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New Records of Humiriaceae Fossil Fruits from the Oligocene and Early

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New Records of Humiriaceae Fossil Fruits from the Oligocene and Early Boletín de la Sociedad Geológica Mexicana / 2018 / 223 New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene of the western Azuero Peninsula, Panamá Nicolas Pérez-Consuegra, Daniel E. Góngora, Fabiany Herrera, Carlos Jaramillo, Camilo Montes, Aura M. Cuervo-Gómez, Austin Hendy, Alejandro Machado, Damian Cárdenas, German Bayona ABSTRACT Nicolas Pérez-Consuegra ABSTRACT RESUMEN [email protected] Department of Earth Sciences, Syracuse Uni- versity, Syracuse, New York 13244, USA. Understanding the origin of the di- Para entender el origen de la diversidad de los Smithsonian Tropical Research Institute, versity in Central American forests bosques de América Central, se necesita inte- Balboa, Ancón, Panamá. requires the integration of both ex- grar estudios de plantas actuales y fósiles. En Daniel E. Góngora tant and fossil taxa. Here, we provide este trabajo, describimos fósiles de Humiria- Aura M. Cuervo-Gómez a description of Humiriaceae fossils ceae, excavados de dos nuevas secuencias Departamento de Geociencias, Universidad from two new sedimentary sequenc- sedimentarias en la Península de Azuero, de los Andes, Carrera 1 No. 18A-12, Bogotá, es in the Azuero Peninsula, Panamá. Panamá. Los fósiles fueron encontrados en Colombia. Fossils were recovered from Oligo- depósitos marinos-marginales del Oligoce- Fabiany Herrera cene (one locality) and Early Mio- no (una localidad) y del Mioceno tempra- Chicago Botanic Garden, 1000 Lake Cook cene (two localities) marginal marine no (dos localidades). Describimos nuevos Road, Glencoe, Illinois 60022, USA. deposits. We describe new specimens especímenes y aumentamos la descripción Carlos Jaramillo and augment the generic description morfológica de Lacunofructus cuatrecasana Alejandro Machado of Lacunofructus cuatrecasana Herrera, Herrera, Manchester et Jaramillo para las Damian Cárdenas Manchester et Jaramillo, and present localidades del Oligoceno y Mioceno tempra- Smithsonian Tropical Research Institute, a new record of Sacoglottis sp. Our no y presentamos un nuevo registro de Saco- Balboa, Ancón, Panamá. results expand the temporal and geo- glottis sp., para el Mioceno temprano. Estos Camilo Montes graphical distribution of both taxa nuevos registros expanden los rangos de dis- Geology, Universidad del Norte, Km. 5 Vía and show that Humiriaceae was an tribución geográfica y temporal para ambos Puerto Colombia, Barranquilla, Colombia. important constituent of early Cen- taxones, y además demuestran que la familia Departamento de Geociencias, Universidad de los Andes, Carrera 1 No. 18A-12, Bogotá, tral American forests, and that this Humiriaceae fue un importante constituyente Colombia. family was dispersed in between Cen- de los primeros bosques en América Central. tral and South America before the La evidencia presentada también sugiere que Austin Hendy final closure of the Central American hubo dispersiones de especies de Humiriaceae History Museum of Los Angeles County, Los Angeles, CA 90007, USA. Seaway. Further studies are necessary entre Centro y Sur América mucho antes del to establish whether the origin of cierre del Istmo de Panamá. Se necesitan German Bayona this family was in Central or South más estudios para establecer con claridad si Corporación Geológica ARES, Bogotá, America. el origen de Humiriaceae se dio en América Colombia. Central o Sur América. Keywords: Central America, fossil endocarps, Lacunofruc- Palabras clave: América Cen- tus, Neotropical forests, Isth- tral, endocarpos fósiles, mus of Panama, Sacoglottis. Lacunofructus, bosques Neo- tropicales, Istmo de Panamá, Sacoglottis. BOL. SOC. GEOL. MEX. 2018 VOL. 70 NO. 1 P. 223 ‒ 239 http://dx.doi.org/10.18268/BSGM2018v70n1a13 Manuscript received: October, 1, 2016. Corrected manuscript received: August 25, 2017. Manuscript accepted: September 1, 2017. New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene 224 / Boletín de la Sociedad Geológica Mexicana / 2018 1. Introduction ple Humiriaceae, Juglandaceae, Malvaceae, Ca- lophyllaceae, Connaraceae and Chrysobalanacea Southern Central America (i.e., Panamá and (Herrera et al., 2010, 2012, 2014a, 2014b; Carval- Costa Rica) is one of the regions with the highest ho et al., 2013; Rodríguez-Reyes et al., 2014, 2017; INTRODUCTION extant angiosperm diversity in the world (Barth- Jud et al., 2016; Jud and Nelson, 2017; MacFadden lott et al., 1996). This plant diversity flourishes in et al., 2017; Nelson and Jud, 2017). varied forest types that grow over a complex geo- We report new permineralized and carbonized logic landscape. The emergence of the Isthmus of fossil endocarps and seeds of Humiriaceae from Panamá started during the Eocene (Herrera et al., three new localities on the Pacific coast of Pana- 2012; Montes et al., 2015, 2012a, 2012b; Jaramil- má, near the town of Torio, province of Veraguas lo, 2016; O’Dea et al., 2016; Jaramillo et al., 2017); (Figure 1). The fossils were recovered both from thus, this region provides an excellent opportuni- Oligocene (1 locality) and Early Miocene (2 locali- ty to understand how the tropical vegetation of a ties) marginal marine deposits (figures 1 and 2). We newly created land evolved in relation to tectonic describe these specimens and augment the gener- and climatic changes over time (Jaramillo et al., ic diagnosis of Lacunofrucutus cuatrecasana Herrera, 2014; Jaramillo, 2016). Paleofloras of peninsular Manchester et Jaramillo, and present a new record Central America during the Cenozoic are import- of Sacoglottis sp. We also provide a sedimentological ant for reconstructing the exchange of plant spe- description and a biostratigraphic analysis based cies between Central America and South America on fossil mollusks, calcareous nannoplankton, and (Jaramillo et al., 2014; Bacon et al., 2015) and also palynology for each of these localities. for understanding paleoenvironmental changes (Jud and Dunham, 2017). 1.1. HUMIRIACEAE FAMILY Most paleobotanical studies about floristic compo- sition in Central America came from palynological Extant Humiriaceae includes eight genera of analyses (i.e., fossil pollen and spores) (Burnham mostly woody, evergreen plants (Cuatrecasas, and Graham, 1999; Jaramillo et al., 2014). Recent 1961; Kubitzki, 2014). Trees in this family range reports have shown that by the Early Miocene, from tall canopy trees to medium sized shrubs Panamanian forests were already dominated by (Kubitzki, 2014). Humiriaceae are typical mem- Gondwanan-Amazonian taxa, suggesting that bers of the rainforests, but are also found in sa- long-distance dispersal across the Central Amer- vannahs up to an elevation of 1400 m (Kubitzki, ica seaway occurred much earlier than previous- 2014). The geographic distribution of the family is ly suggested (Jaramillo et al., 2014; Bacon et al., predominantly Neotropical with ~50 species, with 2015). The palynological record is important for the exception of the species Sacoglottis gabonensis reconstructing the paleocommunities in Pana- (Baill.) Urb. that grows in the west African tropics má (Jaramillo et al., 2014); however, this record is (Herrera et al., 2010; Kubitzki, 2014). limited given the relatively low taxonomic resolu- Humiriaceae fossil endocarps and wood remains tion that the pollen offers for the identification of are some of the most common elements of the plants. Plant macrofossils (e.g., fruits, leaves, and Eocene to Pleistocene floras of Central Ameri- wood) in Central America are necessary to better ca (Herrera et al., 2010, 2012, 2014b; Lott et al., understand the dispersal and evolution of particu- 2011; Jud and Dunham, 2017), which show that lar plant groups. In recent years, excavations along this group has been present in the region since the the Panamá Canal, Lake Alalajuela and Azuero land started to emerge in the Eocene (Herrera et Peninsula have allowed the discovery of various al., 2012; Montes et al., 2012b). Eocene Panama- Cenozoic localities with abundant fruits, seeds, nian fossils (Herrera et al., 2012) are also the old- leaves, and wood from several families; for exam- est record of the Humiriaceae family, suggesting a Neotropical origin, but it is not clear whether New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene Boletín de la Sociedad Geológica Mexicana / 2018 / 225 the center of origin was Central America or South whether Humiriaceae had a single dispersal event, America. or there were multiple dispersal events across the Fossil endocarps, wood and pollen of Humiriaceae Central American Seaway between Central and are also relatively common in Eocene – Pliocene South America. Previous studies on other Neo- sedimentary rocks of northern South America tropical families have found evidence of diversi- INTRODUCTION (Berry, 1924; Wijninga, 1996a, 1996b; Manches- fication after long-distance dispersal events (e.g., ter et al., 2012). By the Early Miocene, this family Chrysobalanaceae; Bardon et al., 2013; Jud et al., was widely distributed in the Neotropics (Herrera 2016). et al., 2010, 2012, 2014b). However, it is not clear Figure 1 General map of Panamá and geological map of the area near the town of Torio. New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene 226 / Boletín de la Sociedad Geológica Mexicana / 2018 INTRODUCTION Figure 2 Stratigraphic columns of the studied sedimentary sequences in the Palo Seco River (A, Oligocene) and Torio Beach (B, Early Miocene) showing the fossil horizons and sample localities. 1.2. GEOLOGY OF THE AZUERO PENINSULA marine
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