Boletín de la Sociedad Geológica Mexicana ISSN: 1405-3322 Sociedad Geológica Mexicana, A.C.

Rodríguez-Reyes, Oris; Estrada-Ruiz, Emilio; Gasson, Peter Evidence of large from the - early Santiago Formation, Azuero, Panama Boletín de la Sociedad Geológica Mexicana, vol. 72, no. 2, 00006, 2020, May-August Sociedad Geológica Mexicana, A.C.

DOI: https://doi.org/10.18268/BSGM2020v72n2a300719

Available in: https://www.redalyc.org/articulo.oa?id=94366150006

How to cite Complete issue Scientific Information System Redalyc More information about this article Network of Scientific Journals from Latin America and the , Spain and Journal's webpage in redalyc.org Portugal Project academic non-profit, developed under the open access initiative This is an open access article under the CCBY-NC-SAThis Universidad NacionalAutónoma de México. Peer Reviewing under the responsibility of Manuscript July accepted: 30,2019 Corrected manuscript received: July 20,2019 Manuscript received: 26,2019 April org/10.18268/BSGM2020v72n2a300719 Mexicana 72(2),A300719.http://dx.doi. Panama: Boletín de la Sociedad Geológica early MioceneSantiago Formation, Azuero, Anacardiaceae treesfrom the Oligocene– large P., 2020,Evidenceof Gasson, E., O., Estrada-Ruiz, Rodríguez-Reyes, How tocitethisarticle: [email protected] *Corresponding author:(O. Rodrguez-Reyes) rey TW93AB, UnitedKingdom. 4 11340, CDMX,. Plan deAyala y Prolongaciónde Carpio s/n, Biológicas, Instituto Politécnico Nacional, de Ecología, Escuela Nacional de Ciencias 3 ama, Panama. Pan 0843-03092, Balboa,Republic Ancón of 2 Panama. versidad de Panamá. 00017, 0824, Apartado Tecnología, de Botánica, Uni Departamento 1 Oris Santiago, Azuero,Panamá Evidencia deárbolesgrandesAnacardiaceaedelOligoceno-MiocenotempranoenlaFormación Formation, Azuero,Panama Evidence oflargeAnacardiaceaetreesfromtheOligocene–earlyMioceneSantiago (https://creativecommons.org/licenses/by-nc-sa/4.0/) Royal Botanic Gardens, Kew, Richmond,Sur de Zoología, LaboratorioDepartamento Smithsonian Tropical Research Institute, Box Facultad de Ciencias Naturales, Exactasy Rodríguez-Reyes 1,2,* , Emilio - - - Estrada-Ruiz Santiago Formation, Panama. Oligocene–Miocene, wood, Keywords: the Anacardiaceae. Mexico) beingadivergence center of supports Central America (including and thefamily ical biogeographyof thehistor of to theunderstanding bythe Oligocene to Mioceneadds these Anacardiaceae in Panama of with large occurrence crystals. The pitting and raysuniseriatemostly vessels (>200 µm), simplevessel-ray Anacardiaceae woods, such as:large share diagnosticfeatures withseveral extant material. two These specimens extensivewith fossil comparisons and wood anatomical characters and Anacardiaceae, using fossilgenus of described and identified this new Among the samples collected we have the Azueroin Peninsula, Panama. cene–Miocene Santiago Formation, lected the fossil woods inthe Oligo Central America to date. We col Anacardiaceae foundPanamain and a fossilwood of record of formal sandovalii Anacardiaceae. Oligocene and Mioceneepochsis record worldwide, especially for the the families with a rich fossil of America duringthe Cenozoic. One that inhabited Central We the have poor knowledge of ABSTRACT Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín gen. et sp. nov. the is first 3 , Peter Anacarcadiaceae, Llanodelacruzoxylon Gasson 4 - - - con un abundante registro fósil a nivel con unabundante fósil a nivel registro zoico. Anacardiaceae esunadelas familias habitaron América Central durante el Ceno Tenemos poco conocimiento de las floras que RESUMEN Formación Santiago, Panamá.Formación Oligoceno–Mioceno, Madera fósil, Palabras clave: Anacarcadiaceae, deAnacardiaceae.divergencia tral (incluyendo México) como un centro de familia y soporta la idea de América Cen de la a entenderlahistoria biogeográfica Panamá para elOligoceno-Mioceno, ayuda grandes. La presencia de Anacardiaceae en simples y radios uniseriados con cristales (>200 µm),punteaduras vaso-radiales de Anacardiaceae, comoson: vasos grandes con maderascaracterísticas diagnósticas Estos dos especímenes comparten moderno. comparacionesextensas de material fósil y caracteres anatómicos de lamadera y especímenes como Anacardiaceae, usando Panamá. Describimos e identificamos estos Formación Santiago, Península de Azuero, Recolectamos las maderas fósiles en la en Panamá y América Central hasta ahora. madera fósil de Anacardiaceae encontrada gen. etsp. nov. de una eselprimer registro Mioceno. mundial, especialmente para el Oligoceno y /72(2)2020 Llanodelacruzoxylon

sandovalii 1 - -

Large Anacardiaceae trees from the Oligocene–early Miocene, Panama ABSTRACT INTRODUCTION/MATERIALS Large Anacardiaceae trees from the Oligocene–early Miocene, Panama AND METHODS 1. Introduction 2 2 and climatic adaptation in angiosperm evolution and climatic adaptation angiosperm in movement tion andthe relative importance of diversifica tropical of history biogeographic the provide anexcellent opportunity for investigating Anacardiaceae () and Burseraceae Anacardiaceae. quently, we erect a new fossil- and of any genus.specific extant foundin arenot Conse they although the Anacardiaceae, to affinity an date. wood The hasseveral features that support eredPanama in andprobably, Central America, to Ocú town (Jud andDunham,2017). the Santiago Formation, found inthe surrounds of fromceae/Burseraceae the Oligocene-Miocene and one fossil wood that resembles Anacardia Tonosí Formation (Herrera Peninsula: ceae from Panama, both discovered in the Azuero (Weeks these records are from South America and al. al. Cabrera and Cevallos-Ferriz, 2004;Gregory 1954; Wheeler andManchester, 2002;Martínez- associatedspecimens (e.g.,with thefamily Kruse, (Stevens, 2001onwards), withapproximately 80 Anacardiaceae extensiveis worldwide record of diaceae (Estrada-Ruiz Anacar based onthe oldest fossil record of the divergence occurred approximately 73 Ma, evolutionary history. Xie habitats (Weeks of species, theAnacardiaceae occupya widerrange families have approximately the same number of 1988; Pennington tropical forests andtropical wet forests (Gentry, temperate, seasonallydry are majorelements of continent except Oceania and Antarctica and (Weeks / , 2017;Woodcock , 2009;Estrada-Ruiz Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Here we report the largest fossiltrunk discov fossilAnacardia There aretwo only reports of Anacardiaceae/Burseraceae have a long et al. et al. Dracontomelon , 2014). , 2014). These families occur onevery, 2014). These et al. et al. et al. , 2010). Although these two , 2014). et al. L. endocarps from the L.endocarps et al. 07.Temjrt f majorityof , 2017).The et al. , 2010;Pérez-Lara , 2010). The fossil, 2010). The (2014) suggested (2014) suggested http://dx.doi.org/10.18268/BSGM2020v72n2a300719 et al. , 2012) et et ------/72(2)2020 2. Materialsandmethods shows the Holotype collection locality shows theHolotypecollection Figure 1 ure paratype 2Ato2C). The STRI 45789, was in length and 80cminpreserved diameter (Fig gitude 80°53’11.2”W)(Figure 1). in LlanodeLaCruz(latitude N; lon 08° 09’4.7” a large44038B is trunk collected on aprivate farm lected in the Veraguas province inPanama. STRI tworeportedThe specimens herein were col theAnacardiaceae. was adivergence centerof dence that Central America (includingMexico) new record This presented is asadditional evi wide and8.5cminlength. approximately in 9.3 cm paratypea size has of 08° 13’46.4”N;longitude 80° 51’45.1”W). The collected in Boquerones, San Francisco(latitude A)MapofPanamaand theAzuero Peninsula,and B)It The preservedThe holotype fossiltrunk is~ 20m Article A300719 - - - - Article A300719 2.1 GEOLOGICAL CONTEXT preserved totaltreelength. Followarrowstoseecontinuityofthefossil extent.C,Preservationof of thefossil theexternal part wood Figure 2 ferential weathering.area been This hasmapped vegetation,resistantthe most as from ‘clasts’ dif with covered farms cattle on specimen float a as woods we have collected from thisarea, is found fossil all STRItional 45789,asthe system. others deposi main the as environments fluvial indicate parallel to bedding. Finingupward sequences sandy mudrock breccia,and sedimentary of ers in Panama to date. trunk is lyingwithinlay The STRI the 44038 is largest fossilized trunk found The “big ” original position inthefield.A,Preservedand length its ofthe“big tree” (arrows). B,Logs splitfromtheoriginal http://dx.doi.org/10.18268/BSGM2020v72n2a300719 Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín - - - opeiy ako goodoutcrops,complexity, andabsence lackof due to its stratigraphic been clearly determined the specimenswaspoor. early Miocene age, although the preservation of concluded that theysupport alate Oligocene to from theMacaracasBasinincentral Azuero and , foraminifera, and nannofossils samples of (Buchs Formation that outcrops inthe Macaracas Basin Formation, sometimes referred to as Macaracas the Oligocene to Miocene Santiago as part of h g fthe Santiago Formationnot has age of The et al. , 2011).Kolarsky / 72(2)2020 et al. (1995)reported 3 3 .

Large Anacardiaceae trees from the Oligocene–early Miocene, Panama MATERIALS AND METHODS MATERIALS AND METHODS/ Large Anacardiaceae trees from the Oligocene–early Miocene, Panama RESULTS 2.3. IAWA FEATURE DESCRIPTIONANDCODING IMAGING 2.2 ACCESSION DATA,SPECIMEN PREPARATION, AND 4 4 biostratigraphy. rently testing detrital zirconsand studying the radiometric elements for dating. Weof are cur feature isvariable inoccurrence. presentis to indicate or absent and “v” that the that there is uncertainty as towhether the feature code numbers. We use the‘?’to indicatesymbol the IAWAdescriptions, we give character a list of vessel element length, and ray height). In the diameter, vessel-rayparenchyma pit diameter, surements (meanvessel diameter, intervessel pit other quantitative 25 mea For features, area. we obtained a minimum of of mm2 1 of fields different 10 in measurements made we grouping, vessel frequency,data of ray density, and vessel tion (IAWA Committee, 1989).For quantitative FeaturesIdentifica Hardwood for (IAWA) of List Wood Anatomists Association of the International We described the fossilwood following specimens 2017). 1994, 1999;León, 2003, 2014;Woodcock edu; Wheeler, 2011)andliterature (e.g., Terrazas, Inside Wood Database (IWD; insidewood.lib.ncsu. andfossilwoods inthe modern able images of MRc5 camera. with anAxioCam V16, photographed Zoom AXIO 10.5MpixandaZeiss CMOSsensor of BX53 anddigital camera Olympus SC100 with Material was observed and imaged using an balsam. Canada with affixed were coverslips and μm, ~30 of thickness a to ground , EpoFix Sections wereslides using mountedonglass (RLS) and tangential longitudinal (TLS) sections. prepared in transverse (TS), radial longitudinal fossilmaterial were Petrographicof thinsections / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín The fossilThe woods were compared with the avail http://dx.doi.org/10.18268/BSGM2020v72n2a300719 et al. - - - - /72(2)2020 , 3.1. SYSTEMATIC PALEOBOTANY 3. Results simple; intervessel polygonal pitting alternate; a fewshortradial multiples; perforation plates with combined solitary are vessels porous; diffuse Diagnosis: ures 2to4). guez-Reyes, Estrada-Ruizsp. et Gasson, nov. (Fig Species — Estrada-Ruiz etGasson,gen.nov. Genus — Family —Anacardiaceae Lindley Order —SapindalesDumortier Description: 99a, 100a,130a,136p, 189p. 11a, 13p, 22p, 31p, 40a, 41a, 66p, 78p, 79p, 98a, Description inIAWA feature numbers: 2p, 5p, 10a 11.2” W. Latitude 08° 09’4.7”Nandlongitude 80° 53’ ero Peninsula). Type locality: Institute, Panama City, Panama. and Archaeology, SmithsonianTropical Research Repository: Paratype: Holotype: large trunkfor thisstudy. Mr. the Carlos Sandoval, who provided samplesof of recognition in is epithet specific The collected. Llano dela Cruz locality, where the holotype was Etymology: Species diagnosis: upright ray cells. veryrhomboidal abundantcrystals in body and solitary large walled, to thick thin fibers, septate lar, mostlyuniseriate and very rarely biseriate, non raysheterocellu vasicentric, andslightlyaliform, to paratracheal scanty diffuse, apotrachealchyma circularmainly withreduced borders, axial paren and mediumto large; vessel-ray parenchyma pits STRI45789 STRI44038B Growth ringsare indistinct; wood is Llanodelacruzoxylon Llanodelacruzoxylon sandovalii Center for Tropical Paleoecology Descriptionbasedontwo samples. The generic name refers to the The Llano de la Cruz, Veraguas (Azu Asfor thegenus Rodríguez-Reyes, Article A300719 Rodrí - - - - - Article A300719 longitudinal section (TLS). Simple perforation plate (arrow). F, (TLS). Alternate intervessel pits (arrow). G, (TLS). Showing parenchyma Showing (TLS). G, (arrow). pits intervessel Alternate (TLS). F, (arrow). perforation plate Simple (TLS). section longitudinal parenchyma. E,Tangential Closeup withvasicentric parenchyma. D,(TS). ofvessel Threewithvasicentric vessels parenchyma. C,(TS). and Solitaryvessels diffuse axial solitaryandfewinradialmultiples.B,(TS). Diffuse porouswoodwith vessels (TS). Transverse section Figure 3 strands Detailof (arrows). (RLS). withreduced thevessel-ray H,Radial Longitudinal borders, parenchyma pits Section round inshape (arrow).

Llanodelacruzoxylon sandovalii Rodríguez-Reyes, Estrada-Ruiz et Gasson gen. et sp. nov.Gasson gen. etA-F,STRI 44038B; G-H,STRI45789.A, Rodríguez-Reyes,Estrada-Ruiz et http://dx.doi.org/10.18268/BSGM2020v72n2a300719 Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín / 72(2)2020 5 5

Large Anacardiaceae trees from the Oligocene–early Miocene, Panama RESULTS Large Anacardiaceae trees from the Oligocene–early Miocene, Panama RESUTLS / DISCUSSION alignment. Crystals D,(RLS). inthesquarebodycells. rays, procumbentand squarebodycellsinradial Heterocellular (RLS). crystals. C, with rays Uniseriate Raysmostlyuniseriate,A, (TLS). somebiseriate. B,(TLS). Estrada-Ruiz etGassongen.sp.nov. HolotypeSTRI 44038B. Figure 4 6 6 element length is365(range= 203 –504,SD= vessel mean 3H); (Figure μm) 12 – 7.5 diameters with reduced borders, round inshape(mean pit pits parenchyma vessel-ray 3F); (Figure μm) 12 – onal and medium to large 8 (mean pit diameters intervessel and crowded, pits are alternate polyg limeter; perforation plates are simple (Figue 3E); (range = 4–9,SD 1.5) vessels per square mil 6 3D); (Figure μm 35.8) = SD 205, – 96 = (range (Figure 3D); vessel mean tangential diameter 154 (Figures 3Ato 3D), circular tooval in outline 2–3(–5) a fewshortradial multiples (34%) of porous; vessels are (66%) combined with solitary Wooddiffuse absent. is to indistinct rings Growth /

Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Llanodelacruzoxylon sandovalii http://dx.doi.org/10.18268/BSGM2020v72n2a300719 Rodríguez-Reyes, - - /72(2)2020 4.1 INSIDEWOOD SEARCH 4. Discussion bent ray are cells (Figure crystals large, 4C). The upright raycells inradial alignment in procum veryrhomboidal abundantcrystals in body and 4B), thin to thick walled. Solitary, (Figure non-chambered, fibers Non-septate 4D). and 4C (Figure bent and upright) cells throughout the ray body mixed (procum (Figureof and 4B),composed 4A quency is19per mm (range = 15 –23,SD= 2.7) rarely biseriate (Figure 4A and 4B). Mean ray fre 6-celled (Figure 3G). parenchyma(Figure 3C).Parenchymaarestrands 3B to 3D). Some vesselsslightly aliform possess and scantyparatracheal to vasicentric (Figures diffuse apotracheal is parenchyma Axial μm. 87) (100a); radial canals absent (130a); prismatic(100a); radial canals ate portion(s) aswide asuniseriate portions absent rays>10-seriate absent (99a); Rayswith multiseri vasicentric (79p);rays4-10 cells wideabsent(98a); (78p), Axialparatrachealscanty parenchyma non-septate (66p); axial paratracheal parenchyma fibers (41a); absent µm -100 50 diameter gential diameter <50 µmabsent(40a); vessel meantan reduced borders (31p);vessel meantangential (22p); vessel-raypits alternate pitswithmuch (11a); perforationplates (13p); intervessel simple absent (10a);vesselmon absent common clusters 4or more com (5p); vessels inradial multiples of rings indistinct or absent (2p); wood diffuse porous IWD includedthe following characters: growth et al. radial (Terrazas,canals 1994;Bell occurrences of fewer and fibers, septate frequent more chyma, tinguished byhaving more abundant axial paren very similar, however Anacardiaceae can bedis and AnacardiaceaeBurseraceae are anatomically (Figures 4Cto4D). width in µm 39 to 22 and length in μm 50 to 43 Rays heterocellular, mostly uniseriate, very The mostrestrictiveThe search we obtained in the , 2010). Article A300719 ------Article A300719 4.2 CO in diagonal pattern andvesselin diagonal pattern grouping common, sil, occasionalreticulate perforation plates, vessels thaceae) severalhas features distinctfrom the fos different from the fossil. and are mainlyshrubs, andnottreesthe fossil. as intervesseldaceae) possesses pitting large (>10µm) Moringaceae.distinguish and more abundant parenchyma. Storied rays growth ring boundaries, vessel grouping common ruled Rhamnaceae out byis distinct its occasional roxylaceae and Malvaceae. result of single The apotracheal parenchymarules outthe Eryth banded thaceae and Rhamnaceae. Abundance of Malvaceae, Moringaceae, Oxalidaceae, Phyllan Erythroxylaceae, Gentianaceae, Juglandaceae, onlyone or two speciesinfamilies such as: of with distinct borders. other results The consisted compared to this fossilwood andvessel ray pitting parenchyma vessels,solitary greater abundance of exclusively Myrtaceae can beruled out becauseof the following Anacardiaceae taxa: to theAnacardiaceae. procumbentin crystals andupright raycells lead largefibers solitary and rhomboidal abundance of uniseriate, very rarely biseriate rays, non septate tracheal to vasicentric, raysheterocellular, mostly para scanty and diffuse apotrachealparenchyma pits mainlycircular with reduced borders, axial intervessel pitting alternate, vessel-ray parenchyma short radial multiples, perforation plates simple, porous, vessels combined with very solitary few diffuse wood indistinct, rings growth specimens, traits described from these two mosaic of The further inthediscussion. identification genus the Wedetail herein. studied thatthe family diaceae is resembles most thefossil and septate fibres present. differentvessel-ray sizes pits inof the same ray cell otyseiso Anacardiaceae and Myrtaceae. mostly speciesof traits led to 18 results that included bination of present (136p);treecrystals com (189p).This MPARISON WITHANACARDIACEAE GENERA The most restrictiveThe search in IWD included weAfter the comparison conducted,Anacar Bridelia micrantha Sarcotheca glauca http://dx.doi.org/10.18268/BSGM2020v72n2a300719 Faguetia falcata Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín (Phyllan (Oxali - - - - - , - - - as growth rings indistinct, wood diffuse porous, diffuse wood indistinct, rings growth as withthe Anacardieae suchacters (sensu Engler), simple perforation plates andnon-septate fibers. has fossil the whereas fibers, septate non- and tate perforation platesscalariform possesses and sep a low ray density(<4 per linear mm). parenchyma andoccasionallyit has and bandsof results and ences between the differ noticeable most The . the in absent parenchyma.All these charactersare of bands like, and gash and occasional tylosescommon sp. Mangifera caesia, Mangifera Rhus, Schinus, Toxicodendron for example are some genera that have pits smaller than 8 µm, ray cells(Dra. Teresa Terrazas com.pers.). There and abundantprocumbentin crystals andupright specieswithuniseriate rays has a good number of vessel-ray pits. of a similar pattern mention Welarger pattern. pits inpalisade and gash-like and small(7.6 to 12.0 µm), most anacards have Although vessel-ray pits are all simple, circular Anacardiaceae.fossilin modern isnot common Moreover, vessel-raypitting in the of the pattern genusthe diagnostic features. possesses modern characters inthisfossiloccur inAnacardiaceae, no that revisionrevealedit was that although all Dra.Teresa Terrazas. of under supervision From UniversidadNacional Autónoma deMéxico, slide collection fromphology Instituto de Biología, consulted the largest Anacardiaceae micromor the taxa match the fossil. We then that none of razas, 1994) Anacardieae compared totheother tribes (Ter on the intervessel pitting size that is larger in the riate to biseriate and raysheterocellular. We stress paratracheal parenchymapresent, butabundant; no raysunise fibres, septate non mm2), per (<5 polygonal andlarge (>10 µm), meanvessel density vessels intervascular withnoarrangement, pitting At tribe level, thisfossilshares several char From the results inInside Wood, we note Faguetia falcata Mangifera LI. sandovalii Cotinus, Dobinea, Haplorhus, Lithraea, hasvessel-ray pitting in palisade a n fthe genera that has asoneof Mangifera , arewider rays basedon , and / 72(2)2020 cf. griffithii species from the Trichoscypha and Mangifera Protorhus Protorhus . These . These also 7 7 ------

Large Anacardiaceae trees from the Oligocene–early Miocene, Panama DISCUSSION Large Anacardiaceae trees from the Oligocene–early Miocene, Panama DISCUSSION 8 8 the Inside WoodDatabase (Wheeler Table 1.Summary of comparisonbetweenAnacardiaceae fossilwoodsand

Tetragastroxylon Tetragastroxylon Sumatroxylon Canarioxylon 06010441703 Burseroxylon Boswellioxylon Terrazoxylon Tapirira Swintonioxylon Sclerocaryoxylon Schinosipxylon Rhus Resinaxylon site Oregon Post Pistacioxylon Melanorrhoeoxylon Maureroxylon Mangiferoxylon Loxopterygium Lanneoxylon Holigarnoxylon Glutoxylon Edenoxylon Dracontomelumoxylon Dracontomeloxylon Mangifera cf Buchanioxylon Bouea Bosquesoxylon Astroniumxylon Anacardium Anacardioxylon /

Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Taxon Comparative

indicum

Tyloses common, septate fibers common, septate Tyloses canals, radial fibers, Septate R common tyloses fibres, Septate 4- rays fibres, Septate canals Vessel f canals radial fibres, Septate T T S canals radial T septatefibres, Woodsemi V bands tangential G thickenings, common,tyloseshelical canalsradial G canals radial rays, T G parenchyma G T S T B W T S A S B S R S S

ossil g ossil eptatefibres, axial eptatefibres, rays 4 eptatefibres, rays 4 canals radial fibres, eptate eptatefibres, rays 4 eptatefibres, vascular/vasicentric tracheids eptatefibres, rays 4 ylosescommon, canalsradial fibres, septate ylosescommon, banded parenchyma, canalsradial ylosescommon, rays4 homocellular parenchyma, banded common, yloses ylosescommon, canalsradial fibres, septate ylosescommon, aliform parenchyma ylosescommon, rays4 fibres, septate adial canals, septate fibers, septate canals, adial anded 50 vessels parenchyma, anded fibres septate canals, adial esselsin diagonal pattern, septate fibres, radial canals rowth rings distinct, wood ring pattern, diagonal in Vessels distinct, rings rowth rowth rings distinct, septate fibres banded parenchyma, aliform distinct, rings rowth parenchyma marginal parenchyma, liform ood semi ood - ray pitting similar to intervessel parenchyma,canalsradial of enera of et al - - ring porous, vessel clusters common, clusters vessel porous, ring canals radial porous, ring

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., 2014 .,

Franco Franco 2006

Article A300719 REMARKS 4.4. PALEOECOLOGICAL ANDBIOGEOGRAPHICAL ANACARDIACEAE 4.3 COMPARISON WITHOTHER FOSSILWOODS OF in thesespecimens. anyfind not the all thatcharacters fossil wood has taxa. We summarize thissurveyin Table 1. We did sandovalii the IWD,using order in toelucidate whether et al. from Europe,and South America (Ramírez Asia et al. Lara Shukla andMehrotra, 2016; Allen,2017;Perez- Cheng Franco, 2009;Pujana, 2009; Gregory been reported worldwide (Franco andBrea, 2008; fossilwoods. More than eighty wood typeshave anacardiaceous There are abundant records of few uniseriate rays aswell asradialcanals. pha genera do nothave parenchyma. aliform aiist nesadmgainpten fof patterns migration families tounderstand vations inthefield. their properties geological and obser because of the Santiago Formation were described aspart of lying was it where sandstones the confirmed We 2A to2C),according toNiklas’(1995)equation. this “bigtree” was probably ~35 m high(Figures length and 2.5 m wide. suggest dimensions The STRI 44038B preserved~ 20min is trunk a newfossil anacard genus andspecies. tracheids, rays1–3 cells wide. Therefore, we erect in having septate fibers, vascular vasicentric 2017). However, Forest Piedra Chamana inPeru (Woodcock ardium incahuasi our fossil woodobservedthe sameas is in We remark that the vessel-ray in pitting pattern We surveyed the Anacardiaceous fossil woods Anacardiaceae are and Burseraceae good has unilateral winged-aliform, but hasunilateral it presents winged-aliform, , 2000). 07.Ago ubro therecordscome , 2017).Agoodnumber of et al. et al. resembles other previously reported fossil , 2017;Wheeler , 2012;Mendez-Cardenas fo al oeeo the Fossil from early Eocene of A. incahuasi et al. fromdiffers our wood http://dx.doi.org/10.18268/BSGM2020v72n2a300719 , 2017;Woodcock Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín et al. et al. Trichoscy , 2009; , 2014; Anac et al. LI. - - - , record. Anacardiaceae diversified during the Cre the during diversified Anacardiaceaerecord. with the fossilreconstructions highlycongruent Weeks (Weeksbiomes thanBurseraceae occupied a wider and hassuccessfullyrange of explain why Anacardiaceae has become more widespread help may difference This Burseraceae. habitats than tolerance to a more diverse range of and morphology a much wider diversity of al. to North and South America (Weeksmigration played2014). Longdistancedispersal akeyrole in Asia and diverged during the Cretaceous (Xie Group,2016). Bothfamiliesprobably originated in Phylogeny families(Angiosperm angiosperm family inthepast. the (including Mexico)a divergence was center of that support and helpsthe theory Central America Anacardiaceae the historical biogeographyof of ama, Central America addsto the understanding Paleogene. Ourreport of New and Old world forests were inthe common and thattions events longtimedispersal between local extinc this could represent an example of Miocene Cucaracha Formation, theyconclude Anacardiaceae genus infossil bedsfrom the lower this Asia andAfricatoday. Based on the absence of alized that endocarps, is a genus onlyinhabiting one Anacardiaceae ( only identified they Formation, this From forest. (Azuero, Panama) that a diverse suggested rain assemblages in the Eocene Tonosí Formation Herrera Anacardiaceae decrease towards theMiocene. the Oligocene inCentral America. Reports of Anacardiaceae was augmented especially during tolerances (Weeks climatic range and likely a diversity of geographic during the Miocene, when ithada widespread steadily colonize Eurasia andtemperate zones thatis suggested Anacardiaceae continued to probably tookplace during the Paleogene. It Fromthere, theroute to conquerSouthAmerica taceous andexpanded intosub-Saharan . , 2014;Xie et al. et al. et al. (2014) performed ancestral area (2014) performed (2012) found endocarp rich (2012)found endocarp et al. , 2014).Anacardiaceae displays Dracontomelon 04.Tefsi eodo fossilrecord, 2014).The of / 72(2)2020 LI. sandovalii ) from perminer et al. , 2014). from Pan et al. 9 9 et - - - - - ,

Large Anacardiaceae trees from the Oligocene–early Miocene, Panama DISCUSSION Large Anacardiaceae trees from the Oligocene–early Miocene, Panama ACKNOWLEDGEMENTS / REFERENCES References Acknowledgements 10 10 Awasthi, N., Srivastava, R.,1990.Some new PhylogenyAngiosperm Group, IV. 2016,An Allen, Bouea in the A. 1988.Occurrence of Agarwal, and CONACyT toE.E.R. (240241) grants Oris J. Rodríguez-Reyes andSIP-IPN (20195100) funded by the SENACyT to ITE15-023 grant the fossils.Thisidentification research of has been ardiaceae wood anatomyfor the on discussions thank Dra.Teresa Terrazas, anexpert on Anac themanuscript. We particularlyearly version of for comments onan California) (University of phy assistance. We thank DoriLynneContreras Instituto de Biología, UNAM, for microphotogra help with the images andto Susana Guzmánat the Lara from Instituto Politécnico Nacional for her Pujana. We are Pérez-to Biól.DianaK. grateful reviewer, Dr. MarianaBrea and Dr. Roberto comments to our manuscript from ananonymous Archaeology), Panama. We really appreciate the CTPA(Center for Tropical Palaeoecologyand Martínez and the Carlos Jaramillo´s labinthe from Judith Callejas-Moreno, César Silva, Miguel field the in help valuable acknowledge We farm. doval for allowing usto collect fossilsprivateon his and Mr. (MICI) for collectionpermits CarlosSan We thankthe Ministerio deComercio eIndustrias / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín palaeoclimate: Palaeobotanist 38,285–292. Kerala Coast and their bearing on of carbonized woodsfrom the doi.org/10.1111/boj.12385 theLinnean Society, 181(1), 1–20.https:// of IV. APG Journal plants: Botanical flowering of families and orders the for classification Phylogeny the Angiosperm Group update of https://doi.org/10.1086/694186 Sciences, 178(9), 689–714. 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