Felger, R.S., S. Rutman, M. Costea, D.F. Austin, and J. Malusa. 2015. Ajo Peak to Tinajas Altas: A flora of southwestern Arizona: Part 9. Eudicots: Convulvulaceae. Phytoneuron 2015-2: 1-22. Published 7 January 2015. ISSN 2153 733X AJO PEAK TO TINAJAS ALTAS: A FLORA OF SOUTHWESTERN ARIZONA PART 9. EUDICOTS: CONVOLVULACEAE - MORNING GLORY FAMILY Richard Stephen Felger Herbarium, University of Arizona Tucson, Arizona 85721 & Sky Island Alliance P.O. Box 41 165 Tucson, Arizona 85717 * Author for correspondence: rfelger@email.arizona.edu Susan Rutman ■■■ ••• I |.."' ii.. i Ajo, Arizona 85321 i|i . i il l' i --I-I- !■ | l' hi Mihai Costea Dept. of Biology Wilfrid Laurier University 75 University Avenue W Waterloo, ON, N2L 3C5, Canada mcostea@wlu.ca Daniel F. Austin Arizona- Sonora Desert Museum .■'.I I ' I II I ii- i. M i.i ' "4. & Herbarium, University of Arizona, Tucson dr_ipomoea@yahoo.com Jim Malusa School of Natural Resources and the Environment University of Arizona Tucson, Arizona 85721 malusata 'em ail. arizona. edu ABSTRACT A floristic account is provided for the morning glory family 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 Sonoran Desert in southwestern Arizona. The family includes 1 1 species in 4 genera (Cuscuta, Evolvulus, Ipomoea. Jacquemontia) in the flora area, all of which are native. This is the ninth contribution for this flora published in Phytoneuron and also posted open access on the website of the University of Arizona Herbarium (ARIZ). The first part of the Flora of southwestern Arizona flora series includes maps and brief descriptions of the physical, biologica l, ecological, floristic, and deep history of the flora area (Felger et al. 2013a) and explanations for the format for the species accounts is given in part 3 (Felger et al. 2013b). These contributions are also posted open access on the website of the University of Arizona Herbarium (ARIZ). Vernacular, or common names, when known, or deemed worthwhile, are given in English, Spanish, and the Hia C-ed O'odham dialect, respectively. The one fossil specimen is marked with a dagger symbol (|). Area designations are: OP = Organ Pipe Cactus National Felger et al: Southwestern Arizona Flora, Convulvulaceae 2 Monument, CP = Cabeza Prieta National Wildlife Refuge, and TA = Tinajas Alias (Figure 1), 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 (Index Herbariorum, Thiers 2014). All microphotographs of Cuscuta are by Mihai Costea and all photos are by Sue Rutman unless otherwise stated. For more images see the galleries provided in the Digital Atlas for the Convolvulaceae of Sonora (Costea & Austin et al. 2012). Descriptions and identification keys pertain to taxa and populations as they occur in the flora area. Figure 1. Flora area in southwestern Arizona. Area designations are: OP = Organ Pipe Cactus National Monument, CP = Cabeza Prieta National Wildlife Relugc ind 1 \ = linens AJta^ Green shading m'taites approximate boundary of federally designated wilderness. CONVOLVULACEAE - Morning Glory Family tpheii.til ti p rennial 1 mo md subshiubs oi herbaceous perennials (those in the flora area); or obligate parasites with little or no chlorophyll and vegetative parts much reduced in Cuscuta. Leaves alternate, simple, entire to deeply parted, and without stipules. Flowers radial, 5-merous (those in the flora area), and often showy. Calyx persistent. Corollas sympetalous, those in flora area (except Cuscuta) with the parts folded within the bud (plicae) being glabrous, whereas those parts exposed in the bud (interplicae) form a star-shaped pattern in the expanded corolla and are often pubescent. Fruits of dry capsules: seeds 1-4 (6). The Convolvulaceae constitute a diverse cosmopolitan assemblage of 58 genera and an estimated 1880 species (Staples 2014). This monophyletic family has greatest diversity in tropical and subtropical regions worldwide and does not generally occur in higher latitudes, and diversity also Felger et al: Southwestern Arizona Flora, Convulvulaceae decreases with aridity. Detailed information for the convolvs of Arizona and Sonora is provided by Austin (1998), Costea, Austin, et al. (2012), Costea, Felger, et al. (2012), and Felger and Austin, et al. (2012). Eleven species in four genera occur in the flora area of southwestern Arizona (Table 1). All occur in Organ Pipe, three extend into Cabeza Prieta, and none range into the hyperarid Tinajas Altas Region. Taxon Region Growth Form Organ — Pipe — Pneta Tinajas Ephemeral Perennial CONVOLVULACEAE OP SU OP CP SU OP SU Cuscuta tuberculata OP SU Cuscuta umbellata var. umbellata OP CP SU : i.i... .-. ■ OP PR Ipomoea barbatisepala OP SU Ipomoea costellata OP SU Ipomoea cristulata OP SU Ipomoea hederacea OP CP SU f Ipomoea sp. (OP) Jacquemontia pringlei OP PR Totals 11 3 0 9 0 2 Table 1. Local distributions and growth forms of Convolvulaceae for the flora in southwestern Arizona, "f = Fossil taxon. OP = Organ Pipe Cactus National Monument OP = Cabeza Prieta National Wildlife Refuge, TA = Tinajas Altas Region. 1. Parasitic twining ephemerals without chlorophyll; stems appearing leafless (leaves reduced to scales) and orange; flowers 5 mm or less in length or width and white Cuscuta 1. Plants green, not parasitic; stems leafy and not orange: flowers 1 cm or more in width or length and not white (except Jacquemontia). 2. Stems not vining; leaves entire, mostly less than 1 cm wide, sessile or very short petioled Evolvulus 2. Stems vining; leaves entire or lobed, more man 1 cm wide and petioled. 3. Ephemeral vines; leaves entire, lobed, or divided; corollas various colors but not while; stigmas lobes globose (the stigma is a 2 -lobed sphere); pollen grains spiny (visible with, a dissecting microscope) Ipomoea 3. Perennial subshrubs with twining upper stems; leaves entire; corollas white, becoming pale lavender-pink as the flower ages during the day; stigma lobes oblong; pollen grains smooth Jacquemontia Cuscuta - Dodder; fideo; vepegi vasai Parasitic twining ephemera Is with orange stems, leaves reduced to minute, alternate scales, and small white flowers mat become creamy when dried (those in the flora). Flowers 5-merous (those in the flora area; some are 4-merous elsewhere). Growing with warm weather, especially during late spring and the summer rainy season. All the species in the flora area are members of subg. Grammica (Costea et al. 2015; Garcia et al. 2014; Stefanovic et al. 2007). The two species with Felger et al: Southwestern Arizona Flora, Convulvulaceae 4 indehiscent (not circumscissile) capsules, C. californica and C. salina, belong to sect. Californicae (Costea et al. 2015; or "clade A", Stefanovic et al. 2007); they have somewhat thicker and more robust stems, and parasitize shrubby perennials or herbaceous plants. The species with dehiscent capsules, C. legitime.., C. tubercuiata, and C. umbeiiata, belong to sect. Umbeiiatae (Costea et al. 2015; previously called "clade L", Stefanovic et al. 2007); they have thinner and more delicate stems, and grow on annuals (ephemerals) and small perennial herbs. Cuscuta is nearly cosmopolitan and comprises ca. 200 species (Costea 2007; Costea et al. 2015). The taxonomy of this large and diverse genus is complicated (Costea et al. 2015; Stefanovic et al. 2007; Yuncker 1932). As in many holoparasitic plants, a number of organs are reduced or absent, and distinguishing features are often minute and not easily seen. Identification during the vegetative stage is possible only by using molecular markers. The minute appendages in Cuscuta flowers called infrastaminal scales are taxonomically useful but require the careful dissection of flowers using a microscope, usually at least 50* power. In the flowers of subg. Grammica, these minute structures likely have the role to protect the ovary against insect herbivory (Riviere et al. 2013). When viewed fresh and with good illumination the floral structures including the scales are quite beautiful. Yet even dried flowers can be hydrated, dissected, and the floral characters discerned. Upon drying at maturity, the dehiscent (circumscissile) capsules open in a clean circle around the capsule towards the base, while the indehiscent capsules may break apart or rupture irregularly. For more information and images of the floral morphology, see the galleries provided in the Digital Atlas of Cuscuta (Costea 2007). Information about pollen, gynoecium, and breeding systems is available in Welsh et al. (2010) and Wright et al. (201 1, 2012). KEY 1. Capsules not needed 1. Perianth, especially the corolla tube papillate; calyx lobes keeled, Vi-Vi as long as the corolla tube; on ephemerals Cuscuta tubercuiata 1. Perianth not papillate (in C. umbeiiata and C. salina papillae or dome-like cells are present only on corolla lobes); calyx lobes not keeled, 3 /4 as long as the corolla tube, or equaling to longer than corolla tube; on ephemerals or perennials. 2. Infrastaminal scales absent or reduced to ridges; Ajo Mountains, often on Calliandra eriophylla Cuscuta californica var. californica _' lull. lM. Illllll.ll sv..iks pusuil. u Jl *kv J<-|kJ 3. Infrastaminal scales slightly shorter than the corolla tube, oblong; styles shorter than the ovary, locally on Suaeda in SW part of Organ Pipe Cuscuta salina 3. Infrastaminal scales equaling or exceeding corolla tube, spatulate to obovate; styles equaling or longer than the ovary; widespread on herbaceous perennials (not on Suaeda) and especially on summer ephemerals. 4. Flowers 4-5.5 (6) mm long 4. Flowers 2-3 mm long Cuscuta legitima Cuscuta umbeiiata var. umbeiiata Felger et al: Southwestern Arizona Flora, Convulvulaceae 5 KEY 2. Capsules needed 1. Capsules not circumscissile, sometimes breaking apart irregularly. 2. Capsules ± globose, not narrowed apically or thickened around the interstylar aperture; 2-4 seeds per capsule; known from the Ajo and Santa Rosa Mountains, often on Calliandra eriophylla Cuscuta californica var. californica 2. C««p«uLs eilii^oiddl, narrowed apically and thickened around the interstylar aperture; 1 seed per capsule (rarely 2); localized on Suaeda at Quitobaquito Cuscuta salina 1. Capsules circumscissile, opening in a clean circle around the capsule towards the base. 3. Calyx lobes keeled, V3-V2 as long as the corolla tube; corolla tube cylindrical; infrastaminal scales conspicuously shorter than the corolla tube, not reaching the filaments C. tuberculata 3. Calyx lobes not keeled, as long as, or longer than the corolla tube; corolla tube campanulate; corolla scales reaching the filaments and often slightly protruding from the corolla tube. 4. Flowers 4-5.5 (6) mm long Cuscuta legitima 4. Flowers 2-3 mm long Cuscuta umbellata var. um bell at a Cuscuta californica Hooker & Arnott var. californica California dodder. Figure 2. Inflorescences loose to dense, paniculate; pedicels 0.5-3.0 mm long. Flowers 5-merous, 3-5 (5.5) mm long, fleshy; papillae absent. Calyx 1.5-2.2 mm long, golden yellow when dried, finely reticulate and shin}', turbinate-campanulate, % to as long as the corolla tube, tube 0.4-0.7 mm long, lobes 0.8-2 mm long, triangular-ovate to lanceolate, overlapping at the base, acute to acuminate. Corolla 3-5 mm long, the tube 1.6-2.4 mm long, cylindric-campanulate to obconic, lobes 2.0-2.6 mm long, narrowly lanceolate, equaling or longer than the corolla tube, initially erect, later reflexed. Infrastaminal scales absent or reduced to ridges. Ovary ovoid to obovoid; styles evenly filiform, 1.2- 2.2 mm long, as long or longer than the ovary. Capsules 1.5-2.2 x 1.2-2.5 mm, indehiscent, ± globose, not narrowed apically or thickened around the interstylar aperture, surrounded (not capped) by the withered corolla. Seeds 2-4 per capsule. References: Costea et al. (2006); Costea & Stefanovic (2009). Plants in the flora area are var. californica characterized by non-papillate flowers and capsules with 2-4 seeds. Often abundant in the Ajo Mountains where it can turn entire slopes orange in April, and visitors to Organ Pipe Cactus National Monument ask about it every year; also in the Santa Rosa Mountains. Often on Calliandra eriophylla but also on a wide range of annuals (ephemerals) and perennials. Washington to Baja California, southern Nevada, and western Arizona. It is unknown from Sonora but is expected in nearby north-central regions of the state. OP: Cement Tank [.Alamo Canyon], [on Calliandra eriophylla., flowers 4.5-5.5 mm, calyx shorter than the corolla tube], 14 Apr 1941, McDougall 97. Alamo Canyon, 28 Aug 1943, Clark 10924 (ORPI). Bull Pasture: 3300 ft, 9 May 1979, Bowers 1712; abundant on Calliandra eriophylla and many other hosts including Calochortus, Cryptantha, Verbena neomexicana, etc, 9 Apr 2005, Felger 05-178. Felger et al: Southwestern Arizona Flora, Convulvulaceae 6 F" -r- E 2: - PM IS 7*1* • ,1 K^l Figui'e 2. Cuscuta californica var. californica. Bull Pasture: (A & B) on Calliandra eriophylla, 7 Mar 2014; (C) 30 Mar 2008; (D) characteristic expansive local population, 11 May 2010. Flower morphology, scale bars = 1 mm: (E) Slower before anthesis; (F) flowers; (G) calyx (not dissected); (H) corolla view r ed from the top; (I) dissected calyx; (J) dissected corolla (note the lack of infrastammal scales); (K) gynoecia. Felger et al: Southwestern Arizona Flora, Convulvulaceae 7 Cuscuta legitima Costea & Stefanovic [C. umbellata var. reflexa (J.M. Coulter) Yuncker] Figure 3. Figure 3. C«KWfe legitima. (A) Mex Hwy 8, 35 km S of Sonoyta, 12 Sep 2014. (B) Carlsbad, Eddy Co., NM, 3 Nov 2011, © photo by Terry Gregston (Costea 2007). Flower morphology, scale bars = 1 mm: (C) flower; (D) calyx, not dissected; (E) dissected calyx; (F) dissected corolla; (G) gynoecium. Inflorescences dense to loose, umbelliform; pedicels 2-10 mm long. Flowers 5-merous, 4- 5.5 (6) mm long, membranous; papillae absent. Calyx 2.5-3.2 mm long, straw-yellow, finely reticulate, slightly shiny, campanulate, longer than corolla lube, divided % the length, the calyx tube 0.6-1 mm long with lobes 1.5-2.2 mm long, not basally overlapping, ovate-lanceolate, not carinate, the apex (calyx lobes) acuminate. Corollas 3.8-5.2 (5.6) mm long, the tube 1.6-2.1 mm long, campanulate, the lobes 1.8-3 mm long, initially erect, later reflexed, longer than the tube, linear- Felger et al: Southwestern Arizona Flora, Convulvulaceae 8 lanceolate, margins entire, apex acuminate. Infrastaminal scales 1.8-2.2 mm long, equaling or slightly longer than the tabs, bridged at 0.2-0.4 mm, spatulate to ob ovate, fimbriae 0.2-0.5 mm long. Ovary globose; styles 0.9-2.5 mm long, longer than the ovary. Capsules 2-3 x 1-2 mm, circumscissile, depressed, irregularly thickened and slightly elevated around the inconspicuous interstylar aperture, translucent, surrounded or capped by the withered corolla. Seeds 2-4 per capsule. Cuscuta legitima is a hybrid species and its putative parents are C. umbellate* var. umbellata on the paternal side and C. odontolepis (or some other unknown related species) on the maternal side (Costea & Stefanovic 2010). This relationship explains the morphological similarity between C. ... .*;. ; ; , \\. jikI ■ --. ........ \ ,\\ --. ........ Known from Organ Pipe by a single record and from Cabeza Prieta by two records, but probably more widespread. On herbaceous hosts. Arizona to Kansas, New Mexico, and Texas, and Mexico in Baja California, Sonora, Chihuahua, Coahuila, and Tamaulipas. OF: Estes Canyon, in wash, on L %onum deflexum 1 Uig 2014 A; tman 20140826-6. CP: 1 mile S of Bat es Well Road on road to Jose J uan Tank at boundary of the Re fuge and Organ Pipe, sandy desert plain, on Boerhavia erecta, B. spicata, Bouteloua aristidoides, B. barbata, Chamaesyce polycarpa, Kallstroemia califon i, K. grandifl i idestrom J m gii < 15 Sep 1992, Felgei ~4 \ < v Daniels Arroyo at Charlie Bell Road, on on Kallstroemia sp and Tidestromia lanuginosa, 24 Aug 2014, Rutman 20140824-6. Cuscuta salina Engelmann Saltmarsh dodder. Figure 4. Inflorescences dense, corymbiform; pedicels (0.5) 1-5 mm long. Flowers 5-merous, 2.5^.5 mm long, fleshy, papillae or dome-like cells present on the corolla lobes. Calyx 1.5-2.5 mm long, glossy yellow, cylindric to narrow campanulate, equaling corolla tube, tube 0.6-1.2 mm long, lobes 0.7-1.5 mm long, ovate-lanceolate to lanceolate, not basally overlapping, acute to acuminate. Corolla 2.2^.0 mm long, the tube 1.2-2 mm long, cylindric-campanulate to obconical, the lobes 1.3-2 mm long, ovate-lanceolate to oblong-lanceolate, equaling the corolla tube, initially erect, later spreading or reflexed, apex acute to acuminate or cuspidate (sometimes appearing tri dentate). Infrastaminal scales 1-1.8 mm long, 80-90% of the corolla tube length, oblong to slightly obovate, bridged at 0. 18- 0.45 mm, fimbriae 0.03-0.20 mm long. Ovary ellipsoid, the styles 0.4-0.9 mm long, shorter than the ovary. Capsules 1.6-2.5 x 1.7-2.2 mm, indehiscent or irregularly dehiscent, ellipsoid, narrowed at the top and thickened around the small interstylar aperture, surrounded or capped by the withered corolla. Seeds 1 (2) per capsule. References: Costea et al. (2006, 2009). Reproductive June-December. Localized along the southwest boundary of Organ Pipe from the vicinity of Quitobaquito to Hocker Well and in adjacent Sonora near Sonoyta, growing on Suaeda nigra [S. moquinii]. Cuscuta salina occurs in Baja California, New Mexico, Texas, Utah, and southern Arizona where the populations apparently are few and widely disjunct and in adjacent northern Sonora, Host plants are Frankenia, Salsola, Suaeda, and Wislizenia. OP: Hocker Well, 20 Dec 2001, Rutman s.n. (ORPI). Quitobaquito: [on Suaeda nigra] 29 Nov 1939, Harbison 26193; 8 Oct 1960, Hevfy s.n,; On. Suaeda moquinii in Suaeda flats, 17 May 1982, Nabhan s.n.; Several large colonies on Suaeda moquinii, 19 Jun 1989, Felger 89-241 (ARIZ, ASU). Felger et al: Southwestern Arizona Flora, Convulvulaceae 9 A ^ ■ Figure 4. Cuscuta salina. (A) On Suaeda nigra, Quitobaquito, 1 1 Sep 2008. Flower morphology, scale bars = 1 mm: (B) fragment of inflorescence; (C) flower; (D) calyx : not dissected; (E) dissected corolla; (F) infrastaminal scale removed from the flower; (G) gynoecium. Cuscuta tuberculata Brandegee Desert dodder. Figure 5. Inflorescences loose, umbelliform or racemiform; pedicels 2-3 (5) mm long. Flowers 5- merous, 2.5-^ mm long, membranous; papillae present especially at the bass of the corolla tube. Calyx 0.5-1.5 mm long, yellow, finely reticulate or not ± glossy, campanulate-angular, X A- X A as long as the corolla tube, the calyx tube 0.2-0.5 mm long, the lobes 1-1.3 mm long, not basally overlapping, triangular to lanceolate, carinate and with multicellular protuberances, acute to acuminate. Corollas 2-3.5 mm long, the tube 1.5-2.2 mm long and cylindrical, the lobes 1.2-2 mm long, Vi the length of the corolla tube, bridged at 0.3-0.5 mm, ovate, uniformly short-fringed, fimbriae 0.05-0.15 mm long. Ovary globose; styles 1.5-3 mm long, longer than the ovary, and evenly filiform. Capsules 1.3-2.2 x 1-2.3 mm, circumscissile, globose, slightly thickened and risen around the small interstylar aperture, translucent, capped by the withered corolla. Seeds usually 4 per capsule. Facultative xenogamous (Wright et al. 2012). Reference: Costea & Stefanovic (2010). Felgeretal: Southwestern Arizona Flora, Convulvulaceae 10 Figure 5. Cuscuta tuberculata. (A <& B) OnBoerhavia, Alamo Canyon, 7 Sep 2013. (C) Gunsight Hills near E boundary of Organ Pipe, 10 Sep 2013. Flower morphology, scale bars = 1 mm: (D) flowers; (E) dissected calyx; (F) calyx, not dissected; (G) detail of calyx lobe carena (carinate keel); (H) corolla (note the papillae on the corolla tube), (I) dissected corolla; (J) gynoecium. Felger et al: Southwestern Arizona Flora, Convulvulaceae 1 1 Widespread in lowland desert areas and extending into mountains in Organ Pipe; on summer ephemerals, especially Boerhavia spp. and Pedis papposa. Southwest and south-central Arizona, New Mexico, northern Mexico including Raja California Sur and western Son or a southward at least to the vicinity of Hermosillo. OF: 16.3 mi by road N of Lukeville, disturbed area, low rocky hills, mostly on Boerhavia coidteri, 26 Sep 1964, Felger 10532. Bull Pasture, on Boerhavia, 5 Nov 1977, Bowers 949. N end Cipriano Hills, on Boerhavia, 1 8 Sep 1988, Wirt s.n. Basalt flat near W-faciiig slopes of Santa Rosa Mts, on Boerhavia spicata, 29 Aug 2001, Rutman s.n. Cuscuta umbellataKunthvar. umbellata Figure 6. Inflorescences dense to loose, umbelliform; pedicels 2-10 mm long. Flowers 5-merous, 2-3 mm long, membranous; papillae sometimes present but only on the adaxial face of corolla lobes. Calyx 0.8-1.4 mm long, straw-yellow, finely reticulate, slightly shiny, campanulate, equaling the corolla tube, the tube 0.25-0.6 mm long, the lobes 0.5-0.9 mm long, not basally overlapping, broadly triangular-o vate, not carinate, the margins entire, the apex obtuse to acute. Corollas 2-2.5 mm long, the tube 0.6-1.2 mm long, campanulate, the lobes 0.8-1.5 mm long, initially erect, later reflexed, equaling or slightly longer man the tube, oblong to lanceolate, the apex obtuse to acute. Infrastaminal scales 0.8-1.2 mm long, equaling or slightly longer than the corolla tube, bridged at 0.1 mm, subspathulate to obovate, uniformly dense-fringed, the fimbriae 0.15-032 mm long. Ovary globose; styles 0.8-1.7 mm, equaling or longer man the ovary, evenly filiform. Capsules dreumscissile, 1-2.5 x 0.5-1.2 mm, depressed, irregularly thickened and slightly risen around the inconspicuous interstylar aperture, translucent, surrounded or capped by the withered corolla. Seeds 4 per capsule. Facultative xenogamous to facultative autogamous (Wright et al. 2011). Reference: Costea & Stefanovic (2010). Lowlands scattered across the flora area, especially areas of dense growth of ephemerals such as washes, floodplains, and sand flats. Parasitic on a wide range of small perennial herbs and spring and summer ephemerals including Allionia, Boerhavia, Bouteloua, B ramus, Daucus, Euphorbia (Chamaesyce), Kallstroemia, Dinebra (Leptochloa panicea), Pectis, Plantago, Senecio, Silene, Tidestromia, and Trianthema. Arizona, Colorado, New Mexico, and Texas to South America and the West Indies. OP: Alamo Canyon, on Boerhavia, 28 Aug 1943, Clark 10924 (ORPI). Armenta Well, on Allionia, 1600 ft, 16 Nov 1974 : Warren s.n. (ORPI). Aguajita, on Bouteloua barb ata, Leptochloa panicea, Trianthema portulacastrum, Tidestromia lanuginosa, 14 Sep 1988, Felger 88-414. Just N of U.S. /Mexico boundary E of Lukeville, 3 mi E of Dos Lomiias, on Tidestromia, 29 Aug 2001, Rutman s.n.. Growler Wash floodplain near Bates Well Ranch, 23 Aug 200 L Rutman s.n.. W braid of Cuerda de Lena on Allionia incarnata, 7 Oct 2006, Rutman 20061007-8. CP: 5 km N of Pmta Sands, [on Pectis papposa], 2 Oct 1964, Simmons s.n. LI mi S of Bates Well Road on road to Jose Juan Tank, at E Refuge boundary, on Kallstroemia, Boerhavia, Allionia, 14 Sep 1992, Felger 92-707. Daniel's Arroyo, on Allionia, 21 Sep 1992, Harlan 329. San Cristobal Wash on E. polycarpa, E. albomarginata, 14 Sep 1992, Felger 92-687B. 1 mi S of Bates Well Road on road to Jose Juan Tank, at E Refuge boundary, on Allionia, Boerhavia erecta, B. spicata, Bouteloua aristidoides, Kallstroemia calijbrnica, K grandiflora, Tidestromia lanuginosa, 15 Sep 1992, Felger 92-740. Evol villus Annuals or perennials, stems not twining. Styles 2, bifid, the 4 stigmas long and thread-like. Native to the New World; about 100 species. Four species occur in Arizona (Austin 1988) and 7 species occur in Sonora (Felger & Austin et al. 2012). Felger et al: Southwestern Arizona Flora, Convulvulaceae 12 Figure 6. Cuscuta umbellata var. umbellata. (A) On Boerhavia, San Cristobal Wash near Camino del Diablo, 26 Sep 201 3. (B) On Boerhavia, near Cuerda de Lena and N boundary of Organ Pipe, 30 Sep 2006. Flower morphology scale bars = 1 mm: (C) calyx (not dissected); (D) dissected corolla; (E) flower; (F) gynoecium; (G) dissected calyx. Evolvulus alsinoides Linnaeus [E. a/sinoides var. acapulcensis (Willdenow) van Ooststroom. E. alsinoides var. angustifolia Torrey] Mouse ears; oreja de ratdn. Figure 7. Small tufted perennials, the herbage densely pubescent with appressed and long spreading hairs. Stems very slender and wiry, usually less than 25 cm long, dying back to near ground level in drought. Leaves 7-23 mm long, lance-linear to elliptic, smaller above, sessile or very short petioled. Pedicels slender, 3-6 mm long; calyx 2-2.5 mm long. Corollas 8-10 mm wide, rotate, pale blue with a white center, gl abrate except the prominent star-like interplicae. Ovary, style, and anthers pure white, the filaments pale blue. Flowering at various seasons; flowers diurnal — opening in mid- morning and closing in the afternoon. Capsules globose, 2.5-3 mm diameter, 1-4-seeded. Flowering in warmer months with sufficient moisture. Widely scattered in Organ Pipe in washes, canyons, and on rocky slopes. Felger et al: Southwestern Arizona Flora, Convulvulaceae 13 Figure 7. Evolvulus alsinoides. (A) Estes Canyon, 21 Sep 2008. (B) Salero Ranch, 8 mi ESE of Tubac, Santa Cruz Co., 19 Feb 2014, photo by Sue Carnahan. (C) Alamo Canyon, 9 Sep 2013. (D) Owl Canyon, Florida Mts, Luna Co., NM, 25 Aug 2008, photo by Patrick Alexander. Native from Arizona to South America, and now a pantropical weed. This species is extremely polymorphic. Van Ooststroom (1934) recognized fifteen varieties around the world, although the variation is often not well defined. Sonora, Arizona, New Mexico, and Texas plants belong to var. angusttfolia Torrey. This variety typically has shorter sepals, narrower leaves, and less densely pubescent stems, and tends to have shorter stems than other varieties, if indeed the variety is worthy of recognition. At least a few populations in Asia and Australia appear to have differentiated Felger et al: Southwestern Arizona Flora, Convulvulaceae 14 since post-Columbian times into potentially new infraspecific taxa, in a manner akin to the English sparrow in North America. OP: Alamo Canyon, Nichol 26 Mar 1938 Sonoyta Hills, 27 Aug 1943, Clark 10885 (ORPI). 2 mi due W of Sweetwater Pass and 0.5 mi SE of Interpretive Marker 1 8, 31°59'20"N. 1 12°4'55"W, 630 m, E-facmg hillside, to 20 cm tall, growing from the bases of shrubs and through them, local but abundant, flowers lavender-blue. Ambrosia deltoidea, Cereus giganteus, Calliandra, Coursetia, Simmondsia, Bursera microphylla, Opuntia acanthocarpa. 1 Mar 1989* Baker 7710 (ASU). Bull Pasture, 9 Apr 2003, Felger (observation). Ip.MII.KM X I . • ■ 1 1 1 1 1 -_- I .1-1 \ . ...... The three morning glories in the flora area are ephemeral vines growing with hot-weather rains. Leaf characters in the key refer to the upper or mid-stem leaves. Fruit a several-seeded capsule. Worldwide, most diverse in tropical and subtropical regions; perhaps 600 species. Arizona has 13 species (Austin 1998) and Sonora has 41 species (Felger & Austin et al. 20 12). 1. Leaves pedately dissected into 5-9 linear or linear-lanceolate segments Ipomoea costellata 1. Leaves entire or shallowly to deeply 3-5 lobed, the lobes broad (not linear or linear-lanceolate) and ii'-l |Kj.ikl l . jiuii-.^J 2. Corollas red-orange; sepals glabrous Ipomoea cristulata 2. Corollas blue; sepals with coarse hairs. 3. Lower leaf surfaces usually with glandular dots; sepals 10-12 mm long; sepal bases and pedicels glabrous Ipomoea barbatisepala 3. Lower leaf surfaces without glandular dots; sepals 12-24 mm long; sepal bases and pedicels pubescent Ipomoea hederacea Ipomoea barbatisepala A. Gray Figure 8. Vines, the stems low-climbing, glabrous. Leaves glabrous, often with glandular dots at least on the lower surfaces, deeply 5-7 lobed, the lobes lanceolate and narrowed toward base, acute to acuminate. Sepals 10-12 mm long, 1-2 mm wide, the bases slightly dilated, hispid-pilose throughout, the apex elongate narrowly linear, typically erect to spreading. Corollas 1.6-2 (2.5) cm long, blue-purple with a white throat. Capsules glabrous. In the flora area known from ephemeral watercourses in Bull Pasture and probably along other drainages in the higher elevations of the Ajo Mountains. The nearest known population is in the Baboquivari Mountains; it extends into the margins of the Sonoran Desert but not into the drier, lower desert elevations. Arizona, New Mexico, Texas; Baja California Sur, Sonora, Sinaloa, and disjunct to Oaxaca. In many ways this species resembles Ipomoea hederacea, which may grow with it, but /. barbatisepala is more delicate, the leaf lobes are narrower, acuminate, and deeply cut, and the corollas are typically smaller. The hirsute pubescence going the length of the sepals and the deeply lobed leaves are diagnostic. OP: Bull Pasture, 19 Sep 201 4, Rutman 20140919- 7. Felger et al: Southwestern Arizona Flora, Convulvulaceae 15 Figure 8. Ipomoea barbatisepala. (A & B) Salero Ranch, Santa Cruz Co., 28 Aug 2013, photos by Sue Carnahan (SEINet). (C & E) Bull Pasture stream, 19 Sep 2014. (D) Brown Canyon, Baboquivari Mts, 5 Sep 2009, photo by Dan Austin. Ipomoea costellata Torrey Crest-rib morning glory. Figure 9. Stems trailing or twining. Leaves pedately dissected into 5-9 linear or linear-lanceolate segments. Sepals conspicuously crested and warty. Corollas 10-12 mm long, 5-10 (14) mm wide, pale lavender to pink. In the flora area known only from the Ajo Mountains; it is common in Bull Pasture. Eastward in Arizona to west Texas, most of Mexico including Baja California Sur, and Guatemala. OF: Bull Pasture, 1 2 Aug 1 990, Wirt s.n. (ORPI). Saddle between Arch Canyon and Boulder canyons, 26 Oct 2003, Rtitman 20031026-17 (ORPI). Bull Pasture area, 3170 ft, temporally wet or moist drainage on bedrock, cryptic annual growing amid dense summer growth [including] Evolvulus, Ipomoea hederacea, I, cristulata, etc., 25 Sep 20\3,Rutman 20130925-5. Figure 9. Ipomoea costellata. Bui! Pasture, seasonally wet stream below the spring, 25 Sep 2013. Ipomoea cristulata Hairier f. [1. coccinea of authors, not Linnaeus. Quamoc lit gracilis Hallier f.] Scarlet morning glory. Figure 10. Delicate to moderately robust but small vines. Leaves variable, with a cordate base, otherwise entire or shallowly to broadly 3 (5) lobed. Calyx glabrous, the lobes linear-subulate. Corollas 1-1.5 cm wide, trumpet-shaped (salviform), bright red-orange. Flowers diurnal and pollinated by butterflies and hummingbirds, however, apparently also capable ofselfLng (facultatively autogamous). In the flora area known from the Ajo and Diablo Mountains. Midwestern United States (where presumably cultivated) to Arizona and Texas, and Mexico south to Veracruz; and in Baja California Sur. OP: Alamo Canyon, 13 Dec 1939, Harbison 26244 (SD). Arch Canyon, Wirt 13 Oct 1990. S fork of Alamo Canyon, 2400 ft, abundant m wash bed and xeroriparian area, 7 Sep 2013, Rutman 20130907- 1. Diablo Mts, 2647 ft, shaded base of N-facing cliff, 22 Sep 2013, Rutman 20130922-13. Felger et al: Southwestern Arizona Flora, Convulvulaceae 17 Figure 10. Ipomoea cristulata. Alamo Canyon: (A & D) 7 Sep 2013; (B) 9 Sep 2013; (C) 17 Oct 2013, (E) By Lucretia Brezeale Hamilton. Ipomoea hederacea Jacquin Ivy-leaf morning glory. Figure 1 1 . Vines; often robust and covering shrubs and small trees, often twining 2-3+ m in mesquites; densely to sparsely hairy throughout. Leaf blades (2) 3.5-13 (15.5) cm. long and about as wide, 3- lobed to 3- or 5-parted. Calyces 15-25 mm long, the lobes lanceolate, narrowed above the base, the expanded, basal portion with long spreading hairs. Corollas opening wide, often 3-3.5 cm wide, light blue with a white base, open in the early morning and visited by white-lined sphinx moths {Hyles lineata) and butterflies, and closing with mid-morning heat. Capsules 9 mm wide. Seeds 4.5-4.8 mm long, dark brown to blackish, wedge-shaped, with minute hairs. Often locally abundant following summer rains, growing through mesquites and other shrubs along large washes and canyon bottoms. Ajo Mountains and the northern part of Organ Pipe, and Felger et al: Southwestern Arizona Flora, Convulvulaceae 18 especially the larger, brushy washes such as the San Cristobal and Growler Wash systems in the eastern part of Cabeza Prieta where it often festoons mesquites in green curtains (Felger 2007). Figure 11. Ipomoea hederacea. (A) Armenta Ranch, 12 Sep 2008. Alamo Canyon: (B) 7 Sep 2013; (D) 9 Sep 2013. (C) Catalina Mts, Pima Co, Sep 1989, photo by Dan Austin. (E) Climbing on Prosoj iveh tin i mldlift tank near N boundary of Organ Pipe, E of Hwy 85, 20 Sep 2013. Temperate North and South America; introduced in the Old World and also likely introduced into South America. OP: Alamo Canyon, 28 Aug 1943, Clark 10939 (ORPI). Bull Pasture Trail, 11 Sep 1988, Wilson 189. Road to Armenta Ranch, 1 Feb 2002, Felger (observation, dry, dead plants). CP: Monreal Well,Monson 19 Sep 1959. San Cristobal Wash: 14 Sep 1992, Felger 92-693. Felger et al: Southwestern Arizona Flora, Convulvulaceae 19 tlpomoea sp. OP: tAlamo Canyon, fruit, 1 150 ybp. Jacquemontia Perennials or occasionally annuals; 80-100 species in the Americas and several in Australia. Four species in Arizona (Austin 1 988) and five in Sonora (Felger & Austin et ai. 2012). Jacquemontia prirsgki A. Gray Desert cluster-vine. Figure 12. Pviern a's sc md^nt ot spuwlins suhvhtubs *\nh -4-6 i -s\ etl stdhte b Mrs the upper stems often twining, growing and flowering during the warmer months. Leaves 2-6 cm long, petioled, the blades ovate with entire margins. Inflorescences of axillary cymes with 1-7 flowers. Outer sepals broadly ovate, the inner sepals narrower. Corollas about 2 cm wide, white, becoming pale lavender as the flower ages during the day. Figure 12. Jacquemontia pringlei. Alamo Canyon, south-facing slope: (A & E) 10 Sep 2008; (Bj 7 Sep 2008; (C&D)7Sep2013. Felger et al: Southwestern Arizona Flora, Convulvulaceae 20 In the flora area known from the Ajo Mountains, at least in Alamo Canyon. This is the most arid and farthest western locality verified for the genus. This species ranges from southern Arizona to northwestern Sinaloa and southwestern Chihuahua; it is widespread through Sonora except the northwestern corner of the state. This genus has its greatest diversity in tropical and subtropical Latin America. An enigmatic specimen collected by A.A. Nichol on 25 April 1938 at the "South end of Gila Mountains" seems out of place and the locality questionable. Before the mid-20th century it was customary to include the Tinajas Altas Mountains in the concept of the Gila Mountains. On that same day Nichol collected Muhlenbergia dumosa in the "Mohawk Mountains" and Aristida purpurea in the "Gila Mountains" (specimens at ARIZ). There are no other records for Jacquemontia or Muhlenbergia dumosa in Yuma Count}' (Felger & Van Devender et al. 2012; SEINet 2014). Richard has looked for them in possible habitats in the Gila-Tinajas Altas Mountains. OP: Ajo Mts, 21 Nov 1934, Goodding 227. Alamo Canyon: 28 Aug 1943, Clark 10945 (ORPI); Van Devender 31 Aug 1978 (ORPI). Ajo Mts, mouth of Alamo Canyon, 2200 ft, 7 Sep 1947, Fonts 198 (NMC). LITERATURE CITED Austin, D.F. 1998. [Vascular plants of Arizona] Convolvulaceae, morning glory family. J. Arizona- Nevada Acad. Sci. 30: 61-83. Costea, M. 2007- onward. Digital Atlas of Cmcuta (Convolvulaceae) Wilfrid Laurier University, Ontario. Costea, M., D.F. Austin, RS. Felger, T.R Van Devender, and J. Sanchez-Escalante. 2012. Convolvulaceae of Sonora, Mexico [Digital atlas]. University of Arizona Herbarium. Iillp .iv ji i i ^Jii Ik i l\u in in n. -vk X" Costea, M, RS. Felger, D.F. Austin, T.R. Van Devender, and J.J. Sanchez-Escalante. 2012. Convolvulaceae of Sonora, Mexico. II. 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Flora of Tinajas Altas, Arizona - A century of botanical forays and forty thousand years of Neotoma chronicles. J. Bot. Res. Inst. Texas 6: 157-257. Garcia, M.A., M. Costea, M. Kuzmina, and S. Stefanovic. 2014. Phylogeny, character evolution, and biogeography of Cuscuta (dodders; ConvoKulaceae) inferred from coding plastid and nuclear sequences. Amer. J. Bot. 101: 670-690. Riviere, S..' C. Clayson, K. Dockstader, M.A.R. Wright, and M. Costea. 2013. To attract or to repel? Diversity, evolution and role of the "'most peculiar organ" in the Cuscuta flower (dodder, Convolvulaceae) — the infrastaminal scales. Plant Syst. Evol. 299: 529-552. SEINet. 2014. Southwest Environmental Information Network, http://swbiodiversity.org/ seinet/index.php> Staples, G.W. (ed.). 2014. Convolvulaceae Unlimited. Stefanovic, S., M. Kuzmina, and M. Costea. 2007. Delimitation of major lineages within Cuscuta subgenus Grammica (Convolvulaceae) using plastid and nuclear DNA sequences. Am. J. Bot. 94: 568-589. Thiers, B. 2014 [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium.