A New Fossil Wood of Peltophoroxylon (Leguminosae: Caesalpinioideae) from the El Palmar Formation (Late Pleistocene), Entre Ríos, Argentina
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Soledad RamosIAWA et al. Journal– Pleistocene 35 (2), Peltophoroxylon 2014: 199–212 from Argentina 199 A NEW FOSSIL WOOD OF PELTOpHOROXYLON (LEGUMINOSaE: CaESaLPINIOIDEaE) FROM THE EL PaLMaR FORMaTION (LaTE PLEISTOcENE), ENTRE RÍOS, ARGENTINa R. Soledad Ramos1,2,*, Mariana Brea1,3 and Romina Pardo4 1Centro de Investigaciones Científicas y Transferencia de Tecnología a la Producción (CICyTTP- CONICET), Dr. Matteri y España SN, E3105BWA, Diamante, Entre Ríos, Argentina 2FONCyT - Agencia Nacional de Promoción Científica y Tecnológica 3CONICET - Consejo Nacional de Investigaciones Científicas y Técnicas 4Cátedra de Dasonomía, Facultad de Ciencias Naturales, Universidad Nacional de Salta, Avenida Bolivia 5150, 4400 Salta, Argentina *Corresponding author; e-mail: [email protected] abstract This paper describes the first record ofPeltophoroxylon (Ramanujam) Müller- Stoll et Mädel 1967 from the late Pleistocene of Argentina. The fossil specimens were recovered from the Colonia Ayuí and Punta Viracho fossil localities of the El Palmar Formation, located in the middle part of the Uruguay Basin, eastern Argentina. The diagnostic features are: growth ring boundaries demarcated by marginal parenchyma, medium-sized vestured intervessel pits, vessel-ray paren- chyma pits similar in size and shape to intervessel pits, vasicentric to lozenge type aliform axial parenchyma, biseriate (70%) and uniseriate (30%) homocel- lular rays, non-septate and septate fibers, and long chains (10+) of prismatic crystals in chambered axial parenchyma cells. These features suggest a relation- ship with Peltophorum (Vogel) Benth. (Leguminosae: Caesalpinioideae). The vessel diameter and vessel density of the El Palmar woods are consistent with the temperate-warm, humid-semiarid climate inferred for this region during the late Pleistocene. Keywords: Wood anatomy, Peltophoroxylon, Pleistocene, Uruguay Basin, Argentina. INTRODUCTION Today, the Fabaceae or Leguminosae includes approximately 730 genera and over 19,400 species worldwide (Lewis et al. 2005). The subfamily Caesalpinioideae, contain- ing approximately 2,250 species in 171 genera (Bruneau et al. 2008), is paraphyletic, and it includes the monophyletic tribes Cercideae and Detarieae and the paraphyletic tribes Cassieae and Caesalpinieae (Lewis et al. 2005). The South American extant flora includes 65 genera of Caesalpinioideae (Zuloaga & Morrone 1999; Ulibarri 2008) and inhabits tropical and subtropical areas, from rain forests, evergreen or deciduous for- ests, to savannas, semi-deserts and high mountains (Ulibarri 2008). In South America there are 31 genera in the tribe Caesalpinieae. Brazil has the highest concentration of endemics at the genus levels (Bruneau et al. 2008; Ulibarri 2008). © International Association of Wood Anatomists, 2014 DOI 10.1163/22941932-00000060 Published by Koninklijke Brill NV, Leiden Downloaded from Brill.com10/08/2021 05:25:38PM via free access 200 IAWA Journal 35 (2), 2014 Figure 1. Location map showing Colonia Ayuí and Punta Viracho fossil localities. South America has an extensive and diverse legume fossil record, and woods with affinities to all subfamilies are reported (see Pujana et al. 2011). Worldwide, the old- est putative fossil wood assigned to the subfamily Caesalpinioideae is from the late Cretaceous of Africa and resembles the modern tribe Cassieae (Wheeler & Baas 1992). South American fossil woods that seem reliably assigned to the Caesalpinioideae are reported from the Miocene onwards (Pons 1980; Martínez & Rodriguez Brizuela 2011; Brea et al. 2012; see Pujana et al. 2011). The El Palmar Formation is well known for its rich fossil wood and phytolith as- semblages. Taxa in these assemblages are interpreted to be trees, shrubs and herbs belonging to the Lauraceae, Fabaceae, Podostemaceae, Myrtaceae, Combretaceae, Anacardiaceae, Poaceae, Cyperaceae and Arecaceae (Zucol et al. 2005; Brea et al. 2010; Brea & Zucol 2011; Patterer & Brea 2011; Patterer 2012; Ramos et al. 2012). The Leguminosae from the El Palmar Formation include five Mimosoideae:Menen - doxylon areniensis, M. mesopotamiensis, M. piptadiensis, Mimosoxylon caccavariae, Prosopisinoxylon castroae, and two Papilionoideae: Holocalyxylon cozzoi and Am- buranoxylon tortorellii (Lutz 1979; Brea 1999; Zucol et al. 2005; Brea et al. 2010). The new fossil wood described herein represents the first report of fossil wood with affinities to Caesalpinioideae from the Uruguay Basin. Downloaded from Brill.com10/08/2021 05:25:38PM via free access Soledad Ramos et al. – Pleistocene Peltophoroxylon from Argentina 201 STUDY AREA AND GEOLOGICAL SETTING Fossil woods were collected in the Colonia Ayuí and Punta Viracho localities of the El Palmar Formation. This fluvial unit is located in the southernmost part of the Uruguay Basin, in eastern Argentina (Fig. 1). The El Palmar Formation consists of medium, reddish and yellowish ochre sands, medium-to-thick sandstone layers with planar stratification and lenses of gravel and pebbles. Sand strata and gravel lenses represent channel facies, while fine sediments represent floodplain facies. This formation, which is 3 to 12 m thick, is found at the modern surface and has not been buried since its deposition (Iriondo 1980; Iriondo & Kröhling 2008; Kröhling 2009). At Federación city in Argentina, this formation was dated to 80,670 ± 13,420 years BP by TL (thermoluminescense dating) (Iriondo & Kröhling 2001), while near to the city of Salto in Uruguay, it was dated to 88,370± 35,680 years BP (Iriondo & Kröhling 2008). MATERIALS AND METHODS Two fragments of wood were found in sediments of the El Palmar Formation (Iriondo 1980; Iriondo & Kröhling 2008; Kröhling 2009). One specimen (CIDPALBO-MEG 111) was collected by Cristina Vassallo de Cettour from the Colonia Ayuí locality. The other specimen (CIDPALBO-MEG 148) was collected by one of the authors (MB) from the Punta Viracho locality, both from Entre Ríos Province, Argentina (Fig. 1). Both specimens are silicified. The CIDPALBO-MEG 111 wood is 27 cm long and 12 cm in diameter and CIDPALBO-MEG 148 wood is 7.6 cm long and 4.2 cm in diameter. Thin sections were made using standard petrographic techniques. Anatomical terms used in this paper generally follow the recommendations of the IAWA List of Microscopic Features for Hardwood Identification (IAWA Committee 1989), with some terms also taken from Chattaway (1932), Kribs (1935) and Carlquist (2001). The bibliographic lists by Gregory (1994) and Gregory et al. (2009) were used. To compare the samples to extant and fossil species, we used the InsideWood website (InsideWood 2004-onwards; Wheeler 2011) and descriptions by Metcalfe and Chalk (1950), Cozzo (1951), Tortorelli (1956), Gasson et al. (2003, 2009) and Evans et al. (2006). Pardo (2012) studied the wood anatomy of extant Peltophorum dubium. The samples she studied are deposited in the Laboratorio de Anatomía Vegetal, Facultad de Ciencias Naturales, Universidad Nacional de Salta, Argentina (LANAVE). Systematic assignment follows APG (2009). Names used are as found in The Interna- tional Plant Names Index (The Plant List 2010) and the Index Nominum Genericorum (ING) (Farr & Zijlstra 1996). The quantitative values in the anatomical description are averages of 25 measure- ments, in all cases the average is cited first, followed by the minimum and maximum values, which are given in parentheses. The material was studied with a Nikon Eclipse E 200 light microscope, and photomicrographs were taken with a Nikon Coolpix S4 digital camera. The fossil specimen and microscope slides are deposited in the Labo- ratorio de Paleobotánica (CICyTTP-CONICET), Diamante, Argentina, under the acro- nym CIDPALBO-MEG for wood specimens and CIDPALBO-MIC for slides. Downloaded from Brill.com10/08/2021 05:25:38PM via free access 202 IAWA Journal 35 (2), 2014 Downloaded from Brill.com10/08/2021 05:25:38PM via free access Soledad Ramos et al. – Pleistocene Peltophoroxylon from Argentina 203 RESULTS FabaLES Bromhead 1838 Leguminosae Jussieu 1789 Caesalpinioideae de Candolle1825 Genus: Peltophoroxylon (Ramanujam) Müller-Stoll et Mädel 1967 Type species: Peltophoroxylon variegatum (Ramanujam) Müller-Stoll et Mädel 1967 Species: Peltophoroxylon uruguayensis Ramos, Brea et Pardo, sp. nov. (Fig. 2 & 3) Specific diagnosis – Growth rings distinct, demarcated by narrow (less than 3 cells wide) marginal parenchyma bands of only a few cells wide; diffuse porous; vessels solitary and in radial multiples of 2–4, rarely in clusters, circular to oval in outline, medium to small in diameter with thin walls; perforation plates simple; intervessel pits alternate, vestured and small to medium; vessel-ray pits similar in size and shape to intervessel pits; gums or other deposits present; axial parenchyma scanty paratracheal, vasicentric, and aliform, diffuse parenchyma scarce; fibers mostly non-septate, rarely septate; rays mostly biseriate and occasionally uniseriate, homocellular composed exclusively by procumbent cells; prismatic crystals in long chains in chambered axial parenchyma cells. Holotype – CIDPALBO-MEG 111 specimen, CIDPALBO-MIC 1368 (three micro- scope slides). Type locality – Colonia Ayuí, Entre Ríos, Argentina. Paratype – CIDPALBO-MEG 148 specimen, CIDPALBO-MIC 1571 (three micro- scope slides). Locality – Punta Viracho, Entre Ríos, Argentina. Horizon – upper part of the El Palmar Formation. Age – late Pleistocene. Etymology – Specific epithet, uruguayensis, refers to the region where the fossil specimens were found. Botanical affinity– Leguminosae-Caesalpinioideae,