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Home , Arecaceae, Brahea, Brahea aculeata, Brahea armata, Corypheae, Coryphoideae

Aliso, l8(1), pp. 1-18 @ 1999, by The Rancho SantaAna Botanic Garden, Claremont, CA 91711'3157

THE PALMS () OF ,

RrcHaRoS. Fslcsn Drylands Institute 2509 North Campbell #405 Tucson, 85719 e- mai I : rfeI g e r @ag. arizona. e du

AND

Elene Jovar-r Department of Arizona State University Tempe, Arizona 85287 e-mail: [email protected]

ABSTRACT

This publication is an account of the palms (Arecaceae) in the state of Sonora in northwestern Mexico. Six native palm in three genera are recorded for Sonora: with four species, and and each with one species. Relationships and within Brahea, especially B. elegans, remain unresolved. Brief botanical descriptions, taxonomic synopses, local names, identification keys, conservation status and recommendations, and distributional and ethno- botanical infbrmation are provided for each species. Basionyms and types are cited. Distributions are documented with citations of nearly all specimens known to us from Sonora. Many palm populations in this arid and semiarid region are threatened by development and in some cases over- exploitation. There is an urgent need for formal management and conservation action for local pop- ulations, although as a whole none of these species in Sonora are rare or immediately threatened.

RESUMEN

Esta publicaci6n es un estudio de las palmas (Arecaceae) en el estado de Sonora, en el noroeste de M6xico. Se registran en Sonora seis especies nativas en tres g6neros: Brahea con cuatro especies, y Srtbttl y Washingtonia ambos con una especie. Quediindo sin resolverse las alinidades y la taxonomia dentro de Brahea, especialmente B. elegans. Para cada especie se proporcionan breves descripciones. sin6psis taxon6mica, nombres locales, claves para la identificaci6n, estado y recomendaciones de conservaci6n e informaci6n sobre la etnobotdnica y distribuci6n. Se presentan basi6nimias e intbr- maci6n sobre los tipos y sfnonimos. Distribuci6n con citas de casi todos los especimenes de herbario de las Areciiceas de Sonora conocidos por noso[os es propocionada. Muchos de las poblaciones de palmas en este regi6n ilrida y semi-ririda son amenzadas por el desarollo y en algunos casos por la sobre-explotaci6n. Hay una necesidad urgente para el manejo formal y acci6n de conservaci6n para poblaciones locales; sin embargo en general ninguna de las especies en Sonora se considera espe- cialmente rara o en peligro inmediato. Key words: Arecaceae, Brahea, Erythea, Mexico, palms, Sabal, Sonora, Washing,tonia.

INTRODUCTION bers of their . Hence, they do not merit the space necessary to describe them." We disagree and devote Writing from in the late 18th century, more this space than a decade after suffering through the Jesuit expul- to describing the magnificent palms of the sion, Ignaz Pfefferkorn (1949:74) stated; "Many palm state of Sonora, in northwestern Mexico. grow in the region near the sea, especially in the The palms of northwestern Mexico have been poor- vicinity of Guaymas. Since these palms neither yield ly known due, at least partly, to their inaccessibility. nor attain the height which is reached by genuine Many Sonoran palm groves remain sheltered in secret palms, they seem to be either degenerate or false mem- canyons and hidden on remote mountains. In addition, there has been a paucity of field studies, field notes I Current address: Depa-rtment of Anthropology, Arizona State accompanying specimens, and "complete" herbarium University, Tempe, Arizona 82587 -2402. collections (few have both and ). Felger and Jolal

(TDF), tropical thornscrub (TTS)' Table 1. Ecological drstribution of palms in Sonora: Sonoran (SD), tropical forest woodland (OAK), and pine-oak woodland or forest (PIN).

OAK

OAK (Brandegee) H.E. Moore TDF Brahea elegans (Franceschi ex Becc.) OAK H.E. Moore SD TTS Brahea dulcis Kunth TDF OAK Brahea nitida Andr6 SD TDF TTS dac4,lifera L. SD OAK Trel. SD TDF H. Wendl. SD

canyons at the The purpose of this work is to provide information border, Brahea nitida occurs in riparian region of the on the natural history, ethnobotany, , dis- uppermost edge of the Arizona Upland Four tributions, and location of principal collections of Son- at its ecotone with oak woodland. tropical oran palms, and to help clarify their taxonomy and species occur in tropical deciduous forest and of nonde- nomenclature. Palms in northwestern Mexico are key- thornscrub (Felger and Joyal 1998). Palms the low- stone species, and we trust that this information will sert in Sonora commonly extend into with aid in efforts to conserve them and their special hab- er oak zones and sometimes even intermingle Al- itats. pines in pine-oak woodland of eastern Sonora. though some Sonoran populations are relatively wide- DISTRIBUTION, DIVERSITY, AND ECOLOGY spread, most are highly localized and niche-specific' Most of the inland groves contain only a single palm Three genera and six species of palms are native to species, although in some places two or three species Brahea with four species, and Sabal and Sonora: may occur together or in close proximity to each other. Washingtonia each with one species (Table 1; Fig- 1). Where these palm species occur sympatrically, the These are among the northernmost native palm genera maximum densities for each species are in somewhat in the Western Hemisphere. All are medium- to large- different niches. Three species, Brohea elegans' Sabal sized fan palms in the tribe of the subfam- uresanct, and Washingtonia robusta, occur together in ily . In addition, the (Phoenix a number of the palm oases near the coast north of dactylfera) is occasionally encountered in nonculti- Guaymas in the Sierra El Aguaje and vicinity, e.g', vated habitats. Cafl6n del Nacapule (Felger 1999)' Los Anegados, La Washingtonla occurs naturally northward to Arizona Huerta (Ojo de Agua), and Cafr6n Las Barajitas. Wash- and , whereas Brahea and Washingtonia oc- ingtonia robusta is the most narrowly restricted, with cur in both states of Baja California (Wiggins 1964; the fewest number of , and concentrated closest Turner et al. 1995). Washingtonia robusta, restricted to water sources. Sabal uresana occupies somewhat to a few riparian canyons at the southern edge of the less restricted habitats and is more numerous, while B' Sonoran Desert, appears to be relictual in Sonora' It elegans is the most u'idespread of the three and is truly occurs naturally otherwise only in Baja California Sur' abundant. Elseghere in Sonora, Sabal sometimesoc- Sabal uresana scar.celyextends beyond the confines of curs tosether $ith B. ttculeata or B. elegans. Sonora, whereas Brahea elegans is endemic to Sonora' Apparent adaptations to aridity and/or winter freez- Brahea aculeata, B. dulcis and B. nitida tange south- ing among palm: of :outhuestern North America in- ward in Mexico and B. nitida extends to ' clutle l;rrger rthickerr trunks and larger with Palm species represent less than 27o of the flora rougher i.rndmore reflective. duller-colored blades' of Sonora, which includes more than 275 species in a The ts'o trull Sonoran Desert species' Brahea armato vascular flora of perhaps 5000 species (Felger and S. r*'ars

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Fig. 1. Distributions of the native palms in Sonora, Mexico. The map of Brahea elegans shows the three major geographic populations. (l) Western Sonora, (2) north-central Sonora, and (3) east-central Sonora. Felger and Joyal ALISO tions of Brahea elegans are larger and more robust palma del taco "fruiting palm" in the case of Sabal, than those farther south, and desert-inhabiting popu- are mostly harvested for thatch in the spring dry sea- lations along the coast tend to have more glaucous son. leaves than those in the mountains of eastern Sonora. In the desert region, especially near lJres, the trunks of Sabal have been used to build corrals and as cross- (vigas) the trunks PALMS AND PEOPLE IN SONORA beams for houses. Traditionally only of dead trees were used to make corrals, but in recent Palms are one of the most highly utilized fam- living trees have been cut, contrary to the policy ilies worldwide, especially in tropical regions. Sonora, of certain local forestry offices. Farther east and south- at the edge of the , is no exception. The most ward in the mountains, palm logs are occasionally u'idely used part of Sonoran palms are the leaves used in construction and as benches, and are also hol- u hich are or have been employed in , thatch- lowed out to make planters. During the 19th century ing. and broom-making. The trunks have been used the trunks of Washingtonia robusta were used in for construction, and the fruit and sometimes the palm Guaymas "as rafters for houses. It is said to be a du- heart for food (e.g., Rose 1899; Standley I92O;Gentry rable ." (Rose 1899: 255). l9Jl. 1963; Pfefferkorn 1949; Hinton 1959, 1969;Di The blackish, sugary mesocarp of the ripe fruit of Peso :rnd Matson 1965; Sobarzo 1966: Nentvig 1977; Sabal, locally called taco, is eaten fresh in northeastern Pennington 1980; Felger and Moser 1985; Joyal and east-central Sonora but is considered unpalatable 1996d). Although much traditional knowledge of in the desert area near Ures. The fruits of Washing- palms in northwestern Mexico has been lost, consid- tonia are sweet and edible, and apparently were used erablc knowledge remains unrecorded as well as cul- for food by the Seri (Felger and Moser 1985). In north- turalll robust (e.g., Joyal 1996a, b). eastern Baja California, the Cocopa people pit-roasted kful uresana, the most widespread Sonoran palm, the large of (Felger and Amadeo is an important nontimber resource, and the most eco- Rea, unpubl. 1980). Some Brahea species, especially nomically valuable of the six regional palm species. B. edulis H. Wendl. ex. S. Watson from Guadalupe Al$ough other palm species and a few palmlike Island off the Pacific coast of Baja California, have a morxrcds are mentioned in the literature, Sabal is the sweet, datelike mesocaq). The fruits of certain Brahea most o(xrunonly and consistently cited Sonoran palm were used for food in Sonora, but the information is in mse than three centuries of historical references fragmentary. For example, the fruits of B. aculeata are tJolal 1996a). eaten by the Guarijio (Felger and David Yetman, un- The rr*'ly emerging, unexpanded palm leaf is publ. 1997), and those of B. elegans are eaten in the called a cojo,r-o(this term is also used as a vulgar term Ures region (Joyal 1996a). However, especially in the for penis). Cojoyos, primarily of Sabal, are highly val- case of the coastal palms north of Guaymas, the lack ued for *'eaving twilled and coiled baskets, mats, and of ethnobotanical information is meaningless since in- other items. and people travel considerable distances digenous people have not lived in the area for nearly to harvest them. Sabal is the only Sonoran palm em- one and one-half centuries and the information was not plol ed in hat-making. The fully expanded leaves recorded (Felger and Moser 1985). called pencas, especially those of Sabal, are widely In the 17th century Nentvig (1672) reported the use used for broom-making. The leaves of Sabal artd Bra- of palm hearts (terminal buds) for food in northeastern hea aculeata are extensively used for . Bra- Sonora. Gentry (1942, 1963) likewise reported that the hea aculeata is also used to make mats by the Guarijio Guarijio ate palm heafi (coraz6n), apparently of Sabal as well as other local people. Strips of leaf blades, and perhaps Brahea aculeata, a practice that kills the especially of Brahea aculeata and to a lesser degree, plant. Palm hearts are apparently no longer harvested those of B. nitida, are woven into a coarse cordage or in Sonora. rope and used for such purposes as tethering animals The fibers that hang between the leaflets of Sabal (Felger and David Yetman, unpublished field notes for uresana were used by the Mountain Pima to weave Guarijio ethnobotany; Gentry 1942; Joyal 1996a). saddle blankets. Orioles. including hooded and Palm leaves, primarily Sabal, Brahea aculeata, and streaked-back. make nests from these fibers. These B. nitida, are traditionally harvested at the time of the nests are sometimes used as sponges. Sabal petioles full moon. The quality and durability is said to be are employed to malie racks for aging cheeses,stirring compromised if the leaves are harvested at other times. sticks, and broom handles. However, harvesting does occur at other times, espe- Writing from Pcipulo in 1692. Adamo Gilg men- cially if for sale without regard for quality. Juvenile tioned that the Seri r*'ove. ". . . mats of palms and 'palm palms, called palma del suelo of-the ground,' reeds" (Di Peso and Matson 1965: 55). P6pulo was a are harvested for cojoyos for weaving mostly during Jesuit mission for the S€ri during the lTth and 18th the monsoon season.Leaves from mature palms, called centuries along thc Rfo Sonora near Ures; the palm VOLUME 18. NUMBER 1 Palms of Sonora. Mexico would have been Sabal uresana, which is common in ing spring drought if no other forage is available. Bra- the region. More than a century after the Seri no longer hea dulcis and B. nitida, at least in Sonora, do not lived in places where palms occur (the coast north of seem to be at risk, largely because they grow on in- Bahfa San Carlos, northwest of Guaymas), the people accessible cliffs and in remote canyons. Individual still had knowledge of palms and their uses from that groves or stands of B. elegans are vulnerable but region (Felger and Moser 1985). Some hint of earlier countless thousands of them occur at higher elevations palm use is found in the modern terms for any wooden on remote, uninhabited mountains and on steep cliffs. box, hehe zamij "palm wood," and the personal car- The palm canyons in the San Carlos-Bahia San Pe- rying box, hehe zamij an iqui ifuicalca "box to put dro region of central-coastal Sonora deserve vigorous belongings in," which have long been made from oth- management and protection. The more accessible, er kinds of wood. Zamij is the generic term for any magnificent palm oases in the vicinity of Bahia San palm including the introduced date palm. Seri oral his- Carlos have been decimated by development, clearing tory of the mid- to late 20th century included two of undergrowth, and removal of plants for landscaping kinds of native palms: zamij cmaam "female palm," (see S. uresana below). The Sabal grove, or Palmar, which most likely refers to Brahea elegans, and Tamij near the present Hotel Club Med, supported perhaps ctam "male palm," which most likely refers to S. ure- as many as 2000 adult palms in the 1960s and early sana and/or Washingtonia robusta. The fruit of zamij 1970s. Today about 500 adult palms remain, but un- ctam was eaten. A hat to provide protection from the dergrowth vegetation has been destroyed and there is sun was woven from tender, inner palm leaves. This virtually no seedling survival. Unless there is a change kind of hat was called zamij yeen haonam "palm-face to conservation management, these palm-dominated hat." "Face" refers to the inner, or youngest leaf or communities are essentially "living dead" sensu Jan- cojoyo. Date palms, Phoenix dactylifura, brought to zen (1988). Other extensive palm groves, such as those Sonora by the Jesuits during Spanish colonial times, in the canyons north of Bahia San Carlos, on the old have long been known to the Seri people. According road to Bahia Algodones, and at the San Carlos golf to oral history, one Seri man planted dates at one or course, have been annihilated during the past few de- more camps south of Bahia Kino in the mid-l9th cen- cades. No vestige remains of their former existence tury. apart from an occasional lone palm. The magnificent palm oasis at Cafl6n del Nacapule. CONSERVATION AND MANAGEMENT. supporting all three genera, is being devastated br of ornamental rock, cattle grazing, and van- Three of the Sonoran palms are listed by the Mex- dalism (Felger 1999). However, many intact palm oa- ican government on its most recent list of species ses remain in this region (Sierra El Aguaje, from the needing protection (SEDESOL 1994). SEDESOL lists vicinity of Bahfa San Carlos to Bahia San Pedro; see Brahea aculeata as "threatened. endemic." B. nitida Johnston 1924; Schnabel 1962, 1964; Felger 1966). as "endangered," and Sabal uresana as "rare, endem- The most extensive palm groves in the region are at ic." SEDESOL did not list B. dulcis, B. elegans, or Cafr6n Las Barajitas (between San Carlos and Bahia Washingtonia robusta. The SEDESOL determinations San Pedro), where all three genera are corunon. apply to a species' status throughout Mexico. Overharvesting has occurred at least since the 19th Based on our field experiences in Sonora, we rec- century (e.g., Gentry 1942). Rose (1899: 255) reported ommend the following IUCN 1997 Red List categories that "At Guaymas a few trees remain of tl:re rare Neo- (Walter and Gillett 1998): Brahea elegans and Sabal washingtonia sonorae t: W. robusta], but most of uresana are endemic or nearly endemic in Sonora and them have been cut out." Practices which ameliorate most closely fit the IUCN criteria for Vulnerable (VU) nonsustainable use of palms in Sonora include limiting status. The remaining four species largely fit the IUCN access to populations on privately controlled property, criteria for Lower Risk (LR), Near Threatened (NT) "sparing" (not cutting living trees in agricultural fields status in Sonora and Data Deficient (DD) in Mexico. and pasture), and restricting harvest according to lunar Both B. aculeata and S. uresana are still abundant cycles. Choice of palm size and leaf age, and control- in many places. Many of their populations are declin- ling harvest times (e.g., season as well as lunar cycle) ing or extirpated due to overexploitation and further regulate leaf harvest (Joyal 1995, I996b). loss, primarily due to clearing of tropical deciduous Throughout Sonora the dry dead leaves, or "skirts," forest habitat for buffelgrass (Pennisetum ciliare lL.l of palms are often purposely burned because they har- Link) pasture (Bfrquez and Martinez-Yrtzar 1997).In bor nests of aggressive paper wasps (Mischocyttarus addition the oases and riparian canyons where palms and Polistes). Palms generally recover from burning, occur are often heavily exploited for cattle grazing, which may be beneficial in destroying insect pests, settlements, and recreation. Cattle eat seedlings of S. opening up oasis vegetation, and increasing palm uresana and probably other species too, especially dur- germination (Hicks 1989). However, many thriving Felger and Joyal palm populations in Sonora occur in remote areas that native only in the Sierra El Aguaje and formerly at Washingtonia are not burned by people, and in the case of Sabal Guaymas uresana statewide onlv about IOTo of trunks show BReHee Mart. ex Endl.. Genera Plantarum 252. 1837. signs of fire. Northern Mexico, including both states of Baja Cal- THE SONORAN PALMS ifornia, to ; about 12 species, all oc- curring in Mexico (Quero 1992). Moore (1.973) and The following treatment of the Sonoran palm spe- Uhl and Dransfield (1987) consider Brahea and Ery- study of herbar- cies is based on our field experience, thea to be congeneric, although Quero (1989, L992) ium specimens, and the literature. We have studied treats them as distinct genera. These taxa are in need palm specimens and photos, especially those of north- of taxonomic study (Uhl and Dransfield t987). western Mexico, from the following herbaria: ARIZ, Brahea sensu stricto includes five species ranging ASU, BCMEX, BH, CAS/DS, DES, FI, GH, HBG, from Central America to northern Mexico excluding MEXU, MO, P, RSA, SD, UC, UCR, and US. These the Baja California peninsula. These palms are report- collections include nearly all known Sonoran palm ed as being restricted to limestone substrates (Bailey Unless otherwise noted, we have seen all specimens. 1943; Quero 1992), although this is not the case in specimens cited. The flrst set of Felger's specimens is Sonora for B. dulcis and some populations of B. nitida. ar ARIZ and the second at MEXU, with additional The two species in Sonora are distinctive and not duplicates when available at BH, FTG, CAS, etc. The closely related. first set of Joyal's specimens is at ARIZ and the second Erythea S. Watson includes seven species (Quero at MEXU, with additional duplicates at BH, US, and 1992). Five of them are confined to northwestern Mex- FTG. Specimen localities are listed by the present-day ico, one occurs farther south in western Mexico, and municipio (mpio.; see Steinmann and Felger 11.9971for another is in southern Mexico and Central America. a map of Sonoran municipios). They grow on soils derived from igneous rocks. In most instances heights of palms are visual esti- Erythea in Sonora is represented by a highly vari- mates. Seed measurement for Brahea and Washing- able and taxonomically ill-defined complex of often tonia include the thin and essentially inseparable en- isolated populations. Plants at the geographic and eco- docarp. The thin endocarp of Sabal uresana does sep- logical extremes are quite distinctive but populations arate and the seeds were thus measured without the showing intermediate features are commonplace. A endocarp. The following keys and brief descriptions comprehensive study of Erythea is sorely needed but are based on palms from Sonora and might not apply is outside the scope of this paper. We choose to treat to plants in other regions. the Sonoran taxa as two species within Brahea, while acknowledging that further research may result in a KEY TO THE GENERA different uurangement and/or the circumscribing of new taxa. l. Plants often branching from the base (suckering); leaves pin- Erythea was "one of the daughters of the Hesperi- nate; dioecious; seeds more than twice as long as wide, con- des, daughters of tlre Evening or the West who, in grooved; date palm, cultivated, rarely surviving spicuously Greek mythology, dwelt on an island on the western at abandoned settlements or marginally naturalized . . Phoenix guarded golden 1' Plants single-trunked; leaves palmate (fan palms); flowers edge of the world and who the apples bisexual; seeds less than twice as long as wide, not grooved; there" (Bailey 1937: t5). Bailey suggested that Wat- native palms. son's choice of thir pocdc name indicated that Watson 2. Petioles always entire, the leaves prominently costapai- meant ta Eryda cfrt (fl. Wendl. ex S. Watson) S. mate ( extending well into blade on the lower sur- Watsoo, rrrdcl b -Lr- b Guadalupe Island (about face), the hastula narrowly triangular, more than twice as long as wide, firm, and glabrous, the margins entire, per- 5m km Gr d 3 tb sparsely inhabited Pacific sistent; shorter than, or sometimes as long Cmd !o s€rve as the type species as the leaves; seeds broader than long, depressed globose fcfu B enuta, which he also trans- @d at that time. Bai- the leaves not costapalmate or 2' Petioles entire or armed, rc far as to designate E. ed- only moderately so, the hastula broadly triangular to ir- regular, broader than wide to less than twice as long s wide, the margins papery to variously pubescent- nc cr- tire, ultimately deciduous; inflorescences often lql THE SPECIES than the leaves; seeds rounded to longer than brod 3. Leaf blades often green above. silvery-blc nall teeth near the base; fruits beneath or tough and dull green or gxay-glEcr l}olva sensu stricto): surfaces: sepals separate.entiret widesprcd . c feltlike patch; flowering branch- 3' Leafblades shiny green on both surfg bases, densely pubescent, and at ed below. the lobes with ragged m-Ft rf,rmeter ... B.dulcis VOLUME 18. NUMBER 1 Palms of Sonora. Mexico

single and sometimes also double marginal teeth 1-6 mm long, often fewer and reduced apically, or some- times young plants and smaller-leaved adult plants with the teeth greatly reduced or essentially absent; hastula small, hyaline; blades often 55-90 cm long, 60-110 cm wide, glaucous bluish-green, especially be- low and slightly greener above, often discoloring with darker blotches upon drying. Inflorescences about equal to, or longer than the leaves, the branches and rachillae sparsely to densely tomentose. Rachillae (3-)5-11 cm long. Flowers white, very fragrant (at least in the late afternoon), the buds 1, 2, or 3 together, usually only 1 developing into a . Sepals I mm long, broadly triangular, green- ish white. Petals 1 mm long. Petals, staminal ring, and anthers white. Ovary green, 3-lobed. Fresh fruits 25- 30 mm maximum diameter, nearly round, the surfaces smooth, yellow-brown and somewhat mottled. Seeds 15.5-20.0 mm maximum diameter, rounded or nearly so. 2n : 36 (Read 1964). Flowering recorded March to May, the fruits often ripe in March of the following .

Distribution -Valleys, hills, mountains, and canyon bottoms and slopes, the distribution decidedly patchy but the plants often locally abundant. Tropical decid- uous forest and lower oak woodland, and sometimes extending into higher elevation zones in oak woodland and lower pine-oak woodland; Rio Mayo and Rio Fuerte drainages in the Municipio de Alamos, south- Frg. l. Bntlrcu utuleattt and B. elegans at Arizona-Sonora De- eastern Sonora, <'n \tus!'um. Tucson. Grown from seed collected in 1970, and ca. 320-1500 m. Populations slightly plantcd :inrultaneously in 1912 from one-gallon containers. Brahea farther north along the Rio Mayo, in the vicinity of .;.u|.'.nd. sniallcr palms on left, fiom the vicinity of Sabinito Sur, Los Bajios, in the Municipio de Quiriego may be B. cJ.t ()i .\lamos. and B. elegart.s,on right, from Cafl6n del Nacapule. aculeata. Likewise a sterile specimen from the vicinity Photob1 R. S. F. April 1998. of La Quema, in the Rio Yaqui drainage, in the Mun- icipio de Y6cora, appears to be B. aculeata. Also I Hastula glabrous; flowering branchlets not floriferous to southwestern , Sinaloa, and Durango. their bases, sparsely or sometimes moderately pubescent, lcss than I mm in diameter . . . . B. nitida Notes and horticulture.-This palm is readily grown I Petioles armed with small to large teeth; fruits globose, l6- from seed and is sometimes cultivated in southern Ar- .10 mm in diameter (.Erythea): izona. Plants more than 25 years old at the Arizona- ,1. Fresh fiuits 25-30 mm maximum diameter . . . B. a.^uleata ' '::'n *::'*l\ Sonora Desert Museum west of Tucson have shown ":'::1'-i'll' ll'-'" .o:"\i:'",""",,no signs of freeze damage. The precise northern limit for B. aculeata and its relationship to the southernmost Bn.lHER ACULEATA(Brandegee) H. E. Moore, Princi- B. elegans remains uffesolved. Both in cultivation and pes 24:91. 1980. Erythea oculeataBrandegee, Zoe in their natural habitats B. aculeata is a much smaller 5: 196. 1905.-TYPE: Mexico, Sinaloa, Cofradia, palm but with larger fruits and usually larger seeds. vicinity of Culiac6n,26 Ocr 19O4, Brandegee s.n. Grown side by side, these palms demonstrate clear dif- (holotype UC 1249991). Fig.2. ferences in size and growth rates (Fig. 2). Vernacular names.-Palmilla. Tajcil (Guarijfo). Specimens eramined.-Mpto. ALelraos: Arroyo Gochico, ca. 9 km E Ta'aco (Mayo). of San Ber-nardo, 860 m, Fishbein 2165 et al. (ARIZ).2.1 mi by road SW of Sabino (NE of Alamos), Felger 5023 (ARIZ). 15.4-15 Description -Tiunks mostly 2-9 m Iall, 12.7-25 cm mi E of Alamos, first drainage E of El Palmarito, 480-500 m, in diameter. Leaves often 10-30 in number, often 95- 27"02-03'N, 108"47'W, Joyal et al. 1426, 1427, 1428 (ARIZ). Los Lianos, NE of Sabinito Sur, ENE of Alamos, Joyal 2401 (ARIZ). 180 cm long; petioles commonly about as long as the E1 Palmarito Canyon, between Sabinito Sur and Santa B6rbara, leaf blade, 39-It6 cm long, 0.7-1.9 cm wide just be- 27'04'N, 108'45"N, 58O m, Jenkins 93-89 (ARIA. 14.6 mi E of low the blade, armed with mostly stout, often yellow, Alamos. E of El Palmarito. 27'02-03'N. 108'47'W. 480-500 m. Felger and Joyal

Vernacular nqas.-pqlrna ceniza "ashy palm," used at Los Bajfos by Guarijfo men "for a palm that grows to the east up in the mountains," apparently this spe- cies.

Description -Solitary palms, often to 5* m tall, the often lo-f cm in diameter, the older leaves fall- ing away. Leaves reaching ca. I25 cm long, the peti- oles 52-80* cm long, usually entire apically and with some broad teeth basally often 1-1.5 mm long, glau- cous, 14-25 mm wide near base, tapering to 8-15 mm just below the blade; hastula as wider or wider than long, to ca. 1.5 cm long, thin, brown, scarious and fraying into fibers; blades often 65-85 cm long, about as wide as long, shiny green above, glaucous-blue be- low, with deciduous, sordid, sparse tomentum on the segment midribs near the base of the blade on both surfaces, this tomentum and the hastula usually wear- ing away with age. Inflorescences arching, usually longer (sometimes twice as long) than the leaves, 4-times branched: flow- ering branches and rachillae crowded, purple-brown, often becoming cream-colored and very densely wool- ly-tomentose; branching angles acute. Rachillae cat- kinlike, floriferous to their bases, 4-10 cm long,2.2- 3.0 mm in diameter. Flowers greenish white, spirally arranged, not paired or clustered (more or less equally spaced from one another), the buds longer than wide, the flower bases submerged in the woolly hairs of the rachillae. Sepals ca. I mm long, overlapping, short woolly (tomentose-canescent)on exposed surfaces,the Fig. 3. Brahea dulcis Rancho Santa Brirbara. photo bv E. J.. margins scarious. Petals 2.3-2.4 mm long, glabrous May 1990. except sometimes sparsely canescent with some small hairs basally, striate (longitudinal lines formed by Joyal et al. 1425 (ARIZ). 1 km S of Rancho Santa Bdrbara, Jenkins prominent veins). Filaments slender from broadly ex- 93-41 (AF.IZ). Ca. 0.5 km N of Las plomosas, 7 km by road east- panded, thick, deltoid bases. Young fruits densely pu- ward from Sabanitas, vicinity 26.59,45,N, 10g"451/2,W,400 m, Fel_ ger 90-682 (ARIZ). Rancho San Pedro, 27"02.5,N, 108"43,W 450 bescent with short hairs. Fruits with a fleshy pericarp, m, 8 Nov 1988, Martin s.n. (ARIZ). 11 mi E of Alamos, 1960_61, smooth, purplish black when ripe, rounded, 10.5-12.5 Weber s.n. (BH). 16.1 mi SE of Alamos along road to Milpillas, mm in diameter, glabrous or sparsely pubescent at Boutin & Kimnach 3617, 3618 (BH).21.7 mi by road NE of Alamos base, wrinkling on drying, 8.5-11.5 mm in diameter. on road to Milpillas, Felger 5568 (ARIZ). 40 mi E of Alamos, Feb Seeds brown. 6-5-9.0 mm long, broadly ovoid. 1961, Nelson & Smith s.n. (BH). Upper crossing of the Rio Cuchu- Flow- jaqui, 7 mi ESE of Alamos, 26"59'N, 108'50,W 1300 ft. ca. 1 mi ering as early as March, but mostly June and July. upstream from road crossing, Sanders lO93A (UCR). Rio Cuchuja- Mature furits found in January and February. qui, upper crossing near Lajitas, 8.9 mi ESE of Alamos, 26"59,N. 108'49'W 320 m, Starr 470 (AF.IZ). Rancho Agua Nueva, 1450 ft. Distribuian--ln Sonora known for certain from Si- Boutin & Kimnach 3711, 3712, 3714, 3715 (BH). Siena de la Ven- erra dc Alre" and two remote areas farther east tana, 3000-4000 ft, oak-nolina-grassland, Gentry 2il79 (.ASlJl. along tb Rb Grchujaqui drainage in the southeastern Guirocoba: Valley slopes and lateral ru.royos with patched woodland parr dtb -r- tl 7m-1400 m. often growing out of formation, Gentry 4919 (ARIZ, DS, GH); Gentry 5959 (ARIZ. BH. roct crwia m pnecipitous cliffs DS, GH); Rancho Giricolii : Guirocoba], and canyon walls, [probably 30 mi S of irt Alamos, 1938, Lindsay s.a. (BH). 7.0 mi by road N of Guirocoba t dfring cliffs, cliff tops, moist slopes, upper edge of short-tree forest, 13 Apr 1989, Salmon s.n. (ARIZ)- dt-bcryons; at the upper elevational limits of lqlf Ad*'ous forest, and oak woodland, or the IAB dflne-oak woodland. The substrates do BnaneaDULCTs (Kunth) Mart., Hist. Nat. palm.3:244. b tc limestone, unlike those of populations 1838. dulcis Kunth, Nov. Gen. Sp. I [qur- b Mcxico. The SonoranB. dulcis is not in tol: 300. 1815.-TYPE: Amerique EquarorialB Bonpland 3938 (holotype P!). Fig 3. d nrtheastern Mexico to Central Ameri- VOLUME 18. NUMBER I Palms of Sonora, Mexico

ca. Probably also in southwestern Chihuahua but not documented from that state. Brahea dulcis is quite var- iable across its entire range and there may be more than one taxon involved.

Specimens examined.-Mpro. ALerraos: Sierra de Alamos: S-facing cliff above Cafl6n Saucito, 1300 m, Fishbein 1610 et al. (ARIZ); Cafl6n de Bolsa (Caff6n las Piedras), 1450 m, Fishbein 1922 et al. (ARZ); La Huerta (abandoned) on "magnolia trail," 1100 m, Jen- kins 88-297 (ARIZ); Above el puente de La Huerta (a saddle above [-a Huerta) on the trail to El Pinal Grande and Aduana Peak, near 26'59'N, 108'58.6'W 1300-1450 m, Sanders 13490 et al. (AF.IZ, UCR)I N side of S. de Alamos, 15 Jun 1996, lttrson s.n. (ARIZ). Vicinity of Puerta de Mano [ca. 1 km S from Choquincahui, N of Guirocobal, 700 m, 26'59'N, 108'39'W 26 Dec 1988, Salmon s.n. (ARIZ). Rancho Santa Biirbara, el valle San Felipe, Rio Cuchujaqui l*ge, ca. 25 km by air ENE Alamos: sffeam bottom below oak- * forest, 27"05.5'N, 108'42.5'W 750-800 m, Joyal et al. 1443 (ARIZ); oak-pine forest, 27"06.8'N, 108"42.2'W ca. l4O0 m, Joyal t192. 1494 (ARIZ).

Bnurre ELEcANS(Franceschi ex Becc.) H. E. Moore, hiley 19: 168. 1975. Erythea elegans Franceschi cf, Bccc., Webbia 2: 138. 1907.-TYPE: U.S.A.. C-alifornia "leaf from the original plant in Los An- geles (this old palm has always been closely shaded by overhanging tree), from original in (the mature fruit fell from this spadix when I cut it)," 13 Sep 1913,J. HarrisonWright s.n. (ho- lotype, FI, not seen; [the specimen is accompanied by correspondencefrom Wright to Beccaril; photos ARIZ!, BH!, MEXU!). Also see additional notes Fig. 4. Brahea elegans. West-central Sonora, Cafl6n Las Bara- and specimens cited below. Fig.4, 5. jitas. The tallest palm is Sabal uresana, the others are B. elegans. Photo by R. S. R, 1995. Erahea clara (L. H. Bailey) Espejo & L6pez-Fenari, Sida l5: 617. 1993. Erythea clara L. H. Bailey, Gentes Herb. 6: I97, Fig. 69. 1943.-TYPE: Mexico, Sonora, Nacapule Canyon 19.0 mm maximum diameter, nearly rounded or with near Guaymas, 14 April 1936, Bailey 263 (holotype, BH!). the maximum diameter IO-2OVo larger than the mini- mum diameter. Mostly flowering in late spring, the Vernacular n6as5.-B,qbiso (north-central Sonora). fruits ripening the following spring. Coguegue (central Sonora). Palmilla (central Sonora). Mahajam stuttum (Pima Bajo; Rea 871). Zamij cmaam Distribution-This palm is endemic to Sonora, and (Seri; Felger and Moser 1985). shows considerable variation across its geographic range. Future studies may reveal geographic variation Description -Medium to large palms, the trunks often that could wrurant infraspecific recognition. 6-12(--ca.20) m tall, 16-33 cm in diameter. Petioles Populations in the northeastern part of the range in often 80-135 cm long, armed with teeth reaching 7.5- Sonora occur at the upper elevations of the Sonoran 8.5 mm long, these usually stout, curved or straight, Desert, while at their southeastern limits these palms solitary or double. Hastula to about 3 cm long, often occur in tropical thornscrub and the northernmost lim- hyaline, the sides unequal, irregularly niurowed or its of tropical deciduous forest. At the higher eleva- pointed. Leaves about 40 or fewer in numbeq the tions, throughout eastern Sonora, this palm extends blades often approx. 1 m long, strongly glaucous with into the lowermost oak zones. much variation in leaf color and indumentum. from We distinguish three populations which are geo- green to silvery glaucous. graphically definable (Fig. 1). There appear to be mor- Inflorescences 10-12 per tree, mostly longer than phological differences in trunk and leaf size, and color the leaves, 2O5-34O cm long. Bractlets 1.0-1.5 mm and indument of leaf blades. These are as follows: long. Flowers (1)2 or 3 in clusters. Sepals 1 mm long, thin, cupped, broad, and scarious. Corollas sympetal- Western Sonora.-Hesper palms occur in three ar- ous below, the lobes 1.6 mm long, thick and broadly eas in westell $sns1a-f1om the Sierra El Aguaje and deltoid. Filaments short and slender. Fruits rounded, vicinity northward to the Sierra Seri. These palms are (15.8-)18.0-19.5 mm maximum diameter. Seeds 14.5- generally large and robust, especially compared with Felger and Joyal ALISO

Fig. 5. Brahea elegans. North-central Sonora, Sierra de Nacozari, upper margin of Sonoran Desert and oak woodland vegetation with many riparian species; collection site for Felger et al. 3288. Photo by R. S. H, June 1960. those in east-central Sonora but seem to be somewhat These palms grow in riparian canyons such as La Pin- less robust than those of northeastern Sonora. The leaf tada at the western base of the range to peak elevation blades are generally tougher and much duller ("whit- at ca. 1160 m. No other palm species is known from er") than those of the east-central and northeastern So- this range. nora populations. The most isolated and undoubtedly the smallest Many thousands or perhaps millions of hesper palm population in Sonora occurs on a north-facing palms occur in and about the rhyolitic Sierra El Aguaje slope below Pico Johnson in the Sierra Seri. There are between Bahfa San Carlos in the south and the vicinity probably fewer than one dozen of these palms, hidden of Bahia San Pedro in the north. These palms range from view and far from any road. The colony includes across many habitats such as canyon bottoms, steep juvenile and adult plants. Known to us only from pho- rock clefts, and cliffs, including sea cliffs, and slopes, tographs taken in 1988, they appear indistinguishable sometimes extend onto upper beaches and beach from the Sierra El Aguaje hesper palms about 185 km dunes. Numerous palms grow out of crevices in oth- to the south. No palms occur farther north in the desert erwise solid rock and on north- as well as south-facing of western Sonora. slopes, and they are especially abundant in the higher During the 1970s a Seri fisherman brought Mary mountain mass (the summit is ca. 860 m). Among the Beck Moser an immature Brahea leaf which he larger groves, such as in Cafr6n del Nacapule and Cafr- claimed he obtained from a single plant on the west 6n Las Barajitas (Fig. 4), there is considerable varia- Other Seri people also knew of tion in leaf color, ranging from green-leaved plants to side of Isla Tibur6n. the much more common dull, more glaucous- or whit- this palm. Repeated attempts by Felger, even with ae- ish-leaved plants. In this region B. elegans often gtows rial searches, have failed to locate this plant. Narrow, alongside Sabal andlor Washingtonia rn riparian can- slotlike canyons on the west coast of this.large island yons and low-elevation palm groves. might shelter a palm, but the region is extremely arid An even larger population occurs in similar habitats and hardly s€ems suitable habitat for palms (see Felger in the Sierra Libre where they are likewise abundant and Moser 1985). However, B. armata does occur in at hieher elevations across the main mountain mass. semiriparian canyons on the not-too-distant Isla \.OLUME 18.NUMBER 1 Palms of Sonora. Mexico de la Guarda. Strangely, the Seri did not tell Moser or The relationships with the geographically neighbor- Felger about the isolated colony in the Sierra Seri. ing hesper palms, B. aculeata, B. armata, and B. bran- degeei remain unresolved. The hesper palm of western N o rt h - c ent ral So n o r a. -The se are the northernmost Sonora is distinguished from B. armata by its more hesper palms in Sonora, occurring within the Rio So- slender trunk, generally greener (not as "white" or nora drainage. They are scattered in localized popu- "silvery") leaves that are more variable in color, and lations in the Sierra Aconchi between Ures and Na- smaller inflorescences. However, some cultivated hes- cozari, and north through the Sierra to the valley bot- per palms at the Huntington Botanical Gardens in toms near Arizpe (Fig. 5). Some are planted in the southern California, grown from seed obtained in the town of Arizpe. They have relatively green leaves and vicinity of Bahfa San Carlos, are exceptionally robust are strikingly large and robust, often reaching 18 m in and approach B. armata in gross appearance. Hender- height. From a distance these palms might be mistaken son et al. (1995) show B. armata in northwestern So- for Washingtonia robusta. They appear to be - nora in a region where there are no native palms. Bra- logically continuous with the increasingly smaller hes- hea armata is endemic to Baja California and Isla An- per palms farther south in east-central Sonora. gel de la Guarda in the Gulf of California. East-central Sonora.-Numerous, mostly widely Brahea brandegeei was considered to be endemic scanered, and often small groups of these palms occur to the Cape Region in southern Baja California Sur in eastern Sonora from the vicinity of Onavas and Nuri prior to the field guide by Henderson et al. (1995). northward to the vicinity of Moctezuma. The region is Wendy Hodgson and Jon Rebman found a large pop- entirely in the Rfo Yaqui drainage. Some of these ulation of hesper palms in the Sierra San Francisco, in paltnr can be seen from Highway 16 (the Hermosillo- northeastern Baja California Sur (Hodgson & Rebman Y6cora road) in the vicinity of El Palmar de Onavas 8171, ASU, BCMEX, DES). Are the Sierra San Fran- and Agua Amarilla. The southernmost of these palms cisco palms conspecific with B. brandegeei s.s. and/or approach B. aculeata in appearance but seem to be the Sierra El Aguaje palms? Henderson et al. (1995) larger in overall size, and bear the smaller fruits char- list B. elegans as a synonym of B. brandegeeibttoffer rteristic of B. elegans. no explanation for their decision. The Sierra San Fran- cisco hesper palms grow among Sonoran Desert veg- Honiculture.-Plants grown from seeds, most often etation in deep semiriparian canyons. Unlike their Son- obtained in the Sierra El Aguaje region, are occasion- oran counterparts, they seem to be confined to canyon ally grown in southern Arizona and Sonora. This palm bottoms and there appears to be a larger percentage of is relatively slow-growing, and thrives under cultiva- very tall palms. Brahea brandegeei from the Cape Re- tion in the Sonoran Desert, tolerating freezing weather gion of Baja California Sur seems to differ from the in Tucson without damage. It is worthy of wide cul- Sierra San Francisco and western Sonora plants by tivation in very hot climates. having generally greener and thinner leaves and other ly'ores.-The source plant for the type of Erythea ele- characters cited by Bailey (1937) including inflores- gans Franceschi ex Becc. was a cultivated palm grown cences only as long as or shorter than the leaves. These from seed said to have been collected about 25 years characteristics, however, have not been quantified or earlier by a miner "in the vicinity of Hermosillo" studied in depth. (Beccari l9O7). The type specimen has petioles, spiny Specimens examined.-CALIFORNIA (cultivated): Buddington's glaucous leaves, and mature fruits 1.7-1.8 cm long. garden, Los Angeles, Wright & Jones .s.n. (BH, photos of source Wright's letter to Beccari pointed out that the source plant). Photographs among papers of M. E. Jones of "Franceschi's plant (the Buddington plant, Los Angeles, CA) and type plant, Buddington's garden, Los Angeles, California," later sent (BH). other plants grown from its seeds were different from to H. Moore by G. H. M. Lawrence, Hunt Botanical Library Mrs. A. Buddington [residence], Alpine St., Los Angeles, supposed the Baja Californian E. armata and E. brandegeei original tree of Franceschi, 25 Aug 1927, 1 May 1928, Braunton, (Purpus) H.E. Moore. .r.n. (BH! [specimen and photographs]). Riverside, J. Harrison Based on its cultivation in southern California in the Wright's grounds, 4 Mar 1927, Bailey 9255 (BH), and 3l May 1921 , late 19th and early 20th centuries, B. eleganq.rwas re- Bailey 9257 (BH).-SONORA: MpIo. BACANoRA:Rancho El Tor- ported to be dwarf in habit and slow-growing. In fact reoncito, W of Bacanora, 950-1000 m, Joyal 2223 (ARIZ)-MPrI. Cuvpas: 10 mi from Cumpas on rd to Bavidcora, Wiggins 7423 (DS, plants from the hot desert of Sonora often are stunted US).-Mrto. L.q.CoLon.e.o.l: E of La Palma, 50 mi N of Guaymas, when cultivated in the nondesert regions of California Sierra Libre, Wiggins 6481 (DS, US).-Mrro. Gunvrrals: Palmar, a (see Bailey 1937). canyon opening eastward from the Gulf of California, 24 mi north Hesper palms in western Sonora have also been of Guaymas near San Pedro Bay, Sabal uresana in same canyon, called B. roezlii Linden (e.g., Wiggins 1964),likewise attractive grove coming down to the beach and extending up the canyon. also on cliffs, Bailey 2 (BH). 14.1 km N of Bahia San poorly based on a documented cultivated specimen. Carlos, ca. 3.2 km from the ocean, steep canyon with spring, stream, Bailey (1943) showed B. roezlii to be a synonym of and palm oasis, 100 m, Boutin 2017, 2018 (BH, HBG). Cafl6n del B. armata of Baia California. Nacapule (Nacapule Canyon), ca. 4 km N of Bahia San Carlos, Felger and Joyal

28"00'56"N, 111"02'58'TV',150 m at entrance: 30 Mar 1934, Bailey a (BIJ); Felger 3j82, 11985 (AF.IZ); Parfin 3027, 3O30 (ASU); Starr 49O (ARIZ); Yatskievych 8248 (ARIZ). S slope in a large grove ofErytheas, 5.5 km NE up a canyon starting 22.1 km NW of San Carlos Bay, ca. 7 km from the ocean, Boutin 2688 (BH, HBG). San Pedro Bay: Moran 4037 (BIJ); Ravines in hills about bay, as- sociated with but much less common than a Sabal sp., Johnston 4340 (CAS, US). Bahia San Carlos (San Carlos Bay): Few scattered trees among a Sabal colony, broad canyon, Johnston 4349 (CAS); Inland side of the main highway, Zona et a/. 258 (RSA). La Huerta, Felger 3069 (ARIZ). Sierra Libre, 32 mi S of Hermosillo, 28"24'N. 110'58'W 3OO m, Starr 753 (ARIZ).-Mpro. Hpnrraosuo: Cafr6n La Pintada, 33 mi on Mex Hwy 15 south of Hermosillo, then 3.9 mi by road E of La Pintada, deep riparian canyon, Felger 3262 (ARIZ). Sierra Seri, N side Pico de Johnson, less than 500 ft from summit, 15 Ap 1988, Caugill (photo: ARIZ, BH, MEXU). Vicinity of Hermosillo, 7 Mar 1910, Rose et aI. 12519 (BH [undoubtedly cultivatedl).-Mpro. NAcozARr: Sierra de Nacozari, 3900 ft, 1l mi by road to Angostura and Oputo east of Esqueda-Nacozari road, then by foot ca. 1 mi S, riparian canyon, Felger et al. 3288 (ARIZ).- Mpro. RosARro Tssoprco: 19.4-19.6 mi (by road) N of river cross- ing at Nuri (Palm Valley), wooded slope, 1840 ft, Reeves et al. L18639 (ASU); Turner et al. 75-62a (ARIZ). 0.7 mi N of Las Cues- tas on road from Rosario to Nuri, 535 m, Zona 261 (RSA).-Mrro. Sar Pelno DE LA CUEVA: 13 mi NW of San Pedro de la Cueva, Wiggins 75I3 (DS, US).-Mpro. Unts: W flank of Sierra Aconchi, ca. 20 km NNE of Ures, Rancho El Estribo, 700-900 m, Joyal 2497 (ARIZ).-Mpro. YEcoRA: 20.7-23.4 mi NE of Nuri, 540 m, Boutin & Kimnach 3396-3699 (BH, HBc). Rancho Los Paredones, W of Tepoca, 28"28'N, 109'18'W 750-800 m, Joyal et al. 1670 (ARIZ). Ca. 20 km E of Onavas: Rancho El Palmar, Joyal 1630 (ARIZ); Punta La Pila [: Atuu Amarilla], drainage above "El Palmar," ca. 800 m, Joyal 1579, 1581, 1607 (ARIZ). NE of Rancho Palmar, arroyo E of Onavas, Rea 871 (ARIZ). Fig. 6. Brahea nitida. North-central Sonora, portion of upper part of El Baviso or Palm Canyon, Sierra Baviso, southeast of Mag- BReHse NITTDAAndr6, Revue Horticole 344. t887.- dalena. Photo by George Ferguson, 1985. TYPE: Illus., Andr6, Revue Horticole 344, Fig. 67, 68.69.70.1887. Figs.6, 7. branchlets (includes rachillae) with flne, whitish hairs. Rachillae floriferous nearly to their bases,20-56 mm long, 0.7-0.9 mm in diameter. Flowers sessile, sub- Brahea prominens L. H. Bailey, Gentes Herb. 6: I92, Fig. 92C, tended by minute, broadly deltoid bractlets 93D, 94D, 96-98. 1943.--:TYPE: Mexico, [Oaxaca], park, Oa- with acute xaca de hdrez, 16 Mar 1940, Bailey 533 (holotype BH!). tips. Calyces1.2-1.5 mm long, cuplike, the sepalsim- bricate, separate,broadly triangular,very thick towards Vernacular names.-Bablso (vicinity of N6cori Chico the base,puberulent with short, thick, white hairs. Pet- and Cucurpe). Jabehui (Rfo Satachic). Palma liza als 2.2-2.8 mm long, with sparse hairs below like "smooth palm" (name given at Los Bajfos "for a palm those of the calyx, valvate in bud, slightly cuppednear that grows to the east up in the mountains"). Metajcu the apex, faintly indented (grooved) to accommodate (Guarijio). the anthers,the filaments only united basally, broadly triangular with the upper portion Description -Graceful palms, the trunk to ca. 10 m slender and free. Young fruits pubescent;mature tall, usually much shorter, and 15.5-ca. 30 cm in di- fruits 11.0-13.0 mm long, 8.5-9.0 mm maximum ameter. Petioles entire or sometimes with a few minute width, broadly ellipsoid, smooth and yellow-brown to purplish teeth near the base, bluish glaucous, about as long as when fresh with a thin fleshy pericarp, wrinkled when the blades, I.6-4 cm wide the near base, tapering to dry, and often with short, sparsehairs near the gla- I.l-2 cm wide near the blade: hastula often 3-5 mm base, otherwise brous. Seeds8.3-11.0 mm long, long, glabrous or ciliate. Leaf blades of larger trees ca. 6.4-:7.2 mm maxi- mum width, ovoid-ellipsoid, 80-100 cm long, slightly wider than long, relatively light brown, smooth. Flowering primarily June and July; fruits flat, shiny green above, bluish glaucous below, the seg- mature late July of the same season,at least ments divided to about half way, with a very slender, among northern So- nora brown, curled fiber in the sinus of each leaf segment. Populations. Inflorescences arching, often reaching ca.2.0-2.3 m Distributiott-Eastern Sonora. 945-1645 m. Moun- long, 4-times branched; branches and tain regio$ often on limestone but sometimesappar- YOLUME 18. NUMBER 1 Palms of Sonora, Mexico

7. Brahea nitida. Northeastern Sonora, Rancho Pinalito, Sierra Torro Muerto. Photo by E. J., August 1993

elso on rhyolite; growing in crevices or among side of this limestone mountain, supports the largest

mac , 27"32'3O'1:t, 108"42'30'W 5400 ft, 18 May 1986, Mar_ in the late seventeenth century. At that time only seeds tin s.n. (ARIZ). Waterfall at Sahuarivo, 2j"2O,N, 108.39.8,W, ca. could be transported. Vegetatively propagated 1500 m, moist pine-oak woodland, barranca drains to San Bernadino varieties : [sic, Bernardo], 17 Mar 7992, Martin et al. s.n. (ARIZ). Ca. 5 mi were first successfully introduced into the southwest- W of Chinipas on rd from Los Tanques, pine forest, 5 May 1996, ern in 1890 (Nixon 1950) and presum- palm l,11ys6n s.n.-l"lpro. Cucunpe: Canyon, 23 km (15_17 mi) SE ably soon taken to Sonora. Date palms are widely cul- of Magdalena on road to Cucurpe, SW flank plata of Cerro Cinta de tivated, sometimes commercially, in the lowlands [Sierra Baviso], 1050+ m: Boutin & Kimnach 3744 (BIF-); Estrada throughout Sonora. The widely cultivated p. et al. 515 (BH); Felger 3275 (AF.IZ); 16 Jul 1977, Van Devender canarien- et al. s.n. (ARIZ).-Mnro. NAconr Cmco: Arroyo Chinalito, Rio sis Hort. ex Chabaud is distinguished by its massive, Sritachi, downstream from Ndcori Chico, 2930 ft, Fetger 3307 solitary trunk, and green leaves. (ARIZ). Rancho Pinalito, Sierra Torro Muerro, E of Buena Vista, 29'38'N, 108'55.5'W ca. 1000 m to 1300 m], Joyal [ranges 2361 Sanel- Adanson ex Guers., Bull. Sci. Soc. philom. par- (ARIZ).-Mno. Necozenr: [Sierra del Tigre]: Cafl6n del Temblor, is 87:205-206. oak-grassland, 23 Aug 1940, E. A. Phillips 735 (GH); Between Las 1804. Tieritas and Cafl5n de las Gallinas, oak-grassland,, 25 Alg 1940, Southeastern United States to northern Phillips 739 (GH). Caff6n del Temblor, oak-grassland, 19 Aug 194O, South Amer- Phillips 590 (AF.IZ, GH).-Mrro. unknown: Las pinitas, 12 Oct ica including most of Mexico except the Baja Califor- 189O, Hartman 140, Lumholtz expedition (GH). Huehuerachi, 7 Dec nia peninsula, most of the , and ; 1890, C.E. Lloyd 469, Lumholrz expedition (GH). 16 species (Quero 1989; Zona 1990). Qfissa,a1i6n.-lvIpro. Guesnses: Cafr6n Cruz de peffasco (E of Guasabas), rock walls of canyon and canyon bottom, Felger, 5 June Sasel URESANATrel., Ann. Rep. Mo. Bot. Gard. 12: 1960. 79, pl. 35-37. l9ol. Inodes uresane (Trel.) O. E Cook, Bull. Torr. Bot. Club 28: 534.1901.-TypE: PsoeNxL., Sp.Pl. 1188.1753. Mexico, Sonora, in the uplands, a few miles to the About 12 speciesnative to the Old World (Barrow N of Ures, 17 Aug I9O0, Trelease 379 (holotype 1998). MO!; photo,25 Oct 1934,BH!). Fig. 8.

Vernacular nqynss.-pqlma (tlrroagh much of Sono- PsoeNrx DACTvLTFERAL., Sp.pl. 1188. 1753.-TypE: ra). Palma grande (Rancho Santa Brirbara, Alamos re- Palma hortensis mas et foemina in Kaempfer, gion). Palma real (vicinities of: Alamos; Guajaray, Amoenitatum Exoticarum 668, 686, t. 1,2 (l7l}); north of Alamos; and Ndcori Chico). palma del suelo (see Taxon 28: 59-10. 1979). (uvenile plants, eastern Sonora near N6cori Chico to Vernacular nomes.-ddrjl (in Sonora also used for var- the vicinity of Alamos). Palma del taco, ot taco (adult ious species of with indehiscent, fleshy fruits); plants, throughout the range in Sonora). Indigenous date palm. languages: Sart, sab6, ta'cu (Gunijfo). Ta'aco (Mayo). Sajavil (Mountain Pima; vicinity of Y6cora to May- Description -Large palms branching (suckering) coba). Maahagam (Pima Bajo; Rea 1997). Zamij ctam from the base. Leaf bases persistent on the trunk. (Seri; Felger and Moser 1985). Leaves glaucous gray-green, several meters long. The genus is "easily distinguished from all other palms by Description.-Large palms, often ca. 5-15 m tall, the the induplicate fieaflets V-shaped in cross-section] pin- trunks often 26-36 cm in diameter, commonly fire- nate leaf with the lower leaflets modified as spines" scarred. Leaves large, reaching 4.4 m long, the petioles (Uhl and Dransfield 1987: 217). Dioecious; inflores- entire, (64-)80-260 cm long, to 9 cm wide near base cence branches orange, much shorter than the leaves. and 2.5-5 cm wide just below the blade, the hastula Male flowers with very short filaments. Pericarp sweet 5-18 cm long, triangular, more than twice as long as and edible, the seeds aa. 1.5-3 cm long, more than wide, persistent and firm with entire margins, the twice as long as wide, and conspicuously grooved. blades often ca. 90-180 cm long (from base ofhastula to apical tip), markedly costapalmate (the petiole Distribution-A few date palms are marginally nat- ex- tends well into the blade on the lower or abaxial sur- uralized, or perhaps only persistent, on Isla Lobos, a face), partly curved, glaucous bluish green, often coastal, sand-bar island of southwestern Sonora. Else- with coarse intersegmental fi bers. where date palms occasionally persist from abandoned Inflorescences including peduncles 96-235 cm long, plantings. In contrast, this palm is naturalized at wide- spreading to arching (not drooping), much branched, ly scattered riparian or semiriparian habitats in Baja shorter California Sur. than, to sometimes nearly as long as the leaves. Flowers s€ssile; sepals united below, the lobes entire, Horticulture.-The date palm is native to the Middle the petals 4.4-5.3 mm long; filaments rather fleshy, East, although truly wild populations are not known unibd below into a tube about as long as the sepals, with certainty to exist. It was apparently first brought 6c frec portion relatively long. Fruits 13-23 mm in to Sonora by the Jesuit missionary Padre Eusebio Kino dirmcler, 12-18 mm in height, globose, brown to \-OLUME 18. NUMBER 1 Palms of Sonora. Mexico t5

7, Fg 8. Sabal uresana. Northeastern Sonora, Municipio de N6cori Chico, Rancho Napopa. Mesquite-grassland-sonoran Desert vege- b' \ote the costapalmate, curved leaf blades of the immature palm in lower right. photo by E. J., August 1993.

; seeds10-16 mm in diameter.4-10 mm in George Lindsay but lacks a specimen. We have seen t. depressed-globose, dark red-brown ("chest- no specimens from Sinaloa, but a large palm exten- " ) to blackish, glossy. Flowering April to June, sively utilized for making hats in the vicinity of Choix y in May; fruits ripening in August in the desert, in northeastern Sinaloa appears to be this species (Bar- September and October in the mountains. 2n : ney Burns, pers. com., 1995). Furthermore, Hermilo (Eichhorn 1957). Quero (pers. comm., 1998) reports that he has docu- /ion.-Sonoran desertscrub,thornscrub, tropi- mented it from northern Sinaloa. There seems to deciduous forest, and lower elevations in oak be north-south and elevational clinal variation in overall plant dland. West-central and eastern Sonora, from near size with plants of the north- ern populations level-t22o m. Mostly on deep, gravelly to sandy larger than those to the south. There is also geographic silty soils. Scattered groves along or near the coast variation in fruit size, with the northernmost populations Bahia San Carlos northward to Bahia San pedro, bearing the largest fruits, while plants y along the bottom of riparian canyons including near Ures and along the coast bear the smallest fruits. e and Barajitas, low-lying swales or depres- and occasionally on coastal dunes and upper Horticulture.-Occasionally cultivated in Sonora and Farther inland on sandy plains, bajios (low- Arizona, S. uresana is recommended as a landscape areas), hills, floodplains, and canyon bottoms. tree for large areas. It is not damaged by freezing tem- low hills in eastern and central Sonora from Opo- peratures at least as far north as Tucson. The plants southward, especially near the desert-thornscrub seldom survive bare-root transplanting and are not Glone (Fig. 8). Seldom on rocky slopes. Sabal palms suited to container cultivation since they quickly de- often left standing in fields, orchards, pastures. and velop deep roots. This speciesalso occurs in southwesternChihuahua rtere it is probably not rare. We do not know of any Notes and ssrnrnsn\s.-B,ruchid beetle larvae in the qrccimens, however. The one record (Zona 1990) is a genus Caryobruchus (Bruchidae: Pachymerinae) feed barium sheet at BH with correspondencefrom on Sabal seeds, and adults feed on the floral nectar. t6 Felger and Joyal ALISO

Seed predation by bruchids can be as high as 5O-9O7o 800-900 m, Jo;*al 167 6ia171. l-os Bajios, ca. 10 km by air S of (Zona 1990; Joyal 1995). The fruits also are eaten by Guisamopa, 28'35'N. lOfO7'W ca. 800 m, Joyal et al. 2478 (ARIZ). Rancho El Palmar de Onavas, on Hwy 16, between Tonichi many animals including coatis, coyotes, foxes, rac- and Tepoca, 28"29-30'N, l@24'W, ca. 700 m, Joyal 2576 (ARIZ). coons, and ringtail cats. Hooded and streaked-backed Rancho Palmar, E of Rio Yaqui on Hwy 16, NE of Onavas, Rea orioles weave their nests from the leaf fibers, attaching 816 (SD). 20 mi (by road) NE Mazocahui, just past Rancho Rodeo, them to the undersides of leaf blades. Marin M72-23 (SD't. The plants and fruits of S. uresanc are among the largest of any species of the genus. Southward in Sin- WasruNcroNrAH. Wendl., Bot. Zeit. (Berlin) 37:. lxi, aloa this species is replaced by S. rosei (Cook) Becc., 68, 148. 1879 (conservedname). distinguished in part by its more slender trunks and Two closely related species (Bailey 1936). Wash- greener leaves. Sabal rosei in turn is replaced by S. ingtonia natiye to waterholesand riparian can- pumos (Kunth) Burret farther south in the Rio Balsas filifera, yons in northeastern Baja California (norte), Basin. These three sabals constitute the phylogenetic south- eastern California, and western Arizona, and natural- arm of the genus in western Mexico (Zona 1990). ized or adventive in southern Nevada (Turner et al. There is a gap of approximately 300 km between the 1995). It is distinguished in part from W. robusta by southernmost documented S. uresana and the north- its conspicuously thicker trunk (especially in cultiva- ernmost S. rosei. However, this intervening area is tion) and dull green leaves. poorly known botanically and much of it was con- verted to agriculture long ago.

Specirnens examined.-Mpro. ALelraos: I km downstream from Sa- We,ssrNcroNrARoBUsra H. Wendl., Gart.-Zeitung bino (NE of Alamos) along the Rio Cuchujaqui, in riverbed, not (Berlin) 2: 198. 1883. Neowashingtonia robusta common, 22 Jan 1961, Felger 5026 (ARIZ). Alcorn's ranch at San Heller, Cat. N. Amer. Pl. 3. 1898.-TYPE: Illus., Pedro, 27"03'N,108"42.2'W,8 Oct 1994, Jenkins 94-136 (ARIZ). Las Cabras, ESE of Alamos, 27'00'N, 108'55'W 340-360 m, Joyal Andr6, Revue Horticole. 404, Fig. 73. 1885. 1328, Joyal 1497 (ARIZ). Rancho Santa Biirbara, E of Alamos, Washingtonia sonorae S. Watson, Proc. Amer. Acad. Arts 24:79. 27'05.5'N, 108'42.5'W 750-800 m, Joyal et al. 1442 (ARIZ). Ran- 1890. I4l. filifera var. sonorae (S. Watson) M. E. Jones, Contr. cho San Pedro, 27'03'N, 108'42'N, 450 m, 9 Nov 1988, Martin s.n. W. Bot. 15:. 48. 1929. Neowashingtonia sonorae (S. Watson) (ARIZ). Near Sabinito Sur, 20 Dec 1989, Martin s.n. (ARIZ).- Rose, Contr. U.S. Natl. Herb. 5:255. 1899.-TYPE: Mexico, MpIo. Guryrrals: Nacapule Canyon, Bailey 26 (BH). Los Anegados, Sonora, in secluded caflon[e]s in the mountains about Guay- north of Bahfa San Caflos, Felger 3096 (ARIZ). Palmar, at Rancho mas, Oct. 1887, Palmer 311 (holotype GH!; isotype CASI). Palmar, NW of Bahia San Carlos, Felger 3100 (ARIZ). La Huerta, Washingtonia gracilis Paish, Bot Gaz. 44: 42O, fi,gs.8-10, 1907. Felger 3070 (ARIZ). Back of San Carlos Bay,27 Mar 1934, Bailey W. robusta var. gracilis Parish ex Becc. Webbia, ii, 197. 1907. 1 (BH). San Carlos Bay, ranch and palm grove in broad flood plain "Described from cultivated trees grown in San Bernadino and just inland from S[an] C[arlos] B[ay] Motel, 3 m elev., Boutin 2010 Riverside, California, probably native to Lower California." (BH). San Carlos Bay, John.ston 4345 (CAS, UC, US). San Carlos Bay, Dec 1960, Schnabel s.n. (BH, photo). N ofSan Carlos Bay and Vernacular names.-Abanico ('fan,' referring to fan Tetas Cove, 10 m elev., Boutin 2013 (BH). Ensenada Grande [= palm, term in Sonora for cultivated plants). Mexican Bahia San Pedrol, Feb-Mar 1947, Lindsay s.n. (BH). San Pedro Bay, Moran 4037 (BH).-Mpro. Hsnvosrllo: Cultivated in plaza prin- (Arizona, for cultivated plants). Zamij ctam cipal, coll. in vicinity of Hermosillo, Rose et al. 12518 (US).-Mpro. (Seri; Felger and Moser 1985). Mocrszuul: 20.6 mi E of Mazocahui on road to Moctezuma, ca. -Trunks 29OO ft, Felger 3347 (ARIZ). Km 128 W of Moctezuma on Hwy Description reaching ca. 15-20 m tall, 98- 14 to Mazocahui, 29'35-40'N, 109'55-60'W <1000 m, Joyal 2044 104 cm circumference, often fire-scarred. Petioles of (ARIZ).-Mpro. NAconr Csrco: 3.8 mi by road NW of Nricori Chi- juvenile to half-grown trees armed with strong, shaq), co, valley bottom, Felger 33O6 (ARIZ). Rancho Los Pescados, ca. orange-brown teeth 10-15 mm long along the full 20 air km SSW of N6cori Chico. 29'29'N. 108'59'W. ca. 780 m. length, some teeth double-pointed; petioles of taller Joyal & Silva 1727 (ARIZ). Rancho La Ciinega Aniba, 29"37- 38'N, 109'04'W 98O m, Joyal 2033 (AF.IZ). Rancho Napopa, S of trees often reaching 1.4 m long, the margins entire or Buena Vista, 29'29'N, 108"54'W 900-1000 m, Joyal & Silva 2316 with some basal teeth to 3.0-3.5 mm long. Hastula (ARIZ).-Mrro. Qurnmco: El Ranchito, 3 km N of Los Bajios, large, papery, more or less triangular, the margins ir- 27'42'N, 108'59'W 450-500 m, Joyal et al. 2417 (ARIZ).-Mpro. regular, fraying with age; leaf blades uniformly shiny Rosnnro TBsop.lco: San Martin, between Nuri and Rosario, 27'55'N, green on both surfaces, nearly flat, the free portion of 109"19-20'W 50O m, Joyal 2495 (ARIZ). El Sauz, 8 km NNW of Rosario Tesopaco, 27'55'N, 1O9"22-23'W,440-460 m, Joyal 2496 the segments usually drooping. (ARIZ). 6 mi S of Cedros, Wiggins 6433 (DS, US).-Mrro. Tern- Flowering branches often reaching ca. 2.6 m long, atr.: 7 .7 mi by road SW of Tepache, 29.4' N, I 09.6'W 545 m, Turne r longer than the leaves, arching, then drooping with et al. 71-53 (BH).-Mpro. Unrs: 10.8 mi N of l]res on road to weight of the fruits. Flowers white, short-pedicelled; Ray6n, Felger 3002 (ARIZ). Rancho La Raja, ca. 15 km S of Ures. sepds united below, the lobes with ragged margins; 29"16'N, 110"20'W 450-500 m, Joyal 2525 (ARIZ). Rancho I-a Noria Aguilarefla, ca.2O km N of Ures, 29"35'N, 110'23'W ca. 530 stanens s€parate, borne on the corolla tube. Flowering m, Joyal 2577 (ARIZ).-INy'.pro. YEcou: Rancho Yerbanis, N of Te- May and June, the fruits ripening in fall to early poca on San NicolSs-Sahuaripa road, 28'30-31'N, 109"10-ll'W. spring. Fmits ellipsoid, 7.5-9.0 mm long, blackish VOLUME 18. NUMBER 1 Palms of Sonora. Mexico 17

when ripe, the thin, fleshy pericarp edible and sweet spines. Cornett's (1986) note on W. filiftra confirms like a date, the fruits mostly falling with the dry, per- Bailey's earlier observations and speculates that there sistent calyx and short pedicel. Seeds 4.7-65 mm might have been a relationship with Pleistocene mas- long,4.5-4.9 mm wide, ellipsoid to globose, dark red- todons which had a vertical reach of about 6 m. How- 'entire brown, smooth or nearly so on the dorsal side, the ever, Sabal has petioles throughout its life his- lower surface (hilum side) wrinkled. 2n : 36 (Ei- tory, and certain Brahea species retain stout petiole chhorn 1957). spines throughout their lifespan.

Distribution -In Sonora native only to canyons and Specimens examined.-Mpro. Gunvuns: Can6n del Nacapule: Bci- oases in the Sierra El Aguaje at ca. (10-) 30-135 m. ley 3, and,262 (BH); Felger 3I18 (ARIZ). Los Anegados, Felger 3095 (ARIZ). La Huerta, Felger 3068 (ARIZ). Pirinola Canyon, Locally dense populations in paft occur the upper of Bailey 16 (BH). Cafr6n del Nacapule and in several very steep, sea- ward-facing canyons and oases in mountains along the ACKNowLEDGMENTS coast northwest of Bahfa San Carlos, e.9., La Huerta. Also locally abundant along canyon bottoms and oases The work by RSF is based on studies supported by in the canyon systems of Los Anegados and Las Ba- the Wallace Research Foundation and in part by the rajitas. In all of these canyons and oases seedlings and National Geographic Society grunt #6063-97. EJ ac- small plants are abundant. These palms are limited to knowledges the National Science Foundation Doctoral immediate margins of streams and at springs with per- Dissertation Improvement Grant #D8893I1035, the manent or near permanent water or wet soil where it Nature Conservancy Small Grant for Graduate Re- bcally outnumbers Brahea and Sabal. search in Conservation, the American Association of The limited distribution and extremely localized University Women Project Renew Grant, Arizona-Ne- tnoves of W. robusla in Sonora make it vulnerable to vada Academy of Sciences Grant-in-Aid of Research, bcal extirpation. Rose (1899: 255) found W. robusta and the Arizona State University Center for Environ- I Guaymas, and reported that "most of them have mental Studies. Many friends and colleagues have pro- bccn cut out and used as rafters for houses." Later vided generous assistance. In this regard we especially make no mention of this species occurring nat- thank Alberto Bfrquez, Gil Gillenwater, Phil Jenkins, at Guaymas. This species is otherwise native Kristen Johnson, Matthew Johnson, Les Landrum, in Baja California Sur. Merv Larson, Linda Leigh, Angelina MartinezYrtza4 Palms on the western slopes of the Sierra del Tigre Lucinda McDade, John Palting, Donald Pinkava, Ana as W. sonoraeby White (1948, Fig. 21) are Lilia Reina G.. Silke Schneider. Victor Steinmann. Na- Brahea, probably B. elegans. talie Uhl, Thomas R. Van Devender, Wilson, and David Yetman. Special thanks to Hermilo .-Although of very local and limited dis- Quero and Scott Zonafor sharing information and manuscript in its native habitats. tosether with the date review. For loans and accessto collections, we extend it is one of the most widely cultivated extra- our appreciation to the curators and staffs at the fol- palms in the world. This tall, stately palm is lowing herbaria: ARIZ, ASU, BCMEX, BH, CAS/DS, oommon street tree and landscape subject in north- DES, FI, GH, HBG, MO, R RSA, SD, UC, UCR, US, Mexico. southern Arizona, and southern Cal- and USON (University of Sonora, Hermosillo). . It is the fastest-growing of the Sonoran palms. f species is raised in containers or field-grown and ,, y transplanted even when large. LITERATURE CITED ll Both species of Washingtonia thrive best in dry or BArLEy, L. H. 1936. Washingtonia. Gentes Herb. 4:53-82. u ly dry climates with some cool winter nights. 1937. Erythea, the hesper palms. Gentes Herb. 4: 85-l 18. r gtonia filifera is much more frost-tolerant than 1943. Brahea, and an Erythea. Gentes Herb. 6: 196-197. la robusta (see Turner et al. 1995). In cultivation I7. BARRow, S. 1998. A monograph of Phoenix L. (Palmae: Coryphoi- r- grows well with hot and either dry or humid deae). Kew Bull. 53: 513-575; ly BECCARI,O. 1907. Le palme americane della tribi delle Corypheae. climates,whereas W. filiftra is susceptibleto Webbia 2: 1-343. of bud rot when cultivated in reqions with a sum- BRo$'N. D. E. (ed.). 1982. Biotic communities of the American rr rainy season or wet, foggy climates such as coast- Southwest-United States and Mexico. Desert Pl. 4: l-342. rg, rl California (Bailey 1936). BuReuEz. A.. AND A. MARTiNEZ-Yp1zrp.. 1997. Conservation and ittr landscape transformation in Sonora, M6xico. J. Southw.39:3'71 d; Notes.---:flte fruits are eagerly eaten by many 398. nd various mammals including badgers, coyotes, and R. S. FELGER.r.No D. YpnaeN. In Press. Biodi- 1rS; versiN at the southern desert edge in Sonora, Mexico. 1z R. Rob- trcoons. Seeds are often found germinating in decay- ng ichaux [ed.]. Ecology and conservation of the Sonoran Desert rly ing raccoons droppings. Bailey (1936) pointed out that plants: a tribute to the Desert Laboratory. University of Arizona ish petioles of mature trees have few or no marginal Press. Tucson. AZ. l8 Felger and Joyal ALISO

ConNsrr, J. W. i986. Spineless petioles in MooRE, H. E. 1973. The ma;c group; of palm and their distdbution. (Arecaceae). Madrofto 33 7 6-78. GentesHerb. ll: 17-l-f l. Dr PESo,C. C., eNn D. S. MersoN. 1965. The "Descripci6n Seri Indians in 1692 NENTVTc,J. 1672. geogrdtica narural y curiosa de la as described by Adamo Gilg, S.J. Arizona and the West 7: 33_56. Provincia de Sonora," rmanuscripn. Transl. bv Germ:in Viveros, ErcnHoRN, A. 1957. Nouvelle contribution e l,6tude caryologique with an introduction. nores. appendrr. and index. Mdxico: Archivo des palmiers. Rdv. Cytol. Biol. Vdg.18: 139_151. General de la Nai6n. l97l Fnr-crn, R. S. 1966. Ecology of the Islands and Gulf Coast of So_ NrxoN, R. W. 1950. Imporred r.arieties of dates in the United States. nora, Mexico. Ph.D. dissertation. University of Arizona, Tircson, U. S. D. A. Circular 83-1.llJ pp. 42.460 pp. PENNrNcroN,C. W. 1980. The marerial culrure. The pima Bajo of 1999. Flora of Cafr6n del Nacapule: a desert_bounded trop_ Central Sonora, Mexico. \'ol. l. Unir.ersitl. of press, Salt ical canyon near Guaymas, Sonora, M6xico. proc. San Diego Soc. Lake City, UT. .110 pp. NaL Hist. no. 35 (in press). PFEFFERKoRN,l.1949. Sonora: A description ofthe province. Tians_ AND M. B. JoHNsoN. 1995. Trees of the Northern Sierra lated by T. E. Treutlein. Coronado Cuarro Centennial publications. Madre Occidental and Sky Islands of southwestern North Amer_ 12. University of press. Albuquerque, NM. 329 pp. ica, pp. 71-83. In L. p F DeBano, F Ffolliott, R. H. Hamre [eds.], Queno, H. J. 1989. Flora gen6rica de Arecdceas de Mexico. ph.D. and management of the Madrean Archipelago: The dissertation. Universidad Nacional Autdnomo de M6xico, M6xico, Sky Islands of the southwestern United States and northern Mex_ D.F. 142 p + 24 figs. ico. Rocky Mountain Forest and Range Experiment Station, U. S. 1992. Current status of Mexican palms. principes 36: 2O3* D. A. Forest Service, Ft. Collins, CO. 2t6. rNn p E. Joynl. 1998. Arecaceae, p. 474.12 S. Martin, D. Rre, A. M. 199'7.At the desert's green edge: an ethnobotany of the Yetman, M. Fishbein, P Jenkins, T, R. Van Devender, and R. K. Gila River Pima. University of Arizona press, Tucson, AZ. 430 Wilson [eds.], Gentry's Rio Mayo plants. University of Arizona pp. Press.Tucson, A2.558 pp. Rsno, R. W. 1964. Palm chromosome studies facilitated by pollen AND M. B. MosER. 1985. people of the desert and sea: culture on colchicinelactose medium. Stain Technol.3g: 99_106. ethnobotany of the Seri Indians. University of Arizona press, Tuc_ Rosp, J. N. 1899. Notes on useful plants of Mexico. Contr. U. S. son, AZ. 435 pp. Natl. Herb. 5:209-259 r.-o M. F WrlsoN. 1995. Northern ScHNeetL, R. O. 1962. Some palms of northwestern Mexico. prjz_ and Its Apachian outliers: A neglected center of biodiversity, pp. cipes 6: 5-9. 36-59. 1964. Palms along princi_ In L. F DeBano, P E Ffolliott, and R. H. 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