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Occurrence and Phylogenetic Significance of Latex in the Malpighiaceae<Link Href= American Journal of Botany 89(11): 1725±1729. 2002. OCCURRENCE AND PHYLOGENETIC SIGNIFICANCE OF LATEX IN THE MALPIGHIACEAE1 ANDREA S. VEGA,2,5 MARIA A. CASTRO,3,5 AND WILLIAM R. ANDERSON4,5 2Instituto de BotaÂnica Darwinion, LabardeÂn 200, C. C. 22, B1642HYD Buenos Aires, Argentina; 3Laboratorio de AnatomõÂa Vegetal, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, C1428EHA Buenos Aires, Argentina; and 4University of Michigan Herbarium, Suite 112, 3600 Varsity Drive, Ann Arbor, Michigan 48108-2287 USA Latex and laticifers are reported for the ®rst time in the genera Galphimia and Verrucularia (Malpighiaceae), with description and illustration of the leaf and stem anatomy of both genera. Those genera and the other two in which latex is known (Lophanthera and Spachea) constitute a single tribe, Galphimieae, that is at or near the base of the family's phylogeny, which suggests that latex in the Malpighiaceae may indicate an ancestor shared with the Euphorbiaceae. Key words: Galphimia brasiliensis; Galphimieae; latex; laticifers; Malpighiaceae; Verrucularia glaucophylla. The angiosperm family Malpighiaceae includes approxi- MATERIALS AND METHODS mately 65 genera and 1250 species in the world and is dis- tributed in tropical and subtropical regions of both hemi- Plant materialsÐFresh and ®xed materials of Galphimia brasiliensis (L.) A. Juss. and Verrucularia glaucophylla A. Juss. were studied. Herbarium spheres (W. R. Anderson, unpublished data). Nearly 950 spe- specimens were deposited in the herbarium of the Instituto de BotaÂnica Dar- cies are endemic to the New World, northern South America winion (SI). Studied materials: Galphimia brasiliensis: ARGENTINA. Prov. being the major center of diversity (Anderson, 1979). The Entre RõÂos. Dpto. ColoÂn: Parque Nacional El Palmar, Los Loros, 14/III/1998, monophyly of the Malpighiaceae is supported by morpholog- MuÂlgura & Vega 1991; 4/XII/1998, MuÂlgura & Vega 1998. Dpto. Concordia: ical characters (Anderson, 1979, 1990) and has recently been Parque Rivadavia, near JoseÂHernaÂndez monument, 15/III/1998, MuÂlgura & con®rmed with molecular data (Chase et al., 1993; Cameron, Vega 1996. Verrucularia glaucophylla: BRAZIL. Bahia. Municipio de Morro Chase, and Anderson, 1995; Wurdack and Chase, 1996). Re- do ChapeÂu, Cachoeira do Ferro Doido, ca. 1000 m, 4/V/2001, Amorim et al. cent molecular studies of the Malpighiaceae did not provide 3662 (CEPEC, SI). convincing bases for dividing it into a small number of mono- phyletic subfamilies or tribes that re¯ect obvious morpholog- Anatomical studiesÐLeaves and stems of G. brasiliensis and V. glauco- ical synapomorphies (Cameron et al., 2001; Davis, Anderson, phylla were ®xed in FAA (formalin : ethanol : acetic acid) and then stored in and Donoghue, 2001), but they did con®rm the monophyly of 70% ethanol. Part of each sample had been previously treated with sodium the tribe Galphimieae Nied. This tribe, as circumscribed by hypochlorite to eliminate contents. All materials were dehydrated in an eth- anol series and embedded in paraf®n after in®ltration in a vacuum oven. Anderson (1978), comprises four genera: Galphimia Cav., Lo- Transverse and longitudinal sections 10 mm thick were cut on a rotary mi- phanthera A. Juss., Spachea A. Juss., and Verrucularia A. crotome. Sections were double stained with safranin-fast green (D'Ambrogio Juss. Anderson (1981, 2001) mentioned the presence of milky de ArguÈeso, 1986). Light microscope studies were made using a ZEISS Phomi latex in the stems of Lophanthera and Spachea, and the pres- III microscope (Zeiss, Oberkochen, Germany) and black and white Kodak ent paper documents the occurrence of latex in the other two Tmax 25 ASA ®lm. genera, Galphimia and Verrucularia. Galphimia includes approximately ten species of tropical RESULTS and subtropical America, with a disjunct distribution (Nieden- zu, 1928). Most of the species grow in Mexico and Central Field workÐDuring ®eld work in Entre RõÂos province (Ar- America and only one, Galphimia brasiliensis (L.) A. Juss., is gentina), the presence of latex in G. brasiliensis was observed. found in South America, in northeastern Argentina, southern The latex is milky and was detected in stems and blades (Figs. Brazil, Bolivia, Paraguay, and Uruguay. This species has leaf 1±2). During the preparation of the present paper, Ch. Ander- son (personal communication) told us about one collection of and calyx glands, and the structural and ultrastructural anato- G. brasiliensis from Bolivia (OrtõÂz S. 7, MICH) whose label my of these glands was studied by Castro, Vega, and MuÂlgura reported it to produce latex. Collectors of herbarium specimens (2001). The genus Verrucularia A. Juss. includes two species, of V. glaucophylla have never reported observing latex in the V. glaucophylla A. Juss. and V. piresii W. R. Anderson. Ver- ®eld, but laticifers were detected by anatomical techniques in rucularia glaucophylla has only calyx glands and is endemic stems and blades of that species. to Bahia, Brazil (Niedenzu, 1928). Anatomical studiesÐGalphimia brasiliensisÐThe blade in transverse and longitudinal sections is shown in Figs. 3±5. The 1 Manuscript received 22 January 2002; revision accepted 21 May 2002. adaxial epidermis is one-layered, the cells quadrangular to The authors thank Andre M. Amorim for his valuable help in collecting rectangular, with a thick outer tangential cell wall and cuticle. materials of Verrucularia glaucophylla and graphic designer TomaÂs E. Aversa for his technical assistance. The papillate abaxial epidermis has a thin outer tangential cell 5 Authors for reprint requests (e-mail: [email protected]; mac@bg. wall. Trichomes are absent. Stomata are present in both sur- fcen.uba.ar; [email protected]). faces and are more abundant in the abaxial epidermis. The 1725 1726 AMERICAN JOURNAL OF BOTANY [Vol. 89 Figs. 1±7. Galphimia brasiliensis (L.) A. Juss. 1±2. Field photographs of latex in petiole and blade, respectively. Figs. 3±4. Blade in transverse section, with laticifers immersed in spongy parenchyma. 3. Cleared section. Bar 5 50 mm. 4. Uncleared section. Bar 5 50 mm. 5. Blade in longitudinal section showing an articulated and unbranched laticifer. Bar 5 50 mm. Figs. 6±7. Stem. 6. Transverse section. Bar 5 50 mm. 7. Longitudinal section. Bar 5 50 mm. ABE, abaxial epidermis; ADE, adaxial epidermis; B, blade; C, cuticle; CH, chlorenchyma; CO, cortex; L, laticifer; LX, latex; P, petiole; PH, phloem; PI, pith; PP, palisade parenchyma; S, stomata; SP, spongy parenchyma; VB, vascular bundles; X, xylem. dorsiventral mesophyll includes one layer of palisade paren- tissue, then secondary phloem and xylem and a conspicuous chyma, composed of axially elongated cells slightly attached pith of parenchyma cells. Laticifers are distributed in the cor- to one another and representing one-third of the total thick- tex and are scarcely recognizable due to their similarity to ness. The spongy parenchyma has conspicuous intercellular parenchyma cells (Fig. 7). spaces in which are immersed abundant laticifers distributed in the proximity of collateral vascular bundles. The laticifers Verrucularia glaucophyllaÐThe blade in transverse and were easily detected in transverse and longitudinal sections of longitudinal sections is shown in Figs. 8±10. The adaxial the blades (Figs. 3±5) and stained intensively in uncleared sec- epidermis is one-layered, the cells quadrangular, with a tions (Fig. 4). In transverse section, laticifers are circular, 21.6± thick outer tangential cell wall and cuticle. The conspicu- 28.3 (±35.1) mm in diameter, numerous, with a parallel and ously papillate abaxial epidermis exhibits one simple or oc- regular pattern in relation to the axis of the leaf and (51.3±) casionally bifurcate papilla on each cell, and a thick outer 126±139 mm apart (Fig. 3±4). In longitudinal section, laticifers tangential cell wall. Trichomes are absent. Stomata are pres- are tubular, articulated, not rami®ed, and not anastomosing ent in the abaxial epidermis overarched by adjacent papil- (Fig. 5). The cells of the laticifers are 64.1±87.7 mm long, late epidermal cells. The mesophyll is dorsiventral, includ- elongated, with dilated extremes like a trumpet and semi-thick ing one-layered palisade parenchyma, which represents terminal walls. about one-half of the total thickness. The cells of typical The stem in transverse and longitudinal sections is illus- palisade parenchyma are axially elongated and attached to trated in Figs. 6±7. The stem is circular in shape and is bound- one another. The spongy parenchyma has conspicuous in- ed by a one-layered epidermis with stomata. The cortex con- tercellular spaces. Abundant laticifers are immersed be- sists of subepidermal spongy chlorenchyma and parenchyma tween the palisade and spongy parenchyma, in the prox- November 2002] VEGA ET AL.ÐLATEX IN MALPIGHIACEAE 1727 Figs. 8±14. Verrucularia glaucophylla A. Juss. 8. Blade in transverse section showing laticifers immersed in the mesophyll. Bar 5 50 mm. Figs. 9±10. Blade in paradermal section. 9. Bar 5 50 mm. 10. Bar 5 100 mm. Figs. 11±13. Stem in transverse section. 11. Bar 5 100 mm. 12. Cuticle, epidermis, incipient periderm, and cortical parenchyma in detail. Bar 5 20 mm. 13. Secondary xylem and central pith of parenchyma showing a peripheral starchy cell zone in detail. Bar 5 20 mm. 14. Stem in longitudinal section showing cortical laticifers (arrow). Bar 5 25 mm. ABE, abaxial epidermis; ADE, adaxial epidermis; C, cuticle; CO, cortex; F, phloem ®bers; L, laticifers;
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