18 LUNDELLIA DECEMBER, 2007 AFIELD STUDY OF HYBRIDIZATION BETWEEN BERBERIS SWASEYI AND B. TRIFOLIOLATA (BERBERIDACEAE) IN HAYS COUNTY,TEXAS Robert T. Harms Plant Resources Center, The University of Texas at Austin, 1 University Station F0404, Austin, Texas 78712-0471 Abstract: The widespread Berberis trifoliolata is sympatric with the narrowly restricted B. swaseyi in central Texas, where apparent intermediates occur. A detailed field study of sympatric populations in northern Hays Co. during 2004 to 2007 clarifies the morphological and phenological differences between the two species and shows that intermediates almost surely arose from hybridization. Limited evidence of introgression is also discussed. Keywords: Berberidaceae, Berberis, hybridization, flora of Texas. Berberis trifoliolata Moric. (Berberida- circumscribed by Ahrendt (1961) and in- ceae) is a widespread and common shrub of cludes both B. trifoliolata and B. swaseyi, open habitats and thickets, ranging from emphasizing the close relationship between central Texas westward to Arizona and south the two species. into northern Mexico (Whittemore, 1997). The range of Berberis swaseyi falls In central Texas, near the eastern edge of its entirely within that of B. trifoliolata,and distribution, it co-occurs with Berberis indeed B. swaseyi appearsalwaystooccurin swaseyi Buckley, a species of far more fairly close association with that species. restricted range that is known from seven This close sympatry and the close relation- counties along a narrow strip of the Edwards ship of the two species present possibilities Plateau immediately west and north of the for hybridization, and although this has southeast edge of the Balcones Escarpment been suggested in the literature (Durand, (Whittemore, 1997; Carr, ined.). 1972, p. 322; Breckenridge, 1983; Whitte- The compound-leaved species of Ber- more, 1997) it has not been documented. beris,includingthesetwo,haveoftenbeen Berberis hybrids have been produced in segregated into the genus Mahonia,aswas cultivation since the early 19th Century, done in the last worldwide revision of the including broad crosses between simple- group by Ahrendt (1961); however, recent and compound-leaved species (Ahrendt, treatments, such as that of Whittemore 1961). Naturally occurring hybridization (1997) for Flora of North America,tendto in wild populations of Berberis species include all species within a more broadly appearstoberelativelycommonasevi- circumscribed Berberis, and a molecular denced by specimens presumably morpho- study by Kim et al. (2004) indicates that logically intermediate between species (e.g., Mahonia wouldbeparaphyleticifitwere Landrum, 1999), but field studies of the treated as a separate genus. Whittemore phenomenon are lacking. notes that a small group of species of the In the late 1970s, Marshall C. Johnston southwestern U.S. and northern Mexico is of the Plant Resources Center (The Univer- morphologically intermediate in many ways sity of Texas) noted the existence of between Berberis sensu stricto and Mahonia, apparent hybrids between Berberis swaseyi and the molecular data of Kim et al. and B. trifoliolata on property owned by the supportthishypothesisaswell.Thissmall author in northern Hays Co., Texas. The group mentioned by Whittemore corre- present phenological and morphological sponds to Mahonia sect. Horridae Fedde as study is the outgrowth of that observation. LUNDELLIA 10:18–31. 2007 NUMBER 10 HARMS: HYBRIDIZATION IN BERBERIS 19 STUDY SITE Before January of 2004, eight BInt were identified, all but one of them along a half- The study site is a 50-acre tract in the mile portion of the bottomland area. These limestone Hill Country of northern Hays eight, along with 21 BTr and 22 BSw County west of Austin, at latitude 30u179200 individuals, were followed in 2004 for N and longitude 98u099570 W; elevations phenology. Starting in the last week of range from 280 to 335 m. It consists of January 2004 (when only a few BTr had a valley of bottomland, largely of oaks and produced floral buds) and continuing until grasses, with a creek down the middle, and early July (when the last fruits of BSw had drier rocky slopes with shallow soils on the disappeared), stages of development were east and west, dominated by Juniperus ashei recorded twice a week for each of the 51 J. Buchholz and grasses plus scattered groves plants. Each plant was visually inspected for of live oak (Quercus fusiformis Small). It is stage of flowering and fruit development and 2 miles upstream from the Pedernales River, an index reflecting that stage was recorded; near which Berberis swaseyi was first discov- e.g., flowering was graded on a scale of 0 ‘first ered by S. B. Buckley in 1866 (Buckley, flower or two just opening’ to 10 ‘plant in full 1870). Over 500 individuals over 18 inches flower, few buds yet to open.’ In addition, tall of Berberis with pinnately compound changes in leaf and stem development were leaves (i.e., B. swaseyi and potential hybrids, recorded through January 2005, while all see below) grow at the site, strongly through the period variation in characters of concentrated in, but not exclusive to, the stated or potential taxonomic importance bottomland area, along with a larger number was studied. Other individuals at the site of B. trifoliolata distributed throughout. were examined to confirm observations or extend quantitative data as needed. The METHODOLOGY survey was repeated with increasing numbers of individuals for the flowering and fruiting A review and comparison of literature, periods of spring 2005 (27 Btr, 19 BSw, 12 herbarium material, and the live specimens at BInt), spring 2006 (28 Btr , 27 BSw, 23 BInt), the site led to morphological characterization and flowering of 2007. At all stages, instances of the two species of Berberis in their relatively of coincidence between intermediate charac- pureformsastraditionallyrecognizedbytaxo- ter states of morphology and phenology were nomists. Character states traditionally used to noted, to serve (if present) as evidence of separate the species were studied and either hybridization. accepted as legitimate differentiae or rejected, Morphological details that did not re- based on observed variation within and quire careful measurement, such as leaflet between the two species. Previously unutilized number and position on the rachis, were characters were also studied, and several were noted in the field in order to determine found that present useful distinctions between ranges of variability. For this purpose, the species. Based on these accepted character hundreds of plants of both species were state differences, individuals at the study site examined for structures that were fully were then divided into three categories: 1) developed and within the normal range for typical B. trifoliolata (referred to as ‘‘BTr’’ the plant, and noted or collected only if they below); 2) typical B. swaseyi (‘‘BSw’’); and 3) extended ranges thus far established. Speci- individuals showing intermediate or mixed mens for detailed measurement were col- character states (‘‘BInt’’). A subset of individ- lected in the field and kept fresh until they uals was studied intensively as to phenology could be imaged for measurement within and morphology throughout the year, espe- 24 hours. These specimens, often dissected cially during the flowering and fruiting period, but not flattened or compressed, were placed as detailed below. on the glass plate of an Epson Perfection 20 LUNDELLIA DECEMBER, 2007 FIG. 1. A, B. Typical leaves of Berberis trifoliolata and B. swaseyi, with rachis and main veins highlighted. C. B. swaseyi shade leaf with acute terminal leaflet base. 2400 flatbed scanner, and scanned at resolu- differences that have been used by one or tions from 300 to 2400 dpi, and saved in more of the above authors to distinguish the TIFF format. Most detailed measurements two species are, for B. trifoliolata: toothless were made with scans at 2400 dpi, with filaments, blue-black fruits, style developed, digital scales approximating measurement middle leaflet sessile; versus, for B. swaseyi: accuracy to 0.01 mm. toothed filaments, yellowish-red fruits, style Voucher specimens of individuals from essentially absent, terminal leaflet stalked. the site representing typical phases of the Study of the two species at the study site two species plus intermediates are deposited indicated that some of these characters are at the Plant Resources Center (TEX, Harms useful in separating the species while others 49 though Harms 60). A web site with many are not. In addition, other useful distin- results, details, and numerous images from guishing characters were noted. Therefore, the study can be found at http://www.biosci. general descriptions of the two species as utexas.edu/prc/DigFlora/HaysBerb.html. found at the study site are given below, emphasizing the characters in which they CHARACTERIZATION OF BERBERIS TRIFO- differ (noted in bold italics). These char- LIOLATA AND BERBERIS SWASEYI acters are illustrated in Figures 1–4 and serve as the basis for the subsequent observations Berberis trifoliolata and B. swaseyi are on hybridization as well as aids to future most easily distinguished by the trifoliolate taxonomic descriptions of these species. leaves of the former as compared to the Although, as will become clear, it cannot pinnately compound leaves of the latter. be assumed that any individual of either Leaflet number in B. swaseyi is generally said species is genetically ‘‘pure,’’ the species as