Albany Thicket Biome
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% S % 19 (2006) Albany Thicket Biome 10 David B. Hoare, Ladislav Mucina, Michael C. Rutherford, Jan H.J. Vlok, Doug I.W. Euston-Brown, Anthony R. Palmer, Leslie W. Powrie, Richard G. Lechmere-Oertel, Şerban M. Procheş, Anthony P. Dold and Robert A. Ward Table of Contents 1 Introduction: Delimitation and Global Perspective 542 2 Major Vegetation Patterns 544 3 Ecology: Climate, Geology, Soils and Natural Processes 544 3.1 Climate 544 3.2 Geology and Soils 545 3.3 Natural Processes 546 4 Origins and Biogeography 547 4.1 Origins of the Albany Thicket Biome 547 4.2 Biogeography 548 5 Land Use History 548 6 Current Status, Threats and Actions 549 7 Further Research 550 8 Descriptions of Vegetation Units 550 9 Credits 565 10 References 565 List of Vegetation Units AT 1 Southern Cape Valley Thicket 550 AT 2 Gamka Thicket 551 AT 3 Groot Thicket 552 AT 4 Gamtoos Thicket 553 AT 5 Sundays Noorsveld 555 AT 6 Sundays Thicket 556 AT 7 Coega Bontveld 557 AT 8 Kowie Thicket 558 AT 9 Albany Coastal Belt 559 AT 10 Great Fish Noorsveld 560 AT 11 Great Fish Thicket 561 AT 12 Buffels Thicket 562 AT 13 Eastern Cape Escarpment Thicket 563 AT 14 Camdebo Escarpment Thicket 563 Figure 10.1 AT 8 Kowie Thicket: Kowie River meandering in the Waters Meeting Nature Reserve near Bathurst (Eastern Cape), surrounded by dense thickets dominated by succulent Euphorbia trees (on steep slopes and subkrantz positions) and by dry-forest habitats housing patches of FOz 6 Southern Coastal Forest lower down close to the river. 541 L. Mucina S % 19 (2006) 1. Introduction: Delimitation and Global Table 10.1 Major correspondence between vegetation Perspective units and STEP vegetation types (Vlok & Euston-Brown 2002). Those STEP units are listed which have at least 80% of their area within the vegetation units and do not necessar- Following the earlier work of Acocks (1953), the structurally unusual vegetation of the semi-arid river valleys of the eastern ily constitute a high proportion of the vegetation unit (these seaboard of South Africa was described as Valley Bushveld. This proportions are given under the headings in the descrip- nomenclature probably followed the colloquial term used by tion of each unit). Only Gouritz Valley Thicket is (much) less the agriculturalists, mainly livestock farmers, who found it a than 80%. largely impenetrable thicket which had to be ‘opened-up’ to allow access to domestic livestock (cattle and goats). According Vegetation Units STEP to Acocks, this comprised ‘a semi-succulent thorny scrub 2–3 metres in height’. AT 1 Southern Cape Valley Thicket Gouritz Valley Thicket Rutherford & Westfall (1986) classified the biomes of South AT 2 Gamka Thicket Gamka Arid Spekboomveld Africa on the basis of dominant life-form combinations Gamka Spekboom Thicket (Raunkiaer 1934, Cain 1950) and climatic features. Therefore, Oudtshoorn Karroid Thicket the areas now considered here to be the Albany Thicket Biome AT 3 Groot Thicket Baviaans Spekboom Thicket were classified by them as Savanna on the basis of the domi- Baviaans Valley Thicket nance of phanerophytes associated with, on average, hemicryp- Bethelsdorp Bontveld tophyte co-dominance (Rutherford & Westfall 1986), although Groot Arid Spekboomveld they acknowledged the existence of a variety of life-form com- Kleinpoort Karroid Thicket binations. This view continued in the work of Scholes (1997), AT 4 Gamtoos Thicket Gamtoos Arid Spekboomveld who mapped the vegetation of the discussed region as a part Gamtoos Bontveld of the broad-leaved Savanna. Following White & Moll (1978) Gamtoos Thicket and Cowling (1983), some evidence for its classification as a Gamtoos Valley Thicket distinct structural and floristic unit was presented, and this pro- Kromme Forest Thicket Otterford Forest Thicket vided justification for the later recognition of the Thicket Biome Vanstadens Forest Thicket (Low & Rebelo 1996, 1998). Rutherford & Westfall (1986) noted that many parts of the areas classified here as Thicket AT 5 Sundays Noorsveld Sundays Noorsveld Biome, namely Spekboomveld and the Fish River Scrub, Addo AT 6 Sundays Thicket Elands Forest Thicket Bush and Sundays River Scrub forms of Valley Bushveld (Acocks Koedoeskloof Karroid Thicket 1988), had vegetation co-dominated by phanerophytes and Kremlin Grassland Thicket chamaephytes. This corresponds to a ‘missing biome’ that they Motherwell Karroid Thicket Sundays Spekboom Thicket suggested may be recognised in the future. This concept is for- Sundays Spekboomveld mally adopted here. Recent analyses, primarily within the STEP Sundays Thicket project (Cowling et al. 2003), have confirmed that the climatic Sundays Valley Thicket uniqueness (Robertson & Palmer 2002) and peculiar vegetation Zuurberg Fynbos Thicket structure resulting from a unique combination of constituent AT 7 Coega Bontveld Grass Ridge Bontveld growth forms (Vlok & Euston-Brown 2002) as well as floristic AT 8 Kowie Thicket Albany Spekboom Thicket diversity (including high regional endemism) justify its recogni- Albany Spekboomveld tion as a biome. Albany Thicket The term ‘Albany Thicket’ is a concept recognised by the WWF Albany Valley Thicket (Olson et al. 2001, Burgess et al. 2004) where this vegeta- Ecca Bontveld tion type (‘ecoregion’) is listed as a part of the broader cat- Salem Karroid Thicket egory (biome) ‘mediterranean forests, woodlands, and scrubs’, Shamwari Grassland Thicket thus, as a part of the Cape Floristic Region—a misplacement Thorndale Forest Thicket that partly reflects the transitional nature of the biome. The AT 9 Albany Coastal Belt Geluk Grassland Thicket same WWF categorisation does not recognise a global thicket Hamburg Dune Thicket biome. Various thicket formations in Africa, Madagascar, Asia, Kiwane Dune Thicket Australia and the Americas are spread over four different Nanaga Savanna Thicket biomes. In Africa, patches of typical thicket are included in two Paterson Savanna Thicket Zuney Strandveld other ‘biomes’: ‘montane grasslands and shrublands’ (e.g. Jos Plateau forest-grassland mosaic), and ‘tropical and subtropi- AT10 Great Fish Noorsveld Fish Noorsveld cal grasslands, savannas, and shrublands’ (e.g. Somali Acacia– AT11 Great Fish Thicket Crossroads Grassland Thicket Commiphora bushlands and thickets). Worldwide, other thicket Doubledrift Karroid Thicket types are classified as ‘tropical and subtropical dry broadleaf Fish Spekboom Thicket forests’ (see Olson et al. 2001). In terms of vegetation struc- Fish Thicket Fish Valley Thicket ture and climate at least two other regions qualify as analo- Hartebeest Karroid Thicket gous to Albany Thicket. These include the thickets of the Chaco straddling the border areas between Argentina and Paraguay AT12 Buffels Thicket Buffels Thicket in South America (Lewis 1991, Cábido et al. 1992, 1994) and Buffels Valley Thicket Kei Thicket the Didiereaceae-rich semideciduous thickets (also called dry Mountcoke Grassland Thicket forests) of southern and southwestern Madagascar (Koechlin et al. 1974, Grubb 2003). Madagascar spiny thickets, largely AT13 Eastern Cape Escarpment Escarpment Thicket Thicket equivalent in vegetation structure to southern Africa’s Albany Thicket, are classified under ‘deserts and xeric shrublands’. AT14 Camdebo Escarpment Thicket Escarpment Spekboom Thicket 542 Albany Thicket Biome S % 19 (2006) Perhaps also the Somali-Masai thickets as described by White the equatorial and tropical regions of Africa, and Addo Bush (1983), the so-called ‘Vine Thickets’ of Australian Queensland (Acocks 1953) is similar to certain thickets of the Arid Lowveld (Webb 1978) and succulent-rich thickets of northern Venezuela (Acocks 1953, 1988) in the Mpumalanga Lowveld, northern and Colombia (Matteuci 1987) could be viewed as part of the KwaZulu-Natal and parts of Swaziland. The Albany Thicket also global subtropical succulent-rich thickets. has a unique phylogenetic and biogeographic origin and is transitional between Nama-Karoo and the subtropical regions Thicket in the valleys from the other side of the Great Kei River of the eastern seaboard of southern Africa. to the Thukela River in KwaZulu-Natal are not recognised here as being part of the Albany Thicket Biome (the Great Kei River Rainfall can occur at any time of the year in Albany Thicket, Valley does contain some mapped Albany Thicket). In fact, this which occurs in a climatic interface between an all-year rainfall is a gradual transition from the characteristic bimodal rainfall, zone in the west and a stronger summer-rainfall zone in the with no seasonal period of pronounced drought of the Albany northeast (Schulze 1997). Thicket vegetation is considerably Thicket to the summer-rainfall areas with dry winters typical fragmented and displaced by renosterveld and fynbos in the of the Savanna Biome. Also in terms of species affinities, an winter-rainfall zone and by grasslands and savanna in the sum- increasing number of typical savanna species occur along this mer-rainfall zone (Vlok & Euston-Brown 2002). The climate in gradient towards the northeast. Woody elements of strand- itself is not the primary selective force for this pattern, but rather veld units to the west, including parts of the west coast, are the fire regimes that are determined by seasonal precipitation. not regarded as part of the biome despite their recognition as Evidence for this lies in the occurrence of outliers of thicket veg- thicket by some authorities (e.g. Cowling 1984). The current etation in fire refugia in the winter- and summer-rainfall zones. delimitation of the Albany Thicket Biome closely follows