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• W3044294522 abstract "In global plant geography, ever since the mid-twentieth century, there are two different types of maps doing different things. On the one hand, there are maps of floral kingdoms, subdivided into regions, provinces, etc. – in one word phytochoria. These are large and contiguous pieces of the world with distinctive floras, that is, high internal similarity in the plant taxa present. Often entire families and genera of plants are characteristic to a kingdom (Good, 1974). Kingdoms and their larger subunits can be continents, islands, or island-like in being somehow isolated from neighbouring units. What this suggests is that their boundaries are in some cases defined by the inability of lineages to overcome historical limitations even when physical environments on the two sides of the divide are fairly similar. Biomes, on the other hand, are first and foremost defined on the basis of the dominant plant growth forms/life forms, which may be determined by climate and other components of the abiotic environment. Biomes often occur patchily within one another, as dictated by climatic or soil patterns (Prentice et al., 1992). The only thing biomes and kingdoms have in common is that they both have to be globally mappable. Biomes can, of course, also be mapped at a much finer scale, and this is increasingly achievable using remotely acquired imagery, with applications in conservation and beyond (Gairola et al., 2013). The recent Research review by Mucina (2019) provides a useful synthesis on what biomes have been understood to be, over the one-century lifespan of the biome concept. The author correctly points out that several factors have been incorporated over time in the definition of biomes. These include plant physiognomy (dominant growth forms/life forms as indicated earlier), climate, as well as other abiotic factors such as soils and fire regimes. The biome concept has been further refined with the inclusion of functional aspects for both plants and whole ecosystems, biogeochemical cycles, etc. Intriguingly, Mucina also includes here evolutionary history. Typically, evolution is mentioned on the historical side of things, and as such is more relevant to kingdoms, regions and other phytochoria. This inclusion is however no accident; Mucina argues that there are good reasons to bring in evolution as a key factor relevant to biomes, and that evolutionary history has already been used in the definition of biomes in several studies. The review also makes brief reference to phytochoria among several other concepts featuring in the history of botanical maps, but not in contrast with biomes. In the final figure of the review, Mucina refers to ‘biogeographic spatial units’, but without any clarity as to the precise nature of such units. While this appears to be an attempt to bring biomes and phytochoria together, this is nowhere clearly articulated. Here I argue that, although evolution plays a key role (actually, more than one; see Fig. 1) in biome assembly and function, it is nevertheless preferable to keep biomes and phytochoria separate, and in fact to contrast them where the role of evolution is concerned. I also contend that, to date, evolutionary history has not been used in the definition of biomes, and probably should not be in the future either. I refer to biomes as mapped by Olson et al. (2001), and floral kingdoms as in Good (1974). The latter are partly congruent with the realms listed by Olson et al. (2001), which are mostly relevant to animal distributions. Despite the differences between kingdoms and realms in these two references and between either of them and other recent ones, most of the points I make later and conclusions I derive should remain valid. Typically, kingdoms and even most regions therein include more than one biome, and all major biomes occur in more than one region. For example, tropical rainforest occurs in both the Neotropical and Paleotropical kingdoms, although with distinct floras in the two cases. World’s smallest, the Cape Floral Kingdom, has been historically defined around fynbos (the local name for the Mediterranean-type biome) but has always had significant components from other biomes (arid, forest), and more recently has been expanded to include the Succulent Karoo (arid) biome on equal footing with the fynbos (Born et al., 2006). The Holarctic Kingdom appears surface-wise to be dominated by boreal forest, but is most diverse elsewhere in terms of floral elements (Milne & Abbot, 2002). In many places, the boundaries between biomes coincide with the boundaries between kingdoms, but not the whole biomes as such. Regarding spatial scale, this is relevant to mapping both biomes and phytochoria, but (at least partly) in different ways. Biomes often occur in small patches, and whether this gets mapped or not depends on the grain size of the mapping exercise. Phytochoria are defined using both endemic and more widespread taxa, but are usually characterized by endemic taxa/lineages, and the higher the level in the phytochorion hierarchy, the higher the taxonomic rank of the characteristic taxa – or the evolutionary age of the characteristic lineages. Sometimes different biomes interdigitating in a given area may be dominated by floristic elements characteristic of different phytochoria. From a vegetation point of view, such an area can be mapped as a patchwork of biomes if the grain is fine enough, or specifically defined as a mosaic if too coarse. However, where phytochoria (and especially high-level phytochoria such as kingdoms) are concerned, contiguity is key (Morrone, 2015), and a vegetation mosaic can be defined as a transition zone. At fine scales, and especially in continental settings where dispersal barriers are not an issue, vegetation types and low-level phytochoria may largely coincide, as both can be defined by using species-level taxa – sometimes dominant and sometimes not. The concept of ‘evolutionary history’ as relevant to vegetation and phytochoria requires some unpacking. When saying that biomes end up somewhat different on different continents depending on what plant lineages are regionally available (Pennington et al., 2004; Moncrieff et al., 2015; Mucina, 2019), one must be primarily referring to the already present evolutionary background of lineages. Lineages also evolve within biomes, occasionally changing their character, although more often they align themselves with the pre-existing character of the biome, and seldom cross into other biomes (Crisp et al., 2009; Donoghue & Edwards, 2014). Where phytochoria are concerned, all taxa present, whether pre-existent or evolved in situ, have an effect on phytochorion definition (Fig. 1). There is a bit of confusion between what Mucina’s predictive and retrodictive models might mean to this question, but the answer should be still, no. Certainly, biomes can be characterized in terms of the evolutionary history of the plants they are made up of, and patterns can be quite divergent across continents. For example, the presence of eucalypts in Australian savanna means that this biome has a slightly different climatic niche here compared to Africa, where the genus is absent (Moncrieff et al., 2015). Even in species-poor and apparently homogeneous biomes such as the boreal forests, the presence of characteristic lineages with different fire ecology results in major discrepancies in, e.g. carbon sequestration (Wirth, 2005; Rogers et al., 2015). Such lineage-driven internal variability can be substantial, but if we are still using the word biome, this is in spite of, and not because of, floristic differences. Contrary to what appears to be suggested in Mucina’s Table 2, Moncrieff et al. (2015) do not use information about the evolution of plant lineages to define biomes. Evolutionary biome assembly is simply mentioned in that study as one of several explanations for the differences between the climatic envelopes characterizing one biome in different parts of the world. In fact, basically none of the references listed under Mucina’s ‘Evolutionary biome assembly’ and ‘Biome ecology and historical biogeography meeting’ headings are relevant to defining biomes. Most of them simply point out that, as would be expected, the plants a biome is made of come with their own evolutionary baggage. Reichstein et al. (2014) do not, as claimed by Mucina, define biomes in terms of functional attributes; Higgins et al. (2016) do, but there is no evolutionary component. One study where lineage presence is indeed used to fine-tune (although perhaps not define) biomes (Ratnam et al., 2011, C4 grasses in savanna) is curiously not mentioned by Mucina. The one study cited by Mucina that does construct biomes based on floristic data (Silva de Miranda et al., 2018) can be discarded because of fine taxonomic and geographic scale – meaning the authors are simply reconstructing phytochoria, but, at that scale, phytochoria and biomes overlap. The distinction between biomes and phytochoria is one relevant globally and for broad plant lineages. No doubt, as Mucina explains, the biome concept has evolved over the years. But has it truly incorporated evolution, the hallmark of phytochoria? At this stage, phytochoria and biomes play different and well-defined roles in our understanding of global plant geography, and maintaining the distinction between the two seems vital, both for a theoretical understanding of global patterns in plant distributions, and for conservation efforts. In a global conservation context, it has been shown that there are substantial mismatches between high-biodiversity areas and high-ecosystem service areas (Girardello et al., 2019). Phytochoria and zoochoria are defined based on their characteristic taxon assemblages, and as such are useful tools for visualizing biodiversity conservation priorities (Giraudo & Arzamendia, 2017), while biomes are more consistent in terms of ecosystem service value. Thus, an integrated approach to conservation should ideally be grounded in mapping exercises that consider the two separately." @default.
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• W3044294522 date "2020-07-21" @default.
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• W3044294522 title "Biomes are nobody’s kingdom: on environmental and historical plant geography" @default.
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