Aura M. http://dx.doi.org/10.18268/BSGM2018v70n1a13 P. 223‒239 VOL. 70NO. 1 BOL. SOC.GEOL.MEX.2018 Colombia. GeológicaARES,Corporación Bogotá, BayonaGerman Angeles, CA90007,USA. LosAngelesCounty, Los Museumof History Austin Hendy Colombia. de losAndes, Carrera 1No. 18A-12,Bogotá, deGeociencias,Departamento Universidad Colombia. Puerto Colombia,Barranquilla, Geology, Universidad delNorte, Km.5Vía Camilo Montes Balboa, Ancón,Panamá. Smithsonian Tropical Research Institute, Damian Cárdenas Alejandro Machado Carlos Jaramillo Road, Glencoe, Illinois60022,USA. Chicago BotanicGarden, 1000LakeCook Fabiany Herrera Colombia. de losAndes, Carrera 1No. 18A-12,Bogotá, deGeociencias,Departamento Universidad Aura M.Cuervo-Gómez Daniel E.Góngora Balboa, Ancón,Panamá. Smithsonian Tropical Research Institute, versity, Syracuse, NewYork 13244,USA. EarthSciences, SyracuseUni of Department [email protected] Nicolas Pérez-Consuegra Nicolas Early MioceneofthewesternAzueroPeninsula,Panamá New recordsofHumiriaceaefossilfruitsfromtheOligoceneand Manuscript accepted: September 1,2017. Manuscript September accepted: Corrected manuscript received: August 25,2017. Manuscript received: October, 1,2016. Pérez-Consuegra Cuervo-Gómez , Austin , Daniel E. - Hendy Góngora u fPanama, mus of tus endocarps, Keywords: Central America, America. this familywasinCentral or South to establish whether the origin of Seaway. Further studiesare necessary the Central American final closure of tral and South America before the inbetweendispersed family was Cen tral American forests, and that this early Cen important constituent of and show that Humiriaceae was an bothtaxa distributiongraphical of results expand the temporal and geo a newrecord of Manchester of and augment the generic description deposits. We describenewspecimens cene (two localities) marginal marine cene (one locality) and Early Mio Fossilswere recovered fromOligo es inthe Azuero Peninsula, Panamá. from two sequenc newsedimentary Humiriaceae a description of tant and fossil taxa. Here, we provide both ex requires of the integration versity inCentral American forests the di the origin of Understanding ABSTRACT Lacunofructus cuatrecasana Lacunofructus , Neotropical, forests,Isth , Alejandro , Fabiany Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et Jaramillo, andpresent Machado Sacoglottis Herrera Lacunofruc Herrera, sp. Our , Damian . , Carlos ------Cárdenas Jaramillo ceae, secuencias de dos nuevas excavados este trabajo, describimos fósiles de estudios de plantas actuales y fósiles.grar En bosques de América Central, se necesita inte Para el origenentender de la diversidad delos RESUMEN Sacoglottis tropicales, IstmodePanamá, Lacunofructus fósiles,tral, endocarpos Palabras clave: AméricaCen Central oSurAmérica. el origen de Humiriaceae se dio en América más estudios para establecer con claridad si del Istmo de Panamá.cierre Se necesitan entre Centro y Sur América mucho antes del hubo dispersiones de especies de Humiriaceae presentadaLa evidencia también sugiere que de los primeros bosques en América Central. Humiriaceae fue unimportante constituyente taxones, yademás demuestran quela familia y temporal paratribución ambos geográfica los rangos expanden de dis registros nuevos glottis de registro no ypresentamos un nuevo localidades del Oligoceno y Miocenotempra Manchester Herrera, morfológica de especímenes yaumentamosladescripción no (dos localidades). Describimos nuevos no (unalocalidad) y del Mioceno tempra del Oligoce marinos-marginales depósitos Panamá. Los fósiles fueron encontrados en sedimentarias en la Península de Azuero, sp., para el Mioceno temprano. Estos , German , Camilo . Lacunofructus , bosques Neo , bosques et /2018 Bayona Montes Jaramillo para las

cuatrecasana , Saco 223 ------

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene ABSTRACT New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene INTRODUCTION 224 224 1. Introduction varied forest types that grow over a complex geo lott diversity inthe worldextant angiosperm (Barth the regionswith the highest Costa Rica) isoneof Central America (i.e.,Southern Panamá and leaves, and wood from several families; for exam Cenozoic localities with abundant fruits, seeds, Peninsula various have allowed the discovery of the Panamá Canal, Lake Alalajuela and Azuero lar groups. In recent years, excavations along particu andevolution the dispersal understand of wood) in Central America are to better necessary . Plant macrofossils( of identification the for offers pollen the that tion limited given the relatively low taxonomic resolu má (Jaramillo reconstructing the paleocommunities inPana palynological record2015). The isimportant for (Jaramillosuggested ly ica seaway occurred much earlier than previous acrosslong-distance dispersal the Central Amer that taxa, suggesting Gondwanan-Amazonian Panamanian forests were already dominated by reports have shown that bythe Early Miocene, Jaramillo1999; Graham, and analyses (i.e., fossilpollen and spores) (Burnham sition inCentral America came from palynological Most paleobotanical studies about floristic compo (Jud andDunham,2017). for paleoenvironmental understanding changes (Jaramillo cies between Central America and South America plant spe ant for reconstructing theexchange of Central America during the Cenozoic are import peninsular of Paleofloras 2016). Jaramillo, 2014; and climatic changes over time (Jaramillo newly created land evolved in relation to tectonic a howty tounderstand the tropical vegetation of thus, thisregion provides anexcellent opportuni lo, 2016;O’Dea 2012; Montes Panamá started duringthe Eocene (Herrera the Isthmusof logic landscape. emergence The of / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. , 1996). This plant diversity flourishes in flourishes diversity plant This 1996). , et al. et al. et al. , 2014; Bacon et al. , 2014);however, thisrecord is , 2015,2012a,2012b;Jaramil , 2016;Jaramillo et al. e.g. , 2014;Bacon et al. , fruits, leaves, and et al. , 2015) and also , 2014). Recent et al. , 2017); et al. et al. et al. ------/2018 , , , 1.1. HUMIRIACEAE FAMILY cdansso of diagnosis ic describe these specimens andaugment the gener ties) marginal marine deposits (figures 1 and 2). We Oligocene (1 locality) and Early Miocene (2 locali (Figure fossilswere 1).The recovered both from Torio, Veraguas provincemá, near the town of of Pana of coast Pacific the on localities new three Humiriaceae from andseedsof fossilendocarps We andcarbonized report newpermineralized et al. Jud ho (Herrera lophyllaceae, Connaraceae and Chrysobalanacea ple Humiriaceae, Juglandaceae, Malvaceae, Ca a Neotropical origin, but it is not clear whether the Humiriaceae family, est record suggesting of nian fossils(Herrera al. land started to emerge in the Eocene (Herrera this group hasbeen present in the region since the 2011; Jud2017), which andDunham, show that Ameri ca (Herrera Central of floras Pleistocene to Eocene the elements themostcommon of are someof and wood remainsHumiriaceae fossil endocarps (Herrera (Baill.) Urb. that growsin the west Africantropics the the exception of predominantly Neotropical with~50species, with the family is distribution geographic of 2014). The 1400m(Kubitzki, vannahs upto an elevation of the rainforests, but are also found insa of bers (Kubitzki, 2014). Humiriaceae are typical mem from tall canopytrees tomediumsized shrubs range family this in Trees 2014). Kubitzki, 1961; woody,mostly evergreen plants (Cuatrecasas, Extant Humiriaceae includes eight genera of theselocalities. for eachpalynology of on fossil mollusks, calcareousand nannoplankton, based analysis description andabiostratigraphic of Manchester , 2012;Montes Sacoglottis et al. et al. , 2017;NelsonandJud, 2017). , 2013; Rodríguez-Reyes , 2016;Jud andNelson,2017;MacFadden et al. et al. sp. We also provide a sedimentological et et al. , 2010;Kubitzki, 2014). , 2010,2012,2014a,2014b;Carval Jaramillo, andpresent a new record Lacunofrucutus cuatrecasana Lacunofrucutus , 2010,2012,2014b;Lott et al. et al. , 2012b).EocenePanama , 2012)are alsothe old Sacoglottis gabonensis et al. , 2014, 2017; Herrera, et al. et ------,

Figure 1 et al. was widelydistributed in the Neotropics(Herrera ter Manches 1996b; 1996a, Wijninga, (Berry,1924; South America northern rockssedimentary of are also relativelyin Eocene – Pliocene common Humiriaceae Fossil wood endocarps, andpollen of America. originwasCentral America orSouth the center of General map ofPanamáand geological mapof theareanear thetown ofTorio. et al. , 2010,2012,2014b). However, it is not clear , 2012).Bythe Early Miocene, thisfamily Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - fication after long-distance dispersal events ( events dispersal long-distance after fication diversi tropical families have found evidence of South America. Previous studies on other Neo Central American Seaway between Central and or there were multiple events dispersal across the event,whether dispersal Humiriaceae had asingle 2016). BardonChrysobalanaceae; et al. , 2013;Jud /2018 et al. 225 225 e.g. - - , ,

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene INTRODUCTION New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene INTRODUCTION 226 226 Miocene) showing the fossil horizons and sample localities. 2 Figure 1.2. GEOLOGY OFTHEAZUERO PENINSULA The forearcThe sediments includeturbidites, shallow overlaid by forearc sediments(Buchs been described as adiverse igneousbasement the Azuero Peninsula has of general geology The / Stratigraphic columns ofthestudiedsedimentarysequences inthePaloSecoRiver(A,Oligocene) and Torio Beach(B,Early Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. , 2011). /2018 larsky larsky Covachón, Tonosi (Ko andSantiago formations – Miocene and have been subdivided into the clastic faciesthat were inthe Eocene deposited silici influenced fluvially and limestone, marine et al. , 1995;Buchs et al. , 2011). - - 2. MaterialsandMethods (al Moee; TI 946 7550 N, 7.54500 990406: STRI Miocene); (Early W -80.95070 N, 7.54510 640015: localities (STRI) Panamá; SmithsonianTropical Research Institute of town the the Azuero Peninsula,Torio near part of inthe western found were floras fossil The group intheTorio Beach outcrops (Figure 1). overlaysthe Azuero proto arc (volcanic basement) 1), and(2)anEarlyunit that Miocene sedimentary Torio (Figure tributaryand extends tothe north of – Paleogene in the Palo Ocú formation Seco River overliesunit that unconformably the Campanian units:(1) an Oligocene sedimentary sedimentary different two identify to us allow Torio of town the the of vicinities cent geological mappingof calcareous nannofossils(Kolarsky been dated as Eocene – Early Miocene based on been assigned to the Tonosi which formation, has Azueronorthwestern Peninsula (near Torio) have sequences that sedimentary cropThe outinthe Specimens were thenwashedwithwater and sub seconds. five for HCl (5%) diluted in etched then were (600) and carbide grit silicon polishedusing (Joy fossilswere prepared using standard techniques the 1200C stereoscope (Figure 3). Acetate peels of Nikon D90 digital camera and with a Nikon DXM saw. specimens were with a The photographed and apex, withalow speed,thin-bladed diamond fossils, sections were cut through the base, equator, Toreveal the anatomy and describethe internal of illosdb/web/login). PaleoDatabase (http://biogeodb.stri.si.edu/jaram locality andsamplescanbeaccessedat the STRI onthe Natural(FLMNH). Information History Museum of specimens are housed at the Florida the Palo Seco River (figures 1 andThe 2). fossil of third localitycropsout inasmalltributary creek the and 2), and 1 figures Beach;(Torio Torio of two localities crop out at the beach near the town first The (Oligocene). W -80.93310 N, 7.63920 -80.94980 W (Early Miocene); and STRI 990413: et al. , 1956; Herrera 1956; , et al. , 2012).Specimens et al. , 1995). Re1995). , Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - 2.2. MICROPALEONTOLOGICAL ANALYSES 2.1. STRATIGRAPHY ANDSEDIMENTOLOGY de e emnlg o h ecito fthe for thedescriptionof added newterminology 2012, 2014b;Kubitzki, 2014);however, we also at the SCZ, for anatomical comparisons: Humiriaceae from Panamanian forests, available fossilembryos. We selected three extant species of the family (Cuatrecasas, 1961; Herrera ing topreviousnomenclature of anddescriptions Humiriaceae were selected accord characters of Tropicalsonian Research Institute (SCZ). Fruit the Smith species housedat the herbarium of We compared the fossilswith fruits from extant balsam mountingmedium. microscopemounted in slides withCanada glass ally to generate the acetate peels. peels were The with airandthenthe acetate removedwas manu placed over the surface.were specimens The dried cellulose acetate was with acetone and aportion of etchedsequently dried. The covered surfacewas on glass slides.on glass Subsequently, the sediment was were extracted from each sample and mounted sediment nannoplankton analysis, of two grams the three fossillocalities (Figure 2). For calcareous collected was fromanalyses eachpalynology of One sample for calcareous nannoplanktonand were found (Figure 2). (Figure 2), to include the strata where the fossils Earlyunits Miocene andOligocene sedimentary the both of base the at staff, Jacob’s a with scale We measured twoat sections stratigraphic a1:50 hypocotyl. cotyledons, and the endocarp, endosperm, of structures cellular specific of identification the for microscope fluorescence E600 research Nikon a microscope (10x,20xand40x),stereoscope, and and the sections weremens observed underalight (Figure fossil 4). The andextant speci endocarps foras thefossilthe samemethodology using carps endo themodern tion, we examinedof sections Cuatrec. and ovicarpa Cuatrec., depleta diguense Humiriastrum McPherson. Inaddi McPherson. /2018 (Cuatrec.) et al. Sacoglottis , 2010, 227 227 ------

INTRODUCTION / MATERIALS AND New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene METHODS New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene MATERIALS AND METHODS 228 228 Scale bar:A–G, I: 5mm;H,J:1mm. ofL, endocarp Thinsection showing abundant lacunaeand twowell-developed seeds, oneindicated bythearrow. K, Lateral viewofendocarp, thearrowshows afurrowcorresponds that totheexterior lingulategermination valve. K –L, peel of1-I showingJ, Acetate abundant lacunae(red arrow) onendocarp wall(i.e.,septae and germination valves). aborted seeds (arrow)andwhich have 2seed loculesfilled withsediment(arrow). ofI, medial Transverse section of part theendocarp showing twoseeds,fiveseptae, and (arrow),threeof fivegermination valves I – J,SpecimenFMNH 68986–STRI40034 (Early Miocene). andH, Detailofgermination valve seed of1-G.Lacunae areclearly visible intheendocarparrow). (white ofG, Transverseof medial section theendocarp part showing twoseeds and twogermination valves. G –H,SpecimenFMNH 68987-STRI40033 (Early Miocene). arrow). (arrow A),four septae (arrow B),and four (arrow germination C).Lacunae valves (cavities)are visiblewithin theendocarp (lower red F, SpecimenFMNH68981-STRI40052 (Early Miocene).Transversesectionoftheupper half of theendocarp, showing four seed locules bundle (arrowA),threeseeds (arrowB),threeseptae, and three germinationvalves. ofE, SpecimenFMNH68982-STRI40060(Earlyof medial Miocene). Transversesection theendocarp part showing centralvascular (arrow B).Thevariationinendocarpvalves shapewithinspecimensdeformation. iscaused bydiagenetic D, SpecimenFMNH 68984-STRI40032 (Oligocene).Dorsalviewof theendocarp showing germination septae (arrowA)and germination C, Lateral viewof theendocarp showing (arrow). germination valves B, Apicalviewoftheendocarp showing septae (arrowA)and (arrowB). germinationvalves A, Dorsalviewof theendocarp showing (arrow). germination valves A –C,SpecimenFMNH68983-STRI38784 (Oligocene). Figure 3 Lacunofructus cuatrecasana / A–H, Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Lacunofructus cuatrecasana fromtheeasternAzuero Peninsula,Tonosi Fm(Herrera fromthewesternAzuero Peninsula. /2018 et al. , 2012). 3.1. STRATIGRAPHY ANDSEDIMENTOLOGY 3. Results h tairpi euneo the Palo Seco River sequence of stratigraphic The Paleoflora, Colombia). Inc.(Bucaramanga, ical procedure (Traverse, 2007) and were done at were prepared following the standard palynolog 2012). For palynological analysis, three samples the Geological Time Scale 2012 (Gradstein and 1971), Worsley(Martini, and Martini of tion Bolli used the parameters proposed by Bown (1998) and ranges, the stratigraphic we of the determination and nannoplankton calcareous of identification polarized light microscope. For the taxonomic slides wereing medium.The observed under a Canada balsam mount the slide with the aid of hot platecover (~40C). A glass mounted on was ally 2–3drops, thenthe slides were dried on a usu processing, for sufficient water distilled and aneedle with the aid of the glass crushed against fragments (dugong), fish teeth, and ex-situ corals. ex-situ and teeth, fish (dugong), fragments fossiliferouslayer vertebrate also contains bone This 990413). (STRI locality endocarp fossil the contains that 9) – 7.5 (meter stratum glomeratic sandstones. layers These are overlayed by a con transitions from thebasal breccia toconglomeratic composition ( a massive breccia at the base with polymictic of measuresposits 48 m(Figure composed 2B).Itis the Torio Beach de section of stratigraphic The from 23.5to26.5m(Figure 2A). the fossilswere excavated from asandstonelayer however,contained cession fossil endocarps; all Several horizonsthe Palowithin SecoRiver suc foraminifera. and wood) but stillwithtracesof more fluvial transported material ( sil material ( careous sublitharenites with abundant marine fos cal sequence transitions to intercalationsThe of igneous rocks)breccia nearthe base(Figure 2A). massive polymictic( a of composed is It 2A). (Figure m 29 measures et al. 18) W flo te egn zona Neogene the follow We (1989). e.g. e.g. , mollusks) to sublitharenites, mollusks) with , chert sequence and basalt). The e.g. , chert, basalt, and other e.g. , leaves, seeds et al. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ------, 3.2. AGE OFTHEDEPOSITS wood. include mollusks The mollusks, crab fragments, and cirripedia, fossilsof horizontal burrows(STRI 640015) andnumerous sandy conglomerates with layer2 –4.5)of (meters stratum isoverlainThis by amassive-bioturbated euneo ~8mo predominantly clasticstra ~38 m of sequence of these fossiliferous levels, a fault separates a second sis Naticidae, Muricidae, of raninidcrabs,echinoderms, andgastropod fossils ing abundant contain 7.5) – 5.5 meters 290950; (STRI stones are calcareous beds andfossiliferous algal lime cf. rea Torio Beachcontained several nannofossilsin Samples from localitySTRI 640015 from the cene agetothisunit. sequence (Figureimentary 1) we assign anOligo sed this of top the on tuff a Kedenburg,of 2016) 25.1 ±0.18Maage (zircon U-Pb geochronology; et al. sample must be younger than 33.76 Ma(Williams datum of atriletes saccolomoides and the spores ramosus sphaera alcicornu characterizedbyis dinoflagellate cysts the assemblage palynological dinoflagellateThe cysts. pollen and spore grains, and low abundancesof terrestrial organic matter,phytoclasts mainly with fossils. However, good recoverysample has this of collected fromthe Palo SecoRiver didnotcontainnanno 990413 STRI locality from sample A conglomerates. sandstones,to medium-grained mudstones and fine- are theretop, the At beds. mudstones minor sandstone and to verywith coarse- coarse-grained interbedded sandstone medium-grained to fine- a massive conglomerate bed, and comprised of are mudstones. meters intermediate The posed of thissequence is com ta (Figure base of 2). The , sp., F. carbasea Conus , 2004). Based on the palynology andonthe , 2004).Based on the palynology , the pollen grain , the pollen grain Ampullinopsis spenceri s. n urde(iue2) ntpo sp., andTurridae (Figure 2B). Ontopof A. alcicornu ntpo thisbasalsequence, there . Ontop of Concavissimisporites fossulatus Thalassinoides , Polysphaeridium zoharyi . Based on the first appearance first the on Based . inequatorial latitudes this Antillophos , Perisyncolporites pokornyi burrows, spatangoid Galeodea Leopecten /2018 aff. sp., and sp., , A. gatunen Spiniferites and Crassost Achomo Ficus 229 229 Stri ------,

MATERIALS AND METHODS / New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS 230 230 with samples cannotbe older than NN4,andtogether bilicus floridanus lithus Torio Beach contained the nannofossils: locality The STRI sample 990406 of also from the cluding F. carbasea sp., 640015) — and ties 290950 associatedMollusks with thissection (STRI locali (2011). Jaramillo and the palynological zonation of South America (21.5Ma;Martínez palynological recordsfrom Central (10.2 Ma)and millo stra, 2000;daSilva-Caminha, 1985; Van1994; Hoorn, Hooghiem and Hammen Der (Graham, Neotropics the in record fossil Miocene, according toitswidespread Neogene both samples isyounger than Early that the age of presence common of The Concavissimisporites fossulatus Echinatisporis muelleri tus Fern spores include microclavatusClavainaperturites pokornyi ae), (Euphorbiaceae?), nacimientoensis iferites ramosus Spiniferites mirabilis Achomosphaera alcicornu cysts dinoflagellate the Taxainclude cysts. gellate dinofla gonyaulacoid abundant relatively a with phytoclasts and sporomorphs, together posed of ter predominantlyis terrestrial com and mainly well-preserved Organic mat palynomorphs. of and localitySTRI 640015yielded a goodrecovery 990406 STRI locality from Palynologicalsamples toTortonian).galian late EarlyMiocene toearly Late Miocene (Burdi sequence is betweensedimentary NN4 to NN7, the Torio Beach younger age of than NN7.The / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (Pteridaceae), Ampullinopsis spenceri Lanagiopollis crassa Lanagiopollis C. floridanus C. ae ntepeec f. Based on the presence of et al. (Malpighiaceae) and the less abundant Reticulofenestra minuta adgsrpdfsiso Naticidae, Mu andgastropod fossilsof , 2011;Martínez (Bombacoidea), . Angiosperms include . Angiosperms , Discoaster deflandrei Discoaster and , Polypodiaceoisporites pseudopsilatus Polysphaeridium zoharyi Echiperiporites estelae , D.deflandrei , Striatriletes saccolomoides (Pelliciera), Polypodiaceoisporites? fossula Operculodonium centrocarpum , Galeodea . C. microclavatus and et al. Leopecten rttiopts finitus Crototricolpites (Hedyosmum). et al. the sample is not thesampleis , 2013),itsoldest and R. pseudoumbilicus sp. and R. pseudoumbilicus Perisyncolporites , 2010;Jara sp., Bombacacidites et al. R. pseudoum (Malvace and Crassostrea Cyclicargo indicates Ficus , 2013), and Spin et al. cf. ------/2018 . , ,

3.3. SYSTEMATICS cunofructus the fossilgenus Here we present a revisionof ure 1). sequence in the Toriothe sedimentary Beach (Fig wean Early assign Mioceneage (Burdigalian?) to (Aquitanian – Burdigalian) age. Based on the data, indicates the succession an Early Miocene of presence of Hendy,and Chucunaque formations; 2013). The strata from Panamá ( Turridae — are similar to other faunas in Miocene ricidae, transverse section (figures 3F, 3I and 3J). Valves and 3I 3F, (figures section transverse enlarging Septa outward,ly arranged. as seen in radial five, to three valves and Septa elliptic. to roundedApex toelliptic (Figure 3B),baserounded (n = 12).Surface width 16.3to8.1mm smooth. and 3K).Length 8.3to21.6mm, 3C, 3D 3A, Description. Manchester Type species. outline. isodiametric andhexagonalto pentagonal cells in approximately line. hypocotyl The of composed is approximately isodiametric and hexagonal in out anticlinal cells, composedof yledons. Endosperm interlocked andelongated sclereids. Twoof cot clusters are wall and septa composed of docarp cavities abundant within valves en The or septa. Valves conspicuous. Seeds one per locule. Small with threehexa-carpellate valvesto six andsepta. Emended generic diagnosis. (2012). the original material described by Herrera andseed that was not present in the endocarp of localities, allows usto observe the cellular structure fossils, from the Oligocene and the Early Miocene Lacunofructus Antillophos h xust rsraino the new exquisite. The preservation of Family HumiriaceaeJussieu et Ampullinopsis spenceri Jaramillo (Figures 3Ato3J). Endocarp elliptic to ovate (figures ovate to elliptic Endocarp millo, here emended.

Lacunofructus cuatrecasana Lacunofructus aff. Herrera, Manchester e.g. A. gatunensis , Culebra, Galique, Gatun inthe lower part Endocarp tri to Endocarp , Conus Herrera, sp., and et Jara et al. La ------

Manchester peel of E, Acetate D, Detailof4-C( showingC, Thinsection endocarp wallcells. C –D,Extant peel of theendocarp;B, Acetate note elongated sclereids cellsinlongitudinalforming section atwistednetwork. A, Acetatepeelof theendocarp; noteelongated cells. Figure 4 Scale bar:100µm. F, Extant A–B, Humiriastrum diguense et al. Vantanea depleta Lacunofructus cuatrecasana Vantanea depleta (2012). Vantanea depleta . Specimen SCZ-15737. . SpecimenSCZ-15737. . Specimen SCZ-15724. Thin section oftheendocarp Thinsection showing. SpecimenSCZ-15724. elongated cells. ). fromtheOligoceneofPerú. Elongateand twistedfibers forming amessynetwork.Imagefrom fromthewesternAzuero Peninsula.SpecimenFMNH 68987-STRI40033(Early Miocene). Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín /2018 231 231

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS 232 232 section (figures 3E, 3F and 3G). without Endocarp imen; pentagonal to ovate as viewed intransverse Seeds one per locule. Up to four locules per spec (Figure 3B). the endocarp the apex orthe baseof running lengthwise, but without reaching fullyto 3D), and 3C 3A, (figures elliptic to lingulate face, conspicuous, withoutamedian furrowon the sur sponds to the sponds Oligocene. Rocksbelong to an un Age andformation. 990413: 7.63920N, -80.93310W). W; STRI 990406: 7.54500 N, -80.94980 W; STRI ro Peninsula (STRI 640015: 7.54510 N, -80.95070 New occurrence. STRI 40033[Early Miocene]). 68987- FMNH and 68982- Miocene] [Early STRI40060 (FMNH permineralizations [Oligocene]) calcite 40047 and 68985-STRI FMNH and [Oligocene], 40032 68984-STRI FMNH cene], [Oligo 38784 40052[Early 68983-STRI FMNH 68981-STRI Miocene], FMNH [Early 40034 Miocene], 68986-STRI FMNH Miocene], [Early 38782 68988-STRI (FMNH bonizations Specimens. cotyledons (figures 5A,5Band5D). a highembryo-to-seed ratio (E:S) and thin,leafy fossilThe seedsof pentagonal cells that vary from 0.017to 0.037 µm. approximately isodiametric hexagonal to posed of section (figures 5B and 5C).The hypocotyl is com radiating from the cotyledons orhypocotyl incross cellsare lines in arranged 5B and5C).Endosperm (figures diameter in µm 0.070 to 0.029 from vary mately isodiametric and hexagonal in outline, that anticlinal cells, approxi composed of Endosperm like texture (Figure 4). Two cotyledons(Figure 5A). very high density, a woody- givingthe endocarps interlocked and elongate sclereids with of clusters (fig width cellsare in arranged uresand 4B).Endocarp 4A in µm 13.2 to 5.4 sclereids elongate are wall and septa composed of endocarp The (Figure 3E). theendocarp tral axisof (figures 3H, 3J and 3L). Vascular strand at the cen ranging fromand septa, in diameter 0.1to0.4mm ities present wall andabundant inthe endocarp apical foramina. Small ellipsoidal to circular cav / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín These are These preserved asbothcar Location: Panamá, west Azue Lacunofructus cuatrecasana Lacunofructus oaiy 943 corre 990413 Locality show ------/2018 eso mens of Peninsula (Herrera the Tonosi Formationthe Eocene of in the Azuero Lacunofructus Humiriaceae. characters diagnosticof of cuatrecasana Lacunofructus characters. of set this has plants flowering among tion valves (Herrera recognizable bytheir distinctive germina dorsal pels, one or two seeds per locule, and are easily withacentral vascular axis,docarps upto 10 car Affinity. nosí Fm.(?) ToEarlyMiocene the to correspond 990406 and Localities 640015 described geological formation. lottis showsthe closest similarity tolivinggenus affinity. Systematic 30 µminwidth(Figure 6B). a woody-like textureendocarps and measure 7 – a twisted and interlocked network,form givingthe the endocarp cells of The not be determined. seedsperloculecould endocarp, orthenumber of the valves, the shape of suchters asthe number of charac Specific mm. 4.7 to from1 ranges ameter 40059 appears to be an isolated valve. Lacunae di seed cavity (Figure 6A), while FMNH 68980-STRI showing two lateralendocarp valves andacentral an of section longitudinal a is 38790 68979-STRI FMNH fossils The 40059. 68980-STRI FMNH Description. have wallcavities endocarp (Herrera between these two genera is that difference major anatomy. The seed and fruit of miriaceae, 4E) (Manchester cipaconensis have the endocarp been reported for of diguense Sacoglottis ovicarpa this study (i.e., ture and anatomy to the extant species analyzed in pce f. Species of [Figure 4F]). The same anatomical details anatomical same The 4F]). [Figure Lacunofructus Lacunofructus Fut fHumiriaceae have woody en Fruits of Lacunofructus rmteOioeeo Perú (Figure from the Oligocene of is a fossil genus first described from described first genus fossil a is MH 87-TI 89 and 38790 68979-STRI FMNH Vantanea depleta Sacoglottis cf. et al. Fgr 6] and 6D], [Figure et al. Sacoglottis et al. are similar in endocarp struc are similar in endocarp , 2012). Among extant, 2012).Among Hu resembles preserves the combination The newfossilThe material , 2012). The new speci , 2012). The are recognized by abun , 2010).Noother family sp. [figures 4C and 4D], Vantaneain Vantanea et al. Humiriastrum Humiriastrum , 2010). Vantanea do not terms terms Sacog ------Scale bar:A,B,D,F:0.3mm;C,E: 0.1mm. showing abundant endosperm tissue(arrowA)and circularhypocotyl(arrowB). F, Extant peel showingE, Acetate ovalhypocotyl,hypocotylcells(arrowA),and endosperm cells(arrowB). of thelowerD, Transversalsection of aseed half showing part theovalhypocotyl. C, Detailof theendosperm cellsof theseed,and anticlinalcells,approximatelyisodiametric hexagonal in outline. of thelowerB, Transversalsection of theseed half showing part thecircularhypocotyl(arrowA)and endosperm tissue(arrowB). endosperm tissue(arrowC). of theupperA, Transversesection halfof part aseed showing twocotyledons(arrowA)divided byamidline(arrow B)and abundant A –D,SpecimenFMNH 68982–STRI40060-11 (Early Miocene). Figure 5 A–E,Epifluorescence imagesof seeds of Sacoglottis ovicarpa . Specimen SCZ-1296. Epifluorescence. SpecimenSCZ-1296. of imageof thelower transversalsection halfof theseed, part Lacunofructus cuatrecasana Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín . /2018 233 233

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS 234 234 Scale bar:A,C:1cm; B,D:10µm. of theendocarpD, Thinsection showing elongated sclereids inamessy and twistednetwork. seed locule(lowerblackarrow). of endocarpC, Transversesection showing (whitearrow),septae(blackabundantvalves five lacunae(lower whitearrow),and C –D, giving theendocarps awoody-like texture. peel of endocarpB, Acetate showing cellular structure. Abundant sclereids arevisible and form atwistedand interlocked network, ofA, Longitudinal endocarp section showing (whitearrows)and twovalves abundant lacunae(black arrow). Figure 6 in shape. valves The areand the inconspicuous 4 valves, whichfrom vary long, elliptical to broad contain only 3 or 5, but sometimes the endocarps valves in number of (i.e., gy ovoid, globose, oblong andelliptic). The wall (Figure 6C) and variable drupe morpholo endocarp the in cavities oil-filled or lacunae dant 40052, andFMNH68982-STRI40060. FMNH 68987-STRI 40033, FMNH 68981-STRI 40034, 68986-STRI FMNH 40032, 68984-STRI Specimens. narrowsepta outwards (Figure 6C). Sacoglottis ovicarpa / A–B,c.f. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Sacoglottis FMNH 68983-STRI 38784, FMNH from Panamá, specimen SCZ-1296 ofHerrera fromPanamá,specimen SCZ-1296 , specimenFMNH 68979-STRI38790 (Early Miocene). Sacoglottis endocarps istypically - /2018 3.4. EXTANT FRUITS et al. ) h rnvredaee fthese cells varies in transverse6). The diameter of 4 and (figures tracts fiber and sclereids of clusters walls are mostly composedof that the endocarp diguense extant anatomicalThe studyof Earlyand formation: MioceneTonosí Fm.? Age W). N,-80.94980 7.54500 W;990406: STRI ro Peninsula (STRI 640015: 7.54510 N, -80.95070 New occurrence. (2010). , Vantanea depleta Location: Panamá, west Azue and Sacoglottis ovicarpa Humiriastrum shows -

4.2. PALEOBIOGEOGRAPHICAL IMPLICATIONS 4.1. SEDIMENTOLOGY 4. Discussion an important clastic input from a continental flu continental a from input clastic important an were in a shallow-marine deposited setting with Paloin SecoRiver that suggest thesesediments thesequencemeasured faciesassociationsof The continentalfluvialsystems. contributions of a shallow-marinesuggest reefal environment with logical ( analysis from the palyno and continental palynomorphs fossilseeds, wood posits,well as the presence as of rowsand talassinoidsfound inthe Torio Beach de corals, shells, echinoids, bur reefs, fragments of from continental terrains. Furthermore, algal basal conglomerate, important input suggesting Seco River (Oligocene) sequences start with a Both theTorio Beach(Early Miocene)andPalo and in in diguense Humiriastrum muchmore than from common NorthAmeri South America into Central America were likely that eventstic compositionsuggests dispersal from natedfloris by This taxa. Gondwana-Amazonian Ma), the Panamanian forests were already domi shownthat atthe Early least since Miocene (~20 Paleobotanical and palynologicalstudies have (Tonosí Fm;Herrera environments Azuero Peninsula inthe southern recordedphases shallow in marine totransitional gocene, which contrastswithearlier exhumation Azuero the western Peninsula atthe Oli least since in terranes the exhumationlocalities suggests of these vironments. Overall, of the sedimentology contributiona suggest from en fluvial continental vironment, and the fossil seeds, leaves, and wood sublitharenitesous support ashallow-marine en foraminifera the calcarein crabs,traces and of fossilmollusks, vial environment. presence The of Vantanea depleta Sacoglottis ovicarpa e.g. from(Figure7.4 –40.1µm 4), , phytoclasts andsporomorphs), from 5.4 – 14.9 µm (Figure 4), et al. from 7–30 µm (Figure 6). , 2012). Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ------(Herrera the family Humiriaceae laas the oldest record of Azuero Peninsu thesouthern the Late Eocene of sil genus. the fos range of temporal and paleogeographical new recordsThe of Azuero Peninsula. the Oligocene and Early Miocene on the western at landscapes least since colonizedformed newly fossillocalities, indicate that Humiriaceae rapidly the study,this of together with thesedimentology Humiriaceae fossilspresentedAmerica. The in known whether the origin was inCentral or South originated in the Neotropics;however, it is still un ca (Jaramillo continental organic matter ( sitional environments with animportant input of glottis of endocarps ied localities (Figure that 2) suggests the fossil the stud and stratigraphy sedimentology of The al. (Lott Rico Puerto of Pleistocene the and 1990), Bogota (Wijninga and Kuhry,the High Plain of al. Cucaracha Fm.inthe Panamá Canal (Herrera Sacoglottis (Herrera Puerto Rico inCentral America the Oligocene of genus this fromcomes oldestrecord2014). The of the Central America Seaway (Jaramillo closure of final the before long place took regions gests that events plant dispersal in between these South America, which sug America and northern several Central other localities from bothsouthern can tropical forests. Central Ameri genus wasanearly component of the genus inCentral America, showing that this Peninsula range of alsoexpand the geographical Azuero the western sp. from the Early Mioceneof cf. as work this in identified fossils The America at leastuntiltheEarly Miocene. ities indicates thatCentral thegenus in persisted occurrence of The likely originatedancestral in Central America. , 2011). , 2010),Middle–Late Miocene toPliocene in sp. were in shallow deposited marine to tran fossilshave alsobeenstudiedfromthe et al. et al. Lacunofructus et al. , 2012), suggesting that, 2012), suggesting Lacunofructus cuatrecasana Lacunofructus , 2014b).Early –Middle Miocene , 2014).Humiriaceae probably Lacunofructus Sacoglottis Lacunofructus was initially describedfrom was e.g. fossilsare in common , seeds, wood, phy inthese new local also expand the /2018 and cf. Lacunofructus Sacoglottis et al. Saco 235 235 et et ------,

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene RESULTS / DISCUSSION DISCUSSION / CONCLUSIONS / New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene AKNOWLEDGEMENTS / REFERENCES 236 236 5. Conclusions ancy and travelancy riversin longdistances and even 1961), Humiriaceae to have high buoy (Cuatrecasas,allows fruits of species some of endocarp the in cavities oil the as well as 6), and 4 (figures a mesh elongated sclereids forming of clusters of withthickwalls,and woody composed endocarps toclasts, observed The andsporomorphs). hard during theOligoceneand Early Miocene. terranes formed newly in systems fluvial active of transitional environments, indicating the presence cord. Fossilswhere inshallow-marine deposited to have aided in their preservation in the fossil re the family, representativesmodern which of could studied havesils similarhistologicalcharactersas and South America duringthe Miocene. fos The sp.not clear, is but across itdispersed both Central cf. until the Early Miocene. origin of The forests during the Eocene and waspresent at least atrecasana the Oligocene and Early Miocene. that during colonized the landscape that formed the early vegetation ro Peninsula arecord of form from Azue fossil new thewestern endocarps The forests by theEocene–Miocene. Panamanian alreadyan important constituent of that2017) suggests theseplantswereDunham, sil wood from the Oligocene – Miocene (Jud and in other Eocene site (Herrera However, Humiriaceae fossilfruits thepresence of ero Peninsula duringtheOligocene – Miocene. Azu the of flora autochthonous an represent not site fromdepositional far away sources andmay the fossils studied here were transported into the Miocene. We note that it is possible that some of Central America duringthe Oligocene and Early landscapesof acrossin the dispersal the forming acteristics couldhave aided Humiriaceae fruits char These 1976). Dennis, and (Gunn sea the in / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín probably originated in Central American et al. , 2012),andfos Lacunofructus cu Lacunofructus Sacoglottis ------/2018

Acknowledgements References provided access to the laboratory of fluorescence of laboratory the to access provided regardingcussions the fossil fruits. Jorge Ceballos Martínez, J. Wright, and C. Zalamea helped in dis extant fruit sections. S.R. Manchester, N. Jud, C. cal discussions. E.Moreno helped the in making geologi in and field the in help their for Monje 2014 – 2015 Azuero Field Camp crew and C. the to thank like would We field. the in support logistical and financial provided also STRI and Universidad de los Andes, Florida, University of project. this PCP-PIRE, fundedpart of suegra Short Term Fellowshipto Nicolas Pérez-Con — Institute Research Tropical Smithsonian The Bardon, L.,Chamagne, J., Dexter, K.G., Sothers, Bacon, C.D., Silvestro, D., Jaramillo, C.,Smith, thisarticle. that improved thequalityof one anonymous reviewer for insightfulcomments Cadena and Antoni Camprubí, as well as Edwin for palynology. We thankIdael Blanco andEditor samples Colombia) forcaramanga, processing of (Bu GustavoRueda Inc. Paleoflora and from ton digitalizing the geological map. We alsothankMil for the STRI-STF.acknowledged C.Ortiz is for the proposal Muller Landau for their revisions of baria. Nicolas PerezA. Cardenas thanks andH. to the materialaccess giving fromthe STRIHer microscopy. C.Galdamei was especially in kind 171, 19–37. the Linnean Society,Botanical Journal of plantsinthe Neotropics: into the evolution of Insights Chrysobalanaceae: and evolution of C.A., Prance, G.T., Chave, J., 2013,Origin Sciences, 112,6110–6115. Academy of theNational Panamá: Proceedings of of the Isthmus and complex emergence of Biological evidence supports anearly B.T., Chakrabarty, P., Antonelli,2015, A., ------on PR, 98 Calcareous Buchs D.M., P.O., Baumgartner Baumgartner- 1998, P.R., Bown, Bolli, H.M., Saunders, J.B., Perch-Nielson, K., Eastern the from FruitsFossil Berry,E.W., 1924, 1996, A., Placke, W., Kauer, W., Barthlott, da Silva-Caminha, S.A.F., Jaramillo, C.A., Revision Taxonomic A 1961, J., Cuatrecasas, Carvalho, M., Londoño, L., Wood, A., Cardenas, Burnham, ,R.J., Graham, A., 1999, The History of Botanical Club, 51,61. theTorrey Colombia: Bulletin of of Andes phytodiversity: Erdkunde, 50,317–327. vascular plants:towardsa world mapof species diversity in Global distribution of by .. 00 egn ayooyo Absy, of M.L.,2010, Neogene palynology 25–214. the United States National Herbarium, 35, theHumiriaceae: Contributionsfrom of America,186–14. of Sociery GSA: Denver, Colorado, Geological The Meeting in Denver: of 125th Anniversary miocene climatic optimum, inGSAAnnual forest ecophysiologyduring themiddle leaf Mummified assemblage provides insightto neotropical 2013, D.L., Royer, C., A., Escobar, J., Herrera, F., Jaramillo, Garden, 86,546–589. Botanical the Missouri and Status: Annalsof Neotropical Vegetation: NewDevelopments 31–46. American margin: Tectonophysics, 512, and subductionerosionalong the Middle (Panamá) and the discontinuous accretion the Azuero area tectonostratigraphy of 2011, Upper to Miocene K.,BandiniA.N.,Mora C., Flores Publishers, 315p. Series, ChapmanandHall, Kluwer Academic Micropalaeontological Society Publication NannofossilBiostratigraphy: British Calpionellids: CUPArchive. Planktic Foraminifera, Calcareous Nannofossilsand 1. Volume 1989, J., Remane, F., H., Jenkins, D.G., Laccarino, S., Rogl, Caron, M.,Toumarkine, M.,Luterbacher, Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Hendy, A.J.W., 2013,Spatial and Stratigraphic to guide World J.V., 1976, Dennis, C.R., Gunn, neotropical in Studies 1985, A., Graham, Gradstein, F.M., Ogg, J.G., Schmitz, M.D., Ogg, Herrera, F., Manchester, S.R., Vélez-Juarbe, Herrera, F., Manchester, S.R., Koll, R., Herrera, F., Manchester, S.R., Jaramillo, C., Herrera, F., Manchester, S.R., Jaramillo, C.,2012, aito fMarine PaleoenvironmentsVariation of 240 p. ​ Quadrangle/ tropical drift seeds andfruits:NewYork, Garden, 72,504–534. the Botanical Missouri Panamá: Annals of of paleobotany. IV. Eocene communities The Boston, Elsevier, 1176p. G.M. Geologic Time Scale: (eds), 2012, The Palaeontographica, 283,1–67. the Solimões Basin,Brazilian Amazonia: 828–840. Plant Sciences, 175, JournalInternational of the Humiriaceae (Part 2): of History J., Jaramillo, C.,2014b, Phytogeographic 128, 124–133. botany from theBotanical Missouri Garden, 80th year:in his in systematic Monographs biogeography:A Festschrift for AlanGraham Raven, P.H.(eds.), Paleobotanyand Panamá,Stevens, in W.D., Montiel, O.M., (Juglandaceae) the earlyin Mioceneof OreomunneaJaramillo, C., 2014a, Fruits of PlantSciences, 171,392–408. Journal of the Humiriaceae: International Phylogeny of S.A., 2010, Phytogeographic and History Silva‐Caminha, da B., MacFadden, and Palynology, 175,10–24. Palaeobotany Central America: Review of forests established early inthe uplift of of the composition Panamá give clues of of Permineralized fruits from the late Eocene 210–227. Panamá: Palaios,Formation, 28, Isthmus of in the Middle-Upper Miocene Gatun New York Times Book Co., /2018 237 237

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene REFERENCES New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene REFERENCES 238 238 aail,C,21,Eouino teItmso the Isthmusof Jaramillo, C.,2016,Evolution of Hoorn, C., 1994, An environmental reconstruction Jud, N.A.,Nelson, C.W., Herrera, F., 2016, Jud, N.A., Nelson, C.W., 2017, A from the Jud, N.A., Dunham, J.I., 2017,Fossil woods from Joy,K.W., Willis, A.J., Lacey, W.S., Rapid A 1956, Jaramillo C., Rueda, M., Torres, V., 2011,A Jaramillo, C.,Moreno, E.,Ramirez, V., da Jaramillo, C., Montes, C., Cardona, A.,Silvestro, / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Panamá: and biological, paleoceanographic, Palaeoecology, 112,187–238. Palaeogeography, Palaeoclimatology, (Middle-Late Miocene, NW, Amazonia): the paleo-Amazon river system of icn fPanamá and the fossil record Miocene of Parinari from the early Fruits andwood of Botany,of 104,685–693. Connaraceae: American Journal record of Panamá and the fossil lower Miocene of Anatomists Journal, 38,366–S2. Wood Association of International reveal an ancient neotropical rainforest: Panamá (Azuero Peninsula) the Cenozoicof Botany,Annals of 20,635–637. Cellulose Peel Technique inPalaeobotany: Palynology, 35,46–84. Colombia: Foothillsand Llanos the Llanos of palynological zonation for the Cenozoic of Missouri BotanicalGarden, 128,134–251. insystematic botanyMonographs from the Festschrift for80 AlanGrahaminhis (eds.), Paleobotany andbiogeography: A Stevens, W.D., Montiel, O.M., Raven, P.H. the last 20 millionyears inPanamá, of S.R., Hoyos, N., 2014,Palynological record Herrera, F., Escobar, J., Koll, R., Manchester, de laBarrera, Sanchez,A., S.,on, C.,Mor Silva-Caminha, S.A.F., de la Barrera, A., Advances, 3,e1602321. Panamá” by O’Dea of theIsthmus “FormationComment (1)on of D., Antonelli, A., Bacon,C.D., 2017, and Biodiversity,. Oxford, John Wiley& Sons C., Antonelli, A., (eds.), Mountains, Climate paleoclimatological implications, inHoorn, et al. : Science th year: in /2018

Kubitzki, K.,2014,Humiriaceae, inKubitzki, K., 1995, S., Monechi, P., Mann, R., Kolarsky, Kedenburg, M.D., 2016, Thermochronological atn, . 17, tnad etay and Tertiary Standard 1971, E., Martini, Martínez, C., Madriñán,S., Zavada, M., Manchester, S.R., Herrera, F., Fourtanier, E., MacFadden, B.J., Jones, D.S., Jud, N.A., Moreno- Lott, T.A., Dilcher, D.D.L., S.S.P., Horn, Vargas, Papers, 295,159–200. America Special Geological Society of marine seismic data and onshore geology: fromPanamá of integration as determined southwestern developmentStratigraphic of M.S. Thesis, Florida, University of american stratigraphyFlorida, and tectonics: Panamá: Implications forcentral south the Azuero Peninsula, exhumation of constraints onCenozoic uplift and Botany, 103,277–289. American Journal Chrysobalanaceae: of of Tecnoscienza, 739–785. theSecondPlanktonic Conference: Rome, of Proceedings 1970, Farinacci,A., in zonation calcareous nannoplankton Quarternary diversification: Grana, 52, 161–180. an oldlineage with recent Neotropical Hedyosmum (Chloranthaceae), record of Jaramillo C., 2013, Tracing the fossil pollen 120, 467–476. Geology,Diatom Biostratigraphy: Journal of North Coastal Peru based on Assemblage of the Belén Classic Fruit and Seed Age of Barron, J., Martínez, J.-N., 2012,Oligocene e0170300. Panamá: PLOS ONE, 12, Late Miocene of theAlajuela Formation, of paleoecology chronology, Integrated and faunas, and flora Perez, V.J., Moran,S.M., Wood, A.R.,2017, J.W.,Bernal, Morgan, G.S., Portell, R.W., Costa Rica: Viejo, Rio Puerto Palaeontologia Electronica, 14,5A. of flora O., Sanford, R.L.Jr., 2011,Pleistocene Springer, 223–228. Plants, Plants, Flowering :Berlin, Vascular Families(ed.), The and Genera of Nelson, C.W., Jud, N.A., 2017,Biogeographic Montes, C.,Cardona,Jaramillo, A., C.,Pardo, Montes, C.,Cardona, A.,McFadden, R.,Morón, Montes, C.,Bayona, G., Cardona,Buchs, A., Rodríguez-Reyes, O., Falcon-Lang, H.,Gasson, Lessios,O’Dea, A., Coates, H.A., A.G., Eytan,R.I., Solid Earth,117,B04105. GeophysicalResearch: seaway:Journal of the Central American Closing of formation: 2012a, Arc-continent collision andorocline Ramírez, D.A., Jaramillo, C.A.,Valencia, V., D.M., Silva, C.A.,Morón,S., Hoyos, N., P.,Jaramillo Collinson,M., C.,2014, e1600883. Panamá: Science Advances, 2, Isthmusof Jackson, the J.B.C., 2016,Formation of E., Soibelzon, L.,Todd, J.A., Vermeij, G.J., N.D., Rachello-Dolmen, P.G., Soibelzon, Leonard-Pingel, J.S., Marko, P.B., Puenson, Keigwin, L.D., Knowlton, N., Leigh, E.G., G.M., Grossman, E.L.,Johnson,K.G., Duque-Caro, S., H.,Finnegan, Gasparini, Budd, A.F., Cozzuol,Coppard, M.A., S.E., Aguilera, O., Aubry, M.-P., W.A., Berggren, R.D., Stallard, R.F., Woodburne, M.O., L.S., de Queiroz, A.,Farris, D.W., Norris, Restrepo-Moreno, S.A., Cione, A.L.,Collins, PlantSciences, 178,241–257. Journal of Calophyllaceae: International Evolution of Panamá andthe from the Lower Mioceneof Implications of 348, 226–229. the Central American Seaway: Science, of V., Nino,2015, MiddleMioceneclosure H., Angel, L.C.,Rodriguez-Parra,Ramirez, L.A., A., Silva, J.C., Valencia, V., Ayala, C., Perez- America, 124,780–799. the Geological Society of closure: Bulletin of centralin Panamá: Implicationsfor Isthmus middle Eoceneandyounger landemergence V., J., Bryan, J.A., Flores, 2012b, Evidencefor D., Bayona, G.A., Jaramillo, C.A.,Valencia, Ramírez, D.A., Hoyos, N., Wilson,J., Farris, S.E., Silva, C.A., Restrepo-Moreno, S., Mammea paramericana sp. nov. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Van DerHammen,T., Hooghiemstra,H., Traverse, A.,2007,Paleopalynology: Netherlands, Rodríguez-Reyes, O.,P., Gasson, Thornton, Williams, G.L., Brinkhuis,Pearce, H., M.A., Wijninga, V.M., Kuhry, P., 1990, A Pliocene Flora and Palynology 1996b, V.M., Wijninga, of ecology Neogene 1996a, V.M., Wijninga, Springer, 813p. Anatomists Journal, 38,1–19. Wood Panamá: Association of International a new Miocene malpighialean tree from Panascleroticoxylon crystallosa C., Falcon-Lang, H.J., Jud, N.A., 2017, Palynology, 209,11–34. Palaeobotany and implications: Review of (Central America) and their biogeographic Panamá the Cucaracha Formation of of Miocene (early to mid-Burdigalian) part Fossilwoods (Malvaceae) from the lower f te ca Diln Porm Scientific Results, 189,1–98. Program, Drilling Ocean the of Late Cretaceous-Neogene: Proceedings cyst events compared: Index events for the dinoflagellate global and Ocean Southern Fensome, R.A.,Weegink, J.W., 2004, and Palynology, 62,249–290. Palaeobotany Oriental, Colombia): Review of From the Subachoque Valley (Cordillera Palynology, 92,329–350. Palaeobotany and implications: Review of and chronostratigraphicalbiostratigraphical Río Frío 17(Cordillera Oriental, Colombia): the Early Pliocene section paleobotany of Palynology, 92,97–156. Palaeobotany and lowland forests: Review of montaneand the paleobotanical record of altitude, Cordillera Oriental, Colombia): the Salto de Tequendama site (2475 m 725–742. Science Amazonia:Reviews, Quaternary 19, vegetation, climate, andplant diversityin of history 2000, Neogene and Quaternary

gen. etsp. nov /2018 239 239 .:

New records of Humiriaceae fossil fruits from the Oligocene and Early Miocene REFERENCES