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Botanical Journal of the Linnean Society, 2013, 172, 500–523. With 1 figure
Phylogenetic position and revised classification of
Acacia s.l. (Fabaceae: Mimosoideae) in Africa, including
new combinations in Vachellia and Senegalia
BRUCE KYALANGALILWA1, JAMES S. BOATWRIGHT2, BARNABAS H. DARU1,
OLIVIER MAURIN1 and MICHELLE VAN DER BANK1,*
1
African Centre for DNA Barcoding, University of Johannesburg, APK Campus, PO Box 524,
Auckland Park 2006, Johannesburg, South Africa
2
Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag
x17, Bellville, 7535, Cape Town, South Africa
Received 9 July 2012; revised 5 October 2012; accepted for publication 27 February 2013
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.
ADDITIONAL KEYWORDS: Acaciella – Australia – Mariosousa – molecular phylogeny – taxonomy.
INTRODUCTION
Numerous phylogenetic studies of Acacia Mill. s.l.
over the last 10 years have shown that Acacia is not
monophyletic and it is now widely agreed that Acacia
s.l. needs to be divided into at least five genera
corresponding to the former Acacia subgenus Phyllodineae (DC.) Ser., Acacia subgenus Acacia, Acacia
subgenus Aculeiferum Vassal section Aculeiferum
Vassal, Acacia subgenus Aculeiferum section Filicinae
(Benth.) Taub. and a group of species from North and
Central America related to A. coulteri Benth. (Luckow
et al., 2003; Miller et al., 2003; Maslin, Miller &
Seigler, 2003a; Maslin, Orchard & West, 2003b;
*Corresponding author. E-mail: mvdbank@uj.ac.za
500
Seigler & Ebinger, 2005; Bouchenak-Khelladi et al.,
2010; Miller & Seigler, 2012).
However, despite the clear resolution of five independent lineages in Acacia s.l., some taxa still remain
unplaced in the molecular phylogenetic analyses. A
good example is Senegalia visco (Lorenz ex Griseb.)
Seigler & Ebinger which, although having been transferred to Senegalia Raf., does not show a close relationship to other members of the former Acacia
subgenus Aculeiferum (Seigler, Ebinger & Miller,
2006). Inclusion of such species could still result in
the recognition of additional genera in the future.
Although the above-mentioned groups correspond
to previously established infrageneric groups in
Acacia, with generic names available for all, the Australian Acacia subgenus Phyllodineae (Acacia s.s.)
comprises the largest number of species (c. 1021
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
species; Thiele et al., 2011). In order to preserve
nomenclatural stability according to the International
Code of Botanical Nomenclature, Orchard & Maslin
(2003, 2005) proposed retypification of the genus from
A. nilotica (L.) Willd. ex Del. ( = Acacia scorpioides
(L.) W.Wight), a species widely distributed from Africa
to India, to A. penninervis Sieb. ex DC., an Australian
species. This proposal was adopted at the 2005 International Botanical Congress (IBC) in Vienna (McNeill
et al., 2006). However, the original proposal and its
subsequent adoption sparked controversy, calling into
question the basis of the proposal and the legitimacy
of the procedures followed (because it was prior to the
Committee’s consideration) and the validity of the
decision (Moore et al., 2010). The 2011 IBC meeting in
Melbourne finally ratified the previous decision,
despite the long-standing controversy, paving the way
for name changes as proposed by Orchard & Maslin
(2005), who recommended recognition of the following
genera: Acacia s.s. (formerly Acacia subgenus Phyllodineae), Vachellia Wight & Arn. (formerly Acacia
subgenus Acacia), Senegalia (formerly Acacia subgenus Aculeiferum section Aculeiferum), Acaciella (formerly Acacia Aculeiferum section Filicinae) and
Mariosousa Seigler & Ebinger (species belonging to
the Acacia coulteri group; Murphy et al., 2010; Thiele
et al., 2011).
Of the five newly recognized genera, Senegalia and
Vachellia are found in Africa (including Madagascar)
and are represented by c. 69 and 73 species, respectively (Lewis, 2005). These are trees and shrubs,
many with wide geographical ranges from South
Africa north to the Mediterranean mainly restricted
to dry savannas and semi-desert scrub habitats (Ross,
1979; Dharani, 2006).
Macromorphological characters important in
assigning African taxa of Senegalia and Vachellia to
the appropriate genus include the presence and type
of prickles and/or stipular spines (Miller & Bayer,
2003; Seigler et al., 2006). Senegalia spp. can be
unarmed or armed with prickles, but always lack
stipular spines. When armed, the prickles are mainly
scattered, sometimes grouped in twos or threes near
the nodes (Ross, 1979). Senegalia spp. possess bipinnate leaves with sessile or stipitate glands in variable
positions on the petioles and leaf rachis. Furthermore, they have porate pollen without columellae,
whereas Vachellia spp. have colporate pollen with
columellae (Miller & Bayer, 2003; Maslin et al.,
2003a; Thiele et al., 2011). Unlike Vachellia, members
of Senegalia do not have a true inflorescence involucre
(Seigler et al., 2006). Inflorescences in Senegalia are
either capitate or spicate. Bouchenak-Khelladi et al.
(2010) reported that the most recent common ancestor of Senegalia probably originated in the late
Oligocene-early Miocene about 27.9-16.8 Mya, most
501
Table 1. Morphological differences between Vachellia and
Senegalia (Maslin et al., 2003b)
Character
Vachellia
Senegalia
Prickles
Stipular spines
Pollen aperture type
Pollen collumelae
Involucre on peduncle
Absent
Present
Colporate
Present
Present
Present
Absent
Porate
Absent
Absent
likely in American forests. They hypothesized that
two separate dispersal events from the Americas to
Africa occurred in the late Miocene, coupled with a
shift from closed to open habitats, probably in Africa.
Vachellia spp. are armed with paired stipular
spines at the nodes that can either be straight,
deflexed or weakly falcate. In some species they are
enlarged, forming ant domatia [e.g. Vachellia luederitzii Engl., V. collinsii (Saff.) Seigler & Ebinger, V. cochliacantha Humb. & Bonpl. and V. drepanolobium
Sjöstedt; Ross, 1979, Maslin et al., 2003a]. The presence of spinescent stipules is the key diagnostic character distinguishing Vachellia from Senegalia. Nearly
all of the species with spinescent stipules have capitate inflorescences (Ross, 1979), with the majority
having pale yellowish–white flowers. However, some
have bright golden or orange–yellow or, rarely,
pinkish or purple flowers (Ross, 1979).
Morphological characters considered important in
defining major groups of Acacia s.l. were reviewed by
Maslin et al. (2003a) and Seigler et al. (2006). Despite
the major macro- and micromorphological differences
between Senegalia and Vachellia (see Table 1), it
remains difficult to demarcate monophyletic lineages
within these genera based only on morphological
traits (Ross, 1979; Miller & Bayer, 2003).
Following the results of the above-mentioned
studies demonstrating the disintegration of Acacia s.l.
into five segregate genera and the recent decision
from the 2011 IBC meeting in Melbourne, there is a
clear need to determine the generic placements of all
the African species and to formalize new name combinations for these species. We expect that a wider
sampling covering the African species would provide a
plausible picture of the true relationships within the
genera. This will facilitate the implementation of the
new classification in Africa, particularly in herbaria,
by providing names for the African taxa of Senegalia
and Vachellia. In this study, we determined the position of the African Acacia spp. in the new generic
classification using a comprehensive sampling covering the African species based on molecular data from
three plastid regions, matK/trnK, trnL-trnF and
psbA-trnH.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
502
B. KYALANGALILWA ET AL.
DNA
EXTRACTION, AMPLIFICATION AND SEQUENCING
MATERIAL AND METHODS
Sequences of the following plastid loci were generated
in this study: trnK/matK (42 taxa); trnL-trnF (57
taxa); and psbA-trnH (44 taxa). These were combined
with existing sequence data from GenBank to represent all major lineages in Mimosoideae, resulting in
36 African Vachellia spp. and 32 African Senegalia
spp. Voucher information and GenBank numbers for
sequences for all taxa used in this study are listed in
the Appendix.
Laboratory protocols for DNA extraction, PCR
amplification and sequencing of the three DNA
regions in this study followed Bouchenak-Khelladi
et al. (2010).
SEQUENCE
EDITING, ALIGNMENT AND
PHYLOGENETIC ANALYSES
Complementary strands of the sequenced genes were
assembled and edited using Sequencher v.4.8 (Gene
Codes Corp., Ann Arbor, MI, USA), aligned using
Multiple Sequence Comparison by Log-Expectation
(MUSCLE v.3.8.31; Edgar, 2004) and the alignment
adjusted manually in PAUP* (v.4.0b.10; Swofford,
2002). We coded gaps (insertions/deletions) as missing
characters, and we excluded sections of ambiguous
alignment from the analysis (265 characters from the
trnL-trnF, 107 from psbA-trnH and 167 from trnK/
matK).
Cladistic analyses for the individual (trnL-trnF,
trnK/matK, psbA-trnH) and combined matrices were
performed using maximum parsimony (MP) in PAUP*
v.4.0b10 (Swofford, 2002). Heuristic tree searches
employed 1000 random sequence additions, keeping
ten trees per replicate using tree bisection–
reconnection (TBR) branch swapping with MulTrees
in effect, with all character transformations treated
as equally likely. Trees generated in the initial 1000
replicates were then used as starting trees for a
second search with no tree limit to ascertain whether
the shortest trees were obtained in the initial search.
Delayed transformation (DELTRAN) character optimization was used to calculate branch lengths instead
of accelerated transformation (ACCTRAN) because of
its reported errors in PAUP v.4.0b10 (http://
paup.csit.fsu.edu/problems.html).
As all three plastid regions could not be amplified
for all taxa included in the study, the effects of the
missing data on patterns of relationships and support
in the combined matrix were investigated by performing two different combined analyses: (1) a subset of
the taxa for which all three loci were sequenced; and
(2) all taxa including those for which some loci were
missing sequences. Thus, the combined analyses were
conducted with all taxa for which any sequence was
available and included.
Internal support was estimated using bootstrap
analyses (Felsenstein, 1985) implemented in PAUP*
v.4.0b10 (Swofford, 2002) based on 1000 bootstrap
replicates performed with equal weights using TBR
branch swapping with ten trees held at each step and
simple taxon addition. The following scale for bootstrap support percentages (BP) was used: 50-74%,
low; 75-84%, moderate; 85-100%, strong.
The individual gene trees were assessed for congruence by visual inspection of the individual bootstrap
consensus trees to look for any areas of strongly
supported incongruence (Seelanan, Schnabel &
Wendel, 1997).
A hierarchical likelihood ratio test implemented in
MODELTEST v.3.06 (Posada & Crandall, 1998) was
used to determine the appropriate substitution model
for each of the three plastid gene sequences based on
the Akaike information criterion (AIC; Sugiura,
1978). The optimal models identified were
GTR + I + G for trnK/matK, TVM + I + G for trnLtrnF and TVM + G for psbA-trnH (Yang, 1994) with
number of rate parameters = 6, rate = gamma, base
frequency = empirical, clock = unconstrained and
number of generations = 3 000 000. The combined
matrix was analysed using Bayesian inference (BI) by
partitioning the sequences according to DNA region to
allow independent estimation of parameters for each
partition. Site-specific rates of substitution were
allowed to vary across partitions as implemented in
MRBAYES v.3.1.2 (Huelsenbeck & Ronquist, 2001).
Two parallel Markov chain Monte Carlo (MCMC)
runs were made for 3 000 000 generations, with trees
sampled every 1000 generations, resulting in 3000
trees. The first 1500 trees were discarded as ‘burn-in’.
The following scale was used to evaluate the posterior
probability values (PP): below 0.95, weakly supported;
0.95-1.0 strongly supported.
RESULTS
The individual plastid gene trees (not shown) were
largely congruent (negligible to zero incongruence) and
were concatenated for a combined analysis of all three
loci. Of the 4718 included characters, 2982 were constant, 1736 (36.8%) were variable and 897 (19%) were
potentially parsimony informative. The combined MP
analysis resulted in 142 equally parsimonious trees
(tree length: 3552 steps; CI = 0.61; RI = 0.83, see
Table 2). The combined MP trees are largely congruent
with the BI trees, and the BI majority rule consensus
tree is presented (Fig. 1) and used as the basis for
interpreting and discussing the results. The tree was
generally well resolved and major clades received
moderate to strong support (Fig. 1).
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
503
Table 2. Results of parameters estimated from maximum parsimony (MP) analyses of individual and combined data sets
Number of included positions in matrix
Number of variable sites
Number of potentially parsimony-informative sites
Number of trees
Number of steps (tree length)
Consistency index (CI)
Retention index (RI)
Average number of changes per variable site
THE SENEGALIA
trnL-trnF
psbA-trnH
trnK/matK
Combined
1293
440 (34.0%)
235 (18.2%)
19
831
0.63
0.86
1.8
751
266 (35.4%)
137 (18.2%)
172
514
0.66
0.86
1.9
2674
1590 (59.5%)
550 (20.6%)
256
2126
0.64
0.84
1.3
4718
1736 (36.8%)
897 (19.0%)
142
3552
0.61
0.83
2.04
CLADE
With the exception of three species (Senegalia visco,
S. muricata Britton & Rose and S. angustifolia
Britton & Rose), Senegalia, although not supported in
the MP analysis (0.85 PP, Fig. 1A), is monophyletic
and sister to a large clade comprising Mariosousa +
Acaciella + Ingeae + Acacia s.s. in Senegalia two welldefined groups are apparent, that are together sister
to S. vogeliana (A and B, Fig. 1A). Several African
Acacia s.l. taxa lacking name combinations in Senegalia are placed in this clade.
THE MARIOSOUSA + ACACIELLA + INGEAE +
AUSTRALIAN ACACIA S.S. CLADE
This clade includes the genera Mariosousa (63 BP/
1.0 PP), Acaciella (75 BP/0.95 PP), and Australian
Acacia s.s. (70 BP/1.0 PP, Fig. 1A), the last of which is
nested in tribe Ingeae. Faidherbia albida (Delile)
A.Chev. forms a clade with Zapoteca tetragona
(Willd.) H.M.Hern. (56 BP/0.63 PP), which is sister to
the Ingeae + Acacia s.s. clade. Albizia Durazz., as
currently
circumscribed,
is
potentially
nonmonophyletic, but the relationships among the three
distinct lineages and other genera in tribe Ingeae are
too poorly resolved and weakly supported to draw any
firm conclusions. Mariosousa and Acaciella are both
robustly supported as monophyletic, as found in previous studies (Clarke, Downie & Seigler, 2000; Miller
et al., 2003; Bouchenak-Khelladi et al., 2010; Miller &
Seigler, 2012) and form successive sister groups to the
Ingeae + Acacia s.s. clade.
THE VACHELLIA
CLADE
The monophyly of Vachellia receives high support
(86 BP/1.0 PP; Fig. 1B) and it is embedded in a larger
grade of former tribe Mimoseae lineages. Vachellia
natalitia (E.Mey.) Kyal. & Boatwr. and V. montana
(P.P.Swartz) Kyal. & Boatwr. form a clade (79 BP/
1.0 PP) that is sister to the rest of Vachellia. Lack of
robustly supported resolution within Vachellia pre-
cludes detailed discussion of relationships and formal
recognition of subclades in the genus. Many of the
African Acacia s.l. taxa currently lacking combinations in Vachellia are placed in this clade. The American taxa (C, Fig. 1B) form a weakly supported
monophyletic group nested in the African taxa.
DISCUSSION
In the interest of clarity, all species discussed here are
referred to by their Vachellia and Senegalia names,
even although in some cases the combinations are
only effected in this paper. In general, the results are
in line with previous studies that demonstrated the
non-monophyly of all five traditionally recognized
tribes (Clarke et al., 2000; Luckow, White & Bruneau,
2000; Miller et al., 2003; Lavin, Herendeen &
Wojciechowski,
2005;
Catalano
et al.,
2008;
Bouchenak-Khelladi et al., 2010; Miller & Seigler,
2012).
THE SENEGALIA
CLADE
Senegalia is represented in Africa, Central and South
America and Asia and is sister to a large clade comprising Senegalia muricata and S. angustifolia +
Senegalia visco + Mariosousa + Acaciella + Ingeae +
Australian Acacia s.s. Our results suggest that the
phylogeny of Senegalia is strongly structured geographically with African taxa found in the two subclades containing Acacia nigrescens Oliv. and
A. brevispica Harms, whereas taxa from America (e.g.
Senegalia wrightii Britton & Rose and S. berlandieri
Britton & Rose) and Asia (S. modesta (Wall.) P.J.H.Hurter and S. catechu (L.f.) P.J.H.Hurter & Mabb.)
are found in separate subclades. However, relationships between the Asian species are still unclear.
Lack of resolution and support for many subclades in
Senegalia and sparse taxon sampling, especially of
Asian and American species, at this point preclude
definitive biogeographical conclusions.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
B. KYALANGALILWA ET AL.
0.58
0.85
70
1.00
100
51
1.00
0.88
92
74 0.63
1.00
1.00
54
0.95
0.69
0.52
0.67
0.72
98
1.00
95
1.00
0.50
92
0.92
1.00
0.99
96
0.99
1.00
0.60
100
91
1.00
1.00
100
0.73
1.00
55
67
0.97
1.00
0.94
0.95
56
0.63
0.99
0.88
0.53
0.99
75 0.64
0.95
96 0.76
63 1.00 100
1.00
1.00
66
0.92
0.93
98
1.00
0.61
0.83
66
1.00
0.98
95
65
1.00
0.98
69
67
0.72
0.92
59
0.82
0.63
1.00
0.89
A
65
62
0.88
0.99
74
1.00
73
1.00
77
1.00
0.52
0.95
68
0.99
100
1.00
0.92
0.55
0.85
0.58
0.85
95
1.00
B 76
1.00
0.63 0.96
0.54
99
1.00
Australia
0.59
67
1.00
Acacia vetista
Acacia pataczekii
Acacia mearnsii
Acacia elata
Acacia boormanii
Acacia platycarpa
Acacia melanoxylon
Acacia colei
Acacia auriculiformis
Acacia translucens
Acacia craspedocarpa
Acacia lycopodiifolia
Acacia adoxa
Acacia spinescens
Acacia pulchella
Acacia drummondii
Acacia ampliceps
Pararchidendron pruinosum
Archidendron hirsutum
Paraserianthes lophantha
Pseudosamanea guachapele
Chloroleucon mangense
Albizia tanganyicensis
Enterolobium cyclocarpum
Enterolobium contortisiliqua
Albizia zygia
Albizia petersiana
Samanea saman
Albizia versicolor
Albizia zimmermannii
Albizia glaberrima
Albizia forbesii
Albizia amara
Albizia sinaloensis
Albizia kalkora
Cathormion umbellatum
Havardia albicans
Havardia pallens
Ebenopsis ebano
Lysiloma tergeminum
Lysiloma acapulcense
Albizia harveyi
Albizia brevifolia
Albizia athlemintica
Zygia lathetica
Inga edulis
Zapoteca tetragona
Faidherbia albida
Acaciella tequilana
Acaciella chamelensis
Acaciella rosei
Acaciella boliviana
Acaciella glauca
Acaciella angustissima
Senegalia visco
Mariosousa sericea
Mariosousa coulteri
Mariosousa acatlensis
Mariosousa usumacintensis
Mariosousa dolichostachya
Mariosousa willardiana
Mariosousa salazari
Senegalia muricata
Acacia scleroxyla
Senegalia nigrescens
Senegalia welwitschii subsp. delagoensis
Senegalia montis-usti
Senegalia galpinii
Senegalia burkeii
Senegalia goetzei subsp. microphylla
Senegalia goetzei subsp. goetzei
Senegalia robysiana
Senegalia cinerea
Senegalia polyacantha
Senegalia senegal var. rostrata
Senegalia senegal subsp. leiorhachis
Senegalia dudgeonii
Senegalia zizyphispina
Senegalia condyloclada
Senegalia modesta
Senegalia somalensis
Senegalia ogadensis
Senegalia mellifera subsp. mellifera
Senegalia senegalensis
Senegalia erubescens
Senegalia caraniana
Senegalia hereroensis
Senegalia chariessa
Senegalia caffra
Senegalia macrostachya
Senegalia eriocarpa
Senegalia ataxacantha
Senegalia catechu
Senegalia wrightii
Senegalia occidentalis
Senegalia roemeriana
Senegalia soraria
Senegalia anisophylla
Senegalia picachensis
Senegalia gaumeri
Senegalia berlandieri
Senegalia schweinfurthii
Senegalia brevispica
Senegalia adenocalyx
Senegalia polyphylla
Senegalia polyphylla
Senegalia kraussiana
Senegalia tenuifolia
Senegalia gilliesii
Senegalia furcatispina
Senegalia vogeliana
America
94
70 1.00
1.00
Africa and Asia
59
0.90
A
Acacia s. s.
Ingeae
Acaciella
unplaced
Mariosousa
unplaced
Senegalia
Africa and America
504
Continued in Fig. 1B
Figure 1. Topologies of the 50% majority-rule consensus tree obtained from Bayesian inference (BI) analyses based on
trnL-trnF, trnK/matK and trnH-psbA plastid DNA sequences generated for Acacia s.l. A few missing taxa were
augmented with sequences from GenBank. Numbers above branches are maximum parsimony (MP) bootstrap support
and those below are BI posterior probabilities.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
505
Continued from Fig. 1A
99
1.00
0.79
87
1.00
1.00
100
1.00
0.97
0.67
0.77 0.55
0.99
77
1.00
80
0.61
68
0.97
0.95
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1.00
58
55
0.99
0.88
54
1.00
0.87
71
93 0.93
62 1.00
1.00
80
0.99
100
0.63
1.00
69
0.99
1.00
1.00
0.93
C 86
73
66 0.97
86
98 0.79
0.93
0.85
76
0.88
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95
1.00
1.00
97
1.00
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88
74
77 0.98 1.00
1.00
0.54
77
1.00
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86
1.00
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1.00
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1.00
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1.00
67
0.98
1.00
1.00
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1.00
55
0.81
99
1.00
0.92
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1.00
0.99
100
57
1.00
0.85
0.72
100
87
1.00
71 1.00
99
1.00
1.00
Figure 1. Continued
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
Africa and Asia
0.80
61
1.00
Piptadenia peruviana
Piptadenia minutiflora
Piptadenia stipulacea
Mimosa tenuiflora
Parapiptadenia rigida
Parapiptadenia pterosperma
Microlobius foetidus
Stryphnodendron rotundifolium
Stryphnodendron porcatum
Pseudopiptadenia suaveolens
Pseudopiptadenia contorta
Vachellia karroo
Vachellia swazica
Vachellia sekhukhuniensis
Vachellia robbertsei
Vachellia ormocarpoides
Vachellia kosiensis
Vachellia dyeri
Vachellia borlea
Vachellia permixta
Vachellia nebrownii
Vachellia exuvialis
Vachellia torrei
Vachellia xanthophloea
Vachellia kirkii
Vachellia hockii
Vachellia flava
Vachellia tortilis subsp. raddiana
Vachellia tortilis subsp. spirocarpa
Vachellia tortilis var. tortilis
Vachellia rehmaniana
Vachellia gerrardii
Vachellia robusta subsp. clavigera
Vachellia grandicornuta
Vachellia luederitzii var. retinens
Vachellia sieberiana var. woodii
Vachellia sieberiana subsp. sieberiana
Vachellia reficiens
Vachellia hebeclada subsp. hebeclada
Vachellia hebeclada subsp. tristis
Vachellia hebeclada subsp. chobiensis
Vachellia stuhlmannii
Vachellia nilotica
Vachellia nilotica
Vachellia nilotica
Vachellia nilotica
Vachellia nilotica subsp. indica
Vachellia nilotica subsp. tomentosa
Vachellia gummifera
Vachellia davyi
Vachellia arenaria
Vachellia amythethophylla
Vachellia cochliacantha
Vachellia campechiana
Vachellia pennatula
Vachellia macracantha
Vachellia cornigera
Vachellia collinsii
Vachellia farnesiana
Vachellia caven
Vachellia oveidensis
Vachellia anegadensis
Vachellia choriophylla
Vachellia vernicosa
Vachellia schottii
Vachellia constricta
Vachellia erioloba
Vachellia erioloba
Vachellia haematoxylon
Vachellia montana
Vachellia natalitia
Parkia speciosa
Parkia biglandulosa
Parkia timoriana
Anadenanthera colubrina
Desmanthus cooleyi
Desmanthus bicornutus
Kanaloa kahoolawensis
Schleinitzia insularum
Prosopidastrum mexicanum
Leucaena retusa
Leucaena leucocephala
Neptunia oleracea
Dichrostachys cinerea subsp. africana
Dichrostachys cinerea
Gagnebina bakoliae
Calliandropsis nervosus
Alantsilodendron pilosum
Neptunia monosperma
Neptunia gracilis
Piptadeniopsis lomentifera
Mimozyganthus carinatus
Prosopis pallida
Prosopis glandulosa var. torreyana
Xerocladia viridiramis
Cylicodiscus gabunensis
Newtonia hildebrandtii
Newtonia buchananii
Fillaeopsis discophora
Piptadeniastrum africanum
Elephantorrhiza elephantina
Pentaclethra macrophylla
Pentaclethra eetveldeana
Adenanthera pavonina
Erythrophleum suaveolens
Pachyelasma tessmannii
Dimorphandra conjugata
Dinizia excelsa
Lemuropisum edule
Delonix elata
Parkinsonia florida
Peltophorum dubium
Bussea perrieri
Jacqueshuberia brevipes
Arapatiella psilophylla
Mimoseae I
Vachellia
America
61
0.78
1.00
0.86
Africa
B
Mimoseae II
Caesalpinieae
Mimoseae III
Caesalpinieae
506
B. KYALANGALILWA ET AL.
Although the majority of Senegalia spp. form a
single clade with moderate support, the placement of
two American species, S. muricata and S. angustifolia
outside the main Senegalia clade represents a novel
finding. These two species lack prickles, which provide
an important diagnostic character for the rest of
Senegalia and are in line with their placement outside
Senegalia. This novel clade is strongly supported as
part of the Mariosousa + Acaciella + Ingeae + Acacia
s.s. clade. These findings, although not affecting the
African species of Acacia s.l., suggest placement of
these two species in a genus separate from Senegalia,
but increased taxon sampling is required to determine
the extent of this new clade; for the time being, this
clade remains unplaced in the generic system of
Mimosoideae.
One other species currently placed in Senegalia,
S. visco, is also placed outside the main Senegalia
clade (Fig. 1B), in line with previous findings (Seigler
et al., 2006), further confirming the non-monophyly of
Senegalia as currently circumscribed. In this study,
S. visco is robustly supported as sister to the
Acaciella + Ingeae + Acacia s.s. clade, rather than the
Mariosousa + Acaciella + Ingeae + Acacia s.s. clade as
suggested by Seigler et al. (2006).
THE
MARIOSOUSA + ACACIELLA + INGEAE + AUSTRALIAN
ACACIA S.S. CLADE
Mariosousa comprises 13 species of unarmed erect
shrubs and small trees (never lianas) with herbaceous
stipules restricted to seasonally dry tropical and subtropical regions of Central America and Mexico.
Flowers are arranged in cylindrical spikes. In Acaciella, most species are shrubs or small trees, except
two taxa (Acaciella tequilana (S.Wats.) Britton &
Rose var. tequilana and A. hartwegii (Benth.) Britton
& Rose), which are perennial herbs. They are
unarmed, have no nectaries on their leaves, and the
inflorescence is typically capitulum-like, which often
elongates into a short raceme (Rico & Bachman,
2006).
THE VACHELLIA
CLADE
Vachellia is monophyletic and sister to a larger clade
comprising Mimoseae I, the rest of Acacia s.l and the
Ingeae, in line with previous studies (Clarke et al.,
2000; Miller et al., 2003; Bouchenak-Khelladi et al.,
2010; Miller & Seigler, 2012). Relationships in Vachellia show high geographical structure, with a robustly
supported New World clade nested in a wider old
World clade, even although it is not yet clear how
many subclades should be formally recognized in the
genus. Increased taxon sampling, especially of New
World species, will be crucial to assess this.
Finally, Ingeae and Australian Acacia s.s. form a
monophyletic group, although only supported in BI.
Acacia s.s. is monophyletic and Ingeae remain
polyphyletic in accordance with previous studies
(Miller et al., 2003; Seigler et al., 2006; BouchenakKhelladi et al., 2010; Miller & Seigler, 2012). This clade
is known to be problematic, but there is as yet not
enough resolution or a large enough sampling to
remedy this. There are certainly several nomenclatural complications looming given that Acacia s.s. (the
type of Acacieae) is embedded in Ingeae with Acacieae
being the older of the two tribal names. The segregate
genera Acaciella, Mariosousa, Senegalia and Vachellia
are essentially left tribally unplaced. The boundaries
of Ingeae should also be investigated to ascertain
which genera should be included therein. The relationships of the different genera in this clade are also not
well resolved (Fig. 1A, B). For example, in Ingeae,
Albizia is not monophyletic as shown previously by
Luckow et al. (2003). In their study, they included six
Albizia spp. (Albizia adinocephala (Donn.Sm.) Record,
A. harveyi Fourn., A. kalkora Prain, A. sinaloensis
Britton & Rose, A. tomentosa Standl. and A. versicolor
Welw. ex Oliv.). Of these, only three species (Albizia
sinaloensis, A. adinocephala, and A. tomentosa) form a
monophyletic lineage. In the present study, although
priority was given to typically African species, the
number of included species was almost doubled (to 13),
with only two species included from previous studies
by Luckow et al. (2003) and Bouchenak-Khelladi et al.
(2010). Bouchenak-Khelladi et al. (2010) suggested
that this polyphyletic state could be interpreted as a
rapid radiation of lineages of Ingeae. It can also, in
part, be attributable to inadequate sampling and specimen misidentification. However, it is important to
increase the sampling size to confirm the status of
Albizia spp.
NOMENCLATURAL IMPLICATIONS:
TAXONOMIC CHANGES
Results presented in this study and previously published studies confirm the recognition of at least five
genera in Acacia s.l., and the placement of the African
taxa in the two reinstated genera Senegalia and
Vachellia. Although some other African botanists
(Smith & Figueiredo, 2011) have suggested the continued recognition of Acacia in Africa, pending wider
sampling and further analyses, there is now overwhelming evidence to support these two clades, and
provision of new nomenclatural combinations in Senegalia and Vachellia in Africa, alongside those for the
New World (Seigler et al., 2006), is much needed by
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
the scientific, conservation, forestry and other applied
research communities. Typification of names did not
fall within the scope of this study and mainly follows
Ross (1979) and those cited in the protologues of taxa
published subsequent to this publication. The names
in the list below have all been reconciled with those in
Ross (1979), Lock (1989) and Roskov et al. (2005) and
all accepted names appearing in these works have
been accounted for. Synonymy (except for basionyms)
is not given in this paper, but the information can be
found in the above-mentioned references. The list also
includes names of accepted taxa that have been published subsequent to those in Ross (1979), Lock (1989)
and Roskov et al. (2005) and the International
Legumes Database System (ILDIS). This compilation
effectively represents a checklist of species of Acacia
s.l. (= Senegalia and Vachellia) in Africa, which we
have endeavoured to make as complete as possible.
The list does not include species from Madagascar,
which will be dealt with in a later publication.
I Senegalia Raf., Sylva Tellur:. 119. 1838. - Type:
Senegalia triacantha Raf. [ = Senegalia senegal
(L.) Britton & P.Wilson].
1 Senegalia adenocalyx (Brenan & Exell)
Kyal. & Boatwr., comb. nov. ≡ Acacia adenocalyx Brenan & Exell, Bol. Soc. Brot., Sér. 2,
31: 115, t. 1 fig. D. 1957 - Type: Tanzania,
Tanga Distr., Kange Estate, Faulkner 855 (K,
holotype; BR, PRE, isotypes).
2 Senegalia andongensis (Welw. ex. Hiern.)
Kyal. & Boatwr., comb. nov. ≡ Acacia andongensis Welw. ex. Hiern., Cat. Afr. Pl. Welw. 1:
314. 1896 - Type: Angola, Cuanza Norte,
Pungo Andongo, Welwitsch 1814 (LISU, holotype; BM, K, isotypes).
3 Senegalia ankokib (Chiov.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia ankokib Chiov., Fl.
Somala 2: 190, fig. 116. 1932 - Type: Somalia,
unlocalized, Robecchi-Bricchetti 529 (F);
Migiurtinia, Bender Merajo, Guidotti 35 [not
found Ross, (1979)].
4 Senegalia asak (Forssk.) Kyal. & Boatwr.,
comb. nov. ≡ Mimosa asak Forssk. Fl.
Aegypt.-Arab.: 176. 1775. ≡ Acacia asak
(Forssk.) Willd. Sp. Pl. 4: 1077. 1806 - Type:
Arabia, Forsskal (C, holotype).
5 Senegalia ataxacantha (DC.) Kyal. &
Boatwr., comb. nov. ≡ Acacia ataxacantha
DC., Prodr. 2: 459. 1825 - Types: Senegal,
Bacle s.n. (G-DC); Senegal, Perrottet s.n.
(G-DC).
6 Senegalia brevispica (Harms) Seigler &
Ebinger, Phytologia 92 (1): 93. 2010. ≡ Acacia
brevispica Harms, Notizbl. Bot. Gart. Berl. 8:
370. 1923 - Type: Tanzania, Lushoto Distr.,
507
Kitivo, Holst 606 (B†, holotype; BM,
drawing).
Two subspecies are recognized:
6.a. subsp. brevispica
6.b. subsp. dregeana (Benth.) Kyal. &
Boatwr., comb. nov. ≡ Acacia pennata
var. dregeana Benth., London J. Bot. 1:
516. 1842 - Type: Transkei, Drège s.n.
(K, holotype; P, isotype).
7 Senegalia burkei (Benth.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia burkei Benth., London J.
Bot. 5: 98. 1846 - Type: South Africa, Transvaal, Magaliesberg, Burke s.n. (K, holotype;
BM, PRE, isotypes).
8 Senegalia caffra (Thunb.) P.J.H.Hurter &
Mabb., Plant Book: 1021. 2008. ≡ Mimosa
caffra
Thunb.,
Prodr.
Fl.
Cap.:
92.
1800. ≡ Acacia caffra (Thunb.) Willd., Sp. Pl. 4:
1078. 1806 - Type: Cape Prov., Thunberg s.n.
sub THUNB-UPS 23132 (UPS).
9 Senegalia caraniana (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia caraniana
Chiov., Fl. Somala 1: 166, t. 18. 1929 - Type:
Somalia, Migiurtini, Behèn, Puccioni & Stefanini 704 (FI, holotype; BM, drawing,
isotype).
10 Senegalia chariessa (Milne-Redh.) Kyal. &
Boatwr., comb. nov. ≡ Acacia chariessa MilneRedh., Bull. Misc. Inform. Kew 1933: 143. 1933
- Type: Rhodesia, Bulawayo, Borle 13 (K, holotype; PRE, isotype).
11 Senegalia cheilanthifolia (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia cheilanthifolia
Chiov., Fl. Somala 1: 168. 1929 - Syntypes:
Somalia, Migiurtini, between Erèri Jellehò e
Martisor Dinsai, Puccioni & Stefanini 663
[727] (FI); valle di Bacba, Puccioni & Stefanini
762 [843] (FI); Bacino del Darror, El Uncùd,
Puccioni & Stefanini 1010 [1115] (FI); Obbia,
Magghiòle, Puccioni & Stefanini 479 [531] (FI);
Obbia, between Uarandi e Scillin-Bilhelli, Puccioni & Stefanini 509 [562] (FI).
Two varieties are recorded:
11.a. var. cheilanthifolia
11.b. var. hirtella (Chiov.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia cheilanthifolia var.
hirtella Chiov., Fl. Somala 1: 169. 1929. Type: Somalia, Migiurtini between Dhur
and Hossa Uein, Puccioni & Stefanini
695 [769] (FI, holotype).
12 Senegalia ciliolata (Brenan & Exell) Kyal. &
Boatwr., comb. nov. ≡ Acacia ciliolata Brenan
& Exell in Consp. Fl. Angol. 2: 288. 1956; Bol.
Soc. Brot., Sér. 2, 31: 132. 1957 - Type: Angola,
Cabinda, Belize, Gossweiler 7579 (BM, holotype; COI, K, LISU, isotypes).
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
508
B. KYALANGALILWA ET AL.
13 Senegalia cinerea (Schinz) Kyal. & Boatwr.,
comb. nov. ≡ Acacia cinerea Schinz in Verh.
Bot. Ver. Prov. Brandenb. 30: 240. 1888, non
Spreng. 1826. - Type: South-west Africa,
Amboland, Omatope, Schinz 252 (Z, holo).
= Acacia fleckii Schinz, Mém. Herb. Boissier 1:
108. 1900 - Type: Botswana, Ghanzi, Fleck
412a (Z, holotype).
14 Senegalia circummarginata (Chiov.) Kyal.
& Boatwr., comb. nov. ≡ Acacia circummarginata Chiov., Ann. Bot. (Rome) 13: 394. 1915 Types: Ethiopia, Ogaden, Paoli 794, 913 bis
920, 1010 (FI). (This species is sometimes
regarded as a synonym of Senegalia senegal).
15 Senegalia condyloclada (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia condyloclada
Chiov., Ann. Bot. (Rome) 13: 391. 1915 - Type:
Ethiopia, Ogaden, between Sassaban and Carbaden, Riva & Ruspoli 1079 (FI, holotype).
16 Senegalia densispina (Thulin) Kyal. &
Boatwr., comb. nov. ≡ Acacia densispina
Thulin, Nordic J. Bot. 8(5): 460. 1989 - Type:
Somalia, Galgaud Region, 5 km south-west of
Dusa Mareb on road to Belet Uen, 9 Dec 1985,
Thulin 5647 fruiting (UPS, holotype; MOG,
isotype).
17 Senegalia dudgeonii (Craib ex Holland)
Kyal. & Boatwr., comb. nov. ≡ Acacia dudgeonii Craib ex Holland, Kew Bull., Addit. Ser. 9,
291. Dec 1911, as ‘dudgeoni’ - Types: Nigeria,
Borgu, Dudgeon 58 (K); Kontagora, Dalziel 41
(K).
18 Senegalia eriocarpa (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia eriocarpa
Brenan, Kew Bull. 12: 360. 1957 - Type: Rhodesia, Chirundu, Goodier 81 (K, holotype; BM;
LISC; SRGH, isotypes).
19 Senegalia erubescens (Welw. ex Oliv.) Kyal.
& Boatwr., comb. nov. ≡ Acacia erubescens
Welw. ex Oliv., Fl. Trop. Afr. 2: 343. 1871 Type: Angola, Mocamedes Distr., between
Bumbo and Bruco, Welwitsch 1826 (LISU, holotype; BM, K, isotypes).
20 Senegalia erythrocalyx (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia erythrocalyx
Brenan, Kew Bull. 32: 546. 1978 - Type:
Nigeria, Kano Prov., Home Distr., near Kano
Gata old motor road, Onwundinjoh FHI 24020
(K, holotype, FHO, isotype).
21 Senegalia flagellaris (Thulin) Kyal. &
Boatwr., comb. nov. ≡ Acacia flagellaris
Thulin, Nordic J. Bot. 8(5): 461. 1989 - Type:
Somalia, Bari Region, escarpment south of
Bunder Murayah, Buraha Dhaxsi, 11°38–39′N,
50°29–32′E, 16-17 Nov 1986, Thulin & Warfa
5844 (UPS, holotype; FT, K, MOG, isotypes).
22 Senegalia fumosa (Thulin) Kyal. & Boatwr.,
comb. nov. ≡ Acacia fumosa Thulin, Nordic J.
Bot. 25: 272. 2007 - Type: Ethiopia, Somali
National Regional State, Harerge, 22 km from
Qarsonney along road to Kebri Dehar, Thulin,
Kaariye & Wilhelmi 11136 (ETH, holotype; K,
UPS isotypes).
23 Senegalia galpinii (Burtt Davy) Seigler &
Ebinger, Phytologia 92 (1): 93. 2010. ≡ Acacia
galpinii Burtt Davy, Bull. Misc. Inform. Kew
1922: 326. 1922 - Type: South Africa, Transvaal, banks of Bad-zyn-loop River, Mosdene
Estate, Naboomspruit, Galpin 483 M (K, holotype; BM, GRA, PRE, isotypes).
24 Senegalia goetzei (Harms) Kyal. & Boatwr.,
comb. nov. ≡ Acacia goetzii Harms in Bot.
Jahrb. Syst. 28: 395. 1900, nom. cons. - Type:
Tanzania, Kilosa Distr., Kidodi, Goetze 387 (B†,
holotype; E, K, isotypes).
Two subspecies are recognized:
24.a. subsp. goetzei
24.b. subsp. microphylla (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia goetzii
subsp. microphylla Brenan, Kew Bull.
11: 204. 1956 - Type: Malawi, Mombera
Distr., Njakwa to Fort Hill, Greenway
6393 (K, holotype; EA, PRE, isotypes).
25 Senegalia gourmaensis (A.Chev.) Kyal. &
Boatwr., comb. nov. ≡ Acacia gourmaensis
A.Chev., Bull. Soc. Bot. Fr. 58, Mém. 8d: 167.
1912 - Type: Benin/Upper Volta, Gourma
Prov., between Konkobiri and Diagapa, Chevalier 24364 (P, holotype; K, isotype).
26 Senegalia hecatophylla (Steud. ex. A.Rich.)
Kyal. & Boatwr., comb. nov. ≡ Acacia hecatophylla Steud. ex. A.Rich., Tent. Fl. Abyss. 1:
242. 1847 - Types: Ethiopia, without locality,
Schimper 628 (BM, FI, P, Z); Schimper 884
(BM, FI, K, OXF, P, Z).
27 Senegalia hereroensis (Engl.) Kyal. &
Boatwr., comb. nov. ≡ Acacia hereroensis
Engl., Bot. Jahrb. Syst. 10: 20. 1888 - Type:
south-west Africa, Karibib Distr., Otjimbingwe,
Marloth 1331 (B†, holotype; GRA, pro parte;
PRE, isotype).
28 Senegalia kamerunensis (Gandoger) Kyal.
& Boatwr., comb. nov. ≡ Acacia kamerunensis
Gandoger, Bull. Soc. Bot. Fr. 60: 459. 1913 Type: Cameroon, between Victoria and Bota,
Winkler 447 (LY, holotype; K, photograph).
29 Senegalia kraussiana (Meisn. ex Benth.)
Kyal. & Boatwr., comb. nov. ≡ Acacia kraussiana Meisn. ex Benth., London J. Bot. 1: 515.
1842 - Type: South Africa, Natal Umlaas
River, Durban, Krauss 198 (K, holotype; BM,
FI, OXF, PRE, TCD, Z, isotypes).
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
30 Senegalia laeta (R.Br. ex Benth.) Seigler &
Ebinger in Phytologia 91 (1): 27. 2009 ≡ Acacia
laeta R.Br. ex Benth., London J. Bot. 1: 508.
1842 - Type: Ethiopia, prope montes Taranta,
Salt s.n. (BM, holotype).
31 Senegalia latistipulata (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia latistipulata
Harms, Bot. Jahrb. Syst. 51: 367. 1914 - Type:
Tanzania, Kwa-Mkopo on the Rovuma River,
Busse 1031 (B†; BM, drawing; EA; isotype);
Uzaramo Distr., Sthulmann 7025 (B†) and
7048 (B†).
32 Senegalia lujai (De Wild.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia lujai De Wild., Bull. Soc.
r. Bot. Belg. 39: 99. 1900, as ‘lujae’ - Type:
Zaire (Democratic Republic of Congo), Kasai,
Bena-Dibele, Luja 273 (BR, holotype).
33 Senegalia macrostachya (Reichenb. ex DC.)
Kyal. & Boatwr., comb. nov. ≡ Acacia macrostachya Reichenb. ex DC., Prodr. 2: 459. 1825
- Type: Senegal, Sieber 44 (K, MEL, OXF, BM,
drawing).
34 Senegalia manubensis (J.H.Ross) Kyal. &
Boatwr., comb. nov. ≡ Acacia manubensis
J.H.Ross, Bothalia 11: 292. 1974 - Type:
Somalia, Manúb, Newbould 1080 (K, holotype).
35 Senegalia mellifera (Vahl) Seigler &
Ebinger, Phytologia. 92 (1): 94. 2010. ≡ Mimosa
mellifera
Vahl,
Symb.
Bot. 2:
103.
1791. ≡ Acacia mellifera (Vahl) Benth., London
J. Bot. 1: 507. 1842 - Type: Arabia, Surdud and
elsewhere, Forsskål (C, holotype).
Two subspecies are recognized:
35.a. subsp. mellifera
35.b. subsp. detinens (Burch.) Kyal. &
Boatwr., comb. nov. ≡ Acacia detinens
Burch., Trav. 1: 310. 1822. ≡ Acacia mellifera subsp. detinens (Burch.) Brenan,
Bull. Misc. Inform. Kew 11: 191. 1956 Type: South Africa, Cape Province,
Prieska Division, Zand Valley, Burchell
1628 (K, holotype; PRE, isotype).
36 Senegalia moggii (Thulin & Tardelli) Kyal. &
Boatwr., comb. nov. ≡ Acacia moggii Thulin &
Tadelli, Willdenowia 17: 125. 1988 - Type:
Somalia, Hiran Region, 320 km on the
Mogadisho-Belet Uein road, Moggi, Tardelli &
Warfa 54 (FT, holotype; B, FT, UPS, isotypes).
37 Senegalia montigena (Brenan & Exell) Kyal.
& Boatwr., comb. nov. ≡ Acacia montigena
Brenan & Exell, Kew Bull. 21: 480. 1968 Type: Uganda, Kigezi Distr., Murole Hill,
Purseglove 2693 (K, holotype, EA, isotype).
38 Senegalia montis-usti (Merxm. & A.Schreiber) Kyal. & Boatwr., comb. nov. ≡ Acacia
montis-usti Merxm. & A.Schreiber, Bull. Jard.
39
40
41
42
43
44
45
46
509
Bot. État Brux. 27: 270, t. 8. 1957 - Type:
Namibia, Brandberg, Welwitsch-Tal, Von
Wettstein 95 (M, holotype).
Senegalia nigrescens (Oliv.) P.J.H. Hurter,
Plant book: 1021. 2008. ≡ Acacia nigrescens
Oliv., Fl. Trop. Afr. 2: 340. 1871 - Type:
Malawi, near Mitonda, Shire River, Kirk s.n.
(K, holotype).
Senegalia ochracea (Thulin & Hassan) Kyal.
& Boatwr., comb. nov. ≡ Acacia ochracea
Thulin & Hassan, Fl. Somalia 1: 373. 1993 Type: Somalia, Gedo and Bay, 7 km on the road
between Awdiinle and Qansaxdheere, Hassan
127 (FHO, holotype; MOG; isotype).
Senegalia ogadensis (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia ogadensis
Chiov., Ann. Bot. (Rome) 13:393. 1915 - Syntypes: Somalia, Ogaden, Robecchi-Brichetti 594
(FI); between Bardera and Marda, Paoli 811
(FI), Paoli 812 (FI).
Senegalia oliveri (Vatke) Kyal. & Boatwr.,
comb. nov. ≡ Acacia oliveri Vatke, Öesterr.
Bot. Z. 30: 274. 1880. excl. descr. leguminis. Type: Ethiopia, Danakil territory, Hildebrandt
729c (BM, drawing).
Senegalia pentagona (Schumach.) Kyal. &
Boatwr., comb. nov. ≡ Mimosa pentagona
Schumach.,
Beskr.
Guin.
Pl.:
324.
1827. ≡ Acacia pentagona (Schumach.) Hook.f.,
Nig. Fl. 331. 1849 - Type: Ghana, Jadofa,
Thonning (C, holotype; K, photograph).
Senegalia persiciflora (Pax) Kyal. &
Boatwr., comb. nov. ≡ Acacia persiciflora Pax,
Bot. Jahrb. Syst. 39: 624. 1907 - Type: Ethiopia, West Shoa, Urga valley, Rosen s.n.
(BRSL?, holotype).
Senegalia petrensis (Thulin) Kyal. &
Boatwr., comb. nov. ≡ Acacia petrensis Thulin,
Kew Bull. 58: 495. 2003 - Type: Somalia,
Thulin 11000 (UPS, holotype; K, isotype).
Senegalia polyacantha (Willd.) Seigler &
Ebinger, Phytologia 91(1): 28. 2009. ≡ Acacia
polyacantha Willd., Sp. Pl. 4: 1079. 1806 Type: Eastern India, Herb. Willdenow 19166
(B, holotype; K; isotype).
Two subspecies are recognized:
46.a. subsp. polyacantha
46.b. subsp. campylacantha (Hochst. ex.
A.Rich.) Kyal. & Boatwr., comb.
nov. ≡ Acacia campylacantha Hochst. ex.
A.Rich., Tent. Fl. Abyss. 1: 242.
1847. ≡ Acacia polyacantha Willd. subsp.
compylacantha (Hochst. ex. A.Rich.)
Brenan, Kew Bull. 11: 195. 1956 - Syntypes: Ethiopia, Mai Dogale, Schimper
639 (BM, E, FI, K, OXF, P, Z); Dschelad-
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47
48
49
50
51
52
B. KYALANGALILWA ET AL.
scheranne, Schimper 893 (BM, E, FI, K,
OXF, P, Z).
Senegalia pseudonigrescens (Brenan &
J.H.Ross)
Kyal.
&
Boatwr.,
comb.
nov. ≡ Acacia pseudonigrescens Brenan &
J.H.Ross, Bothalia 11: 293. 1974. - Type:
Ethiopia, 8 km west of Mustahil on western
track to Kelafo, M.G Gilbert 2129 (K, holotype).
Senegalia robynsiana (Merxm. & A.Schreiber) Kyal. & Boatwr., comb. nov. ≡ Acacia
robynsiana Merxm. & A.Schreiber, Bull. Jard.
Bot. État Brux. 27: 268, t. 7. 1957 - Type:
Namibia, Outjo Distr., Grootberg-Hang, Walter
2/197 (M, holotype).
Senegalia rovumae (Oliv.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia rovumae Oliv., Fl. Trop.
Afr. 2: 353. 1871 - Type: Tanzania or Mozambique, Rovuma Bay, Kirk s.n. (K, holotype).
= Acacia macalusoi Mattei, Boll. Orto Bot.
Giard. Col. Palermo 7: 94. 1908. – Type:
Somalia, Guimbo, Macaluso 65 pro parte
quoad specim. fructifera (?PAL, lectotype).
Senegalia schlechteri (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia schlechteri
Harms, Bot. Jahrb. Syst. 51: 367. 1967 - Type:
Mozambique, Ressano Garcia, Schlechter
11901 (B†, holotype; Z, isotype).
Senegalia schweinfurthii (Brenan & Exell)
Seigler & Ebinger, Phytologia 92 (1): 94.
2010. ≡ Acacia schweinfurthii Brenan & Exell,
Bol. Soc. Brot., Sér. 2, 31: 128 1957 - Type:
Sudan, Gubbiki, Schweinfurth 2206 (BM, holotype; K, P, Z, isotypes).
Two varieties are recognized:
51.a. var. schweinfurthii
51.b. var. sericea (Brenan & Exell) Kyal. &
Boatwr., comb. nov. ≡ Acacia schweinfurthii Brenan & Exell var. sericea
Brenan & Exell, Bol. Soc. Brot., Sér. 2,
31: 131. 1957 - Type: Tanzania,
Mpwapwa, Mrs Hornby 56 (K, holotype;
EA, isotype).
Senegalia senegal (L.) Britton, Sci. Surv.
Porto Rico & Virgin Islands 5: 538.
1930. ≡ Mimosa senegal L., Sp. Pl. 1: 521. 1753
- Type: Senegal, Herb. Adanson 16899 (P,
neotype, designated by Ross, 1979).
Four varieties are recognized:
52.a. var. senegal
52.b. var. kerensis (Schweinf.) Kyal. &
Boatwr., comb. nov. ≡ Acacia senegal
var. kerensis Schweinf., Bull. Herb. Boissier 4, app. 2: 216. 1896 - Syntypes:
Ethiopia, near Keren, Schweinfurth 745
(B†; K); Bogu valley, Schweinfurth 741
(B†); near Djuffa, Schweinfurth 998 (B†).
53
54
55
56
57
58
59
52.c. var. rostrata (Brenan) Kyal. & Boatwr.,
comb. nov. ≡ Acacia senegal var. rostrata
Brenan, Kew Bull. 8: 99. 1953 - Type:
South Africa, Transvaal, Soutpansberg
Distr., Dongola Reserve, Verdoon 2264 (K,
holotype; PRE, isotype).
52.d. var. leiorachis (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia senegal
var. leiorachis Brenan, Kew Bull. 8: 98.
1953 - Type: Tanzania, Tanga Prov.,
Pare Distr., Same, Greenway 2192 (K,
holotype; EA, FHO, isotypes).
Senegalia senegalensis (Forssk.) Kyal. &
Boatwr., comb. nov. ≡ Mimosa senegalensis
Forssk., Fl. Aegypt.-Arab. 176. 1775, non
Houtt. 1774 - Syntypes: from Arabia, Forsskål
(C).
= Acacia hamulosa Benth., Hook., London. J.
Bot. 1: 509. 1842 - Type: Saudi Arabia, Jiddah
(Gedda), S. Fischer 72 (K, holotype).
Senegalia somalensis (Vatke) Kyal. &
Boatwr., comb. nov. ≡ Acacia somalensis
Vatke, Öesterr. Bot. Z. 30: 274. 1880 - Type:
Somalia, near Meid, Hildebrandt 1396 (BM; K,
isotype).
Senegalia tanganyikensis (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia tanganyikensis
Brenan, Kew Bull. 11: 195. 1956 - Type: Tanzania, Shyianga Distr., unlocalized, B.D. Burtt
6427 (K, holotype; BM; isotype).
Senegalia taylorii (Brenan & Exell) Kyal. &
Boatwr., comb. nov. ≡ Acacia taylorii Brenan
& Exell, Bol. Soc. Brot. Sér. 2, 31: 139. 1957 Type: Tanzania, Lindi Distr., 6.5 km N. of
Lindi, Milne-Redhead & Taylor 7588 (K, holotype; BM, isotype).
Senegalia tephrodermis (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia tephrodermis
Brenan, Kew Bull. 32: 549. 1978 - Type: Tanzania, Bagamoyo Distr., Bana Forest Reserve,
Mgaza 779 (K, holotype).
Senegalia thomasii (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia thomasii Harms,
Bot. Jahrb. Syst. 51: 366. 1914 - Types: Kenya,
Kitui Distr., Ikhuta, F. Thomas III 127 (B†,
holotype; BM, drawing, isotype); Machakos
Distr., mile 138 and 129 from Mombasa on
main Nairobi road, near Kenani, Verdcourt
2390 (K, neotype, designated by Ross, 1979;
PRE; isotype).
Senegalia venosa (Hochst. ex Benth.) Kyal. &
Boatwr., comb. nov. ≡ Acacia venosa Hochst.
ex Benth., London J. Bot. 5: 98. 1846 - Type:
Ethiopia, Schire Prov., Dschogarti, Schimper
524 (K, holotype; BM, E, FI, MEL, OXF, P, Z,
isotypes).
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RECLASSIFICATION OF AFRICAN ACACIA SPECIES
60 Senegalia welwitschii (Oliv.) Kyal. &
Boatwr., comb. nov. ≡ Acacia welwitschii Oliv.,
Fl. Trop. Afr. 2: 341. 1871 - Type: Angola,
Luanda Distr., Barra de Bengo, entre Mutolo e
Cacuaco, prox. De Quicuxe, Welwitsch 1806
(LISU, lectotype; BM, K, P, isotypes).
Two subspecies are recognized:
60.a. subsp. welwitschii
60.b. subsp. delagoensis (Harms ex Burtt
Davy)
Kyal.
&
Boatwr.,
comb.
nov. ≡ Acacia welwitschii subsp. delagoensis (Harms) J.H.Ross & Brenan,
Kew Bull. 21: 67. 1967 ≡ Acacia delagoensis Harms ex Burtt Davy, Bot.
Jahrb. Syst. 51: 367. 1914 - Type:
Mozambique,
Umbulezi,
Schlechter
11718 (B†, holotype; BM, K, Z, isotypes).
61 Senegalia zizyphispina (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia zizyphispina
Chiov., Fl. Somalia 1: 167. 1929 - Types:
Somalia, between Garass-Hebla-Aden and
Jesomma, Puccioni & Stefanini 152 (FI; BM,
drawing); between Avorrei and Bulo-Burti,
Puccioni & Stefanini 171 (FI; BM, drawing,
isotypes).
II Vachellia Wight & Arn., Prodr. Fl. Ind. Orient. 1:
272. 1834. - Type: Vachellia farnesiana (L.) Wight
& Arn.
1 Vachellia abyssinica (Hochst. ex. Benth.)
Kyal. & Boatwr., comb. nov. ≡ Acacia abyssinica Hochst. ex. Benth., London J. Bot. 5: 97.
1846 - Type: Ethiopia, near Mendel, Schimper,
Sect. 3, 1813 (K, holotype; BM, M, P, isotypes).
Two subspecies are recognized:
1.a. subsp. abyssinica
1.b. subsp. calophylla (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia abyssinica
Hochst. ex. Benth. subsp. calophylla
Brenan, Kew Bull. 12: 82. 1957 - Type:
Kenya,
South
Kavirondo
Distr.,
Mugunga near Kisii, Greenway 7860 (K,
holotype; EA, PRE, isotypes).
2 Vachellia amythethophylla (Steud. ex
A.Rich.) Kyal. & Boatwr., comb. nov. ≡ Acacia
amythethophylla Steud. ex A.Rich., Tent. Fl.
Abyss. 1: 245. 1847 - Type: Ethiopia, near
Djeladjeranne, Schimper 887 (P, holotype; BM,
K, OXF, isotypes).
3 Vachellia ancistroclada (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia ancistroclada
Brenan, Kew Bull. 13: 412. 1959 - Type:
Kenya, Masai Distr., Amboseli, Knight &
Thomas H 344/58 (K, holotype).
4 Vachellia antunesii (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia antunesii
Harms, Bot. Jahrb. Syst. 30: 76. 1901 - Syn-
511
types: Angola, Huila Distr., Huila, Antunes 28
(B†, BM, LISC); Kamunguo, Dekindt 219 (B†).
5 Vachellia arenaria (Schinz) Kyal. & Boatwr.,
comb. nov. ≡ Acacia arenaria Schinz, Mém.
Herb. Boissier 1: 105. 1900 - Types: south-west
Africa,
Ovamboland,
Olukonda-Oshiheke,
Schinz 2071 (Z); Amboland, ‘Omatope’, Schinz
2072 (Z).
6 Vachellia bavazzanoi (Pichi-Sermolli) Kyal.
& Boatwr., comb. nov. ≡ Acacia bavazzanoi
Pichi-Sermolli, Miss. Stud. Lago Tana, Ric.
Bot. 1: 54, tt.7, 8. 1951 – Type: Ethiopia,
Gorgorà, Pichi-Sermolli 2253 (FI, holotype; K,
isotype).
7 Vachellia borleae (Burtt Davy) Kyal. &
Boatwr., comb. nov. ≡ Acacia borleae Burtt
Davy, Kew Bull. 1922: 325. 1922 - Type:
Mozambique, Maputo (Lourenco Marques),
Borle 271 (PRE, holotype; FHO, isotype).
8 Vachellia bricchettiana (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia bricchettiana
Chiov., Ann. Bot. (Rome) 13: 396. 1915 - Type:
Somalia, Ogaden, Robecchi Bricchetti 533 (FI,
holo).
9 Vachellia bullockii (Brenan) Kyal. &
Boatwr.,
comb.
nov. ≡ Acacia
bullockii
Brenan, Kew Bull. 12: 77. 1957 - Types: Tanzania, Buha Distr., Kaberi mbuga, Bullock
3144 (K, holotype).
Two varieties are recognized:
9.a. var. bullockii
9.b. var. induta (Brenan) Kyal. & Boatwr.,
comb. nov. ≡ Acacia bullockii Brenan
var. induta Brenan, Kew Bull. 12: 78.
1957 - Type: Tanzania, Kigoma Distr.,
Tandala in Uvinza, C.H.N. Jackson 117
(K, holotype; BM, isotype).
10 Vachellia burttii (Bak. f.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia burttii Bak. f., J. Bot. 71:
342. 1933 - Type: Tanzania, Kahama Distr.,
9 km along Shinyanga road, B.D. Burtt 4501
(BM, holotype; EA, FHO, K, isotypes).
11 Vachellia bussei (Harms ex Sjöstedt) Kyal. &
Boatwr., comb. nov. ≡ Acacia bussei Harms ex
Sjöstedt, Schwed. Zool. Exped. Kilimanjaro 8:
117-118 1908 - Syntypes: Tanzania, Lushoto
District, Mazinde, by Kisiwani road, Busse 361
(B†, BM, K); Lushoto/Pare Districts, between
Usambara Mts and Kihurio, Pare Districts,
Engler 1506 (B†, K, drawing); Pare District,
between Kihurio and Gonja, Zimmermann
1758 (B†, EA).
12 Vachellia cernua (Thulin & Hassan) Kyal. &
Boatwr., comb. nov. ≡ Acacia cernua Thulin &
Hassan, Nordic J. Bot. 16: 303. 1996 - Type:
Somalia, Sanaag Region, escarpment S of
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512
13
14
15
16
17
18
19
20
B. KYALANGALILWA ET AL.
Laasqoary, Thulin, Dahir & Hassan 9188
(UPS, holotype; FT, K, isotypes).
Vachellia davyi (N.E.Br.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia davyi N.E.Br., Kew Bull.
1908: 161. 1908 - Syntypes: South Africa,
Transvaal, Houtbosch, Rehmann 6276 (BM, K,
Z); Burtt Davy 5132 (PRE); Soutpansberg,
Junod sub Herb. T.D.A. no. 1323 (PRE), Swaziland, near Manzini, Burtt Davy 3024 (BM,
holotype; FHO).
Vachellia dolichocephala (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia dolichocephala
Harms, R. Ist. Bot. Roma 7: 86. 1897 - Type:
Ethiopia, Galla Sidamo, between Rogono and
Goba Duaya, Riva 599 (FI, holotype; K,
isotype).
Vachellia drepanolobium (Harms ex Sjöstedt)
P.J.H.Hurter,
Plant
book:
1021.
2008. ≡ Acacia drepanolobium Harms ex
B.Y.Sjöstedt, Wissensch. Ergebn. Schwed. Zool.
Exped. Kilimanjaro 8: 116–117. 1908 - Type:
Tanzania, Kilimanjaro, between Kwagogo and
Moshi, Engler 1688 (B†, holotype; K, drawings,
isotype).
Vachellia dyeri (P.P.Swartz) Kyal. & Boatwr.,
comb. nov. ≡ Acacia dyeri P.P.Swartz, CoatesPalgrave, Trees of southern Afr.: 19. 2002 Type: South Africa, Eastern Cape, Butterworth
District, Kei Mouth, Robbertse 871 (PRE, holotype; PRU, isotype).
Vachellia ebutsiniorum (P.J.H.Hurter) Kyal.
& Boatwr., comb. nov. ≡ Acacia ebutsiniorum
P.J.H.Hurter, Bothalia 34(1): 42. 2004 - Type:
South Africa, Mpumalanga, Ebutsini tribal
land, Farm Tothietoe 7 JT, Hurter 133 (PRE,
holotype; K, NBG, PRU, isotypes).
Vachellia edgeworthii (T.Anders.) Kyal. &
Boatwr., comb. nov. ≡ Acacia edgeworthii
T.Anders., J. Linn. Soc. Bot. 5, suppl.: 18. 1860.
- Syntypes: Aden, Edgeworth, Hooker &
Thomson s.n. (K); T. Anderson s.n. (K).
Vachellia elatior (Brenan) Kyal. & Boatwr.,
comb. nov. ≡ Acacia elatior Brenan, Kew Bull.
12: 94. 1957 - Type: Kenya, Tana River,
Garissa, Greenway 8857 (K, holotype; FHO,
isotype).
Two subspecies are recognized:
19.a. subsp. elatior
19.b. subsp. turkanae (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia elatior
Brenan subsp. turkanae Brenan, Kew
Bull. 12: 95. 1957 - Type: Kenya, northwest Turkana, Lodwar, Hemming CFH
250 (K, holotype; EA, FHO, isotypes).
Vachellia erioloba (E.Mey.) P.J.H.Hurter,
Mabberley’s plant book: 1021. 2008. ≡ Acacia
21
22
23
24
erioloba E. Mey., Comm 1: 171. 1836. - Type:
South Africa, Namaqualand [not found by
Ross, (1979)] Transvaal, Wolmaransstad Distr.,
between Kommandodrif and Makwassie, J. W.
Morris 1042 (K, neotype, designated by Ross,
1979; PRE, isotype).
Note: Vachellia erioloba (E.Mey.) Seigler &
Ebinger, Phytologia 92(1): 94. 2010 is an
isonym of Vachellia erioloba (E.Mey.) P.J.H.Hurter and has no nomenclatural status
according to the Code (McNeill et al., 2006),
art. 6.3 note 2 ‘when the same name, based on
the same type, has been published independently at different times by different authors,
then only the earliest of these “isonyms” has
nomenclatural status.’
Vachellia erythrophloea (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia erythrophloea
Brenan, Kew Bull. 12: 76. 1957 - Type: Tanzania, Tabora Distr., Kakoma, Glover 186 (K,
holotype; EA, isotype)
Vachellia etbaica (Schweinf.) Kyal. &
Boatwr., comb. nov. ≡ Acacia etbaica Schweinf, Linnaea 35: 330. 1867–1868 - Types:
Sudan, Soturba Mts, Schweinfurth 1994 (K)
and 1995 (BM, K, P).
Four subspecies are recognized:
22.a. subsp. etbaica
22.b. subsp. uncinata (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia etbaica
Schweinf. subsp. uncinata Brenan, Kew
Bull. 12: 91. 1957 - Type: Somalia,
Erigavo, McKinnon 8/220 (K, holotype).
22.c. subsp. australis (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia etbaica
Schweinf. subsp. australis Brenan, Kew
Bull. 12: 92. 1957 - Type: Tanzania,
Tanga Distr., Ngomeni, Greenway 7034
(K, holotype; EA, FDHO, isotypes).
22.d. subsp. platycarpa (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia etbaica
Schweinf. subsp. platycarpa Brenan,
Kew Bull. 12: 93. 1957 - Type: Kenya,
Northern Frontier Province, Moyale,
Gillett 13641 (K, holotype; BM, EA, isotypes).
Vachellia exuvialis (Verdoorn) Kyal. &
Boatwr., comb. nov. ≡ Acacia exuvialis Verdoorn, Bothalia 6: 154. 1951 - Type: South
Africa, Transvaal, Nelspruit Distr., Kruger
National Park, 25.6 km west of Skukuza, Codd
& Verdoorn 5464 (PRE, holotype).
Vachellia farnesiana (L.) Wight & Arn.,
Prodr. Fl. Ind. Orient. 1: 272. 1834. ≡ Mimosa
farnesiana L., Sp. Pl.: 521. 1753. ≡ Acacia farnesiana (L.) Willd., Sp. Pl. 4: 1083. 1806 -
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RECLASSIFICATION OF AFRICAN ACACIA SPECIES
25
26
27
28
29
Type: Aldinus, Exactissima description rariorum plantarum Romae, Horto Franesiano 4.
1625. (lectotype, designated by Ross, 1979).
Vachellia fischeri (Harms) Kyal. & Boatwr.,
comb. nov. ≡ Acacia fischeri Harms, Bot.
Jahrb. Syst. 51: 365. 1914 - Syntypes: Tanzania, without locality, Fischer 157 (B†, BM);
‘Manjanga Bach’ (probably Manyonga River),
Stuhlmann 672 (B†); Kondoa Distr., near
Salia, B. D. Burtt 1131 (BM, neotype; FHO, K,
isotypes).
Vachellia flava (Forssk.) Kyal. & Boatwr.,
comb. nov. ≡ Mimosa flava Forssk., Fl.
Aegypt.-Arab. 176. 1775 - Type: Arabia, Forsskal (C, holo).
= Acacia ehrenbergiana Hayne, Arzneyk. Gebr.
Gewächse 10: t. 29. 1827 - Type: Sudan,
Dongola, Ehrenberg & Hemprich [not found by
Ross, (1979)].
Vachellia gerrardii (Benth.) P.J.H.Hurter,
Mabberley’s plant book: 1021. 2008. ≡ Acacia
gerrardii Benth., Trans. Linn. Soc. London,
Bot. 30: 508. 1875 - Type: South Africa, Natal,
without locality, Gerrard 1702 (K, holotype;
BM, TCD, isotypes).
Three varieties are recognized within the
African V. gerrardii subsp. gerrardii:
27.a. subsp. gerrardii var. gerrardii
27.b. subsp. gerrardii var. latisiliqua
(Brenan) Kyal. & Boatwr., comb.
nov. ≡ Acacia gerrardii Benth. subsp. gerrardii var. latisiliqua Brenan, Kew Bull.
12: 369. 1958 - Type: Kenya Machakos
Distr., Sir F. Wilson’s farm near Machakos, Trapnell 2215 (K, holotype).
27.c. subsp. gerrardii var. calvescens
(Brenan) Kyal. & Boatwr., comb.
nov. ≡ Acacia gerrardii Benth. subsp.
gerrardii var. calvescens Brenan, Kew
Bull. 12: 370. 1958 - Type: Tanzania,
Mbulu Distr., near Mbulu, Eggeling 6689
(K, holotype; EA, isotype).
Vachellia grandicornuta (Gerstner) Seigler
& Ebinger, Phytologia 92(1): 94. 2010. ≡ Acacia
grandicornuta Gerstner in J. S. African Bot. 4:
55. 1938 - Syntypes: South Africa, Natal,
‘Flowered at Emkunzana and Mkuzi Drift
between Nongoma and Magidu’, 6 Jan 1936,
Gerstner 2870 (BOL); ‘fruits found at same
place and at lower Pongola’, 13 May 1936,
Gerstner 2870 (BOL).
Vachellia gummifera (Willd.) Kyal. &
Boatwr., comb. nov. ≡ Acacia gummifera
Willd. Sp. Pl. 4: 1056. 1806 - Type: Morocco,
near Mogador, Broussonet s.n. sub Herb.
Willdenow 19125 (B, holotype).
513
30 Vachellia haematoxylon (Willd.) Seigler &
Ebinger, Phytologia 92(1): 94. 2010. ≡ Acacia
haematoxylon Willd., Enum. Hort. Berol. 1056.
1809 - Type: South Africa, Cape Province,
Lichtenstein s.n. sub Herb. Willdenow 19186
(B, holotype).
31 Vachellia hebeclada (DC.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia hebeclada DC., Cat. Hort.
Monsp. 73: 1813 - Type: South Africa, Cape
Province, Kuruman Distr., between Kuruman
and the Matlowing River, Burchell 2267 (G,
holotype; K, PRE, isotypes).
Three subspecies are recognized:
31.a. subsp. hebeclada
31.b. subsp. chobiensis (O.B.Miller) Kyal. &
Boatwr., comb. nov. ≡ Acacia stolonifera
var. chobiensis O.B.Miller, J. S. African
Bot. 18: 25. 1952. ≡ Acacia hebeclada
subsp. chobiensis (O.B.Miller) A.Schreiber, Mitt. Bot. Staatssamml. München
6: 251. 1966 - Type: Botswana, Serondela, O.B. Miller B/1069 (K, holotype).
31.c. subsp. tristis (A.Schreiber) Kyal. &
Boatwr., comb. nov. ≡ Acacia hebeclada
DC. subsp. tristis A.Schreiber, Mitt. Bot.
Staatssamml. München 6: 251. 1966 Type: Angola, Huila Distr., between
Lopolo e Ferrão da Sola, Welwitsch 1829
(LISU, holotype; BM, K, isotypes).
32 Vachellia hockii (De Wild.) Seigler &
Ebinger, Phytologia 92(1): 94. 2010. ≡ Acacia
hockii De Wild., Reprium nov. Spec. Regni veg.
11: 502. 1913 - Type: Zaire (Democratic Republic of Congo), Katanga, Luafu valley, Hock s.n.
(BR, holotype).
33 Vachellia horrida (L.) Kyal. & Boatwr.,
comb. nov. ≡ Mimosa horrida L., Sp. Pl. 1:
521. 1753. ≡ Acacia horrida (L.) Willd., Sp.
Pl. 4: 1082. 1806. non sensu auct. mult. - Type:
Plukenet, Phytographia t. 121.1962 – holo:
backed by the specimen drawn by Plukenet,
Herb. Sloane: 95, fol. 3 (BM).
Two subspecies are recognized:
33.a. subsp. horrida
33.b. subsp. benadirensis (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia horrida
(L.) Willd. subsp. benadirensis (Chiov.)
Hillcoat & Brenan, Kew Bull. 13: 40.
1958 ≡ A. bussei Harms ex Sjöstedt var.
benadirensis Chiov., Miss. StefaniniPaoli Bot. 72. 1916. - Type: Somalia,
Mogadishu [Mogadiscio], Paoli 94 (FI,
lectotype; K, photograph, isotype).
34 Vachellia karroo (Hayne) Banfi & Galasso,
Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat.
Milano 149(1): 149. 2008. ≡ Acacia karroo
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37
38
39
B. KYALANGALILWA ET AL.
Hayne, Arzneyk. Gebr. Gewächse 10: t. 33.
1827 - Type: South Africa, Cape Province,
locality unknown, Herb. Willdenow 19184 fol. 2
(B, lectotupe; PRE, isotype).
Vachellia kirkii (Oliv.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia kirkii Oliv., Fl. Trop. Afr.
2: 350. 1871 - Type: Zambia, Southern Prov.,
Batoka country, Kirk s.n. (K, holotype).
Two subspecies are recognized, and two varieties in the typical subspecies:
35.a 35.a.1. subsp. kirkii var. kirkii
35.a.2. subsp. kirkii var. sublaevis
(Brenan) Kyal. & Boatwr., comb.
nov. ≡ Acacia kirkii subsp. kirkii
var. sublaevis Brenan, Kew Bull.
12: 363. 1958 - Type: Uganda,
Acholi Distr., Aswa River, GuluKitgum road, Eggeling 775 in
F.H. 1161 (K, holotype; EA,
isotype).
35.b. subsp. mildbraedii (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia mildbraedii Harms, Zentr. Afr. Exped. 1907-1908,
2: 234. 1911. ≡ Acacia kirkii subsp. mildbraedii (Harms) Brenan, Kew Bull. 12:
364. 1958 - Types: Rwanda, between
Gisenyi and Mpororo, Mildbraed 343
(B†, BM, drawing); Zaïre (Democratic
Republic of Congo), Kwenda, Mildbraed
1887 (B†); Tanzania, Bukoba Distr.,
between Itara and Kakindu, by the
Kagera R., Holtz 1712 (B†).
Vachellia kosiensis (P.P.Sw. ex Coates
Palgr.) Kyal. & Boatwr., comb. nov. ≡ Acacia
kosiensis P.P.Sw. ex Coates Palgr., CoatesPalgrave’s Trees of southern Afr.: 19. 2002 Type: South Africa, Kwa-Zulu Natal, Lake
Sibayi, Vorster 2720 (PRE, holotype; K,
isotype).
Vachellia lahai (Steud. & Hochst. ex. Benth.)
Kyal. & Boatwr., comb. nov. ≡ Acacia lahai
Steud. & Hochst. ex. Benth., London J. Bot. 1:
506. 1842 - Type: Ethiopia, Tigré, near Adua
(Adowa), Schimper 119 (K, holotype; BM, FI,
OXF, P, Z, isotypes).
Vachellia lasiopetala (Oliv.) Kyal. &
Boatwr., comb. nov. ≡ Acacia lasiopetala Oliv.,
Fl. Trop. Afr. 2: 346. 1871 - Type: Malawi,
Mpemba Mt, Kirk s.n. (K, holotype).
Vachellia latispina (J.E.Burrows & S.M.Burrows) Kyal. & Boatwr., comb. nov. ≡ Acacia
latispina J.E.Burrows & S.M.Burrows, Bothalia 39: 222. 2009. – Type: Mozambique, Cabo
Delgado Province, 14.8 km from the main
Pemba–Metoro road, on road to Mecufi,
13°11′13″S, 40°33′10″E, 23 December 2006,
40
41
42
43
44
45
46
47
J.E. Burrows & S.M. Burrows 9764 (PRE,
holotype; BNRH, K, LMA, isotypes).
Vachellia leucospira (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia leucospira
Brenan, Kew Bull. 13: 407. 1959 - Type:
Somalia, near Galkayu (Galcaio), Bond &
Pechanec 65 (EA, holotype; K, isotype).
Vachellia luederitzii (Engl.) Kyal. &
Boatwr., comb. nov. ≡ Acacia luederitzii Engl.,
Bot. Jahrb. Syst. 10:23, t. 3B (July 1888) pro
parte quoad specim. Marloth 1328 - Type:
South West Africa, Otjimbingwe, Marloth 1328
(PRE, lectotype; GRA, M, OXF, isotypes).
Two varieties are recognized:
41.a. var. luederitzii
41.b. var. retinens (Sim) Kyal. & Boatwr.,
comb. nov. ≡ Acacia retinens Sim, For.
Fl.
P.E. Afr.
157,
t. 40
fig. A.
1909. ≡ Acacia luederitzii Engl. var. retinens (Sim) J.H. Ross & Brenan, Kew
Bull. 21: 72. 1967 - Type: Mozambique,
‘Umbeluzi and Lebombo’, Sim 6391 [not
found by Ross, (1979) presumed lost].
Vachellia macrothyrsa (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia macrothyrsa
Harms, Bot. Jahrb. Syst. 28: 396. 1900 - Type:
Tanzania, Iringa, Goetze 653 (B†, holotype;?BM, E, K, isotypes).
Vachellia malacocephala (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia malacocephala
Harms, Bot. Jahrb. Syst. 51: 364. 1914 - Type:
Tanzania, Shinyanga Distr., between Samuye
and Kizumbi, Holtz 1548 (B†, holotype; BM,
drawing, K, fragment and drawing, isotypes).
Vachellia mbuluënsis (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia mbuluënsis
Brenan, Kew Bull. 12: 79. 1957 - Type: Tanzania, Mbulu Distr., Ufana, between the rift
wall and Dongobesh, B.D. Burtt 4936 (K, holotype; BM, isotype).
Vachellia montana (P.P.Swartz) Kyal. &
Boatwr.,
comb.
nov. ≡ Acacia
montana
P.P.Swartz in Coates-Palgrave, Trees of southern Africa: 19. 2002, nom. illegit., non Benth.
1842. ≡ Acacia theronii P.P.Swartz, Bothalia
33: 164. 2003 – Type: South Africa, KwaZuluNatal, Hlabisa District, Swartz 178 (PRE,
holotype; PRU, isotype).
Vachellia natalitia (E.Mey.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia natalitia E.Mey., Comm.
1: 167. 1836 - Types: South Africa, KwaZuluNatal, Durban and Umgeni, Drège s.n. (K, P);
South Africa, Eastern Cape, between Umgazana and Umzimvubu, Drège s.n. (P).
Vachellia nebrownii (Burtt Davy) Seigler &
Ebinger, Phytologia. 92(1): 95. 2010. ≡ Acacia
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RECLASSIFICATION OF AFRICAN ACACIA SPECIES
nebrownii Burtt Davy, Kew Bull. 1921: 50.
1921. pro parte excl. specim. Burtt Davy 3045
et 5230. - Syntypes: Botswana, Kwebe Hills,
Mrs E. J. Lugard 14 (K) and 16 (K).
48 Vachellia negrii (Pichi-Sermolli) Kyal. &
Boatwr., comb. nov. ≡ Acacia negrii PichiSermolli, Miss. Stud. Lago Tana, Ric. Bot. 1:
55, tt.9, 10. 1951 - Syntypes: Ethiopia, Gande
Cabanna, Negri 335 (FI); Addis Ababa, Senni
383 (FI); Senni 1589 (FI, K).
49 Vachellia nilotica (L.) P.J.H.Hurter & Mabb.,
Plant Book: 1021. 2008. ≡ Mimosa nilotica L.,
Sp. Pl. 1: 521. 1753. ≡ Acacia nilotica (L.) Willd.
ex Del., Fl. Aegypt. Ill. 79. 1813 - Type: Egypt,
Herb. Linnaeus 1228.28 (LINN, lectotype).
Seven subspecies are recognized:
49.a. subsp. nilotica
49.b. subsp. indica (Benth.) Kyal. & Boatwr,
comb. nov. ≡ Acacia arabica var. indica
Benth., London J. Bot. 1: 500.
1842. ≡ Acacia nilotica subsp. indica
(Benth.) Brenan, Kew Bull. 12: 84. 1957
- Types: India, ‘East India’, Roxburgh
(K); Oungein, collector unknown in Herb.
Bentham (K).
49.c. subsp. tomentosa (Benth.) Kyal. &
Boatwr., comb. nov. ≡ Acacia arabica
var. tomentosa Benth., London J. Bot. 1:
500. 1842., Acacia nilotica subsp. tomentosa (Benth.) Brenan, Kew Bull. 12: 84.
1957 - Type: Senegambia, Heudelot s.n.
(K, lectotype).
49.d. subsp. adstringens (Schumach. &
Thonn.) Kyal. & Boatwr., comb.
nov. ≡ Mimosa adstringens Schumach. &
Thonn.,
Beskr.
Guin.
Pl. 327.
1827. ≡ Acacia nilotica subsp. adstringens (Schumach. & Thonn.) Roberty,
Candollea 11: 150. 1948 - Type: ‘Guinea’,
without locality, Thonning 239 (C, holotype).
49.e. subsp. subalata (Vatke) Kyal. &
Boatwr., comb. nov. ≡ Acacia subalata
Vatke, Öesterr. Bot. Z. 30: 276.
1880. ≡ Acacia nilotica subsp. subalata
(Vatke) Brenan, Kew Bull. 12: 85. 1957 Type: Kenya, Teita Distr., Ndi, Hildebrandt 2589 (B†?, holotype).
49.f. subsp. leiocarpa (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia nilotica
subsp. leiocarpa Brenan, Kew Bull. 12:
84. 1957 - Type: Kenya, Lamu Distr.,
Patta Island, Dale 3832 in C.M. 13988
(K, holotype; EA, isotype).
49.g. subsp. kraussiana (Benth.) Kyal. &
Boatwr., comb. nov. ≡ Acacia arabica
50
51
52
53
54
55
515
var. kraussiana Benth., London J. Bot. 1:
500. 1842. ≡ Acacia nilotica subsp.
kraussiana (Benth.) Brenan, Kew Bull.
12: 84. 1957 - Type: South Africa,
KwaZulu-Natal, Durban, Krauss 69 (K,
holotype; FI, TCD, isotypes).
Vachellia nubica (Benth.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia nubica Benth., London J.
Bot. 1: 498. 1842. – Type: Sudan, Kordofan,
Kotschy 407 (K, holotype; FI, OXF, P, Z, isotypes).
Vachellia oeforta (Forssk) Kyal. & Boatwr.,
comb. nov. ≡ Mimosa oeforta Forssk, Fl.
Aegypt.-Arab.: 177. 1775. ≡ Acacia oeforta
(Forssk) Schweinf., Bull. Herb. Boissier 4,
app. 2: 213. 1896 - Types: Yemen, Dahi, Forsskal s.n. (missing from C, comment based on
Ross, 1979).
Two varieties are recognized:
51.a. var. oeforta
51.b. var. brevifolia (Boulos) Kyal. &
Boatwr., comb. nov. ≡ Acacia oerfota
(Forssk.) Schweinf. var. brevifolia
Boulos, Kew Bull. 50: 334. 1995 - Type:
Yemen, J.R.I. Wood 3089 (K, holotype;
BM, isotype).
= Acacia sarcophylla Chiov., Fl. Somala
1: 161. 1929 - Type: Somalia, Migiurtini,
near Hordio, Puccioni & Stefanini 630
(FI, holotype; BM, isotype).
Vachellia ormocarpoides (P.J.H.Hurter)
Kyal. & Boatwr., comb. nov. ≡ Acacia ormocarpoides P.J.H.Hurter, Bothalia 35(2): 167. 2005 Type: Limpopo, Zwemkloof 283-KT, Hurter 1983
(PRE, holotype; NBG, PRU, isotypes).
Vachellia origena (Hunde) Kyal. & Boatwr.,
comb. nov. ≡ Acacia origena Hunde, Nordic J.
Bot. 2(4): 337. 1982 - Type: Ethiopia, Eritrea
West, Ad Rassi, Pappi 4946 (FT, holotype; K,
isotype).
Vachellia paolii (Chiov.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia paolii Chiov., Ann. Bot.
(Rome) 13: 395. 1915 - Types: Ethiopia,
Ogaden, between Bardera and Mansur, Paoli
578 (FI, K); Heima, Paoli 611 (FI, BM, drawing
K, photograph).
Two subspecies are recognized:
54.a. subsp. paolii
54.b. subsp. paucijuga (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia paolii
Chiov. subsp. paucijuga Brenan, Kew
Bull. 17: 165. 1963 - Type: Kenya,
Northern Frontier Prov., Mt Akoret,
Pratt MS 720 (K, holotype).
Vachellia permixta (Burtt Davy) Kyal. &
Boatwr., comb. nov. ≡ Acacia permixta Burtt
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516
56
57
58
59
60
61
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B. KYALANGALILWA ET AL.
Davy, Kew Bull. 1922: 330. 1922. prop parte
excl. var. glabra. - Type: South Africa, Transvaal, Potgietersrust, Burtt Davy 5230 (PRE,
holotype; K, isotype).
Vachellia pilispina (Pichi-Sermolli) Kyal. &
Boatwr., comb. nov. ≡ Acacia pilispina PichiSermolli, Miss. Stud. Lago Tana, Ric. Bot. Bot.
1: 205, t. 43. 1951 - Type: Ethiopia, Atghebà
Ghiorghis, Pichi-Sermolli 2696 (FI, holotype).
Vachellia prasinata (Hunde) Kyal. &
Boatwr., comb. nov. ≡ Acacia prasinata
Hunde, Nordic J. Bot. 2(4): 341. 1982 - Type:
Ethiopia, Shewa region, Awash Nat. Park,
Thulin, Hunde & Tadesse 3865 (UPS, holotype;
ETH, K, isotypes).
Vachellia pseudofistula (Harms) Kyal. &
Boatwr., comb. nov. ≡ Acacia pseudofistula
Harms, Bot. Jahrb. Syst. 51: 363. 1914 - Syntypes: Tanzania, Tabora Distr., Goweko, Holtz
2801 (B†, BM, K); Dodoma Distr., Kilimatinde,
Holtz 1358 (B†).
Vachellia qandalensis (Thulin) Kyal. &
Boatwr., comb. nov. ≡ Acacia qandalensis
Thulin, Nordic J. Bot. 18: 513. 1998 - Type:
Somalia, Bari Region, Cal Miskaat in Bahaya
area, c. 20 km south-west of Qandala, Thulin,
Abdi Dahir & Ahmed Osman 9419 (UPS, holotype; K, isotype).
Vachellia quintanilhae (Torre) Kyal. &
Boatwr., comb. nov. ≡ Acacia quintanilhae
Torre, Bol. Soc. Brot., Sér. 2, 36: 1, t. 1. 1962 Type: Angola, Mocamedes Distr., km 10 do
Apeadeiro do C.F. de Dois Irmãos, Torre 8274
(LISC, holotype; BM, LUA, LUAI, isotypes).
Vachellia reficiens (Wawra) Kyal. & Boatwr.,
comb. nov. ≡ Acacia reficiens Wawra, Sitzungsber. Akad. Wiss. Wien, Math.-Naturwiss.
Kl., Abt. 1 38: 555. 1859 - Type: Angola,
between Benguela and Catumbela, Wawra 248
(W, holotype; K, fragment, isotype).
Two subspecies are recognized:
61.a. subsp. reficiens
61.b. subsp. misera (Vatke) Kyal. & Boatwr.,
comb. nov. ≡ Acacia misera Vatke,
Öesterr. Bot. Z. 30: 275. 1880. ≡ Acacia
reficiens Wawra subsp. misera (Vatke)
Brenan, Kew Bull. 12: 90. 1957 - Type:
Somalia, Meid, Hildebrandt 1394 (B†,
holotype; BM, K, isotypes).
Vachellia rehmanniana (Schinz) Kyal. &
Boatwr., comb. nov. ≡ Acacia rehmanniana
Schinz, Bull. Herb. Boissier 6: 525. 1898 Type: South Africa, Transvaal, Streydpoort,
Makapansberge, Rehmann 551 (Z, holotype).
Vachellia robbertsei (P.P.Swartz) Kyal. &
Boatwr., comb. nov. ≡ Acacia robbertsei
64
65
66
67
P.P.Swartz, Coates-Palgrave’s Trees of southern Africa: 19. 2002, as ‘robbertsii’ - Type:
South Africa, Mpumalanga, Lydenburg District, 5 miles north-west of PO Morone, Codd
10483 (PRE, holotype).
Vachellia robusta (Burch.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia robusta Burch., Trav. 2:
442. 1824 - Type: South Africa, Northern Cape
Province, Kuruman Distr., Takoon, Burchell
2265 (K, holotype).
Three subspecies are recognized:
64.a. subsp. robusta
64.b. subsp. clavigera (E.Mey.) Kyal. &
Boatwr., comb. nov. ≡ Acacia clavigera
E.Mey., Comm. 1: 168. 1836. ≡ Acacia
robusta Burch. subsp. clavigera (E.Mey.)
Brenan, Fl.Zambesiaca 3, 1: 104. 1970 Type: South Africa, KwaZulu-Natal, near
Durban, Drège (K, isotype; P, fragment).
64.c. subsp. usambarensis (Taub.) Kyal. &
Boatwr., comb. nov. ≡ Acacia usambarensis Taub., Pflanzenw. Ost-Afrikas C:
195, t. 20H. 1895. ≡ Acacia robusta
Burch. subsp. usambarensis (Taub.)
Brenan, Fl. Zambesiaca 3, 1: 104. 1970 Types: Tanzania, Lushot Distr., Simbili,
Holst 2362 (B†, K); Mashewa, Holst 8820
(B†, K).
Vachellia sekhukhuniensis (P.J.H.Hurter)
Kyal. & Boatwr., comb. nov. ≡ Acacia sekhukhuniensis P.J.H.Hurter, Bothalia 34(2): 109.
2004 - Type: South Africa, Limpopo, Sekhukhuneland, Farm Schlickmannskloof 258KT,
Mukoma & Hurter 17 (PRE, holotype; NBG,
PRU, isotypes).
Vachellia seyal (Del.) P.J.H.Hurter, Mabberley’s plant book: 1021. 2008. ≡ Acacia seyal
Del., Fl. Egypt. Expl. Planches 286. 1813 Syntypes: Egypt, between Nile and Red Sea,
near Syene, Delile s.n. (?MPU); Medynet-Abou,
Delile s.n. (?MPU).
Two varieties are recognized:
66.a. var. seyal
66.b. var. fistula (Schweinf.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia fistula Schweinf.,
Linnaea
35:
344:
tt
11-14.
1867-68. ≡ Acacia seyal var. fistula (Schweinf.) Oliv., Fl. Trop. Afr. 2: 351. 1871 Syntypes: Sudan, Gedaref region, and
Mt Gule in the Sennar Prov., Schweinfurth (B†).
Vachellia sieberiana (DC.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia sieberiana DC., Prodr. 2:
463. 1825. - Type: Senegal, Sieber 43 (G, holotype; K, MEL, isotypes).
Three varieties are recognized:
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RECLASSIFICATION OF AFRICAN ACACIA SPECIES
68
69
70
71
72
73
67.a. var. sieberiana
67.b. var. villosa (A.Chev.) Kyal. & Boatwr.,
comb. nov. ≡ Acacia sieberiana DC. var.
villosa A.Chev., Bull. Soc. Bot. France
74: 959. 1927 - Type: Haute Volta
(Burkina Faso), Ouré, Chevalier 700 (P,
holotype; K, isotype).
67.c. var. woodii (Burtt Davy) Kyal. &
Boatwr., comb. nov. ≡ Acacia woodii
Burtt Davy, Kew Bull. 1922: 332.
1922. ≡ Acacia sieberiana DC. var.
woodii (Burtt Davy) Keay & Brenan,
Kew Bull. 5: 364. 1951 - Type: South
Africa, KwaZulu-Natal, Estcourt Distr.,
between Estcourt and Colenso, Wood
3528 (K, holotype; MEL, NH, isotypes).
Vachellia stuhlmannii (Taub.) Kyal. &
Boatwr., comb. nov. ≡ Acacia stuhlmannii
Taub., Pflanzenw. Ost-Afrikas C: 194, t.21E, F.
1895 - Syntypes: Tanzania, Dar es Salaam,
Stuhlmann 6755 (B†, EA); Pangani, Stuhlmann 282 (B†); Tanga, Volkens 189 (B†);
Amboni, Holst 2202 (B†, K, Z); Tanzania/
Kenya, Lake Jipe, Volkens 2383 (B†).
Vachellia swazica (Burtt Davy) Kyal. &
Boatwr., comb. nov. ≡ Acacia swazica Burtt
Davy, Bull. Misc. Inform. Kew 1922: 332. 1922
- Type: Swaziland, near Manzini, Burtt Davy
3045 (PRE, holotype; K, fragment, BM,
isotype).
Vachellia tenuispina (Verdoorn) Kyal. &
Boatwr., comb. nov. ≡ Acacia tenuispina Verdoorn, Bothalia 6: 156. 1951 - Type: South
Africa, Transvaal, Waterberg Distr., Hoogbult
Farm, Naboomspruit, Galpin 475 M (PRE,
holotype; K, isotype).
Vachellia tephrophylla (Thulin) Kyal. &
Boatwr., comb. nov. ≡ Acacia tephrophylla
Thulin, Nordic J. Bot. 18: 515. 1998 - Type:
Somalia, Bari Region, Cal Miskaat, north of
Dasan, Thulin, Abdi Dahir & Ahmed Osman
9482 (UPS, holotype; K, isotype).
Vachellia torrei (Brenan) Kyal. & Boatwr.,
comb. nov. ≡ Acacia torrei Brenan, Kew Bull.
21: 480. 1968 - Type: Mozambique, Manica e
Sofala, between Inhaminga and Rio Urema,
Torre 4068 (LISC, holotype; BM, K, PRE, isotypes).
Vachellia tortilis (Forssk.) Galasso & Banfi,
Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat.
Milano 149(1): 150. Jan. 2008. ≡ Mimosa tortilis Forssk., Fl. Aegypt-Arab 124: 176.
1775. ≡ Acacia
tortilis
(Forssk.)
Hayne,
Arzneyk. Gebr. Gewächse 10: t. 31. 1827 Type: Arabia, ‘Mons Soudân prope Hás’, Forsskål (C, holotype; K, isotype).
517
Note: Vachellia tortilis (Forssk.) P.J.H.Hurter
& Mabb., Pl.-Book 1021. 2008 [1 PubMed May
2008] is an isonym of Vachellia tortilis
(Forssk.) Galasso & Banfi and has no nomenclatural status according to the Code (McNeill
et al., 2006), art. 6.3 note 2 ‘when the same
name, based on the same type, has been published independently at different times by different authors, then only the earliest of these
“isonyms” has nomenclatural status.’
Four subspecies and three varieties are
recognized:
73.a. subsp. tortilis
73.b. subsp. raddiana (Savi) Kyal. &
Boatwr., comb. nov. ≡ Acacia raddiana
Savi, Alc. Acazie Egiz. 1. 1830. ≡ Acacia
tortilis subsp. raddiana (Savi) Brenan,
Kew Bull. 12: 87. 1957 - Type: Egypt,
Raddi (K, isotype).
73.b.1. var. raddiana
73.b.2. var. pubescens (A.Chev.) Kyal.
& Boatwr., comb. nov. ≡ Acacia
tortilis (Forssk.) Hayne subsp.
raddiana (Savi) Brenan var.
pubescens A. Chev., Bull. Soc.
Bot. France 74: 960. 1927 - Syntypes: Mali, Tombouctou (Timbuktu), Chevalier 1186 (K, P);
1187 (P); 1188 (P).
73.c. subsp. spirocarpa (Hochst. ex. A.Rich.)
Kyal. & Boatwr., comb. nov. ≡ Acacia
spirocarpa Hochst. ex. A.Rich., Tent. Fl.
Abyss. 1: 239. 1847. ≡ Acacia tortilis
subsp. spirocarpa (Hochst. ex. A.Rich.)
Brenan, Kew Bull. 12: 88. 1957 - Syntypes: Ethiopia, near Djeladjeranne,
Schimper 502 (BM, FI, K, M, P, Z);
Schimper 612 (BM, FI, K, M, P); Schimper 658 (BM, K, M, OXF, P, Z).
73.c.1. var. spirocarpa
73.c.2. var. crinita (Chiov.) Kyal. &
Boatwr., comb. nov. ≡ Acacia
tortilis (Forssk.) Hayne subsp.
spirocarpa (Hochst. ex. A.Rich.)
Brenan var. crinita Chiov., Res.
Sci. Miss. Stefanini-Paoli 1: 71.
1916 - Type: Somalia between
Doriànale and Oneiátta, Paoli
907 (FI, holotype; K, isotype).
73.d. subsp. heteracantha (Burch.) Kyal. &
Boatwr., comb. nov. ≡ Acacia heteracantha Burch., Trav. 1: 389. 1822. Acacia
tortilis (Forssk.) Hayne subsp. heteracantha (Burch.) Brenan, Kew Bull. 12:
88. 1957 - Type: South Africa, Northern
Cape Province, Hay Distr., Spuigslang-
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B. KYALANGALILWA ET AL.
fontein, between Griquatown and the
Orange River, Burchell 1710 (K, holotype).
Vachellia turnbulliana (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia turnbulliana
Brenan, Kew Bull. 12: 370. 1958 - Type:
Kenya, Northern Frontier Prov., 23 km northeast of Wajir, Gillett 13364 (K, holo; EA,
isotype).
Vachellia walwalensis (Gilliland) Kyal. &
Boatwr., comb. nov. ≡ Acacia walwalensis Gilliland, Kew Bull. 6: 140, t. 5. 1951 - Type:
Ethiopia, Ogaden, between Wardere and
Walwal, Glover & Gilliland 392 (K, holotype;
BM, FHO, PRE, isotypes).
Vachellia xanthophloea (Benth.) P.J.H.Hurter, Mabberley’s plant book: 1021.
2008. ≡ Acacia xanthophloea Benth., Trans.
Linn. Soc. London, Bot. 30: 511. 1875 - Syntypes: Malawi, E. end of Lake Shirwa, Meller
s.n. (K); Mozambique, Sena, Kirk s.n. (K).
Vachellia zanzibarica (S.Moore) Kyal. &
Boatwr., comb. nov. ≡ Pithecolobium? zanzibaricum
S.Moore,
J.
Bot.
15:
292.
1877. ≡ Acacia zanzibarica (S.Moore) Taub.,
Pflanzenw. Ost Afrikas C: 195. 1895 - Type:
Kenya, Mombasa, Hildebrandt 1939 (K, holotype; BM, isotype).
Two varieties are recognized:
77.a. var. zanzibarica
77.b. var. microphylla (Brenan) Kyal. &
Boatwr., comb. nov. ≡ Acacia zanzibarica (S.Moore) Taub. var. microphylla
Brenan, Kew Bull. 12: 75. 1957 - Type:
Kenya, Northern Frontier Prov., Turbi,
Gillet 13803 (K, holotype; EA, isotype).
Species insufficiently known:
1 Acacia callicoma Meisn., London J. Bot. 2: 104.
1843 - Type: unknown.
2 Acacia balfouri G.M.Woodrow, J. Bombay Nat.
Hist. Soc. 11: 420–430. 1898. – Type: not seen. This
species is listed in Lock (1989), but not in Ross
(1979); it is provisionally accepted in Roskov et al.
(2005).
3 Acacia leucophaea Willd. is listed in Lock (1989)
but not in Ross (1979); it is also listed in Roskov
et al. (2005). We were unable to find any literature
on the original publication of this name.
Unvalidated names:
1 Acacia firozei Najma Dh. in Dharani N. 2006. Field
guide to acacias of the East Africa.
2 Acacia kenyensis Najma Dh. in Dharani N. 2006.
Field guide to acacias of the East Africa.
3 Acacia tirion Najma Dh. in Dharani N. 2006. Field
guide to acacias of the East Africa.
Excluded names:
1 Acacia purpurea Bolle, Reise Mossamb. Bot. 1: 6.
1861 - Types: Mozambique, Chupanga, Sena, Tete,
etc., Peters (B†). Acacia mauroceana DC., Cat.
Hort. Monsp. 74: 1813 - Type: grown from seed
collected by Broussonet in Morocco. (G,
holotype). = Painteria leptophylla (DC.) Britton
& Rose (D. Seigler & J. Ebinger, pers. comm.).
2 Acacia redacta J.H.Ross, Bothalia 11: 231. 1974 –
Type: South Africa, Cape Province, Namaqualand,
22 km N. of Stinkfontein on way to Jenkinskop,
Werger 1518 (PRE, holotype; K, isotype). = Calliandra redacta (J.H.Ross) Thulin & Asfaw Nordic J.
Bot. 1(1): 29 (1981).
ACKNOWLEDGEMENTS
The authors thank the University of Johannesburg,
the National Research Foundation of South Africa
and the Royal Society of London for financial support.
The DNA Bank of the Jodrell Laboratory Molecular
Department at the Royal Botanic Gardens, Kew (UK)
is acknowledged for supplying materials used in this
study. Three anonymous reviewers provided useful
comments on an earlier version of the manuscript.
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APPENDIX
LIST OF TAXA WITH VOUCHER INFORMATION AND
GenBank ACCESSION NUMBERS FOR EACH DNA
REGION
Species, voucher specimen, herbarium, trnK/matK
GenBank accession, psbA-trnH GenBank accession,
trnL-trnF GenBank accession. *Sequences not
included in Bouchenak-Khelladi et al. 2010.
Acacia s.l. Miller: *A. adenocalyx Brenan & Exell,
OM2439, JRAU, JQ230127, —, JQ230284/JQ230285;
A. adoxa Pedley, Archer 50994, MEL, AF523076,
AF195716,
AF195703/AF195684;
A. ampliceps
Maslin, J. Miller 583, CANB, AF523074, AF525003,
AF522983; *A. amythethophylla Steud ex. A.Rich.,
MSB171878, K, JQ230136, JQ230187, JQ230294;
A. arenaria Schinz, OM1048, JRAU, GQ872212,
GQ872302, GQ872257; A. ataxacantha DC., RL1326,
JRAU, GQ872213, —, —; *A. auriculiformis Benth.,
MSB57484, K, JQ230110, JQ230158, JQ230305/
JQ230306; *A. boormanii Maiden, Chase 17932, K,
JQ230111, —, —; A. borlea Burtt Davy, RL1308,
JRAU, GQ872214, GQ872303, GQ872258; A. brevispica
Harms,
RL1333,
JRAU,
GQ872215,
GQ872304, GQ872259; A. burkeii Benth., RL1283,
JRAU, GQ872216, GQ872305, GQ872260; *A. carani-
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
520
B. KYALANGALILWA ET AL.
ana Chiov., M. Thulin et al. 10431, K, —, —,
JQ230256/JQ230257; A. chariessa Milne-Redh., M.
Steyn 2, JRAU, GQ872218, GQ872307, GQ872262;
*A. cochliacantha Willd., J. Miller 246, CANB, —, —,
AF522968; A. colei Maslin & L.A.J.Thomson, J. Miller
871, CANB, AF274215, AF525007, AF522987;
*A. condyloclada Chiov., C.F. Hemming76 and R.M.
Watson 30, K, —, —, JQ230258/JQ230259; *A. craspedocarpa F.Muell., Chase 17931, K, —, JQ230162,
JQ230250/JQ230251; A. davyi N.E.Br, RL1319,
JRAU, GQ872219, GQ872308, GQ872263; A. drummondii Lindl., 2034627, MEL, AF523106, AF195714,
AF195704/AF195685;
*A. dudgeonii
Craib
ex
Holland, MSB125505, K, JQ265930, JQ230180,
JQ230307/JQ230308; *A. dyeri P.P.Sw. ex Coates
Palgr., RL1309, JRAU, JQ230137, JQ230188,
JQ230221/JQ230222;
*A. ehrenbergiana
Hayne,
MSB99862, K, JQ230138, JQ230189, JQ230295/
JQ230296; A. elata A.Cunn ex Benth., D. Murphy
234, MEL, AF274149, AF195709, AF195702/
AF195683; A. eriocarpa Brenan, M. Steyn 1, JRAU,
GQ872220, GQ872309, GQ872264; A. erubescens
Welw. ex Oliver, RL1318, JRAU, GQ872222,
GQ872311, GQ872266; A. exuvialis I.Verdn., RL1284,
JRAU, GQ872223, GQ872312, GQ872267; *A. fleckii
Schinz., MSB83043, MSB, JQ230130, JQ230181,
JQ230309/JQ230310; *A. furcatispina Burkart, J.
Miller
1214,
CANB,
EU812018,
EU811960,
EU439992; A. galpinii Burtt Davy, RL1304, JRAU,
GQ872224, GQ872313, GQ872268; A. giraffae Willd.,
RL1300, JRAU, GQ872226, GQ872315, GQ872270;
*A. goetzei subsp. goetzei Harms, RL1322, JRAU,
JQ230131, JQ230182, JQ230229/JQ230230; *A. goetzii subsp. macrophylla Brenan, RL1320, JRAU, —,
JQ230183, JQ230227/JQ230228; A. grandicornuta
Gerstner, RL1286, JRAU, GQ872227, GQ872316,
GQ872271; *A. gummifera Willd., MSB84062, K, —,
—, JQ230311; *A. haematoxylon Willd., OM1069,
JRAU, JQ230141, JQ230192, JQ230275/JQ230276;
*A. hamulosa Benth., J.C.W. Fagg & B.T. Styles 39,
K, —, —, JQ230262/JQ230263; A. hebeclada DC.
subsp. chobiensis (O.B.Miller) Schreiber, OM1034,
JRAU, GQ872228, GQ872317, GQ872272; *A. hebeclada subsp. hebeclada DC., MSB104986, K,
JQ230143, JQ230193, JQ230312/JQ230313; A. hebeclada subsp. tristis A.Schreib., RL1301, JRAU,
GQ872229, GQ872318, GQ872273; *A. hereroensis
Engl., RL1332, JRAU, JQ230132, JQ230184,
JQ265939/JQ265940;
*A. hockii
De
Willd.,
MSB369514, K, JQ230144, JQ230194, JQ230301/
JQ230302; A. kirkii Oliv., RL1307, JRAU, GQ872231,
GQ872319, GQ872275; A. kosiensis P.P.Sw. ex Coates
Palgr., RL1305, JRAU, GQ872232, GQ872320,
GQ872276; A. kraussiana Meisn. ex Benth., RL1287,
JRAU, GQ872233, GQ872321, GQ872277; A. luederetzii Engl. var. retinens (Sim) J.H.Ross & Brenan,
RL1285, JRAU, GQ872234, GQ872322, GQ872278;
A. lycopodiifolia Hook., M.F. Duretto 1063, MEL,
DQ371879, AF195715, AF195705/AF195686; A. macrostachya Rchb. ex G.Don, CANB632225, CANB,
DQ371881, —, DQ371856; *A. mammifera Schltdl.,
Chase 8247, K, JQ230112, JQ230163, —; A. mearnsii
De Wild., D. Murphy 200, MEL, AF523110,
AF195707, AF195694/AF195675; A. melanoxylon
R.Br., J. Miller 748, CANB, AF274166, AF195712,
AF195699/AF195680; A. mellifera (Vahl) Benth.
subsp. mellifera, OM1030, JRAU, GQ872235,
GQ872323, GQ872279; A. montis-usti Merxm. & Schreiber, OM1065, JRAU, —, GQ872324, GQ872280;
*A. natalitia E.Mey., RL1330, JRAU, —, JQ278603,
JQ230233/JQ230234; A nebrownii Burtt Davy,
OM1050, JRAU, GQ872236, GQ872325, GQ872281;
*A. ogadensis Chiov., S. Bidgood et al. 4991, K, —, —,
JQ230264; JQ230265; A. ormocarpoides P.J.H.Hurter,
RL1293, JRAU, GQ872239, GQ872327, GQ872284;
*A. pataczekii D.I.Morris, Chase 16092, K, JQ230113,
JQ230164, JQ230248/JQ230249; A. permixta Burtt
Davy, Hurter J.2, JRAU, GQ872240, GQ872328,
GQ872285; A. platycarpa F.Muell., D. Murphy 327,
MEL, AF274223, AF525005, AF522985; A. polyacantha Willd., RL1323, JRAU, GQ872241, GQ872329,
GQ872286; A. pulchella R.Br., D. Murphy 268, MEL,
AF523100, AF195724, AF195692/AF195673; A. reficiens Wawra, RL1297, JRAU, GQ872242, GQ872330,
GQ872287; A. rhemaniana Schinz, RL1288, JRAU,
GQ872243, —, —; A. robbertsei P.P.Sw. ex Coates
Palgr., RL1289, JRAU, GQ872244, GQ872331,
GQ872288; A. robusta Burch. subsp. clavigera
(E.Mey.) Brenan, RL1316, JRAU, GQ872245,
GQ872332, GQ872289; *A. robusta Burch. subsp.
usambarensis, OM2458, JRAU, JQ230146, —,
JQ230286; JQ230287; *A. robysiana Merxm. & Schreiber, OM1066, JRAU, JQ230133, JQ230186,
JQ230273/JQ230274; A. schweinfurthii Brenan &
Exell, RL1299, JRAU, GQ872246, GQ872333,
GQ872290;
A. sekhukhuniensis
P.J.H.Hurter,
RL1296, JRAU, GQ872247, GQ872334, GQ872291;
A. senegal (L.) Willd. var. leiorachis Brenan, RL1324,
JRAU, GQ872248, GQ872335, GQ872292; A. senegal
(L.) Willd. var. rostrata Brenan, RL1331, JRAU,
GQ872249, GQ872336, GQ872293; A. sieberiana DC.
subsp. sieberiana, OM1029, JRAU, —, GQ872337,
GQ872294; A. sieberiana DC. var. woodii (Burtt
Davy) Keay & Brenan, RM02, JRAU, GQ872250,
GQ872338, GQ872295; A. somalensis Vatke, M.
Thulin 10823, K, —, —, JQ230266/JQ230267;
A. spinescens Benth., D. Murphy 246, MEL,
AF523082, AF195725, AF195706; A stullmannii
Taub., RL1294, JRAU, GQ872251, GQ872339,
GQ872296; A. swazica Burtt Davy, RL1327, JRAU,
GQ872252, GQ872340, GQ872297; *A. theronii
P.P.Sw., RL1313, JRAU, —, JQ230196, JQ230223/
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
JQ230224; *A. torrei Brenan, OM2429, JRAU,
JQ230147, —, JQ230282/JQ230283; A. translucens
Cunn. ex Hook., D. Murphy 302, MEL, AF523087,
AF525004, AF522984; A. tumida F.Muell ex Benth.,
J. Miller 872, CANB, AF523111, AF525006,
AF522986; *A. vetista Ker Gawl., Chase 15949, K,
JQ230114, JQ230168, JQ230246/JQ230247; *A
zizyphispina Chiov., A.S. Hassan 53, K, —, —,
JQ230269/JQ230270; A. welwitschii Oliv. subsp. delagoensis (Harms) J.H.Ross & Brenan, RL1325, JRAU,
GQ872254, GQ872342, GQ872299; *Vachellia nilotica
(L.) P. J. Hurter & Mabb.subsp. tomentosa (Benth.)
Kyal. & Boatwr., MSB132963, K, JQ230151,
JQ230200, JQ230316; *V. nilotica (L.) P. J. Hurter &
Mabb. subsp. indica (Benth.) Kyal. & Boatwr.,
MSB61070, K, JQ230150, JQ230199, JQ230314/
JQ230315; V. tortilis (Forssk.) Galasso & Banfi,
RL1290, JRAU, GQ872253, GQ872341, GQ872298;
*V. tortilis (Forssk.) Galasso & Banfi subsp. tortilis,
MSB151463, K, JQ230154, JQ230203, JQ230320/
JQ230321; *V. tortilis (Forssk.) Galasso & Banfi
subsp. heteracantha (Burch.) Kyal. & Boatwr.,
MSB82839, K, JQ230153, JQ230201, JQ230317/
JQ230318; *V. tortilis (Forssk.) Galasso & Banfi
subsp. spirocarpa (Hochst. ex. A.Rich.) Kyal. &
Boatwr., MSB26381, K, JQ230152, JQ230202,
JQ230303/JQ230304; *V. tortilis (Forssk.) Galasso &
Banfi subsp. raddiana (Savi). Kyal. & Boatwr.,
MSB69065, K, —, —, JQ230319; Acaciella Britton
& Rose: A. angustissima (Mill.) Britton & Rose var.
angustissima, DS15993, DS, DQ371887, AF195715,
DQ371872; A. boliviana Rusby (= A. angustissima), D.
Murphy 248, MEL, AF274144, AF525001, AF522981;
*A. chamelensis (L.Rico) L.Rico, L. Rico 8236, K, —,
JQ230160, —; A. glauca (L.) L.Rico, 96-12580
(DLEG), DQ371880, —, DQ371857; *A. rosei (Standl.)
Britton & Rose, Lott 9535, K, JQ265929, JQ230165,
JQ230241/JQ230242;
*A. tequilana
(S.Watson)
Britton & Rose, Rico 1206, K, —, JQ230167,
JQ230237/JQ230238. Adenanthera L.: A. pavonina
L., Major Howell Seeds, BH, AF521808, —, —. Alantsilodendron Villiers: A. pilosum J.-F.Villiers, M.
Luckow 4301 (BH), AY125844, —, AY125844. Albizia
Durraz.:
*A. amara
Boiv.,
OM2136,
JRAU,
JQ230117, JQ230170, JQ230279; *A. athlemintica
Brongn, OM363, JRAU, JQ230118, JQ230171,
JQ230209; *A. brevifolia Schinz, OM826, JRAU,
JQ230119, JQ230172, JQ230214/JQ230215; *A. forbesii Benth., OM331, JRAU, JQ230120, JQ230173, —;
*A. glaberrima (Schum. & Thonn.) Benth., OM2605,
JRAU, JQ230121, JQ230174, JQ230288/JQ230289;
*A. harveyi Fourn., OM1402, JRAU, JQ230122,
JQ230175, —; *A. petersiana Oliver, OM745, JRAU,
JQ230123, JQ230176, JQ230212/JQ230213; A. kalkora Prain, J. Miller 877, CANB, AF523083,
AF524965, AF522945; A. sinaloensis Britton & Rose,
521
J. Miller 878, CANB, AF274121, AF524966,
AF522946; *A. suluensis Gerstner, SA 156, JRAU,
JQ230124, —, JQ230235; *A. tanganyicensis Baker f.,
OM1972, JRAU, —, —, JQ230278; *A. versicolor
Welw. ex Oliver, RL1214, JRAU, JQ265933,
JQ230177, JQ230218; *A. zimmermannii Harms,
OM2363, JRAU, JQ230125, JQ230178, JQ230280;
JQ230281; *A. zygia J.F.Macbr., OM1820, JRAU,
JQ230126, JQ230179, —. Anadenanthera Speg.:
A. colubrina (Vell.) Brenan, R.T. Pennington 845, E,
AF278481, —, AF278481. Arapatiella Rizzini &
A.Mattos: A. psilophylla (Harms) R.S.Cowan, Carvalho 6095, K, EU361859, —, EU361738. Archidendron F.Muell.: A. hirsutum I.C.Nielsen, Douglas 625,
MEL, EU361860, —, AF365042. Bussea Harms:
B. perrieri R.Vig., Randrianasolo 527, P, EU361896,
—, EU361757. Calliandropsis H.M.Hern. &
P.Guinet: C. nervosus (Britton & Rose) H.M.Hern. &
P.Guinet, Hernandez 2365, BH, AF278520, —,
AF278520. Cathormion Hassk.: C. umbellatum
(Vahl) Kosterm., J. Miller 882, CANB, AF274122,
AF524968, AF522949. Chloroleucon Britton &
Rose ex Record: C. mangense (Jacq.) Britton &
Rose, J. Miller 527, CANB, AF523072, AF524969,
AF522950. Ebenopsis Britton & Rose: E. ebano
(Berland.) Barneby & J.W.Grimes, J. Miller 529,
CANB, AF274123, AF524970, AF522951. Cylicodiscus Harms: C. gabunensis Harms, M.S.M. Sosef
645A, BH, AF521819, —, AY125845. Delonix Raf.:
D. elata Gamble, Herendeen 20-XII-97-1, US,
EU361928, —, AF365106. Desmanthus Willd.: D. bicornutus S.Watson, 615637, CANB, AF523108, —,
AF522939; D. cooleyi (Eaton) Branner & Coville,
Wojciechowski 1018, ASU, AY386916, —, —.
Dichrostachys Wight & Arn.: *D. cinerea (L.) Wight
& Arn., OM256, JRAU, JQ230155, JQ230204,
JQ230207; JQ230208; *D. cinerea subsp. africana
Brenan & Brummitt, RBN359, JRAU, JQ230156,
JQ230205, JQ265941; D. richardiana Baill., Luckow
4261, BH, AF521823, —, —. Dimorphandra Schott:
*D. conjugata Sandwith, Breteler 13800, WAG,
EU361934, —, AF365099. Dinizia Ducke: D. excelsa
Ducke, Sergio de Faria s.n., —, AF521827, —,
AF278479. Elephantorrhiza Benth.: E. elephantina
Skeels, Sergio de Faria s.n., —, AF521828, —,
AF278484. Enterolobium Mart.: E. contortisiliqua
(Vell.) Morong, J. Miller 888, CANB, AF274124,
AF524971, AF522952; E. cyclocarpum (Jacq.) Griseb.,
D. Murphy 355, MEL, AF521831, AF524972,
AF278518. Erythrophleum Afzel. ex R.Br.: E. suaveolens (Guill. & Perr.) Brenan, Herendeen 17-XII97-3, US, EU361949, —, AF365103. Faidherbia
A.Chev.: F. albida (Delile) A.Chev., RM 01, JRAU,
GQ872256, GQ872344, GQ872301. Fillaeopsis
Harms: F discophora Harms, D. Harris 4111, E,
AF521832, —, AF278508. Gagnebina Neck.: G. ba-
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
522
B. KYALANGALILWA ET AL.
koliae Luckow & Du Puy, Lockow 4243, BH,
AY125848, —, AY125848. Havardia Small: H. albicans (Kunth) Britton & Rose, J. Miller 881, CANB,
AF523085, AF524975, AF522956; H. pallens (Benth.)
Britton & Rose, J. Miller 615547, CANB, AF274125,
AF524974, AF522955. Inga Mill.: I. edulis Mart.,
2066677, MEL, AF523078, AF524976, AF522957.
Jacqueshuberia Ducke: J. brevipes Barneby,
Redden 1240, US, EU361984, —, EU361815.
Kanaloa Lorence & K.R.Wood: K. kahoolawensis
Lorence & K.R.Wood, D. Lorence s.n., NTBG,
AF521839, —, AF278489. Lemuropisum H.Perrier:
L. edule H.Perrier, Du Puy M1033, K, EU361991, —,
EU361818. Leucaena Benth.: L. leucocephala
(Lam.) de Wit, J. Miller 615639, CANB, AF523094,
—, AF522942; L. retusa Benth., Boke & Massey 419,
UC, AY386858, —, —. Lysiloma Benth.: L. acapulcense (Kunth) Benth., J. Miller 885, CANB,
AF274126, AF524977, AF522958; L. tergeminum
Benth., J. Miller 532, CANB, AF523089, AF524978,
AF522959. Mariosousa Seigler & Ebinger:
M. acatlensis (Benth.) Seigler & Ebinger, DS16002,
DS, DQ371890, —, DQ371874; M. coulteri (Benth.)
Seigler & Ebinger, DS15953, DS, DQ371893,
AF525008, DQ371868; M. dolichostachya (S.F.Blake)
Seigler & Ebinger, DS16035, DS, DQ371892,
AF525009, DQ371866; M. salazari (Britton & Rose)
Seigler & Ebinger, DS15978, DS, DQ371888, —,
DQ371865; *M. sericea (Martens & Galeotti) Seigler
& Ebinger, Chase 19849, K, JQ230115, JQ230166,
JQ230252/JQ230253; M. usumacintensis (Lundell)
Seigler & Ebinger, DS16025, DS, DQ371889, —,
DQ371863; M. willardiana (Rose) Seigler & Ebinger,
89-0143, DLEG, AY386898, —, DQ371862. Microlobius C.Presl: M. foetidus (Jacq.) M.Sousa & G.Andrade, J. Miller 435, CANB, AF523095, —,
AF522960. Mimosa L.: M. tenuiflora (Willd.) Poir.,
615541, CANB, AF274120, AF524963, AF522943.
Mimozyganthus Burkart: M. carinatus (Griseb.)
Burkart., F. Fortunato 7575, BAB/BH, AY944556, —,
DQ344604. Neptunia Lour.: N. gracilis Benth., J.
Grimes 3168, BH, AF521845, —, AF278494; N. monosperma F.Muell. ex Benth., B. Jackes s.n., BH,
AF274209, —, AF522944; *N. oleracea Lour., RBN162
(JRAU), JQ230157, JQ230206, JQ230216/JQ230217.
Newtonia Baill.: N. buchananii (Baker) G.C.C.Gilbert & Boutique, BNBG 69-6494, BR, AF521847, —,
AF278501; N. hildebrandtii (Vatke) Torre, BNBG
73-2891, BR, AF521848, —, AF278502. Pachyelasma Harms: P. tessmannii Harms, Harris 3972, K,
EU362013, —, AF365105. Pararchidendron I.C.Nielsen: P. pruinosum Koorders, 615549, CANB,
AF274127, AF524980, AF522961. Parapiptadenia
Brenan: P. pterosperma (Benth.) Brenan, E. Tameirao 2458, NY, DQ784651, —, DQ784651; P. rigida
(Benth.) Brenan, A. arambarri s.n., BH, AF278505,
—, AF278505. Paraserianthes I.C.Nielsen: P. lophantha (Willd.) I.C.Nielsen, 615550, CANB,
AF274128, AF524981, AF522962. Parkia R.Br.:
P. biglandulosa Wright & Arn., Banana Tree Nursery,
BH, AF521850, —, AF278498; P. speciosa Hassk.,
Bruneau 931, BH, AF521851, —, AF278499; P. timoriana (DC.) Merr., DM 265, MELU, AF523091,
AF195719, AF195682. Parkinsonia Plum. ex L.:
P. florida S.Watson, Salywon 919, ASU, AY386856, —,
EU361827. Peltophorum (Vogel) Benth.: P. dubium
Taub., No. 90.2705, Wojciechowski 892, ASU,
AY386846, —, EU361828. Pentaclethra Benth.:
P. eetveldeana De Wild. & T.Durand, BNBG 65-6191,
BR, AF521852, —, AY125850; P. macrophylla Benth.,
BNBG 87-1143, BR, AF521853, —, AF278485. Piptadenia Benth.: P. minutiflora Ducke, CM Leme 6, NY,
DQ790624, —, DQ784667; P. peruviana (J.F.Macbr.)
Barneby, M. Nee 38898, NY, DQ790627, —,
DQ784670; P. stipulacea Ducke, L.P. de Queiroz et al.
3115, NY, DQ790634, —, DQ784675. Piptadeniastrum Brenan: P. africanum (Hook. f.) Brenan, D.
Harris 4319, E, AF521857, —, —. Piptadeniopsis
Burkart: P. lomentifera Burkart, M. Luckow 4476,
BAB/BH, AY944559, —, AY944541. Prosopidastrum
Burkart: P. mexicanum (Dressler) Burkart, Rebman
4021, DES, AY386919, —, —. Prosopis L.: P. glandulosa Torr. subsp. torreyana (L.D.Benson) A.E.Murray, Wojciechowski 875, ASU, AY386851, —, —;
P. pallida Kunth, M. Lavin 3088, BH, AF521860, —,
—. Pseudopiptadenia Rauschert: P. contorta (DC.)
G.P.Lewis & M.P.Lima, L.P. de Queiroz et al. 3366,
NY, DQ784676, —, DQ784676; P. suaveolens (Miq.)
J.W.Grimes, S.A. Mori et al. 24790, NY, DQ784677,
—, DQ784677. Pseudosamanea Harms: P. guachapele (Kunth) Harms, D. Murphy 350, MEL,
AF523079, AF524983, —. Samanea Merr.: S. saman
Merr., D. Murphy 357, MEL, AF523073, AF524984,
AF522965. Schleinitzia Warb.: S. insularum (Guill.)
Guinet, Waimanalo Res. Station, PI282460, BH,
AF521862, —, AF278491. Senegalia Raf.: *S. anisophylla (S.Watson) Seigler & Ebinger, Chase 14817, K,
JQ230134, JQ230159, JQ230244/JQ230245; S. berlandieri (Benth.) Britton & Rose, J. Miller 501,
CANB, AF274145, AF524998, AF522978; S. caffra
(Thunb.) P.J. Hurter & Mabb., RL1335, JRAU,
GQ872217, GQ872306, GQ872261; S. catechu (L.f.)
P.J. Hurter & Mabb., 615594, CANB, AF274141, —,
DQ371870; S. gaumeri (Blake) Britton & Rose,
DS16042, DS, DQ371895, —, DQ371858; S. gilliesii
(Steud.) Seigler & Ebinger, DLEG94-0167, DLEG,
DQ371882, —, DQ371860; S. glomerosa (Benth.)
Britton & Rose, 249, CANB, AF274147, AF525000,
AF522980; S. modesta (Wall.) P.J.Hurter, 615595,
CANB, AF274142, AF524995, AF522975; *S. muricata (L.) Britton & Rose, DS14548_JM1606, CANB,
EU812032, EU811974, EU440008; S. nigrescens
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523
RECLASSIFICATION OF AFRICAN ACACIA SPECIES
(Oliv.) P.J.Hurter, OM255, JRAU, GQ872237, —,
GQ872282; *S. occidentalis (Rose) Britton & Rose, J.
Miller
1219,
CANB,
EU812055,
EU811991,
EU440032; S. picachensis (Brandegee) Britton &
Rose, DS15981, DS, DQ371895, —, —; S. polyphylla
(DC.) Britton & Rose, 910150, DELEP, AF274147,
AF525000, AF522980; S. roemeriana (Scheele)
Britton & Rose, J. Miller 517, CANB, AF523099,
AF524997, AF522977; S. sororia (Standl.) Britton &
Rose, DS16067, DS, DQ371876, —, DQ371859;
*S. tenuifolia (L.) Britton & Rose, W. Thomas 9537, K,
—, —, JQ230243; *S. visco (Griseb) Seigler &
Ebinger, Conicet (s.n.), K, JQ230116, JQ230169,
JQ230239/JQ230240; *S. vogeliana (Steud.) Britton &
Rose, J. Miller 1603, CANB, EU812025, EU811969,
EU440001; S. wrightii (Benth.) Britton & Rose,
DLEG900444, DLEG, AF274148, —, DQ371854.
Stryphnodendron Mart.: S. porcatum D.A.Neill &
Occhioni f., D. Neill 14001, MO, AY944547, —,
AY944547; S. rotundifolium Mart., B.M.T. Walter
et al. 2913, NY, DQ784685, —, DQ784685. Vachellia
Wight & Arn.: *V. anegadensis (Britton) Seigler &
Ebinger, MH49, K, JQ265931, —, JQ230254/
JQ230255; V. campechiana (Mill.) Seigler & Ebinger,
MH81, MH, AF274133, —, AY574113; V. caven
(Molina) Seigler & Ebinger, J. Miller 247, CANB,
AF274131, AF524987, AF522967; *V. choriophylla
(Benth.) Seigler & Ebinger, J. Miller 1419, CANB,
EU812041, —, EU440017; V. collinsii (Saff.) Seigler &
Ebinger, DS16041, DS, DQ371884, —, DQ371869;
V. constricta (Benth.) Seigler & Ebinger, J. Miller 505,
CANB, DQ371883, AF524989, DQ371861; *V. cornig-
523
era (L.) Seigler & Ebinger, J. Miller 1344, CANB,
EU812045,
EU811981,
EU440021;
V. erioloba
(E.Mey.) P.J.Hurter, RL1298, JRAU, GQ872221,
GQ872310, GQ872265; V. farnesiana (L.) Wight &
Arn., T.J. Entwisle 2708, MEL, AF523115, AF195723,
AF195688/AF195669; V. gerrardi (Benth.) P.Hurter,
RL1321, JRAU, GQ872225, GQ872314, GQ872269;
V. karroo (Hayne) Banfi & Galasso, RL1282, JRAU,
GQ872230, —, GQ872274; *V. macracantha (Humb.
& Bonpl. ex Willd.) Seigler & Ebinger, J. Miller 1346,
CANB, EU812053, EU811989, EU440030; V. nilotica
(L.) P.J.Hurter & Mabb., RL1302, JRAU, GQ872238,
GQ872326, GQ872283; *OM626, JRAU, JQ230148,
—, JQ230210/JQ230211; *OM1063, JRAU, JQ265932,
JQ230197, JQ230271/JQ230272; *OM2607, JRAU,
JQ230149, JQ230198, JQ230290/JQ230291; *V. oviedoensis (R.Garcia & M.Mejía) Seigler & Ebinger, J.
Miller 1601, CANB, EU812029, —, EU440005; V. pennatula (Schltdl. & Cham.) Seigler & Ebinger,
DS16053, DS, DQ371878, —, DQ371855; V. schottii
(Torr.) Seigler & Ebinger, J. Miller 520, CANB,
AF274136,
AF524991,
AF522971;
V. vernicosa
(Britton & Rose) Seigler & Ebinger, J. Miller 265,
CANB, AF523113, AF524990, AF522970; V. xanthophloea (Benth.) P.J.Hurter, RL1291, JRAU,
GQ872255, GQ872343, GQ872300. Xerocladia
Harv.: X. viridiramis Taub., Krosnik 8244, BH,
EU000438, —, EU004653. Zapoteca H.M.Hern.:
Z. tetragona (Willd.) H.M.Hern., J. Miller 615626,
CANB, AF523097, AF524986, AF522966. Zygia
P.Browne: Z. lathetica Barneby & J.W.Grimes, D.
Neill 14002, MO, AY94456, —, AY944550.
© 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 500–523