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Non-Congruent Fossil and Phylogenetic Evidence on the Evolution of Climatic Niche in the Gondwana Genus Nothofagus

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Non-Congruent Fossil and Phylogenetic Evidence on the Evolution of Climatic Niche in the Gondwana Genus Nothofagus See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/284030463 Non-congruent fossil and phylogenetic evidence on the evolution of climatic niche in the Gondwana genus Nothofagus ARTICLE in JOURNAL OF BIOGEOGRAPHY · NOVEMBER 2015 Impact Factor: 4.59 · DOI: 10.1111/jbi.12650 READS 166 11 AUTHORS, INCLUDING: Felipe Hinojosa Mirta Quattrocchio University of Chile National Scientific and Technical Research … 43 PUBLICATIONS 780 CITATIONS 90 PUBLICATIONS 1,010 CITATIONS SEE PROFILE SEE PROFILE Kazuhiko Uemura Atsushi Yabe National Museum of Nature and Science National Museum of Nature and Science 22 PUBLICATIONS 266 CITATIONS 11 PUBLICATIONS 51 CITATIONS SEE PROFILE SEE PROFILE All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Mirta Quattrocchio letting you access and read them immediately. Retrieved on: 29 March 2016 Journal of Biogeography (J. Biogeogr.) (2015) ORIGINAL Non-congruent fossil and phylogenetic ARTICLE evidence on the evolution of climatic niche in the gondwana genus Nothofagus Luis Felipe Hinojosa1,2*, Aurora Gaxiola2,3, Marıa Fernanda Perez2,3, Francy Carvajal1,2, Marıa Francisca Campano1,2, Mirta Quattrocchio4, Harufumi Nishida5, Kazuhiko Uemura6, Atsushi Yabe6, Ramiro Bustamante1,2 and Mary T. K. Arroyo1,2 1Facultad de Ciencias, Universidad de Chile, ABSTRACT Santiago, Chile, 2Instituto de Ecologıa y Aim We used fossil and phylogenetic evidence to reconstruct climatic niche Biodiversidad (IEB), Santiago, Chile, 3 evolution in Nothofagus, a Gondwana genus distributed in tropical and temper- Departamento de Ecologıa, Facultad de Ciencias Biologicas, Pontificia Universidad ate latitudes. To assess whether the modern distribution of the genus can be Catolica de Chile, Santiago, Chile, explained by the tropical conservatism hypothesis, we tested three predictions: 4Departamento de Geologıa, Universidad (1) species from all Nothofagus subgenera coexisted under mesothermal cli- Nacional del Sur, Bahıa Blanca, Argentina, mates during the early Eocene; (2) tolerance to microthermal climates evolved 5Department of Biological Sciences, Chuo during the Eocene–Oligocene cooling from an ancestor that grew under University, Tokyo, Japan, 6Department of mesothermal conditions; and (3) the climatic niche in Nothofagus is phyloge- Geology and Palaeontology, National netically conserved. Museum of Nature and Science, Tsukuba, Location Australia, New Zealand, New Caledonia, Papua-New Guinea and Japan South America. Methods We estimated the palaeoclimate of the Early Eocene, fossil-bearing Ligorio Marquez Formation (LMF, Chile), using coexistence and leaf physiog- nomic analysis. We reconstructed ancestral climatic niches of Nothofagus using extant species distributions and a time-calibrated phylogeny. Finally, we used the morphological disparity index and phylogenetic generalized least squares to assess whether climatic variables follow a Brownian motion (BM) or an Ornstein–Uhlenbeck (OU) model of evolution. Results Our palaeoclimatic estimates suggest mesothermal conditions for the LMF, where macrofossils associated with subgenera Lophozonia and possibly Fuscospora, and fossil pollen of Brassospora and Fuscospora/Nothofagus were recorded. These results are not supported by our phylogenetic analysis, which instead suggests that the ancestor of Nothofagus lived under microthermal to marginally mesothermal conditions, with tolerance to mesothermal conditions evolving only in the subgenus Brassospora. Precipitation and temperature dimensions of the realized climatic niche fit with a gradual BM or constrained OU model of evolution. Main Conclusions Our results suggest that the use of phylogenetic recon- struction methods based only on present distributions of extant taxa to infer ancestral climatic niches is likely to lead to erroneous results when climatic requirements of ancestors differ from their extant descendants, or when much extinction has occurred. *Correspondence: Luis Felipe Hinojosa, Facultad de Ciencias Universidad de Chile & Keywords Instituto de Ecologıa y Biodiversidad (IEB), Eocene, Gondwana, into the tropics, niche modelling, Nothofagaceae, palaeo- Santiago, Chile. E-mail: [email protected] climate, phylogenetic signal, tropical conservatism hypothesis. ª 2015 John Wiley & Sons Ltd http://wileyonlinelibrary.com/journal/jbi 1 doi:10.1111/jbi.12650 L. F. Hinojosa et al. Nothofagus, Drimys, Weinmannia) (Arroyo et al., 1995; INTRODUCTION Villagran & Hinojosa, 1997; McGlone et al., 2001). Over the course of geological time, lineages either adapt to Nothofagus is considered an emblematic plant genus for new conditions, track their ancestral habitats or become understanding biogeographical processes of Gondwana extinct (Hoffmann & Sgro, 2011). In the case of plants, phy- lineages with modern distributions in both tropical and tem- logenetic and evolutionary studies suggest that related species perate regions (Linder & Crisp, 1995). Nothofagus comprises tend to grow in similar climates, latitudes and biomes, 43 species grouped into four monophyletic subgenera (Hill indicating that ecological requirements are phylogenetically & Read, 1991; Hill, 2001a). The tropical subgenus Bras- conserved (Crisp et al., 2009; Hawkins et al., 2011; Perez sospora (25 species), endemic to New Guinea and New Cale- et al., 2014). Consequently, habitat tracking, associated with donia, is sister to the temperate subgenus Nothofagus, niche conservatism, is the main explanation for the persis- endemic to southern South America, and together they are tence of lineages through geological climate change (Wing & sister to the other two temperate subgenera, Lophozonia and Greenwood, 1993). The hypothesis that lineages track Fuscospora (Romero, 1986; Swenson et al., 2000; Hill, 2001a; favoured climates has strong support, to the extent that it Sauquet et al., 2012). The majority of the literature on fossil tends to be assumed that ancestral climatic conditions are and extant Nothofagus uses the taxonomy erected by Hill & similar to those under which extant species of a lineage Read (1991), but Heenan & Smissen (2013) recently sug- thrive (Wing & Greenwood, 1993; Mosbrugger & Utescher, gested that morphological and molecular differences between 1997). Furthermore, it is assumed that ancestral climate the four subgenera are sufficient for these to be recognized niches may be inferred from a consideration of the ecological as separate genera. We do not adopt this revision in the pre- and climatic characteristics of modern taxa (Yesson & Cul- sent work, in order to avoid confusion between the modern ham, 2006; Evans et al., 2009). However, comparisons taxonomy and the fossil record. between phylogenetic inferences and those obtained from The fossil record of Nothofagus pollen dates back to the fossil records are scarce (Quental & Marshall, 2010). The fos- Late Cretaceous (80 Ma) at high latitudes in Antarctica sil record provides evidence about the past ecological and (Dettman et al., 1990; Hill & Dettman, 1996), but the lin- abiotic conditions under which species lived and thus can eage only became widespread and diverse after the Mid- contribute to our understanding of the constraints associated Eocene, under a mesothermal climate with little thermal with a phylogenetic approach. seasonality and high precipitation throughout the year Climatic niche conservatism has been invoked to explain (Romero, 1986; Hill & Dettman, 1996; McGowran et al., the well-documented global trend, whereby species diversity 2000; Hinojosa & Villagran, 2005; Macphail, 2007; Read in most groups declines from the tropics to the poles [e.g. et al., 2010). The low percentage of pollen and absence of tropical conservatism hypothesis (TCH; Wiens & Donoghue, macrofossils of Nothofagus in the fossil record for the warm 2004; Donoghue, 2008)]. TCH links both historical and evo- Early Eocene suggests that the genus was mostly restricted to lutionary assumptions (Kerkhoff et al., 2014). Historical high latitudes or high-altitude areas during that period explanations assume that lineages from low latitudes origi- (McGowran et al., 2000; Read et al., 2010). However, recent nated and underwent diversification under tropical climatic micro- and macrofossils of Nothofagus found at mid-lati- conditions (Donoghue, 2008), while evolutionary explana- tudes in southern South America from the Early Eocene sug- tions suggest that dispersal into the temperate zone is limited gest that Nothofagus lived at mid-latitudes during the Eocene by the capacity of organisms to adapt to new, colder and (Suarez & de la Cruz, 2000; Okuda et al., 2006). This fossil more seasonal climates, due to niche conservatism (Dono- record can be used to reconstruct the past climatic condi- ghue, 2008; Kerkhoff et al., 2014). Palaeoclimatic reconstruc- tions under which Nothofagus species from different subgen- tions have shown that near-tropical conditions dominated at era coexisted, and thus to assess whether Nothofagus was high latitudes (i.e. 60° S) in the Early Eocene (Carpenter capable of growing under a warm climate during the Early et al., 2012; Pross et al., 2012), when global temperatures Eocene. were 12 °C warmer than today (Zachos et al., 2001). During We examined the climatic history of Nothofagus using the Eocene–Oligocene glaciation and presence of ice-sheets phylogenetic models and fossil evidence obtained from the over the poles, global temperatures decreased by 7 °C mid-latitude, Early Eocene, Ligorio
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