1 What Are Relicts

Dictionaries define a "relict" as something that has survived, usually as a trace, from the past. In biology, relicts are distinctive populations or species that typically are small in size or severely restricted in geographic range. Biologists distinguish between taxonomic and biogeographic relicts. Taxonomic relicts are a few or sole survivors of a once diverse taxonomic assemblage, whereas biogeographic relicts are descendants of once widespread taxa (or populations) that now have a narrow geographic distribution (Lomolino et al. 2006). Both categories sometimes coincide, as for example in the case of "living fossils" (such as ginko, lungfishes, crossopterygians, or marsupials) that closely resemble their ancient ancestors in overall phenotype (Futuyma 2005; Lomolino et al. 2006; Beierkuhnlein 2007). In the following, we focus on biogeographic relicts.

Climatic and other large-scale environmental changes can have fundamental impact on the distributional patterns of species and alter the composition of communities and ecosystems (e.g. Hewitt 1999; Schweiger et al. 2009). Species react by range shifts, or by local or regional extinctions when their adaptive capacity is exceeded (Agarwal 2001). The origins and distributions of modern-day relicts can often be related to environmental changes of the past. Glacial periods and warm interglacial periods of the Quaternary, in particular, often help us to understand how relict species and

Musée national d'histoire naturelle, Section Zoologie des Invertébrés, L-2160 Luxembourg, Luxembourg e-mail: Janchristianhabel@gmx.de J.C. Habel and T. Schmitt

Department of Biogeography, Trier University, D-54286 Trier, Germany T. Assmann

Institute of Ecology and Environmental Chemistry, Leuphana University Lüneburg, D-21335 Lüneburg, Germany

J.C. Avise

University of California, CA 92697 Irvine, USA

J.C. Habel and T. Assmann (eds.), Relict Species: Phylogeography and Conservation Biology, 1 DOI 10.1007/978-3-540-92160-8_1, © Springer-Verlag Berlin Heidelberg 2010

populations arise and sometimes have served as important sources for re-colonizing much larger areas (de Lattin 1967). For example, many thermophilic species survived the last ice age in relatively small refugia on Europe's Southern peninsulas. During interglacial periods, these species often colonized large areas. Thus, many temperate species that are widespread today existed as small relict populations just a few thousand years ago (cf. Schmitt 2007). Conversely, many cold-adapted species show evidence of formerly wide distributions, followed by severe range restrictions during postglacial warming. Prominent examples of the latter are the butterflies Proclossiana eunomia pictured on this book (front side) and Lycaena helle (small picture on the back side), which currently resides only in high-elevation mountain enclaves and at more Northern latitudes that have been shown to coincide with distinctive genetic clusters in this species (Habel et al. 2009). Although such species today (and during the other warm periods between glacial periods) are restricted to small areas, they may have the potential to recolonize larger areas if and when the climate cools again. In general, the assumption that relict species inevitably are on their way to extinction may be erroneous; at least some relict populations undoubtedly retain the potential to adapt to a broad spectrum of environmental conditions (Hampe and Petit 2005).

Range restrictions and expansions are also known to result from human-mediated landscape changes. For example, woodlands covered large areas of Central Europe prior to anthropogenic deforestation. Especially during the Middle Ages and early modern times, these extensive woodlands were degraded to a few small remnants. After the establishment of modern forestry and changes in land use, woodland coverage again increased enormously. The former woodland remnants preserved forest-inhabiting plants and animals, some of which still show relict-like distributions but others of which were able to re-colonize large areas (e.g. Assmann 1999; Drees et al. 2008). Comparable changes are known for numerous habitat types and regions. These alternating scenarios of range expansions and restrictions remind us that species with relict-like versus wide distributions are not necessarily contradictory indicators of past environmental changes.

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