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A Paleogene Trans-Antarctic Distribution for Ripogonum (Ripogonaceae: Liliales)?

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A Paleogene Trans-Antarctic Distribution for Ripogonum (Ripogonaceae: Liliales)? Palaeontologia Electronica palaeo-electronica.org A Paleogene trans-Antarctic distribution for Ripogonum (Ripogonaceae: Liliales)? Raymond J. Carpenter, Peter Wilf, John G. Conran, and N. Rubén Cúneo ABSTRACT An impressive and growing list of biogeographically interesting plant and animal taxa occur in Paleogene sediments of both southern Australia and southern South America, indicating trans-Antarctic distributions during the warm past. Here, we pro- vide the first evidence that the living, woody, Australasian monocot Ripogonum was present during the early Eocene of Patagonia, Argentina. Two fossil leaves are suffi- ciently well preserved in overall shape, size, and fine venation details to be assigned to the genus, being closely comparable to leaves of the extant species R. album and R. scandens and the recently described early Eocene species R. tasmanicum from Tas- mania, Australia. The new species, R. americanum, is described. Overall, this evi- dence suggests that Ripogonum had a significantly larger past range, including Antarctica, during the early Paleogene, when Antarctica was much more closely con- nected to both Tasmania and South America and high latitude climates were warm and wet. Ecologically, the South American Ripogonum was probably a scrambling vine in mesic forest and grew in association with a rich understory flora of ferns and angio- sperms. Raymond J. Carpenter. School of Earth and Environmental Sciences, Benham Bldg DX 650 312, University of Adelaide, South Australia 5005, Australia and School of Plant Science, University of Tasmania, Private Bag 55, Hobart, Tasmania 7001, Australia. Peter Wilf. Department of Geosciences, Pennsylvania State University, University Park, Pennsylvania 16802, United States of America. John G. Conran. Australian Centre for Evolutionary Biology and Biodiversity, School of Earth and Environmental Sciences, Benham Bldg DX 650 312, University of Adelaide, South Australia 5005, Australia. N. Rubén Cúneo. Museo Paleontológico Egidio Feruglio, Consejo Nacional de Investigaciones Científicas y Técnicas, Trelew 9100, Chubut, Argentina. Keywords: Argentina; Australasia; Eocene; Liliales; new species; Ripogonum PE Article Number: 17.3.39A Copyright: Palaeontological Association October 2014 Submission: 8 February 2014. Acceptance: 29 September 2014 Carpenter, Raymond J., Wilf, Peter, Conran, John G., and Cúneo, N. Rubén. 2014. A Paleogene trans-Antarctic distribution for Ripogonum (Ripogonaceae: Liliales)? Palaeontologia Electronica Vol. 17, Issue 3;39A; 9p; palaeo-electronica.org/content/2014/921-early-eocene-ripogonum CARPENTER ET AL.: EARLY EOCENE RIPOGONUM INTRODUCTION eastern mainland Australia, New Zealand, and Papua New Guinea (Conran and Clifford, 1986), Interest in Southern Hemisphere biogeogra- and the Philesiaceae consists only of the Chilean phy has accelerated in recent years, in large part national flower, Lapageria rosea, and Philesia because of the discovery of new fossils and the magellanica, which both are species of the moist increased use of molecular data to estimate the temperate Valdivian rainforest (Conran and Clif- timing of lineage diversifications. During the early ford, 1998; Chileflora, 2012). Paleogene, Antarctica was largely ice-free, and Recently, Conran et al. (2009) described there were no significant water gaps between this Ripogonum tasmanicum from the early Eocene of continent and Australia and South America Tasmania. These authors also noted that the (Lawver et al., 2011). Moreover, early Eocene cli- potential for fossilization of Ripogonum leaves was mates at high latitudes were warm and wet enough good because the living leaves are robust, abun- to support complex vegetation types (Francis et al., dantly produced, and deciduous at senescence. 2008; Carpenter et al., 2012; Pross et al., 2012; Indeed, Ripogonum fossils are often locally abun- Contreras et al., 2013). The Antarctic region during dant in New Zealand. These records are mostly this interval can thus be seen as a portal for biotic Miocene (Holden, 1983; Pole, 1993, 1996, 2007b; interchange over vast distances, and the legacy of Lee et al., 2012), but a recent Eocene discovery this past can still be seen in the distributions of has been made (Conran et al., 2013). Here, we numerous modern taxa, famously including provide evidence that Ripogonum was also present Nothofagus (Veblen et al., 1996). in Patagonia, Argentina, during the early Eocene. Some plant taxa provide striking direct evi- dence for trans-Antarctic paleodistributions MATERIALS AND METHODS because they have Paleogene fossil records in both Australia and southern South America, even The two fossils presented here were recov- though they may be extinct in one or both modern ered 29 November and 8 December, 2002, from a regions, or more circumstantially because they single quarry (quarry LH13 of Wilf et al., 2003) of have a fossil record in just one area but survive in the Laguna del Hunco caldera-lake deposit, the other (Hill and Carpenter, 1991). Knowledge of Chubut Province, Argentina, which has been the living Australasian taxa that are present as fos- described and discussed by numerous authors sils in South America but extinct there now, in par- (e.g., Berry, 1925; Aragón and Mazzoni, 1997; Wilf ticular, has dramatically increased in recent years, et al., 2003, 2005, 2013; Zamaloa et al., 2006; and a large number of these contributions have Hermsen et al., 2012). The 170 m stratigraphic come from the early Eocene (ca. 52.2 Ma) Laguna section at Laguna del Hunco has 27 fossil plant del Hunco caldera-lake beds of Chubut, Patagonia, localities distributed vertically and laterally, concen- Argentina (see Wilf et al., 2013 for a review). trated in a densely fossiliferous interval of ca. 60 m These elements of the Laguna del Hunco fossil thickness; there are two paleomagnetic reversals assemblage are accordingly considered to repre- and three dated tuffs within the section (Wilf et al., sent the western reaches of a vast, trans-Antarctic 2003). One of these tuffs, within the fossiliferous paleorainforest flora (Wilf et al., 2013). This flora interval, was further subjected to multiple 40Ar-39Ar not only included large tree species, but also a analyses of sanidines, yielding an age of 52.22 ± range of understory species, notably including sev- 0.22 Ma (early Eocene, Ypresian; M. Smith in Wilf eral genera of tree and ground ferns (Carvalho et et al., 2005 and Wilf, 2012), which is the working al., 2013) and probable small-statured Laurales age for the entire floral assemblage. Laguna del trees (Knight and Wilf, 2013). Hunco preserved vertebrates and insects as well One small lineage of the monocot order Lilia- as more than 200 species of land plants, most of les that shows a classic extant Gondwanan distri- them angiosperms (Wilf et al., 2005, 2009). Since bution is Ripogonaceae + Philesiaceae. This clade 1999, ca. 6000 fossil plant specimens from Laguna is supported by numerous multigene analyses del Hunco have been recovered by expeditions (Patterson and Givnish, 2002; Davis et al., 2004; from Museo Paleontólogico Egidio Feruglio (MEF, Janssen and Bremer, 2004; Givnish et al., 2005; Trelew, Chubut), where they are curated (reposi- Fay et al., 2006; Kim et al., 2013; Petersen et al., tory acronym MPEF-Pb). 2013). The species of both families occupy mostly Among these large collections at MEF are two wet forests, where they occur as woody understory compression specimens of the informal leaf mor- shrubs or vines. The Australasian Ripogonaceae photype TY171 (Wilf et al., 2005), which is charac- comprise only six extant species of Ripogonum in terised by having a single primary vein with 2 PALAEO-ELECTRONICA.ORG divergent secondary vein pairs (marginal and lami- arising from the primary vein ~4 mm from the leaf nal), with the laminal pair suprabasal. These speci- base at ~30°, decurrent; submarginal vein pair mens were photographed with a Nikon (Tokyo, emerging from the leaf base at ~60°; all secondary Japan) D700 digital SLR camera, and the images veins weakening apically, the basal pair forming were adjusted to optimise brightness and contrast part of a series of exmedial loops formed by per- using Adobe Photoshop Elements® software. The current tertiary veins that connect with the supra- leaf surfaces of both fossils were closely examined basal secondaries most obviously from the top of under epifluorescence, but no useful cuticular fea- the basal third of the leaf to the apex. Tertiary veins tures were preserved. Nevertheless, the excellent between primary and suprabasal secondaries preservation of the leaf architecture showed com- opposite percurrent to random reticulate. Marginal pellingly clear similarities to living Ripogonum. ultimate veins looped. Also, considering the rarity of these fossils (only Comments two specimens among thousands, from more than a decade of field work in a remote location) and Conover (1983) and Conran et al. (2009) dis- their unusual geographic interest, we felt it import- cussed leaf architecture and venation in Ripogo- ant to present the specimens as is. num and for taxa with similar leaves. The latter An image of a leaf from a herbarium specimen concluded that the presence of divergent second- of Ripogonum held at the Queensland Herbarium ary vein pairs (marginal and laminal) with the lami- (BRI) was captured using a Nikon (Tokyo, Japan) nal pair suprabasal (Figure 1.2) is typical only of D5000 SLR camera. Venation patterns were also Ripogonum species. Related and superficially simi-
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