflora (Sw.) DC, Leucaena leucocephala (Lam.) de Wit (Mimosaceae), Azadirachta indica A. Juss. (Meliaceae), and others. In this paper, we mainly assess the impacts of invasive Prosopis species in Africa, particularly in Kenya and South Africa. Introduced Prosopis species have invaded over 4 million hectares in Africa, threatening crop and pasture production, reducing underground water reserves, and displacing native plant and animal species. This has major implications for millions of people who depend on natural resources for their survival. It is therefore suggested that known invasive or potentially invasive plant species not be introduced to countries or regions for biofuel production. If (after a stringent cost–benefit analysis) the introduction of a potentially invasive species is deemed critical for economic development and the benefits clearly outweigh the potential costs, countries should endeavour to abide by the Principles and Criteria for Sustainable Biofuel Production developed by the Roundtable on Sustainable Biofuels.     

The effect of silicon on the growth of Prosopis juliflora growing in saline soil

A decrease in whole plant dry weight was observed when Prosopis juliflora (Swartz) DC. was treated with saline irrigation water for 24 days which was partially alleviated by the addition of 0.47 mM SiO₂ to the irrigation water. The plants treated with high salinity and SiO₂ showed a greater distribution of dry material to the leaves at the expense of the stems and roots compared to control plants. The possible use of SiO₂ to grow plants may be beneficial in areas of high soil salinities.     

Analysis of the genetic similarities among seven species of Prosopis (Leguminosae:Mimosoideae)

Summary Prosopis (mesquite) is a commercially promising plant genus that has received increased attention lately. Allelic frequencies at 25 enzyme loci in seven species from the section Algarobia were transformed in matrices of genetic distance using three methods. The indices give highly correlated results; only in the cluster Prosopis alba -P. hassleri were minor discrepancies evident. The phenetic relationships observed agree with other biochemical evidence (chromatography of phenol compounds, electrophoresis of seed proteins, etc) but not with morphological groupings. The present data support the hypothesis that the species belonging to the section Algarobia would be equivalent to sub- or semispecies; the community of such sympatric subspecies constitutes a syngameon.     

Germination of Prosopis juliflora (Sw) DC seeds after scarification treatments

Invasive plant species are the second most important threat to global biodiversity loss after land‐use change. Invasive species can modify native community composition, deplete species diversity and affect ecosystem processes. The Caatinga is one of the most human‐affected Brazilian ecosystems owing to non‐sustainable use of its natural resources. Prosopis juliflora is an important invasive plant species in the Caatinga ecosystem. Seed germination is a critical stage in plant life cycles and is a major factor in the establishment and success of invasive plant species. Among the factors that affect seed germination and dormancy, coat‐imposed seems to be the most important for P. juliflora. In Prosopis species, the ingestion of fruits by wild and domestic animals may promote and accelerate germination, enhancing the dispersal of seeds and fruits of these species. We investigated the germination capacity of P. juliflora seeds after artificial mechanical and chemical scarification and analyzed the changes in seedling vigor caused by the scarification treatments. Germination rate, germination time (TMG) and germination synchrony (E) differed significantly with the length of the scarification treatments in H₂SO₄ for both seeds with endocarps and seeds without endocarps (non‐endocarp seeds). Sulfuric acid affected plant survival more strongly than germination rate, particularly in non‐endocarp seeds.     

Classification of Prosopis juliflora invasion in the Lake Baringo basin and environmental correlations

The spread of Prosopis juliflora in the Baringo basin, Kenya, has led to severe changes in the ecosystem with negative socio‐economic impacts. The drivers that foster the invasiveness of Prosopis are not fully understood. Thus, a method to quantify the degree of infestation will support the determination of environmental preferences and the risk assessment of future Prosopis invasion. We developed a methodology for characterising and classifying degrees of Prosopis infestation in vegetation stands and propose its application in environmental correlation models. The relative cover was identified as the most suited attribute for assessing and monitoring the invasion of Prosopis. The distance of invaded stands from original plantations and environmental attributes related to water availability (ground water table, rainfall and soil water–holding capacity) have potential to predict potential or future invasion risks.     

Unexpected low genetic differentiation among Allopatric species of section Algarobia of Prosopis (Leguminosae)

The Chaqueña Biogeographic Province, in South America, is the main diversity centre of Prosopis. A group of sympatric species of Section Algarobia in this region constitutes a syngameon, characterised by frequent hybridization and introgression. These processes have been postulated as responsible for the low genetic differentiation observed among species within this group. In this study genetic variability and differentiation among geographically isolated species of the same section was analysed through isoenzyme electrophoresis. Variability parameters and fixation indices were estimated to determine the genetic structure of populations. Two Argentinean, P. ruscifolia (‘vinal’) and P. flexuosa (‘algarrobo amarillo’), and one North American species, P. glandulosa (honey mesquite) were studied. All of them showed, similarly to other species of the same section, high genetic variability and exhibit homozygote excess, probably due to population substructure and low rates of selfing. In contrast to our hypothesis, genetic similarity among species is not related to geographic distance. Genetic distances between P. glandulosa and South American species are similar to those observed among species of this subcontinent. The results obtained suggest that the high genetic similarity among the species of the section Algarobia studied is not due to hybridization.     

Modulation of Rubisco activity in leaves of <Emphasis Type="Italic">Prosopis juliflora</Emphasis> in response to tropical conditions in north India

The success of P. juliflora, an evergreen woody species has been largely attributed to temperature acclimation and stomatal control of photosynthesis under wide range of environmental conditions prevalent in India. We studied the contribution of the enzyme ribulose-1,5 bisphosphate carboxylase/oxygenase (Rubisco) in diurnal and seasonal photosynthesis changes in P. juliflora. The changes observed in photosynthesis under natural conditions could be effected by the growth temperatures, which ranged from 10–30 °C in winter to 30–47 °C in summer. However, the Total Rubisco activity displayed a constant diurnal pattern and showed a maximum at 1200 in all seasons namely spring, summer, monsoon and winter irrespective of the changes in temperature. The Total Rubisco activity from two cohorts of leaves produced in spring and monsoon appeared to be down-regulated differentially at low PPFD during the evening. The in vivo and in vitro measurements of carboxylation efficiency of Rubisco showed wide variation during the day and were correlated with the photosynthesis rate. The light activation of Rubisco showed the acclimation to moderately high temperatures in different seasons except in summer. The exceptionally high temperatures (>45 °C) in summer, though not affecting Total activity, severely inhibited the light activation of Rubisco and also modulated the recovery process for the activation of Rubisco. Our studies suggest that the modulation of Rubisco driven by Rubisco activase and not Rubisco per se was crucial for the diurnal regulation of photosynthesis. NBRI Publication No.: 528     

Ecological implications of seed characteristics of the native Prosopis cineraria and the alien P. juliflora

Seed weight, percentage germination, seedling growth, and nutrient concentrations (Mg, Na, K, Zn, Cu and P) of whole seeds, and of seed coats and embryos separately of two tree species, the native Prosopis cineraria and the invasive alien P. juliflora from semi-arid and arid areas of north and north-west India, were analysed to understand the differences in their ecology. Seeds of P. cineraria were heavier than those of P. juliflora. Percent germination was similar in the two species, but seedling growth was faster in P. juliflora than in P. cineraria. Nutrient concentrations of seeds of the two species were similar (except Cu). Nutrient concentrations in the embryo were higher in P. cineraria, while those in the seed coat were higher in P. juliflora. The relative allocation of nutrients to seed coat was higher in P. juliflora than in P. cineraria. Nutrient-rich embryos and slow growth, along with a staggered seed germination pattern in the native P. cineraria could be linked to delayed establishment as well, in the substratum. Faster growth of the nutrient-poor embryos in P. juliflora along with its simultaneous seed germination pattern, and creation of a favourable microenvironment through leaching of nutrients from a nutrient-rich seed coat can facilitate immediate and successful establishment of this alien species in the invaded habitats.     

Prosopis glandulosa and the nitrogen balance of rangelands: extent and occurrence of nodulation

Summary We report the recovery of root nodules from P. glandulosa var. glandulosa in the eastern portion of its range, where the species reaches its greatest vegetational development. Single cores 4.7 cm in diameter and up to 250 cm deep yielded from 0 to over 250 nodules. Nodules were found at all depths below 10 cm, with the highest concentration often around 100 cm. Detailed studies of three trees revealed relatively small volume densities of about 0.02 nodules cm^–3, high surface area densities of 2–4 nodules cm^–2, and high nodule biomass of 8–23 g m^–2, when compared to cultivated legumes. Nodules are small, weakly attached to roots that are seldom over 0.5 mm in diameter, and not easily observed under field conditions. No nodules were recovered from cores from the more arid western portion of P.glandulosa's range, although seedlings nodulated readily in these soils in the glasshouse as well as in most unamended soils from throughout mesquite's geographical range. Local differences in nodulating potential of soils included a negative association with mesquite canopies and a positive association with depth. These results suggest a significant role for biological fixation in the nitrogen regime and vegetation dynamics of Prosopis-dominated ecosystems.