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An Unusual New Cheilanthoid Fern from California1

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An Unusual New Cheilanthoid Fern from California1 738 AMERICAN JOURNAL OF BOTANY [Vol. 44 --, AND --------. (in press) Apomixis in Bothriochloa. HUBBAIlD, J. 19050. A study of Andropogon ischaemum. 1. Dichanthium, and Capillipedium. Phytomorphology. (Masters thesis.) A. & M. College of Texas. GARBER, E. D. 1944. A cytological study of the genus KIlISHNASWAMY, N. 1941. Untersuchungen zur Cytologic Sorghum: subsections Para-Sorghum and Eu-Sor­ und Systematik der Gramineen. Bot Centralhl, 60: ghum. Amer. Naturalist 78: 89-94. 1-056. ---. 19050. Cytotaxonomic studies in the genus Sorghum. KIiWADA, Y. 1919. On the number of chromosomes in Univ. California Publ. Bot. 23: 283-362. maize. Jour. Col. Sci. Imp. Univ. Tokyo. 39: 1-148. GOULD, F. W. 19053. A cytotaxonomic study of the genus MElIIlA, K. 1. 190505. Chromosome numbers in the tribe Andropogon. Amer. Jour. Bot. 40: 297-306. Andropogoneae, Gramineae. Indian Jour. Gen. and ---. 19056. Chromosome counts and cytotaxonomic notes Plant Br. IS: 144. on grasses of the tribe Andropogoneae. Amer. Jour. STEBBINS, G. 1. 19.50. Variation and evolution in plants. Bot. 43: 39.5-404. New York. AN UNUSUAL NEW CHEILANTHOID FERN FROM CALIFORNIA1 W. H. Wagner, J1'. and Elizabeth F. Gilbere RECENT STLI>ENTS of phylogeny in Filicineae involved, the "California lace fern," interpreted by have been interested in two general problems espe­ Copeland as Aspidotis californica Nuttall, has had cially-the value of the conventional characters almost as diverse a generic career: it was put in used in taxonomy, and the effects of hybridization Hypolepis by Hooker (1851) and more recently in in producing species. The traditional "key" char­ Cheilanthes by most authors (Jepson, 1923). Al­ acter of the sorus does not always confirm relation­ though the taxonomists have disagreed on exactly ships. Wholly unrelated plants may have similar what the generic assignment of these two ferns sori; and conversely, ferns demonstrated hy their should be, many of them do agree that, whatever constellation of other characters to have near the case, they should he kept in separate genera. affinity may differ from one another in such ex­ Only Maxon (1918) seems to have been entirely tremes as marginal vs. dorsal sor i, or coenosori convinced that the two plants are congeneric. vs. discrete sori. The other general problem con­ In the comparative study of the new, interrne­ cerns particularly the numerous intermediate taxa diate species to be described here, it was discovered which have been described among ferns. Cytologi­ that there are but few good characters that really cal and experimental work now suggests that in the separate "Onychiuni densum" and "Aspidotis cali­ future most if not all of those intermediate ferns fornica," and that at least one of the alleged differ­ will be shown to he species hybrids. These two ences is actually invalid. Accordingly, they will conclusions bear on the new plant to he descrihed both be treated in this paper as belonging to the here. same genus, Cheilanthcs. The intermediate fern is The description of a presumably lH'W species of known at present only from a narrow range, ill fern from the United States is a rare event in the San Luis Obispo, San Benito, and Marin counties. present day. The fern discussed here also possesses the few collections scattered over an area of some unusual taxonomic significance in that it seems to 180 miles along the coast of California, at distances be an exact intermediate between two species often inland from 10 to 60 miles. The late Mrs. Carlotta treated as helonging to separate gpnera. The sori C. Hall, a student of western ferns over many years, of the generically separated ferns are sharply dis­ was the first to recognize the intermediate plant as similar. Copeland (1947) placed them, both fa­ possibly distinct. The purpose of this report is to miliar cheilanthoid ferns of the west coast, in describe the new species, to compare it with its two distinct genera of his Pteridaceae, One of them is immediate relatives, and to discuss its possible evo­ the "Oregon cliffbrake," Unychiuni densurn Brack­ lutionary significance. enridge, which has been variously treated by other CHEILANTHES ear'Intta-halfiae" Wagner and authors as a Pellaea (for example, JI'pson, 192~ I, Gilhert, sp. nov. A small tufted fern of dry, rocky­ a Cheilanthes (as C. siliquosa Maxon, 1918; and soil habitats, with 5-10 photosynthetic leaves, 6-27 Fernald, 1950), and a Cryptograrnrna (Dir-ls, em. tall, and numerous persistent dark-pigmented 1899). Fernald (1950) wrote that "by its dimor­ old petioles. Stem creeping, dying off posteriorly phic fronds and aspect [it could he] ahout as well and leaving a rhizome 1-5 em. long and 0.2-0.5 placed in Cryptograrnrna." The other of the spec'es mm. thick, completely clothed with old petiole- 1 Received for publication April 24, 19057. 3 A C. cali/ornica et C. siliquosa stipitibus atrocastaneis 2 We are indebted to the directors of the University of l Lf (4.0-19.0) em. longis, laminis triangularibus 4-pinnatis', California and University of Michigan herbaria for the loan 4.05 (2.8-8.2) em. longis, 2.7 (1.4-5.05) em. latis, pinnis of herbarium specimens. We wish also to thank Mr. Robert infimis 3 (1-05) -jugis basaliter atrocastaneo-tinctis, coeno­ S. Whitmire for making a spore germination test for us, soris rnarginalibus valde interruptis vel soris discretis varie and Mr. Conrad V. Morton for his help in various ways. elongatis, sporis 46.7 (23-61) fJ. diam. differt. November, 19S7] WAGNER AND GILBERT-A NEW CHEILANTHOID FERN 739 bases and roots that increase its apparent diameter tically straight, finely toothed flap (fig. 1,A, and to 1-2 em.; roots abundant, wiry, 0.4 mm. in 2,D). By Copeland (1947) the long sorus was diameter, and extending S-15 em. or more in soil; considered to be subtended by a connecting vein paleae covering the sides of the rhizome, shoot along the laminar margin. The other species, C. apex, and petiole bases, linear-attenuate, 0.1-0.3 californica has a much narrower range, and is mm. broad at base, 1.5-2.0 mm. long, shiny, with known at present only from the state of California, an opaque black central strip and translucent where it is probably an endemic. The habitat of brown margins; leaves wiry and coriaceous, 15 this fern is also on rocks or rocky ground, but it (6-27) em. tall; petiole atrocastaneous, lustrous, does not seem to be so specifically limited to ser­ 11 (4-19) em. long, 0.7-1.0 mm. thick, devoid of pentine or limestone. The leaf-blades are very scales in the mature state except at base; leaf­ finely divided, usually 4-5-times pinnate. The ulti­ blade deltoid, four times divided, 4.5 (2.8-8.2) mate divisions tend to be broad, either lanccolate em. long, 2.7 (1.4-5.5) em. broad, the basal pinna or triangular, and the sori are always short and pairs triangular, 2.3 (1.5-3.0) em. long, 1.5 (1.0­ entirely discrete structures, less than 1 mm. in 2.0) broad at base; midrib atrocastaneous and breadth and well separated from adjacent sori on lustrous (like petiole) except for the green apical the same segment margin (fig. I,C; 2,F). No com­ third or fourth; pinna axes mostly pale green, but missural vein has been reported. In discussing the atrocastaneous basally on the abaxial surface of relationships of C. carlotta-halliae, each of its the lower 3 (1-5) pinna pairs; ultimate segments distinctive features will be considered in terms of linear-acuminate to lanceolate-acuminate. 2-8 mm. the corresponding characters of these two other long, 1.0 (0.8-1.5) mm. broad. the margins more species. or less shallowly dentate; sorus a strongly inter­ The comparative data of leaf and soral charac­ rupted marginal coenosorus or divided completely teristics given below are based on 100 dried plants into discrete sori of various lengths; indusium of C. siliquosa, 71 of C. carlotta-halliae, and 50 of varying from an irregularly lobed and cut coenoin­ C. californica. Although use of herbarium speci­ dusium to an irregular series of discrete indusia, mens is not ideal, the collections had probably been the indusiaI flap 0.3-05 mm. broad; spores accorded much the same treatment, making reason­ smooth, spherical, 46.7 (23-61) microns in diame­ ably valid comparisons possible. A total of 11 ter, pale-brown, with darker triradiate lines extend­ characters of the three taxa involved were measured ing nearly one-fourth of the circumference. in the expectation of establishing differences. Of TYPE SPECIMEN.-Locally abundant in dry rocky these, however, only seven yielded quantitative re­ soil below picnic ground, Bootjack Camp, Mt. Tam­ sults of recognizable comparative value. alpais, Marin Co., California, W. H. Wagner 4604, The ratio of blade length to stipe length is differ­ February 10, 1948 (MICH). ent among these plants: C. siliquosa has relatively COLLECTIONS EXAMINED.-CALIFORNIA: Marin the longest petioles, its stipe/blade ratio averaging County: Tiburon Hills along gully, Feb. 23, 1930, 3.1 (1.1-8.1); in C. carlotta-halliae this value is Alice Eastwood (UC); Bare hillsides, Tiburon, 2.6 (1.5-4.9); and in C. cali/ornica, 2.1 (0.5-3.8). Katherine Brandegee, 1907 (UC); "Rhizopogon The atrocastaneous pigmentation of the midrib growing among roots," Mt. Tamalpais, H. E. Parks varies in the amount to which it extends to the 3086 (UC); Bootjack Camp, Mt. Tamalpais, Feb­ pinna axes, and it was found that an average of ruary, 1948, W. H. Wagner 4604 rue, MICH-­ 1.7 (1-4) of the lower pinna pairs of C.
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