Phylogenetic position and revised classification of Acacia s.l. (Fabaceae: Mimosoideae) in Africa,including new combinations in Vachellia and Senegalia

Previous phylogenetic studies have indicated that Acacia Miller s.l. is polyphyletic and in need of reclassification. A proposal to conserve the name Acacia for the larger Australian contingent of the genus (formerly subgenus Phyllodineae) resulted in the retypification of the genus with the Australian A. penninervis. However, Acacia s.l. comprises at least four additional distinct clades or genera, some still requiring formal taxonomic transfer of species. These include Vachellia (formerly subgenus Acacia), Senegalia (formerly subgenus Aculeiferum), Acaciella (formerly subgenus Aculeiferum section Filicinae) and Mariosousa (formerly the A. coulteri group). In light of this fragmentation of Acacia s.l., there is a need to assess relationships of the non‐Australian taxa. A molecular phylogenetic study of Acacia s.l and close relatives occurring in Africa was conducted using sequence data from matK/trnK, trnL‐trnF and psbA‐trnH with the aim of determining the placement of the African species in the new generic system. The results reinforce the inevitability of recognizing segregate genera for Acacia s.l. and new combinations for the African species in Senegalia and Vachellia are formalized. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 500–523.     

Implications of the changing phylogenetic relationships of Acacia s.l. on the biological control of Vachellia nilotica ssp. indica in Australia

Plant relationships have implications for many fields including weed biological control. The use of DNA sequencing and new tree building algorithms since the late 1980s and early 1990s have revolutionised plant classification and has resulted in many changes to previously accepted taxonomic relationships. It is critical that biological control researchers stay abreast of changes to plant phylogenies. One of the largest plant genera, Acacia, has undergone great change over the past 20 years and these changes have ramifications for weed biological control projects in a number of countries. Vachellia nilotica (prickly acacia) is a major weed in Australia, originating from the Indian subcontinent and Asia, and it has been a target for biological control since 1980. Once a member of Acacia, a large (>1,000 spp.) and iconic group in Australia, prickly acacia is now part of the genus Vachellia. Current knowledge suggests that Vachellia is more closely related to mimosoid genera than it is to Acacia s.s. There has also been a recent reclassification of legume subfamilies with subfamily Mimosoideae now part of subfamily Caesalpinioideae, and four new subfamilies. In this paper we review the changes that have occurred to this group since the prickly acacia biological control project began and discuss the implications for the project. A new host test list for quarantine testing is proposed. Developed following the modernisation of the centrifugal‐phylogenetic method, it is shorter than past lists, containing 46 species, although still lengthy because of the expectations of regulatory bodies, which are slower to accept advances in scientific knowledge. The list includes five Vachellia species, six “Mimoseae” species and 26 Acacia species. The number species from legume subfamilies other than the new Caesalpinioideae is greatly reduced.     

Advances in clarifying the phylogenetic relationships of acacias: Relevance for biological control

Biological control of invasive Australian acacias will benefit from recent advances in resolving the phylogenetic relationships of Acacia s.l. and Acacia s.s. (“Australian acacias”) within the subfamily Mimosoideae. Some of the phytophage taxa associated with Acacia s.s. display fidelity to a derived clade within the genus. This derived clade contains most of the Acacia s.s. species that have become problematic around the world. Phytophages that are demonstrably restricted to species within the derived clade pose essentially no risk to species outside Acacia s.s.. In contrast, prospective agents able to develop on species in the basal lineages of Acacia s.s. would require more-expansive testing because Acacia s.s. is closely related to the Ingeae, and then sequentially to the genera Acaciella, Mariosousa and Senegalia. Importantly, Vachellia is distantly related to Acacia s.s., being nested in basal Mimoseae lineages, and is thus less likely to be at risk than previously envisaged. Elucidation of these trends shows the benefits of having a comprehensive knowledge of the phylogeny of plants and phytophages under consideration for biological control.     

Natural rhizobial populations and nodulation status of woody legumes growing in diverse Kenyan conditions

The rhizobial populations and nodulation status of both indigenous (mainly Acacia species) and some introduced woody legume species were assessed under glasshouse conditions in soils collected from 12 sites located in different ecological zones of Kenya. The populations among the sites, as estimated by the MPN technique, varied from <3.6 to>2.3×10⁵ cells g^-1 of soil. There were some intrasite variations in population estimates depending on the trap host species, date of soil collection and the method used in sampling the soils. Nodulation in whole soil also varied across the sites with test species frequently showing higher nodulation ability in native soils. Sesbania sesban (L.) Merr. was the most prolific nodulating species while Acacia tortilis (Forsskal) Hayne was very erratic in nodulation. Nodulation of most species showed interplant and intraspecific variability within a single soil source.     

Cyanogenesis in Acacia pachyphloia

The cyanogenic glycoside, proacacipetalin, is reported from Acacia pachyphloia (Acacia subgenus Acacia). This represents the first record of a glycoside with an aliphatic aglycone from a species of Acacia indigenous to Australia. This finding reinforces the taxonomic distinctions between subgenus Acacia and subgenus Phyllodineae.     

Interspecific variation in the resprouting responses of Acacia species following simulated herbivory in a semi‐arid southern African savannah

Plants have evolved a diverse suite of tolerance traits against herbivory, including compensatory growth, increased photosynthesis and activation of dormant meristems. We studied the responses of five Acacia species to simulated herbivory in a semi‐arid southern African savannah. We clipped terminal shoots of five juvenile Acacia species (Acacia rehmanniana, A. nilotica, A. karroo, A. arenaria and A. gerarrdii) to simulate herbivory. We then determined biomass change after 5 months and also counted the number of resprouts and measured their length and diameter. All clipped shoots produced resprouts, with all the Acacia species compensating for the lost biomass. We found considerable interspecific variation in the compensation for biomass lost to herbivory in the five Acacia species. Resprouts biomass ranged from two times in A. arenaria to four times that removed in A. karroo. Acacia karroo produced many resprouts, while A. arenaria produced very few resprouts (4 vs 15 resprouts). The relationship between the number of resprouts and their growth also varied among the different Acacia species. We conclude that the response of Acacias to herbivory ranges from prolific resprouters (such as A. karroo) to poor resprouters (e.g. A. arenaria).     

A new species of Parapanteles Ashmead, 1900 (Hymenoptera: Braconidae: Microgastrinae) parasitic on Charaxes athamas (Drury) (Lepidoptera: Nymphalidae) in India

A new species of gregarious endoparasitoid, Parapanteles athamasae n. sp. (Hymenoptera: Braconidae), parasitising caterpillars of Charaxes athamas (Drury) (Lepidoptera: Nymphalidae) on the host plant Senegalia catechu (=Acacia catechu) (L.f.) Hurter & Mabb., is described from Maharashtra, India. Diagnostic characters of the new species include: propodeum with areola 0.93× longer than wide, legs yellow, hind tibia 4.30× as long as ovipositor, ovipositor sheaths exerted, first metasomal tergal plate 1.24× longer than wide, with coarse sculpture merging with longitudinal striations at 3/4 of the apical region. This is the first time a species of the family Nymphalidae Rafinesque is recorded in association with Parapanteles Ashmead, 1900. A key to the Indian species of Parapanteles based on females is also provided.     

The population of Acacia tree species producing gum arabic in the Karamoja region, Uganda

The study aimed at determining the population status of the different Acacia tree species producing gum arabic in the undisturbed, grazed and cultivated habitats in the Karamoja region, Uganda. A total of 135 sample plots each measuring 20 × 20 m² (0.04 ha) with each habitat having 45 plots were selected and established in the seven counties using a simple random sampling technique. The tree species present, their abundances and sizes were recorded. Twelve Acacia species were identified and a total of 5535 recorded in the sampled area. Out of these, five were gum‐producing acacias. Acacia senegal dominated the acacias in all the seven counties and in all habitats of Karamoja with Acacia nilotica (72.3%), Acacia seyal (13.4%), Acacia sieberiana (4%) and Acacia gerrardii (2.6%). Non‐gum‐producing acacias constituted 7.19% of the total abundance. The tree densities increased with increase in tree size in the undisturbed and grazed habitats but decreased in the cultivated habitat. Most Acacia trees were of large size, an indication of old age and poor regeneration that could affect their future population status. It is recommended that further investigations be carried out into the causes of poor regeneration of Acacia species.     

Phylogenetic analysis of Acacia (Mimosaceae) as revealed from chloroplast RFLP data

 Chloroplast DNA of 22 species of Acacia (Tourn.) Miller was digested with ten restriction endonucleases, Southern-blotted and probed with cloned fragments covering the chloroplast genome of tobacco (Nicotiana tabacum L.). Phyletic and phenetic analyses of the resulting 176 polymorphic bands recorded among the 22 species were performed. The phylogram was reconstructed using heuristic search and Wagner parsimony. The resulting most parsimonious consensus phylogram displayed three major phyletic lineages, consistent with the previously established three subgenera of Acacia. The 10 species of subgenus Acacia and the 6 species of subgenus Heterophyllum formed two monophyletic sister clades. The 5 species of subgenus Aculeiferum studied and Acacia albida (Syn. Faidherbia albida) grouped together and were basal to the clades of subgenera Acacia and Heterophyllum. The phylogram indicated that subgenus Heterophyllum diverged earlier from subgenus Aculeiferum than did subgenus Acacia; however, the phenogram indicated the reverse. The study indicated that A. nilotica and A. farnesiana are sister species, though A. nilotica is Afro-Asiatic and A. farnesiana is American. The phenogram separated the three subgenera in agreement with the phylogram, but the two dendrograms differed regarding the topologies of the species and the distance of evolution between subgenera Acacia and Heterophyllum. Received: 8 July 1998 / Accepted: 24 July 1998     

Assessment of woody species encroachment in the grasslands of Nechisar National Park,Ethiopia

Ecological survey was executed to assess woody species encroachment into the grassland plain of Nechisar National Park (NNP). Forty‐one woody species were recorded. Dichrostachys cinerea Wight & Arn., Acacia mellifera (Vahl) Benth., Acacia nilotica (L) Willd., Acacia senegal (L.) Willd., Acacia seyal Del. and Acacia tortilis (Forssk.) Hayne were among the major encroaching woody species. The majority of the woody species were found to be highly aggregated in their pattern of distribution, while only few species showed some degree of randomness. The mean woody species density was ca. 1995 woody plants ha^?1. Mean cover of woody, grass, unpalatable forbs and total herbaceous species were 31%, 58%, 68% and 121%, respectively. The woody species density and cover, unpalatable forbs and bare land cover were significantly higher in the highly grazed and fire‐suppressed part of the grassland plain. Pearson correlation coefficient matrix indicated that woody species cover and density were negatively correlated with total herbaceous and grass cover. The high woody, unpalatable forbs and bare land cover indicated the progressively increasing perennial grass species diversity deterioration in the grass plain of the Park. Decline in the grassland condition, unless reversed, will jeopardize the biological diversity as well as the aesthetic value of the NNP.     

THE PHOTOPERIODIC RESPONSE OF AMERICAN PROSOPIS AND ACACIA FROM A BROAD LATITUDINAL DISTRIBUTION

Prosopis and Acacia from North and South America were studied in the laboratory under constant and increasing night length under two temperature regimes. The Prosopis juliflora (Sw.) DC. complex showed strong populational differentiation; Acacia farnesiana (L.) Willd. showed minimal populational differentiation. In Prosopis a population from Oklahoma (36° N) showed the greatest inhibition of stem height under increasing night length. A population from Peru (6° S) showed least control of stem height by night length. In Acacia the response of the latitudinal series was relatively uniform within each of the 6 sets of light period-temperature conditions. In general the height increase for Acacia was two to three times greater under the warmer regime than under the cooler regime at a given night length. Since Acacia farnesiana occurs sympatrically with the Prosopis juliflora complex over a broad range of latitudes, it is of interest to find that the distributional mechanisms of these two leguminous trees are basically different.     

Effects of ectomycorrhizal and vesicular-arbuscular mycorrhizal fungi on drought tolerance of four leguminous woody seedlings

Seedlings ofAcacia auriculiformis A. Cunn. ex. Benth.,Albizia lebbeck (L.) Benth.,Gliricidia sepium (Jac.) Walp andLeucaena leucocephala (Lam.) de Wit. were inoculated with an ectomycorrhizal (Boletus suillus (l. ex. Fr.) or indigenous vesicular-arbuscular mycorrhizal (VAM) fungi in a low P soil. The plants were subjected to unstressed (well-watered) and drought-stressed (water-stressed) conditions. InGliricidia andLeucaena, both mycorrhizal inoculations stimulated greater plant growth, P and N uptake compared to their non-mycorrhizal (NM) plants under both watering regimes. However, inAcacia andAlbizia, these parameters were only stimulated by either ectomycorrhiza (Acacia) or VA mycorrhiza (Albizia). Growth reduction occurred as a result of inoculation with the other type of mycorrhiza. This was attributed to competition for carbon betweenAcacia and VA mycorrhizas and parasitic association betweenAlbizia and ectomycorrhiza. Drought-stressed mycorrhizal and NMLeucaena, and drought-stressed mycorrhizalAcacia tolerated lower xylem pressure potentials and larger water losses than the drought-stressed mycorrhizal and NMAlbizia andGliricidia. These latter plants avoided drought by maintaining higher xylem pressure potentials and leaf relative water content (RWC). All the four leguminous plants were mycorrhizal dependent. The higher the mycorrhizal dependency (MD), the lower the drought tolerance expressed in terms of drought response index (DRI). The DRI may be a useful determinant of MD, as they are inversely related.     

Convergence Between Dung Beetle Assemblages of a Post-Mining Vegetational Chronosequence and Unmined Dune Forest

Abstract In Maputaland, South Africa vegetative and microclimatic changes on mined dunes drive the composition of the dung beetle fauna toward convergence with that in natural dune forest on unmined dunes. We assessed the pattern of these changes using a 23‐year vegetational chronosequence on mined dunes, which passes from grassland (approximately 1 year) to open Acacia shrubland thicket to Acacia karroo‐dominated woodland (approximately 9 years). Across this sequence, which represents successional stages in the restoration of dune forest, there was a sequential trend toward convergence in dung beetle species composition in both the entire species complement and, particularly, in shade specialist species. However, species abundance patterns showed a trend toward convergence only in early chronosequence Acacia woodland, followed by a decline in similarity between dung beetle assemblages of older Acacia woodland and unmined natural forest. This trend toward divergence was common both to the entire species complement, which includes widespread taxa, and to species endemic to Maputaland or the east coast. These trends in similarity and dissimilarity between dung beetle assemblages closely parallel the greater physiognomic and microclimatic similarity between early Acacia woodland and natural forest and the relative dissimilarity of older Acacia woodland. In conclusion, although percentage similarities between dung beetle assemblages of approximately 12‐year woodland and natural forests were comparable with those between each natural forest stand, decline in similarity in older woodland stands suggests that lasting convergence in dung beetle species abundance will only be attained once the Acacia woodland is replaced by secondary natural forest.     

The sugar and nitrogen content of the gums of Acacia species in the Mountain Ash and Alpine Ash forests of central Victoria and its potential implications for exudivorous arboreal marsupials

Abstract Chemical analysis of the total sugar and total nitrogen content of Acacia dealbata, Acacia obliquinervia and Acacia frigescens gum exudate was completed. These trees were located within stands of 53 year old Mountain Ash, Eucalyptus regnans and Alpine Ash, Eucalyptus delegatensis forest in the Central Highlands of Victoria, southeastern Australia. Values for sugar content ranged from 24 to 68% per sample. Gum samples that were collected in E. regnans forests had a significantly lower (P<0.05) sugar content than those from stands dominated by E. delegatensis. Statistical analyses using Scheffe's S-test indicated that there was a significant difference in the sugar content of gums between A. dealbata and A. frigescens but not between A. dealbata and A. obliquinervia or A. obliquinervia and A. frigescens. Values for the nitrogen content of Acacia gum varied from 0.2 to 0.7% per sample. Statistical analyses revealed that nitrogen content was significantly influenced (P<0.05) by a combination of three interacting factors: (i) Acacia species; (ii) tree diameter; and (iii) forest type. Therefore, the findings of this study indicate that the sugar and nitrogen content of Acacia gum may vary between forest types and tree species. The gum of Acacia species is an important source of food for several species of arboreal marsupials, and differences in sugar and nitrogen content could be a factor potentially influencing the distribution and abundance of these animals.     

Cross-amplification of nonnative Acacia species in the Hawaiian Islands using microsatellite markers from Acacia koa

Thirty-one microsatellite markers, developed from Acacia koa, were evaluated for cross-amplification of seven nonnative species of Acacia in the Hawaiian Islands. The number of alleles per locus ranged from one to seven, with 16 of the 31 markers showing polymorphism in at least two of the nonnative Acacia species. Of the Acacia species examined, A. melanoxylon showed the highest percentage of amplified loci.     

The 5S DNA units ofAcacia species (Mimosaceae)

The DNA sequence structure of 5S DNA units inAcacia species, including representatives from the three subgenera ofAcacia, have been determined. The data was interpreted to suggest that at least three lineages of 5S DNA sequences exist inAcacia and the proposal was made that the lineages be named5S Dna-1, 5S Dna-2, and5S Dna-3. The5S Dna-1 lineage was represented by units fromA. boliviana andA. bidwilli, the5S Dna-2 lineage by units fromA. melanoxylon, A. pycnantha, A. ulicifolia, A. boliviana, A. bidwillii, andA. albida, and the5S Dna-3 lineage by units fromA. bidwillii, A. boliviana, andA. senegal. Based on this interpretation of the sequence data, the Australian species of subg.Phyllodineae grouped together as a cluster, quite separate from the subgeneraAculeiferum andAcacia. As expected from the analyses of morphological characters, the 5S DNA units fromAcacia albida (syn.Faidherbia albida) were quite separate from the otherAcacia spp.     

The importance of species identity in the biocontrol process: identifying the subspecies of Acacia nilotica (Leguminosae: Mimosoideae) by genetic distance and the implications for biological control

Aims A molecular genetic distance study has been used in an initial survey to identify subspecies and genotypes of the weed Acacia nilotica in Australia, information needed to find suitable biocontrol agents. We use patterns of DNA sequence variation (in two DNA fragments) from each of the nine described subspecies of Acacia nilotica (L.) Delile (Leguminosae: Mimosoideae) that is to determine their genetic similarity, to verify if the Australian populations are A. nilotica ssp. indica (Benth.) Brenan, and to establish if any other subspecies are present in Australia. Location Australia and southern Africa through the Arabian peninsular to the Indo‐Pakistan subcontinent. Methods Representative specimens from the global distribution of the nine A. nilotica subspecies were sourced primarily from herbaria sheet specimens where available, and secondarily from field collections. These specimens together with related outgroups from Mimosoideae were genetically analysed using the DNA fragments trnL and internal transcribed spacer one (ITS1). We calculated a similarity index as set out in paup * using upgma (Unweighted Pair‐Group Method Arithmetic average) methods to cluster taxa to produce a genetic distance phenogram. Results Sequence results from ITS1 and trnL DNA fragments identified seven of the described subspecies of A. nilotica. Acacia nilotica ssp. cupressiformis (J. Stewart) Ali & Faruqi and A. nilotica ssp. adstringens (Schumach. & Thonn.) Roberty were not found to be genotypically distinct from A. nilotica ssp. indica and A. nilotica ssp. nilotica, respectively, based on the two DNA fragments. Subspecific ITS1 genotypes were geographically distributed similarly to previous reports that were based on morphology, with the exception that the hemispherica ITS1 genotype also occurred in Somalia. We confirmed that the Australian A. nilotica populations are mostly comprised of subspecies indica, but in addition, some individuals were found to be genetically identical to an unidentified Pakistan genotype not previously reported as occurring in Australia. Main conclusions Australian A. nilotica populations originated from India and Pakistan and we recommend further analysis to determine the complete genetic diversity profile and origins of the Australian populations. We highlight the importance of determining any hybridization between Australian populations of A. nilotica and native subgenus Acacia species. This study demonstrates the importance of genotyping weed species targeted for biocontrol and/or listed host specificity test species that may be easily misidentified. Biocontrol practitioners can justify genetic studies by considering the costs should a project fail through misidentification.     

Contrasting Cloud Forest Restoration Potential Between Plantations of Different Exotic Tree Species

The role of exotic tree plantations for biodiversity conservation is contested. Such plantations nevertheless offer various ecosystem service benefits, which include carbon storage and facilitation of indigenous tree species regeneration. To assess forest restoration potential in tropical exotic tree plantations, we assessed native cloud forest tree regeneration in 166 plots in ca. 50‐year‐old plantations of five timber species that are widely used in tropical plantations (Pinus patula, Eucalyptus saligna, Cupressus lusitanica, Grevillea robusta and Acacia mearnsii). Differences in species abundance, diversity and composition were compared among plantations, and between plantations and disturbed and undisturbed indigenous Afromontane cloud forest (southeast Kenya) relicts after controlling for environmental variation between plots (i.e. altitude, distance to indigenous forest, soil depth, slope, aspect) and for environmental and stand structural variation (i.e. dominant tree height and basal area). Regenerating trees were mostly early‐successional species. Indigenous tree species regeneration was significantly higher in Grevillea plantations, where the seedling community also included late‐successional tree species. Regeneration under Eucalyptus was particularly poor. Acacia had a strong invasive nature, reducing its potential role and usefulness in indigenous forest restoration. Our study underlined that exotic tree plantations have differential effects on native tree species regeneration, with high potential for Grevillea plantations and low potential for invasive exotic species.     

The response of fast- and slow-growing Acacia species to elevated atmospheric CO2: an analysis of the underlying components of relative growth rate

In this study we assessed the impact of elevated CO₂ with unlimited water and complete nutrient on the growth and nitrogen economy of ten woody Acacia species that differ in relative growth rate (RGR). Specifically, we asked whether fast- and slow-growing species systematically differ in their response to elevated CO₂. Four slow-growing species from semi-arid environments (Acacia aneura, A. colei, A. coriacea and A. tetragonophylla) and six fast-growing species from mesic environments (Acacia dealbata, A. implexa, A. mearnsii, A. melanoxylon, A. irrorata and A. saligna) were grown in glasshouses with either ambient (˜350 ppm) or elevated (˜700 ppm) atmospheric CO₂. All species reached greater final plant mass with the exception of A. aneura, and RGR, averaged across all species, increased by 10% over a 12-week period when plants were exposed to elevated CO₂. The stimulation of RGR was evident throughout the 12-week growth period. Elevated CO₂ resulted in less foliage area per unit foliage dry mass, which was mainly the result of an increase in foliage thickness with a smaller contribution from greater dry matter content per unit fresh mass. The net assimilation rate (NAR, increase in plant mass per unit foliage area and time) of the plants grown at elevated CO₂ was higher in all species (on average 30% higher than plants in ambient CO₂) and was responsible for the increase in RGR. The higher NAR was associated with a substantial increase in foliar nitrogen productivity in all ten Acacia species. Plant nitrogen concentration was unaltered by growth at elevated CO₂ for the slow-growing Acacia species, but declined by 10% for faster-growing species. The rate of nitrogen uptake per unit root mass was higher in seven of the species when grown under elevated CO₂, and leaf area per unit root mass was reduced by elevated CO₂ in seven of the species. The absolute increase in RGR due to growth under elevated CO₂ was greater for fast- than for slow-growing Acacia species. Received: 21 December 1998 / Accepted: 31 May 1999