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Genetic and Taxonomic Studies in Gilia: VIII. the Cobwebby Gilias Alva Grant

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Genetic and Taxonomic Studies in Gilia: VIII. the Cobwebby Gilias Alva Grant Aliso: A Journal of Systematic and Evolutionary Botany Volume 3 | Issue 3 Article 2 1956 Genetic and Taxonomic Studies in Gilia: VIII. The Cobwebby Gilias Alva Grant Verne Grant Follow this and additional works at: http://scholarship.claremont.edu/aliso Part of the Botany Commons Recommended Citation Grant, Alva and Grant, Verne (1956) "Genetic and Taxonomic Studies in Gilia: VIII. The oC bwebby Gilias," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 3: Iss. 3, Article 2. Available at: http://scholarship.claremont.edu/aliso/vol3/iss3/2 EL ALISO VOL. 3. No. 3, pp. 203-287 }UNE 1, 1956 GENETIC AND TAXONOMIC STUDIES IN GILIA VIII. THE COBWEBBY GILIAS1 ALVA GRANT AND VERNE GRANT HISTORY OF THE SECTION The section Arachnion of Gilia, including the species G. tenuiftora, G. latiftora, G. ochroleuca, G. sinuata, and their relatives, is the largest single group in the genus, comprising some 44 taxa distributed among 17 species. This section has its principal center of distribution in California, where all but three of the species (G. mexicana, G. tweedyi and G. crassifolia) and all but four of the taxa (the aforenamed species plus G. ophthalmoides ftavocincta) occur. The Cobwebby Gilias2 have evolved the largest number of taxa and attained the greatest abundance of indi­ viduals in the Mojave Desert and the mountain ranges on its western border. The group extends also into various peripheral areas, particularly the South Coast Range of California, Baja California, the Sonoran Desert of Arizona, the Organ and Franklin Mountains region of New Mexico and western Texas, the northern Sierra Nevada, the Great Basin, and the Rocky Mountains. In South America a secondary center of distribution exists in the Chilean Desert, the Argentine Andes, and Pata­ goma. Not only does the section Arachnion include the largest number of entities in any subdivision of Gilia, but it also comprises the taxonomically most complex and critical group in the genus and perhaps in the entire family Polemoniaceae. In conse­ quence of numerous intergrades between the species, resulting from the widespread occurrence of natural hybridization, and in consequence of the presence of several pairs of sibling species separated by elusive distinguishing characters, the develop­ ment of a taxonomic treatment serving the practical purposes of identification and classification has been beset with serious difficulties and has been greatly retarded as compared with many other plant groups. No existing regional flora or revisional treatment permits the correct identification of more than a small fraction of the Cobwebby Gilias. This situation is reflected in the state of chaos which is found in the Cobwebby Gilia folders in every herbarium, where several standard and well­ known names have been applied more or less indiscriminantly to a variety of speci­ mens. The charting of the variation pattern in the section Arachnion may for convenience be divided into the three stages of exploration, surveying, and analysis, which ac­ cording to Camp and Gilly (1943) and Stebbins (1950, ch. 1) describe the course of development of taxonomic research in general. The exploratory stage has con­ sisted of the finding and describing of new forms. This process began with the first botanical expeditions to western North America and southern South America and has continued to the present day. Among the botanists who have contributed to the taxonomic exploration of the Cobwebby Gilias are Bentham ( 188 3) 3 , Douglas (1845), F. Philippi (1857-1860), Gray (1870-1886), R. A. Philippi (1895), Jones (1898-1908), Parish (1900), Eastwood (1904-1943), Rydberg (1904), 'This study was aided by a research grant from the National Science Foundation. 2 Also referred to as the Woolly-leaved Gilias in earlier publications. 'Literature citations corresponding to these dates of publication of new taxa will be found in the lists of synonyms in the formal taxonomic section of this paper. [203] 204 EL ALISO [VoL. 3, No. 3 Milliken (1904), Brand (1907-1913), A. Nelson (1934), Munz (1935), Jepson (1943), Mason and A. Grant (1948), and numerous others who collected without publishing new names. Nor is the exploratory work on the section Arachnion ended yet. In the course of our own studies embodied in the present work it has been neces­ sary to describe 4 new species and 15 new subspecies, and some additional described entities will almost certainly be added by future students. Attempts to organize the scattered knowledge about the Cobwebby Gilias into a system of classification were begun, rather prematurely as it now appears, by Gray in the Synaptical Flora of North America ( 1886), Milliken in her Review of Cali­ fornian Polemoniaceae ( 1904), and Brand in Das Pflanzenreich (1907). These early efforts led up to Jep:;on's treatment of the genus Gilia in his Flora of California ( 1943), with which the surveying "stage of the taxonomic work may be properly said to have begun. This very competent treatment of the Cobwebby Gilias was soon superseded by the treatment of Mason and A. Grant in Abrams' Illustrated Flora of the Pacific States ( 1951). The latter treatment has served as a stepping­ stone for the revision of Gilia § Arachnion which is presented in the following pap:::r. In Jepson's Manual (1925) the entire group was placed in six taxa, and in Jepson's Flora (1943) in nine taxa (cf. Table 1). Munz (1935) recognized eight taxa, and Mason and A. Grant ( 1951) fourteen taxa. The present treatment recog­ nizes 39 taxa in California. Until the treatment of Mason and A. Grant it was customary to include G. splendens, now regarded as belonging in the section Saltu­ gilia, as a variety of G. tenuiftora. The latter authors segregated G. splendens as a species, but failed to exclude it and several other extraneous elements (G. stellata, G. scopulorum, G. leptomeria) from the section. In the present paper the section Arachnion is circumscribed and named; a new system of classification of the constituent entities is proposed; new formal descrip­ tions and illustrations of the species and subspecies are given; their geographical areas are mapped; and the nomenclature is straightened out. The system of classifi­ cation adopted here is outlined in Table 2. This treatment is most reliable for the large-flowered diploid species, to which the closest attention has been paid. The small-flowered diploid and tetraploid species of the Great Basin and Rocky Mountains, viz. G. inconspicua, G. ophthalmoides, G. tweedyi, G. transmontana, require much further study; the treatment proposed here should be regarded as a framework within which the future studies can be carried out. Gilia crassifolia and its allies in South America can not be dealt with in any satisfactory way for the present and are included in the treatment merely to indicate their general relationships to the North American members of the section. The third or analytical stage in the mapping of the taxonomic landscape, designed to elucidate the evolutionary relationships and history of the various entities, has been fairly commenced by a series of cytotaxonomic and genetic studies underway since 1948. The systematic revision of Gilia § Arachnion which follows is both a summary of the surveying work which has been carried out up to this point and a preface to future contributions to the evolutionary genetics of the group. METHODS Herbarium and Field Studies The present study is based primarily on a combination of herbarium studies and field studies. The herbarium work revealed the critical problems and areas which required attention, and the field work in those areas made it possible to interpret more intelligently the herbarium material. Each winter the Cobwebby Gilia collec- JUNE, 1956] COBWEBBY GILIAS 205 6 G. I. excellens • G. I. elongata * 5 -:: '0 ... ·· '* 'i 4 l/ .. / "' ~ 0 f ~t ..................... * .ca. *-·1···..... \*'/ •* \ . * c:: 3 * * ... · .!! ! ·· .. :.·· ... ,.... ...... ···········*·········· 0e -:: . t, i .. / * + / /0 "' 2 .... ~ .. Ee • 0 (.) • • • I I 2 4 5 6 Corolla length : Corolla lobe length Fig. 1. Range of variation in Gilia latiflora excel/ens and G. l. elongata. tions in the Rancho Santa Ana Botanic Garden and Pomona College Herbaria were restudied in the light of the field experience of the preceding spring and summer; each spring and summer we made a series of field trips which clarified problems raised by the herbarium studies of the winter before. This coordinated herbarium and field approach, carried on during the past seven years, has been indispensable for gaining an understanding of the section. In the final analysis, nearly all of the western North American herbaria, and several eastern ones, were consulted. Specific and subspecific names were applied after consultation of type specimens or photographs thereof. The steps followed in working out one particular taxonomic problem may be described as an illustration of the method of analysis. Preliminary observations in nature and in the herbarium indicated that there were two similar but markedly different forms of long-tubed large-flowered Cobwebby Gilias in the El Paso Mt. area of the northwestern Mojave Desert. A series of herbarium specimens including population samples collected by the authors was measured for thirteen carefully chosen characters in an effort to determine precisely the taxonomic differences. Two ratios using four of these characters proved consistently useful in separating the two forms. These two composite characters involved the proportions of the corolla: the ratio of corolla length to throat width, and the ratio of corolla length to lobe size. 206 EL ALISO [VoL. 3, No. 3 TABLE 1. PREVIOUS CLASSIFICATIONS OF THE COBWEBBY GILIAS IN CALIFORNIA. Jepson ( 1925) Munz (1935) Jepson (1943) Mason&A.Grant (1951) 1. G. ochroleuca 1. G. ochroleuca 1. G. ochroleuca 1. G. ochroleuca 2. G. tenuiflora 2. G. tenuiflora 2. G. tenui/lora subsp. transmontana var. purpusii 3. G. exilis var. arenarra 2. G. abramsii var.
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