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Taxonomic Revision of the Genus Fendlera (Hydrangeaceae)

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Taxonomic Revision of the Genus Fendlera (Hydrangeaceae) NUMBER 4 TURNER: REVISION OF FENDLERA TAXONOMIC REVISION OF THE GENUS FENDLERA (HYDRANGEACEAE) Billie L. Turner Section of Integrative Biology and Plant Resources Center, The University of Texas, Austin, Texas 79812 Abstract: Fendlera is treated as having five closely related species, four of which have been recognized by previous authors, and F. tamaulipana B. L. Turner, sp. nov., a newly described taxon from northeastern Mexico. Except for the latter, the species are all to some extent syrnpatric but their populations are relatively uniform and their diagnostic characters do not suggest that intergradation in a populational sense occurs. The taxa do not normally occur together, although they may occur in close proximity and prob­ ably do occasionally hybridize, this perhaps confounding taxonomic interpretations of the genus by previous workers. The five species recognized are: F. rupicola, so far as known, confined to three small populations in central Texas; F. falcata, occurring in northwestern Mexico and the southwestern United States; F. wrightii, having a distri­ bution similar to that of the foregoing; F. linearis (including F. rigida), occurring mostly in northeastern Mexico and closely adjacent Trans-Pecos, Texas; and the newly de­ scribed, F. tamaulipana, known by three collections from the vicinity of Miquihuana, Tamaulipas, Mexico. Distribution maps are provided for all of the species concerned. Keywords: Fendlera, Hydrangeaceae. The genus Fendlera was erected by En­ (e.g., Soltis et al., 1995; Holmgren and gelmann and Gray in 1852 with the descrip­ Holmgren, 1997) where its relationships ap­ tion of F. rupicola in Plantae Wrightianae pears to be with or near Philadelphus (in my from collections made by Lindheimer along opinion, based largely upon habital, floral the Guadalupe River in Comal County, Tex­ and ovarian features as well as ecogeograph­ as. At the same time they also recognized a ical considerations). I hypothesize that the var. wrightii, this based upon collections present day, largely xerophytic, Fendlera was made by Charles Wright in western Texas. derived from the same ancestor as members The latter was elevated to specific rank by of the large, mainly mesophytic genus, Phil­ Heller in 1895. Thornber, in 1913, added adelphus, becoming progressively modified two additional species, F. falcata and F. to­ as to stamen number (reduced from nu­ mentosa. The latter taxon is treated as syn­ merous to a set number, mostly eight), and onymous with F. wrightii in the present filament modification (from terete to flat­ treatment. Rehder, in 1920, added F. linearis, tened with extended apical appendages). Sol­ a species largely confined to Mexico, the type tis et al. (1995), however, using DNA (rbcL from near Monterrey, Nuevo Leon. John­ sequence data), suggest that Fendlera and Ja­ ston, in 1941, proposed an .additional Mex­ mesia are sister taxa to the rest of the 16 or ican species, F. rigida, the latter treated here­ more genera of the Hydrangeaceae; follow­ in as synonymous with F. linearis. Four of ing Fendlera and Jamesia, they found the re­ the above proposed taxa are recognized as maining hydrangeoids. to cluster into two valid species. A fifth species, F. tamaulipana, large subclades that closely parallel the tra­ is proposed in the treatment that follows. ditional division of the family into the tribes Philadelpheae and Hydrangeae. GENERIC RELATIONSHIPS CHROMOSOME NUMBERS Fendlera was initially placed in the large family Saxifragaceae but most recent work­ Chromosome counts are available for ers position the genus in the Hydrangeaceae only a single species of Fendlera, this re- LUNDELLIA 4:1-11. 2001 2 LUNDELLIA DECEMBER, 2001 ported as F. wrightii (Fedorov, 1969). Two Intricately branched F. finearis coup.ts are reported in the compendium, one by Sax in 1931, listing a count of 2n = '----"No=!'"'- 0 - F. tamaufipana 22, and another by Hamel in 1953, also re­ ~--- F.rupicola porting a count of2n = 22. Sax (1931) pro­ ~-- F. falcata vided a camera lucida drawing of meta­ '---- F. wrightii phase meiotic chromosomes of F. wrightii showing n = 11 pairs. He also noted that the plants concerned were obtained from FIG. 1. Hypothetical phyletic relationships among species of Fendlera based upon ecomor­ the grounds of the Arnold Arboretum. I phogeographical considerations. Numbers refer have not encountered vouchers for any of to character states as follows: 1) upper leaf sur­ these early counts. faces glabrous; central Texas; 2) upper leaf sur­ faces pubescent, west of the Devil's River (Val SPECIES CONCEPTS Verde and Crockett Counties, Texas); 3) lower leaf surfaces with a single layer of vestiture; 4) lower leaf surfaces with a double layer of ves­ I am an avid proponet of the biological titure. Gaps between the clades represent in­ species concept as elucidated by Mayr ferred divergences. (1969, 1992), Levine (1981), and numerous other workers. While some botanical work­ ers believe that "Plant species lack reality, sonably sure that these relationships will be cohesion, independence, and simple evolu­ shored up by appropriate DNA studies. tionary or ecological values" (Levin, 1979), my many years of field work has led me to FENDLERA Engelm. & A. Gray the conclusion that plant species, for the most part, are very much like animal spe­ Much-branched, usually thorny or cies and with approiate field work can be "spinescent" shrubs (except for F. tamau­ recognized as populational units having lipana) mostly 1-3 m high. Leaves simple, morphoecogeographic integrity. Mayr opposite, short-petiolate to nearly sessile, (1992) applied the biological species con­ entire, the blades 1-3 nerved, ovate to fal­ cept to plant populations at the local flora cate or linear, mostly 1-3(4) cm long, 1-10 level and concluded that the concept ap­ mm wide, variously pubescent above and plied equally well to plant taxa. I concur below, rarely not. Flowers showy, mostly in with Mayr's assessment. clusters of 2 or 3, the pedicels 2-25 mm I freely admit that there are alternative long; hypanthium narrowly to broadly tur­ ways to treat the five taxa recognized here­ binate; sepals 4, deltate to narrowly trian­ in. For example, one might recognize two gular; petals 4, white or tinged with pink or widespread species: 1) F. rupicola, with red, clawed, the blades broadly ovate to three varietal taxa; and 2) F. linearis with rhombic and having erase margins; stamens but two varietal taxa. I have given reasons mostly 8, the filaments flattened and having for their treatment as biological species un­ lateral lobes which extend beyond the an­ der the taxa concerned. For a more extend­ thers, the latter apically appendaged; ovary ed account of my views on infraspecific cat­ about half inferior, 4-celled; styles 4, dis­ egories and their heirarchical ranks see tinct or somewhat united below. Fruit a Turner and Nesom (2000). septicidal capsule, the seeds relatively few in The phyletic relationships of the five each cell (mostly 1-6). Base chromosome taxa of Fendlera recognized here, as inferred number, x = 11. from morphoecogeographical consider­ Type species, Fendlera rupicola Engelm. ations, is summarized in Fig. 1. I am rea- & A. Gray NUMBER4 TURNER: REVISION OF FENDLERA 3 Map of Texas ,-,--, 0 100 200 scale km FIG. 2. Distribution of Fendlera rupicola. The genus is named for Augustus Fen­ 3. Upper surface to some extent pubescent; lower dler (1813-1883), German-born naturalist surfaces moderately to densely pubescent; widespread in the southwestern United States of both North and South America, and one and northern Mexico of the earliest plant collectors in the south­ 4. Undersurfaces of leaves with a single layer western U.S.A. (mostly in New Mexico dur­ of coarse, appressed or ascending hairs ... ing the period 1846-1850). 2. F. falcata 4. Undersurfaces of leaves with two layers of vestiture, a lower layer of minute frizzly or crinkled hairs, this overlain by a much KEY TO SPECIES coarser vestiture of appressed or ascending hairs . 3. F. wrightii 1. Leaves linear, 1-2 mm wide, the margins com­ pletely enrolled 2. Intricately branched "spinesent" or thorny 1. FENDLERA RUPICOLA Engelm. & A. Gray shrubs mostly 0.5-2.0 m high; leaves mostly in A. Gray, Pl. Wright. 1: 77. 1852. (Fig. 2) 1.0-1.5 mm wide; northcentral Mexico and closely adjacent U.S.A . 4. F. linearis Fendlera rupicola var. lindheimeri A. Gray, 2. Strict, sparsely branched, shrubs mostly 1.5-4.0 m high; leaves 1.5-2.0 mm wide; Tamaulipas, Pl. Wright. 1: 77. 1852. Mexico, vicinity of Miquihuana ......... 5. F. tamaulipana TYPE: United States. Texas: Comal Co., 1. Leaves, narrowly lanceolate or falcate to ovate, among "perpendicular rocks" along the up­ mostly 2-10 mm wide, the margins incompletely per Guadelupe River, ca. six miles above enrolled, if at all 3. Upper surface ofleaves glabrous; lower surfaces New Braunfels, May 1850, F. Lindheimer sparsely strigose; known from only three pop­ 257 (HOLOTYPE: GH!; ISOTYPES: MOl, ulations in Central Texas ..... 1. F. rupicola NYl, USl). 4 LUNDELLIA DECEMBER, 2001 Lindheimer apparently first collected near the Kerr Co. line; and in central Ban­ this species (in fruit only) in May of 1850. dera Co. by the late Benny Simpson ( 1928- However, it was noted that the taxon con­ 1996) in April of 1991. The latter collector cerned flowered "in April," apparently a was well aware of his significant find, noting surmize provided by Engelmann, who on the label itself, "not collected in over 150 transmitted the plants, since the authors years," (being unaware of the aforemen­ note in the protologue itself that the illus­ tioned Reverchon collection). tration provided with the original descrip­ Fendlera rupicola appears to be most tion was comprised of fruiting material of closely related to F. falcata of far western var. rupicola ( = var. lindheimeri) and flow­ Texas, New Mexico and closely adjacent ering material of var.
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