The Allioniaceae of the United States, with Notes On
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THE ALLIONIACEAE OF THE UNITED STATES, WITH NOTES ON MEXICAN SPECIES. liy I'ai i, (Staniii.ky. INTRODUCTION. Of all the families of North American plants none, prohnblv. have been more neglected than the Allioniaceae. In the last fifty-five years no monograph of the American representatives of the family has ap- peared. Linmrus in the Species Plantarum published two North American genera of this family—Mirabilis, with one species, and Iioerhaavia with four species, only two of which, however, occur in the region under consideration. Other genera and species of the family were soon published, all of them in scattered publications. The first treatment of the familv as a whole was that of Ohoisv in «- * I)e Candolle's Prodromus. In that work, 10 North American genera were described and, under them, 31 species, not including several species of Pisonia. Choisv's work is interesting and at times help- ful. but the author laljored under the difficulty of not having seen some of the plants of which he wrote, as a result of which some seri- ous mistakes were made. The next work of any importance dealing with the family was that of Asa Gray, in the Hot any of the Mexican Boundary Survey;" that paper is a very brief one and includes de- scriptions of but few species, although Gray described at various times a considerable number of new genera and species in the Al- lioniaceae. I)r. Anton Heinierl, of Vienna, probably the foremost student of this group of plants, contributed to Engler and PrantPs Natiirlichen Pflanzen-Kami lien h the section dealing with the Allioniaceae, a paper valuable for the excellent discussion it contains of the various genera. The work is exceedingly conservative, and the family is treated as nA. Gray in Torrey, Hot. Mex. Bound. 172-175. .1859, (Emory, Rep. U. S. & Mex. Homul. Snrv. Vol. II. Pt. 1.) i»Teil III, Abt. 1 t>, pp. 14-32. 1889. 303 304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM. European botanists so commonly treat groups of American plants. The genus Allionia, for instance, is made a mere section of Mirabilis, and other adjustments of the same kind are made which, although they may be the easiest way of disposing of genera, are certainly not conducive to clearness. In 1902 Mr. M. E. Jones published in his Contributions to West- ern Botany a a paper dealing with the family as it is represented in the Great Plateau region, an area in which are found almost all the species at that time known to occur in the United States. In the same year there appeared in the Bulletin of the Torrey Botanical Club h a paper by Dr. Per Axel Kydberg dealing with the Allioniaceae of the Rocky Mountains and containing descriptions of a number of new species, which is undoubtedly the most critical and valuable publication dealing with any group of the American representatives of the familv. # The work, the results of which are here discussed, was carried on at the New Mexico Agricultural College during the years 1 DOT and 11)08. The writer had the privilege of examining all the material of the Allioniaceae to be found in the herbaria of the following insti- tutions and individuals: National Herbarium; Missouri Botanical Garden, including the Engelmann and Bernhardi herbaria; Field Museum of Natural History: University of California, including the Brandegee Herbarium; University of Wyoming; University of Nevada; Universitv of Arizona; Mr. A. A. Heller. Mr. Iv. Iv. Mac- keuzie, Prof. E. O, Wooton, and the New Mexico Agricultural Col- lege. He wishes here to express his obligations to the curators or owners of these collections; also to Mr. G. E. Osterhout, who furnished material for examination. It was only through the kindness of those who have charge of these various collections that this work was made possible. The author is under special obligations to Prof. E. O. Wooton, under whose direction the work was begun and completed. The present paper is intended to cover all the representatives of the family occurring within the United States and most of those found in Mexico and the West Indies, with the exception of the genus Pisonia. The drawings are by the author, with the exception of Plates XXXIV and XXXV, which are by the German artist, W. Liepoldt. The author wishes especially to express his indebtedness to Dr. Anton Heimerl, who forwarded to him the two latter drawings and the descriptions which accompany them, with permission to use them here. Doctor Heimerl's notes attached to the sheets of the National Herbarium have also in several instances been of great help in the preparation of this paper. "10: 6 29: (ISO-tiiW. STANDLEY—ALLIONIACEAE OF THE UNITED STATES 305 SYSTEMATIC TREATMENT. ALLIONIACEAE Iteichenb. Oonsp. sr>. 182S, yyvtuginaccuc Limit. Nat. Syst. ed. 2. 213. 1S3(S. Annual or i>erennial herbs, often shrubs or trees, with brandling or dichoto- nious-forkJng stems; stems usually with swollen joints, sometimes armed with spines; leaves opposite or alternate. simple, entire, or sometimes re pa ml, exstipulate; inflorescence various; flowers regular, perfect or sometimes uni- sexual, often subtended by bracts which form a calyx-like involucre; perianth consisting of a calyx only, this often showy and corolla-like, tubular, funnel- form, or campanulate, usually deciduous above the ovary; stamens 3 to many: filaments filiform, distinct or united at the base, often unequal in length, ex- ported or included; anthers 2-ceIled. opening by longitudinal fissures; ovary l-celled, superior but surrounded by the calyx tube, sessile or short-stalked; style slender: stigma usually capitate; ovule solitary, erect, sessile; fruit an nnthocarp. indehiscent. fleshy, leathery, or hard, angled, ribbed, grooved, or winged; seed erect, with a hyaline testa which is free from or adnate to the pericarp; endosperm variable; embryo straight or curved. The fhmily consists of about 2t! genera and 2,10 species. Most of tbe genera and species are confined to the Western Hemisphere. In the Old World there are found one species of Allionia, several of Hoerhaavia and Pisonia, and the mono- typic South African genus Phaeoptilon. Of these only one, a species of Itoer- haavia. occurs in Europe (in southern Spain), the others being confined to Africa, soul hern and eastern Asia, and the islands of the Pacific. Doctor Heiinerl mentions the fact that one or two American species have become naturalized at various places in Europe. In the Western Hemisphere there seem to be two centers of distribution, one in tropical and subtropical South America and the West Indies, characterized by such genera as Pisonia, Xeea, Bougainvillea, and others: the other in Texas, New Mexico, Arizona, California, and northern Mexico, especially characterized by such genera as Boerhaavia, Abronia, Aeleisanthes, Allionia, but presenting several others. Of the entire number of genera included in the family 1ft occur in the latter region embracing more than 100 species. It is the region about this center that this paper attempts in cover. KEY TO THE OKNKKA. Flowers involucrate. Involucre polyphyHons, composed of 5 to 15 bracts which surround a few-flowered or many- flowered head. i'ruit winged or at least with rudimentary wings: bracts few: stamens and pistil In- cluded. Wings not completely encircling the fruit but interrupted above and below I. Auronia ( p. 300). "J. THIPTKKOCALYX ( p.327 ) Wings completely encircling the fruit.. Fruit not winged but merely 10-ribbed; bracts more numerous; stamens and pistil exserted X. \ YlT A(i IN [A (p. 3Ii0). Involucre giiniophyllous; flowers 1 to several. Fruit with prominent lateral wings which are often toothed; with 2 rows of glands along the dorsal surface 4. Wkiiklia (p. t. ■% 306 CONTRIBUTIONS FROM THK NATIONAL UKRBAltirM. Fruit not winged. Fruit witli % prominent ribs; involucre en l;i rgei1 and membranous in fruit--. .".Ai.i.hima (]i. HIM). Fruit smooth or somewhat 5-angled but not ribbed; involucres not nieinbra- nous and usually not enlarged in fruit. Involucre rotate, slightly enlarged in fruit, .'{-flowered U, Ai,uoniei.i,a i p. Involucres campniiulnte, not en- larged in fruit. Involucres containing several flowers which have a rather thick tube of medium length or sometimes rather long T. Qi:amo<i,ii»iox (p. H57). Involucres 1-flowered. I'erianth canipaimlate s. UrcsruKoNiA ip. HtiO), I'erianth funnelforin with a Ions, slender tube n. Mikakh.ik (p. HiS(t). Flowers without an involucre or each flower sub- tended by 1 to H liraets. 1 Fruit with conspicuous, thin, membranous wings- Id. SKMNocAWTstp. IJN7). Fruit not conspicuously winged: wings when present thick and coriaceous. Flowers large, 2 cm. long or usually more. I'erianth with a long slender tube and broad limb, each flower subtended by l! or H small, narrow bracts to. Aci.i;isantim:s( p. 11(11)). I'erianth campanulate, subtended by a large, ovate, leaf-like bract. H. IIkumidii \i < p. XT2). Flowers small, 2 cm. long or usually much less. Fruit 10-angJed or 10-ribbed. Fruit asymmetrical, Mowers in ra- cemes. 12. Sknkkxhkkui a I p.H72). Fruit symmetrical, flowers not in racemes. Fruit with conspicuous, mucila- ginous glands : climbing or re- clining plants with thin leaves; flowers in umbels 12. Sfnkknri'Kcia f p. H72i. Fruit without conspicuous glands; erect plants with very thick leaves; flowers ir- regularly clustered, not in umbels 14. ANrr.oc\ri.i,s( p.H74 i. Fruit iVangled. r>-ribbed, or sometimes with low, thick wings; perianth cam pauulate _ 15. Hokkhaavia (p. 375). 1. ABRONIA Jnss. Abronia Juss. (Jen. 448. ITS!). 'f'rici alHn L'ller.; Willd. Sp. l'l. X: SOT. 1799, Annual or perennial herbs, erect or prostrate, glabrous or pubescent ; leaves opposite, petloled.