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Evolution of the New Zealand Vascular Flora: Regional and Provincial Patterns of Richness, Radiation, and Endemism

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Evolution of the New Zealand Vascular Flora: Regional and Provincial Patterns of Richness, Radiation, and Endemism New Zealand Journal of Botany ISSN: 0028-825X (Print) 1175-8643 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzb20 Evolution of the New Zealand vascular flora: Regional and provincial patterns of richness, radiation, and endemism Geoffrey Rogers & Susan Walker To cite this article: Geoffrey Rogers & Susan Walker (2005) Evolution of the New Zealand vascular flora: Regional and provincial patterns of richness, radiation, and endemism, New Zealand Journal of Botany, 43:2, 381-414, DOI: 10.1080/0028825X.2005.9512963 To link to this article: http://dx.doi.org/10.1080/0028825X.2005.9512963 Published online: 17 Mar 2010. Submit your article to this journal Article views: 214 View related articles Citing articles: 4 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tnzb20 Download by: [Dept of Conservation] Date: 28 March 2017, At: 18:31 New Zealand Journal of Botany, 2005, Vol. 43: 381-414 381 0028-825X/05/4302-0381 © The Royal Society of New Zealand 2005 Evolution of the New Zealand vascular flora: regional and provincial patterns of richness, radiation, and endemism GEOFFREY ROGERS trast, mainland botanical provinces all show average Science & Research Unit levels of regional endemism after correcting for spe- Science, Technology and Information Services cies richness. With the exception of gymnosperms, Department of Conservation the most New Zealand-endemic plant groups have P.O. Box 5244 the most narrow regional distributions. We discuss Dunedin, New Zealand latitudinal patterns of endemism among plant groups SUSAN WALKER in terms of dispersal and adaptation. Land protec- Landcare Research tion measures will achieve different phylogenetic Private Bag 1930 outcomes in different landscapes across the New Dunedin, New Zealand Zealand botanical region. Keywords New Zealand; species richness; bioge- Abstract Analysis of species richness, radiation, ography; endemism; radiation and endemism in 171 list-regions and 19 botani- cal provinces of the New Zealand botanical region indicates that the Late Cenozoic radiation of the INTRODUCTION New Zealand vascular flora has produced diver- gent phylogenetic trends in regional and provincial Discussion of the biogeography and evolution of the floras. Floristic richness at all taxonomic levels is New Zealand vascular flora has focused on regional strongly determined by land area. After accounting patterns of endemism and substantial disjunctions in for area, most exceptionally species-rich list-regions species ranges. There are strong claims for endemic- occur immediately east of the South Island main rich latitudinal centres in northern North Island, divide, on calcareous substrates in West Nelson, or northern South Island, and southern South Island, on several central North Island ranges. Depauperate with depauperate zones in southern North Island floras are predominantly in the south on Auckland and the waist of the South Island (Cockayne 1928; and Campbell Islands, and four Central Otago and Burrows 1965; Wardle 1963, 1991; Druce 1984; two Southland ranges or basins. Family and genus McGlone 1985; McGlone et al. 2001). Reported pat- richness decrease with latitude, but it appears that terns of species disjunctions more or less reinforce there has been compensatory radiation from a few this predominant pattern (McGlone 1985; Wardle herbaceous families in southern island and upland 1988; Rogers 1989; Heads 1997, 1998). However, floras, so that species richness shows no latitudinal when the zones are analysed in detail, their degree trend. We present a measure of aggregate regional of endemism and disjunction vary in terms of bound- endemism that is comparable between list-regions ary position (Rogers 1989; Williams & Courtney and provinces of different floristic richness. Offshore 1998; Connor 2002). Biogeographic hypotheses for island groups are the most insular floras, with excep- these patterns have emphasised just one or permu- tionally high endemism across different taxonomic tations of tectonics, glaciation, and plant dispersal. ranks; northern island floras have high degrees of Cockayne (1926), Willet (1950), Wardle (1963), family and genus endemism and high phylogenetic and Burrows (1965) suggested that plants tended to diversity, while the subantarctic island floras have become concentrated in refugia due to the disruption high species endemism and high radiation. In con- of glacial ice and harsh climates. Climo (1975) was first to make extensive use of tectonism to explain biogeographic distributions of, in particular, snails. Panbiogeographic explanations have emphasised B02069; Online publication date 5 May 2005 tectonic movement of geological terranes (Craw Received 24 October 2002; accepted 20 January 2005 382 New Zealand Journal of Botany, 2005, Vol. 43 1989; Heads 1989,1998; Craw et al. 1999). Further as surrogate measures of ecosystem diversity, phy- work has debated the biogeographic origins of both logenetic diversity, biotic and ecological distinctive- the New Zealand flora and the patterns of endemism ness, and spatial heterogeneity. Assessment of New and disjunction within New Zealand (Raven 1973; Zealand biodiversity is advancing on all of these Wardle 1978; Pole 1994; Burrows 1998; Lee et al. fronts. Recent advances in spatial modelling tech- 2001; McGlone et al. 2001). Lee et al. (2001) char- niques to quantify and map environmental pattern acterised the Cenozoic evolution of the flora as a and distinctiveness (Overton & Leathwick 2001; filtering out of characteristically Australian lineages, Leathwick et al. 2003) will need validation against with compensatory radiation of an upland flora. Mc- databases of biological richness. Investigations Glone et al. (2001) favoured Pleistocene extinctions, of taxonomic richness and endemism at regional speciation, and dispersal and repeated glaciation to scales (e.g., Wardle 1988; McGlone et al. 2001) explain endemic and disjunct patterns. Using the are continuing to generate hypotheses regarding alpine flora, they have shown that less endemic ecological and evolutionary processes operating at plant groups have the widest distributions within large geographic scales, which exert control over New Zealand. Trewick & Wallis (2001) have used regional species pools (Rosenzweig 1995; Karlson invertebrate phylogeography and genetic distance to & Cornell 2002). Understanding of the origins of show that patterns of endemism are not consistent the New Zealand biota is also being advanced by with ancient vicariant processes, preferring instead studies of the fossil record (e.g., Pole 1994; Lee et al. the explanation of disparate responses to late Ceno- 2001) and elucidation of taxonomic and evolutionary zoic mountain building and glacial dislocation. relationships through molecular studies (e.g., Wag- There are several ways to gauge the biodiversity staff & Wardle 1999; Trewick & Wallis 2001). For value of floras, including species richness, richness the flora, molecular work is especially useful since adjusted for area, the relative abundance of species, Tertiary and Quaternary records are incomplete and the number of rare or endemic species, species rich- generally biased to woody plants (Wardle 1991). ness at higher taxonomic rank, and phylogenetic The debate on New Zealand plant biogeography diversity. New Zealand's insular biota shows marked needs to be advanced beyond numbers of endemic variations in the ratio of species to higher taxonomic and disjunct species in large, latitudinally defined ranks (Fenner et al. 1997), with particularly strik- zones, not least because endemism is a function of ing contrasts between the offshore islands and the sampling area and floristic richness. Divergent pat- mainland, and between coastal or lowland regions terns of richness, endemism, and radiation have been and those in the uplands (Rogers & Overton 2000). used previously to extend the concept of regionalism Fenner et al. (1997) promoted the measurement of in the flora of southern New Zealand (Rogers & phylogenetic diversity (i.e., the distribution of spe- Overton 2000). We now seek to typify gamma-scale cies among higher taxonomic ranks) for assessing biodiversity across the New Zealand botanical re- priorities and allocating resources for conserva- gion, at regional and provincial scales, by examin- tion, on the basis that if species are spread across ing measures of richness, endemism, and radiation numerous genera and families, a flora is likely to at various taxonomic ranks and across gradients of contain more genetic and character diversity than land area, species richness, and latitude. We intro- one where species are concentrated in few genera. duce a new measure of endemism that goes beyond Polasky et al. (2001) suggested that taxon diversity that of the simple number of species confined to an may be a good surrogate for phylogenetic diversity, area, by compiling measures of the relative ranges but Rodrigues & Gaston (2002) cautioned against of the entire floras of different regions. We focus this generalisation if the phylogenetic tree is highly particularly on the question of whether the Late unbalanced (highly ramified branches as opposed Cenozoic radiation of the New Zealand flora pro- to monophylic branches in different areas). Since duced divergent patterns of richness, radiation, and detailed phylogenies for all New Zealand plant line- endemism, particularly between the offshore islands ages will not be available
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