The Status of Fraxinus Papillosa (Oleaceae): Sem Study of Epidermal Features
Total Page:16
File Type:pdf, Size:1020Kb
14 LUNDELLIA DECEMBER, 2010 THE STATUS OF FRAXINUS PAPILLOSA (OLEACEAE): SEM STUDY OF EPIDERMAL FEATURES Justin K. Williams1 and Guy L. Nesom2 1Department of Biological Sciences, Sam Houston State University, Huntsville, TX 77341-2116 22925 Hartwood Drive, Fort Worth, TX 76109, USA www.guynesom.com Abstract: The abaxial leaf surfaces of Fraxinus in the southwestern USA and northwestern Mexico are sometimes white-glaucous and have been describedas ‘‘papillose.’’ Individuals exhibiting this morphology have been identified as F. papillosa, a species reported from Arizona, New Mexico, Texas, and Mexico. In the original description of F. papillosa, its leaf epidermal structure was compared to F. americana L., but the abaxial surface morphology of F. papillosa is distinctly different from that of F. americana. In the latter, a cuticular reticulum overlays the epidermis and partially obscures it, while the abaxial surface of F. papillosa is similar to that of F. velutina in its lack of a cuticular overlay. In F. papillosa each epidermal cell is abruptly convex or folded upward into a papilla-like structure, compared to the relatively flat-surfaced cells of F. velutina—intergradation occurs but whether the two are appropriately considered conspecific remains to be determined. SEM photographs of the abaxial leaf surface morphology of F. papillosa, F. velutina, and other species that are similar to F. papillosa are provided. Keywords: Fraxinus papillosa, F. velutina, F. americana, Scanning Electron Microscopy, leaf epidermis. Fraxinus papillosa Lingelsh. has been accounts. It is among the names on the widely recognized as a species of the south- Flora of Arizona checklist posted on the western United States and northwestern Mex- Southwest Environmental Information Net- ico (e.g., Lingelsheim, 1920; Standley, 1924; work (SEINET, 2009), but there is no Sargent,1933; Little, 1952, 1976; Miller, 1955; voucher specimen identified as F. papillosa Kearney and Peebles, 1960; Martin and Hutch- for any state in any of the herbaria associated ins 1980; Elias 1980; Powell 1988; USDA, with the SEINET database. NRCS 2009). The original description of the Correll and Johnston (1970) included species (Lingelsheim, 1907) was based on only Fraxinus papillosa in the Texas flora on the the type collection from northwestern Chihua- basis of identifications by Little: ‘‘Specimens hua. Its attribution to the USA was first from Presidio Co. have been tentatively reported by Rehder (1917), who cited Mearns identified as this species (fide E.L. Little, 2533 from the San Luis Mountains in Grant Jr.). Additional material is needed for Co., New Mexico; Sargent (1933) repeated the complete verification.’’ Little’s Atlas map same citation. Little’s account of the species (1976) unequivocally (though without doc- (1952) was the first to describe a wider umentation) added trans-Pecos Texas to the geographic range and was accompanied by overall range of F. papillosa. Simpson (1988) voucher citations (from MICH and US) from mapped F. papillosa in Presidio County, Arizona, New Mexico, Chihuahua, and Sonora. noting (p. 156) that it occurs ‘‘in the Chinati The citations and map of Fraxinus Mountains … and is also thought to occur papillosa by Little (1952, 1976) apparently in the Bofecillos Mountain range to the east have been the basis for most subsequent and the Sierra Vieja range just north of accounts and summaries of its range. In fact, Chinati Peak.’’ The Texas distribution of F. it appears that the recognition of the species papillosa later was seemingly corroborated in in the USA stems mostly from Little’s the Atlas by Turner et al. (2003) through a LUNDELLIA 13:14–22. 2010 NUMBER 13 WILLIAMS AND NESOM: EPIDERMAL FEATURES OF FRAXINUS PAPILLOSA 15 dot in Presidio County, based on a TEX feature, which has been mentioned under F. voucher. americana, is found also in F. texensis [5 F. The feature generally emphasized in albicans] and suggests a relationship with distinguishing Fraxinus papillosa is the those species in addition to the similarity ‘‘papillose’’ nature of the abaxial leaf surface. with F. velutina (or F. standleyi). These A papillose abaxial surface also has been minute papillae, though not shown by a noted as a constant feature of the F. hand lens, are revealed by a high power americana L. complex (including F. albicans (40x) dissecting microscope or by low power Buckley [see Nesom 2010a], the diploid, (50x to 100x) of a compound microscope tetraploid, and hexaploid entities of F. through direct examination of the leaves or americana, and F. pauciflora Nutt. [see collodion leaf peals.’’ Nesom 2010b]). Photos of this feature by Wright (1944, p. 88) noted that ‘‘Leaf Wright (1944) and line drawings by Miller peels of the Mexican Fraxinus papillosa (1955) are remarkably similar to SEM Lingelsh. are similar to those of white ash,’’ photos by Hardin and Beckmann (1982). but he did not show a photo of a F. papillosa At high resolution, the minute areolae sample or cite a voucher. Wallander (2008) formed by the cuticular reticulum that also observed that ‘‘Like the eastern and overlays the epidermis are equally as con- much larger F. americana [F. papillosa] has a spicuous as the papillae in leaves with the distinctly papillose lower leaf epidermis.’’ ‘‘papillose’’ morphology—the whole reticu- The observations of Miller (1955, p. 54), lum appears minutely, shallowly, and evenly however, were different—she noted that ‘‘reticulate-perforate,’’ or ‘‘micro-foveolate.’’ Fraxinus papillosa ‘‘is perhaps more easily The term ‘‘coronulate’’ was used by Soler- confused with F. velutina than with any eder (1908, ‘‘coronulate papillae united by a other species. The elliptical leaflets and the network of ridges’’) and preferred by Wilk- fruits are often very ‘velutina-like;’ however, inson (1979), Hardin and Beckmann (1982), it is distinguished by the papillose condition and Kremer et al. (2008) to describe this of the leaflets. It may be closely related to F. aspect of the abaxial epidermis. Once one americana ssp. texensis; but the papillose becomes familiar with this morphology, pattern in F. papillosa is a very close one, the nature of the abaxial epidermis of F. quite unlike the typical network pattern of americana and F. texensis can be readily the typical white ash complex.’’ identified, making it easy to differentiate the Regarding Fraxinus papillosa,Felgeretal. F. americana group from the other North (2001, p. 262) noted that its ‘‘[geographic] American ash species by a quick inspection. distribution is wholly within the range of F. In the original description of Fraxinus velutina.’’ Although most authors have treated papillosa, Lingelsheim (1907) described the F. velutina and F. papillosa as distinct, leaflets as having a network of papilla Standley (1924: 1136) expressed the opinion beneath and noted that the new species that they ‘would have to be united ultimately’ was related to F. americana. Rehder (1917, and Powell (1988) likewise cast doubt on their p. 211) noted that F. papillosa differs from F. distinctiveness. We also have found that the velutina Torr. ‘‘chiefly in the glaucous distinctions lack clarity. papillose under surface of the leaflets and The present study investigates the abax- from F. americana in its sessile smaller ial foliar features of Fraxinus papillosa and leaflets.’’ Little (1952, p. 377) observed the those of potentially and putatively closely abaxial epidermal features of F. papillosa to related species. The null hypothesis, as its be similar to those of F. americana. ‘‘As its epithet suggests, is that F. papillosa is most specific epithet indicates, F. papillosa is closely similar in morphology, due to its characterized by the papillose, glaucous or pappilose epidermis, to the F. americana whitish under surface of the leaflets. This complex. 16 LUNDELLIA DECEMBER, 2010 MATERIALS AND METHODS Each abaxial epidermal cell of a Fraxinus papillosa leaf is abruptly convex (Fig. 1a, the Seven putative species are included in type of F. papillosa, from Chihuahua), or the study: F. papillosa, F. velutina, F. perhaps better described as folded upward, berlandieriana A. DC., F. pennsylvanica presenting the papillose aspect, and the Marsh., F. profunda Bush, F. albicans, and whole surface is prominently white. These F. americana—including diploid, tetraploid leaves have often been described as ‘‘glau- (F. smallii Britton), and hexaploid (F. cous,’’ but the white color apparently results biltmoreana Beadle) variants, as identified from increased reflectivity of the raised cells, from chromosome counts and morpholog- as there is no obvious evidence of deposition ical features (Nesom in prep.). Leaf samples of any kind of material. Trees with such from the seven taxa were collected from leaves occur in northwestern Mexico and dried herbarium specimens housed at BRIT, into New Mexico and Arizona. They inter- LL, MO, and TEX. Vouchers with brief grade to some degree with trees of F. localities are given in Table 1. In order to velutina, which produce leaves with green provide unambiguous identifications, sam- abaxial surfaces, the individual epidermal ples were taken from collections with mature cells without abruptly convex cell surfaces fruits, except for Powell 3148 and Sanders (Figs. 1c, e, f, g, h) but the details of the 2316. Vestiture on the abaxial surface is biological interaction between the two taxa variable; for this study, we selected leaflets remain to be delineated. with reduced vestiture in order to simplify A Texas collection, as the basis for the views of the epidermal features. All samples inclusion of Fraxinus papillosa in the Texas were taken from the distal one-fourth of the Atlas (Turner et al. 2003), has proved to be a leaflets. The samples were not rehydrated, specimen of F. velutina: Presidio Co: Arroyo acetolyzed, or critical-point dried. The Primero, a tributary of Fresno Creek on the specimens were mounted onto an SEM stud Big Bend Ranch, common tree, 14 Jun 1975, with conductive graphite tape and coated Butterwick & Strong 954 (TEX). This plant with gold using a LADD sputter coater.