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 . 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 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.

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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: 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.

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~ - IAWA 232 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. Fig. 14) all rays composed of procumbent 12,13), oval in outline, mean tangential diam­ cells (Fig. 17), 10 (range 9-12) rays per mil­ eter 141 11m (range 93-210 11m), mean radial limetre. Vessel to ray parenchyma pits similar diameter 99 11m (range 40-190 11m), average to intervascular pits. Chains up to 14 cells vessel element length 320 11m (range 210-450 long of thin-walled parenchyma cells contain­ 11m). End walls of vessel elements oblique ing a rhomboidal crystal each scattered among (40°), with simple perforation plates (Fig. 16); fibres (Fig. 15). intervascular pits oval, alternate, medium (4- 7 11m), and vestured (Fig. 18). Fibres thick­ Subfamily: CAESALPINIOIDEAE walled (Fig. 13), oval in transverse seetion, non-septate, mean tangential and radial diam­ Genus: Copaijeroxylon Müller-Stoll & Mädel eter 10 11m and 7 Ilffi (range 9-12 Ilffi and 6-9 1967 11m), respectively. Up to four cells thick vasi­ Species: Copaiferoxylon matanzensis Ceval- centric parenchyma (Figs. 12, 13), some dif­ 10s-Ferriz & Barajas-Morales, spec. nov. fuse apotracheal parenchyma; parenchyma - Figs. 19-29 cells mean tangential diameter 12 11m (range 7-13 11m), mean radial diameter 6 11m (range Holotype: IGM-LPB 179-194. 3-7.5 11m), mean celllength 78 11m (range Paratype: IGM-LPBI95-208.

Figs. 2-8. Mirrwsoxylon tenax. - 2: Transverse seetion showing vessel and parenchyma distri­ bution. IGM-LPB 973, x 57. - 3: Transverse section showing detail of vasicentric parenchyma and thin-walled fibres. IGM-LPB 973, x 138. - 4: Radial section showing vessel element with simple perforation plates and alternate intervascular pits. IGM-LPB 977, x 367. - 5: Radial sec­ tion showing alternate intervascular pits. IGM-LPB 981, x 885. - 6: Tangential seetion with uni­ and multiseriate rays, vasicentric parenchyma, and vessel elements with alternate intervascular pits. IGM-LPB 974, x 138. - 7: Tangential seetion showing uni- and multiseriate rays. IGM-LPB 974, x 690. - 8: Radial seetion showing homocellular rays. IGM-LPB 971, x 126.

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Material: Two sm all wood sam pies known, 325-978 11m), heterocellular, composed of 18 cm long by 5 cm in diameter and 10 procumbent cells in their main body with up cm long by 4 cm in diameter, respectively, to 3 marginal rows of upright cells (Fig. 26), with the same anatomical pattern. 7 (range 5-9) rays per millimetre. Vessel to Etymology: The specific epithet matanzensis ray parenchyma pits similar to intervascular is after the nearby ranch Rancho Matanzas. pits (Figs. 24, 28). Chains up to 10 cells long of thin-walled parenchyma cells con­ Diagnosis: Growth rings and axial paren­ taining a rhomboidal crystal scattered among chyma visible to the naked eye, limit between fibres (Fig. 29). Wood without storied ele­ one growing season and the next one is mark­ ments. ed by a band, 1- 6 cells wide, of marginal pa­ renchyma (Fig. 19). Around the axial canals Discussion the parenchyma bands are up to 10 cells wide Within the sam pies, there is little variation (Fig. 19). Diffuse-porous, on average 5 ves­ in the amount and type ofaxial parenchyma, sels/mm 2 (range 2-6; Figs. 19, 20). Ves­ ray structure, and dimensions and frequen­ sels solitary (84%; Fig. 19) accompanied by cy of tracheary elements, all of which sug­ radial multiples of two (11%), and three gests mature wood structure (Carlquist 1975, (3%), or clusters (2%), alm ost circular in 1988). However, it is not possible to say if outline, mean tangential diameter 104 11m the sampies are trunk or branch wood, al­ (range 87-128 11m), mean radial diameter though their small diameter suggests that they 128 11m (range 70-150 11m); average vessel are branches. element length 291 11m (range 178-321 11m). An important character in determining af­ End walls of vessel elements oblique (25°), finities of the fossil material is presence of simple perforation plates (Fig. 22); intervas­ vestured pits. Although it is difficult to rec­ cular pits oval, small (5-7 11m), alternate and ognise vestured pits in permineralised wood, vestured (Fig. 23). Non-septate, thin-walled intervascular pit apertures of nine undescribed fibres (Figs. 19, 20); many-sided in trans­ woods from the same localities from which verse section, mean tangential diameter 16 these Leguminosae woods were collected 11m (range 5-20 11m), mean radial diameter have smooth pit apertures in contrast with the 19 11m (range 13-25 11m), small simple pits tufted or verrucose pit apertures in the woods in the radial walls (Figs. 27-29). Paratra­ reported here. The presence of vestured pits cheal parenchyma aliform to confluent and eliminates many families (e.g., Rhamnaceae, marginal, and apo tracheal parenchyma diffuse Rutaceae) from further comparisons. How­ (Figs. 19,20). Aliform parenchyma winged, ever, there were a few taxa in other families individual parenchyma cells 18 x 23 x 98 11m. with vestured pits (e.g., Vochysiaceae and Uniseriate rays common, up to 12 cells tall Dipterocarpaceae) with anatomical structure (Fig. 21), heterocellular rays (Figs. 25, 26) similar to the Baja California Sur material. composed of procumbent cells in their main The absence of marginal parenchyma dis­ body and one row of upright cells in the mar­ tinguished Vochysia and Qualea (Vochysia­ gins. Multiseriate rays 2- or 3-seriate (Fig. ceae) from Copaijeroxylon matanzensis (Met­ 21), with a mean height of 528 11m (range calfe & Chalk 1950). Shorea (Dipterocarpa-

Figs. 12-18. Bajacalifornioxylon cienense. - 12: Transverse section showing vessel and paren­ chyma distribution. IGM-LPB 1238, x 60. - 13: Transverse section showing detail of parenchyma and thin-walled fibres. IGM-LPB 1211, x 85. - 14: Tangential section with multiseriate rays, and vessel elements with simple perforation plates. IGM-LPB 1212, x 80. - 15: Tangential sec­ tion with multiseriate rays, and crystals. IGM-LPB 1212, x 210. - 16: Tangential section show­ ing vessel elements with simple perforation plates. IGM-LPB 1213, x 228. - 17: Radial section with multiseriate homocellular ray. IGM-LPB 1210, x 92. - 18: Tangential seetion showing de­ tail of alternate pits. IGM-LPB 1213, x 428.

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Figs. 26-29. Copaijeroxylon matanzensis. - 26: Radial seetion showing detail of heterocellular ray.IGM-LPB 191, x 270. - 27: Radial seetion showing detail of fibre pits and crystals in paren­ chyma. IGM-LPB 191, x 190. - 28: Radial seetion showing vessel-ray pits. IGM-LPB 191, x 220. - 29: Radial section showing detail of pitted fibres and crystals. IGM-LPB 191, x 305.

ceae) is also comparable to C. matanzensis Members of the three Leguminosae subfam­ and Bajacalifornioxylon cienense, but it has ilies (Caesalpinioideae, Mimosoideae, Papil­ enlarged vessel element to ray parenchyma ionoideae) are comparable to the wood struc­ pits, while in the Mexican woods these pits ture of the EI Cien Formation plants (Metcalfe are more or less the same size (Awasthi 1975 & Chalk 1950; Müller-Stoll & Mädel 1967; [1977]). The absence of apotracheal paren­ Baretta-Kuipers 1981; Wheeler & Baas 1992). chyma in Qualea contrasts with the presence However, each wood from Baja Califomia of diffuse parenchyma in Mimosoxylon tenax Sur has a set of characters that relates them to a (Metcalfe & Chalk 1950). particular small number of Leguminosae taxa.

Figs. 19-25. Copaijeroxylon matanzensis. - 19: Transverse seetion showing vessel, axial canals and parenchyma distribution. IGM-LPB 179, x 50. - 20: Transverse seetion showing detail of aliform parenchyma, axial canals, and thin-walled fibres. IIJM-LPB 181, x 110.- 21: Tangential seetion with uni- and multiseriate rays, and vessel with alternate intervascular pits. IGM-LPB 185, x 55. - 22: Radial seetion showing vessel element with simple perforation plate, alternate intervascular pits, and parenchyma cells with vessel element-parenchyma pits. IGM-LPB 191, x 180. - 23: Radial seetion showing detail of intervascular pits. IGM-LPB 191, x 1150. - 24: Radial seetion showing detail of vessel-ray pits. IGM-LPB 190, x 430. - 25: Ra­ dial seetion showing general view of heterocellular rays. IGM-LPB 186, x 90.

Downloaded from Brill.com10/08/2021 03:21:38AM via free access N W Table 1. Comparison of Mimosoxylon tenax from Baja California Sur and Oaxaca with extant Mimosa arenosa, Chlorophora and Haematoxylon. 100

M. tenax M. tenax M. tenax Mimosa Chlorophora Haematoxylon arenosa tinctoria brasiletto IGM-LPB 970-981 IGM-LPB 148-177, 1202, 1203

(Baja Califomia Sur) (Baja California Sur) (Oaxaca) (Jalisco) (Jalisco) (Jalisco)

Vessels per sq.mm 7 12 8-24 19 6 11

Vessel grouping solitary, solitary, solitary, solitary, solitary, sOlitary, radial multiples radial multiples radial multiples radial multiples radial multiples radial multiples (2-5) (2-4) (2-4) (2-4) (2-5) (2-3)

Mean vessel element length in JlIIl 189 213 178 228 223

Mean tangential diameter of vessels in !lIIl 160 152 88 121 144 89

Intervascular pit size in JlIIl 4-5 2-3 4-5 4-6 3-7 (coalescent) ( coalescent) (coalescent) (coalescent) 1- >- ,:E! Downloaded fromBrill.com10/08/2021 03:21:38AM Axial parenchyma arrangement vasicentric, vasicentric, vasicentric, vasicentric, vasicentric, aliform, >- confluent aliform, aliform, aliform, aliform, confluent, ..... confluent, confluent, confluent, marginal marginal c0 marginal marginal 00nded 3 ~ Rays per linear mm 7 7 7 9 8 < f?...... Ray width in cell number up to 4 up t04 up to 3 up to 3 up to 3 up to 4 Vl

Mean ray height in lUD 620 403 460 320 675 900 ~ ......

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Mimosoxylon millimetre in M. calpocalycoides Lemoigne, Mimosoxylon tenax Müller-Stoll & Mädel high rays in M. grandiporosum Pons, very compares positively to some species of Pip­ small pores and 2-seriate rays in M. mbakaou­ tadenia, Leucaena (Mimosoideae) and Caes­ ense Duperon-Laudoueneix, and distinct alpinia (Caesalpinioideac; Wheeler et al. 1986; growth rings, septate fibres and ray width in LaPasha & Wheeler 1987). The main differ­ M. santamariensis Lutz distinguish them from cnce among these taxa is thc size of the inter­ the plants of Baja California Sur (Kramer vascular pits, which are smaller in thc woods 1974; Lemoigne 1978; Pons 1983; Lutz 1987). of Baja California Sur. Therc is, however, a vcry close similarity of the fossil to Mimosa Bajacalifornioxylon arenosa (Willd.) Poir., Chlorophora and Hae­ Although woods of Melanoxylon brauna matoxylon (Table 1; Chehaibar & Grether (Caesalpinioideae) and Myroxylon balsamum 1990). Although diffcrences from Chloro­ (Papilionoideae) have so me similarity with phora and Haematoxylon are quantitative, it Bajacalifornioxylon cienense, the presence of is clear from Table 1 that M. tenax is most storied elements (rays and axial parenchyma) similar to M. arenosa. Among extant Legu­ in these extant species contrasts with the un­ minosae, the absence of storicd elements and storied wood structure of the fossil plant thc presence of homocellular rays (Baretta­ (Wheeler et al. 1986; LaPasha & Wheeler Kuipers 1981) in Mimosoxylon tenax sup­ 1987). The wood of Prioria copaijera (Caes­ port a relationship to Mimosoideae. Within alpinioideae) is also similar to that of B. cie­ this subfamily, Mimosa appears most similar nense, however, the presence of heterocellu­ to the fossil plants (Chehaibar & Grether lar rays and axial canals in the former contrast 1990). In the absence of other fossil organs with the homocellular rays and the absence supporting this relationship it would be pre­ ofaxial canals in the wood of Baja Califor­ mature to assign this fossil wood to M. are­ nia Sur. Among the wood of extant mimo­ nosa; however, a close relationship to Mimo­ soid plants that compare closely to B. cie­ soideae, especially to Mimosa is weH sup­ nense are: Ebenopsis flexicaulis, Havardia ported. sp., Klugiodendron laetum, Marmoxylon The EI Cien Formation Mimosoxylon is sp., Pithecellobium sp. (Wheeler et al. 1986; quantitatively and qualitatively very similar to LaPasha & Wheeler 1987), and Acacia me­ M. tenax supponing its identification as a lanoxylon (De la Paz Perez-Olvera et al. member of this taxon, previously described 1980; Cevallos-Ferriz & Carmona-Valdo­ from Tertiary sediments near TIacolula, Oaxa­ vinos 1981). There is, however, a tendency ca, Mcxico (Fe1ix & Nathorst 1899; Müller• in the wood of the first five p1ants to have Stoll & Mädel 1967; Table 1). The genus either uniseriate rays and/or aliform paren­ Mimosoxylon includes 9 species (Gros 1992). chyma. In contrast, B. cienense has up to The main differcnce between Mimosoxylon 4-seriate rays and one cell thick vasicentric tenax and other Mimosoxylon species is the parenchyma. Acacia melanoxylon is most absence in the Baja California Sur wood of a similar to B. cienense, but the wood of the tendency to storied axial parenchyma (MüHer• former has more abundant parenchyma and Stoll & Mädel 1967). However, in revicwing sm aller vessel elements than the wood of EI the descriptions of the species belonging to Cien Formation (De la Paz Perez-Olvera et al. Mimosoxylon, this tendency is not always 1980; Cevallos-Ferriz & Carmona-Valdovi­ present, and Baretta-Kuipers (1981) suggests nos 1981). that storied wood is not typical of Mimosoi­ Some taxa of Leguminosae fossil woods deae. Large intervascular pits in M. ducis­ are comparable to B. cienense. However, the aprutii (Chiarugi) Müller-Stoll & Mädel, het­ presence of weIl developed marginal paren­ erogeneous rays in M. piptadenioides (Da chyma (e.g., Albizia vantagiensis Prakash Silva Curvello) Müller-Stoll & Mädel, 1- or- & Barghoorn, Dichrostachyoxylon zirkelli 2-seriate rays in M. acacioides Kramer and (Felix) Müller-Stoll & Mädel, Paraalbizioxy­ M. krameri Lemoigne, weIl developed ali­ Ion bavaricum (Selmeier) Gros, Isoberlini­ form parenchyma and numerous rays per oxylon congoense Lakhanpal & Prakash, and

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Table 2. Leguminosae with axial canals. Data from Metcalfe & Chalk (1950), Baretta-Kuipers (1981), and Gasson (1994).

Genus Canals Canals Parenchyma Rays diffuse (D) / normal (N) or paratracheal (S) or uniseriate (U) tangential (T) traumatic (T) aliform (A) and/ar storied (S)

Daniella D T A Gossweilerodendron D T A S Kingiodendron D T A U/S Oxystigma D T A S Prioria D T A S Bathiaea T N S Copaijera T N S U Detarium T T A Pseudosindora T N S U Sindora T N S Sindoropsis T N S Eperua T N S

Tetrapleuroxylon limagnense Prive), as well renchyma. Metacacioxylon was introduced as septate fibres, heterogeneous rays, and by Gros (1981, 1984, 1988) to include woods confluent parenchyma, contrasts with B. cie­ similar to Acacia but with rays being (1-)7-12 nense. (-18)-seriate. In contrast, rays in B. cienense Acacioxylon is the form-genus proposed are 1-3(-4)-seriate. While similarity of the by Schenk (1883) to include fossil woods wood of Bajacalijornioxylon cienense to that similar to the secondary xylem of Acacia. of extant Acacia is dose, recognition of a new However, reassignment of most fossil spe­ taxon is supported by the quantitative and cies in this taxon to Euacacioxylon and Par­ qualitative differences found among wood acacioxylon by Müller-Stoll & Mädel (1967) form-genera comparable to the wood of Aca­ and its recent use to name some fossil woods cia and by the similarities shared by the EI from Chile (Nishida 1981, 1984) has created Cien wood and Acacia melanoxylon. nomenclatural problems (Gros 1992). Both Euacacioxylon and Paracacioxylon have con­ Copaijeroxylon fluent and vasicentric parenchyma with a ten­ The presence ofaxial canals in Legumi­ dency to become aliform, their rays are 1-10- nosae is a rare character, restricted to Caesal­ seriate, and the latter has septate fibres (Mül• pinioideae (Metcalfe & Chalk 1950; Baretta­ ler-Stoll & Mädel 1967). In contrast, B. cie­ Kuipers 1981; Gasson 1994; Table 2). These nense has only vasicentric parenchyma, 1-3 can be normal canals arranged in tangential (-4)-seriate rays, and no septate fibres. Baja­ bands, as in Copaijeroxylon matanzensis, or calijornioxylon cienense, although similar to scattered, or traumatic canals in tangential extanl Acacia, differs from Acacioxylon and bands (Metcalfe & Chalk 1950; Gasson 1994). an Eocene Acacia in axial parenchyma distri­ Five genera in Caesalpinioideae have normal bution (Kräusel 1939; Navale 1962 [1963]; axial canals in tangential bands, Bathiaea, Gregary 1970; Gros 1992; Nishida 1984). Copaijera, Eperua, Sindora, and Sindoropsis Whilc Acacioxylon and the Eocenc Acacia (Baretta-Kuipers 1981; Gasson 1994; Tab1e havc paratracheal (vasicentric, aliform and 2). All of these genera, except Detarium, are cOnnUenl) and uniseriate apotracheal bands, characterised by having mostly scanty para­ B. cienense lypically has up to 4 cells thick tracheal parenchyma, although sometimes it vasicentric and some diffuse apo tracheal pa- tends to be aliform (Metcalfe & Cha1k 1950;

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Baretta-Kuipers 1981), in contrast with the (Fessler-Vrolant 1977). They are similar to mostly aliform and marginal parenehyma of C. matanzensis not only in the presence and the new fossil wood. Detarium lacks uniseri­ distribution ofaxial canals, but also in the ate rays, but has vasicentric, mostly aliform, type ofaxial parenchyma, and vessel element and also clearly demarcated lines of paren­ distribution and grouping. There are, how­ chyma on growth ring boundaries, while ever, minor differences among them. Vasi­ Pseudosindora further differs from the fossil centric parenchyma is more common in C. plant in having uniseriate rays (Baretta-Kui­ migiurtinium and C. copaiferoides (Fessler­ pers 1981). Eperua is also distinct in having Vrolant 1977); in contrast, aliform parenchy­ banded parenchyma (Metcalfe & Chalk 1950). ma is more common in C. matanzensis. Rays Distinction between the other genera, Bath­ tend to be slightly wider in C. migiurtinium iaea, Copaifera, Sindora, and Sindoropsis, (1-5-seriate) than in the new specics (1-3- with qualitative characters is difficult; how­ seriate); ray parenchyma cells in the former ever, there is a closer qualitative similarity of are erect while in the latter these are mainly the fossil plant to Copaifera (Metcalfe & procumbent; rays are homocellular in the Chalk 1950; Moens 1955; Wheeler et al. 1986; Somalian material and heterocellular in the LaPasha & Wheeler 1987). Baja California Sur specimens (Müller-Stoll Among the fossil Leguminosae genera & Mädel 1967). Copaiferoxylon copaiferoi­ known to the authors, eight species have des is also distinct from C. matanzensis by axial canals and aliform and marginal paren­ having radial canals and crystals in parenchy­ chyma: Acada gregorii Gregory, Caesl1lpin­ ma cells (Fessler-Vrolant 1977). Similarity oxylon moragjonesiae Crawley, Copaifer­ between these woods is close enough as to oxylon copaiferoides Fessler Vrolant, C. mi­ include the new wood in the same genus. giurtinium (Chiarugi) Müller-Stoll & Mädel, However, differences in age and 10cality, as Detarioxylon aegyphacum (Unger) Louvet, weIl as the qualitative and quantitative differ­ Erythrophloeoxylon scholleri (Boureau) Mül• ences between the wood of the three plants, ler-Stoll & Mädel, Hopeoxylon indicum Na­ support the designation of a new species, vale, and Kingiodendron prebinnatum Awas­ C. matanzensis Cevallos-Ferriz & Barajas­ thi & Prakash. However, axial canals in Aca­ Morales. da gregorii, Caesalpinoxylon moragjonesiae, and Erythrophloeoxylon scholleri are of the Subfamily ajfinities traumatic type (Gregory 1971 [1973]), while Müller-Stoll and Mädel (1967) reviewed in Copaiferoxylon matanzensis they are nor­ the problems of classifying fossil woods with mal. Kingiodendron prebinnatum has normal affinity to the Leguminosae. Our study sup­ axial canals, but their diffuse arrangement ports their idea of classifying Leguminosae contrasts with the banded pattern of C. matan­ fossil wood using taxa that compare to one or zensis (Awasthi & Prakash 1986 [1987]). more extant taxa, sometimes even from dif­ Detarioxylon aegyphacum differs in having ferent subfamilies. Reviews such as Gros's broader rays (4-1O-seriate vs 1-3-seriate), (1991, 1992) for Mimosoideae are important some enlarged vessel element to ray paren­ in evaluating the status of fossil Leguminosae chyma pits (vs similar pits), and more vasi­ woods. However, new and more complete centric parenchyma than C. matanzensis. wood descriptions of extant plants will help Hopeoxylon indicum also has some enlarged in reviewing the limits of fossil taxa. There­ vessel element to ray parenchyma pits (vs fore, we suggest that there be more systematic similar pits), and more vasicentric parenchy­ studies of extant woods of Leguminosae. ma, but differs further from C. matanzensis Comparison of the generic diagnoses of in having more vessels per square millimetre Müller-Stoll and Mädel (1967) for fossil (Awasthi 1975 [1977]). Lcguminosac woods and the systcmatic rc­ Copaiferoxylon is known from two spe­ view of Mimosoideae wood by Gros (1991, eies, C. migiurtinium from the Miocene of 1992) show litde difference between some Somalia (Müller-Stoll & Mädel 1967) and C. taxa. Both reviews suggest that the amount copaiferoides from the Oligocene of Tunisia and distribution ofaxial parenchyma, the

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relative size of vessel element to ray paren­ Califomia Sur will be discussed in a later chyma pits, presence of septate fibres and paper, along with other woods from the EI storied elements are important characters in Cien Formation. However, the ineonspieu­ delimiting fossil Leguminosae genera. Al­ ous growth rings, relatively abundant axial though these characters may in fact be useful parenehyma, and relatively wide vessels sug­ to identify fossil Leguminosae woods, cau­ gest that the fossil flora inhabited a relatively tion is needed. The reviews of the family by humid environment without pronouneed sea­ Cozzo (1950), Reinders-Gouwentak (1955) sonality (cf. Barajas-Morales 1985). and Baretta-Kuipers (1981) point out impor­ tant differences between Mimosoideae and Acknowledgments the combined Caesalpinioideae and Papilion­ This work was supported in part by Diree­ oideae, suggesting that additional characters, ei6n General de Asuntos dei Personal Aeade­ like number of cells in a parenchyma strand mieo (DGAPA-UNAM) projeet IN201291 and ray structure, mayaiso be useful in de­ and Consejo Naeional de Ciencia y Tecno­ termining relationships among fossil and ex­ logfa (CONACYT) projeet 0450T to SRSCF. tant woods of Leguminosae. Field work was partly sponsored by the Uni­ versity of Baja Califomia Sur. We thank Dr. Biogeography Shelton P. Applegate, Instituto de Geologfa, Extant plants with wood similar to Mi­ UNAM, for information on wood loealities mosoxylon (Acacia, Mimosa, Hardwickia, and helpful diseussion on the geology of the Bussea and Sesbania) are mainly restricted to area, and Laboratorio de Mieroeine, Faeultad tropieal South America, although some taxa de Ciencias, UNAM, and Mr. Antonio Alta­ are weil represented in Mexico (e.g., Acos­ mira for photo graphie assistanee. Authors mium, Acacia, and Mimosa), while others are gratefully aeknowledge review of the manu­ found in India (e.g., Hardwickia) or tropical seript by Dr. Elisabeth Wheeler and two anon­ Africa (e.g., Bussea; PoihilI & Raven 1981; ymous reviewers. Gros 1992). Presence of South American plants in the EI Cien Formation is further suggested by the presence of Copaiferoxylon References in Baja Califomia Sur. Extant taxa with wood Applegate, S. 1985. The EI Cien Formation; similar to Copaijeroxylon are restrieted to strata of Oligoeene and early Mioeene age tropical Afriea, tropical South Ameriea and in Baja Califomia Sur. Rev. Inst. Geo!. Asia; and Copaifera, which is most similar to UNAM 6: 145-162. Copaiferoxylon matanzensis grows in tropical Awasthi, N. 1975 (iss. 1977). Revision of South America and Africa (Polhill & Raven Hopeoxylon indicum Navale and Shoreo­ 1981). Although Bajacalifornioxylon is sim­ xylon speeiosum Navalc from the Cudda­ ilar to Acacia, a pantropieal genus, it could be lore Series near Pondieherry. The Palaeo­ related to Acaciella, a group represented in botanist 24: 102-107. Mexico, the Caribbean and South America, Awasthi, N. 1992. Indian fossillegumes. In: and whose wood anatomy remains to be P. S. Herendeen & D. L. Dilcher (eds.), studied. In this regard it is significant that the Advanees in Legume Systematies: Part 4. Oligocene flower from Simojovel, Chiapas, The Fossil Record: 225-238. Royal Bot. may be Acacie lla rather than Acacia (Miranda Gard., Kew. 1963; Sousa & Delgado 1993). Awasthi, N. & U. Prakash. 1986 (iss. 1987). These Leguminosae woods support the Fossil woods of Kingiodendron and Bau­ idea of an influence of the South American hinia from the Namsang beds of Deomali, flora in the EI Cien Formation; however, Arunaehal Pradesh. The Palaeobotanist 35: wh oie plant reconstruetions are needed in 178-183. order to fully understand the phylogenetie Barajas-Morales, J. 1985. Wood structural and phytogeographie importance of these differences between trccs 01' two tropical woods. The functional-ecological significance forests in Mexico. IAWA Bull. n.s. 6: of the fossil Leguminosae plants from Baja 355-364.

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Baretta-Kuipers, T. 1981. Wood anatomy in Gros, J.P. 1981. Nouveaux bois du Ceno­ Leguminosae: Hs relevance to . zoique d' Autriche et d'Ethiopie. These In: R.M. PolhilI & P.H. Raven (eds.), 3eme Cycle, n. 1068, 2 t., 143 pp. Univ. Advances in Legume Systematics: Part 1: Claude Bemard Lyon. 677-705. Royal Bot. Gard., Kew. Gros, J.P. 1984. Etude comparative de 4 Carlquist, S. 1975. Ecological strategies of echatillones de bois fossiles de l'Eggen­ xylem evolution. Univ. Califomia Press, burigien d' Autriche, rapportes au nouveau Berkeley. genere Metacacioxylon n. g. et aux es­ Carlquist, S., 1988. Comparative wood anat­ peces M. marglii n. sp. et M. lemoignei omy. Systematic, ecological and evolution­ n. sp. Rev. Gen. Bot. 91: 35-80. ary aspects of dicotyledon wood. Springer­ Gros, lP. 1988. Nouveau specimen de bois Verlag, Berlin. de l'Oligocene d' Autriche rapporte a l'es­ Cevallos-Ferriz, S. R. S. & T. F. Carmona­ pece Metacacioxylon lemoignei Gros 1981 Valdovinos. 1981. Banco de informaci6n emend. Nouv. Arch. Mus. Hist. Nat. Lyon de estudios tecnol6gicos de maderas que 26: 19-27. vegetan en Mexico. Inst. Nac. Investig. Gros, J.P. 1991. Les bois fossiles de Mimo­ Forestales, Catalogo No. 3. 150 pp. soideae. Inventaire systematique. Revue Chehaibar, T. & R. Grether. 1990. Anatomfa de Paleobiologie 10: 47-60. de algunas especies de Mimosa (Legumi­ Gros, lP. 1992. A synopsis ofthe fossil rec­ nosae). Bol. Soc. Bot. Mexico 50: 3-17. ord of mimosoid legume wood. In: P. S. Cozzo, D. 1950. Anatomfa deI leno secun­ Herendeen & D.L. DiIcher (eds.), Ad­ dario de la leguminosas mimosoideas y vances in Legume Systematics: Part 4: cesalpinoideas Argentinas. Rev. Inst. Nac. 69-83. Royal Bot. Gard., Kew. Invest. Ci. Nat. II-2. Herendeen, P.S., W.L. Crepet & D.L. Dil­ De la Paz Perez-Olvera, C., T.F. Carmona­ cher. 1992. The fossil history of the Legu­ Valdovinos & M.A. Rogel-G6mez. 1980. minosae: Phylogenetic and biogeographic Estudio anat6mico de la madera de 43 es­ implications. In: P.S. Herendeen & D.L. pecies tropicales. Instituto Nacional de In­ Dilcher (eds.), Advances in Legume Sys­ vestigaciones Forestales, Boletfn Tecnico tematics: Part 4. The Fossil Record: 303- No.43. 276 pp. 316. Royal Bot. Gard., Kew. Felix, l & A. Nathorst. 1899. Versteinerun­ Herendeen, P.S. & D.L. DiIcher. 1991a. gen aus dem mexikanischen Staat Oaxaca. Fossil mimosoid legumes from the Eocene Fossile Hölzer von Tlacolula. In: J. Felix and Oligocene of southeastern North Arn­ & A. Lenk, Beiträge zur Geologie und erica. Rev. Pa1eobot. Palyn. 62: 339-361. Paläontologie der Republik Mexiko 2: 46- Herendeen, P.S. & D.L. DiIcher. 1991b. 51 Leipzig. Caesalpinia subgenus Mezoneuron (Legu­ Fessler-Vrolant, C. 1977. Sur la presence minosae, Caesalpinioideae) from the Ter­ d'un bois de Legumineuses dans l'Oligo­ tiary of North America. Arner. J. Bot. 78: cene des Monts de Teboursouk (Tunisie). 1-12. Notes Servo Geol. Repub. Tunisie 43: IAWA Committee. 1989. IAWA list of mi­ 139-151. croscopic features for hardwood identifi­ Gasson, P. 1994. Wood anatomy ofthe tribe cation. IAWA BuH. n.s. 10: 219-332. Sophoreac and related Caesalpinioideae Kramer, K. 1974. Die tertiären Hölzer süd• and Papilionoideae. In: LK. Ferguson & ost-asiens (unter Ausschluss der Diptero­ S. Tucker (eds.), Advances in Legume carpaceae) 1. Palaeontographica Abt. B, Systematics: Part 8. Structural Botany. 144: 45-181. Royal Bot. Gard., Kew. Kräusel, R. 1939. Ergebnisse der Forschungs­ Gregory, L 1970. An ancient Acacia wood reisen Prof. E. Stromer's in den Wüsten from Oregon. The Ore Bin 32: 205-210. Ägyptens. IV. Die fossilen Floren Ägyp­ Gregory, L 1971 (iss. 1973). An ancient Aca­ tens, 3. Die fossilen Pflanzen Ägyptens. cia wood from Oregon. The Palaeobota­ Abh. boyer. Akad. wiss., math-nat., Abt., nist 20: 19-21. N.F. 47: 1-140.

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LaPasha, c.A. & E.A. Wheeler. 1987. A mi­ Nishida, M. 1984. The anatomy and affinit­ crocomputer based system for computer­ ies of the petrified plants from the Tertiary aided wood identification. IAWA Bull. of Chile. III. Petrified woods from Mocha n.s. 8: 347-354. Island, Central Chile. In: M. Nishida (ed.), Lemoigne, Y. 1978. Flores tertiaires de la Contributions to the Botany of the Andes: haute vallee de rOmo (Ethiopie). Palaeon­ 96-110. Academia Scientific Book Inc., tographica Abt. B, 165: 89-157. Tokyo. Lutz, A.I. 1987. Estudio anat6mico de ma­ PolhilI, R.M. & P.H. Raven (eds.). 1981. deras terciarias deI valle de Santa Matia Advances in Legurne Systematics: Part 1. (Catamarca-Tucuman), Argentina. Facena, Royal Bot. Gard., Kew. Corrientes Argentina 7: 126-143. Pons, D. 1983. Contribution a l'etude paleo­ MagaIl6n-Puebla, S. & S. R. S. Cevallos­ botanique du Mesozoique et du Cenozo­ Ferriz. 1993a. Fossil legurne pods from ique de Colombie. These Dr. Etat, 655 the Upper Cenozoic Pie de Vaca Forma­ pp., Univ. Pierre et Marie Curie. Paris. tion, Puebla, Mexico. Abstracts XV In­ Reinders-Gouwentak, c.A. 1955. The sto­ ternational Botanical Congress, Tokyo: ried-structure features and the taxonomie 240. rank of the leguminous taxa. Acta Bot. MagaIl6n-Puebla, S. & S. R. S. Cevallos­ Neerl. 4: 460-470. Ferriz. 1993b. Fossil mimosoid pods from Schenk, A. 1883. Fossile Hölzer. In: K. Zit­ the Upper Cenozoic Pie de Vaca Forma­ tel, Beiträge zur Geologie und Paläonto• tion, Puebla, Mexico. Abstracts XV In­ logie der Lybischen Wüste. II Abt. 1. Pa­ ternational Botanical Congress, Tokyo: laeontographica 30: 1-19. 240. Sousa, M. & A. Delgado. 1993. Mexican Le­ Metcalfe, C.R. & L. Chalk. 1950. Anatomy guminosae: Phytogeography, endemism, of the dicotyledons. 2 Vols. Clarendon and origins. In: T.P. Ramamoorthy, R. Press, Oxford. Bye, A. Lot & J. Fa (eds.), Biological di­ Miranda, F. 1963. Two plants from the am­ versity of Mexico. Origins and distribution: ber of Simojovel, Chiapas, Mexico, area. 459-511. Oxford Univ. Press, New York. 1 Paleontol. 37: 611-614. Unger, F. 1845. Synopsis Plantarum Fos­ Moens, P. 1955. Les formations secretrices silium. Leipzig. des capiliers Congolais. Cellule 57: 35- Unger, F. 1857. Beiträge zur näheren Kennt­ 64. nis des Leithakalkes. Denksehr. Akad. Müller-Stoll, W.R. & E. Mädel. 1967. Die Wiss. 14: 13-38. fossilen Leguminosen-Hölzer. Eine Revi­ Wheeler, E.A. & P. Baas. 1991. A survey 01' sion der mit Leguminosen verglichenen the fossil wood record for dicotyledonous fossilen Hölzer und Beschreibung älterer wood and its significance for evolution­ und neuer Arten. Palaeontographica 119B: ary and ecological wood anatomy. IAWA 95-174. Bull. n.s. 12: 275-332. Navale, G.K.B. 1962 [iss. 1963]. Fossil Wheeler, E.A. & P. Baas. 1992. Fossil wood woods of Leguminosae from the Tertiary of the Leguminosae: A case study in xy­ rocks of the Cuddalore series near Pondi­ lem evoution and ecological anatomy. In: cherry, India. The Palaeobotanist 11: 54- P. S. Herendeen & D. L. Dilcher (eds.), 65. Advances in Legurne Systematics: Part 4. Nishida, M. 1981. Petrified woods from thc The Fossil Record: 303-316. Royal Bot. Tertiary of Mocha Island (a preliminary Gard., Kew. report). In: M. Nishida (ed.), Areport of Wheeler, E. A., R. Pearson, C. A. LaPasha, the paleobotanical survey of southern Chile W. Hatley & T. Zack. 1986. Computer­ by a Grant-in-Aid for Overseas scientific aidcd wood identification: rcfercnce man­ survey: 31-33. Chiba Univ., Yayoicho, ual: North Carolina Agricultural Research Japan. Service Bull. 474: 1-160.

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APPENDIX

List of extant taxa of Leguminosae compared with the fossil material.

Acacia angustissima, A. cochliacantha, A. dolychostachya, A. hindsii, A. macrocantha, A. me­ lanoxylon - Albizia sp. (Mexu 507), A. oceidentalis, A. policephala - Apoplanesia paniculata - Caesalpinia caladenia, C. coriaria, C. eriostachys, C. platyloba, C. pulcherrima, C. sclero­ carpa - Cassia atomaria, C. emarginata - Cereidium torreyanum - Cynometra oaxacana - Dalbergia sp. (Mexu 143), D. cogeitijlora, D. granadillo - Dialium guianense - Diphysa ocei­ dentalis, D. thurberii - Dussia mexicana - Enterolobium cyclocarpum - americana, E. lanata - polystachya - Glirieidia sepium - Haematoxylum brasilelto, H. cam­ pechianum - Leucaena lanceolata, L. leucocephala - Lonchocarpus atropurpureus, L. castilloi, L. cochleathus, L. constrictus, L. cruentus, L. eriocarinalis, L. guatemaltensis, L. honduren­ sis, L. lanceolatum, L. magallanesii - Lysiloma acapulcensis, L. bahamensis, L. divaricata - Mimosa sp. (Eitelma 33), M. arenosa - Myrospermum Jrutescens - Pithecellobium arborum, P. dulce - Piseidia communis, P. lanceolatum, P. seleri - Pityrocarpa constricta - Platymis­ eium lasiocarpum - Poeppigia procera - Prosopis julijlora - Pterocarpus amphymenium, P. rohrii - Schizolobium parahybum - Swartzia guatemalensis - Vatairea lundellii - Wil­ lardia mexicana.

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