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IAWA Journal, Vol. 15 (3), 1994: 229-245 FOSSIL WOODS FROM THE EL CIEN FORMATION IN BAJA CALIFORNIA SUR: LEGUMINOSAE by Sergio R. S. Cevallos-Feniz1 and Josefina Barajas-Morales2 Summary Three types of fossil woods with similari­ Wheeler & Baas 1991, 1992) from around the ties to the Leguminosae are described, Mimo­ world. The confidence with which Legumi­ soxylon tenax (Felix) Müller-Stoll & Mädel, nosae fossil wood can be identified to generic Bajacalijomioxylon cienense Cevallos-Ferriz or specific levels contrasts with the confidence & Barajas-Morales, gen. et sp. nov., and Co­ with which fossil reproductive organs or paijeroxylon matanzensis Cevallos-Ferriz & leaves of this family are compared to extant Barajas-Morales, sp. nov. These woods are plants. While differentiation of the three sub­ from the EI Cien Formation in Baja Califor­ families (Caesalpinioideae, Papilionoideae, nia Sur, Mexico, which is dated as Zemor­ and Mimosoideae) in Leguminosae occurred rian-Saucesian, i.e., late Oligocene-early by the Eocene (e.g., Herendeen & Dilcher Miocene. Although two of the names of the 1991a, 1991b; Herendeen et al. 1992), the fossil woods suggest affinity with a particu­ assignment of wood remains to a particular lar extant taxon, differences in some quanti­ subfamily often is difficult. Even Pliocene or tative and qualitative features preclude their Pleistocene woods with affinities to the Legu­ identification with a single extant taxon. The minosae usually cannot be assigned to a sin­ similarity among wood of some groups of gle extant genus (e.g., Müller-Stoll & Mädel extant Leguminosae and limited knowledge 1967; Gros 1991, 1992). of character variability in woods of this fam­ The first putative Leguminosae wood col­ ily explains this taxonomie uncertainty. These Iected from the Tertiary of Mexico was Haurea fossil woods from Baja California underscore americana Unger from sediments near Papant­ the need for an extensive systematic study of la, Veracruz (Unger 1845, 1857). However, the wood anatomy of Leguminosae, add to its preservation is poor and its affinities are the poorly known plant history of the Penin­ questionable, so it is considered a nomen sula, suggest a tropical South American in­ nudum (Müller-Stoll & Mädel 1967). Later, fluence in the fossil flora of Baja Califomia, Mimosoxylon tenax (Felix) Müller-Stoll & and indicate that the climate during the Zemor­ Mädel from Tertiary sediments near Tlacolula, rian-Saucesian was different from the xeric Oaxaca, was described (Felix & Nathorst conditions that prevail today in the area. 1899; Müller-Stoll & Mädel 1967; Gros 1991, Key words: Wood anatomy, Leguminosae, 1992; Wheeler & Baas 1991, 1992). Repro­ fossil wood, Oligocene, Miocene, Baja ductive organs of fossil Leguminosae collect­ California Sur, Mimosoxylon, Bajacalijor­ ed from Mexico include six different fruits nioxylon, Copaijeroxylon. from Tepexi de Rodriguez, Puebla, similar to Lysiloma, Mimosa, Prosopis, and Sophora Introduction (Magal16n-Puebla & Cevallos-Ferriz 1993a, b) Over 100 Tertiary Leguminosae woods among others, and Oligocene-Miocene leaflets have been reported (e.g., Müller-Stoll & of Acacia included in amber have been report­ Mädel 1967; Awasthi 1992; Gros 1991, 1992; ed from Simojovel, Chiapas (Miranda 1963). 1) Instituto de Geologia, Universidad Nacional Aut6noma de Mexico, Ciudad Universitaria, DeI. Coyoacan, 04510 Mexico, D.F. 2) Instituto de Biologia, Universidad Nacional Aut6noma de Mexico, Ciudad Univcrsitaria, DeI. Coyoacan, 04510 Mexico, D.F. Downloaded from Brill.com10/08/2021 03:21:38AM via free access 230 IAWA Journal, Vol. 15 1994 Fig. 1. Map with loeation of fossiliferous outerops. In the present study, three types of fossil fossil wood are known. Woods reported here legurne woods, Bajacalifornioxylon cienense eome from near 'Rancho Matanzas' and 'Ca­ Cevallos-Ferriz & Barajas-Morales gen. et fiada EI Canelo' loealities (Fig. 1). The former sp. nov., and Copaijeroxylon matanzensis is about 5 km northeast from the town of EI Cevallos-Ferriz & Barajas-Morales sp. nov., Cien, Baja California Sur, while the latter is and Mimosoxylon tenax (Felix) Müller-Stoll approximately 3.5 km south (Fig. 1). The EI & Mädel, are deseribed and eompared to ex­ Cien Formation has been dated as Zemorrian­ tant and fossil woods assigned to the Legu­ Saueesian or late Oligoeene-early Mioeene minosae. (27-17 million years before present) based on paleontologieal, stratigraphical and radio­ Materials and Methods metrie data (Applegate 1985). The EI Cien Formation outerops around These fossil woods are preserved as siliea and to the east of the town of EI Cien, Baja permineralisations. All wood sampies were California Sur, and eonsists of alternating cut into small slabs and transverse, tangen­ shales, sandstones, and limestones with tial, and radial seetions prepared using the tuffaeeous and diatomaeeous interealations standard thin seetion teehnique. Cell dimen­ (Fig. 1). At least 15 different wood types are sions given in the descriptions are based on known from the EI Cien Formation in Baja 25 measurements for eaeh eharaeter, eell mea­ California Sur, Mexico. Of the over 50 wood surements include cell walls, and terminology sampies eolleeted so far, five have Legumi­ follows the IAWA feature list (IAWA nosae affinity. Several loealities eontaining Committee 1989). All fossil speeimens are Downloaded from Brill.com10/08/2021 03:21:38AM via free access Cevallos-Ferriz & Barajas-Morales - Fossil Leguminosae from Baja California Sur 231 housed in the Museo de Paleontologfa of the Paratracheal parenchyma vasicentric to ali­ Instituto de Geologfa, UNAM (IGUNAM). form and confluent, and in thin marginal Holotype numbers refer to slides produced bands (Figs. 2, 3), parenchyma cells with a from a single sample. mean tangential diameter of 17 Jlm (range Extant Leguminosae woods (see Appen­ 10-20 )lm), mean radial diameter of 11 )lm dix) were compared anatomically with the (range 6-15 Jlm), mean celliength of 60 Jlm fossil material. Sections 15-25 Jlm thick (range 26-125 Jlm). Rays homocellular were cut on a sliding microtome and stained (Figs. 6-8), uniseriate rays short, up to 21 with safranin-fast green. Slides are deposited cells tall (Fig. 7), multiseriate rays 2-4-se­ in the Instituto Nacional de Investigaciones riate (Figs. 6, 7), with a mean height of 403 Forestales y Agropecuarias (INIFAP), Mexi­ Jlm (range 246-598 )lm), all rays composed co, and in the National Xilotec in the Instituto of procumbent cells (Fig. 8), 7 (range 5-10) de Biologfa, UNAM. For identification of the rays per millimetre. Vessel to ray pits similar fossil woods, besides the direct comparison to intervascular pits (Figs. 10, 11). Chains to woods of extant plants, the Guess program, up to 19 cells long of thin-walled parenchyma OPCN database and a fossil angiosperm wood cells each containing a rhomboidal crystal, database (Wheeler et al. 1986; LaPasha & scattered among other parenchyma cells (Fig. Wheeler 1987; Wheeler & Baas 1991) were 9). Wood without storied elements. used. The second specimen is quantitatively and qualitatively very similar (Table 1). Their dif­ Systematic description ferences may reflect variation within a plant or between different plants. Class: Magnoliopsida Order: Fabales Genus: Bajacalijornioxylon Cevallos-Ferriz Family: LEGUMINOSAE & Barajas Morales, gen. nov. - Type spe­ Subfamily: MIMOSOIDEAE cies: Bajacalijornioxylon cienense Ceval­ los-Ferriz & Barajas-Morales, spec. nov. Genus: Minwsoxylon Müller-Stoll & Mädel Species: Mimosoxylon tenax (Felix) Müller• Diagnosis: Diffuse-porous wood, growth Stoll & Mädel- Figs. 2-11 rings indistinct; vessels mainly solitary, in ra­ dial multiples and clusters, intervascular pits Specimens: IGM-LPB 970-981; IGM-LPB medium, alternate and vestured, oblique sim­ 148-177 and IGM-LPB 1202, 1203. ple perforation plates; non-septate libriform Material: Two small sampies, 4 cm in diam­ fibres; paratracheal parenchyma vasicentric, eter and 9 cm long are known. apotracheal parenchyma diffuse; homocellular Distinct growth rings marked by marginal rays 1-4-seriate, vessel element to ray parenchyma (Fig. 2). Diffuse-porous, on av­ parenchyma pits similar to intervascular pits; erage 8 vessels/mm 2 (range 4-12). Vessels crystals in chambered axial parenchyma cells. mainly solitary (77%) and in radial multiples W ood without storied elements. of 2 to 4 (23%; Fig. 2), circular in outline, mean tangential diameter 152 )lm (range 88- Bajacalijornioxylon cienense Cevallos-Ferriz 198 )lm), mean radial diameter 138 )lm & Barajas-Morales, spec. nov. - Figs. (range 120-230 )lm), mean vessel element 12-18 length 213)lm (range 18-305 )lm). End walls of vessel elements oblique (30°), simple per­ Holotype: IGM-LPB 1210-1213, IGM-LPB foration plates (Fig. 4); intervascular pits oval, 1238. alternate, minute (2-3 Jlm; Figs. 4, 5), and Material: A small sampie, 5 cm in diameter vestured. Thin-walled tyloses occasional in and 12 cm long. vessel elements. Non-septate thick-walled Etymology: The generic name is after the Baja fibres, oval in transverse section, mean tan­ California Peninsula where the sample gential and radial diameter 15 )lm and 12 )lm was found. The specific epithet cienense (range 6-18 Jlm and 7-15 Jlm) respectively. is after the nearby town of EI Cien. Downloaded from Brill.com10/08/2021 03:21:38AM via free access - 232 IAWA~ Journal, Vol.lS (3), 1994 Downloaded from Brill.com10/08/2021 03:21:38AM via free access Cevallos-Ferriz & Barajas-Morales - Fossil Leguminosae from Baja California Sur 233 Figs. 9-11. Mirrwsoxylon tenax. - 9: Tangential section showing axial parenchyma cells where crystals were contained. IGM-LPB 971, x 400. - 10: Radial seetion showing vessel-ray pits. IGM-LPB 971, x 500. - 11: Radial seetion showing vessel-ray pits. IGM-LPB 979, x 350. Diagnosis: Indistinct growth rings, diffuse­ 40-110 11m). Rays homocellular (Figs. 14, porous, on average 6 vessels/mm 2 (range 17), uniseriate rays common, short, up to 12 4-8). Vessels mainly (78%) solitary and in cells tall; multiseriate rays 2-4-seriate, with a radial multiples of 2 (19%), occasionally in me an height of 383 11m (range 250-730 11m; radial multiples of 3 or more (up to 7; Figs.
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