A Felger, R.S. and S. Rutman. 2015. Ajo Peak to Tinajas Altas: flora of southwestern Arizonaa. Part 15. Eudicots: Fagaceae to Lythraceae. Phytoneuron 2015-59: 1-53. Published 20 Oct 2015. ISSN 2153 733X AJO PEAK TO TINAJAS ALTAS: A FLORA OF SOUTHWESTERN ARIZONA PART EUDICOTS: FAGACEAE TO LYTHRACEAE 15. Richard Stephen Felger Herbarium, University of Arizona Tucson, Arizona 85721 & Sky Island Alliance Box P.O. 41165 Tucson, Arizona 85717 *Author for correspondence: [email protected] Rutman Susan 90 West 10th Street Ajo, Arizona 85321 com tjt@tabletoptelephone. ABSTRACT A floristic and natural history account is provided for 11 eudicot families as part of the vascular plant flora of the contiguous protected areas of Organ Pipe Cactus National Monument, Cabeza Prieta National Wildlife Refuge, and the Tinajas Altas Region in the heart of the Sonoran Desert in southwestern Arizona: Fagaceae, Fouquieriaceae, Gentianaceae, Geraniaceae, Grossulariaceae, Koeberliniaceae, Krameriaceae, Lamiaceae, Linaceae, Loasaceae, and Lythraceae. This is the fifteenth installment of our flora in southwestern Arizona and includes 1 1 eudicot families: Fagaceae (1 species), Fouquieriaceae (1 species); Grossulariaceae (1 species); Koeberliniaceae (1 species); Linaceae (1 species); Lythraceae (1 species); Krameriaceae (1 genus, 2 species); Gentianaceae (2 genera, 2 species); Geraniaceae (2 genera, 3 species); Loasaceae (3 genera, km2 9 species), and Lamiaceae (8 genera, 13 species) (Table 1). The flora area covers 5141 (1985 2 mi of contiguous protected areas in the heart of the Sonoran Desert (Figure This publication 1). is ) also available open access on the website of the University of Arizona Herbarium (https ://ag. arizona.edu/herbarium). The first article in this series includes maps and brief descriptions of the physical, biological, ecological, floristic, and deep history of the flora area (Felger et al. 2013a). This flora includes the modem, present-day taxa as well as fossil records from packrat middens. Explanation of the format APG for the flora series is provided in part 3 (Felger et al. 2013b). Family designations follow III (Angiosperm Phylogeny Group 2009) and the Angiosperm Phylogeny Website (Stevens 2012). Erodium cicutarium and Linum bienne non-native taxa established in the flora area, are marked with , an asterisk (*) and two non-natives not established in the flora area, Salvia reflexa and Punica granatum are marked with double asterisks (**). Fossil specimens are indicated with a dagger , symbol and fossil specimens of the one species no longer present in the flora area {Salvia (f) mohavensis are marked with two dagger symbols (ID- ) All specimens cited are at the University of Arizona Herbarium (ARIZ) unless otherwise indicated by the abbreviations for herbaria at Cabeza Prieta National Wildlife Refuge (CAB), Organ Pipe Cactus National Monument (ORPI), and the standardized abbreviations for herbaria (Thiers 2014). All photos and scans are by Sue Rutman unless otherwise stated and botanical illustrations are by Lucretia Breazeale Hamilton (1908-1986) unless otherwise stated. Descriptions and keys pertain to taxa and populations as they occur in the flora area. & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae Table Local distributions and growth forms of Fagaceae, Fouquieriaceae, Gentianaceae, Geraniaceae 1. = Grossulariaceae, Koeberliniaceae, Krameriaceae, Lamiaceae, Linaceae, Loasaceae, and Lythraceae. t = Modem * = species and fossil specimen(s); fossil no longer present; non-native species established in the j't ** = OP = flora area; non-native species not established in the flora area. Organ Pipe Cactus National CP = TA = SU = Monument; Cabeza Prieta National Wildlife Refuge; Tinajas Altas Region. Summer/warm- = WI = NS = AP season ephemerals; winter-spring/cool-season ephemerals; non-seasonal ephemerals; PR = facultative annuals or perennials; perennials. Growth Form Region Ephemerals Facultative Species annuals or Organ Cabeza Perennials Summer Winter perennials Pipe Prieta Altas seasonal | FAGACEAE OP PR Quercus turbinella f FOUQUIERIACEAE OP CP TA PR Fouquieria splendens f GENTIANACEAE OP PR Eustoma exaltatum OP NS Zeltnera arizonica GERANIACEAE OP CP TA WI *Erodium cicutarium OP CP TA WI Erodium texctnum f Geranium OP WI carolinianum GROSSULARIACEAE OP PR Ribes quercetorum KOEBERLINIACEAE OP CP TA PR Koeberlinia spinosa f KRAMERIACEAE OP CP TA PR Krameria bicolor OP CP TA PR \Krameria erecta LAMIACEAE OP PR Agastache wrightii \Condea OP CP TA PR albida Hedeoma nana OP CP PR f OP PR \Monardella arizonica fTA ' OP CP WI Salvia columbariae OP TA PR Salvia mohavensis ft OP PR Salvia pinguifolia t OP WI **Salvia reflexa CP PR Salvia vaseyi OP CP TA PR mexicana Scutellaria OP CP PR Tetraclea coulteri OP CP NS Teucrium cubense & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae OP CP TA PR Teucrium glandulosum LINACEAE OP WI *Linum bienne LOASACEAE OP CP NS Eucnide rupestris OP CP WI Mentzelia affinis CP TA WI Mentzelia desertorum OP CP WI Mentzelia involucrata OP SU Mentzelia isolata OP CP TA WI Mentzelia hngiloba pubemla CP TA AP Mentzelia OP TA PR Petalonyx linearis OP CP PR Petalonyx thurberi LYTHRACEAE OP PR **Punica granatum FAGACEAE - Beech Family Nearly worldwide; 7 genera, 670 species. - Quercus Oaks Trees and shrubs. Mostly northern hemisphere, not in deserts except sometimes at desert edges; 400 species. Quercus turbinella Greene & O. afoensis C.H. Muller. O. turbinella subsp. ajoensis (C.H. Muller) Felger C.H. Lowe] | Scrub oak; encinillo. Figure 2. m Shrubs or trees to 8 tall. Bark gray, rough and fissured on trunk and larger limbs. Leaves evergreen or ultimately deciduous in late spring-early summer during extreme drought. Leaves cm alternate, often 1-3.5 (5) long, broadly oblong to ovate or rounded, tough, often glaucous bluish- green, mostly glabrous above and glabrate below; margins with spine-tipped teeth or leaves sometimes entire, especially in shade. Flowers unisexual, wind pollinated, small and inconspicuous, without petals; male flowers many in pendulous catkins; female flowers solitary or in small clusters. Flowering in April, the small acorns ripening in summer. Acorns partially surrounded by a cup, cm annual, and on stalks (peduncles) often to 1.5+ long. Ajo Mountains in Alamo and Arch canyons, Bull Pasture, and widespread at higher elevations. In Alamo Canyon and along the Bull Pasture Trail there are small groves of trees with m well-formed, often single trunks. One tree in Alamo Canyon had a trunk nearly 1 in diameter, and Some in such places there is extensive oak-leaf litter. of these trees died as a result of the extended drought of 200 1-2002. At higher elevations on the mountain these oaks are shrubs, often growing among rocks. The nearest known population to the north in the Sand Tank Mountains. is This oak has been in the Ajo Mountains for at least 32,000 years. There are numerous late Wisconsin and early Holocene records for Quercus turbinella throughout sub-Mogollon Arizona mountains (Betancourt et al. 1990). This oak ranges from California to LTah, Texas, and northern Mexico including Baja California and northernmost Sonora. As the common name “scrub oak” indicates, it is generally a shrub. The tree-sized stature of O. turbinella in Alamo Canyon is notable. Elsewhere tree-sized individuals of O. turbinella are usually rare, e.g., near the San Pedro River near . & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythr Winkelman in Pinal County (Phil Jenkins, pers. comm, to Felger, 2005). There are only a few places, New such as the Ajo Mountains, and in the Lower Gila Box along the Gila River in southwestern m Mexico, where more than an occasional individual reaches tree size (more than 5 tall with a well- formed trunk). Like so many oaks, these giants likely share genes with other, related white oaks = Quercus Leucobalanus). (sect. Figure Quercus turbinella. (A) By Lucretia Breazeale Hamilton. Alamo Canyon above Alamo Well: (B) 2. New shoots and flowers, 2 Mar 2005; (C) Galls, 2 Mar 2005 (D) 28 Feb 2009; (E) 7 Feb 20 3 1 1 1 ; Quercus turbinella in the Ajo Mountains was a potential source of edible acorns, which probably were ground into meal and consumed as cakes or gruel. & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae The scrub oak in the Ajo Mountains and nearby southern Arizona mountains was named as a distinct species, Quercus ajoensis, and also as a subspecies of Q. turbinella. Nixon (1997) in the Flora of North America recognized Q. ajoensis as a distinct species, as did Spellenberg (2001) for the oaks of the U.S.-Mexico border region. For the oaks of Arizona, Landrum (1993) treated Q. ajoensis as a synonym of Q. turbinella. The distinctions are technical, involving pubescence, and are not easy Lowe to decipher. Treating Q. ajoensis as a subspecies of Q. turbinella as Felger and (1970) did, , may make sense biologically but is not practical. Among the 400 species of oaks around the world, too often one finds continuous, indistinct boundaries between the species. Distinguishing subspecies or varieties among oaks can be a futile exercise, like constructing a taxonomic house of cards and inviting dissension among botanists. Botanical taxonomy can be contentious and political (see Acacia, Fabaceae, in the present flora series), but it is the essential poetry of the diversity of life. OP: Alamo Canyon, 19 Mar 1933, Shreve 6200. Along wash in main fork of Alamo Canyon, tree with & scaly thin gray bark and sprawling branches, 2675 15 Aug 1952, Muller 9519 Tucker (isotypes of Q. ft, NMC, ajoensis, ARIZ, NY, ORPI). Bull Pasture Trail, 3000 Bezy 25 Oct 1964. Arch Canyon, canyon ft, bottom, 3 May 1978, Bowers 1303. Upper reaches of Alamo Canyon, bottom of ravine, 30 Mar 1989, Baker 7601 (ASU). fAlamo Canyon, twigs, leaves, acoms, 8130 to 32,000 ybp (6 samples), fMontezuma’s Head, twigs, leaves, acoms, 13,500 to 21,840 ybp (4 samples). FOUQUIERIACEAE - Ocotillo Family A family of one genus. - Fouquieria Ocotillo Spiny shrubs or trees. Dry regions of Mexico, 11 species, with 1 extending into the USA. Includes the boojum tree or cirio, Fouquieria columnaris. The system of branching and spine formation in Fouquieria is unusual but not entirely unique. For example, convergence occurs among Sesamothamnus in the Pedaliaceae, native to eastern and southern Africa, and Fouquieria. Some Sesamothamnus and Fouquieria have similar leaf-spine formation, and some of the species of Sesamothamnus approach some Mexican Fouquieria species in gross morphology including growth Fouquieria splendens Engelmann subsp. splendens Ocotillo; ocotillo melhog. Figure 3. ; m Unique long-lived desert “shrubs” often 2-3+ with slender, few-branched, wand-like tall, spiny branches arising from a very reduced trunk or more often trunkless. Long shoots produced during warmer weather and high soil moisture, such as during summer rains and especially in the wake Long of a fall hurricane-fringe storm, or occasionally in spring. shoots with alternate, widely spaced nodes, these leaves with well-developed petioles, which form a rigid spine from the petiole and leaf-blade midrib (Figure 3C). Short shoots in axils of the spines, their intemodes extremely reduced and not apparent, with leaves clustered and sessile, appearing at almost any time of year following a ground-soaking rain except during freezing weather, quickly becoming yellow and 3A shedding as the soil dries or with freezing weather (Figure and F). Inflorescences of densely 19-24 cm flowered panicles at branch tips, often (6) long, with conspicuous leafy bracts falling at anthesis. Flowers bright red-orange, the corolla fused below into a tube, with 5 reflexed lobes. Flowering mostly late February-March, attracting hummingbirds; fruits ripening in late spring. Fruit a capsule, with 6-15 flat, papery-winged seeds. Ubiquitous across the region on desert flats and rocky slopes to the summits of most of the desert peaks, including the Crestline of the Ajo Mountains, and also on the Pinta Sands and locally on sand flats. 6 Widespread across the Sonoran, Mojave, and Chihuahuan deserts to elevations well above the deserts; two other subspecies occur in the Chihuahuan Desert in Mexico (Henrickson 1972). The oldest Sonoran Desert records for ocotillo are from the late Holocene. Considering the ecological amplitude (range of habitats) and wide geographic range of this species, its apparent late arrival and rarity in packrat midden assemblages are somewhat surprising. Nectar can be sucked from the flower base for a trailside snack (Barrows 1900; Felger et al. & 1992). The flowers were steeped in water for a beverage (Bean Saubel 1972). Nectar pressed from & the flowers hardens when dry and was chewed as a delicacy (Castetter Bell 1942). The Cahuillas & parched and ground the seeds to make a gruel or cakes (Bean Saubel 1972). Ocotillo was used as a framework for traditional Hia-Ced brush houses (Betty Melvin in Zepeda 1985: 34). Living fences & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae & & made from the stems (Bean Saubel 1972; Castetter Bell 1942) continue to be popular in the region. OP: Growler Well, Nichol 27 Apr 1939 (ORPI). Growler Mts, foothills, 16 Apr 1952, Parker 7986. Quitobaquito, 29 Mar 1988, Felger_88-132. Trail from The Cones to Mount Ajo, 4090 10 Apr 2005, Felger ft, & (observation). fPuerto Blanco Mts, on ridge, spines, 130 3440 ybp. CP: Observations: Agua Dulce Mts, Cabeza Prieta Tanks, Charlie Bell Pass, Granite Pass Tank, Half- NE m N ENE way Tank, of Tule Well, 12-15 Jun 1992, Felger. 100 of Camino del Diablo 5.5 mi of Papago Well, 300 m, granitic hill with crude rock-walled circle at summit used by smugglers as a lookout, many stems with leafy, young long-shoots, flowers bright red-orange, common and widespread, 26 Mar 2010, Felger 10- 119. TA: Camino del Diablo, SE of Raven Butte, 25 Oct 2004, Felger 04-15. Tinajas Altas Mts, near the summit, 26 Oct 2004, Felger (observation), fButler Mts, stem fragment with spines, 740 ybp. GENTIANACEAE - Gentian Family Annual to perennial herbs, mostly glabrous (those in the flora area). Leaves mostly opposite, simple, usually entire, often clasping the stem; stipules none. Flowers usually bisexual and radial, often showy, 5-merous (those in the flora area), the calyx persistent, the calyx and corolla each united below. Fruit a capsule; seeds minute, and numerous. Worldwide; 88 genera, 1675 species. Mostly in moist, cool, often montane, arctic or alpine habitats; few in deserts and hot climates. Apart from two other Zeltnera species at the desert edge, ours are the only gentians in the Sonoran Desert. cm Leaves glaucous blue-green; corollas white, the tube 2-3 long, anthers not twisting 1. Eustoma 1. Leaves green, not glaucous; corollas bright pink, the tube 1-2 cm long, the anthers twisting after the flowers open Zeltnera - Eustoma Prairie Gentian, Catchfly Gentian This genus has a single species. Eustoma exaltatum (Linnaeus) Salisbury ex G. Don subsp. exaltatum hawan Catchfly prairie gentian; ta:tad. Figure 4. Annuals or short-lived perennials. Stems erect and leafy, 30-75 cm tall; herbage glaucous blue-green, winter deciduous. Leaves opposite, 2-6 cm long, the lowest leaves short petioled, the stem leaves sessile, clasping, and often fused basally, obovate to broadly elliptic or oblong, relatively cm thick and firm, the margins entire. Inflorescences of panicles. Flowers showy, 2.5^1 long, with mostly long pedicels. Calyx deeply cleft, the lobes slender and keeled. Corollas shiny cream-white cm or very pale blue, with 5 large, erect to partially spreading lobes. Anthers yellow. Capsules 1-1.5 mm long, oblong, thickened and tough, rough-surfaced, and many-seeded. Seeds 0.5 wide, rounded, deeply pitted, gray, and iridescent. Flowering during the warmer months. common Locally in alkaline and saline wet soil at Burro, Quitobaquito, and Williams springs. Not known elsewhere in the flora area. USA Three subspecies, southern to northern South America, and the West Indies. Across much of range the corollas of Eutoma exaltatum are variously blue to purple. Subspecies its exaltatum with white petals is known as forma albiflorum Benke. Selections of subsp. exaltatum and E. exaltatum subsp. russellianum are grown as garden plants and there are a number of horticultural selections. & Felger Ri ' Arizona Flora, Pt. 15, Fagaceae to Lythraceae OP: Quitobaquito: Mearns 7 Feb 1894 (US); Harbison 27 Nov 1939 (SD); 17 Mar 1945, Darrow 2401 Wet slopes near springs, flowers whitish or very pale blue, 25 Nov 972, Pinkava 9995 (ASU). Williams 1 ; common Spring, locally at springs seeping from edge of low hills, 13 Sep 1986, Felger_86-268. & Figure Eustoma exaltatum. Quitobaquito: (A) Sep 2008; (B) 10 Sep 2007; (C D) 13 Sep 2006. 4. 1 1 - Zeltnera Centaury A Annuals. North and South America; 25 species. genus segregated from Centaurium. G Zeltnera arizonica (A. Gray) Mansion [Centaurium arizonicum (A. Gray) A. Heller. C. calycosum (Buckley) Femald var. arizonicum (A. Gray) Tidestrom] Arizona centaury. Figure 5. Ephemerals, 12-80 cm tall, the stems erect, single to much-branched; herbage green. Leaves 1-4 cm sessile, long, lanceolate or oblanceolate to ovate or obovate, with entire margins. Inflorescences of terminal cymes. Flowers showy; pedicels often about as long as the calyx tube. mm Calyx lobes linear, separate nearly to the base. Corollas bright pink, 5-lobed, the lobes 7-12 long and spreading, the throat yellow-green (elsewhere in southern Arizona the corollas are sometimes white). Filaments and style white, the anthers and stigma yellow; anthers twisting spirally after mm cm dehiscence. Capsules long, oblong, the walls thin and smooth. Seeds 0.35-0.45 long, 1 ovoid, minutely reticulate and pitted, gray, and iridescent. Flowering March-November. & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae Figure 5. Zeltnera arizonica. Quitobaquito: (A) 3 May 2008; (B-D) 5 Mar 2005. Locally abundant in alkaline and saline wet soil at seeps and springs from Quitobaquito to Williams Spring. Not known elsewhere in the region. USA Alkaline wetlands in southwestern and northern Mexico. OP: Quitobaquito: Fonts 15 Jun 1949 (ORPI); 4 May 1978, Bowers 1308\ 3 Sep 1980, Nabhan 146\ May 23 Jul 1986, Felger_86-212m 23 1991, Baker 8357 (ARIZ, ASU). Williams Spring, 13 Sep 1986, Felger , GERANIACEAE - Geranium Family Ephemerals (those in the flora area; perennials and shrubs elsewhere). Leaves opposite, with stipules. Inflorescences umbellate or cymose. Flowers radial, the sepals and petals 5 and separate. & SW Felger Rutman: Arizona Flora, Pt. 15, Fagaceae to Lythraceae IQ Fruit a capsule, beaked and distinctive by its elastic dehiscence and separation of the segments (mericarps). Both hemispheres, largely tropical and temperate, especially Africa; 14 genera, 730 species. The garden and greenhouse geraniums belong to the genus Pelargonium. Ornamental geraniums are sometimes cultivated in Sonoran Desert gardens but they suffer from the extreme heat of summer and are frost sensitive. 1. Flowers with 5 functional stamens (with anthers); fruits including the beak (2.6) 3-5 cm long; Erodium widespread cm 1. Flowers with 10 functional stamens; fruits including the persistent style column to 2.5 long; Ajo Mountains Geranium - Erodium Stork’s bill Winter-spring ephemerals. First leaves in a basal rosette. Flowers lavender-pink. The long, corkscrew-like beak on each fruit segment uncoils when moistened and screws the sharp-pointed and heavier seed-bearing end into the ground. Mostly Old World; 75 species. Several are important forage plants and some Old World Young perennials are cultivated as ornamentals in temperate regions, especially in rock gardens. plants of the two Sonoran Desert species were eaten as greens, fresh or cooked, by the Cahuillas and & others (Bean Saubel 1972; Hodgson 2001). Leaf blades pinnately dissected, much more than twice as long as wide, the petiole shorter than the 1. Erodium blade cicutarium Leaf blades ovate to 3-lobed or 3-parted, less than twice as long as wide, the petiole longer than 1. Erodium texanum the blade * Erodium cicutarium (Linnaeus) L'Heritier ex Aiton Filaree, heron's bill, stork's bill; alfilerillo', hoho'ibad. Figure 6. Stems sometimes to 30+ cm, usually much shorter or essentially stemless (dwarfed rosette plants). Herbage with glandular and non-glandular white hairs, the glands minute. Leaves pinnatifid, mostly twice divided, much longer than wide, the blades 2-15+ 0.8-4 cm, the petioles 8-7.5 cm 1. mm long. LTmbels (1) 3-7-flowered. Flowers 11-14 wide, the petals lavender-pink, readily falling, mm slightly longer than the sepals. Fruiting sepals 5-6.5 long. Beak of fruits 2. 7-3. 8 cm long. m During highly favorable conditions the plants occasionally reach 2 across with lower leaves to 27 cm long, peduncles 8-14 cm long, and fruit beaks 3.5M.5 cm long. Widely scattered and thoroughly established across the flora area; washes, gravelly flats, bajadas, rocky slopes to higher elevations, and especially disturbed habitats such as roadsides. Widespread in the Sonoran Desert. Native to the Mediterranean region, widely naturalized and weedy in non-tropical regions worldwide. OP: Alamo Canyon. Nichol 14 Mar 1939. Growler Canyon (E of Bates Well), 19 Mar 1975, Lehto L18312 (ASU). Aguajita Wash, 3 Mar 1992, Felger 92-121 (ARIZ, DBS). Quitobaquito, 6 Apr 1988, Felger m 88-279. Trail above Bull Pasture, 15 below Crestline, 4090 10 Apr 2005, Felger 05-296. ft, CP: San Cristobal Wash. 20 Mar 1992, Harlan 19. Charlie Bell Pass, 3 Apr 1992, Whipple 391 7. Redtail Tank, 24 Feb 1995 (CAB). Griffin TA: Coyote Water, 21 Feb 2005, Felger 05-143. Tinajas Altas Mts, above the tinajas, 19 Mar 1998, Felger (observation).