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Global Ecology and Biogeography Volume 18 Number 4 July 2009 Pages 393–520 ., Pp geb_v18_i4_ofbcover 6/4/09 9:06 AM Page 1 Global Ecology and Biogeography ISSN 1466–822X Global Ecology Volume 18, Number 4, July 2009 Global Ecology and Biogeography Volume 18 Number 4 Contents July 2009 and Biogeography Research Papers A Journal of Macroecology H.Verbruggen, L. Tyberghein, K. Pauly, C.Vlaeminck, K.Van Nieuwenhuyze,W.H. C. F.Kooistra, F.Leliaert and O. De Clerck: Macroecology meets macroevolution: evolutionary niche dynamics in the seaweed Halimeda 393–405 Editor-in-Chief: R.A. Pyron and F.T.Burbrink: Can the tropical conservatism hypothesis explain temperate species richness patterns? David J. Currie An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini 406–415 B. Oberle, J. B. Grace and J. M. Chase: Beneath the veil: plant growth form influences the strength of species richness–productivity relationships in forests 416–425 J. Belmaker: Species richness of resident and transient coral-dwelling fish responds differentially to regional diversity 426–436 18Volume Number 4 July 2009 Pages 393–520 S. Normand, U.A. Treier, C. Randin, P.Vittoz,A. Guisan and J.-C. Svenning: Importance of abiotic stress as a range-limit determinant for European plants: insights from species responses to climatic gradients 437–449 S. M. Shirley and S. Kark: The role of species traits and taxonomic patterns in alien bird impacts 450–459 E. Batllori, J. J. Camarero, J. M. Ninot and E. Gutiérrez: Seedling recruitment, survival and facilitation in alpine Pinus uncinata tree line ecotones. Implications and potential responses to climate warming 460–472 P.Prieto, J. Peñuelas, Ü. Niinemets, R. Ogaya, I. K. Schmidt, C. Beier,A. Tietema,A. Sowerby, B.A. Emmett, E. K. Láng, G. Kröel-Dulay, B. Lhotsky, C. Cesaraccio, G. Pellizzaro, G. de Dato, C. Sirca and M. Estiarte: Changes in the onset of spring growth in shrubland species in response to experimental warming along a north–south gradient in Europe 473–484 J. C. Linares, J. J. Camarero and J.A. Carreira: Interacting effects of changes in climate and forest cover on mortality and growth of the southernmost European fir forests 485–497 M.A. Beketov: The Rapoport effect is detected in a river system and is based on nested organization 498–506 Research Review D. Corenblit, J. Steiger,A. M. Gurnell and R. J. Naiman: Plants intertwine fluvial landform dynamics with ecological succession and natural selection: a niche construction perspective for riparian systems 507–520 Front cover: Joint analysis of macroecological and phylogenetic data sets permits inference of the evolutionary dynamics of the macroecological niche and assists interpretation of biogeographical patterns. Study of the evolutionary niche dynamics of the calcified green seaweed genus Halimeda (Verbruggen et al., pp. 393–405) shows niche conservatism for nutrient-poor, tropical habitats in most of the genus except one section that has invaded colder, more nutrient-rich waters multiple times during the Cenozoic. Interpretation of global niche models and the species phylogeny provides evidence that species occupy only part of the suitable habitat as a consequence of vicariance events. Niche models also identify understudied areas that can be targeted for the discovery of species of special biogeographical interest. The front cover shows a sea surface temperature map (top), a representative of the genus Halimeda on a coral reef in the Maldives (bottom left), and the inferred evolution of sea surface temperature affinities in a lineage of the genus (bottom right). World list abbreviation: Global Ecol. Biogeogr. www.blackwellpublishing.com/geb www.blackwellpublishing.com This journal is available online at Wiley InterScience. Visit www3.interscience.wiley.com to search the articles and register for table of contents e-mail alerts. Global Ecology and Biogeography, (Global Ecol. Biogeogr.) (2009) 18, 393–405 Blackwell Publishing Ltd RESEARCH Macroecology meets macroevolution: PAPER evolutionary niche dynamics in the seaweed Halimeda Heroen Verbruggen1*†, Lennert Tyberghein1†, Klaas Pauly1†, Caroline Vlaeminck1, Katrien Van Nieuwenhuyze1, Wiebe H.C.F. Kooistra2, Frederik Leliaert1 and Olivier De Clerck1 1Phycology Research Group and Center for ABSTRACT Molecular Phylogenetics and Evolution, Ghent Aim Because of their broad distribution in geographical and ecological dimensions, University, Krijgslaan 281 S8 (WE11), B-9000 seaweeds (marine macroalgae) offer great potential as models for marine biogeo- Ghent, Belgium, 2Stazione Zoologica ‘Anton Dohrn’, Villa Comunale, 80121 Naples, Italy graphical inquiry and exploration of the interface between macroecology and macroevolution. This study aims to characterize evolutionary niche dynamics in the common green seaweed genus Halimeda, use the observed insights to gain under- standing of the biogeographical history of the genus and predict habitats that can be targeted for the discovery of species of special biogeographical interest. Location Tropical and subtropical coastal waters. Methods The evolutionary history of the genus is characterized using molecular phylogenetics and relaxed molecular clock analysis. Niche modelling is carried out with maximum entropy techniques and uses macroecological data derived from global satellite imagery. Evolutionary niche dynamics are inferred through applica- tion of ancestral character state estimation. Results A nearly comprehensive molecular phylogeny of the genus was inferred from a six-locus dataset. Macroecological niche models showed that species distribution ranges are considerably smaller than their potential ranges. We show strong phylogenetic signal in various macroecological niche features. Main conclusions The evolution of Halimeda is characterized by conservatism for tropical, nutrient-depleted habitats, yet one section of the genus managed to invade colder habitats multiple times independently. Niche models indicate that the restricted geographical ranges of Halimeda species are not due to habitat unsuitability, strengthening the case for dispersal limitation. Niche models identified hotspots of habitat suitability of Caribbean species in the eastern Pacific Ocean. We propose that these hotspots be targeted for discovery of new species separated from their Caribbean *Correspondence: Heroen Verbruggen, Phycology Research Group, Krijgslaan 281 S8, siblings since the Pliocene rise of the Central American Isthmus. B-9000 Ghent, Belgium. Keywords E-mail: [email protected] †These authors contributed equally to this Geographical information systems, Halimeda, historical biogeography, macroecology, work niche evolution, niche conservatism, niche modelling, phylogenetics. survival, and the realized niche, which is a subset of the funda- INTRODUCTION mental niche that a species actually occupies. Species tolerances Various interacting features influence the distribution of a are determined by their morphological, reproductive and species. The niche of a species is commonly defined as the set of physiological traits, which are in turn susceptible to evolutionary biotic and abiotic conditions in which it is able to persist and forces. Hence, niche characteristics can be interpreted as evolutionary maintain stable population sizes (Hutchinson, 1957). Further phenomena. Understanding niche evolution yields valuable insights distinction is made between a species’ fundamental niche, which into biogeography, biodiversity patterns and conservation biology consists of the set of all conditions that allow for its long-term (Wiens & Graham, 2005; Rissler et al., 2006; Wiens et al., 2007). DOI: 10.1111/j.1466-8238.2009.00463.x © 2009 Blackwell Publishing Ltd www.blackwellpublishing.com/geb 393 H. Verbruggen et al. The niche concept provides a conceptual framework to predict identified, questions about what causes them remain (Kooistra geographical distributions of species. Niche models establish the et al., 2002; Verbruggen et al., 2005b). Are species restricted to macroecological preferences of a given species based on observed one ocean basin because of habitat unsuitability in the other distribution records and a set of macroecological variables, and basin or should the limited distribution ranges be attributed to these preferences can subsequently be used to predict geographical dispersal limitation? It is also not known with certainty which areas with suitable habitat for the species (e.g. Guisan & Thuiller, vicariance event may be responsible for the phylogenetic 2005; Raxworthy et al., 2007; Rissler & Apodaca, 2007). The separation of Indo-Pacific and Atlantic lineages. So far, two availability of macroecological data, either in the form of geological events have been implied: the Miocene closure of the remotely sensed or interpolated measurement data, is increasing Tethys Seaway in the Middle East and the Pliocene shoaling of the and has already provided many biological studies with environ- Central American Isthmus (Kooistra et al., 2002; Verbruggen mental information (Kozak et al., 2008). To date, most ecological et al., 2005b). niche modelling studies have focused on terrestrial organisms. The first goal of the present study is to investigate the A notable exception is the study by Graham et al. (2007), which evolutionary niche dynamics of the seaweed genus Halimeda, used a synthetic oceanographic and ecophysiological model to focusing on niche dimensions relevant to global geographical identify known kelp populations and predict the existence of distributions rather than local distributional issues such as undiscovered
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