AGAVE DELAMATERI (AGAVACEAE) AND ITS ROLE IN THE SUBSISTENCE PATTERNS OF PRE-COLUMBIAN CULTURES IN ARIZONA
WENDY C. HODGSON AND LIZ SLAUSON
Desert Botanical Garden, 1201 N. Galvin Parkway, Phoenix, AZ 85008
Summary. Agave delamateri from central Arizona was first discovered by Susan McKelvey but never formally described by William Trelease. Rick DeLamater rediscovered the agave almost 60 years later, and it is now formally described as a new species. Throughout its range in central Arizona it is found in direct or indirect association with pre-Columbian agricultural artifacts and other features associated with habitation sites. These plants may represent living archaeological assemblages, possibly persisting for more than five centuries after their presumed cultivation.
Introduction and Taxonomic History
Susan D. McKelvey collected numerous Agave specimens in central Arizona during the late 1920's and early 1930's. Among these were a series of specimens (McKelvey 786, 787, 937, 938, 1075 [photo 17-2], 1016, 1080, 1081; GH) of an unknown agave collected in and around the Sierra Ancha Mountains, Globe, and Coolidge Dam, Gila County. Unsure of the identity of these specimens, McKelvey sent them to William Trelease for study. In a letter to McKelvey, William Trelease (1929) tentatively recognized her collections as a new species, calling it "Agave repanda" based on the obvious sigmoid, or s-shaped leaves. In response to Trelease's letter, McKelvey (1929) noted that collections 1074 and 1075 were obviously very different from "A. repanda" and instead represented A. couesii Engelm. ex Trel. (A. parryi Engelm. var. couesii (Engelm. ex Trel.) Kearney and Peebles). In 1964 Gentry annotated McKelvey 1074 as A. parryi var. parryi (GH). However, McKelvey 1075 is a mixed collection including two sets of photographs; one photograph (17-1) appears to represent A. parryi var. parryi, while the other (17-2) clearly depicts a different plant, "A. repanda". McKelvey also collected a "wider-leaf form with large marginal teeth" in the Pinal and Mazatzal mountains (McKelvey 1038,1039; GH). Trelease called this form "A. repanda var. pinalensis" suggesting that it linked "A repanda" with A. couesii (A. parryi var. couesii). Gentry annotated these specimens as A. parryi var. parryi (GH), but based upon our examination they suggest A. parryi strongly introgressed with A. chrysantha.
Even though McKelvey suggested that Trelease publish "A. repanda" (McKelvey, 1929), he did not do so. In 1964, Howard S. Gentry annotated McKelvey 937, 938, 1080, and 1116 as A. aff. palmeri (GH), and listed these specimens as A. palmeri Engelm. in his monograph (Gentry, 1982). In our opinion, he mistakenly listed "A. repanda" under synonymy with A. chrysantha
Almost 60 years later, in 1988, while hiking in Tonto Basin in Gila County, Rick DeLamater, a graduate student of botany at Arizona State University, rediscovered Trelease's "A. repanda", although he was unaware of this at the time. Subsequent surveys by DeLamater and the authors have resulted in the documentation of over 90 sites. As the name "A. repanda" was misapplied to three different taxa and never published, we choose not to recognize Trelease's epithet. We name this species A. delamateri in honor of our dear friend Rick DeLamater (1954— 1989). We also propose the common name, Tonto Basin agave, for this species.
Fig. 1. Agave delamateri, Hodgson s.n., near Punkin Center, Arizona. Plant, x 1/12; leaf, x 2/3; flower cluster, x 3/5; flower, longitudinal section, x 1 3/5. Note the relatively deep and wide tube and equal insertion of the filaments.
Affinities and Relationships
A. delamateri is most appropriately placed within the Group Ditepalae sensu Gentry (1982) based on its deep floral tube with short, leathery, persistently erect, and unequal tepals (Figs. 1, 2). It appears to be most closely related to A. fortiflora Gentry and A. palmeri and is distinguished by its numerous rhizomatous offsets, easily cut leaves, and 1- , not 2-seriate filaments. It further differs from A. fortiflora in having glaucous purple-tinged leaves, greenish-ochroleucous, apically rusty-maroon tepals and slightly flattened, maroon rather than strap-shaped, yellow filaments. In addition, it differs from A. palmeri in having broadly lanceolate, apically incurved rather than lanceolate or linear-acuminate straight leaves, more numerous lateral branches in the inflorescence, and in its wider perianth tube and longer tepals (Figs. 1-5; Table 1).
A. delamateri also differs from known hybrids between A. chrysantha and A. parryi, taxa which may occur with A. delamateri. Specimens which suggest hybridization between A. chrysantha and A. parryi include (McKelvey 1038, 1039, GH; Hodgson 7373, 7374, 7375, 7376, 7377, 5815, 6329, 6330, 6331, 6332, DES; Ecker 0159, 0158, 0157, 0156, 1055 and 0160, DES). A delamateri differs from these variable specimens in its consistently lanceolate, acuminate leaves which are erect and inwardly flexed at the apex, more numerous teeth, generally taller and more narrow inflorescence with fewer lateral branches, and consistently cream-colored, erect, rugose-tipped tepals.
Distribution
The distributions of A. delamateri, A. palmeri, and A. fortiflora are allopatric. A. delamateri is known only from a small geographic area in central Arizona, while A. palmeri is known from southern Arizona and southwestern New Mexico into northern Sonora; A. fortiflora is only known from small, isolated mountain ranges in northwestern Sonora (Gentry, 1982).
A. delamateri is known from approximately 90 separate sites, occurring in Gila County from near Coolidge Dam and Globe north to Young and west to Sunflower in Maricopa County, and near Oak Creek in Yavapai County (Fig. 6), just south of the Mogollon escarpment, at elevations ranging from. 725 to 1554 meters. The greatest concentration of sites occurs along the south end of Tonto Creek near the northwest end of Roosevelt Reservoir in an area referred to as Tonto Basin, situated between the Sierra Ancha and Mazatzal Mountains. The vast majority of sites occur in the Arizona Upland subdivision of the Sonoran Desert as defined by Shreve (1951). Only a few sites are found in Interior Chaparral and Great Basin Conifer Woodland associations as defined by Brown (1982). Plants are generally found on open, level to moderately steep slopes of alluvial fans or dissected old terraces, often overlooking major drainageways (Hodgson, 1994b). Soils are commonly cobbly and gravelly, deep, and well-drained (Hendricks, 1985).
Fig. 2. Floral ideographs of A. delamateri and other agaves, based on Gentry (1982). All measurements are taken from flowers during or soon after anthesis; D = measurements taken from dried, pressed flowers, all others are from fresh flowers. Numbers are Hodgson collection numbers except D, H = DeLamater and Hodgson and G = Gentry. Key: a = tube width, b = tube height, c = outer tepal width, d = outer tepal length, and e = bar indicating filament insertion, all in mm.
Agave delamateri Hodgson and Slauson, sp. nov.
A Agave fortiflora Gentry et A. palmeri Engelmann similis, sed ab ambabus surculis rhizomatosis numerosis et filamentis 1-, nec 2-seriatis, ulterius ab A. fortiflora foliis glaucis purpureo-tinctis, tepalis ochroleuco-virentibus apice purpureo-rubiginosis, filamentisque purpureis nec luteis, ulterius ab A. palmeri foliis late lanceolatis apice incurvis, nec lanceolatis vel lineari-acuminatis rectis, necnon perianthii tubo latiori tepalisque longioribus diversa.
Type. ARIZONA. Gila County: foothills of Sierra Ancha Mountains, above Tonto Basin, 5.25 miles E of State Route 188 and Punkin Center, 160-250 m SE of Forest Service Road 71, T6N, RUE, S 5, NE1/4., 33°53'45"N latitude, 111°14'55"W longitude, ca. 975 m; Hodgson 5478, 5 July 1989 (holotype, DES; isotype, ASU).
Fig. 3. Agave delamateri near Punkin Center, Arizona, showing habit, rhizomatous offsets and characteristic leaf shape and color.
Plants medium to large, acaulescent, rosettes caes-pitose, ca. 1 m high, 1 m wide. Leaves erect, inwardly arcuate at apex, mostly 50-63(-74) cm long, 7.5-9 cm wide (excluding marginal teeth), broadest near or just below middle, lanceolate, acuminate, guttered above, bluish-gray glaucous with purple-maroon tinge and green cross-banding, easily cut near base, margins straight to repand; terminal spine 28-35(-49) mm long, brownish-gray, decurrent for 1/5-1/6 of leaf, 1-1.5 mm wide, brownish-red; marginal teeth variable, with additional interstitial teeth usually below middle of leaf, dark glossy brown to gray and pruinose (especially towards leaf apex), 1.5 (near leaf base)-l 1(—40) mm apart, the larger teeth 3.5-5 mm long, usually reflexed, becoming porrect near leaf base, the smaller teeth 1-1.5 mm long, mostly porrect to upwardly flexed. Inflorescence paniculate, broad, open, 4.5-6 m tall, pruinose but purple-maroon beneath, with 12-17 widely spaced, long, lateral branches in upper 3/5-5/8 of shaft, perpendicular to shaft; pedicels 4-5 mm long. Flowers in clusters of 14-20, large, durable, 47-67 mm long, pale cream tinged with light green; ovary 21-29 mm long, 7-10(-13) wide, slightly angled longitudinally, cream-green; neck 1-3.5 mm long, slightly constricted; tube 11-16 mm deep and wide; outer tepals ovate, longer and narrower than inner tepals, 14-18 mm long, 6-7 mm wide, light cream-green with maroon-rust, rugose, hooded tips; inner tepals broadly ovate, 9-15 mm long, 5-7 mm wide, broadly ovate, strongly keeled, tips white-villous, slightly hooded; tepals clasping filaments during and after anthesis; stamens 6, filaments 36-53 mm long, equally inserted 5-8 mm below apex of tube, maroon; anthers attached to filament off-center. 11-20 mm long, bright yellow; pistil cream-colored, maroon-flecked, becoming more maroon towards stigma; stigma cream tinged with maroon, trigonous, with whitish pubescence along slits. Capsules short-stipi-tate, beaked, oblong, 35 mm long, 8-12 mm wide. Seeds unknown.
Fig. 4. Agave delamateri near Punkin Center, showing habit and characteristic leaf shape.
Fig. 5. Agave delamateri; note the inflorescence with widely spaced lateral branches perpendicular to the main shaft.
Ethnobotany
Agaves have been an important source of food, fiber, and beverage for many indigenous cultures. Recently, extensive prehistoric agave cultivation in south-central Arizona was deduced from the strong association of roasting pits and mescal knives with water and nutrient enhancement features such as numerous rock piles, man-made terraces, and checkdams covering tens of thousands of hectares of desert bajada slopes (Fish et al., 1985, 1990, 1992). A. delamatenalso persists in or near indigenous cultural landscapes, rather than landscapes unmodified by past cultivation (Hodgson, 1994a, 1994b). Plants are found in direct or indirect association with archaeological features including single and multi-room foundations, agave knives, pot sherds, linear alignments (Fig. 7), rock piles, and check-dams (Figs. 8, 9).
We believe A. delamateri was grown for food or fiber by pre-Columbian cultures (Hohokam-Salado-Mogollon) inhabiting the area northeast of the Verde River and south of the Mogollon Rim from approximately 1000 A.D. to 1450 A.D. Its role in peoples' subsistence patterns appears to be similar to the situation described for A. murpheyi Gibson and the Hohokam along the Gila and lower Salt River valleys (Hodgson et al., 1989; Hodgson, 1994a, 1994b). Following the breakdown or fragmentation of these pre-Columbian cultures during or after the 15th century, plantings of A. delamateri were presumably no longer cared for and left to grow "wild". Individual plants or clones may have been harvested later for food and fiber by roving Apache or Yavapai tribes until the 1870's, when these groups were relegated to military control.
Fig. 6. Agave delamateri in Arizona. A single dot may represent one clone or more than one clone nearby (Arizona map based on Kearney and Peebles 1964: Yuma County is now divided into La Paz and Yuma County).
Fig. 7. Pre-Columbian linear alignment in Haufer Wash below Hodgson 5486, 5487, 5488, 3.2 miles north of Punkin Center, Arizona.
The topography and climate of an area influenced not only the type of agave planted by pre-Columbian cultures but also how these agaves were managed. Agave murpheyi occurs in the Lower Colorado Valley subdivision of the Sonoran Desert, an area which is characterized by gently sloping bajadas and an annual precipitation of only 80-250 mm. Here it is found on low-gradient slopes and benches in direct association with runoff features (such as linear rock alignments, rockpiles, and checkdams), which presumably were used to increase soil moisture and organic matter retention. In contrast, A. delamateri is primarily found in the Arizona Upland subdivision of the Sonoran Desert, an area largely characterized by small mountains, hilly plains, and old dissected terraces. Annual precipitation varies between 250-410 mm in Tonto Basin and the surrounding areas. In these habitats, A. delamateri is found only on top of the alluvial terraces overlooking drainage systems; apparently there was little or no effort to construct runoff and channeling systems for the cultivation of A. delamateri. Agricultural techniques may have been altered due to more favorable amounts of precipitation and the highly dissected nature of the terrain.
As a result of increased populations, competition for farm land on the floodplain may have led to exploitation of areas for farming on higher terraces by pre-Columbian cultures (Wood and McAllister, 1984). Numerous extensive linear alignments and checkdams were constructed in the drainages above the floodplain, presumably for less drought-tolerant annual crops, such as maize, beans, and squash (Hodgson, 1994a, 1994b). Agaves, which can tolerate less water and nutrients, were grown above the drainages and farther from the floodplain.
There are several present-day agave distributions that may have been influenced by man. Gentry (1982) suggested that A. applanata was disseminated from Veracruz and Puebla as far north as Durango and Chihuahua "by man's hand in historic or prehistoric times". Sporadic localities in the northern part of A. applanata's distribution are found along the old Native American trail between central Mexico and Casas Grandes, Chihuahua (Gentry, 1982). Localities of A. murpheyi are also sporadic and in direct association with archaeological features in central Arizona (Hodgson et al., 1989; Hodgson, 1994a). We believe this agave may have originated farther south than its present range (Hodgson, 1994a) and was transported, transplanted, or traded by the Hohokam. An unknown agave (Hodgson 8357; DES) was recently found growing amidst archaeological features in Deer Creek Canyon, Grand Canyon National Park, Coconino County. This area was inhabited and farmed extensively by the Anasazi (Robert Euler, pers. comm., August 1994) and possibly historically by the Southern Paiutes (Helen Fairly, pers. comm., August 1994). Whether or not this unknown agave represents an offset from a recently planted clone or from clones planted long ago after being dispersed northward via trade is unknown. We hypothesize that A. delamateri may have also originated elsewhere, perhaps farther south in Mexico, and was transported to its present day range. Planted and grown for prized attributes, such as its robust size, easily cut leaves, and profuse offset production, pre-Columbian peoples may have gradually dispersed A. delamateri northward through migration. The rhizomatous offsets could have been easily carried or traded, then planted.
Pre-Columbian peoples who utilized agaves for food or fiber probably managed plants to favor vegetative regeneration by "castrating" them, or cutting out the embryonic reproductive structures to prevent flowering, similar to the present day practices of some mescaleros (Tello-Balderas and Garcia-Moya, 1985;
Fig. 8,9. Pre-Columbian checkdams in unnamed drainage below Agave delamateri near jeep trail 67. 0.5 miles north of Sycamore Creek, 3 miles south of Punkin Center, Arizona. Drawings and photos by W. Hodgson.
Reproductive Biology
Inflorescences of A. delamateri begin to emerge in May and early June, and mature in late June through July with flowers on the lowermost lateral branches opening first. Flowering within any given population or clone is synchronous. Anther dehiscence and peak nectar production occur during early morning hours, when the air is saturated with a heavy musky fragrance. Flowers usually abort early, or infrequently remain on the pedicels and become hardened and woodlike.
Fruit production has been observed in only one clone of A. delamateri (Hodgson 8889; DES), although plants produce profuse rhizomatous offsets. A recent collection (Hodgson 6038; DES) consists of flowers with enlarged ovaries containing immature small white seeds. Two specimens of McKelvey's (938 and 7075; GH) have notes that read, "see fruit specimen". McKelvey 938 includes a photograph of a plant with an old flowering stalk with the upper three umbels bearing dried perianth parts on immature fruits. McKelvey 1075 is a mixed collection with two photos, one depicting A. parryi var. parryi, the other depicting A. delamateri. The "fruit" in question may well belong to A. parryi var. parryi.
Several factors may contribute to the low frequency of fruit set in A. delamateri. Self-sterility is one possibility, as agaves are primarily obligate outcrossers (Gentry, 1982; Sutherland, 1982; Slauson, unpubl. data). If A. delamateri at a single site represents plants derived from a single genetic clone, these plants would be largely incompatible and produce little or no fruits/seeds. If individual plants from different sites represent different genotypes, the spatial isolation (large distances between the majority of plants) and temporal isolation (flowering in different years) would significantly decrease the chances of cross-pollination occurring between sites (different genotypes). Another agave with a limited and sporadic distribution with low fruit set is A. parryi var. huachucensis (Baker) Little ex Benson. Sutherland and Delph (1984) found that fruit set in this taxon was only 3.7% and suggested that it is probably a clonal inbreeding population (Sutherland, 1982).
Lack of pollinators may also reduce sexual reproduction, but it is most likely not a factor in A. delamateri. Large numbers of insects and birds have been observed to visit flowers and contact stigmas. Although the musty odor of flowers is suggestive of bat pollination, recent work has shown that adequate pollination and subsequent fruit set does occur in A. palmeri and A. chrysantha (species with similar musty floral scents) when visited by diurnal animals only (Slauson, in press). However, pollination studies may shed light on other problems, such as whether distances between plants inhibit cross-pollination.
Climatic factors may affect the mode of reproduction in agaves and are not well understood. Growth and reproductive strategies in cultivated agaves are often altered by environmental conditions different from those of their natural habitats. Certain species which readily offset in habitat have been observed to remain solitary' in cultivation, and vice versa, at the Desert Botanical Garden. Some species which normally flower, produce fruit, and rarely, if at all, offset in their native environments have been observed to abort flowers and produce offsets and/or bulbils from vegetative shoots in cultivation. If A. delamateri originated from a site further south in Mexico and was dispersed northward by pre-Columbian people, it may require different temperature and/or moisture regimes for flower and fruit development. Gentry (1982) notes that flower and seed development is inhibited for a number of agaves due to the intensity of summer temperatures when grown in desert gardens in Arizona. Stamens may fail to expand and dehisce due to insufficient turgor pressure to extend the filaments (Gentry, 1982). In A. delamateri, filaments were observed to remain folded while anthers dehisced within the tenals It is possible that flower and fruit development are inhibited due to climatic conditions where A. delamateri is presently known to occur.
The amount of viable pollen produced influences the degree of successful fertilization events, and pollen stainability percentages can provide an indication of pollen viability. In A. delamateri, pollen stainability percentages were low, ranging from 5% to 18%. Sterility (or partial sterility) may be a problem, as both non-viable micro-pollen grains (7-46%) and viable macro-pollen grains (12-26%) were observed. These findings suggest that A. delamateri may be of hybrid origin or a polyploid. Hybridization and polyploidy are common in Agave and appear to play significant roles in the evolution of the genus (Pinkava and Baker, 1985). Polyploidy can arise and become established more easily among species which are perennial and reproduce vegetatively, as they are better able to survive the period of partial sterility that occurs before polyploids become stabilized (Stebbins, 1950). Further cytological work is needed to evaluate meiosis in A. delamateri.
Despite the fact that A. delamateri has produced little fruit or seed, it appears to have persisted for centuries through asexual reproduction. During this time it has also played a significant role in the subsistence patterns of pre-Columbian people in the arid Southwest.
Additional Specimens Examined
The following were collected by Susan McKelvey (M), Gila County, Arizona, in May and June, 1929, deposited at GH: Sierra Ancha Mountains, M 786, 787, photos; road to Carr's Ranch: M 1075, photo 17-2; M 937, lf, photo; M 938, lf, photo; M 1116 lf. Road from Globe to Coolidge Dam, M 1080, lf, photo, M 1081, lf, photo.
The following were collected by Rick DeLamater (D) or Wendy Hodgson and Liz Slauson (H), Tonto Basin/Roosevelt Reservoir area, Gila, Yavapai, and Maricopa counties, from primarily May through July, 1988 to 1993; all are deposited at DES with some duplicates at ASU (D 325, D 307, D 306, D 305, D 335, D 356, D 101, D 102, D 103, D 104, H 5642, H 5644, H 5652, H 5653, H 5641, H 7371, H 5486, H 5473, H 5477, H 5538), UTEP (H 5736, H 5474), and NY (H 5007): GILA CO: TIN R15E, SE1/4 of S 9, 3300 ft. H sn. lf, photo; NE1/4 of S 16, 3300 ft, H sn, lf, photo; T2N R15E: SW1/4 of S 18, 3180 ft, H 6088, lf, photo; just S of Murray Wash. 2900 ft, H 6087, lf, photo; NW1/4 of S 20, 3200 ft, H 6083, lf, photos, 6083, lf, photo; S 29, ca. 1/3 mile E of State Route (SR) 88, 3260 ft, H 6069, lf, immature fr, photo; extreme SE1/4 of S 29, 3200 ft, H 6084, lf, photo, H 6085, lf, photo, H 6086, lf, photo; NE1/4 of S 30, 3180 ft, H 6090, lf, photo, H 6089, lf, photo; T3N R14E: NW1/4 of S 4, 3/4-1 mile NW of Salt River bridge, 2300 ft, H 6694, lf; T4N RUE: NW1/4 of S 5, off Forest Service Road (FSR) 445 A, N side of road, 3250 ft, D 101, lf, photo; T5N R10E: S ?, 0.5 mile E of milepost 257 along SR 188, 2500 ft, D 103, lf, D 102, lf, D 104, lf, photo; T5N R11E: SE1/4 of S 6, 5.5 miles S of Punkin Center, 1 mile SE of Sycamore Creek/SR 188 junction, 2400 ft, H 5725, lf; T5N R12E: NE1/4 of S 9, 4200 ft, D 309, lf; NE1/4 S 13, 3800 ft, D 305, lf, old fls, photo; S 21, 3400 ft, D 325, lf, infl, photo; SE1/4 of S 24, 6.4 miles N of SR 88 along SR 288, 4420 ft, H 6768, lf, fls, NE1/4 of S 25, just NE and upslope from SR 288, 4900 ft, H 7371, lf; S 33, 2900 ft, D 306, lf, photo, D 307, lf, photo; T6N R10E: SW1/4 of S 10, 2200 ft, D 302, lf, photo; NE1/4 of S 15, 2520 ft, H 5474, lf, fls, H 5473, lf, fls; NW1/4 of S 26, 3/4 mile SW of Tonto Basin Post Office, 2600 ft, H 5736, lf; T6N RUE: SW1/4 of S 1, 3960 ft, H 5477, lf, fls; S 2, 3800 ft, D 331, lf, old infl, photo; S 3, 3500 ft, D 335, lf, old infl, photo; center of S 5, 2760 ft, H 5556, fls, photo, H 5557, lf, fls, photo; NE1/4 of S 6, 0.7 miles W of FSR 71, one mile E of FSR 60, 2920 ft, H 5482, fl, fls, photo; center of S 6, 0.7 mile W of FSR 71, 1/3 mile E of FSR 60, 2760 ft, H 5483, lf, photo; center of S 8, 2600 ft, H 5007, lf; south of center of S 15, 3400 ft, D 329, lf, photo; NW1/4 of S 12, 3960 ft, H 5475, lf; NE1/4 of S 22, 3300 ft, D 330, lf, photo; T7N R10E: SE1/4 of S 3, 2720 ft, H 5723, lf; SE1/4 of S 4, 2800 ft, D 298, lf, old infl, photo; SE1/4 of S 4, ca. 1 mile W of SR 188, along jeep trail above Cottonwood Creek, H 5481, lf, fls; center of S 9, 1/3 mile S of Cottonwood Creek, 3/4-1 mile W of SR 188, 2840 ft, lf, 5642, lf, 5644, lf; center of S 12. 2700 ft, H 5020, lf, fls; S 12, 2400 ft, D 356, lf; SW1/4 of S 22, ca. 1 mile E of SR 188, just S of Slate Creek, 2540 ft, H 5630, lf; SE1/4 of S 27, ca. 1 mile N of milepost 265, E side of SR 188, 2560 ft, H 5538, lf, fls; SW1/4 of S 27, 3.3 miles N of Reno Creek-SR 188 junction at Punkin Center, 2500 ft, H 5735, lf; S 28, along Haufer Wash, 2700 ft, D 332, lf, old infl, photo; NE1/4 of S 28, ca. 1 mile E of SR 188, just south of Slate Creek, 2600 ft, H 5631, lf; SE1/4 of S 29, 3300 ft, D 299, lf, old fls, old infl, photo; NE1/4 of S 34, ca. 1/2 mile W of SR 188, N of Buena Vista Creek, milepost 265, 2600 ft, H 5487, lf, fls, photo, H 5488, lf. fls, photo; SW1/4 of S 35, just W of SR 188, N of Buena Vista Creek, 2600 ft, H 5486, lf, fls, photo; T7N RUE: N edge of S 5, 3000 ft, D 338, lf, fls, old infl, 339, lf, photo; S 5, 3300 ft, D 341, lf, old infl, D 342, lf, photo; S 34, 3400 ft, D 336, lf, old infl; T7N R14E: SW1/4 S 8, ca. 1.9 miles S of SR 288 along FSR 203 (Cherry Creek Road), 5160 ft, H 6343, lf; T8N R10E: SE1/4 of S 2, ca. 5 miles N of SR 188 and Jakes Corner, 2 miles NE of FSR 184, 100 yards S of jeep trail, just W of Indian Farm Well, 3080 ft, H 5652, lf; NE1/4 of S 11, ca. 4.5 miles N of SR 188 and Jakes Corner, 1/2 mile SW of Indian Farm Well, 1/6 mile W of Tonto Creek, 2880 ft, H 5653, lf; NW1/4 of S 17, ca. 1/3 mile N of SR 188 and 3/4 mile E of SR 87, 2900 ft, H 5847, lf, fls; SW1/4 of S 33, ca. 1/2 mile SW of Reed Gulch, 3340 ft, H 5645, lf; NE1/4 of S 34, ca. 300 yards W of SR 188, just N of drainage which enters Gold Creek, 2840 ft, H 6038, lf, older fls; SE1/4 of S 35, ca. 1 mile E of SR 188, 2.5 miles SE of Jake's Corner, 3120 ft, H 5629, lf; T9N R12E: SW1/4 of S 24, ca. 7 miles W of Young, 5200 ft, H 6097, lf, photo.
YAVAPAI CO: 2.5 miles W of Interstate 17, just south of Stagecoach Historical Marker, adjacent to Dry Beaver Creek, 4150 ft, H 8889, lf, cap, photo.
MARICOPA CO: T6N R9E NE1/4 of S 18, ca. 1/2 mile NW of Sunflower, 3500 ft, 11 April 1990, H 5728, lf, fls, photo.
Acknowledgments
The authors thank Emily Wood, Jean Boise and Judith Warnement, Harvard University Herbaria, and Rupert Barneby, New York Botanical Garden, for their kind assistance; also Donald Pinkava, Jon Rebman, Tony Burgess, Richard Felger, Joe McAuliffe, and Edward Anderson for their encouragement, suggestions, comments and thought-provoking discussions. We are grateful to the Desert Botanical Garden for providing support for this research.
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Haseltonia, 1995