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DENDRON » No/Nr: 43 » November 2011 36 review review CLASSIFICATION AND NOMENCLATURE OF THE GENUS ACACIA Classification and nomenclature of the genus Acacia (Leguminosae), with emphasis on Africa Abstract The widespread and well-known genus Acacia has experienced a huge amount of taxonomic turmoil over the past couple of decades. In this contribution the nomenclatural history of Acacia is outlined. Evidence for the recent split of the genus into five segregate genera is provided and the retypification from the African species A. nilotica to the Australian A. penninervis is discussed critically. The International Code for Botanical Nomenclature (ICBN) is introduced and discussed as a set of guidelines for taxonomic change. Lastly, some ideas are provided for plant name users of how to deal with this nomenclatural inconvenience. “No other taxonomic proposal has ever had the potential for disruption to nomenclature on a continental scale as the one to split Acacia.” – Orchard & Maslin 2005 1. Introduction Convincing evidence has been accumulating over the past two to three decades that the large, well-known and charismatic genus Acacia Mill. is polyphyletic (Maslin et al. 2003; Orchard & Maslin 2003). Polyphyletic refers to a group derived from different common ancestors. The ideal in plant classifcation is to recognize monophyletic groups. In the case of a monophyletic group all members are derived from the same recent common ancestor. Many taxonomic workers now agree that Acacia senso lato (as traditionally defined), meaning Acacia in a broad sense, should in fact be five segregate genera, because five separate monophyletic groups can be recognised within the genus (Maslin et al. 2003). In 2005 the genus was split and retypified in accordance to Proposal 1584 by Orchard & Maslin (2003). This caused a plant nomenclatural uproar, the likes of which had never been seen and hopefully will never be seen again. The aim of this contribution is to review the developments in Acacia s.l. phylogeny and its impacts on the classification and the nomenclature of the group, with emphasis on Africa. I will further attempt to critically evaluate the split and retypification of the genus and provide William a. Haddad some thoughts on how to cope with this great nomenclatural inconvenience. 2. Background on the genus Acacia Although the palaeobotanical evidence is inconclusive, it is generally believed that the genus Acacia s.l. evolved in the tropical lowland forests of West Gondwana, but there is no agreement as to which section should be considered ancestral. The various sections in existence today must have evolved from the original form in secondary centres where the climatic or edaphic factors were different (Ross 1979). The oldest Acacia s.l. fossil was found in Tanzania and dated at 46 mya (Bouchenak-Khelladi et al. 2010). The highly complex nomenclatural history (outlined in Table 1) of the genus Acacia s.l. started with its formal description by Phillip Miller in 1754. It was not until 1842, however, that the generic limits were clearly defined by George Bentham by restricting the name to mimosoid plants with numerous free stamens (Maslin et al. 2003). The genus was only typified in 1913 by Britton and Brown to the African species A. scorpioides (L.) W.Wight (=A. nilotica (L.) Delile). Since that time many authors have proposed different ways to subdivide the genus, making use of various different characters (Bentham 1875; Ross 1979; Vassal 1981; Pedley 1986; Orchard & Maslin 2003). Acacia, in the broad sense, is the second largest genus in the family Leguminosae (Fabaceae s.l.) or the largest genus in the family Mimosaceae, if considered under an alternate classification system, with about 1540 living taxa (specific & intra-specific)(Orchard & Maslin 2005). It is widely distributed across the world’s tropics and sub-tropical zones with centres of diversity in Australia (± 1000 species), the Americas (±180 species), Africa (± 160 species), Asia (±100 species) and a couple of species on the Pacific Islands (Thiele et al. 2011) (Figure 1). As of yet, there are no explanations for this remarkably disjunct geographic distribution (Bouchenak- Khelladi et al. 2010). Ross (1979) describes the ecological role of Acacia s.l. very satisfactorily: “There is probably no group of trees or shrubs in Africa that can rival Acacia in the combined importance of its ecology and extent of its geographical range. Almost everywhere from South Africa northwards to the Mediterranean, where the climate is reasonably dry, there will CLASSIFICATION AND NOMENCLATURE OF THE GENUS ACACIA Year Nomenclatural Event Key Reference 1754 Genus Acacia described. Miller (1754) 1842 Generic limits defined 1875 Considered one genus with six series. Bentham (1875) 1913 Genus Acacia typified to A. nilotica. 1981 Considered as one genus with three subgenera. Vassal (1981) 1986 Proposal to raise subgenera to three individual genera. Pedley (1986) 2003 Proposal 1584 to split genus into five genera and to move the type of name Acacia to A. penninervis. Orchard & Maslin (2003) 2004 The IAPT Committee for Spermatophyta recommends proposal 1584. Brummitt (2004) 2005 2011 Proposal 1584 is controversially accepted at the nomenclature section of the 16th IBC in Vienna. The type is moved and five different genera are now recognized. An attempt is made at the 17th IBC in Melbourne to overturn the decisions made in Vienna. The attempt failed and the type remains an Australian species. TABLE 1.– Simplified nomenclatural history of the genus Acacia. 180 160 Acacia be found, usually frequently and often in dominant abundance. Only in the evergreen forest regions of Africa do Acacia play a comparatively subordinate role”. 100 1000 review review FIGURE 1.– Distribution map showing the range of Acacia s.l. with approximated numbers of endemic taxa per continent. Image from www.worldwidewattle.com and data from Thiele et al. (2011). 37 DENDRON » No/Nr: 43 » November 2011

DENDRON » No/Nr: 43 » November 2011<br />

36<br />

review<br />

review<br />

CLASSIFICATION AND NOMENCLATURE OF THE GENUS ACACIA<br />

Classification and nom<strong>en</strong>clature<br />

of the g<strong>en</strong>us Acacia (Leguminosae), with emphasis on Africa<br />

Abstract<br />

The widespread and well-known g<strong>en</strong>us Acacia<br />

has experi<strong>en</strong>ced a huge amount of taxonomic<br />

turmoil over the past <strong>co</strong>uple of decades. In this<br />

<strong>co</strong>ntribution the nom<strong>en</strong>clatural history of Acacia is<br />

outlined. Evid<strong>en</strong>ce for the rec<strong>en</strong>t split of the g<strong>en</strong>us<br />

into five segregate g<strong>en</strong>era is provided and the<br />

retypification from the African species A. nilotica to<br />

the Australian A. p<strong>en</strong>ninervis is discussed critically.<br />

The International Code for Botanical Nom<strong>en</strong>clature<br />

(ICBN) is introduced and discussed as a set of<br />

guidelines for taxonomic change. Lastly, some ideas<br />

are provided for plant name users of how to deal<br />

with this nom<strong>en</strong>clatural in<strong>co</strong>nv<strong>en</strong>i<strong>en</strong>ce.<br />

“No other taxonomic proposal<br />

has ever had the pot<strong>en</strong>tial for<br />

disruption to nom<strong>en</strong>clature on a<br />

<strong>co</strong>ntin<strong>en</strong>tal scale as the one to split<br />

Acacia.” – Orchard & Maslin 2005<br />

1. Introduction<br />

Convincing evid<strong>en</strong>ce has be<strong>en</strong> accumulating<br />

over the past two to three decades that the large,<br />

well-known and charismatic g<strong>en</strong>us Acacia Mill. is<br />

polyphyletic (Maslin et al. 2003; Orchard & Maslin<br />

2003). Polyphyletic refers to a group derived from<br />

differ<strong>en</strong>t <strong>co</strong>mmon ancestors. The ideal in plant<br />

classifcation is to re<strong>co</strong>gnize monophyletic groups.<br />

In the case of a monophyletic group all members<br />

are derived from the same rec<strong>en</strong>t <strong>co</strong>mmon ancestor.<br />

Many taxonomic workers now agree that Acacia<br />

s<strong>en</strong>so lato (as traditionally defined), meaning Acacia<br />

in a broad s<strong>en</strong>se, should in fact be five segregate<br />

g<strong>en</strong>era, because five separate monophyletic groups<br />

can be re<strong>co</strong>gnised within the g<strong>en</strong>us (Maslin et al.<br />

2003). In 2005 the g<strong>en</strong>us was split and retypified<br />

in ac<strong>co</strong>rdance to Proposal 1584 by Orchard &<br />

Maslin (2003). This caused a plant nom<strong>en</strong>clatural<br />

uproar, the likes of which had never be<strong>en</strong> se<strong>en</strong> and<br />

hopefully will never be se<strong>en</strong> again. The aim of this<br />

<strong>co</strong>ntribution is to review the developm<strong>en</strong>ts in Acacia<br />

s.l. phylog<strong>en</strong>y and its impacts on the classification<br />

and the nom<strong>en</strong>clature of the group, with emphasis<br />

on Africa. I will further attempt to critically evaluate<br />

the split and retypification of the g<strong>en</strong>us and provide<br />

William a. Haddad<br />

some thoughts on how to <strong>co</strong>pe with this great<br />

nom<strong>en</strong>clatural in<strong>co</strong>nv<strong>en</strong>i<strong>en</strong>ce.<br />

2. Background on the g<strong>en</strong>us Acacia<br />

Although the palaeobotanical evid<strong>en</strong>ce is<br />

in<strong>co</strong>nclusive, it is g<strong>en</strong>erally believed that the g<strong>en</strong>us<br />

Acacia s.l. evolved in the tropical lowland forests<br />

of West Gondwana, but there is no agreem<strong>en</strong>t as<br />

to which section should be <strong>co</strong>nsidered ancestral.<br />

The various sections in exist<strong>en</strong>ce today must<br />

have evolved from the original form in se<strong>co</strong>ndary<br />

c<strong>en</strong>tres where the climatic or edaphic factors<br />

were differ<strong>en</strong>t (Ross 1979). The oldest Acacia s.l.<br />

fossil was found in Tan<strong>za</strong>nia and dated at 46 mya<br />

(Bouch<strong>en</strong>ak-Khelladi et al. 2010).<br />

The highly <strong>co</strong>mplex nom<strong>en</strong>clatural history<br />

(outlined in Table 1) of the g<strong>en</strong>us Acacia s.l.<br />

started with its formal description by Phillip<br />

Miller in 1754. It was not until 1842, however, that<br />

the g<strong>en</strong>eric limits were clearly defined by George<br />

B<strong>en</strong>tham by restricting the name to mimosoid<br />

plants with numerous free stam<strong>en</strong>s (Maslin et<br />

al. 2003). The g<strong>en</strong>us was only typified in 1913<br />

by Britton and Brown to the African species A.<br />

s<strong>co</strong>rpioides (L.) W.Wight (=A. nilotica (L.) Delile).<br />

Since that time many authors have proposed<br />

differ<strong>en</strong>t ways to subdivide the g<strong>en</strong>us, making use<br />

of various differ<strong>en</strong>t characters (B<strong>en</strong>tham 1875;<br />

Ross 1979; Vassal 1981; Pedley 1986; Orchard &<br />

Maslin 2003).<br />

Acacia, in the broad s<strong>en</strong>se, is the se<strong>co</strong>nd largest<br />

g<strong>en</strong>us in the family Leguminosae (Fabaceae s.l.)<br />

or the largest g<strong>en</strong>us in the family Mimosaceae,<br />

if <strong>co</strong>nsidered under an alternate classification<br />

system, with about 1540 living taxa (specific<br />

& intra-specific)(Orchard & Maslin 2005). It is<br />

widely distributed across the world’s tropics and<br />

sub-tropical zones with c<strong>en</strong>tres of diversity in<br />

Australia (± 1000 species), the Americas (±180<br />

species), Africa (± 160 species), Asia (±100<br />

species) and a <strong>co</strong>uple of species on the Pacific<br />

Islands (Thiele et al. 2011) (Figure 1). As of yet,<br />

there are no explanations for this remarkably<br />

disjunct geographic distribution (Bouch<strong>en</strong>ak-<br />

Khelladi et al. 2010). Ross (1979) describes the<br />

e<strong>co</strong>logical role of Acacia s.l. very satisfactorily:<br />

“There is probably no group of trees or shrubs<br />

in Africa that can rival Acacia in the <strong>co</strong>mbined<br />

importance of its e<strong>co</strong>logy and ext<strong>en</strong>t of its<br />

geographical range. Almost everywhere from<br />

South Africa northwards to the Mediterranean,<br />

where the climate is reasonably dry, there will<br />

CLASSIFICATION AND NOMENCLATURE OF THE GENUS ACACIA<br />

Year Nom<strong>en</strong>clatural Ev<strong>en</strong>t Key Refer<strong>en</strong>ce<br />

1754 G<strong>en</strong>us Acacia described. Miller (1754)<br />

1842 G<strong>en</strong>eric limits defined<br />

1875 Considered one g<strong>en</strong>us with six series. B<strong>en</strong>tham (1875)<br />

1913 G<strong>en</strong>us Acacia typified to A. nilotica.<br />

1981 Considered as one g<strong>en</strong>us with three subg<strong>en</strong>era. Vassal (1981)<br />

1986 Proposal to raise subg<strong>en</strong>era to three individual g<strong>en</strong>era. Pedley (1986)<br />

2003<br />

Proposal 1584 to split g<strong>en</strong>us into five g<strong>en</strong>era and to move the type of<br />

name Acacia to A. p<strong>en</strong>ninervis.<br />

Orchard & Maslin<br />

(2003)<br />

2004 The IAPT Committee for Spermatophyta re<strong>co</strong>mm<strong>en</strong>ds proposal 1584. Brummitt (2004)<br />

2005<br />

2011<br />

Proposal 1584 is <strong>co</strong>ntroversially accepted at the nom<strong>en</strong>clature<br />

section of the 16th IBC in Vi<strong>en</strong>na. The type is moved and five differ<strong>en</strong>t<br />

g<strong>en</strong>era are now re<strong>co</strong>gnized.<br />

An attempt is made at the 17th IBC in Melbourne to overturn the<br />

decisions made in Vi<strong>en</strong>na. The attempt failed and the type remains an<br />

Australian species.<br />

TABLE 1.– Simplified nom<strong>en</strong>clatural history of the g<strong>en</strong>us Acacia.<br />

180<br />

160<br />

Acacia be found, usually frequ<strong>en</strong>tly and oft<strong>en</strong> in<br />

dominant abundance. Only in the evergre<strong>en</strong> forest<br />

regions of Africa do Acacia play a <strong>co</strong>mparatively<br />

subordinate role”.<br />

100<br />

1000<br />

review<br />

review<br />

FIGURE 1.– Distribution map showing the range of Acacia s.l. with approximated numbers of <strong>en</strong>demic taxa<br />

per <strong>co</strong>ntin<strong>en</strong>t. Image from www.worldwidewattle.<strong>co</strong>m and data from Thiele et al. (2011).<br />

37<br />

DENDRON » No/Nr: 43 » November 2011

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