NEW BOERHAVIA COULTERl PALMERl A VARIETAL VAR. COMBINATION FOR BOERHAVIA (NYCTAGINACEAE) OF SOUTHWESTERN NORTH AMERICA Richard Spellenberg Department Biology NM Las Cruces, 88003-8001, U.S.A. Boerh aviacoult^;n (Hook. Wats, va f.)S. Chois from y var. pal-tleri S.Wats., the VI, ;ico, is proposed for a Boerh m from var iffers the coulter, US' Bend )rnia to the Big regie Texas near the Uo Grande, and very I is symp and theymtergradetoash atric, Boerh avia with:spicate mflorescences, Ti(Hookf)SWatsva, Boerh a..acoulU Chois painieriS.Wats.,delavecin y var. deBo.•rhavia que difierede variedadc la eninf lorescenci asespiciformesLavari, iende desde las Islas Canales de California hasta la muy comun regior del "Big Bend" del Rio Bravo del > <as, y es en el Desierto Sonorense. Las 1 menudo variedades a son simpatricas, y se intergradan un poco. Se presenta una clave para distinguir The species of Boerhavia with spicate inflorescences have proven taxonomi- cally difficult for the past century, the variation presented by the complex re- sulting in very different taxonomic treatments. At one extreme that of is where Standley (1911) nine species were recognized. In 1918 Standley submerged synonym B. watsoni as a of B. coulteri, but maintained the other taxa he had recognized. At the other extreme is Reed's (1969) treatment, in which five of the species Standley recognized coulteri (Hook, Wats., rosei Standi, [B. f.) S. B. B. Choisy spicata in D.C., B. torreyana (S. Wats.) Standi., B. watsoni Standi] were synonyms placed as in one variable species, spicata. Most authors of floras B. that include or part of the Sonoran Desert region have followed Standley all (1918) in recognizing B. coulteri, and also B. spicata, and B. torreyana if they were & present in the area of concern (Spellenberg Martin Hutchins 1993; 1980; & & Munz Kearney Keck Wiggins Peebles 1964; 1968; 1964). In contrast, Felger some (2000) recognized plants included broadly constructed in coulteri, as B. by those authors, as B. spicata var. palmeri S. Wats. It is this taxon that is recog- nized as a variety of coulteri in this paper. B. While studying the complex for an upcoming treatment for the Flora of was North America North of Mexico, noted that torreyana conspicuously B. I papillate in the sulci between the ribs of the fruit, a character previously un- None complex published. of the other Boerhavia taxa in the are at all papillate. when name Watson (1889) cited no specimens he coined the Boerhavia spicata torreyana but specimens from western Texas collected during the Wats., var. S. mid-19th century Boundary Survey, very likely seen by him, have papillate fruits (specimens from Torrey Herbarium, NY, have been relevant specimens seen; GH ENCB from on loan to have not been seen). The character of papillae on the m New consistent populations from western Texas, Mexico, southeastern fruit is much Arizona at higher elevations, throughout of Arizona north of the Mogollon On Rim, and extreme south-central Utah (= B. spicata by Welsh et al. 1987). very similar appearing plants from the Sonoran Desert region, some of which are which identified as torreyana in collections, but actually represent coulteri B. B. much var palmeri (as proposed below), papillae are absent and sulci are narrower and rugose. That torreyana clearly distinguishable from other spicate less B. is Boerhavia has helped to clarify the complex. This particularly so in Arizona is and Sonora where three or four very similar taxa occur within the region. Watson and In 1889 transferred Senkenhergia coulteri Hook, to Boerhavia, f . he stated that the collection upon which the name was based was Coulter M25. mm) He noted that the perianth was one line in length (= 1/12", ca. 2 and that were and Only one such fruits clavate, 1-1.5 lines long, truncate, acutely angled. come specimen Hooker Herbarium Kew. having exists in the at labeled as It is from "Aha Sonora, Mexico," and here considered the holotype (photo and is NMC). photocopies fruits are clearly in irregularly fasciculate clusters in at Its On and same page the spicate inflorescence, they are truncate at the apex. the Watson new of his 1889 publication, stated that his variety, Boerhavia spicata Ml palmeri, based on the collection Palmer from Guaymas, Sonora, in 1887, var. had perianths 0.5 line long, and fruits clavate-oblong, a line long, obtuse, ob- He and tusely angled. also noted the inflorescence to be "open very slender." Palmer Ml represented by two specimens GH! (on loan ENCB, lectotype at to is not selected; fragment at DS!). In raising this taxon to specific status, Standley new name (1909) had to adopt a watsoni) because the binomial palmeri (B. B. S. Wats, was occupied. He noted that Arizona plants were placed there somewhat geography would them with doubtfully, suggesting that place coulteri. Rela- B. tive to other plants in the complex, the plants he identified as watsoni have B. more widely and seem smaller, spaced, less clustered fruits (if at all clustered), have had stems were decumbent-spreading. that to proposed here that the populations Standley recognized as watsoni, It IS B. and Watson that described as spicata var palmeri, are part of the coulteri B. B. complex and can be recognized at the varietal level. The following combina- tion proposed: is Wats. palmeri Wats.) comb. var. Spellenb,, nov. (S. .. The differences the two varieties of B. coulteri that Watson, and later Standley, noted by their classifications are very apparent in the field in their extreme but intergradient plants occur Occasionally var and states, coulteri coulteri B. & var palmeri are intermixed in populations Spellenherg Zucker 12954 = (e.g., SW km var palmeri, 12955 = var AZ, Pima 10 Marana, coulteri, Co., ca. air of km Anway Rd. S of Avra Valley Rd., 16 Aug. 1999; NMC). In the two mixed 1 populations have seen, the taxa differed in habit, the stems of var coulteri de- I cumbent immediate at the base, then strongly ascending or erect, contrasting to the more decumbent-spreading habit of var palmeri, but habit very likely is to be influenced by ecological conditions and density of populations. Some mono- varietal populations of the var coulteri have decumbent-spreading stems. The var coulteri also has larger flowers, a denser inflorescence, and fruits in irregular fasciculate whereas the var palmeri usually has minute flow- clusters, much and ers, well-spaced, smaller fruits, the clusters less pronounced. Plants most monomorphic are often present in populations representing only one The variety. flower of B. coulteri var. palmeri is illustrated in Spellenberg (2000, fig. but is labelled as B. torreyana; the complex was not understood at the 1), writing of that article. Inflorescences of all the spicate species (B. coulteri included) have observed I when by morning open and are visited various insects in the flowers are pro- ducing moving from and from nectar, the insects inflorescence to inflorescence, more plant to plant. Larger-flowered species are visited frequently (Spellenberg mixed 2000). In populations, particularly, inter-taxon pollination very is likely, and presumed hybridization Both weedy, possible. varieties of coulteri are is B. human responding positively to disturbance, and with the advent of increased may come impact in arid areas of the Southwest, have into increased contact. more Boerhavia coulteri var coulteri geographically the restricted of the is two varieties in the southwestern United States (ranges in Mexico have not been from determined), occurring the eastern to the western borders of Arizona, A New south of the Mogollon Rim. population discovered in south-central may Mexico and in a well-collected area be the result of a recent introduction km Ana (N.M., Doiia Co., e Las Cruces along University Ave. e of Telshor Drive, 1 Aug e of IH-25, in gravel planting strip between sidewalk and road, 24 2001, MX and DUKE, NMC). The Spellenherg Brouillet 13267, var palmeri occurs throughout the Sonoran region in the United States, from southeastern Califor- on Channel nia to southeastern Arizona. occurs Santa Catalina Island in the It Islands of California (Standley 1909, cited as watsoni, Orcutt 12 Sep. 1889, B. s.n., may known where have been introduced. In Texas the var palmeri US!)., it is from a few collections on or near the Rio Grande, the eastern most being from muddy the eastern side of present day Big Bend National Park (M. Young, 5. s.n., SW Aug TEX bank of Rio Grande, near San Vicente mi of Boquillas], 26 19f |5 5, On LL one of these Texas collections, all identified originally as B. [2], [I]). torreyana, pencilled, "but looks just like spicata Choisy Arizona," the writer is B. probably alluding to similarity to coulteri var palmeri of Arizona, as here its B. recognized. Because this the report of this taxon for Texas, the two other is first Canyon Hinckley Co, Capote collections seen are cited here: L.C. 1505, Presidio mi. below 28 May f94f, US; CM. MuUer8438, Presidio Co., along washes 1 falls, of rolling Rio Grande Plain, 4 mi SE of Ruidosa, 28 Jul 1945, LL. In southwest- m New ern Mexico the var palmeri occasional and weedy; the vicinity of Las is may Cruces it is always in disturbed areas. It be introduced there, but if so, it is m & NM, Ana Wooton long-established the region Standley3905, Dofia Co., (e.g., W mesa Organ 4000 Oct TEX, of Mts., 5 1907, LL, US). ft, may The spicate taxa of Boerhavia in the United States be distinguished by the following key. In this key the range of fruit length-to-width ratios is given in the format (length/diam. = 1.3-2.1). = Fruits usually 4-angled(rar ely 5-angled), bPoadly obovoid (length/diam. 1,3-2.1); 1. branches of inflorescence usually densely glandular villous, witinout sticky intern- B.wriqhtiiAGrav obovo = 1. Fruits 5-angled, broadly to narrowly id (length/diam. 1.; inflorescence glabrate to Duberulentbutr glandular, usually with sticky intern- lot 1 smooth orsomewhatrug- Surface of sulci of matijre fruit papillose otherwise 2. ;, ose.the edges of the dges along sulci strongly rugose B. torreyana ri( edges Surface of not instead rugose, the 2. sulci pafDillose, sr ofthe ridges along sulc not rugose to trongly rugose s :i broadly obovc = 1.7-2.1),usuallyoveriapping 3. Fruits in inflo- rescence; sulci and lbs slightly rugo^ sulci usually about 0.5 times as wide ,e; r spicata 6. narrowly obovoid or obpyramic (length/diam. = -3.1),overlapping 3. Fruits iai 2.1 in inflorescence or re>mote; sulci and lbs slightly rugose to smooth;sulci0.1- r wide 0.2 times as as often ove rlapping50-100%of their length, corn- fruits 1 monly±fascicula groups of 2 Tim long.truncate. ite in round-truncate, b luntlyconicorrc)undapically iO% Inflorescence with of their length, 4. mm infrequently 2-3 nacluster.eachf 2-2.4 long,o ccasionally longer, ruit ii rounded palmeri usually e'Pically B. coulteri V,ar ACKNOWLEDGMENTS who acknowledge with gr.ititude the he Ip of Brian Stanrlard (K), seritphoto- I m and photograp leholotypeofSer Hook. copies a hic print of tf ikenhergiacoulti w also thank curators.of herbaria ho loaned specimens for this stud and to f. I iy, Richard Felger for varicDus e-mail "cflats" regarding tfns taxon. REFERENCES Felger, R.S. 2000. Flora of the Gran Desierto and Rio Colorado of northwestern Mexico. Univ. Arizona Press, Tucson. and Kearney,T.H. R.H.Peebles. 1964. Arizona flora, 2nd. ed., with supplement. Univ. Calif Press, Berkeley. W.C New and A Martin, C.R.Hutchins. 1980. flora of Mexico, vol. Cranner, Vaduz. 1. J. MuNZ, PA. and D.D. Keck. 968. A California flora, with supplement. Univ. Calif Press, Berke- 1 Reed, CF. 969. Nyctaginaceae. In: C.L. Lundell, ed. Flora of Texas, vol. 2(1 ).Texas Research 1 Foundation, Renner. -220. Vol. 2(1 pp. 5 1 1 ), Spellenberg,R. 1993. Nyctaginaceae. Hickman, ed.TheJepson manual: higher plants of In: J. 768-774. California. Univ. Calif Press, Berkeley. Pp. SpELLENBERG,R.2000.Blooming"behavior"in five species of fioer/iowo (Nyctaginaceae). Sida 19:311-323. Contr. Herb. 12:303-389. U.S. Natl. 3:377-430. 1 New Standley, PC. 91 8. Allioniaceae. North America flora 21 71 -254. York Botanical Gar- 1 :1 Watson,S. 1889. VI. Contributions to American botany. Upon a collection of plants made 1. by Dr. E. Palmer, in 887, about Guaymas, Mexico, at Muleje and Los Angeles Bay in 1 Lower California, and on the island of San Petro Martin in the Gulf of California. Proc. Amer. Acad. 24:36-87. Arts LC Welsh, S.L, N.D. Atwood, Goodrich, and Higgins 987. A Utah Great Basin S. (eds.). flora. 1 Mem. Brigham Young Naturalist University, Provo. 9. Wiggins, 964. Flora of the Sonoran Desert. Shreve and L.Wiggins, Vegetation and I.L In: 1 F. I. flora of the Sonoran Desert. Stanford Univ. Press, Stanford. 80-1 669. Pt. pp. 2, 1