Eocene Journal Anacardiaceae 38 (4), 2017: 543–552From Chiapas, Mexico 543

Pérez-Lara et IAWAal. – Eocene Journal Anacardiaceae 38 (4), 2017: 543–552from Chiapas, Mexico 543

A new genus of Anacardiaceae fossil wood from El Bosque Formation (Eocene), Chiapas, Mexico

Diana K. Pérez-Lara1, Carlos Castañeda-Posadas1, and Emilio Estrada-Ruiz2,* 1Benemérita Universidad Autónoma de Puebla, Facultad de Ciencias Biológicas, Blvd. Valsequillo y Av. San Claudio, Edificio BIO-1, Ciudad Universitaria, 72570 Puebla, México 2Laboratorio de Ecología, Departamento de Zoología, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional, Prolongación de Carpio y Plan de Ayala s/n, 11340 Ciudad de México, México *Corresponding author; e-mail: [email protected]

Abstract We describe a new fossil wood from the El Bosque Formation (Eocene) in Chiapas, southern Mexico. It has a combination of features found in the Anac- ardiaceae, including distinct growth rings, diffuse porosity, vessels solitary and in radial multiples of 2–3, simple perforation plates, medium to large alternate intervessel pits, vessel-ray parenchyma pits rounded and elongate with reduced borders, septate and non-septate fibers, axial parenchyma scanty paratracheal, vasicentric, apotracheal diffuse, Kribs heterogeneous rays type IIA, and multiseriate rays with radial canals. The mosaic of features of this wood supports the erection of a new genus, Bosquesoxylon Pérez-Lara, Castañeda-Posadas et Estrada-Ruiz. This new genus of anacardiaceous fossil wood extends our knowl- edge of this family’s history and offers hints on the possible relationships with floras from other localities worldwide, especially North America and Asia. Keywords: Anacardiaceae wood, Bosquesoxylon, Eocene, Chiapas, Mexico.

Introduction Anacardiaceae is a family with around 75 genera and 1000 species, represented by trees, shrubs and some woody climbers, with a wide distribution in tropical and warm-tem- perate regions (Mabberley 2008). In Mexico, there are ~62 extant species distributed in 20 genera, including some trees of economic importance, e.g., Spondias purpurea L. (jocote) (Martínez-Millán & Cevallos-Ferriz 2005; Medina-Lemus & Fonseca 2009). Anacardiaceae have an abundant fossil record. Fossils of the family have been reported worldwide, principally in America and Asia. In a literature review, Ramírez- Garduño (1996) listed ~120 fossil species in 30 genera, and that number has recently in- creased. In Mexico, several anacardiaceous macrofossils have been described, including flowers, leaves and woods. An Oligocene wood resembling Loxopterygium Hook. f. from Coayuca de Andrade, Puebla (Méndez-Cárdenas et al. 2014); Oligocene-Miocene flower and wood of Tapirira (Miranda 1963; Martínez-Cabrera & Cevallos-Ferriz 2004, respectively). Leaves and/or leaflets of Pseudosmodingium Engl., Haplorhus

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Engl., Rhus L., Comocladia L., and Pistacia L. from the Oligocene Los Ahuehuetes locality Tepexi de Rodríguez in Puebla (Ramírez et al. 2000; Ramírez & Cevallos- Ferriz 2002). The record of anacardiaceous fossil woods is rich, with approximately 78 wood types reported worldwide (e.g., Awasthi 1966; Gregory et al. 2009). The majority of the records are from Cenozoic sediments of Europe, Asia, and North and South America (Martínez-Cabrera & Cevallos-Ferriz 2004; Gregory et al. 2009). The oldest known anacardiaceous woods are Anacardiaceoxylon semecarpoides Prakash & Dayal (1964), and Dracontomeloxylon palaeomangiferum Prakash from the latest Cretaceous-earliest Paleocene Deccan Intertrappean Beds of India (Bande & Khatri 1980; Wheeler et al. 2017). The Deccan sample of Lanneoxylon grandiosum that Srivastava and Guleria (2004) described has features of both the Anacardiaceae and Burseraceae clade of the Sapindales (Wheeler et al. 2017). Recently, Estrada-Ruiz et al. (2010) described wood resembling Anacardiaceae /Burseraceae from the late Campanian of the Olmos Formation. We describe a new genus and species of Anacardiaceae from the El Bosque Formation (Eocene). Bosquesoxylon represents the oldest record of radial canals in Mexico. This new fossil wood further supports the hypothesis that Mexico has been a diversification center of the Anacardiaceae since the Paleogene, as suggested by its extant and fossil diversity in Mexico.

MATERIALS AND METHODS Fossil wood was collected from a single outcrop locality of the El Bosque Formation. The locality is known as “Las Maderas Acala” and is located approximately 10 km southeast of the county seat Acala, Chiapas, Mexico (92°43'47" N and 16°30'41" W) (Fig. 1).

-93° -92° -92° -92° -92° 16° 16°

16° 16°

Figure 1. Location of the fossiliferous outcrop. -93° -92° -92° -92° -92°

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The Bosque Formation has been dated as Eocene. This formation consists of a sequence of ~37 meters of strata and is a rhythmic sequence of siltstones, shales, calcareous sandstones, and occasional polymictic conglomerates. The Bosque Formation has yielded invertebrate fossils such as gastropods, bivalves, echinoderms, some verte- brates (e.g., sharks and turtles), calcareous algae, leaf impressions, and permineralized woods (Juárez-Hernández 2014). These sediments belong to a continental sedimentary environment marked by marine transgressions and regressions (Juárez-Hernández 2014). In some studies, this formation is divided as the El Bosque (lower Eocene) and San Juan (middle Eocene) Formations (Frost & Langenheim 1974; Müllerried 1982; Ferrusquía et al. 2000). The wood described herein was found in sediments within the El Bosque Formation. We prepared transverse (TS), tangential (TLS), and radial (RLS) sections using standard thin section techniques. The descriptions use terminology from the IAWA Hardwood List (IAWA Committee 1989). Affinities were determined by consulting the literature (e.g., Metcalfe & Chalk 1950; Détienne & Jacquet 1983; Ilic 1987, 1991; Terrazas 1994, 1999; Terrazas & Wendt 1995), and by using the multiple entry key of InsideWood, a wood anatomy website (InsideWood 2004-onwards; Wheeler 2011). For the classification of ray types, we employed the terminology proposed by Kribs (1935). Anacardiaceae has mostly heterogeneous rays (Kribs’s Types IIA and IIB). In the heterogeneous Type IIA, the uniseriate rays are composed of upright cells, and the multiseriate portions of multiseriate rays are composed mostly of procumbent cells with short uniseriate wings of upright cells. In the heterogeneous Type IIB, the uniseriate rays are of two types: composed of upright cells or composed of procumbent cells. Multiseriate portions are composed mostly of procumbent cells and have very short uniseriate wings composed of upright cells (Kribs 1935; Carlquist 2001). The speci- mens described here are housed in the Colección de Paleontología, from Benemérita Universidad Autónoma de Puebla, Puebla, Mexico.

systematiC description Core Eudicots Malvids – Eurosids II Order – Sapindales Family – Anacardiaceae Genus – Bosquesoxylon Pérez-Lara, Castañeda-Posadas et Estrada-Ruiz, gen. nov. Species – Bosquesoxylon chiapiasense Pérez-Lara, Castañeda-Posadas et Estrada- Ruiz, sp. nov. Etymology – The generic name refers to the Formation, where the material was collected. The specific epithet refers to Chiapas State, Mexico, where the El Bosque Formation is found. Holotype hic designatus – BUAPALV 1536 A, B and C. Age – Eocene. Material – Description based on a single mature wood sample of permineralized float wood, about 7.7 cm in width and 13.6 cm long. Estimated original axis diameter of ~60 cm.

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Figure 2. Bosquesoxylon chiapasense Pérez-Lara, Castañeda-Posadas et Estrada-Ruiz, gen. et sp. nov. (BUAPALV 1536). – A: Diffuse porous wood (TS). Scale bar = 300 μm. – B: Solitary vessel, scanty paratracheal parenchyma, and growth ring boundary marked by latewood fibers (arrows) (TS). Scale bar = 100 μm. – C: Simple perforation plate; polygonal and alternate intervessel pits (TLS). Scale bar = 92 μm. – D: Axial parenchyma strand (arrows; F = fibers) (RLS). Scale bar = 16 μm. – E: Multiseriate rays (TLS). Scale bar = 100 μm. – F: Vessel-ray parenchyma pits circular, angular, and horizontally elongated with reduced borders (RLS). Scale bar = 23 μm. – G: Septate fibers (arrows) (TLS). Scale bar = 20 μm. – H: Heterocellular rays and a radial canal (TLS). Scale bar = 100 μm. – I: Heterocellular rays (RLS). Scale bar = 100 μm. – J: Showing three radial canals in a ray (TLS). Scale bar = 100 μm. – K: Prismatic crystals (arrow) (RLS). Scale bar = 30 μm. →

Diagnosis – Distinct growth ring boundaries marked by radially narrowed latewood fibers; vessels solitary and in radial multiples of mainly 2 to 3; intervessel pitting alternate; vessel-ray and vessel parenchyma pits with reduced borders, horizontally elongated to round; non-septate and septate fibers, 1–2 septa per fiber; axial parenchyma mostly scanty paratracheal, also apotracheal diffuse, vasicentric; heterocellular rays, Kribs type IIA, ray body composed entirely of procumbent cells, and usually one marginal row of erect or square cells; radial canals in some multiseriate rays. Description in IAWA feature numbers: 1, 5, 13, 22, 23, 26, 27, 31, 42, 47, 56, 65, 66, 76, 78, 79, 97, 98, 106, 115, 130. Description – Growth rings distinct, the rings marked by four rows of radially flattened latewood fibers (Fig. 2A,B, arrows). Diffuse porous wood. Solitary vessels (36%) and in radial multiples of two to three (up to 6). Vessel elements round to oval in outline (Fig. 2A,B). Mean tangential vessel diameter 131 (range = 55–184) μm; 8 (range = 5–14) vessels per square millimeter; vessel walls in transverse sections of 8 (range = 5–11) μm thick; simple perforation plates inclined 73° (range = 30°–99°) (Fig. 2C); intervessel pitting alternate, crowded, polygonal in outline, 7 (range = 4–19) μm in horizontal diameter, with elliptical apertures (Fig. 2C) with a mean diameter of 3 (range = 2–4) μm; vessel-ray parenchyma pitting with reduced borders, horizontally elongate to round, elongate, 13 (range = 9–23) μm in diameter (Fig. 2F). Mean vessel element length 567 (range = 286–738) μm. Bubble-like tyloses common (Fig. 2C, J). Fibers septate and non-septate, septate with 1–2 septa per fiber (Fig. 2G), thick to very thick walls, pitting not observed. Axial parenchyma scanty paratracheal, vasicentric and apotracheal diffuse (Fig. 2A,B,D), with 5 cells per parenchyma strand (Fig. D, arrows). Rays mostly 2- to 3-seriate (up to 4), uniseriate rays rare (Fig. 2E,H). Rays heterogeneous type IIA, with procumbent body cells and usually one marginal row with erect or square cells (Fig. 2 I). Multiseriate rays averaging 19 (range = 7–36) cells and 492 (range = 277–904) μm high and 5 (range = 2–7) cells and 70 (range = 28–120) μm wide (Fig. 2E). Rays 6–8(–10) per millimeter. Radial canals in some multiseriate rays (Fig. 2H,J); mean tangential diameter of 56 (range = 44–80) μm, delimited by a layer of 1–2 epithelial cells (Fig. 2H), occasionally two or three canals per ray (Fig. 2 J). Prismatic crystals present in the erect or square ray cells (Fig. 2K, arrow).

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Discussion Affinities This El Bosque Formation wood has a combination of features that indicate affinity with the Anacardiaceae and Burseraceae. However, members of Burseraceae generally have higher percentages of uniseriate rays, less abundant axial parenchyma, exclusively septate fibers, and indistinct growth rings (Terrazas 1994; Martínez-Cabreraet al. 2006).

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Table 1. Comparison of Androtium, Buchanania, Myracrodruon, Rhodosphaera, Spondias, and Bosquesoxylon. Hetero- radial Layers of Septa Axial Genera VEG V/mm2 geneous canals epithelial per fiber parenchyma ray type (µm) cells

Androtium 2– 6 3–6 1–2 sp and Vc IIA 31 ± 9 2 Buchanania 1 7–10 5–15 1–2 sp and Vc IIA 34 ± 12 2–3 Myracrodruon up to 6 10–18 2–3 Sp and Vc IIB 33 ± 9 2–4 Rhodosphaera 1 2– 6 12–24 1–2 absent IIB 54 ± 24 2–5 or Sp Spondias 1 up to 5 3–11 2–5 Vc IIA 31 ± 4 2–3 Bosquesoxylon up to 6 5–14 1–2 Dif, Vc, IIA 44–80 1–2 and Sp

VEG = vessel element grouping; V/mm2 = vessel elements per square mm; Sp = scanty paratracheal; Vc = vasicentric; Dif = diffuse. 1Terrazas (1994).

The presence of well-defined growth rings and diffuse porosity (1p, 5p), vessels solitary and in radial multiples, simple perforation plates (13p), large alternate intervessel pits that are polygonal in outline (22p, 23p, 27p), vessel-ray parenchyma pits with reduced borders, pits horizontally elongate to round (31p), septate (65p) and non-septate fibers (66p), axial parenchyma diffuse (76p), scanty paratracheal (78p), and vasicentric (79p), heterocellular rays (106p), and radial canals (130p) is a combination of features that does not exist in any extant genus of Anacardiaceae. The presence of diffuse apotracheal parenchyma and septate and non-septate fibers, further support its inclu- sion in a new genus. An InsideWood search for the features listed above, except for diffuse parenchyma, returned five anacardiaceous genera. Table 1 compares their features with Bosqueoxylon. Differences include: Androtium Stapf and Buchanania Spreng. have smaller canals with up to three ensheathing cells, rarely tyloses; Myracrodruon Allemão and Rhodosphaera Engl. have heterogeneous rays, Kribs type IIB, canals with up to four ensheathing cells; lastly, Spondias L. has fibers with two to five septa, and paratracheal and vasicentric parenchyma to weakly lozenge-aliform (Terrazas 1994).

Comparisons with Eocene anacardiaceous woods Approximately 10 anacardiaceous woods have been described from the Eocene of America and Europe (InsideWood 2004-onwards; Gregory et al. 2009). Rhus crystal- lifera from the Eocene of Yellowstone National Park, Wyoming, USA differs from Bosquesoxylon in that it has ring-porous wood and lacks radial canals (Wheeler et al. 1978). Schinoxylon actinoporosum Kruse and Edenoxylon parviareolatum Kruse are from the Eocene of Eden Valley, Wyoming, USA (Kruse 1954). Schinoxylon actinoporosum differs in having vessels arranged in radial rows and rare to absent axial parenchyma; Edenoxylon parviareolatum differs in having a higher vessel frequency (56–70 per sq.mm) and rays with one to four rows of upright/square marginal cells (Kruse 1954; Boonchai & Manchester 2012). Lastly, three genera from the Middle Eocene Clarno Nut Beds of Oregon have been described: Maureroxylon, Tapirira, and

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Terrazoxylon (Manchester 1977; Wheeler & Manchester 2002). Maureroxylon crystal- liphorum has diffuse-in-aggregates axial parenchyma, exclusively septate fibers, and lacks radial canals. Tapirira clarnoensis differs in having exclusively septate fibers and mostly biseriate rays. Terrazoxylon ductifera Wheeler et Manchester differs because it has indistinct growth rings, higher vessel frequency, and exclusively septate fibers (Manchester 1977; Wheeler & Manchester 2002). Brett (1966) described two species of Edenoxylon from Herne Bay, England: E. aemulum differs in its higher vessel frequency and biseriate rays; Edenoxylon? atkinsoniae Crawley (1989) differs in lacking growth ring boundaries and radial canals.

Comparison of Bosqueoxylon with other anacardiaceous fossil woods Several species of Anacardioxylon Felix have been reported from different localities (e.g., Europe, Asia and Africa), but Anacardioxylon differs in lacking growth ring boundaries. The exception is Anacardioxylon shardai Prakash et Tripathi but its axial parenchyma is aliform (e.g., Schönfeld 1947; Prakash & Dayal 1965; Prakash & Tripathi 1975; Crawley 2001). Astroniumxylon Brea, Aceñolaza et Zucol from the Paraná Formation (Middle Miocene) in Argentina differs in having confluent axial parenchyma (Brea et al. 2001; Franco & Brea 2008). Several genera attributed to the Anacardiaceae have been described from the Neogene of India, viz. Buchanioxylon, Dracontomeloxylon, Holigarnoxylon, Mangiferoxylon, Swintonioxylon, Glutoxylon, Bouea, and Melanorrhoeoxylon, all differ in having more abundant axial parenchyma, usually aliform and banded (e.g., Awasthi 1965; Prakash & Tripathi 1968; Prakash & Awasthi 1969; Prakash 1978; Roy & Ghosh 1980; Agarwal 1988). Coahuiloxylon terrazasiae from the Olmos Formation (upper Campanian) of Mexico resembles both the Anacardiaceae and Burseraceae (Estrada-Ruiz et al. 2010), it differs in lacking radial canals. Méndez-Cárdenas et al. (2014) described an Oligocene species of Loxopterygium from Coayuca de Andrade, Puebla; it differs in having heterogeneous type IIB rays. Martínez-Cabrera and Cevallos-Ferriz (2004) described Tapirira peninsularis from the early Miocene of Baja California Sur, differing in having 1–3 septa per fiber, heterocellular rays with uniseriate marginal extensions of 1–12 (mostly 1–6) upright or square cells. This new genus from the Eocene of southeastern Mexico provides more evidence for and reinforces the assumption that the Northern Hemisphere was a center of diversification of Anacardiaceae (Martínez-Millán 2000), and supports the hypothesis that Mexico was one of centers of diversification of Anacardiaceae because of their extant endemism and extensive fossil record from the Late Cretaceous onwards (Martínez- Millán 2000; Ramírez & Cevallos-Ferriz 2002; Estrada-Ruiz et al. 2010).

Acknowledgments

We are grateful to Jon D. Richey (UC, Davis) for help with the English review. The authors give thanks to the anonymous reviewers for useful comments on this manuscript. We would also like to thank Dr. Rosa Andrés-Hernández from BUAP for her helpful comments, and Dr. Angélica Trujillo- Hernández from BUAP and Dr. Gabriel Villegas from IPN for providing some of the equipment we used. This research has been funded by the CONACyT (240241) and the SIP-IPN (20161352 and 20170872) grants to E.E.R.

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Accepted 16 January 2017

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