?t* iir *«>' 7tu OCCASIONAL PAP OF THE CALIFORNIA ACADEMY OF SCIENCES No. 130, 119 pages, frontis. +6 pis., 62 figures, 11 tables December 28, 1978 The Agaves of Baja California By Howard Scott Gentry Desert Botanical Garden, Papago Park, Phoenix. Arizona 85010 ^O^DEl^y^^ SAN FRANCISCO PUBLISHED BY THE ACADEMY The Agaves of Baja California Frontispiece. The wonderful vegetation north of Punta Prieta with Sierra San Luis in the background. Agave shawii goldmaniana. Yucca valiJa, Fouquieria splendens. and Idria columnaris are all prominent here. Photographed, 21 February 1968. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES No. 130, 119 pages, frontis. +6 pis., 62 figures, 11 tables December 28, 1978 The Agaves of Baja California By Howard Scott Gentry Desert Botanical Garden, Papugo Park. Phoenix. Arizona 85010 WoED\^''^^ SAN FRANCISCO PUBLISHED BY THE ACADEMY COMMITTEE ON PUBLICATIONS Laurence C. Binford, Cliain>uin Tomio Iwamoto. Editor Paul H. Arnaud, Jr. William N. Eschmeyer George E. Lindsay us ISSN 0068-5461 The California Academy of Sciences Golden Gate Park San Francisco. California 94 11 8 PRINTED IN THE UNITED STATES OF AMERICA BY ALLEN PRESS. INC.. LAWRENCE, KANSAS TABLE OF CONTENTS Page List of Color Plates vi Acknowledgments vii Abstract viii Introduction 1 Environment and Evolution 1 Indigenous Uses of Agaves 3 Agave Measurements and Floral Ideographs 7 Genus Agave 10 Key to the Groups 10 Group Deserticolae 13 Agave deserti 15 Agave mckelveyana 25 Agave subsimplex 27 Agave cendata 35 Agave margaritae 48 Agave sobria 48 Agave moranii 58 Agave aveUanidens 61 Agave gigantensis 63 Agave vizcainoensis 67 Group Campaniflorae 70 Agave capensis 72 Agave aurea 78 Agave promontohi 81 Group Umbelliflorae 82 Agave shawii 86 Agave sebastiana 96 Group Datyliones 97 Agave datylio 97 Appendix: Exsiccatae 101 Glossary of Special Terms 114 Literature Cited 116 Index 117 LIST OF COLOR PLATES Plate L (Left) Agave deserti. La Rumerosa. (Righi) Agave margaritae Bahi'a Santa Maria 28 Plate 2. (Top) Agave gigantensis on volcanic top of Sierra de las Palmas. (Bottom) Agave cerulata dentiens. Isla San Esteban 29 Plate 3. (Left) Agave cerulata dentiens in natural cluster and habitat on Isla San Esteban. (Right) Agave promontorii. La Aguja 32 Plate 4. (Top) Agave aiirea 19 km southeast of La Paz. (Bottom) Agave aurea at southwest end of Mesa de San Geronimo 33 Plate 5. (Top) Agave shanii. Inflorescence of plant. (Bottom) Agave shawii with Simmondsia chinensis along the Pacific shore north of Ensenada 38 Plate 6. (Left) Agave shawii spp. goldmaniana near Punta Prieta. (Right) Agave sebastiana on West San Benito Island 39 ACKNOWLEDGMENTS Of first mention is Carl O. Erlanson. former Chief of Plant Exploration and Introduction. Agricultural Research Service, who first assigned me to formulate a taxonomic and eco- nomic monograph of the genus Agave. I am grateful to him for such a challenging assign- ment. Others in that unique institution who assisted in my early efforts were Arthur S. Barclay, Frederick J. Hermann, Howard L. Hyland, Ernest Imle, Quentin Jones, Fred- erick G. Meyer, Bernice Schubert, and C. Earle Smith, Jr. Field assistants who were especially helpful in Baja California included Juan Arguelles of San Bernardo, Sonora, and Frank L. Cech of San Diego, California. I am indebted to the fine artists Regina Hughes, Lucretia Hamilton, and Wendy Hodg- son, whose drawings enhance this book. I wish also to thank Bruce Gentry, John McClure, and Reid Moran for the photographs they provided for illustrations and for study. Donald Pinkava and Reid Moran reviewed the script and suggested many changes, many of which I gratefully accepted. Thomas Howell has competently edited my Latin diagnoses. The color illustrations in this book are due to the generous support of E. R. Le Roy of San Francisco, California. Furthermore, through correspondence, he has given the author interest and encouragement during the past decade. Final efforts in field and laboratory were provided for by the National Science Foun- dation with their fund Grant No. BMS74-24553. With this and the facilities so magnani- mously provided by the Desert Botanical Garden and Director W. Hubert Earle, I could do no less than my best to make this a useful and enduring work. Thank you all, including the many others not called by name! Howard Scott Gentry June 5, 1975 ABSTRACT Gentry, Howard S. The agaves of Baja California. Occasional Papers of the California Academy of Sciences . no. 130. 119 pages, frontis. + 62 figures, 11 tables, 1978. — This is a systematic and economic account of plants of the genus Aguie. family Agavaceae, that are indigenous to Baja California and the adjacent areas of the Gulf of California. Ta.xonomically, it is a revision of the three generic groups established by William Trelease in 1912: Deserticolae, Campaniflorae, and Umbelliflorae. Of the 23 taxa described, 4 are proposed as new species: Agave capensis. A. gigantensis. A. moranii. and A. vizcainoensis. Eight other new taxa proposed are: A. deserti ssp. pringlei: A. deserti ssp. simplex: A. cerulata ssp. dentiens; A. cerulata ssp. nelsonii: A. cendata ssp. subcerulatu: A. sobria ssp. frailensis: A. sabria ssp. roseana: A. shawii ssp. goldmaniana. Keys to assist in identification are provided for groups, species, and subspecies. Maps show the distributions of all taxa and nearly all are copiously illustrated with line drawings and photographs. The economic aspects of the plants are represented by historical notes on Indian uses. Under various species there are reports of modern attempts to exploit agaves for fiber and chemical constituents. Tables on food con- sumption, chemical analyses, and fiber tests summarize the information. Sustaining documentation is provided in a bibliography, a glossary of special terms, and extensive citations of the specimens reviewed in many U.S. herbaria. The latter have been organized in an appendix of exsiccatae listed alphabetically by species, states, and collectors with respective herbaria. This report is the culmination of a field study of the wild populations of these plants made intermittently by the author over a 30-year period. INTRODUCTION This work is primarily a taxonomic revision of three groups of the large genus Agave in the family Agavaceae: Deserticolae, Campaniflorae. and Umbelliflorae. Its scope has been enlarged beyond the taxonomic skeleton in order to in- clude the historic uses of the plants and to report investigations of their chemistry and fiber. These investigations were made by the Agricul- tural Research Service of the U.S. Department of Agriculture, the procurement of the samples being done by the writer. The results are con- densed in tabular form with brief textual com- ments. For the purpose of brevity, many special articles in the press and journals have not been repeated or reviewed here. A great part of this study is based on extensive observations of the wild agave populations. These observations form a basic part of the writer's taxonomic con- cepts, and for this reason, many field notes are given space in the accounts of species. Trelease's (1912) account of the agaves of Lower California included descriptions of 23 species. It was based on a relatively small scat- tering of specimens from field collections made principally by J. N. Rose, T. S. Brandegee, E. W. Nelson, and E. A. Goldman. The present account recognizes 16 species, 3 of which are proposed as new, 8 subspecies, and 1 variety. Several taxa are outside the confines of Baja California, but they complete this revision of the group Deserticolae. Trelease arranged this spe- cies in groups. I regard these groups as equiv- alent to sections, but am not taxonomically for- malizing them as such in this revision. In addition to the three groups listed above, there remain two Californian taxa that Trelease put into his small group Datyliones. I am in- cluding them in this work also so that all known wild agaves of Baja California are accounted for. As further preamble, the environmental back- ground is discussed below. The harmony that naturally exists between agave and other life forms of the peninsula is clearly apparent in the frontispiece. ENVIRONMENT AND EVOLUTION The California Gulf region is physiographical- ly unique with its long slender peninsula sepa- rated from the Sierra Madrean mainland by a rather deep seaway. The geologic structures, especially the widespread volcanic formations. all indicate a tectonically active history during much of the Tertiary Period. The dynamic San Leandro fault system of upper California does not stop at the U.S. -Mexico border, but contin- ues in the Gulf of California along the length of the peninsula. The escarpments on the eastern side of the peninsular mountain axis are the result of upthrusts along this fault zone, at least rela- tive to the "graben"" or trough where the sea lies. Until recently the reading of the older geologic reports (as those of Darton 1921, and Schuchert 1935), left me with the physiographical impres- sion that the present peninsula is actually a chain of land segments gradually united by volcanic magmas and land emergence. While this still ap- pears true in most instances, modern students of marine geology, equipped with cunningly de- vised electronic equipment, are developing high- ly illuminative information about earth crustal movements. The proponents of tectonic plate movements (growing out of the continental drift hypothesis) now regard the peninsula as part of a land raft slipping northward along the western edge of the North American continent. The Sci- entific American with its "Continents Adrift" (Wilson et al. 1972) has put together a fine series of articles on this large subject. Other articles have been carried in Science during the 1960's. One recent study by Larson, Menard, and Smith (1968) concludes that southern Baja California was torn from the Mexican mainland about 4 million years ago and since has been rafted 260 km to the northwest at a rate of 3 cm per year. This is in conflict with earlier geologic studies that generally place the opening of the Gulf in the Miocene some ten to fifteen or more million years ago. While such discrepancies are being adjusted by further work, we can consider the possible effect of land movements on our plant subjects. It would be presumptuous to try to correlate Agave evolution with land changes in the Cali- fornia Gulf region, but nevertheless, I will essay it in a suggestive and tentative way. Several of the Agave complexes are limited to definite land areas, and, in some cases, these land areas ap- pear to have been separated by seaways during much of the upper Tertiary and Pleistocene pe- riods. In Figure 1. I have drawn a map of pen- insular land areas that were apparently separat- ed from one another during much of these geologic periods. On these separated land areas OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 .»• \ \ .J \ «. 1 k^ 7 '■ \ \ £^^ ^^ -"4. ^ ^•>^ ^ ^ r ii V( -^j "^ \ A \ \ L^ ti { J .A y K \^' (^ \fi \ % \ - FiGLiRE \. Physiographic staging and some agave distri- butions in the California Gulf region. are written the respective agaves growing there today. It all suggests that many of the distinctions among the species appear to have been fostered by the seaway separations. Starting in the north, various situations can be outlined as follows: 1 . The A^«v'^ deserti complex was split into two groups by the Gulf: A. deserti simplex on the mainland northeast; A deserti deserti and A. deserti pringlei on the peninsular side. 2. Agave sohria is set off rather sharply mor- phologically and anatomically from A. ceru- lata, the two complexes also being separated by a postulated mid-peninsular sea portal. 3. Agave vizcainoensis, resident upon what was obviously a western island aligned with Ced- ros Island, with its deep floral tube, is quite distinct from any of its relatives in the De- serticolae. 4. Agave margaritae, A. sohria roseana, A. cer- uiata dentiens are all island endemics still limited, or nearly so. to their respective is- lands. 5. The Campaniflorae are limited to the south- ern part of the peninsula (south of the midpeninsular sea portal); A. aurea is wide- spread along the Sierra de la Giganta volca- nics and is also scattered over the Cape Dis- trict granitics. Agave promontorii and A. capensis are endemic to the Cape mountains. If their progenitors were originally based on the Mexican mainland 4 million years ago, one would expect to find near relatives along the Nayarit-Jaliscan coast, but no such rela- tives have been found there. The Campani- florae are quite distinct from both mainland and peninsular agaves. This suggests long isolation, longer than the 4 million years al- lowed by Larson et al. (1968). The overall picture suggests some coincidence between Ag«ir evolution and land evolution and that xmny Agave species do not migrate readily. On the basis of fossil records for many flowering plant families, we can assume that agaves and their ancestors have been on the North Ameri- can continent throughout the Tertiary Period. But without agavoid fossils this is uncertain. Nor do we really know how long agaves have inhabited the peninsula. Just a few fossil impres- sions of spines, seeds, or capsule sections would be of great historical value, and I hope the pa- leontologists will be looking for them. Most of the California Gulf region has been well characterized as the Sonoran Desert (Shreve 1951), and the agaves living there are successful succulent xerophytes. Most of the desert species make do with only 90-250 mm (4-10 inches) of rain annually, and some of them, such as A. deserti and A. ceridata, can survive several years with little or no rainfall. A. promontorii, a large fleshy mesophytic species living on the Cape mountains, receives an an- nual average of ca. 750 mm (25 inches). A. .s77 0 J. 91 deserti subsimplex J Carrizal, Sonora 197 cerulata subcerulato San Ignacio 2O0t gigantensis lOO" 0*- 97 San Javier 317 deserti J Borrego Park, Calif. 2oot margarltae 100.. OA 90 Bahia magdalenae 81 GENTRY: AGAVES OF BAJA CALIFORNIA mckeiveyana 200t i D 100 + Kingman, Ariz 170 0 A sobria f railensis San Jose de Cabo 266 morani i Valle Trinidad 180 200t lOO" 0 J- sobria roseana La Paz 185 Comondu 156 200- vizcainoensis 100". 0 A Punta Abreojos 103 Figure 2. Rainfall (silhouettes) and flowering (bars) perimeters of the Deserticolae. Relevant meteorological stations with average annual rainfall given in millimeters. Data from Atlas Meteorologico de Mexico (Servicio Meteorologico Mexicana 1939), Hastings (1964). Hastings and Humphrey (1969). and U.S. Weather Bureau. Flowering periods based on herbarium specimens and field observations, supplemented by plants in cultivation. Uncertainty expressed by broken lines. from 1738 to 1768. After his return to Spain in the latter year, he wrote a lengthy account on the natural history of ancient California (Barco 1973). Other historians, such as Clavigero (1789). made use of Barco's report. Barco's de- scriptions of the uses of agaves by the native peoples, especially the Cochimi Indians inhab- iting the Sierra de la Giganta region, is one of the most accurate and revealing I have read. Because of its excellence, I have translated from the Spanish to quote much of it here. The Californians did not use, nor do they now use the mezcal for making beverage. They employ it only for eating when the plant is ready or mature. They know when it has reached this stage, as the bud. which had its leaves tightly folded together, has spread out with its leaves open, sep- arated one from another. In this stage, the last leaves to open are much smaller than the rest, and there is ready to sprout in place of the leaf bud. a shoot which, if the mezcal is not cut. will come out as thick as a man's arm. and which will rise to four to six varas. At its tallest it forms a great panicle composed of many small branches of yellow flow- ers which produce its seeds. When ripened, the whole plant dries up and the mezcal is worthless. The mezcal is not eaten raw but only roasted, and the whole operation, which is carried out by women, is done in the following way. Three or four women or more leave their rancheria or settlement in the morning, each provided with a carrying net on the shoulder over which it is maintained by some coarse cords which pass to the forehead of the woman. In this net. they carry the mezcales as well as whatever else needs to be carried all year. This net the Cochimi women call uani. and the Spaniards call it aparejo, because it serves to support and carry cargo. Over the forehead they put a piece of deer hide, doubled so that the cords do not cut into their foreheads when they are loaded. On such occasions they go bent over, lowering not only the head, but also inclining the shoulders and back because this rein- forces the head against the weight of the cargo which it must resist. Besides the net, each woman carries a large heavy knife. In place of a machete to cut the trunk of mez- cal, they have a little slab of hardwood, three to four fingers wide and two or three palmos [.'51-64 cm] long, in the form of a shovel blade but without point, but they bevel the end of it to cut the mezcal. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Figure 3. Agave shawii goldmaniana showing the old reclining trunks radiating out from a common mother rosette, long gone. Arriving at whatever locality they are going, they sepa- rate and each one goes to look for mezcales. Finding a mature one, they take in hand their hardwood tool and apply the beveled end to the stem of the plant. The upper part of the plant having the leaves is the thickest, juicy and tender; this they call the head of mezcal and is what is eaten. Applying then the beveled end of the hardwood tool to the base of the head (where the green leaves begin), they strike heavily the upper end of the tool with a rock. Little by little they behead the mezcal. Then with their knives they cut off the leaves near the head of the plant, leaving a small basal portion of the leaves, about two fingers, which here are about a finger and a half thick, stuck to the head. When the head is so trimmed, it remains the size of that of a man. Then they look for more, and in the afternoon, each woman returns with eight or nine mezcales carried in the uani. which is a good load! Sometimes they travel one or two leagues [about 4-9 km), adding on top some wood to roast them. Near the rancheria, they make fire and add to it some stones that are not too large. When the wood is consumed and the stones as hot as red coals, they spread the coals and stones a little with poles, and then, one by one, place the mezcales in accommodation onto the fire according to GENTRY: AGAVES OF BAJA CALIFORNIA their judgement. Altogether it forms a mound, and they cover it with nearby hot earth, which reconcentrates the heat and delays its dissipation. Thus, it is left for twenty- four hours and. more frequently, for two nights and one day; and they dig it out all well cooked. The Mexicans and also all the Spaniards of New Spain call this method of roasting tatemar, and the thing that is roasted they call tatema. They use it to roast various things, principally heads of sheep and cows, etc The meat remains tender, juicy, mild, and gustatory. When the mezcales are taken out and left to cool, the woman has food for her family for three days, more or less, according to the number of persons. The bases of the leaves (the white growing tissue) which were left on the stems are also eaten, but we could better say they are chewed, as they have little more than a sweet juice and tough fibers which are not eaten. They spit out this bagasse like "tacos de escopete" (shotgun wads). This bagasse is not always lost, because the old men and women (who, although given food, are always hungry) recover these tacos that were ejected on the ground. When they are well dried, they ground them to powder between stones and eat them. The remainder of the mezcal, freed of its leaves, is better esteemed as solid food. It is cut with a knife into slices and eaten with gusto; it is almost as sweet as conserves made with syrup. However, it leaves the mouth asperous, as with those of delicate palate who are not accustomed to it. In some regions, much finer mezcales are found that contain little or no fiber. This is the most common food from October inclusive until April. In May they cease to eat it because the time of ripening is past or the new shoots are past and dry, and they will not start to ripen until the beginning of autumn. Then also they have no appetite for a hot food like mezcal in the warm season. When these plants are not cut, they produce a flowering shoot, as we stated above, and in its flowers a good quan- tity of nectar is contained in the calyx of each flower. In order to collect this, they cut each small branch of flowers separately, turning them upside down over an adda. Each flower is emptied of its contents, and in a short while, the adda or wooden bowl is filled. This nectar is sweet but rather nauseous, but when boiled and the spume skimmed off, it becomes much better. For the rest of the account, there are other species of mezcales that are not eaten, some because they are bitter and others because they cause bellyache. Finally, from the leaves of mezcales they extract fiber, with which they make both fine and coarse cords for making their nets and for other uses. . . . The roots of the mezcal are good for noth- ing, at least in California. [Barco 1973: 122 (transl. from Spanish)] Additional notes on the indigenous uses of agaves are given below under appropriate group and species headings. AGAVE MEASUREMENTS AND FLORAL IDEOGRAPHS Measurements Measurements of organs and parts have been widely employed in the classification of both plants and animals. Measurements provide basic data for direct and analytical comparisons be- tween species and other taxonomic categories or comparison of individual specimens. Total length of leaf is measured from leaf axil to tip of spine. Width is generally given at its widest in mid-leaf, as in dried, pressed specimens. Teeth measurements include the broad corneous base and are only approximate because most teeth are curved or flexed in various ways, making precise measurements impractical. Height and width of rosettes, as well as height of panicle (overall length of inflorescence from ground lev- el to tip), are also usually only approximate or estimates. Unusual variations are not given in descriptions. The ones given should be regarded as a general mean or average prevailing in the subject population or species. Figure 4 shows the flower organs selected for measurement and how the measurements are made. However, my experience has shown that to be reliably comparable, the measurements must be made in a standardized way subject to certain qualifying conditions imposed by the de- veloping flower. The flowers used should be nor- mal, healthy, and precisely in the stage of anther dehiscence. The flower bud grows slowly but opens and expands very rapidly during anthesis (expansion of the flower) within one day. Therefore, mea- surements for comparative purposes must be taken at about the time of anther dehiscence, which occurs during a few daylight hours. How- ever, flowering begins at the base of the inflo- rescence and continues to move upwards in the inflorescence for two to four weeks, depending upon individual plants and species. One can usually find a series of flowers in late-bud stage, at anther dehiscence, and after dehiscence, all in one flower cluster. I prefer to have measure- ments of flowers taken just before, during, and right after stamen dehiscence so that maturation is bracketed. For this purpose, I collect and im- merse the flowers in a pickling solution of one part formaldehyde, three parts 95 percent alco- hol, and six parts water. Pint-size canning jars in cartons of one dozen are suitable for field work and accommodate all except a few of the very largest flowers. For permanent storage, they must be transferred to laboratory jars with noncorrosive resinoid tops. Labels are put in- side the jars. The pistil continues to grow after anther de- hiscence, usually for two or three days, OCCASIONAL. PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Table 2. Representative flower measurements (in mm) of the genus A^ave, Group Deserticolae. (* — measure- ments from dried flowers, relaxed by boiling; ** — measurements from fresh or pickled flowers.) Taxon & locality Ovary Tube Tepal Fil. ins. & length Total Anther length Coll. No. cerulata cerulata Catavinya Laguna Chapala San Euis Gonzaga Bay Punta Prieta Valley cerulata nelsonii S of San Miguel S of San Miguel cerulata subcerulata W of San Ignacio E of San Ignacio Isla San Marcos deserti deserti Pinyon Flats Pinyon Flats San Felipe, Calif. San Felipe, Baja Calif. deserti pringlei San Matias Pass San Matias Pass deserti simplex Harquahala Mt. Harquahala Mt. Providence Mt. mckelveyana Black Mountains Hualapai Mt. S of Wikieup subsimplex Libertad Desemboque sobria sobria Comondu Comondu sobria roseana Espiritu Santo La Paz sobria frailensis Punta Frailes Punta Frailes Punta Frailes 24 4.5 X 12 16 X 6 4-4.5 30 14 45** 23159 31 6 X 12 20 X 6 6 32 19 57** 19973 26 3 X 13 17 X 7 3-4 35 16 45** 23298 24 4 X 13 18 X 8 5 42 18 45** 23302 35 3 X 12 17 X 4 3 40 17 55** 23324 27 5 X 13 21 X 7 5 30 54** 11179 28 2.5 X 12 18 X 5 3-2 34 18 48** 23170 24 5 X 13 20 X 6.5 5 36 20 48** 23175 27 3 X 12 19 3 33 20 50** 11892 33 4 X 12 17 X 6 5 36 17 53** 19759 31 3 X 11 15 X 4 3 37 16 50** 23326 26 4 X 15 17 X 8 6 28 15 48** 19940 38 4.5 X 12 18 X 7 4.5 40 18 60** 23286 24 6 X 12 14 X 5 6.5 20 14 43** 19959 22 7 X 15 16 X 7 7 21 15 44** 19959 25 6 X 13 20 X 3^ 5-6 38 21 50** 23404 26 7 X 12.5 17 X 4.5 5-6 40 20 50** 23404 28 8 15 5-6 17 52** 323IA 17 3 X 8 12 X 3 2-3 28 12 30** 21979 21 4.f — f 15 X 5 4 25 16 40** 22312 21 4 X 12 12 X 4 2-3 29 12 36** 23000 23 5 X 11 17 X 8 4-5 30 16 45** 3880 25 3 X 10 14 X 7 2 28 15 43** 14171 25 3 X 9 21 X 3 3 47 ~ii 48** 1989 29 4 X 10 17 X 3 3 35 20 49** 11882 40 5 X 12 22 X 5 4-5 42 23 66** 11277 26 3.5 X 11 18 X 5 3.5 29 19 46** 11869 37 3.5 X 12 19 X 4 3 40 20 58** 11264 31 2-3 X 12 24 X 6 2-3 38 23 57** 11257 24 4 X 11 17 X 4 4 32 18 43** 11858 Bajada of Sierra San Pedro Martir Bajada of Sierra San Pedro Martir Bajada of Sierra San Pedro Martir 41 6 X 13 23 X 7 40 5 X 13 24 X 6 26 4 X 13 19 X 7 6-5 46 2! 70** 23287 5 43 21 68** 23287 4.5 32 17 49** 23287 GENTRY: AGAVES OF BAJA CALIFORNIA Table 2. Continued. Taxon & locality Ovary Tube Tepal Fil. ins. & length Total Anther length Coll. No. avellanidens Paraiso Mesquital Mesquital Calmalli gigantensis Sierra Falmas Sierra Falmas Sierra Falmas vizcainoensis Cerro Tordillo Picachos Santa Clara 31 6 X 15 16 X 4.5 6 38 18 52** 23187 34 5 X 14 19 X 6 5 46 18 57** 23186 18 4 X 12 15 X 5 4 35 15 37** 11932 35 6 X 15 22 X 7 6 45 23 65** 23184 30 5 X 13 22 X 5 5 44 21 56** 23320 25 4 X 12 23 X 5 4 25 24 51** 23320 27 5 X 12 18 X 5 5 37 20 SO** 10324 39 13 X 15 21 X 4 9&8 50 21 72* 7469 35 8 X 15 26 X 5 6 63 25 68* 7713 pedicel brocteole ovory -'^-)^v,4_ i. Hi^fj^ A 0 - ovory body length, n - neck of ovory length, t - tube length, fi- filament insertion (measured to bottom of t-tube), s- tepal lengths, f - filament length, a - anther length, tl-total length Figure 4. Cross section of agave flower with parts measured and a tuhe/tepal ideogram, x. The white column represents the tepal, the black the tube, and the black square the insertion of the filament in the tube. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 when the stigmas open and are receptive. Be- cause of this later growth, measurements of pis- tils at anther dehiscence have no comparative value and have not been made. Floral Ideograph The measurements can be analytically applied to form the symbolic floral ideograph. The var- ious forms of ideographs and their applications were very ably described and applied by the late Edgar Anderson (1949). The form ideographs may take is more or less determined by the or- gans which the taxonomist finds best represen- tative of the variability that expresses specia- tion, introgression, or developmental features of a related group of organisms. The ideograph here selected shows the tube/tepal ratio, togeth- er with the insertion of the filament within or on the rim of the tube and, more rarely, on the base of the tepal. The ideograph is a kind of taxo- nomic shorthand that throws into perspective the selected characters, so strikingly apparent in Figures 5 and 6. Old dried flowers are shrunken and distorted by the growing ovary, and measurements of them are unreliable. They are little improved by relaxing. Berger, at La Mortola Botanical Gar- den, pressed split flowers under great pressure and, so treated, their proportions remained fair- ly reliable. So are split flowers from the pickling jar. The junction of tube and tepals is a particu- larly critical level, as the length of both organs derive from this. In measuring the length or depth of tube, one must note the various tepal sinuses because some may be deeper than others in the same flower. The upper limit of tube is taken at the median level of the sinuses. Where there is marked overlap of tepal bases in the sinuses, the median point of the overlap is taken as the upper limit of tube. There is, thus, a built- in indefiniteness about many tubes that may amount to a 1-mm variance or a margin of about 10 percent inexactness for tube measurement. Also, one must watch for damage by insects and birds, which sometimes split the sinuses. With measurements of three or four flowers, made after splitting the flower lengthwise, these diffi- culties are largely overcome, and the mean or median measurement is suitable for compari- sons with other collections. Diameter of tube is its widest extent, usually at about the level of filament insertion. Width of outer tepal is the median width just above the basal widening, or at about V3 to V2 tepal length above the tepal sinuses. This is bound to be a general-impression kind of mea- surement as it depends on degree of wilting or degree of inrolling and cannot, therefore, always be precise. Where there is a marked difference in length of outer and inner tepals. the length of the outer is given first. Width of the inner tepal is not giv- en, but its form, shape, or other qualifying char- acteristics are indicated in the description of species. Floral ideographs and the measurements on which they are based are given for each group, following the group description. GENUS AGAVE Agave Linnaeus, Species Plantarum 1:323. 1753. Succulent rosettes, monocarpic or polycarpic, perennials or multiannuals with long-lived leaves, frequently suckering at base and occa- sionally bulbiferous in the inflorescence; roots hard fibrous, radiately and shallowly deployed; stems thick, very short, usually shorter than the terminal bud, simple or branched; leaves large, generally succulent, spine-tipped, margin armed or unarmed with teeth; inflorescence tall, brac- teate, scapose. spicate, racemose, or paniculate with flowers in umbellate clusters; flowers most- ly large, generally proterandrous; perianth tu- bular to shallowly funnelform, the six segments erect to variously curved, similar or dimorphic, imbricate in the bud; stamens six. exserted; fil- aments long, inserted in tube or on tepal bases; anthers versatile; ovary inferior, three-celled, succulent, thick-walled with numerous axile ovules in two rows; pistil elongate, filiform, tu- bular; stigma three-lobate, papillate glandular; fruit a dehiscent, loculicidal capsule; seeds flat- tened, black. Subgenus Agave: inflorescence paniculate, flowers in umbellate clusters on lateral branch- es. Type species of genus and subgenus: Agave americana L. Sp. PI. 1:323. 1753. described from cultivated European plants. KEY TO THE GROUPS la. Leaves ensiform, 10-20 times longer than wide at mid-blade; rosettes suck- ering widely with elongate rhizomes _. Datyliones (p. 97) GENTRY: AGAVES OF BAJA CALIFORNIA DESERTICOLAE subslmplex Felger & Bezy 14171 Figure 5. Flower sections representative of the Deserticoiae. OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 DESERTICOLAE deserti 19759 19759 23326 23326 19741 19940 23285 23286 deserti simplex desert i pringlei U i:! d^k U U Ji 23404 23404 23404 23404 Eng. 3231A 22990 22990 19959 cerulata cerulata 23159 19973 19973 Jar. 23298 23298 23302 23302 23306 cerulata nelsonii cerulata subcerulata 23324 11179 11179 11679 11185 1991 23170 23175 11892 mckelveyana subsimplex IL Ji Jl Ji Ji fl 21979 22312 23000 23000 286 sobria sobria sobr j a f railensis 4486 3880 14171 sobria rosea na H JIIL 1989 11882 HBG JlJl 11257 11858 11869 GENTRY: AGAVES OF BAJA CALIFORNIA 13 DESERTICOLAE avellanidens JlJiJiJiJiJiJliiJl 19948 Bge6 23187 23186 11929 11932 23184 23184 23184 gigantensis JlJljlJlJl 23320 23320 23320 23320 23320 10327 10324 10324 JlJlJl 7693 23287 23287 vlzcoinoensis A 7469 Figure 6. Floral ideographs of the group Deserticolae, showing relative proportions of the tube (black) to outer tepal (white column), and level of insertion of filament (black square). Measurements are listed in Table 2. lb. Leaves not ensiform, only 2-10 times longer than wide; rosettes not suckering or bearing suckers appressed to the base __ _ _ 2 2a. Stems short, usually about as wide (leaf axil to leaf axil) as long (except A. ca- pensis): branches of the panicle not borne in large succulent bracts, the um- bels small- to m.edium-sized; flowers smaller (40-70 mm long), tubes shallow or broad and open 3 2b. Stems elongate, longer than broad; branches of the panicle usually borne in large succulent bracts, the umbels broad and massive; flowers large (70-100 mm long), tubes deep Umbelliflorae (p. 82) 3a. Plants generally green, nonsurculoseor branching from leaf axils; leaves rela- tively smooth; flowers red to purplish in bud, opening to light orange, campanu- late with broad open tubes and curved tepals Campaniflorae (p. 70) 3b. Plants light gray to yellowish green, sur- culose or single, not branching from leaf axils; flowers greenish yellow, funnel- form with short tube and ascending te- pals Deserticolae (p. 13) Group Deserticolae Trelease, Missouri Bot. Gard. Rep. 22:45. 1912. Plants small- to medium-sized, glaucous gray to greenish, freely suckering, or medium-sized to large, green, nonsurculose, the rosettes acau- lescent to short caulescent; leaves rigid, coarse- ly fibered, with thick cuticle, narrowly lanceo- late and with weak, easily detached teeth, or broader and with firmer teeth; panicle narrow with short lateral branches, dry scarious pedun- cular bracts, and small umbellate flower clus- ters; flowers small with very short open tube, the tepals about equal and three to five times as long as the tube; spring flowering; capsules small to medium, freely seeding. Sonoran Desert re- OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No, 130 gion in southeastern California. Arizona. Baja California, and Sonora. The leaf surface in the Deserticolae is gener- ally light colored, glaucous gray or yellowish or light green, minutely sculptured, the cuticle rather thick and rough with variable papillae. The stomata are relatively dense, ranging from 30 to 50 per mm^ on the upper leaf surface. The stomata are generally depressed and frequently so close together that the stomatal depressions are confluent, forming grooves in the cuticle, transverse to the long axis of the leaf, as in A. deserti and A. cerulata, or aligned with the leaf axis as in A. sobria sobria. The margin of the stomatal pore is generally irregular and is with- out the stomatal rim. However, among the sev- enteen taxa of this group, there are many ex- ceptions to the above generalities. The intrasectional variability is reflected in the epi- dermal morphology even as it is in the gross morphology of this complex group. For further particulars on epidermal morphology, the reader is referred to the companion study (Gentry and Sauck 1978). The Deserticolae do not display close mor- phological relation to any other Agave group. Certain other taxa, such as A. neomexicana and A. gracilipes of the Parryanae, are also small- leaved xerophytes, and the latter has an ideo- graph reflecting its short tube, much like the Deserticolae. However, the geographic position and other morphological characters of A. gra- cilipes recommend its inclusion in the Parry- anae. The Marginatae are also similar to the Deserticolae with their very short tubes and rel- atively long tepals, but they belong in the sub- genus Littaea and have other important distin- guishing characters, such as their marginate leaves and filament-clasping tepals, while their geographic distribution is distinct. Perhaps the closest relatives of the Deserti- colae are the anomalous A. iitahensis group, oc- cupying an adjacent area. Similar characters in the two groups are the short flower tubes, the short branches of the panicle, the weakly at- tached teeth, and the prolific stooling rosettes. However, other characters of flower, such as its connivent form, and of fruit and leaves demon- strate a wide divergence between the groups. The common group ancestor, assumed to have existed on general biological theory, was remote in biologic time and in physiographic if not geo- graphic space. Agave deserti deserti occupies a centric po- sition in Deserticolae, certainly in physiographic position, and interpretively in morphological re- lations. Several of the taxa, such as A. pringlei and A. deserti, are too close to separate specif- ically, while others, such as A. sobria and A. moranii. are quite distinct. The latter, along with A. avellanidens and A. gigantensis, form a dis- tinctive group to themselves, again based mainly on rosette and habit characters. They could well be treated as a subsection, as Berger 1915 and most other German taxonomists would probably have done upon cognizance of the groups. How- ever, I prefer to keep the groups simple wherever possible. The Deserticolae forms a natural phylo- genic group on both morphological and histori- cal grounds. Flower structure is quite stable or uniform throughout the Deserticolae (Figs. 5, 6). The differences or variations in the floral organs are not always critical for defining species, as I had hoped before the respective series of flowering specimens were assembled. Variation in flower structure was sometimes found to be as great within local populations as it was between pop- ulations judged to be separate species on other criteria. This is the case, for instance, with A. deserti and A. cerulata. These two complexes are difficult to separate by morphological char- acters, although respective populations appear distinct when viewed overall in the field. The chemical compositions of sapogenins appear to substantiate their specific separation (Tables 3 &4). However, there are some cases where flower size, proportions, and shapes were correlative with vegetative characters and were useful for separating species, as with the small flowers and narrow tepals of A. sobria and A. mckelveyana. Leaf form, size, habit, and other variation pat- terns, combined with geographic distribution, are the principal criteria used in separating the taxa. Key to the Species of the Deserticolae la. Plants surculose; leaves small, usually narrow, less than 10 cm wide, light glau- cous gray to yellowish; teeth frequently fragile; panicles short, mostly with only 8-15 umbels 2 lb. Plants not surculose; leaves broad, 10- 25 cm wide, green to glaucous gray; GENTRY: AGAVES OF BAJA CALIFORNIA 15 teeth firmly attached; panicles elongate with 20^0 lateral umbels 8 2a. Plants of the northern sonoran Desert; leaves generally gray to light green, broader, less acuminate 3 2b. Plants of peninsular Baja California; leaves generally yellowish to gray, nar- rower, more acuminate; teeth with a brown ring around base, or leaves larg- er, frequently cross-zoned 5 3a. Leaves larger, mostly 25-40 x 5-10 cm; teeth very feebly attached; spine decurrent for several cm and confluent with uppermost teeth; flowers 40-60 mm long; filaments inserted on base of tepals or in orifice of tube deserti (p. 15) 3b. Leaves smaller, 3-5 cm broad at mid- blade; teeth feebly or firmly attached; spine very shortly decurrent and not confluent with uppermost teeth; flowers 30-50 mm long; filaments inserted in ori- fice of tube 4 4a. Leaves 18-30 x 3-5 cm; teeth firmly attached; flowers 30^0 mm long; flow- ers yellow; tepals 3^ mm broad in mid- dle, conduplicate, strongly cucullate; ovary constricted below tube. North- western Arizona mckelveycma (p. 25) 4b. Leaves 15-25 x 3-6 cm; teeth weakly attached; flowers 40-50 mm long; flow- ers partly pink or red; tepals 6-8 mm broad in middle, plane or rounded, slightly cucullate; ovary not constricted below tube. Coastal Sonora subsiniplex (p. 27) 5a. Leaves small, mostly narrow, long-acu- minate, 3-6 cm broad (rarely 7-8 cm); yellowish to light glaucous gray; teeth relatively small, mostly 3-5 mm long, very weakly attached and ringed by a brown margin below base. Mid-penin- sular region cerulata (p. 35) 5b. Leaves larger, or at least broader, ovate to linear-lanceolate, mostly 5-12 cm broad at mid-blade, green to gray glau- cous; teeth usually large, 8-20 mm, var- iously flexed, firmly attached, not ringed by a brown basal margin 6 6a. Flower tube shallow, 3-5 mm deep; leaves mostly 40-60 cm long, sometimes conspicuously cross-zoned. Sierra de la Giganta region sohria (p. 48) 6b. Flower tube 8-12 mm deep; leaves short, usually less than 50 cm long, not cross-zoned. Outer coastal region 7 7a. Leaves short, less than 25 cm long, ovate to oblanceolate, the margin prom- inently mammillate and with long (8-15 mm) slender teeth margaritae (p. 48) 7b. Leaves 25-40 cm long, lanceolate, the margin nearly straight to undulate, with shorter, more flattened teeth vizcainoensis (p. 67) 8a. Leaves 70-120 cm long, triangular long- lanceolate, deeply guttered, the margin nearly straight; terminal spine subulate, light gray; peduncle sometimes swollen below the panicle. South and east slopes of the Sierra San Pedro Martir nioranu (p. 58) 8b. Leaves 40-70 cm long, broadly lanceo- late, plane to concave above, margin lightly to deeply sinuate; spine coarser, dark brown to dark gray, frequently sin- uous; peduncle not swollen below pan- icle 9 9a. Leaves green, broadly linear-lanceo- late, not or scarcely narrowed towards the base; panicle generally occupying one-half the shaft with 25-35 laterals; flowers yellow to orange. Central pen- insula avellanidens (p. 61) 9b. Leaves light green to glaucous gray, ovate acuminate to spatulate, conspic- uously narrowed toward the base; pan- icle in upper one-third of shaft with 18- 25 laterals; flower buds whitish or pale green, opening pale yellow. Southern peninsula gigantensis (p. 63) Agave deserti Agave deserti Engelm. ssp. deserti (Figures 7, 8. 9 (upper); Tables 2. 3) A^ave deserti Engelm. Trans. Acad. Sci. St. Louis 3;310, 370. 1875. Aguve consocicita Trel. Missouri Bot. Gard. Rep. 22:53. 1912. Medium-sized, light gray, sparingly or prolif- ically suckering rosettes mostly 30-50 cm tall, 40-60 cm in diam.; leaves variable, mostly 25- 40 x 6-8 cm, lanceolate to linear-lanceolate, scarcely narrowed above the broad clasping base, acuminate, gray glaucous to bluish glau- cous, often cross-zoned, thick, rigid, concave OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Agave desert! desert! _ Agave desert! simplex _ _ Agave desert! pr!ng!e! „ _ Agave mckelveyana Agave subs!mplex * Figure 7. Distribution of Agave deserti. A. mckelveyana. and A. suhsimplex . based on herbarium specimens listed in the Exsiccatae (Appendix). above, convex below, usually regularly armed with slender-cusped teeth, from small, the larger 2-3 mm long, to large, the larger 6-8 mm long, gray, loosely attached, mostly 15-30 mm apart; spine strong, generally 2^ cm long, light brown to grayish, openly grooved above, decurrent to the first or second tooth above; panicles gener- ally 2.5-4 m tall, on slender shafts with scarious triangular bracts 8-15 cm long and 6 to 15 short laterals with small umbels in upper V4 to V5 of shaft; flowers yellow, 40-60 mm long; ovary 22- 40 mm long with a slightly narrowed neck 4-6 mm long, greenish; tube shallow, spreading, 4- 6 mm deep, 12-15 mm wide, lined with a thick nectiferous disk; tepals equal, 14-20 mm long, 6-8 mm wide, light yellow, broadly linear. spreading at anthesis, rounded and abruptly hooked inward at apex; filaments 25-35 mm long, inserted at base of tepals; anthers 13-18 mm long, yellow; capsules ovoid to oblong or obovoid, mostly 3.5-5 cm long, 1.5-1.8 cm wide, thick walled, short-stipitate; seeds black, 4 X 5 m. Type. — Based on Emory in 1846 and Hitchcock and Palmer in 1875. collected on Rancho San Felipe, San Diego County, California. Distribution, Variability, and Relationships . — Agave deserti is a large variable complex, the limits of which are hard to define. The geograph- ic distribution is mapped in Figure 7 and docu- mented in the Exsiccatae. While the extensive population of A. deserti at the type locality along San Felipe Creek (Fig. 8) is morphologically homogenous, there are other localities showing wide variability in leaf form. One of these lo- calities is Pinyon Flats by the northwest slopes of Santa Rosa Mountain along Route 74 in southern California. Some of the variability appears attributable to hybridization, such as those populations on Pin- yon Flats and in the San Matias Pass in Baja California. The population from the latter area is discussed below under A. deserti pringlei. Variability in the Pinyon Flats population ap- pears to be a relic condition of geographically past introgression between two disparate geno- types, which are not recognizable today. Cave ( 1964: 166) has reported a collection (Hutchinson 710) from Baja California with chromosomes /; = 59. This is a polyploid condition, as the ba- sic number \n Agave is n = 30. The genetic vari- ability in the Agave complex indicates a high potential for species evolution, as examples are the populations of A. deserti pringlei and A. cer- ulata nelsonii in our present Recent Period. Ivan Johnston (1924) included several of Tre- lease's taxa as synonyms under A. deserti, viz. A. pringlei, A. nelsonii, A. consociata , and A. dentiens, thus forming a kind of superspecies category. After considerable study and vacilla- tion, I am using Trelease's name, A. pringlei, as subspecies to designate scattered populations bordering the main complex on the south and west of the deserti area. Another group in and adjacent to Arizona is segregated as subspecies simplex, the name referring to its more simple habit. A. cerulata Trel. is closely related, but ap- GENTRY: AGAVES OF BAJA CALIFORNIA 17 Figure 8. Agave deserti near the type locality along the San Felipe arroyo, San Diego County, California. (Upper) The desert habitat; (lower) detail of drought-shrunken leaves. Otherwise, these are robust plants in deep sandy soil. 18 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Table 3. Sapogenin Content in Af-ave deserti. Collated from Wall (1954) and Wall et al. (1954a. 1954b, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin; git. = gitogenin: man. = manogenin; tig. = tigogenin: w.pl. = whole plant; fl. = flower; dead = leaf of dead plant. Month Plant Sapogenins Coll. 9c 7c % 9c % no. Source locality coll. part total hec. git. man. tig- Agave deserti deserti California 10041 San Felipe Narrows Dec. leaf 0 10041 San Felipe Narrows Dec. stem 0 10044 San Felipe Narrows Dec. w.pl. 0.1 50 50 10051 Pinyon Flats Jan. fl. 0 10051 Pinyon Flats Jan. fruit 0.6 X 10051 Pinyon Flats Jan. seed 0 10051 Pinyon Flats Jan. fruit 0.6 70 5 5 20 11650 Pinyon Flats Apr. leaf 0 11652 Pinyon Flats Apr. leaf 0.75 20 14 66 12436 Pinyon Flats Agave deserti pringlei Baja California Dec. leaf (dead) 0.6 X X X 10287 S of San Pedro Martir Mar. leaf 0 10287 S of San Pedro Martir Agave deserti simplex Arizona Mar. stem 0 9947 Harquahala Ml. Nov. leaf tr. 9947 Harquahala Mt. Nov. stem 0 pears to be a separate specific complex, difficult to separate morphologically. Reasons and char- acters for separating it from the deserti complex are given under A. ceridata below. A. cerulata does not belong with A. sobria Edge, where Johnston placed it. The following outline sepa- rates three main A. deserti geographic popula- tions, none of which is in all individuals mor- phologically distinct. Key to Subspecies oi Agave deserti la Leaves mostly 25-40 cm long. 4-7 times longer than broad, moderately acumi- nate, the margins usually straight; teeth weakly attached; spine decurrent as a corneous margin only to the upper first or second pair of teeth. Tube 3-8 mm deep lb. Leaves mostly 40-70 cm long, 8-12 times longer than broad, long-acumi- nate, the margin straight; teeth firmly attached; spine conspicuously decur- rent in a corneous margin frequently to the mid-blade or even below. Tube 5-8 mm deep. San Matias Pass and vicinity pringlei 2a. Rosettes copiously surculose, forming large clones; flower tube 3-5 mm deep; filaments inserted on base of tepals. Western side of the Gulf of California deserti 2b. Rosettes generally single, rarely with 1- 3 offsets; flower tube 5-10 mm deep; fil- aments inserted in orifice of tube below tepal bases. NE of the Gulf of California simplex Uses and Chemistry . — Agave deserti is among the more edible of the agaves. Castetter et al. (1938) wrote an excellent account of uses of agave by the Indians of the American South- west. Their map of "mescal finds"" (ibid.:37). including "mescal pits,"" shows that A. deserti was eaten and its fibers used throughout the de- serti area. "Mescal pits"" are very common and have been traced as far south as the latitude of Punta San Fermin in the Gulf of California (ibid.:60). This is in Baja California at the south- ern margin of the known distribution of A. de- serti (e.g.. Gentry & McGill 23286 on the San GENTRY: AGAVES OF BAJA CALIFORNIA 19 Figure 9. {Upper) Af^ave deserti deserti on the desert plain southeast ol ( ciio Hmagd H.ii'i Calitoinia. The high small panicles are characteristic in this region. (Lower) A. deserti priiiglei in San Matias Pass with green and pale-glaucous forms growing side by side. Pedro Martir bajada). Barrows (1967:59) stated that the Coahuilan Indians of southern Califor- nia made much use of A. deserti for food and fiber. They called the plant "a-mul," sections of the flowering stalk "u-a-sil/' the leaves "ya- mil," and the yellow blossoms "amul-sal-em," all of which were cooked in various ways and eaten. Bairows regarded the agaves as one of 20 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 the principal plant resources of southwestern Indian tribes. Other tribes known to use A. deserti. as well as other species in their respective areas, are the Pimas and Papagos of both Arizona and Sonora. the Yumans. Kamias, Chemehuevis, and Ya- vapais along the Rio Colorado, and the Diguefios and Cocopahs of northern Baja California. Other adjacent tribes, such as the Utes and Apaches, received cooked A. deserti in trade. The earliest missionaries in the 17th century reported the uses of agave in the deserti area, for example. Padre Kino (Bolton 1919). Other early reports, such as that of the German missionary Baegert (Rau 1864). who was stationed near the 25th par- allel on the western slope of the Sierra de la Giganta in the 1770"s, and Barco (1973), who was stationed at San Javier between 1738 and 1768, corroborate the widespread use of agaves in Baja California. The Indians of the peninsula were hunters and gatherers; they did not prac- tice any agriculture, and the wild agaves were of major importance for their survival in this aridly inhospitable land. The absence of mescal pits south of Sierra San Pedro Martir does not mean that agaves were not eaten south of this latitude. The account of Miguel del Barco (1973:123). tells us why there are no mescal pits southward. It just was not the custom of the Cochimi to dig cooking pits. They roasted their mescal upon the ground "en forma de monton." The Cochimi ranged through a large central section of the peninsula and spoke a different language than the northern Digger Indians. The Cochimi guides on Link's expedi- tion of 1766 (Burrus 1966) found they could not communicate with the San Pedro Martir people. The southern end of this sierra marks a division margin between the two language-and-cultural groups; so also do the southern limits of mescal pits. Agave deserti is important to wildlife; birds visit the flowering stalks for nectar and insects. Small rodents live about the plants and are pro- tected by the armed rosettes. The pack rats, Neotoma, make nests among the clumps of ro- settes, gnaw through the leaves, and eat the flowers and perhaps the seeds. Wild bighorn sheep, like cattle, eat the new-flowering shoots. As a wildlife resource growing in our most arid deserts. Agave deserti deserves conservation and perhaps should be established on desert mountains where it is now lacking. Table 3 gives the sapogenin content found in Agave deserti by chemists of the Agricultural Research Service in Philadelphia, Pennsylvania (Wall 1954; Wall et al. 1954a, 1954b, 1955, 1957). Hecogenin is the leading steroid. The steroid content is erratic to absent in leaves, absent in the edible stems, and appears to run about 0.6 to \7c in the fruits. Agave deserti Engelm. ssp. pringlei (Engelm. ex Baker) Gentry, stat. nov. (Figures 7. 9 (lower). 10; Tables 2. 3) Agave pringlei Engelm. ex Baker. Handbook .'\marillid. 182. 1888. Green or whitish, cespitose, rather strict ro- settes, offsetting closely from root crown, 4-7 dm tall, 5-8 dm wide; leaves narrowly triangular lanceolate, very long acuminate, mostly 40-70x 5-7 cm, thick and widest at base, deeply cres- centic in cross-section, guttered and thinner above, green to yellowish green or light glaucous gray; spine 3-4 cm long, aciculate, reddish brown to light gray, usually narrowly grooved for short distance at base, decurrent as a cor- neous margin to upper teeth or to the mid-blade; teeth rather regularly spaced 1-2 cm apart, mostly 5-10 mm long, slender and little curved, more rarely smaller or more broadly flattened and irregularly flexed; panicle 3-6 m tall with 10-15 lateral branches, very narrow or more am- ple with longer laterals and larger umbels; flow- ers 40-60 mm long, yellow; ovary 20-35 mm long, fusiform, roundly angulate, with furrowed neck; tube ample, 5-8 mm deep, bulging, strong- ly furrowed from tepal sinuses; tepals nearly equal, 15-20 x 5-7 mm, spreading, linear, rounded over finely cuculate apex, the inner wider than outer and with broad low keel; sta- mens small, filaments inserted on rim of nectary in orifice of tube; capsules 3.5-5.5 x 1.2-1.5 cm, mostly oblong, beaked, short-stipitate; seeds as for species. Type. — "Central mountains of Lower California, alt. 6000 ft.. Orcutt! Described from a dried specimen sent to Kew by Mr. C. G. Pringle." Probably from the Sierra Juarez in 1882, when Orcutt collected there. The type, perforce, is only nomenclatorial in scope. The type specimens are but a pitifully small expression of the peninsular highland pop- ulations. In the description above, I have drawn this taxon's outline from the listed specimens (see Exsiccatae) to include the apparent in- GENTRY: AGAVES OF BAJA CALIFORNIA 21 Figure 10. Agave descni pnngUi m San Matia^ Pass. Baja California. {Upper) Detail of ro,sette and cross section of leaf bases where specimens were cut. (Lower) Rosette of a green clone in flower. April 1963. 22 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 trogressions with A. deserti desert i and with A. moranii as well. Without including some mor- phological characters of A. moranii, A. deserti pringlei would appear as a green, highland eco- type of v4. deserti. The variable population in and about San Ma- tias Pass shows relation to A. deserti deserti in the tall, very narrow panicle and the shorter white glaucous leaf, which are present in differ- ing combinations in some of the clones. Relation to A. moranii appears in the denser broader pan- icles and yellowish-green leaves of other clones, as well as leaf elongation in both green and whit- ish forms. Other specimens on the western slopes of the Juarez-Martir mountain axis are very similar to A. deserti deserti in leaf size and form, but have atypically green leaves. The type collections of A. pringlei appear to be of this green form. The population in and about San Matias Pass appears to be a "hybrid swarm," and individual genotypes here, as well as else- where, reflect genetic factors of one or the other "parent" in varying degree and combinations. Subspecies pringlei, with its surculose habit, slender panicles, and smaller size is more closely related to deserti than it is to the larger, nonsurculose, and diffusely paniculate moranii. The deeper tube with its deeper insertion of fil- aments is perhaps the most unique character distinguishing A. deserti pringlei from its neigh- bors, but incidence of this character appears to be sporadic, judging from the limited flower specimens available. Agave deserti ssp. simplex Gentry, ssp. nov. (Figures 7, 11. 12: Tables 2. 3) Small- to medium-sized, mostly single, light green to light glaucous gray, compact, short- stemmed rosettes; leaves mostly 25-40 x 6.5- 10 cm, rarely to 50 cm, lanceolate, firm, rigid, mildly concave above, with undulate to crenate margins, broadest in mid-blade; spine stout, subulate, 3^ cm long, broadly grooved above, dark brown to light gray, decurrent for several centimeters to upper second or third pair of teeth; teeth in mid-blade mostly 5-8 mm long, 1-3 cm apart, brown to pruinose gray, frequent- ly brown and gray-ringed about base, variously curved or reflexed; panicles 4-6 m tall with 8 to 15 short umbellate branches in upper third of shaft, the peduncles stout, large reflexed bracts below rapidly decreasing in size upward and persisting erect, scarious; flower clusters small, congested, pale yellow to ferruginous in bud; flowers 40-60 mm long, yellow with pale-green ovary; ovary 20-30 mm long, fusiform, with constricted grooved neck 4-6 mm long; tube short. 5-8 mm long, open funnelform, grooved, angular; tepals subequal, 15-20 x 4-5 mm, erect to spreading, rather thin, linear, condupli- cate wilting and incurved, apiculate-cucullate, the outer slightly longer, flat to rounded on back, the inner with broad low keel; filaments 30^2 mm long, unequally inserted below tepal bases high in tube, slender, elliptic in cross-section, pale yellow; anthers 15-21 mm long, yellow, regular, centric; capsules 3.5^.2 x 1.5-2 cm, oblong to obovoid, stipitate, rounded to rostrate at apex, rather thin-walled; seeds 5-6 x 4-4.5 mm, lunate to lacrimiform, dull or shiny black, the margin with a raised, irregularly fluted wing. Type. — Gentry 23404. N slope of Harquahala Mountain. 12 miles [ca. 19 km] W of Aguila. Yuma Co., Arizona. 12 June 1974: deposited in US, isotypes in DES. MEXU. ARIZ. Planta parva vel media plerumque simplex tandem paucisurculosa; folds plerumque 25-40 cm longis 6-10 cm latis ad medium raro ad 50 cm longis, linearibus vel lanceolatis glaucoviri- dibus rigentibus undulato-marginatis; spinis marginalibus ad medium foliorium plerumque 5-8 mm longis, 1-3 cm separatis, debilibus flex- uosis acicularibus bruneis vel albicantibus; spina terminali 3-4 cm longa subulata castanea vel grisea supra late canaliculata: inflorescentia paniculata angusta 4-6 m alta cum scapo, fer- enti 8-15 ramos umbelliformes parvos; floribus 40-60 mm longis viridi-luteis; ovario 20-30 mm longofusiformi apice constricto sulcato; tubo 5- 10 mm longo 10-13 mm lato infundibuliformi; segmentis subequalibus luteis 15-20 mm longis 3.5-5 mm latis, linearibus involutis apiculati-cu- cullatis; filamentis 35-40 mm longis ad apice tubi inaequaliter insertis; antheris 15-20 mm longis, luteis; capsulis 3.5-4.2 cm longis 1.5-2 cm latis, oblongis stipitatis bruneis; seminibus 5-6 mm longis 4^.5 mm latis lunatis vel lacrim- iformibus margine alatis undulatis. Relationships and Comparisons . — Agave deser- ti simplex is distinguished from A. deserti deserti by its predominantly solitary habit, deeper flow- er tube, tepals being only ca. 2-3 times longer than the tube (vs. 4-5 times longer), filaments GENTRY: AGAVES OF BAJA CALIFORNIA 23 Figure II. Agave deserti ssp. simplex drawn from type, Genliy 23404: leaf, flower cluster, capsule xi/2; flower section X Wi; seed x2. A. deserti ssp. deserti, flower section from Gentry' 19940. 24 OCCASIONAL PAPERS OF THF CALIFORNIA ACADEMY OF SCIENCES. No. LW Figure 12. Agave deserti simplex in the Silver Bell Mountains, Pima County, Arizona. The plants are frequently depau- perate, resemble ,4. mckeheyana with small flowers, 33-46 mm long, but the tube is characteristically deep. 5-7 mm. being inserted in the tube (vs. inserted on the base of the tepals), and the disjunct distributions of the two taxa (Fig. 7). It is not possible to separate the two taxa on leaf specimens alone, even when accompanied by photographs. How- ever, if labels state the origin of the specimen and whether the population from which it was selected consists of cespitose plants or generally of singles, identification of leaf specimens be- comes relatively certain. Large plants frequently offset, especially at maturity, but the offsets may wither and die. Another close relative of A. deserti simplex is A. mckelveyana, which occupies higher mon- tane elevations northeast of the simplex area. As in A. deserti simplex, the filaments of A. mckel- veyana are inserted in the orifice of the tube, but the latter is a smaller plant with small narrow leaves, slender wandlike inflorescence, and small flowers. It is possible that these two taxa will be found in contact, and if so, it will be interesting to see if there is introgression. In Arizona the scattered populations of sim- plex rosettes sucker much more sparingly than the California colonies, and the large cespitose California clones are absent. In the Lechuguilla Desert of southwestern Arizona, one of our most arid regions, the subspecies is represented by small drought-depauperized, scattered indi- viduals, with leaves commonly only 20-25 cm in length. Other than for their small size, the characters of the plants are quite like those of GENTRY: AGAVES OF BAJA CALIFORNIA 25 Figure 13. iM). Agave mckelveycma from type: leaf, flower cluster, and capsules (D).A. deserti flower section slightly enlarged. '4; flower section slightly enlarged. the species — rather broad, gray, cross-banded, thick rigid leaves and slender small panicles. However, also found in the region is a long- leaved robust form in the Mohawk Mountains {Wiggins 8643), which I have not seen in habitat. The native uses of Agave deserti simplex are essentially what has been reported above for the species. Agave mckelveyana (Figures 7. 13-15: Table 2) Agave mckelveyana Gentry. Cactus Succulent J. (U.S.) 42:225. 1970. Small, single or suckering, rather few-leaved rosettes 20^0 cm tall; leaves 20-35 x 3-5 cm, linear or lanceolate and broadest in the middle, light glaucous green or yellowish green, firmly spreading, the margin nearly straight or undulate with low teats; spine 1.5-4.0 cm long, shortly decurrent, subulate, rounded except for shallow groove above at base, castaneous to gray; teeth small to medium, the larger at mid-blade 4-8 mm long, mostly 1-3 cm apart and downflexed, gray- ish with reddish tips, rather friable; panicle small, narrow 2-3 m tall with 10-19 laterals in 26 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 , >rfC^ ^'^ ^ ■ * ■ ■'•^^'■^':\- ^- '■'<^''% %^^m-^M^r ■ ^ ■■■ ■ '" • •«>•■' ' i - »^ Figure 14. Agave mckelveyami near Sitgreave Pass in the Black Mountains. June l%6, with highland dispersed desert shrub at ca. 3.500 ft ( 1 ,070 m) altitude, and Linnea Gentry. upper half of shaft with small compact umbels; pedicels short-bracteolate; flowers small. 30^0 mm long, the perianth and stamens openly spreading, yellow; ovary light green, 16-22 mm long including constricted neck faintly grooved below tepal sinuses, fusiform or cylindric: tube shallow, open, 3.0-4.5 mm deep, 8-9 mm broad; tepals 12-13 x 3-4 mm spreading, thin, with open sinuses, linear conduplicate, the outer lon- ger and narrower than the round-keeled inner. both abruptly hooded at tips; filaments 25-30 mm long, inserted in orifice of tube, flattened towards base; anthers 12-16 mm long, yellow; capsules 30^5 x 10-14 mm, oblong, narrowly stipitate. obtuse to apiculate, thin-walled, striate; seeds 5.0-6.5 x 4.0^.5 mm. obovate from apical hilum or half-moon, the faces ru- gose, completely margined with a low wing. Type. — Gentry 21979. Sitgreave Pass in Black Mountains, ca. 4 miles [ca. 6 km] NE of Oatman. Arizona. 26 June 1%6, GENTRY: AGAVES OF BAJA CALIFORNIA 27 US. It occurs here as a small scattered population on rocky volcanic slopes between 3.000 and 4.000 ft (900 and 1.200 m) elevations. Agove mckelveyana occupies a central part of western Arizona (Fig. 7). Its habitat is with the chaparral and juniper associations on rocky slopes between 3,000 and 6,000 ft (ca. 900 and 1,800 m) elevations. I have not seen it growing with other Agave species, but McKelvey col- lected it in contiguous numbers with A. utah- ensis in the Aquarius Mountains and with A. parryi var. couesii in the Juniper Mountains. It appears to occupy an ecologic niche of its own, quite distinct from that of its near relative, A. deserti simplex, which is confined generally to the Sonoran Desert at lower elevations. The habitat of A. mckelveyana is better watered and is cooler than that of A. deserti. Marcus Jones collected a cutting of inflores- cence in 1930 near Oatman (Jones 25167) and mixed it with his specimens which he described as A. utahensis var. discreta. Trelease annotat- ed some of McKelvey's earlier collections as A. aquariensis , but the name was never published. The slender panicles, small flowers with shallow tubes and spreading tepals place it in the section Deserticolae of Trelease. Its nearest relatives appear to be A. deserti and A. subsimplex Trel. The small leaves of A. mckelveyana with vari- ably flexed teeth resemble those of A. subsim- plex, from which it can hardly be separated ex- cept by the flower differences. The flowers of A. mckelveyana differ from these two, and other relatives, in smaller size, in the narrow spread- ing, thin, linear, conduplicate tepals with sharp- ly inflexed cucullate tips, and small ovary with constricted neck. Some observers of A. mckelveyana have re- garded it as a depauperate form of A. deserti. The plants in some of the drought-inhibited pop- ulations of A. deserti simplex, as in the Lechu- guilla Desert, are about the size of A. mckelvey- ana, but they still have the essential, if subtle, characters of A. deserti. The latter are generally more robust than A. mckelveyana. The pattern of variation in A. mckelveyana is phenotypically homogenous and does not intermesh with the variability of the polymorphic A. deserti. A leaf series of A. mckelveyana is shown in Fig. 15; flowers are shown in Fig. 13. I find no inter- grading between the two groups and believe them to be biologically distinct species. They probably separated from a common ancestor long ago. There is nothing particularly remarkable about this small Agave mckelveyana, and its habit of growing among shrubbery keeps it ob- scure. The flowering season appears to be during May, June, and July. However, flowering time will vary from year to year and according to elevation. The life span is not known. Mature flowering and fruiting rosettes observed had only 30-40 leaves, but as the rate of leaf growth is unknown, the relatively few leaves do not necessarily indicate a short-lived rosette. Suck- ering offsets were observed on other plants on the Black Mountains and again on Hualapai Mountain. In the Hualapai Mountains, Abert's tree squirrel, Sciurus abertii, has been observed gathering green capsules (word of Rodney En- gard). In one instance the squirrel was observed cutting and carrying the fruits down. Other fruit- denuded panicles in the same locality were at- tributed to the work of this animal. Agave subsimplex (Figures 7, 16. 17; Table 2) Agave subsimplex Trel. Missouri Bot. Gard. Rep. 22;60. 1912. Small, single or cespitose, glaucous, colorful, low-spreading rosettes, 20-35 cm tall, 50-70 cm broad; leaves variable, 12-35 x 3-5 cm, lanceo- late to ovate, long-acuminate to short-acumi- nate, but little narrowed towards the base, thick, rigid, rounded on back, hollowed in inner face, gray glaucous or light yellow-green, or some- times purple-tinged, the margin nearly straight or strongly teated; teeth variable, friable, the larger 3-15 mm long, straight or variously flexed, rarely bicuspid, brown or more often yel- lowish gray; spine subulate, 2-4 cm long, not or but little decurrent, frequently sinuous, shallow- ly grooved above, glaucous gray; panicle slen- der, narrow, 2-3.5 m tall, with 5-8 short laterals; flowers in small umbels, yellow to pink 40^5 mm long; ovary about 25 mm long with an un- constricted long (5 mm) neck; tube shallow, spreading, 3^ mm deep, 10 mm wide; tepals 12-15 X 6-7 mm, equal, ascending, elliptic, plane, widest at the middle, apiculate and scarcely hooded; filaments 25-28 mm long, round in cross section, inserted below base of tepals 3 mm above bottom of tube; anthers 13- 28 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 ^ S a. o GENTRY: AGAVES OF BAJA CALIFORNIA 29 Plate 2. (Top) Agave gigantensis on the volcanic top of Sierra de las Palmas. Photograph by the author, April 1952. (Bottom) Agave cerulata Jentiens. Isla San Esteban. Photograph by Reiil V. Moran. 17 April 1975. 30 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. I 30 FiGLRL l.V AMcne- nukelveyana in the Black Mouniains. [Upper) Mature leaves from 6 plants, the leaf at right depauperate trom a poor site, the leaf at left from the rosette shown below. GENTRY: AGAVES OF BAJA CALIFORNIA 31 Figure 16. Agave subsimplex: larger leaf drawn from Gentry 10217. the smaller from Gentry 10221; flowers from Felger & Bezy 14171 . section x 1, cluster x-Zs; capsules x Wi, and seeds x 1% from Gentry 10221. 15 mm long, centrically attached; capsules 40 X 15 mm to 35 x 10 mm, variable, oblong, sometimes narrowly so, light glaucous (Gentry 10221), bluntly apiculate, narrowly or broadly stipitate, the valves thick and striate-nerved; seeds (10221) mostly 4.5 x 3 mm, sooty black, roughly lunate, hilum notch narrow and the op- posite corner frequently apiculate, edges rimmed with a sharp, winglike flange. Type .—SONORA: Seal Island, just off Tiburon Island. 13 Apr. \9\\.Rose 1681 1 .[JS. Known only from coastal Sonora. where it is thinly scattered in small colonies on the outwash slopes of the granitic and volcanic mountains and the adjacent islands. Reid Moran in 1966 found Agave subsimplex common on Turner's Island, or Isla Datil, as it is also called, just south of Tiburon Island in the Gulf. Moran's (1967) account provides a short botanic history of the plant, a diagnostic de- scription, and his field observations. This small xerophytic agave is closely related to A. deserti and A. cerulata Trel.. as expressed by their common characters of small variable leaves, nearly tubeless flowers, and narrow 32 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. HO -2; o GENTRY; AGAVES OF BAJA CALIFORNIA 33 F 4 :fe. *'*.. Plate 4. (lo;?) A^tjif aurea 19 km southeast of La Paz. Photograph by Reid V. Moian. 25 January 1959. (Bottom) Agave aurea "southwest end of Mesa de San Geronimo, northerly from Rancho Viejo (on road from Loreto to San Javier). Alt. ca. 1,100 m. Lat. 20°58'N, long. 1 1 l°32.5-34'W."" Photograph and quotes by Annetta Carter, 8 May 1%6. 34 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 17. Agave subsimplex along the arid Sonoran coast. Feb. 1951. small panicles, which characterize the group Deserticolae. I think the characters outlined in the key to species should be sufficient to distin- guish A. subsimplex from A. deserti. but depau- perate plants of the latter in the Lechuguilla Desert of Arizona resemble A. subsimplex. The two species are not known to cohabit, but it is possible that they may be found together on some unexplored mountains in northwestern So- nora. such as the coastal Sierra del Alamo. Agave subsimplex may be more closely related to the polymorphic A. cerulata of Baja Califor- nia, as indicated by the small, variable leaves and the narrow, oblong, wa.xy capsules. How- ever, the Sonoran plants have more spreading rosettes, gray not yellow, with thicker, wider. GENTRY: AGAVES OF BAJA CALIFORNIA 35 less acuminate leaves, and the filaments are in- serted below the base of the tepals. In A. sub- simplex the nectarious innerliner of the tube, from the rim of which the filaments ascend, does not completely fill the tube, as it does in A. cer- iihita. The flowers of A. subsimplex are smaller than those of A. cerulota, 40-45 mm vs. 45-60 mm. The pink to red color frequently occurring in the pistil, filaments, and corolla (see notes of collectors in Exsiccatae) is distinctive of A. sub- simplex. Such color is not known in the flowers of A. deserti and A. cerulata. The Seri Indians, a maritime hunting-and- gathering people of the Sonoran coast, called this agave "ahmmo," as nearly as I could render the sound, and stated that it was gathered for cooking and eating. Whiting noted the Seri name as "den; kl." Small amounts of sapogenin (0.07- 0.14 percent) were found in the leaves. Felger and Moser (1970) render the name phonetically as "?aamXW." They reported that the pit- baked stems are eaten in several ways: cubed and cooked with sea turtle, or made into flat cakes or patties, which may be stored for later use or taken on long trips. These cakes could also be soaked in water and consumed as a sweet drink. The various agave viands all had Seri names and indicate an habitual, important resource for survival by a people living on the outer edge of human environment. Agave cerulata Agave cerulata Trel., ssp. cerulata (Figures 18-21: Tables 2. 4) Agave cerulata Trel. Missouri Bot. Gard. Rep. 22:55. 1912. Small yellow to pale green, rarely light glau- cous gray, abundantly surculose, few-leaved ro- settes 25-50 cm tall; leaves mostly 25-50 x 4- 7 cm, long acuminate, narrowly lanceolate to triangular-lanceolate, yellow to light green, sometimes cross-zoned, the margins nearly straight to mildly undulate with teated teeth; teeth small, 1-4 mm long, irregularly spaced, sometimes lacking through much of the blade, grayish brown, bordered with a brown ring at base, weakly attached; spine 3-6 cm long, acic- ular, light gray to dark gray, decurrent only to uppermost teeth or less; panicle slender narrow, 2-3.5 m tall, with mostly 6-12 small lateral um- bels, the whole white waxy glaucous in bud stage; bracts small scarious triangular; flowers waxy white in bud, opening pale yellow, mostly 45-60 mm long; ovary 22-32 mm long, fusiform, narrowly tapered toward base; tube 3x11 mm to 5 X 14 mm, broadly funnelform or discoid with thick nectary and prominent bulges oppo- site filament insertions; tepals 16-22 mm long, ascending to spreading, equal, elliptic, the inner wide with low broad keel and broadly over- lapped at base by outer; filaments inserted at base of tepals on rim of nectary, variable in length, usually incurved at anthesis, 30^0 mm long; anthers 15-20 mm long, centric to excen- tric; capsules 3-5 x 1.2-1.3 cm, narrowly ob- long, waxy light gray, narrowly stipitate, bluntly apiculate; seed 5x3 mm, lunate, sooty black, the hilar notch open or obscure, the marginal wings pronounced on both sides around the cur- vature. Type. — Nelson & Goldman 7180. Calmallf, Baja California, 29 Sept. 1905. US. Characteristics and Habitat . — Most plants of A. cerulata are characterized by slender, yellow, long-acuminate, lanceolate leaves with brown eyelets ringing the weakly attached, moderate to small teeth, small narrow panicles whitened from peduncle to capsules with a waxy bloom, and light-yellow spreading tepals. A. cerulata has been confused with its near relative A. de- serti, but the latter is more robust, the leaves light gray-green rather than yellow-green, 4-7 times longer than broad (vs. 5-12 times longer than broad). A. deserti generally lacks the dis- tinctive brown eyelets ringing the teeth and the capsules are generally broader and, although glaucous, lack the waxy white bloom. The geo- graphic distributions of the two species are dis- tinct (cf. Figs. 7 and 21), although they may meet on the peninsula south of Sierra San Pedro Mar- tir in an area I have not explored. Agave cerulata is primarily a product of the upland maritime environment where frequent fogs and ocean breezes temper the desert cli- mate, while A. deserti is a product of the hot arid continental desert with high insolation and more extreme circadian temperature changes. Figures 9 and 20 show the two species in their characteristic respective habitats. There are sig- nificant chemical differences between these two species, as may be inferred from Tables 3 and 4. Chemistry. — The data in Table 4 on the sapo- genin contents of Agave cerulata are the most detailed for any Agave species and is assembled 36 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Table 4. Sapogenin content in Agave ceruUito. Collated tVoni Wall (1954) and Wall et al. (1954a, 1954h, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin: git. = gitogenin; man. = manogenin; tig. = tigogenin; fl. = flower; dead = leaf of dead plant. Baja California Month Plant Sapogenins Coll. ' r % % % % no. source locality coll. part total hec. git. man. tig- Agave cerulata ccrulatci 10346 Calmalli May leaf (white) 1.2 25 68 7 10346 Calmalli May leaf (yellow) 1.1 65 35 10346 Calmallf Dec. leaf (white) 1.1 X X X X 10346 Calmalli Dec. leaf (yellow) 0.5 X X X 10346 Calmalli Dec. leaf (dead) 1.2 X X 11322 Laguna Chapala Apr. leaf 1.1 20 45 X 11322 Laguna Chapala Dec. leaf 1.0 X X X X 11322 Laguna Chapala Dec. leaf (dead) 0.5 X X 11322 Laguna Chapala Dec leaf 0.4 50 50 11322 Laguna Chapala May leaf- 1.2 35 65 11324 Laguna Chapala Apr. leaf 0.1 X X 11188 Laguna Chapala Sep. leaf 0.8 69 31 11740 Sierra San Luis Apr. leaf 0.6 80 20 11741 Sierra San Luis Apr. leaf 0.72 60 40 11744 Sierra San Luis Apr. leaf 0.9 7 93 11746 Sierra San Luis Apr. leaf 0.6 20 70 10 11919 Calmalh May leaf 0.6 40 60 11926 Calmalli May leaf 0.6 45 55 11924 Calmalli May leaf 0.8 55 25 20 11962 Laguna Chapala May leaf 0.1 100 11962 Laguna Chapala Dec. leaf 3.4 X X X X 11953 E of Punta Prieta Agave cerulata nelsonii May leaf 0.4 30 70 10370 San Fernando Sept. leaf 1.2 100 10370 San Fernando Sept. stem 0.2 70 15 10370 San Fernando Apr. leaf 0.7 5 95 10370 San Fernando Apr. leaf 0.25 100 10370 San Fernando ec. leaf 1.3 100 10370 San Fernando Dec. stem 0.6 100 10370 San Fernando Dec. leaf (dead) 1.0 X X X X 10370 San Fernando May leaf 1.1 X X X X 11160 San Fernando Sept. leaf 0.77 75 25 11160 San Fernando Apr. leaf 0.94 70 30 11162 San Fernando Sept. leaf 1.0 90 10 11162 San Fernando Apr. leaf 0.6 50 30 20 11164 San Fernando Apr. leaf 0.5 75 25 11170 San Fernando Dec. leaf 2 2 X X X 11665 San Fernando Apr. leaf 0.3 50 50 11665 San Fernando Dec. leaf 2.5 X X X 11665 San Fernando Dec. leaf 1.3 X X X 11666 San Fernando Dec. leaf 0.5 55 11669 San Fernando Agave cerulata subcerulala Dec. fl. 0.25 60 10330 San Ignacio Apr. leaf 0.35 64 8 11 17 10330 San Ignacio May leaf 0.3 45 30 25 10330 San Ignacio Dec. leaf 1.5 X X X X 11892 Isia San Marcos May leaf 0.7 X X 12393 E of San Ignacio Dec. leaf (dead) 2.0 X X GENTRY: AGAVES OF BAJA CALIFORNIA 37 Figure 18. Agave cerulata spp. cerulata: C. drawn from topotypic Gentry 23185, 11924; leaves x'/i, flower cluster xVs, capsules x'/2, one unusually elongate; flower section x I'/i. Smaller leaf shows a recurrent, partly toothless form. (/V) Flower section of A. cerulata ssp. nelsonii. xl'/2. 38 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Plate 5. (Top) Agave shawii. Inflorescence of plant photographed about 16 km southeast of EI Rosario, Baja California. Photograph by E. S. Ross, 17 November 1977. (Bottom) Agave shawii with Simmondsia chinensis along the Pacific shore north of Ensenada, spring 1952. Many such stands have been cleared away since that date. Photograph by the author. GENTRY: AGAVES OF BAJA CALIFORNIA 39 - 8 I a y d 40 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Figure 19. {Upper) Agave cerulata cenilata near Laguna Seca Chapala. The dense clusters of rosettes with long acuminate leaves are characteristic. (Lower) A. cerulata cerulata east of Punta Prieta along road to Bahia de Los Angeles, May 1952; a robust, almost white-glaucous form with cross-zoned leaves. GENTRY: AGAVES OF BAJA CALIFORNIA 41 III .-. - {Upper) Fairy ring of Aijuvc nrnUiui ccrnlata on calcaicoiis pchlilc pavement in northwest corner of San Andres Valley, spring, 1973. Bruce Gentry photograph. (Lower) A. cerulata ceruluia on roiling plain near Laguna Seca Chapala. May 1952. 42 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 116* 114* 112* 110* \ 4 I > 34* .^ 32 ^ '^ ^ s s \^ N \ 30 n ^^t>. \ '^ 1 H^i ^ \ 28 K T 1 ?fi Agave cerulata cerulata • 1, t Agave cerulata subcerulata • Agave cerulata dentiens ^ _ _ _i> t^ 1^ \ ^ 1 1 1 1 1 r Figure 2\. Distribution of At'^iv ceriiUita and its subspe- cies based on herbarium specimens, see Appendix — Exsicca- from U.S.D.A. sources (Wall 1954; Wall et al. 1954a, 1954b, 1955, 1957). Of the four genins present, hecogenin and manogenin are the more prevalent. Hecogenin occurs in 89%, manogenin in 82%. tigogenin in 24% of the 46 plant samples. These four compounds shift about quite at ran- dom, as there is little apparent correlation with time of year, locality, plant part, or stage of plant maturity. The three cases in the table of "leaf, dead"" represent dry leaves from rosettes that had flowered and died. However, it is ap- parent that total sapogenin content is higher in December than during other months. The aver- age content of the 13 leaf samples of December is 1.5%, or about double the average content of 0.66% of the 28 other seasonal leaf samples. The average sapogenin content oi Agave cer- ulata cerulata is 0.87%, of A. cerulata nelsonii is 0.96%, and of A. cerulata subcerulata is 0.98%. This is nearly 1% and contrasts strongly with 0.25% for A. deserti. Although samples of the latter are few, they show that there are chemical and physiological differences between A. deserti and A. cerulata as well as morpho- logical and geographic ones. Populations. — The populations o^ Agave ceru- lata are vast, extending through northern and central Baja California from just above latitude 30°N to ca. 27°N (Fig. 21). The clustering clones must number over a million and the individual rosettes many millions more — a wild species population probably exceeded only by Agave lechuguilla of the northern desert region of Mex- ico. A. cerulata is polymorphic, showing several forms at many localities. An example of this was recorded on 17 May 1952 near Calmalli, the type locality. The forms noted here were itemized as follows, with Gentry collection numbers in pa- rentheses: A. Rosettes large, leaves 45-60 cm long; fig. stalk to 6-7 m (11919). B. Size medium, glaucous-yellow leaves (11921). C. Size small, leaves thick (11922). D. White glaucous with slender narrow leaves (11923). E. Thick-headed glaucous rosettes with part of the margins toothless (11924). F. Short, thick-leaved, yellow-green with nu- merous proximal teeth (1 1926). Corresponding forms as well as other striking variants were noted in other localities. The ex- tensive collections and detailed study necessary to organize a rational taxonomy of these genetic forms has not been attempted. However, the three geographic subspecies set forth below will clarify some relationships. Agave nelsonii Trel. has long been considered conspecific with A. deserti (Johnston 1924; Shreve and Wiggins 1964) and indeed is very similar in habit, in the gray glaucous leaves, and in other characters. However, the ceriferous in- florescence of A. nelsonii, the brown eyelets ringing the teeth, its chemistry and its geograph- ic distribution relate it with the A. cerulata com- plex. I am accordingly treating it as a subspecies of A. cerulata. This and two other variants are separable according to the following key and descriptions. GENTRY: AGAVES OF BAJA CALIFORNIA 43 Key to Subspecies of Agave cendata la. Leaves yellowish or light glaucous, long-acuminate, mostly 6-12 times as long as broad, the margins nearly straight to mildly undulate; teeth small, fragile; spines acicular 3-6 cm long 2 lb. Leaves mostly light-gray to bluish glau- cous over green, short-acuminate, mostly 3-6 times as long as broad, the margins undulate to mammillate; teeth larger, firmer; spines subulate, 2—4 cm long 3 2a. Leaves generally yellowish, 25-50 cm long; panicles narrow, the lateral branches 15-20 cm long on upper V2, to !4 of shaft c. cendata 2b. Leaves light glaucous gray, 40-55 cm long; panicles broad, the lateral branch- es 30—40 cm long on upper Vi of shaft. Isla San Esteban c. dentiens 3a. Leaves larger, 25-40 cm long, the mar- gins undulate with small teats or nearly straight. Sierra San Miguel to San Agus- tin c. nelsonii 3b. Leaves smaller, 15-28 cm long, the mar- gins conspicuously crenate with prom- inent teats. Vicinity of San Ignacio to San Marcos Island c. subcendata Edibility of A. cerulata. — Considering the rela- tively high sapogenin contents. Agave cendata is probably not among the best for eating. How- ever, it was eaten by the Cochimi Indians, a fairly large tribe inhabiting the central part of the peninsula. They ate the thick stem or ca- beza, which contains less sapogenins than the leaves. Wenceslaus Linck, a Jesuit missionary, founded the Mision de San Francisco Borja dur- ing the 5th decade of the 18th century. In 1766 he set out with thirteen soldiers and a band of missionized Indians to explore northern Baja California to the mouth of the Colorado River (Burrus 1966). He traveled over 480 km through the heartland of Agave cendata. Of the second day northwest of San Borja at Vimbet he wrote, "A native chieftain and his entire settlement of Nuestra Sonora de Guadalupe had prepared for us thousands of mescales , a food which is their daily bread and their favorite sustenance"" (ibid.:45). He mentioned mescales several times as the most common food the Indians gave to them along his trail, in exchange for his friendship and blessings. When crossing the southern end of the Sierra San Pedro Martir on March 13, he noted, "Our great pagan friend regaled us here with ten loads of ground mescal. Many uncon- verted natives gathered here, accompanied by a pagan savage whom I cured."" His account is the first we have on these wild people of north- ern Baja California and leaves no doubt of the importance of agave as food in this desert land. Further confirmation of this is given for more southern Indians in the wonderful account writ- ten by Padre Barco of the Mision de San Javier in the Sierra de la Giganta (see Introduction). Today, except in the Sierra San Pedro Martir region, there are no Indians left to eat the agaves. However, cattle, including horses, bur- ros, cows, and goats, crop the young flowering shoots extensively. Horses will bend over the flowering stalks and eat the flowers. Near Tres Virgines Mountain, I observed where some large animal had learned to break out the terminal bud of A. cendata subcendata and eat the soft cen- tral tissue of stem, bud, and leaf bases. The young flowering stem contains little or no sa- pogenin. The heads or stems of A. cendata den- tiens were eaten by the Seri Indians, who plied across the Gulf to reach them (see below). Agave cerulata ssp. dentiens (Trel.) Gentry, stat. nov. (Figure 21; Tables 2. 4) Agave dentiens Trel. Missouri Bot. Gard. Rep. 22:.SI. 1912. Medium-sized, acaulescent, surculose, open, few-leaved, green to light glaucous gray ro- settes, 50-70 cm tall, 8-15 dm wide, forming dense clumps; leaves 40-70 cm long, 4-7 cm wide in the middle, wider below, triangular-long- lanceolate, thickly crescentic at base, concave above towards apex, rigid, light gray glaucous, sometimes bluish, cross-zoned, the margins straight, usually set with small, weak, friable teeth 1-2 mm long, or margin nearly toothless; spine acicular. 3-5 cm long, brown to gray, nar- rowly, shortly channeled above, shortly decur- rent; panicle 3^ m tall, open, with 8-18 wide- spreading lateral branches in upper half of shaft; buds ceriferous, pale yellow, in small congested umbels; flowers pale yellow, slender, 49-53 mm long; ovary fusiform, 32-35 mm long with slen- der neck, constricted at base; tube shallow, open, 3 X 10 mm, swollen with nectaries below filament insertions, grooved; tepals ca. equal. 44 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. \M) 15-16 mm long, 4-5 mm wide, ascending- spreading, occasionally recurving, conduplicate and involute with anthesis, rounded and shortly hooded at apex; filaments slender, 28-30 mm long, bulbous at base with nectaries; anthers 15- 16 mm long, yellow, centric; capsules glaucous ceriferous, 40-50 x 15-20 cm. oblong, short- stipitate, short-beaked. (Flower description from Felger s.n. San Esteban Island. 20 June 1977). Type.— Rose 16819, San Esteban Island. Baja California 12 Apr. I9II, MO, US. There is little doubt that Agave dentiens is closely related to A. cerulata. The old frayed capsules on the type specimen show a waxy bloom like that of A. cerulata, while the long, narrow, acuminate leaf-form and small, weak, brown-eyeleted teeth are additional evidence for cerulata affinity. The random occurrence of toothless margins is also a cerulata character. A long-leaved, light-glaucous variant {Gentry & Fox 11953) of cerulata (Fig. 19) found on the peninsula between Punta Prieta and Bahi'a de Los Angeles looks like the island dentiens. The satellitic-island position of dentiens in relation to the peninsular cerulata populations is very similar to that of A. sohria roseana, here also treated as a subspecies. Moran's color photo- graphs of collection 4079 on San Esteban Island show a generally uniform population with rather consistent characters of a wide-branching pani- cle and tall, long-leaved, light gray-glaucous, narrow rosettes. This is in obvious distinction from the narrow panicles and usually small, yellowish rosettes of peninsular cerulata. But this difference seems to be at the subspecific level. It is possible that the agaves on Angel de la Guarda Island should be aligned with A. cerulata dentiens , but until flowering specimens of the re- spective populations are assembled, the dispo- sition would be uncertain. Altogether, ssp. dentiens looks like an isolated genome undergo- ing evolution and, perhaps, on its way to com- plete speciation. The Seri Indians called Agave cerulata den- tiens "?emme" (Felger and Moser 1970). The Seri visited San Esteban Island to collect the plant and found the more fibrous plants the most savory, distinguishing also a white-and-green variety. Large baking pits were observed on the island, and the Seri reported that as many as 100 heads were cooked at a time. They also relieved thirst with juice of the macerated leaves after they were cooked in a fire. The juice was also drunk after fermenting for several days with warm water added. Felger reports (personal communication) that the green-leaved form gen- erally occupies the higher elevations of the is- land, where fogs or clouds tend to hang and re- duce insolation. Agave cerulata Trel. ssp. nelsonii (Trel.) Gentry, Stat. nov. (Figures 21-23; Tables 2, 4) Active nelsonii Trel. Missouri Bot. Gard. Rep. 22:61. 1912. Short-Stemmed, rather small, cespitose, com- pact, glaucous green rosettes, 50-75 cm in di- ameter at maturity; leaves mostly 20-35 x 6-8 cm, rarely larger, lanceolate to triangular-lan- ceolate, short-acuminate, usually a little nar- rowed above a broad base, thick rigid, light green with a gray to bluish glaucous bloom; teeth in the mid-blade 3-9 mm long, frequently on small teats, brown-ringed at base, grayish brown, closely regular-spaced 1-2 cm apart, rel- atively firmly attached; spine 2-4 cm long, strongly subulate, flat or openly grooved above, grayish brown, decurrent to first or second pair of upper teeth; inflorescence slender, 2.5-4 m tall, with 15-20 ascending to arching laterals in upper half of shaft, the yellow flower umbels compact, globose; flowers slender, light yellow, 45-55 mm long; ovary 25-35 mm long, slightly angled, the neck long, grooved, slightly con- stricted; tube small, 3-5 mm deep, openly fun- nelform; tepals subequal, linear, erect, incurved at tip, hooded, the inner with prominent keel; filaments slender, 30-40 mm long, inserted at base of tepals on nectary disk; anthers centric, 15-18 mm long; capsules 4^.5 x 1.5 cm, ob- long or smaller and pyriform, stipitate, with acute beak; seeds 4x5 mm with hilar notch at apex and thin marginal wing. Type. — Nelson & Goldman 7111 . San Fernando (Sierra San Miguel). Baja California, 4 Sept. 1905. US. Goldman's photo (Treiease I9I2:pl.65) shows very well the habit of this plant and its habitat on the igneous highlands with open cover of cirio, ocotillo, cactus and low shrubs. Agave cerulata Trel. ssp. subcerulata Gentry, ssp. nov. (Figures 21. 23 (lower), 24. 25: Tables 2. 4) Small, acaulescent. few-leaved, open, glau- cous green cespitose rosettes 15-30 cm tall. 30- GENTRY: AGAVES OF BAJA CALIFORNIA 45 Figure 22. Af^ave cenilata nelsonii on igneous rocky slope of Sierra San Miguel near San Fernando. 22 June 1973. Pho- tograph by John McClure. 50 cm in diameter; leaves 15-30 x 2.5-7 cm, lanceolate to triangular lanceolate, thickly fleshy, arching upward, guttered at maturity, white glaucous to glaucous green, margins cren- ate with prominent tests under well developed teeth; teeth at mid-blade 3-8 mm long, variously flexed, weakly attached, grayish brown, 1-3 cm apart; spine 2^ cm long, subulate, usually sin- uous, grayish brown; panicles 2-3 m tall, slen- der, waxy glaucous, with mostly 8-10 small lat- eral umbels of light yellow flowers; flowers 44- 55 mm long; ovary 23-30 mm long with grooved neck 4-7 mm long; tube 3-6 mm long, shallow, openly spreading; tepals equal, 17-22 x 6-7 46 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Figure 23. {Upper) Af^avi' ccnilaiu nclsonii. detail of rosetle showing the typical short-acuminate leaves. {Lower) A. cerulaia suhcerulata near San Ignacio; the few-leaved rosettes with short thick leaves are characteristic. GENTRY: AGAVES OF BAJA CALIFORNIA 47 Figure 24. Agave cenilala ssp. siihcerulata. leaf drawn from type Gentry 10330; flowers from topotypic Gentry 23170. Leaf, flower cluster, capsule x'/i: flower section xWi. mm, smooth, thick, oblong, apex broadly round- ed, slightly hooded, the inner with broad low keel; filaments 33^0 mm long, inserted on rim of nectary disk at base of tepals; anthers 18-21 mm long; capsules waxy glaucous, mostly pyr- iform or obovoid, 3.5-4 x 1 cm. Type. — Gentry 10330, San Ignacio, Baja California. 3 Apr. 1951, "northern slope with volcanic rocks; Cardon-Fouquier- ia-Jatropha, etc..' deposited in US. Duplicates in DES. MEXU. Planta parva surculoso-caespitosa paucifol- iata: foliis plerumqiie 15-30 cm longis 2.5-7 cm 48 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 lafis ad medium, crassis lanceolatis condupli- ciitis ^lauco-pallidis, marline valde creiuita, dentibus supra mammas plerumque 3-8 mm longis flexuosis pallidogriseis dehilibus; spina terminali 2-4 cm longa suhulata leviter flexuosa pallido-castanea; inflorescentia paniculata cum scapo 2-3 m alta gracili omnino cerasea,fercnti 8-10 ramos parvos remotos; floribus pallido-lu- teis 44-55 mm longis: ovario cum apice sulcata 23-30 mm longo; tuba vadoso 3-6 mm longo expanse: segment is 17-22 mm longis 6-7 mm latis aequalibus, pallido-luteis crassis oblongis apice obtusis: filamentis 33-40 mm longis ad basim segmentorum insertis: antheris 18-2 1 mm longis luteis: capsulis 3.5-4 cm longis 1 cm latis pyriformibus vel obovatis . Agave cerulata subcerulata strongly resem- bles A. subsimplex across the Gulf along the Sonoran coast, but the relationship is clearly with A. cerulata. The depauperate, gray to yel- lowish rosettes growing on the gypsiferous is- land, San Marcos, may reach only 10 cm in stat- ure on open exposures. On northward slopes, the rosettes are larger and the panicles reach 2.5-3 m in height with 5 to 8 lateral branches (Fig. 25). Johnston (1924:998) made a similar observation. The ovary and neck of the San Marcos Island flowers collected are more slen- der than the peninsular ones, but otherwise the flowers are similar, including the waxy bloom on ovaries and pedicels. The short, thick, rela- tively broad leaves with long teeth on prominent teats provide characters for recognizing this subspecies both in the native habitats and in bo- tanical garden collections. The distribution as far as known is shown in Fig. 21, while habit and habitat appear in Figures 23 {lower) and 25. Agave margaritae (Figure 32) Agave margaritae Brandegee. Proc. Calif. Acad. Sci. ser. 2. 2:206. 1889. Agave connochaerodon Trel. Missouri Bot. Gard. Rep. 22:58. 1912. Small, compact, cespitose. glaucous gray to yellowish green rosettes with 40-50 leaves; leaves 25 x 10 cm to 12 x 7 cm, ovate to broad- ly lanceolate, short acuminate, thick, fleshy rig- id, concave above, narrowed above the base, the margin crenate with moderate to prominent teats; teeth at mid-blade 4-5 mm or to 8-15 mm long, variously curved or flexed. 1-1.5 cm apart, weakly attached, reddish brown to grayish; spine 2-3 cm long, subulate, shortly and shal- lowly grooved above, shortly decurrent; pani- cles 2-3.5 m tall, slender, with 6 to 12 short lateral branches in upper l/i of shaft; flowers light yellow. 45-50 mm long; ovary 25-30 mm long, fusiform; tube ca. 10 mm deep; tepals 15 X 5 mm. attenuate; filaments 25 mm long, adnate to the base of tepals (Brandegee) or in- serted in the throat of tube (Trelease); capsules 30-50 X 15-20 mm. not stipitate. oblong or pyr- iform; "seeds 3—4 mm in diameter, smooth" (Brandegee). Type. — Brandegee s.n.. 14 Jan. 1889, Magdalena Island, Baja California, UC. The type consists of two leaves, flowers, and a section of a lateral branch with capsules. Trelease separated his species A. connochae- todon from A. margaritae on the basis of longer, more flexuous teeth on larger teats, as he wrote, '"with considerable hesitancy that, because of the very different arming of the specimens col- lected, a second species is recognized for these islands." The population on Isla Margarita shows considerable variation in the size, cur- vature, teating, and the color of the teeth. Tre- lease's segregate, in the writer's opinion, is only one of the variants of A. margaritae and is ac- cordingly treated as a synonym here. The deep flower tube, the short broad leaves, together with other leaf characters described above dis- tinguish this species from all other peninsular taxa. The relatively deep tube of A. margaritae indicates relationship with A. vizcainoensis , but on leaf characters they appear specifically dis- tinct, as discussed in the account of A. vizcain- oensis. Agave margaritae is known with cer- tainty only from Magdalena Island, just off the outer coast in the southern part of the peninsula. Agave sobria Agave sobria Bdge. ssp. sobria (Figures 26-28: Tables 2. 5) Agave sobria Brandegee. F*roc. Calif. Acad. Sci. ser. 2. 2:207. 1889. Agave affinis Trel. Missouri Bot. Gard. Rep. 22:56. 1912. Agave carminis Trel. Missouri Bot. Gard. Rep. 22:55. 1912. Agave slevinii I. M. Johnston. Proc. Calif. Acad. Sci. ser. 4. 12:1000. 1924. Small- to medium-sized, usually cespitose, open, few-leaved, bright glaucous-gray rosettes with short stem, or appearing stemless, 50-150 cm in diameter; leaves linear to lanceolate, long- GENTRY: AGAVES OF BAJA CALIFORNIA 49 Figure 25. Agave cerulatu subceruUita on the Island of San Marcos. May 1952. acuminate, variable, but mostly 80 x 10 cm to 45 X 5 cm, frequently cross-zoned, straight to curved, sometimes twisted, plane to somewhat concave above, thick and concave below to- wards base, the margin undulate to mammillate; teeth remote, mostly 5-10 mm long with broad, flattened, gray bases, reddish towards apex, var- iously flexed or straight, mostly 3^ cm apart; spine acicular, mostly 3-6 cm long, narrowly grooved above; panicles slender, sometimes arching, 2.5-4 m tall with 12-20 short lateral branches, the umbels of flowers compact, nearly globose; peduncular bracts small, triangular, chartaceous; flowers pale yellow. 45-55 mm long, slender; ovary 25-35 mm long, tapering at base, the neck short, scarcely constricted; tepals 50 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 26. Agave sobria sobria. drawn from topotypic Barclay & Arguelles 1989: leaf, flower cluster and capsules xVi. flower section xl.3, seed x2. about equal, 17-22 x 3-4 mm, very narrow, lin- ear, rather thin, involuting at anthesis, rounded at tip, cucullate, the inner keeled; tube short, spreading. 3-4 mm long, 9-11 mm wide; fila- ments inserted at base of tepals on nectary disk. 35^7 mm long, slender; anthers 18-23 mm long, centrically attached; capsules 5-6.5 x 1.5-1.8 cm. oblong, thick-walled, short stipitate. apic- ulate; seeds 7-8 x 5-6 mm, lunate, the margins narrowly winged. Figure 27. A^uve sohria at the type locality on the lava rocks above San Miguel de Comondu, October 1951. 52 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Agave tobria sobria _ Agave sobria frailensis Agave sobria roseana _ Figure 28. Distribution of Agave sobria and subspecies, based on herbarium specimens (see Appendi.x — Exsiccatae). Type. — Brandegee 2. 28 Mar. 1889, Comondii mesas (Fig. 27), Baja California, UC. DS. Agave sobria is distinguished by its slender flowers with long narrow tepals and its very light-glaucous, long, lanceolate leaves with re- mote teeth. It is widely scattered but common on both sides of the Sierra de la Giganta moun- tain axis, from near sea level to 3,500 ft (1,070 m) elevation (Fig. 28). It appears to make its best growth on shadier or northern slopes in canyons, such as below the volcanic rim rock on the talus slope above Comondii. Figure 27 shows the plants at home at the type locality. As examined there in 1951, the rosettes com- monly throw out one or more branches from above the base, thus forming cespitose plants. When small, these shoots are readily lifted out of the dry trunk tissue as separate plants with young hard roots started downward. Large ro- settes develop when the plants are single or only double. The plants are scattered and do not form dense colonies. On the eastern foot of the Sierra de la Giganta the plants are generally smaller and drought depauperate. Edibility and Use . — The people at Comondii call this plant "mescal pardo"" or "pardito,*" saying also that it is good for making mescal. This is contrary to what Brandegee (1889) has reported and contrary to his use of the name sobria (for sober, or not given to alcohol); so I questioned the inhabitants closely about its edibility and suitability for mescal. They were unanimous in pronouncing it good and that Agave aurea, growing in the vicinity, was the one not suitable for mescal. Hence, I believe that Brandegee confused these two agaves on this point in his notes or memory. However, the Comondii informants, consist- ing of several older men, said that the agave best for eating and for distilling is called "lechuguil- la" and grows in the Sierra de la Giganta. A guide was appointed among them, and I dis- patched with him Juan Arguelles, my field as- sistant. They went by horse and returned with several plants (Gentry & Arguelles 10324, 10327) collected on the highland near the base of the mountains above the Llano San Julio. The plants were not flowering, but leaf specimens were pressed, young plants kept for planting, and samples sent into the USDA laboratory in Phil- adelphia for analysis. With this local information and our present series of specimens, it is now possible to orient the detailed description of agave uses made by Miguel del Barco in the 1770"s. He was located for about 30 years (1735- 1768) at the Mission of San Javier in the Sierra de la Giganta, and his remarks apparently apply mainly to A. sobria and A. gigantensis . both of which grow around the mission and were used extensively by the Cochimi Indians. There was a kind that was not eaten, wrote Barco, probably what is now called Agave aurea. Most of his remarks I have translated and quoted in the In- troduction. Chemistry. — Table 5 summarizes the sapogenin content of Agave sobria as determined by the United States Department of Agriculture on their cortisone-source survey during the 1950*s (Wall 1954: Wallet al. 1954a, 1954b, 1955, 1957). The four genins — hecogenin, tigogenin, mano- GENTRY: AGAVES OF BAJA CALIFORNIA 53 Table 5. Sapogenin Content in Agave sohria. Collated from Wall (1954) and Wall et al. (1954a, 1954b, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin: git. = gitogenin; man. = manogenin; tig. = tigogenin; dead = leaf of dead plant. Baja California Month Plant Sapogenins Coll. '^'f '"f % % % no. source locality coll. part total hec. git. man. tig- Agave sobria sobria 11291 Comondii May leaf 1.6 55 45 11291 Comondii May fruit 0.6 100 11291 Comondii Nov. leaf 0.5 X X X 11291 Comondii Nov. leaf (dead) 0.4 X X X 11811 Sierra de Palmas May leaf 1.3 100 12384 Sierra de la Giganta Dec. leaf 0.0 12384 Sierra de la Giganta Dec. leaf 0.6 X X X 12387 Bahfa Concepcion Dec. leaf 0.2 100 12387 Bahia Concepcion Dec. stem 0.0 12387 Bahia Concepcion Agave sobria roseana Dec. leaf 0.2 X X X 11274 Islota Gallo Oct. leaf 0.7 40 15 10 35 11274 Islota Gallo Oct. leaf 0.2 60 40 11277 Espi'ritu Santo Is. Oct. leaf 0.6 57 42 11869 La Paz Agave sobria frailensis May leaf 0.45 25 10 65 11264 Punta Frailes May leaf 1.1 20 10 70 11264 Punta Frailes Nov. leaf 1.4 X X 11257 Punta Frailes Oct. leaf 1.7 10 90 11257 Punta Frailes Nov. leaf (dead) 1.3 X X X 11858 Punta Frailes May leaf -> -> 100 12367 Punta Frailes Nov. leaf 1.7 X X 12368 Punta Frailes Nov. stem 0.0 12369 Punta Frailes Nov. leaf (dead) 1.3 X X X genin, and gitogenin — continue as in the ceru- lata complex to chemically characterize the De- serticolae, both in quantity and in random distribution. However, manogenin is lacking in the subspecies /ra/7(^//5/5 . nearly so in ssp. ro- seancu but frequent in ssp. sobria. Sapogenin content is conspicuously higher in frailensis , av- eraging 1.5% in leaves, compared to 0.6% in sobria and 0.5% in roseana. Early summer shows a higher average content of sapogenins in leaves (May 1.3%) than for winter (Nov. -Dec. 0.7%), but the data are scarce and not always strictly comparable. This is the reverse of what was obtained for A. cerulata, which is more in the winter-rainfall region, while A. sobria frai- lensis is in the summer-rainfall region. Is higher genin content correlated with dry seasons rather than with temperature? The lack of sapogenin in the edible stems or "cabezas" confirms the statement of the Comondii people that Agave sobria is a good source for mescal. Agave sobria, although it reproduces asexu- ally. is a freely seeding sexual species. It shows a lot of variability among individuals in certain localities, especially in the southern part of its area and on adjacent islands, where isolation has had time to foster genetic divergence. Trelease's A. roseana, which Johnston reduced to a vari- ety, appears to represent a natural segregate of sobria, and accords with my use of subspecies in this treatise. I am separating it and another geographic variant, hitherto unrecognized, ac- cording to the following key and descriptions. Key to the Subspecies of A. sobria la. Leaves linear lanceolate, mostly 50-80 cm long, 6-10 times longer than wide. 54 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. L^O the margin merely undulate; tepals very narrow, 3—4 mm wide sobria lb. Leaves broadly lanceolate. 25-50 cm long, 4-5 times longer than wide, the margins prominently mammillate; te- pals 4-6 mm wide 2 2a. Rosettes few-leaved, openly spreading, yellow-green; leaves larger, 40-50 cm long, with large teeth few and remote on mammillate margins. Isla Espiritu Santo and adjacent peninsula roseana 2b. Rosettes with more leaves, compact, urceolate, bluish-gray glaucous; leaves smaller, 25-30 cm long, with smaller teeth closely spaced along mammillate margin. Punta Frailes and vicinity. Cape District frailensis Agave sobria Edge. ssp. roseana (Trel.). stat. nov. (Figures 28. 29. 30 [riglit and upper): Tables 2. 5) Agave roseana Trel. Missouri Bot. Card. Rep. 22:59. 1912. Agave sobria var. roseana I. M. Johnston. Proc. Calif. Acad. Sci. ser. 4. 12:1002. 1924. Small, open, few-leaved, cespitose or single, yellow-green rosettes; leaves 35-50 x 7-10 cm, broadly lanceolate, acuminate, concave above, frequently twisted, the margin prominently mammillate, teats 1-1.5 mm high, with remote large flexuous teeth, sometimes bicuspid, the larger 10-25 mm long; spine 5-7 cm long, sin- uous to contorted, acicular, narrowly grooved above, castaneous to gray; panicles 2.5-3.5 m tall, slender, with 8 to 12 compact, globose um- bels of light yellow flowers in upper third of shaft; peduncular bracts small, triangular; flow- ers 45-65 mm long; ovary 30-40 mm long, cy- lindric; tube 4-5 mm long, spreading; tepals 20- 22 X 4-5 mm. long-linear, erect-ascending, in- curved at apex and cucullate. narrowing with anthesis. inner strongly keeled; tepal sinuses at first closed, wilting open; filaments 35-42 mm long, inserted at base of tepals on rim of tube; stamens 18-20 mm long; capsules 4^.5 x 1.5- 1.8 cm, oblong, short-stipitate. Type.— Rose 16854. 18 Apr. 1911. Espiritu Santo Island. Baja California. US. Agave sobria roseana is distinguished by its open, few-leaved rosettes, its short broad leaves with prominent mammillate margins with large irregularly flexed teeth, and its restriction to the southeastern portion of the sobria area. It owes its distinction apparently to long isolation on Espiritu Santo Island and the adjacent islets. Is- lote Gallo and Islote Gallina. It also occurs near La Paz on the peninsula along with plants that are typically sobria. but I have had no trouble in separating the few specimens collected there. I did not find evidence of crossing between them, but such may occur. A. sobria roseana appears morphologically consistent both on the islands and about La Paz. Variations noted were differences in size of plants and in the size and number of teeth. The population on Islote Gal- lina showed cespitose plants as common, but all plants on Islote Gallo were singles. Islote Gallo had a rather uniform and dense colony, while elsewhere the plants were widely scattered. As an ornamental this plant is striking because of its bizarre teeth on big teats. Agave sobria ssp. frailensis Gentry, ssp. nov. (Figures 28. 29. 30 {lower left): Tables 2. 5) Small, compact, sometimes urceolate, glau- cous green to bluish glaucous, sparingly cespi- tose rosettes; leaves mostly 20-35 x 6-8 cm, conduplicate, lanceolate, acuminate, markedly narrowed below, the margin mammillate and with numerous, flexuous, castaneous to graying teeth mostly 6-10 mm long; spine subulate 3^ cm long, frequently sinuous or contorted, de- current to upper teeth; panicle slender, 3^ m tall, with deltoid chartaceous peduncular bracts and 10-15 compact yellow umbels in upper '/3 to '4 of shaft; flowers 45-63 mm long, slender; ovary 25-40 mm long, cylindric, with short un- constricted neck; tube 2.5^ mm deep, small, sulcate from tepal sinuses; tepals 17-23 x 4-6 mm, linear-lanceolate, incurved and cucullate at apex, the outer dark-tinged at tip, the inner with prominent keel; filaments slender 30-40 mm long inserted on rim of tube; anthers 18-22 mm long, centric or excentric; capsules short ob- long. 3-4 cm X 1.5-1.7 cm. sessile to short-stip- itate. rounded at apex. Type .—Gentry & Cech 11264. 4 miles [ca. 6 km] N of Punta Frailes, Baja California, 7 Oct. 1951, on granitic slopes. US. Transplant flowered in Murrieta. California. July 1%2. Planta parva surculoso-caespitosa: species foliorum glaucedine cyaneis insignis; foliis ple- rumque 20-35 cm longis 6-8 latis ad medium, rigescentibus condiiplicatis lanceolatis, margine valde crcnata. dentibus supra mammas pleruin- GENTRY: AGAVES OF BAJA CALIFORNIA 55 Figure 29. Agave sobria ssp.frailensis: from type. Gentry & Cech 11264, leaf, flower cluster, capsule xi/2. flower section xl. Agave sobria ssp. roseana from topotypic Gentry & Cech 1 1277, flower section x 1. 56 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. KM) hiGL Rt 30. {Linvci icji I Ai;uvc uihna i,iis castcmcis tandem pallido- cinerasventihus: spina lerminali aciculata 3-4 cm longa ftexuosa castanca \el canescenli de- currenti supra hrevi-canaliculata; inflorescentia paniculata gracili, cum scapo 3^ m alta.ferenti 10-15 ramos congestos; floribus 45-63 mm longis pallido-luteis; ovario cylindrico 25^0 mm longo: tubo brevi 2.5^ mm longo: segmentis 17-23 mm longis lineari-lanceolatis apice incurvatis cucul- latis; filamentis gracilibus 30-40 mm longis ad tubi orificium insertis: anlheris 18-22 mm longis luteis: capsulis 3^ cm longis 1.5-1.7 cm latis breviter oblongis sessilibus vel bievi-stipitatis apice rotundatis . Agave sobria frailensis differs from its near relative A. sobria roseana in its smaller size, more numerous leaves, lighter gray-glaucous color, softer flesh of the leaves, and more nu- merous, more regular, and smaller teeth. In 195 ! GENTRY: AGAVES OF BAJA CALIFORNIA 57 Figure 31. Agave moranii. drawn from type. Geiiiry & McGill 23287: leaf, flower cluster, capsule xi/2. flower section, slightly enlarged. and 1952 it was found in typical form scattered upon the granitic coastal cerros 2 to 8 miles (3- 13 km) northwest of Punta Frailes in the Cape District on the peninsula. This is the only area in which it was observed. Because of its compact urceolate rosette form and the glaucous, sometimes bluish, color of the leaves, it makes an attractive ornamental. A transplant (Gentry & Cecil 11264) flowered in Murrieta after 1 1 years, but numerous offsets 58 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. LW 116° 114" 112° 110° \ ! \ 1 ; 1 1 Agave avellanldens • Agave gigantensis _ __ _ O Agave vlzcalnoensi* _ _ _* Agave margaritae _ — * Agave moranii _ _ _ 19 V \ — ?^\ \ V^-s^ ~'~'~-'-^ ) \ ^^ _\^L: k \>!i\n ^■. IcY \ 28° ^, .N ^ i s. 3.\.^ ^o^ i ^ h ^ \ > sT Figure 32. Distribution of Aguve uvellanidens. A. gigan- tensis, A. margaritae. A. vizcainoensis . and A. moranii. based on herbarium specimens listed in Appendix — Exsiccatae. from the base of the plant subsequently failed to root. Agave moranii Gentry, sp. nov. (Figures 31-35: Table 2) Large, single, short-stemmed, mostly light- green rosettes with symmetrical sword-leaved habit. 1-1.5 m tall. 2 m broad; leaves triangular- long-lanceolate. 70-120 X 8-12 cm. deeply gut- tered, straightly ascending to spreading, rigid, deeply rounded beneath, light green to yellowish green, sometimes glaucous, strongly armed; spine stout 4-6 cm long, nearly white with cas- taneous tip, broadly grooved above, decurrent to mid-blade; teeth sinuously flexed or curved, flattened, 6-12 mm long through the mid-blade and below, 2-4 cm apart, reduced and more re- mote towards apex on the white corneous leaf margin, light gray, the base broad or continuous with the margin; panicle 4-5 m tall, stout, fre- quently swollen and closely bracteate below base of panicle, with 20-30 closely spaced, com- pact, large, umbellate branches subtended by prominent, lanceolate, reflexing. strong bracts; flowers 50-70 mm long, bright yellow, on slen- der bracteolate pedicels 1-3 cm long; ovary 25- 40 mm long, fusiform, tapered at base and with short, grooved, thick neck; tube shallow, 4-6 X 12-13 mm. discoid-spreading, the nectary ridged by decurrent filaments; tepals 18-24 x 6- 7 mm, erect to ascending, broadly linear, acute, with papillate tufted beak, the inner with prom- inent bulbous keel and 2 costae within, slightly lobed at base; filaments very slender, 35-46 mm long, inserted on rim of nectary and partly on base of tepals; anthers centrically affixed; cap- sules 5-7 X 1.6-2 cm, long-oblong, stipitate, short-beaked, yellowish to brown; seeds 7-8 x 5-6 mm, lacriform, shiny, the marginal wing smooth, variable in width. Type.— Gentry- & McGill 23287, 2-3 miles [3-5 km) SE of Rancho Agua Caliente on E bajada of Sierra San Pedro Martir, Baja California, elev. ca. 1.500 ft [450 ml, 13 June 1973 [1. f, cap] US, isotypes in DES, SD. Planta grandis hreviter caulescens, rosula pa- tiilus simplex 1-1.5 m alta 2 m lata: foliis 70- 120 cm longis 8-12 cm latis ad medium, anguste irianguli-lanceolatis conduplicatis rigentibus viridibus vel pallidi-viridibiis: margine fere recto cornea; spinis marginalibus variabilibus flex- liosis ad medium foliorum 6-12 mm longis 2-4 cm separatis, pallido-brunneis vel albis; spina terminali robust a 4-6 cm longa, alba apice cas- tanea supra late canaliculata; inflorescentia paniculata cum scapo 4-5 m alta robusta, fer- enti 20-30 ramos approximatos umbelliformes congestos; bracteis grandis siccis triangularibus ad apice aliquando congestis; floribus clarolu- teis 50-70 mm longis; ovario fusiformi 25^0 mm longo ad basim angusto; tubo breri 4-6 mm longo 12-13 mm lato; segmentis 18-24 mm lon- gis 6-7 mm latis, linearibus acutis apice papil- latis cucullatis; filamentis gracilibus 35^6 mm longis ad basim segmentorium insertis; antheris 17-21 mm longis luteis ad medium affixis; cap- sidis 5-7 cm longis 1 .6-2 cm latis, oblongis stip- itatis brevirostratis; seminibus 7-8 mm longis 5- 6 mm latis lacrimiformibus nigris. GENTRY: AGAVES OF BAJA CALIFORNIA 59 Figure 33. Agave moranii in fine array on northeastern bajada of Sierra San Pedro Martir. near Rancho Parras, June 1973. Photograph by John McClure. Agave moranii is distinguished from all other Deserticolae by its large single rosettes of large, long, rigid leaves with white corneous margins apically, by its large-bracteate, stout peduncles, sometimes swollen below the panicles, and by its relatively congested panicles. The only agave with which it may be easily confused, especially from leaf specimens alone, is A. deserti pringlei in the same region. As discussed in the descrip- tion of A. deserti priiiglei. these two taxa appear to hybridize; without complete specimens they may not be satisfactorily identified. This is the case with Moran 15256 from Rancho San Pedro Martir on the western slope of Sierra San Pedro Martir. A photograph of the plant, however, shows that it is probably surculose — the large panicle, small-bracteate with well-spaced later- als, leading me to place it with A. deserti prin- glei. The respective habitats appear distinct: A. moranii is desertic, while A. deserti pringlei is at home in the chaparral zone on the Pacific slopes of the mountains. Agave moranii occu- pies a small area of the southern Sierra San Ped- ro Martir and its eastern bajadas, between ele- vations of 1,500 and 5,000 ft (450 and 1,850 m). The specific name is in honor of Reid Moran, the first botanist to make recognizable collec- tions of the species. Agave moranii appears aligned phylogenet- ically with A. avellanidens and A. gigantensis, all having separate areas along the sierran axis of the peninsula (Fig. 32). As stated above, the three are distinct from other Deserticolae by their non-surculose habits, and their broader and greener leaves. Near a fork in the road 7 miles ( 1 1 km) south of Rancho Agua Caliente near Rancho Parras, there is a fine population of Agave moranii (Fig. 33). It is most concentrated on low ridges run- ning out from the sierra side, but it is also scat- tered on sandy slopes and in the arroyos. Leaf color varies from a light green or glaucous green to yellow-green; a few are lightly zoned. Leaf form (Fig. 33) and armature are fairly constant, but there is some variation in teeth. All healthy plants have outstanding radiating leaf attitudes. Leaf margin is white with a dark brown border along the green. Some peduncles are distinctive with congestion of bracts below the lowest lat- erals where the peduncle is perceptibly swollen 60 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 34. Leaves of Agave moranii. Photograph by John McClure. (Fig. 34). This suggests the same flowering habit as A. inacrocidinis , A. murpheyi, and A. par- rasana, which have the peduncle initiated in the fall, arrested over winter, and a second elonga- tion with the warmer days of spring. The people living at Agua Caliente have no name to distinguish A. moranii from A. deserti, which is common in the same vicinity. They stated that the flowers were good to eat. The leaf has an abundance of strong coarse fibers and probably was employed by the Cocopah In- dians who formerly inhabited the region. Moran GENTRY: AGAVES OF BAJA CALIFORNIA 61 Figure 35. A flowering plant of Agave niorami and detail of a thickened peduncle with congested bracts. Photograph by John McClure. found a single outpost plant of this species 10 miles (16 km) or so towards the east, Moran 18295, "only one seen, SE side of San Felipe Valley (Sierra Santa Rosa?)/' Agave avellanidens {Figures 32. 36. 37; Table 2) Agave avellanidens Trel. Missouri Bot. Gard. Rep. 22:60. 1912. Single, medium to large, multi-leaved, green rosettes, 6-12 dm tall, 10-15 dm broad, with stems to 5 dm long; leaves 40-70 x 9-14 cm, broadly linear-lanceolate to ovate, little or un- narrowed base, short-acuminate, thick-fleshy, rigid, smooth, green, the margin straight or un- dulate and frequently corneous; teeth rather reg- ularly spaced, mostly 1-3 cm apart, but variable in size and curvature, 5-15 mm long, straight or 62 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OE SCIENCES. No. 130 Figure 36. Agave avcllanidcns west ot Calmalli. April IV72. I he deep narrow panicles and small bracts distinguish it from Agave shawii goldmaniuna . Photograph by Bruce Gentry. GENTRY: AGAVES OF BAJA CALIFORNIA 63 variously flexed, flattened, dusky gray over brown; spine strong, conical, 2.5-4.5 cm long, broadly grooved above, brown to grayish, strongly decurrent as a corneous margin; panicle 4-6 m tall with 25-35 lateral branches of dense, large, globose umbels of small flowers; flowers pale yellow, drying orange-yellow, 40-70 mm long, slender; ovary 20^0 mm long, fusiform, tapering to base, the neck sometimes constrict- ed, pale yellow; tube 4-6 mm deep, 13-15 mm wide, wide-spreading, ridged within and grooved without from overlapping tepal sinuses; tepals ca. equal, 16-24 mm long, linear-lanceolate, ob- tuse, the inner with thick keel and sometimes lobate within an inlapping base; filaments slen- der, 30-45 mm long, inserted on rim of tube; stamens 15-22 mm long, yellow, centric; cap- sules dark brown, broadly oblong, 20 x 35 mm, not stipitate and scarcely beaked; seeds un- known. Type. — Bnuidegee 6. Paraiso [S of San Borja). Baja Cali- fornia. I May 1889. UC [1. f. cap]. Agave aveUanidens is a shcmii-\\V.Q member of the Deserticolae. The elongated many-leaved green rosettes look like A. shawii goldmaniana where the two meet and mingle southeast of Punta Prieta. This relationship has been dis- cussed under goldmaniana, group Umbelli- florae. When in flower, A. aveUanidens is dis- tinguishable by the long, narrow, small-bracted panicles of small flowers with short open tubes. These inflorescence characters align it with the Deserticolae rather than with Umbelliflorae, re- gardless of the shawiian appearance. The distribution of A. aveUanidens is poorly known. So far as known, it is limited to a small area in mid-peninsula between latitudes 29° and 28° N. The type locality, Rancho Paraiso, is sev- eral miles south of the Mision San Borja in the mountains and apparently has not been visited by any botanist except Brandegee, who was fol- lowing mule trails. The species is abundant along the old auto road from Calmalli to Rancho Mesquital (Fig. 36). Agave shawii goldmaniana is scattered abundantly along this old road north of Rancho Mesquital to Punta Prieta, and oc- casional elongated panicles appear to represent A. aveUanidens. Certain sightings were also made along the road from San Borja to Rancho Rosalito. Atypical, small-bracted panicles on single rosettes of A. shawii goldmaniana . com- mon in the region, made reliable sightings diffi- cult south and east of Punta Prieta. Agave avel- lanidens and its associates of Fouquieria, Yucca , and Pachycormus comprise many exotic and beautiful plantscapes. Perhaps A. aveUani- dens is extensive in that large unknown area about and south of the Sierra Calmalli east of the main road. Agave gigantensis Gentry, sp. nov. (Figures 32. 38^0: Table 2) Single, acaulescent, green to glaucous green, rather open, few-leaved, medium-sized rosettes 5-10 dm tall. 8-12 dm broad; leaves 40-75 x 11-16 cm. plane, rigid, thick-fleshy, sclerophyl- lous, smooth, broadly lanceolate, acuminate, widest in the middle, markedly narrowed at base, green to glaucous, turning red to purplish with flowering, and depressed, the margin un- dulate to prominently mammillate; teeth large, sometimes grotesque, mostly 10-20 mm and even longer, the bases thick, frequently 2-3-cus- pidate and confluent along upper leaf margins, brown to light grayish, variously flexed and curved, generally remote, up to 6-8 cm apart; spine 3-6 cm long, strongly subulate, deeply sul- cate above, straight or sinuous, gray, long de- current as a heavy corneous margin, sometimes to mid-leaf; inflorescence 4-5 m tall, slender, the peduncular bracts narrowly lanceolate, thickly chartaceous; panicles rather narrow, in upper V3 to '4 of shaft, with 15 to 25 rather small umbels of bright pale-yellow flowers; buds waxy white; flower 48-60 mm long, slender, with spreading tepals; ovary slender, fusiform, with short, con- stricted, slightly grooved neck; tube short dis- coid, spreading, bulbous with tepal bases, 4-5 mm deep, 11-13 mm broad; tepals 18-25 x 5-6 mm, linear, rounded and briefly hooded at apex, the inner with prominent keel, grooved within and strongly overlapped at base by outer; fila- ments 30-45 mm long, variable in length, slen- der, inserted on rim of tube with base of tepals; anthers 18-25 mm long, somewhat excentric; capsules 3.5-4 x 1.2-1.5 cm. oblong, non-stip- itate, non-beaked; seeds unknown. T\pe. — Gentry & McGill 23320. above Rancho San Sebas- tian, Sierra de las Palmas, 31 miles [ca. 50 km] by road W of San Bruno. Baja California, elev. 3.800-5.000 ft [1.150-1.520 m]. 20 June 1973, US. Isotypes DES. SD. MEXU. Plant a simplex nonsurculosa, rosida 5-10 dm alia, 8-12 dm diametro lata glaucifolia vel vir- 64 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 37. Agave avellanidens: upper, a series of '■normar' leaves; lower, a form with bizarre armature of 2- to 3-cuspid teeth. Photographed west of Calmalli. GENTRY: AGAVES OF BAJA CALIFORNIA 65 Figure 38. Agave gigantensis, drawn from type, Gcniry & McGill 23320: leiif. flower cluster 2. flower section x 1%. idifolia aliquando ad maturitatem sanguiuea: crenoto: dentihiis supra mammas plerumque foliis 40-75 cm longis 11-16 cm latis ad medium, 10-20 mm longis saepe 2-3-cuspuiatis flexuosis late kmceolatis planis rigidulis crassi-succulen- vel curvatis, remotis usque ad 6-8 cm separatis; tis laevibus, ad basim angustatis; margine valde spina termmali 3-6 cm longa robusta subulata 66 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 <%■ * ^, Figure 39. Agave gigunlensis: (Upper) a transplant flowering at Murrieta, California, with geotropic flowers and a hum- mingbird. (Lower) A mature rosette on Sierra de las Palmas. GENTRY: AGAVES OF BAJA CALIFORNIA 67 recta vel sinuosa albocastanea, supra prof uncle canaliculata; inflorescentia paniculata cum sca- po 4-5 m alta gracili ferenti 15-25 rainos um- belliformes congestos parvos; bracteis anguste triangulis crassi-chartaceis; floribus pallide lu- teis fere albis 48-62 mm longis; gemmis albis; ovario fusiformi 25-35 mm longo apice constric- to; tuba vadoso 4-5 mm longo 11-13 mm dia- metro; segmentis 18-25 longis 5-6 mm latis ad medium linearibus, apice brevi-cucullatis; fila- mentis 30-45 mm longis variabilibus gracilibus ad apicem tubi insertis; antheris 18-25 mm lon- gis luteis excentricis: capsulis 3.5^ cm longis 1.2-1.5 cm latis, oblongis non stipitatis: semi- nibus ignotis . The small waxy pale flowers with short spreading tube in a slender panicle places A. gigantensis in the Deserticolae. As with other members of this alliance, the habit and leaves provide specific distinction. The leaves are re- markable in their broad form above a narrowed base, their singular quality of color, bud print- ing, and firm sclerophyllous smoothness, and bizarre variable large teeth on mammillate mar- gins (Figs. 38, 39). The nearest phyletic relatives appear to be the disjunct A. avellanidens and A. moranii to the north. The name gigantensis is given because the distribution is coincident with the Sierra de la Giganta range of the southern peninsula (Fig. 32). On Sierra de las Palmas (also known locally as Sierra Campana), Agave gigantensis is closely associated with an oak woodland com- munity that mantles the craggy tops of these vol- canic mountains (Fig. 40). Nolina beldingii is a conspicuous member of this community. South- ward, A. gigantensis has been observed and col- lected on the brushy slopes west of the Cerro Giganta itself and along the scarp-rim country above Loreto (see Exsiccatae citations). Alto- gether, it is known to range from elevations be- tween 2,000 and 5,000 ft (600 and 1,520 m) be- tween latitudes 27° N and 25° N. This is a small area, and A. gigantensis shapes up as another rare endemic plant. Fortunately it occupies a very rugged and largely inaccessible habitat, which in this age of destructive man will favor its survival. The rancher at San Sebastian, who guided us upon the mountain, stated that this agave is more abundant on "Sierra del Potrero," northward in this same mountain range. It is a symmetrical, beautiful plant that could well be valued and cultivated by the sophisticated agave fanciers. It does not reproduce vegetatively, however, and seeds would be needed to keep it generating. The name 'iechuguilla" is applied to this agave by the people of Comondu. It is consid- ered as one of the best for distilling into mescal. Of the five samples of leaves sent in for analysis, only two showed sapogenin in minor amounts, 0.3% hecogenin. Undoubtedly, this is one of the more edible agaves to which Miguel del Barco referred in his early account (quoted in the In- troduction). Agave vizcainoensis Gentry, sp. nov. (Figures 32, 41; Table 2) Plants acaulescent, small to medium, surcu- lose or single, the rosettes 30-50 cm tall, 50-90 cm wide, few-leaved and open in habit; leaves lanceolate, 25-40 x 6-10 cm, glaucous gray to green, sometimes reddish, broadest in middle, narrowed above base, thick-fleshy, rather rigid, the margin undulate, corneous above with de- current spine; teeth largest along mid-blade. 5- 10 mm long. 1-3 cm apart, slender or broadly flattened, dark brown to grayish, nearly straight or curved, spine 2.5-4 cm long, stoutly subulate, mostly rather straight, brown to grayish, shal- lowly grooved above, long decurrent; panicle 2- 3 m tall with 8 to 15 spreading lateral umbellate branches in upper half of shaft; flowers yellow 65-75 mm long, on stout small-bracteolate ped- icels 8-12 mm long; ovary green. 36^1 mm long, in fresh flower neck scarcely constricted but drying very narrow, 6-8 mm long; tube fun- nelform, 8-12 mm deep, 15 mm broad, bulging at filament insertion; tepals 21-26 x 4-5 mm, linear-lanceolate, involute above spreading base, obtuse, cucullate. strictly erect at anthe- sis; filaments 55-70 mm long, inserted somewhat unequally 7-9 mm above base of tube; stamens 21-26 mm long, excentric; pistil to 70-85 mm long with broad lobate stigma; capsules 5-7 x 2 cm, oblong, stipitate, rostrate, striate, brown; seeds 6-7 x 4.5 mm, lacrimiform, black, mar- ginal wing narrow. Type. — Gentry 7469. Cerro Tordillo and vicinity. Sierra Viz- caino, Baja California, 12-13 March 1947; elev. 400-800 ft [120-250 m]; desert of dispersed succulent trees and suffrutes- cent shrubs, UC; isotypes, DS, ARIZ, MEXU, DES. 68 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 «s^ ^i:*-* .4 f^i^ Figure 40. Agave giganiensis in native habitat on Sierra de las Palmas. Solina hcldingii in background. Photographed by John McClure, June 1973. Planta acaulis simplex vel surculosa grisea, 3-5 dm alta 5-9 dm lata, paucifoliata: foliis lan- ceolatis 25-40 cm longis 6-10 cm latis, pallidis vel griseo-viridibus carnosis; margine undulata. dentibus plerumque 5-10 mm longis 1-3 vel 4 cm separatis, surrectis vel flexuosis gracilibus aliquando complamitis, atrobninneis vel ciner- ascentibus: spina terminali 2.5-5 cm longa ro- biista recta vel flexuosa. brunnea vel cineras- centi, supra canaliculata longe decurrenti: inflorescentia paniculata 2-3 m alta diffusa, fer- enti 8-15 ramos laterales: floribus luteis 65-80 GENTRY: AGAVES OF BAJA CALIFORNIA 69 'Gentry ■ by Howard Scott U«ntrv PLANTS OF BJU* (;«I!hoi!\n m, xi, Figure 41. T\pe cnllectuMi c)t\4i,'<;i(' vizcuiruH'usis . Gentry 7469. an isotype sheet in the Gentry Herbarium. mm longis; ovario viridiflavo; tubo infundibidi- formi 8-12 mm longo, apice 15 mm diametro: segmentis 21-26 mm longis 5-6 mm latis lineari- lanceolatis, involutis cucidlatis, rectis sub an- thesi; filamentis 55-70 mm longis gracilibus, 7- 9 mm supra basim tubi insertis, antheris 21-26 mm longis, excentricis; pistillo usque ad 70-85 mm longo stigmate irilobato; capsulis 5-7 cm longis 2 cm latis oblongis stipitatis rostratis brunneis; seminibus 6-7 mm longis, 4.5 mm latis lacrimiformibus nigris. marginc in alum angiiste e.xpansa . The relatively deep flower tubes of this spe- cies suggests affinity withy4. margaritac of the 70 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 42. Distribution of the group Campaniflorae. based on herbarium specimens hsted in .'Kppendi.x — Exsicca- tae. same geographic region, but the latter, with its short, broad, long-toothed leaves and small size, appears distinct. Neither species is well known; the collections are few and incomplete. The spe- cies do not show close relation with other mem- bers of the Deserticolae, but they are assigned here for want of a more suitable section to re- ceive them. Their small size, surculose habit, shape of leaf and armature, and the small scar- ious bracts of the peduncle exclude them from the A. shawii complex. The deepening tube of subspecies simplex in the pivotal complex of A. deserti, at almost opposite ends of the Deserti- colae area, seems also to permit their inclusion in the Deserticolae. The plants growing about the Picachos de Santa Clara are more robust and less surculose than those in the Sierra Vizcaino proper and re- semble A. gigantensis. Gentry 7713 agrees well with the type in leaf morphology, but the tepals are relatively long in relation to the shorter tube (Table 2). Another specimen (Gentry 7693). also from the Picachos de Santa Clara, is more ex- treme with tepals to 35 mm long. Group Campaniflorae Trelease. Missouri Bot. Card. Rep. 22:44. 1912. Large to small, perennial or multiannual plants with rather open, short-stemmed rosettes; leaves green, rather soft succulent, long-lanceo- late, frequently sigmoid towards apex, with reg- ular, close-set, moderate teeth on noncorneous margins. Panicles large, diffuse, in upper half of shaft subtended by small scarious bracts; flow- ers campanulate with deep, ample tubes, deep- set filaments, red to purple on outside, yellow within; capsules short-oblong to ovoid, rather thin walled. Southern half of peninsular Baja California. The leaf surface in the Campaniflorae is mod- erately roughened but noticeably thickened. The stomates are slightly depressed and without rims and with overlapping polar lips in the montane A. promontorii. The number of stomata per mm' ranges from 18 to 31 on the upper leaf surface, a relatively low density, setting them apart from the Deserticolae and Umbelliflorae. Further par- ticulars of epidermal structure are given in the companion study of Gentry and Sauck (1978). The Campaniflorae occupy one of the most isolated areas of the continental Agavaceae (Fig. 42). The pre-Cretaceous granitic pediment on which they grow has long been isolated from the contemporary formation of the Sierra Madrean mainland. This isolation seems to be reflected by the Campaniflorae in their consistent confor- mation to the distinct campanulate flowers with broad tubes and thin-margined tepals, all with a certain succulent texture difficult to describe. The regular, moderate, close-set teeth on the soft succulent leaves are also consistently main- tained through the group. The flower propor- tions appear unusually variable (Fig. 43 and Ta- ble 6). I have no explanation for this. They do not appear useful for species separation. The connective gland with the filament-anther fixa- tion in A. capensis has not been observed else- where (Fig. 45). The habit of axillary budding with resultant large clusters of many rosettes. GENTRY: AGAVES OF BAJA CALIFORNIA 71 aurea CAMPANIFLORAE 12375 23182 23182 1988 12341 capensis promontori 11247 11676 ^O LJEl 11676 1987 1986 1986 promontori i 1986 Figure 43. Floral ideographs of the group Campaniflorae. showing relative proportions of the tube (black) to outer tepal (white column), and level of inseertion of filament (black square). Measurements are listed in Table 6. fouiKJ in A. capensis, is regarded as a relic char- acter, widely scattered in the genus from the UmbeUiflorae of the northwest to the Striatae and Polycarpae of central and eastern Mexico. The rainfall silhouettes in Figure 44 show that the Campaniflorae occupy moister habitats than other agave groups of the Peninsula. Rain falls predominantly in summer-fall from storms of tropical origin. The large growth-form o^ Agave promontorii appears to reflect the higher rainfall. 72 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 CAMPANIFLORAE a urea San Javier 317 Sierra Lacuna 747 Cabo San Lucas 200 ^ 100 250 Figure 44. Rainfall (silhouettes) and flowering (bars) perimeters of the Campanifiorae. Relevant meteorological stations with mean annual rainfall in millimeters. Data from Hastings 1964. Flowering periods based on herbarium specimens and field observations, supplemented by plants in cultivation. It is notable also that flowering occurs in the dry spring months (Fig. 44), when birds or other pol- linators would be. by thirst, more attracted to the liquid flower nectar. The Campaniflorae are free-seeding outbreeders, not given to vegeta- tive reproduction. The prehistoric uses of the Campaniflorae plants are unknown. Agave aurea alone is known to have been exploited for fiber for a short period late in the 19th century, and a large population on the lavas about Comondu was probably decimated. The following tables (7 through 9) of chemical analyses show that some steroids are common in all three species, with hecogenin being the most common. As with oth- er peninsular species, however, the incidence of the sapogenins appears fugitive within species, by seasons, and by physiological plant stages. Until further investigations can show how three percent (3%) or more sapogenin content can be consistently produced for harvest, the cultiva- tion of Agave for sapogenin alone is not eco- nomically feasible. Key to Species of the Campaniflorae la. Rosettes large, nonsurculose multian- nuals; leaves 60-150 cm long; panicles broad; filaments without apical gland. __ 2 lb. Rosettes smaller, perennials by axillary budding; leaves mostly 35-60 cm long; panicles narrow; filaments with an api- cal gland at anther fixation. Cape Dis- trict A. capensis (p. 72) 2a. Rosettes generally 0.7-1 m tall; leaves smaller, more pliant, 7-12 cm broad. Sierra de la Giganta and Cape District A. aurea (p. 78) 2b. Rosettes 1.5-2.3 m tall; leaves larger, thicker, relatively rigid, 11-17 cm broad. Sierra Laguna, Cape District , A. promontorii (p. 81) Agave capensis Gentry, sp. nov. (Figures 42-47; Tables 6-7) Agave brandegeei in hort. & herb. Plants perennial by axillary budding, even- tuating in large clusters with short-stemmed, small, open rosettes. 6-8 dm tall, 8-12 dm broad; leaves mostly 30-60 x 4-7 cm, narrowly lanceolate, straight to arching, commonly sig- moid towards apex, concave above, convex be- low, light glaucous green, soft, brittle, succu- lent, with undulate noncorneous margin; teeth regular, 4-5 mm long, mostly 1-2 cm apart, mildly curved, on short mammillate bases, reddish brown to grayish; spine 1.5-3 cm long. GENTRY: AGAVES OF BAJA CALIFORNIA 73 Table 6. Flower Measurements (in mm) of the Group Campaniflorae. (* — measurements from dried flowers; measurements from fresh or pickled flowers.) Fil. insert Total Coll. Taxon & locality Ovary Tube Tepals & length Anther length no. aurea Comondii 32 8 X 14 16 X 6 5 17 18 55** 12375 29 8 X 14 16 X 6 6 35 19 54** 12375 20 9 X 14 16 X 6 5 38 20 45** 12375 30 11 X 17 19 X 5.5 7-8 44 23 67** 23182 34 11 X 16 19 X 5 8-7 45 23 63** 23182 36 15 X 15 16 X 5 10-9 46 20 66** 23182 39 14 X 15 17 X 5 9-8 45 21 70** 23182 Todos Santos 30 9 X 18 17 X 9 7&5 41 21 55** 1988 30 10 X 18 15 X 9 7& 5 40 20 54** 1988 20 7 X 8 17 X 10 6-5 45 19 43** 12341 21 6 X 18 15 X 10 5-t 43 19 43** 12341 Cape Region 32 7 X 8 14 X 7 4-5 40 19 52* Bdge. capensis Cabo San Lucas 34 12 X 20 20 X 7 6 38 28 65** 11247 31 10 X 20 23 X 7 7 43 27 63** 11247 27 8 X 13 13 X 6 6-7 26 15 48** 19676 24 8 X 14 13 X 6 5-6 27 16 44** 1%76 25 11 X 16 12 X 7 5-6 32 19 48** 1987 28 11 X 16 13 X 7 5-6 40 20 52** 1987 promontorii Sierra Laguna 42 14 X 20 16 X 9 8 &6 54 26 72** 1986 36 14 X 20 14 X 9 8&6 50 24 64** 1986 41 14 X 20 15 X 9 8&6 50 24 70** 1986 subulate, dark brown, short decurrent for 1-2 cm; panicles mostly 2.5-3.5 m tall with 15-24 lateral branches in upper % to Vi of shaft, the laterals up to 30 cm long, ascending: umbels small; bracteoles very small, 2-3 mm long, nar- row-triangulate; flowers in bud reddish brown or purplish, opening yellow inside. 50-65 mm long; ovary green, 25-35 mm long, thick. 3-angled, furrowed in unconstricted neck; tube 8-14 mm deep. 15-20 mm broad. 6-bulged. 6-furrowed from tepal sinuses; tepals equal. 13-23 x 6-7 mm, lanceolate, incurved and rather thin; fila- ments 30-43 mm long, yellow, inserted 5-7 mm above base of tube, the apex swollen with col- orless glands; anthers 15-27 mm long, excentric, yellow or bronze; capsules 25-35 x 15-17 mm. ovoid, scarcely stipitate, apiculate; thin walled; seeds not seen. Type.— Gentry & Fox 11247. Cabo San Lucas & vicinity, Baja California. Young plant collected 5 Oct. 1951; flowered in Murrieta. California. July 1964; deposited in US. Planta caespitosa perennis per geminas ro- sulatas axillares; foliis angustis plerumque 30- 60 cm longis 4-7 cm latis linearibus vel lanceo- latis canaliculatis arcuatis vel sigmoideis fnigi- libus molliter succiilentis glauco-viriclihus, mar- gine undulatis; dentibiis ad medium foHonim 4- 5 mm longis plerumque separatis 1-2 cm. pau- lum curvatis castaneis vel griseis; spina termin- ali 1.5-3 cm longa subulata alrobrunnea. de- currenti usque ad 1-2 cm; inflorescentia paniculata cum scapo 2.5-3.5 m altaj'erenti 15- 24 ramos umbelliformes parvos: floribus 50-65 mm longis in alabastro rubris vel purpureis sub anthesiferrugineis intra luteis; ovario 25-35 mm longo viridi rotundato-triquetro: tubo 8-14 mm longo 15-20 mm diametro. campanulato 6-sul- cato ferruginescenti; segmentis 13-23 mm lon- gis 6-7 latis infra medium, lanceolatis cucullatis tenuibus incurvatis aequalibus; Jilamentis luteis 30-43 mm longis 5-7 mm supra tubi basim in- sertis. glanduloso-tumidis ad apicem: antheris 15-27 mm longis luteis vel ferrugineis excentri- cis: capsulis 2.5-3.5 cm longis 1.5-1.7 cm latis. ovoideis subsessilibus apiculatis valvis tenuibus: seminibus ignotis . 74 OCCASIONAL PAPFRS OF THE: CALIFORNIA ACADEMY OF SCIENCES, No. 130 Figure 45. (C), Agave capensis drawn from type: leaL flower cluster, capsule xi2. flower section x2'/i, apex of filament enlarged to show gland. {P) A. promoniorii. flower section x2'/3. GENTRY: AGAVES OF BAJA CALIFORNIA 75 t •^-'"^ Figure 46. Agave capensis. a large axillary-branching clone in Huntington Botanical Gardens, California, January 1951. Agave copensis appears to be an islandic en- demic of the Cape District. It grows with the sparse open shrubbery on the arid, neutral, well- aerated soils on granitic slopes from near sea level to 1,000 ft (ca. 300 m) or more above sea level. Scattered single rosettes northwest of Punta Frailes appeared similar to those of A. capensis, but their immaturity at time of sighting leaves their identity doubtful. This is also true of a collection {Gentry 11301) in the shady can- yon above La F^irisima, which probably repre- sents a narrow-leaved shade form of A. aurea. The flower structure o^ Agave capensis is very similar to that of its sectional relatives,/!, aurea and A. promontorii , but it is distinguished by its small narrow leaves and clustered growth habit. Figure 46 shows a large clump in Huntington Botanical Gardens in 1951, which Mr. Hertrich, the superintendent, stated was about 40 years old. The photographs in Figure 47, taken in the same year at San Jose del Cabo, show both sin- gle and branching rosettes and are both very similar to the Huntington clone. A specimen transplanted from San Jose del Cabo in 1951 flowered at Murrieta in July 1964 and is selected as the type. Subsequent to the July flowering, some of the axillary rosettes also flowered, but none of the flowers at Murrieta developed fertile fruits. The connective gland or swelling found on the stamen where the anther is affixed to the fila- ment (Fig. 45) is a structure peculiar to the Cam- paniflorae. While well developed in A. capensis, it was also noted in minor form on a putative hybrid, x A. aurea. These two species appear to be close, even genetically compatible, as dis- cussed under A. aurea, but the origin of the fil- ament gland appears to lie with A. capensis. 76 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. \iO Figure 47. Agave capensis at Cabo San Lucas; (upper) a cluster of rosettes showing axillary branches, and (lower) a single rosette. GENTRY: AGAVES OF BAJA CALIFORNIA 77 Figure 48. Agave aiirea, leaf from Gentry 11295. flower cluster from Barclay & Argiielles I9m. capsules xVi. flower section xWi. Leaf is incurled from drought. 78 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 In the Huntington Botanical Gardens in San Marino, and in other California gardens, this species has been known as Agave hrandegeei Trel. I had hoped to maintain this name, but examination of the type specimen. "Cape Re- gion mountains. Brandegee Sept. 20. 1899. " in the herbarium of the Missouri Botanical Garden shows that this name must be abandoned. The type specimen consists of a partial leaf and flow- ers of A. aurea Brandegee and flowers oi A. sobria Brandegee. Furthermore, in describing the species. Trelease excluded flowers of A. au- rea and described those of A. sobria to accom- pany the leaf specimen. The two parts of the type, therefore, are synonymized under A. au- rea and A. sobria. The Cape species must be recognized and described as a new species with a new name. It is a curious coincidence that both the Brandegee and Purpus collections from the Cape region are of the same mixture. Obviously, there was some mishandling of specimens after the collections were made, but by whom and when is unknown. Agave aurea (Figures 42-44. 48^9: Tables 6. 8) Agave aurea Brandegee. Proc. Calif. Acad. Sci. ser. 2. 2:207. 1889. Single short-stemmed, rather open, green, graceful rosettes 10-12 dm tall. 15-20 dm broad, with widely arching leaves; leaves 63 x 7 cm to 110 X 12 cm. average 86.3 x 8.6 cm. linear to long-lanceolate, pliant, guttered, rounded be- low, thickly fleshy towards base, green to some- what glaucous, the margin straight to undulate: teeth moderate, regular, mostly 4-7 mm long. 1- 2 cm apart, straight or moderately curved cusps from low angular bases, dark brown to light brown: spine subulate. 25-35 mm long, dark brown or grayish red. with a short narrow groove above, shortly decurrent or decurrent as a dark corneous margin through 8 to 10 teeth bases: panicles 2.5-5 m tall, broad, the peduncle reddish and with remote, quickly drying lanceo- late bracts: umbels broad, congested. 15 to 25 in upper half of shaft, red to purplish in bud and fruit: flowers opening yellow to orange-yellow. 43-70 mm long, campanulate: ovary reddish. 25-35 mm long, slender, angular-cylindric. with constricted furrowed neck 6-10 mm long: tube 8-14 mm deep. 14-18 mm broad, evenly grooved from tepal sinuses: tepals 16-19 mm long, lanceolate, acute, incurving, the outer larger with tufts of white papillae below distinct hood, the inner with long keel and yellow, thin margin involuting at anthesis: filaments stout, 35-45 mm long, those of outer tepals inserted 5- 10 mm above base of tube, slightly higher than those of inner tepals: anthers 19-23 mm long: capsules oblong. 45 x 17 mm to 35 x 15 mm, non-stipitate, rounded apiculate, reddish, drying light brown: seeds 7-8 x 5-5.5 mm, irregularly lunate, dull black. Type. — Brandegee s.n.. Purisima. Baja California. 13 Feb. 1889. UC. Consists of marginal sections of a large leaf, a small leaf. 5 flowers, pieces of 2 capsules. The plant does not grow in the sandy valley at La Purisima: doubtless collected on the volcanic mesas near that settlement. Characteristics and Distribution . — Agave au- rea is the most abundant species in the group Campaniflorae. It is easily recognized by the long narrow lanceolate green leaves arching out to form an open, spreading rosette, by the broad, rather diffuse, reddish panicles, and by the bright yellow flowers from reddish buds and ovaries. The teeth are moderate and regularly spaced (Figs. 48. 49). It forms scattered and sometimes massive populations on the extensive lava fields of the western slopes of the Sierra de la Giganta. mostly between elevations of 1.000 and 3,500 ft (ca. 300 and 1,070 m) above sea level. It is also widely scattered on the granitic lower slopes of the Cape District at lower ele- vations. It is lacking or rare on the sandy Mag- dalena plain. In the latitude of Todos Santos, an A. aurea population shows forms similar to both A. pro- montorii and A. capensis . Here there are forms like aurea in the size of their rosette and in the color and form of their leaves, but with the soft- er leaves and more proximal teeth of promon- torii. Here also are small cespitose-rosette forms with softer narrower leaves of low fiber content like those of capensis at Cape San Lucas. Large inflorescences with well-developed fruits and ovules are among these forms, indicating inter- fertility. It appears that A. aurea may have in- vaded the islandic Cape District after it was con- nected to the peninsula in Pleistocene times and is now receiving genes from the insular promon- torii and capensis populations. Fiber of A. aurea and A. promontorii. — The fi- ber of Agave aurea was at one time harvested in the vicinity of Comondu. where this agave grew in abundance upon the lava beds. Accord- GENTRY: AGAVES OF BAJA CALIFORNIA 79 Figure 49. Agave aurea on the lava beds north of Comondii. April 1973. Photograph by Bruce Gentry. ing to Comondii informants, this was towards the end of the 19th century. In 1951 some adobe ruins several kilometers back in the canyon be- low Comondii marked the site of the decorticat- ing mill. The operation was extensive, and cart trails may still be traced through the lava fields where the leaves were hauled in. Nelson (1921:38) in 1905 noticed these same roads when approaching Comondii by horse from La Puri- sima. It is not now known just why this opera- tion terminated; was the accessible supply ex- hausted, or did the operation not profit? The fiber oi Agave aurea is fine and of excel- lent spinning quality, but yield of fiber per leaf 80 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. L^O Table 7. Sapogenin Content in Agave capensis. Collated from Wall (1954) and Wall et al. (1954a. I9.'i4h. 1955. 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin: git. = gitogenin: man. = manogenin: tig. = tigogenin. Baja California Month Plant Sapogenins Coll. 'T'c 9?- 9r 9c 9c no. source locality coll. part total hec. git- man. tig- 11247 Cabo San Lucas Oct. leaf 0.4 35 65 11247 Cabo San Lucas Oct. stem 0.0 11247 Cabo San Lucas Mav leaf 1.0 100 11247 Cabo San Lucas Nov. leaf 0.0 11250 Cabo San Lucas Oct. leaf 0.8 80 20 112.S0 Cabo San Lucas May leaf 0.55 20 80 11250 Cabo San Lucas Nov. leaf 3.6 X X X 11847 Cabo San Lucas May fruit 0.3 20 30 50 is low. and compared with henequen or sisal could not be competitive for twine and rope. In 1952 leaf samples of A. aiirea and A. promon- torii were sent to the Mexican Fibre Company in Tampico. Tamaulipas, for testing. Grady Venable. in charge of that operation, made the following report. 1) A^ave promontorii. The fiber is excellent and, it is believed, would be acceptable to the market. The yield of fiber per leaf, however, is too low for commercial operation, but it could probably be made to produce more fiber under proper planting conditions. If the chemical prop- erties are more valuable than the henequen now grown on El Carrizal. Tamaulipas. it might be cultivated commercially. 2) Agave aurca. This fiber, while of excellent texture, cannot be produced commercially un- less it is found that the yield can be increased. It cannot be commercially produced unless its chemical content should prove exceptionally valuable. weight, approx. 2.000 1. Agave promontorii: A. 4 leaves — total grams. Average weight per leaf — 500 grams. Yield of fiber — total 65 grams. Percentage yield per gross — 3.25%. E. Yield per leaf — average 16 grams. F. Milling characteristics — good. G. Texture of fiber — fine, softer than hene- quen. H. Spinning factor — rates as excellent. I. Break test — our estimate 20% weaker than henequen. J. Length recovered fiber — 80 cm. good. B. C. D. K. Bleaching — color (sun), good. L. Pulp adherence — minimum, good. 2. Agave aurea: A. Approximately as above. B. Approximately as above. C. Yield — approximately 26 grams. D. Percentage yield — 1.21%. E. Yield per leaf — average 6 grams. F. Same as above. G. Same as above. Better ihan promontorii . H. Same as above. Better {ha.n promontorii . 1. Same as above. J. Length recovered fiber — 60 cm. short commercially. K. Same as above. L. Same as above. 1 supplied the test samples and can add some additional qualifying notes. The A. aurea sam- ples sent were collected near Todos Santos in the Cape District from plants having atypically little fiber in the leaves (see discussion above). Leaves decorticated by hand from Comondii plants were found to have a stronger and more abundant fiber than the sample sent. Transpor- tation difficulties and working conditions pre- vented getting Comondu samples to Tampico for analyses. Chemistry. — The tables (7, 8 and 9) giving the sapogenin contents of the Campaniflorae. en- able one to judge further any commercial pos- sibilities. The selection of individual plants with abundant fiber together with high sapogenin content for breeding could materially raise the commercial potential. GENTRY: AGAVES OF BAJA CALIFORNIA 81 Table 8. Sapogenin Content in Aguve aun-a. Collated from Wall (1954) and Wall et al. (1954a. 1954b, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin; git. = gitogenin; man. = manogenin; tig. = tigogenin; dead = leaf of dead plant. Baja California Month Plant Sapogenins Coll. % % % % % no. source locality coll. part total hec. git. man. tig. 11198 La Paz Sep. leaf 0.7 100 11253 San Jose del Cabo Oct. leaf 0.5 70 20 11253 San Jose del Cabo May leaf 0.4 100 11255 Las Cuevas Oct. leaf 0.0 11283 NW of La Paz Oct. leaf 0.0 11295 Comondii Oct. leaf 0.1 100 11295 Comondii May leaf 0.0 100 11295 Comondii Oct. leaf (dead) 0.3 11297 Comondii Oct. leaf 0.55 90 10 11297 Comondii May leaf 0.34 87 13 11297 Comondii Dec. leaf 1.4 X X X X 11823 Todos Santos May leaf 0.0 11823 Todos Santos May fruit 1.6 45 40 15 11886 Comondii May fruit 0.5 5 30 65 12383 San Javier Dec. leaf 1.2 100 12338 La Paz Nov. leaf 1.2 X X X X 12339 La Paz Nov. leaf 1.1 100 12342 Todos Santos Nov. leaf 0.0 12341 Todos Santos Nov. leaf 0.7 X X 12420 Comondii Dec. leaf 0.1 X X Agave promontorii (Figures 42^4, 50; Tables 6, 9) Agave promontorii Trel. Missouri Bot. Gard. Rep. 22:50. 1912. Large, single, green, open rosettes, 1-2 m or more tall, 2-2.5 m broad, with thick stems; leaves 10-15 dm long, 1 1-17 cm wide, lanceolate fleshy succulent, stiff, usually concave above, thick at base, straight to arching, green to soft glaucous green, the margins about straight and closely set with regular straight to curved teeth, mostly 4-8 mm long, 5-10 mm apart, reddish brown; spine 3-5 cm long, conic-subulate, dark brown, short-decurrent, narrowly sulcate above; inflorescence 5-9 m tall, massive, the peduncle reddish and with conspicuous deltoid bracts; panicles large, broad, with 25-30 diffuse umbels on recurving laterals in upper half of shaft, the buds and ovaries red to purplish; flowers short- pedicellate, 60-75 mm long, campanulate; ovary 36^2 mm long, 3-angled-cyIindric, scarcely nar- rowed at base, 6-furrowed and narrowed in neck; tube 14-15 mm long, 20 mm broad, cup- shaped, 6-bulged, with deep furrows between bulges; tepals 14-16 x 9 mm, triangular-lanceo- late, equal, thin, reddish purple on outside, yel- low within, the outer plane and overlapping in- ner at base, the inner with yellow margins and low, broad, 4-lined keel, with inner costae pro- nounced and wide apart; filaments 50-55 mm long, inserted unequally at 8 and 6 mm, oval in cross-section, yellow; anthers 24-26 mm long, yellow; capsules shortly pyriform-oblong, 15-20 X 30-35 mm, rather stipitate, beaked; seeds narrow, 4-5 x 6-9 mm. (Fl. description from Barclay and Arguelles 1986; cap. and seed de- scription from Trelease 1912). Type. — Nelson & Goldman 7437, Sierra de la Laguna. Baja California, 21 Jan. 1906. US. Agave promontorii is a large handsome plant, becoming massive with thick stems and thick juicy leaves in fertile situations (Fig. 50). The arching attitude of the leaves combined with their soft glaucous green and regular close-set teeth make it an attractive ornamental. It is, however, rare in gardens because it does not off- set and must be renewed with seedlings. The plants growing in Balboa Park, San Diego, and my plants near Murrieta, California, did not set seed. Night freezes of 24 F (-44 C) injured the leaves causing apical dieback. The species is known only from the granitic 82 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. L^O Table 9. Sapogenin Content w Afiaw pnmiontohi. Collated from Wall (1954) and Wail et al. (1954a, 1954b, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin; git. = gitogenin; man. = manogenin; tig. = tigogenin; dead = leaf of dead plant. Month Plant Sapogenins Coll. % % % % % no. Source locality coll. part total hec. git. man. tig. Baja Cal PORN I a 11218 Sierra Laguna Oct. leaf 0.3 55 55 11218 Sierra Laguna Oct. stem 0.0 11229 La Burrera Oct. leaf 0.75 90 10 11229 La Burrera May leaf 0.0 11218 Sierra Laguna Nov. leaf 3.4 X X X 12346 Sierra Laguna Nov. leaf (dead) 0.4 X X 12347 Sierra Laguna Nov. inflo. 0.2 X X 12353 Sierra Laguna Nov. leaf 0.8 X X 12349 Sierra Laguna Nov. leaf 0.7 X X 12351 Sierra Laguna Nov. leaf (dead) 0.9 X X 12352 Sierra Laguna Nov. leaf 0.9 X X 12350 Sierra Laguna Nov. leaf 0.5 100 12356 Sierra Laguna Nov. leaf 0.5 X X X 12344 Burrera Nov. leaf 0.4 100 12345 Burrera Nov. leaf 0.8 X X X X California 3233 Balboa Park, San Diego Nov. leaf 0.0 mountains of the Cape District. It attains best development in the higher elevations from 3,000 to 6,000 ft (ca. 900-1,800 m) above sea level, coincident with the better moisture conditions about the Sierra de la Laguna. It flourishes, for instance, on the open, precipitous, rocky, south- west slopes above the Rancho Burrera. Al- though it was observed in the shade of trees, it does not thrive in the crowded chaparral thick- ets common on the Sierra Laguna. Strays from the montane populations are common along the rocky arroyos issuing from the canyons on the west, as at Rancho La Burrera. Due to its lim- ited and remote range, A. promontorii is another vdre Agave scarcely known to man. Except for variations in color and teeth. Agave promontorii appears well-knit morpho- logically. The flowers show it to be closely re- lated to the other members of the Campaniflor- ae. However, its large size in both rosette and leaf distinguish it clearly from both A. aurea and the still smaller cespitose A. capensis. It was noted earlier that some of the aurea population around Todos Santos bears resemblance io pro- montorii, but no individuals of the latter were observed in the vicinity. The apparent introgres- sion there was probably only between A. aurea and A. capensis, whose populations meet in the low coastal elevations there and near Cabo San Lucas. I failed to record any local uses o^ Agave pro- montorii. nor have I seen any account on this subject by other field botanists. The thick, suc- culent, young, flowering shoots look tempting as a primitive sweet, and quite possibly they were eaten by the original Amerindians, who, because of their susceptibility to European diseases, dis- appeared from the Cape District soon after Spanish colonization in the 18th century. The leaf fiber is fine, readily twillable, and of good length, strength, and quality, but leaf yield rel- ative to the heavy succulent bulk or weight is low. Analysis of the leaf fiber is given and dis- cussed above under A. aurea. The sapogenin content is indicated in Table 9, but here, as else- where in the peninsular A^'ait', the percentage content of sapogenin is rather fugitive. The com- mercial possibilities of the combined fiber and chemical products remain but little investigated. Group Umbelliflorae Trelease, Missouri Bot. Card. Rep. 22:44. 1912 Small to large, freely seeding, long-genera- tion, and frequently long-stemmed plants com- GENTRY: AGAVES OF BAJA CALIFORNIA 83 Figure 50. Agave promontoiii in Balboa Park, San Diego, California, with large hut sicnic mtmctescences. monly branching from leaf axils, eventuating in fragmented supine clones; leaves broad, thickly succulent, stiff, well armed, mostly bright green: inflorescence stout, compact, with large, many- flowered, umbellate branches subtended by large, succulent, sheathing bracts; flowers large, fleshy, with ample tubes and strong divergent stamens; ovaries thick; capsules large, thick walled. Baja California. The shawii complex is distinguished primarily by perennial plants that flower repeatedly, ac- cording to the development of axillary branches. Flowering is not at regular seasons, but occurs only as the branching rosettes mature. This ap- pears, from the few records available and from field observations, to require twenty to forty years. Records show that a specimen of ssp. golclmaniana (Desert Botanical Garden No. 42) 84 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. \30 UMBELLIFLORAE shawii J Ensenada 283 200t 100-- QJ- sebastiano Isia Cedros 85 shawii goldmaniana J Punta Prieta 200 T lOO" O-*- 95 Figure 51. Rainfall (silhouettes) and flowering (bars) of the Umbelliflorae. Relevant meteorological stations with average annual rainfall given in millimeters. Data from Alias Meteorold^ico de Mexico (Servicio Meteorologico Mexicana 1939) and Hastings 1964. Flowering periods based on herbarium specimens and field observations, supplemented by plants in gardens: uncertainty expressed by broken line. collected as a small plant by George E. Lindsay near San Andres, Baja California, in the mid- thirties, required more than 35 years to mature and flower in 1972. The trunk at that time was about 2 m long, had declined upon the ground, and is presently being continued by two large branch rosettes. Further exposition of this unique habit is given in the Introduction. The individual rosettes, especially when they have declined upon the ground, rooted, and become separate or independent, may be regarded as multiannuals, a common reproduction habit among agaves. Moran (1964) reported an age of 31 years to maturity for a plant of ssp. gold- maniana at the Mina Desengafio in Central Baja California. It should be noted that this perennial habit is not maintained throughout the various popula- tions. Numerous individuals have been ob- served that mature and die without leaving any branched rosettes to follow. This is strikingly apparent, for instance, in Ihe goldmaniana pop- ulations south of Punta Prieta, where single ro- settes far outnumber the clustered rosettes (Fig. 58). There are no other consistent morphological features to distinguish these single plants from clustered plants, and they are regarded as intra- subspecific. The causes of this variation in habit, whether environmental or genetic, are not known. It may be regarded as a growth-habit potential that is not always individually fulfilled. The Umbelliflorae are further distinguished by their green, smooth leaves with relatively thin and unchanneled cuticle, by their rimmed sto- mata without polar lips, and entire surface-pore margins. The number of stomata on the upper leaf surface ranges from 40 to 47 per mm^. Il- lustrations and further particulars of epidermal structure are given in Gentry and Sauck (1978). The closest group relation of the Umbelli- florae is not clear. The Campaniflorae of the southern half of the peninsula are well separated by their narrower, nonrigid. fine-fibered leaves with regular teeth, and by their shorter, thinner, campanulate flowers in deep, small-bracteate panicles. One species of the Campaniflorae, A. capensis, like A. shawii. branches from the leaf axils. But I do not believe this constitutes close affinity because other, more distant relatives in the subgenus Littaea also have this growth hab- it. The Umbelliflorae and Campaniflorae occupy distinct geographic regions with no sympatric members. The only observed possibility of gene interchange by members of Umbelliflorae with other group representatives is that between A. shawii goldmaniana and A. avellanidens of the Deserticolae. The Umbelliflorae occupy an arid climate, but GENTRY: AGAVES OF BAJA CALIFORNIA 85 are unique among agave groups in being con- fined to a Mediterranean type of climate with winter-spring rainfall and dry summers. The rainfall silhouettes of Figure 51 show only 85 & 95 mm (3 & 4 inches) of annual rainfall for .v(^- bastiana and ^oldinaniana, respectively, yet they are robust plants with broad leaves. How- ever, the rainfall lack is greatly ameliorated by frequent fogs that condense and bathe the leaves, and reduce insolation and temperatures. Even the more inland populations of ^oUiman- iana are regularly visited by inland-moving fogs. The fogs increase and dilute the nectar in the cupullate flowers, which is frequently copious in early morning hours. The flowering seasons are also indicated in Figure 51. The flowering seasons of A. shawii and ssp. goUlmaniana are relatively long and commonly start with the spring rains, but they appear to vary from year to year and have been observed to flower in the fall. The Deserticolae diverged with the increasing aridity of continental climates, while Umbelli- florae remained in the more equable, conserva- tive, maritime environment. In concert with western mountain forming and diversification of climate into isolated habitats, the Deserticolae evolved into relatively numerous species and subspecies, while in the Umbelliflorae specia- tion remains indistinct. In our present state of knowledge, it is not possible to state when, in geologic or biologic time, the group divergence began . The axil-branching growth habit, a type of pe- rennialism among the agaves, is found generally in the more equable tropical climates and ap- pears to be an ancient characteristic. Its disjunct occurrence on the maritime California coast ap- pears as a relic survival. In the Agavaceae, axil- branching is a homologous permagene charac- ter. Agcive shawii ssp. shawii and A. sebastiana look very similar in habit, as they both have elongate, tightly imbricated rosettes, short thick peduncles, and wide congested panicles of very similar, large, succulent, bright-yellow flowers. They are the maritime ecotypes of the species, whi\e goUlmaniana is the desertic ecotype. His- torically, one can speculate that in pre-Pleisto- cene times, shawii and sebastiana forerunners were continuous along the coast from San Diego to Cedros Island. With the orogenic changes in the Pleistocene, there was also a drying of cli- 1 \'^. \ \ \ 1 1 1 Agave shawii shawii _ — — • Agave shawii goldmanlana „'* Agave sebastiana — — _ * ^- ■ \ I, v^ ?^ 1 J^ "S "^ 1 • \ I \ ^l* * ^ .)} - *4' ( \ ^ ^ ;\ \ I IM, n\ / V 0 I 4 1 vi ^ 1 \ \ ^ Y Figure 52. Distribution of the Umbelliflorae, based on herbarium specimens listed in Exiccatae (Appendix). mate. Desert shrub moved over to the coast be- low Rosario, and A. goldmaniana moved with the shrub and displaced and separated/!, shawii into two populations. A. goUlmaniana became contaminated with shawii genes and remained or regressed to a subspecies. A. sebastiana sur- vived as a disjunct on the Cedros Island group and, as of the present, has reached a specific level of taxonomic divergence. During the early 1950's, samples of leaves, stems, and inflorescences were collected of Agave shawii to determine the sapogenin con- tent. The results of this chemical survey of the Agricultural Research Service are brought to- gether in Table 10. The sapogenin content was found insufficient as practicable sources for fab- ricating cortisone and other steroidal drugs. Other economic aspects are noted below. 86 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Table 10. Sapogi-nin Content in Af;civc slwnil. Collated from Wall (19.S4) and Wall et al. (19.S4a. 19.'i4h, 1955, 1957). Sapogenins are given in percentages on dry-weight basis: hec. = hecogenin; git. = gitogenin; man. = manogenin: tig. = tigogenin: wh.pl. = whole plant. Month Plant Sapogenins Coll. % % % Vc % no. Source locality coll. part total hec. git. man. tig- Agave shawii shawii Baja California 10282 Km 57 S of Tijuana Mar. leaf 0.0 10283 Km 57 S of Tijuana Mar. stem 0.0 10281 Km 57 S of Tijuana Mar. infl. 0.0 10285 San Telmo Mar. leaf 0.38 10379 Socorro Apr. fruit 0.35 10379 Socorro Apr. leaf 0.0 11180 Socorro Apr. infl. 0.56 11694 Rosario Apr. fruit 0.2 97 11698 Rosario Apr. leaf 0.25 55 35 10 11699 Rosario Apr. leaf 0.15 100 10383 Rosario California Apr. fruit 0.0 Huntington Bot. Card. Agave shawii goldmanianu Baja California leaf 0.0 10349 Marmolito Apr. leaf 0.0 10355 Tinaja Yubay Apr. infl. 0.14 100 10355 Tinaja Yubay Apr. leaf 0.06 50 50 10355 Tinaja Yubay Apr. stem 0.1 10355 Tinaja Yubay Apr. infl. 0.15 60 27 10359 Tinaja Yubay Apr. leaf 0.3 63 10 20 11317 Marmolito Oct. fruit 0.25 55 28 17 11318 Punta Prieta Oct. leaf 0.3 X 11319 Punta Prieta Oct. leaf 0.0 11964 Mina Desengano Oct. fruit 1.3 70 30 11936 Mesquital Oct. wh.pl. 0.7 X X 12402 Punta Prieta Dec. wh.pl. 0.8 X 12401 Punta Prieta Dec. leaf 0.9 X X 12400 Punta Prieta Dec. leaf 0.6 X Key to Taxa of the Umbelliflorae la. Panicle about as broad as long, subtend- ed by a cluster of large, succulent, sheathing bracts below lowest laterals; laterals 8 to 15; leaves short, generally only 20^0 cm long, short-acuminate, usually less than 3-3.5 times longer than broad 2 lb. Panicle longer than broad, the bracts below lowest laterals smaller and not closely clustered; laterals mostly 20 to 30; leaves longer, generally 40-60 cm long, long-acuminate. Mid-peninsula __ shawii goldmaniana (p. 93) 2a. Panicle with a rounded crown, the sub- tending bracts large, succulent, red to purple; leaves bright green. Northern peninsular coast __._ shawii shawii (p. 86) 2b. Panicle with a flat crown, subtending bracts small, scarious, pink to yellow; leaves glaucous gray to green. Cedros Island sebastiana (p. 96) Agave shawii Agave shawii Engelm. spp. shawii (Figures 51-57; Tables 10, II) Agave shawii Engelm. Trans. Acad. Sci. St. Louis 3:314, 370. 1875. Agave onuiiicinu Tree. Missouri Bot. Card. Rep. 22:47, 1912. Agave pachyacantha Tree. Ibid. p. 48. Single or cespitose, compact, green, small to medium rosettes with short to long stems (to 2 GENTRY: AGAVES OF BAJA CALIFORNIA 87 Figure 53. (5), Agave shawii ssp. shanll: leaf, flower cluster x |'/2. flower section x I , from Gentry 23190: larger leal" from Gentry 23154, capsule xVi, seeds x IVi. (G), A. shawii ssp. golJmaniana: flower section x 1 from Gentry 19974. 88 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No, 130 Table 11. Flower Measuremenls (in mm) of ihe Group Umbellielorae. (* ** — measurements from fresh or pickled flowers.) measurements from dried flowers: Fil. nsert. Total Coll. Taxon & locality Ovary Tube Tepals & 1 sngth Anther length no. shawii shawii La Mision 37 12 X 19 25 X 8 8 73 25 74** 23190 37 16 X 18 23 X 7 8-7 72 26 77** 23190 37 14 X 18 24 X 7 9-7 72 25 77** 23190 42 14 X 18 26 X 6 9-10 68 25 81** 23190 41 16 - 19 25 X 7 10-9 70 26 82** 23190 Arroyo Seco 38 16 X 16 23 X 5 11-12 55 21 78** 11716 29 16 X 16 24 X 5 11-12 52 20 67** 11716 Socorro 47 12 X 20 28 X 7 10 30 88** 11714 56 14 X 20 26 X 7 9 65 30 96** 11714 39 13 X 20 30 X 5 7-8 53 27 82** 11967 53 15 X 18 25 X 6 9-10 40 27 92** 19%7 55 13 X 18 27 X 5 9-iO 60 26 94** 19967 E of Rosario 53 15 X 20 30 X 7 10 64 28 97** 23154 55 16 X 21 30 X 7 10-8 70 28 101** 23154 50 14 X 19 28 X 5 10-8 70 25 92** 23154 54 14 X 17 25 X 7 10 55 26 92** 19968 55 13 X 18 27 X 5 10 60 27 95** 19%8 shawii galdmaniana Punta Prieta 40 13 X 24 25 X 8 9-10 65 26 78** 19974 51 14 X 24 27 X 9 10-11 65 27 92** 19974 50 13 X 20 29 X 8 9-10 53 33 92** 19975 40 9 X 14 25 X 7 8 52 21 76** 19975 37 11 X 14 18 X 6 8 45 22 67** 19975 Tinaja Yubay 34 9 X 15 22 X 6 7 45 26 65** Harb. 35 11 X 15 24 X 6 8 42 26 71** Harb. Punta Prieta 50 10 X 21 25 X 8 8-9 47 24 85** 23161 49 11 X 20 25 X 9 9-10 58 24 84** 23161 34 9 X 17 21 X 8 7 50 26 65** 1985 34 9 X 18 TT X 8 6-8 50 25 65** 1985 W of San Borjas 57 16 X ->-> 24 X 10 12-13 55 24 96** 23166 41 12 X 23 23 X 9 10-11 70 24 78** 23166 45 12 X 23 26 X 9 8-9 75 26 82** 23166 36 10 X 18 24 X 8 8-6 46 23 70** 23166 San Andres 51 12 X 20 34 X 9 9-10 70 26 96** 42 50 11 X 20 29 X 8 8-9 70 21 90** 42 51 10 X 20 32 X 8 7-8 58 27 93** 42 Mesquital 33 11 X 17 19 X 7 9-10 42 20 63** 11935 34 10 X 17 21 X 7 8-10 42 20 64** 11935 29 9 X 17 18 X 7 6-8 45 19 56** 11937 26 9 X 17 15 X 7 5-7 42 18 50** 11937 sehastiana San Benito Is. 38 15 X 18 16 X 5 8-10 42 21 69* 17449 42 15 X 19 23 X 5 10-11 50 20 80* 17430 35 20 X TT 20 X 6 14-15 42 21 75* 17431 44 15 X 20 20 X 6 12 45 20 80* 17431 Natividad Is. 52 14 X 20 21 X 5 8-9 60 18 86* 15142 m). erect to decumbent, frequently branching from leaf axils; leaves mostly 20 x 8-10 cm to 50 X 20 cm, ovate to linear-ovate, short acu- minate, glossy light to dark green, cuticle slight- ly asperous, thickly fleshy, rigid, plane to slight- ly hollowed above, about as wide in mid-blade as at base; spine 2^ cm long, acicular, broad at base, openly grooved above, dark reddish brown to gray, straight or sinuous, decurrent for 8-10 cm or along entire leaf as corneous margin; GENTRY: AGAVES OF BAJA CALIFORNIA 89 UMBELLIFLORAE shawii shawii 23190 23190 11716 11714 19967 19967 23154 23154 19968 shawii goldmaniana 19974 19975 19975 Marb. 23161 1985 23166 23166 shawii goldmaniana 23166 DBG 1193S 11937 441 sebast iana shawii shawi 17449 17430 17431 15142 23190 shawii goldmaniana Figure 54. Floral ideographs of the Umbelliflorae. showing relative proportions of the tube (black) to outer tepul (white column), and level of insertion of filament (black square). Measurements are listed in Table II. teeth very variable in size, shape, in the mid- blade from 5 or 6 to 10-20 mm long, decreasing below, straight or variously flexed, rarely con- fluent or usually 1-2 cm apart, reddish to dark brown or dark gray; panicle 2^ m tall, the pe- duncle stout, closely imbricated by large, pur- ple, succulent bracts, closely investing the lat- eral umbellate branches; lateral branches commonly 8 to 14, horizontal to ascending, broad, stout, elliptic to oval in cross section, greenish red to purplish; flowers 75-100 mm long, sessile to short-pedicellate, bracteolate, closely clustered; buds frequently puiplish or red; ovary greenish 35-50 mm long, thick- fleshy, cylindric, with grooved, scarcely con- stricted neck 6-15 mm long, shortly tapered at base; tube amply funnelform, 12-16 mm deep, 16-22 mm wide, thick-fleshy, knobby at filament insertions with furrows between, light yellow, smooth, shiny, sinuses sometimes at unequal levels; tepals unequal, broad and thick at base, becoming linear-lanceolate, thinner, yellow or 90 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Figure 55. Agave shawii shawii near La Mision: (Left) Flowering rosette and {right) detail of the large, purplish, clasping bracts. Photographs by Bruce Gentry. reddish, soon involute and wilting while stamens extend, the outer tepal longer, 25 x 5 to 38 x 8 mm, rounded to plane on outer face, the inner with fleshy keel and thin involuting margins; fil- aments thick, strong, 60-70 mm long when ex- tended, inserted near mid-tube 8-12 mm above base of tube, tapered towards apex; anthers large, 20-28 mm long, usually centrically af- fixed; capsule variable in shape and size, 5.5 x 1.5 cm to 7 x 2.3 cm, obovoid or pyriform to oblong, slightly attenuate to rounded at base, thickly succulent when green and drying as a scurfy exocarp, with a short to long (1.5 cm) beak; seeds 7 x 5-6 mm, lunate to cuneiform, dull black to lustrous black. Type. — Hitchcock s.n. 20 miles [32 km] S of San Diego on the mesa, near the ocean, at the initial monument of the Mex- ican boundary, July 1875, MO [4 Ivs. fls.. 2 sheets]. Habitat and Characteristics . — The habitat of ssp. shawii is a narrow one along the maritime northwest coast of the peninsula. Subspecies shawii is a frequent component of the coastal sagebrush community with regular associates of Siminondsia chinensis , Rosa minutifolia , Opun- tia , Franseria , Atriplex , and other endemics like Aescidus parryi and Bergerocactus emoryi. Ten to twenty miles [16 to 32 km] inland shawii also contacts the higher chaparral community, where open slope and dispersed cover allow light for it to grow. Annual average rainfall ranges from GENTRY: AGAVES OF BAJA CALIFORNIA 91 ,^v«;ii<^4 ■^ a ^ ca 92 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 Figure 57. (Upper) Comparison of leaf series of (left) A^'iji f sliawii iioUtmuniana from the mid-peninsula, and (x\g\\l) Agave shawii shawii from near La Mision and east of El Rosario, the two larger indicating genes oi goldmciniana . (Lower) Agave shawii goldmaniana in massive population north of Punta Prieta in mid-peninsula. GENTRY: AGAVES OF BAJA CALIFORNIA 93 5 to 10 inches [130-250 mm], fogs and dew are frequent, solar intensity is reduced, and tem- peratures are equable, with mild and infrequent frosts (Fig. 52). The bright and gleaming rosettes are especially conspicuous on the low terraces and slopes overlooking the sea (Fig. 55). This habitat contrasts strongly with the more arid and interior habitat of ssp. goldmaniana (Fig. 58). These two subspecies are regarded as ecotypes of their respective habitats, slowly and presently evolving independently in their re- spective centers of area. The goldmaniana plants around Punta Prieta, for instance, are eas- ily separable from those of shawii about La Mi- sion north of Ensenada. They are not treated as species here because of my inability to separate the two along the southern perimeters of the shawii area, where the more robust forms of goldmaniana mingle with the more compact and dwarflike forms of shawii. This blending of forms is viewed as a natural and long-sustained potential o{ shawii to respond robustly to envi- ronmental factors in its southern area. The se- rious student should note, however, that this judgment is based on gross morphological con- siderations, not on direct genetic evidence. Se- ries of leaves of the two subspecies are shown in Figure 57. Exploitation of A. shawii. — Agave shawii fibers were probably used, and its meristem eaten, by the Indians (Barrows 1900). During the 1950"s there was a brief attempt to use the coarse fibers commercially. This project was featured in a San Diego newspaper, and I later observed where plants had been bulldozed out in piles for hauling to a decorticating machine. The fibers were rel- atively short and of poor quality; the oper- ation was apparently not commercially success- ful, and it was discontinued. Before 1950, plants were rarely seen as ornamentals in resident gar- dens, but since that time they are more com- monly planted in the beach developments. South of Ensenada thousands of acres with large stands of Agave shawii have been de- stroyed, along with the rest of the coastal sage- brush community, in clearing for new agricul- tural lands made possible by modern well drilling. Such developments together with the spread of urban growth along the beaches and environs will continue to eliminate the wild agaves. When the human density in the A. shawii area reaches the dimension now existing in coastal southern California, A. shawii may be considered an "endangered species." A flow- ering colony of A. shawii is particularly spec- tacular and pleasing to the human eye. This alone is sufficient to give modern man pause and to provide means of conserving this beautiful plant for the future. Agave shawii Engelm. ssp. goldmaniana (Trel.) Gentry, stat. & comb. nov. (Figures 3, 51-53. 54^, 57. 58; Tables 10, II) Af^ave goldmaniana Trel. Missouri Bot. Gard. Rep. 22:49. 1912. Like subspecies shawii. but rosettes medium to large; leaves longer, generally from 40 x 10 cm to 70 X 18 cm, more acuminate, lanceolate rather than linear-ovate; teeth also variable, commonly to 10-15 mm long in upper part of blade, sometimes paired or forked at tip, "fish- tailed." 1-3 cm apart and usually joined by a heavy corneous dark margin: spine very stout 3-4 cm long, very shallowly grooved above, brown to weathered gray; inflorescence 3-5 m tall with 18 to 25 massive laterals forming a deep wide panicle, pyramidal at top rather than rounded, peduncular bracts small and more re- mote than in ssp. shawii \ flowers greenish yel- low to red in bud, opening yellow, 65-96 mm long; ovary green, 30-50 mm long, the neck 5- 12 mm long and scarcely constricted; tube large, funnelform, thick walled, bulging and furrowed from tepal sinuses; tepals subequal, the outer 20-30 mm long, 7-10 mm wide in the middle, thick, linear, ascending, evenly rounded on out- er face, blunt and cucullate at tip, the inner thick- ly keeled, ca. 2 mm shorter, low-costate within; filaments inserted above middle of tube, strong, elliptic to oval in cross-section, reaching 45-70 mm in length; anthers 20-27 mm long; capsule large, thick walled, short-stipitate and short beaked, 1.6 x 5.5-6 cm to 2 x 8 cm; seeds 4 X 6 to 5 X 7 mm, with small hilar notch and narrow- winged margin (Gentry 10355). Type. — Nelson <& Goldman 7151 . Tinaja Yubay, Baja Cal- ifornia, 1905, US. This locality, about 20 miles [32 km] north of Punta Prieta, is a natural water trap, as the term "tinaja" implies, in an arroyo that floods through granitic rock. I was guided there in 1949 by a native resident of the area, whose name I have forgotten. It is a watering place known only to a few, and I found no indications of dwellings, recent or pre- historic. The name has various spellings in chronicles and on maps — "Ubi," "Yubay."" My guide refeired to it as Tinaja Yubay. The sand-scoured pockets in the hard bedrock were nearly filled with sand, and except for brief periods after heavy 94 OCCASIONAL PAPERS OK THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 C ON '^ o GENTRY: AGAVES OF BAJA CALIFORNIA 95 rains, one had to dig out the sand to reach the water. Nelson and Goldman camped and watered their stock there on Sep- tember 17-19, 1905, one of the few sources of sweet water in this beautiful area of the peninsula. Distribution and Habitat. — Agave shawii gold- maniana occupies a central part of the penin- sula, between latitudes 30°30'N and 28°30'N, or from Laguna Chapala to the vicinity of Mesqui- tal (Fig. 52). Elevations occupied range from 5 to 700 m. Except for a few tributaries that flow into the Bahfa de Los Angeles, it is limited to the Pacific slopes. Although there are gaps and irregularities, the population, by virtue of con- necting strands, is nearly continuous and very large for agaves generally. It occurs on both ig- neous and sedimentary rocks. The most concen- trated stands and best growth development ob- served are in the deep granitic soils south of Sierra Calamujue in the highland valley north and east of Punta Prieta, called by Goldman (1951:98) the Valley of San Andres. Here gold- maniana is a co-dominant form in the vegetation along with spectacular forests of Idria . Pachy- cereus. Yucca, a.nd Pachycormus . The aridity of this central highland is ameliorated by heavy nocturnal fogs moving in from the southwest from fall to spring, and which are indicated by the epiphytic lichens and Tillandsia that festoon the desert trees. The following notes were made among \\\i% Agave population on April 8, 1951, not far from the type locality, Tinaja Yubay. Populations and Variability . — Agave shawii goldmaniana is abundant, forming extensive colonies which are dense (up to 200-300 per acre) or thinly scattered (10-20 per acre). Large individual elongate heads or stems with leaves weigh up to 200-300 lbs (ca. 90-140 kg). Plants are caulescent with stems 3-8 dm long, with old leaves deflexing and covering stems. They are usually single, but frequently form rounded clus- ters, the older central rosette as much as 15 dm or more tall, the whole mound 2-3 m broad. They develop with branching from leaf axils. The whole in large specimens must weigh over 1,000 lbs (450 kg). They commonly form recli- nate trunks that branch repeatedly through the years from leaf axils. Old clones become rather open and cover several square meters (Fig. 3). The old dead trunks point the way back to the original plant or to where it was. The growing terminals, each tending to continue in one di- rection, now appear to be separate plants. Trunks in some clones are traceable back for 6-8 m, the axes being prone except for the as- cending leafy terminals. The plant is thus ac- tually tree size, but decumbent. Leaf is variable in form, but compared to A. shawii, is consistently more acuminate — a dis- tinguishing character. The leaf may be small or large, relatively narrow or broad, narrowed to- ward the base or scarcely so. Teeth are also highly variable and the spine only somewhat less so. The following leaf forms outline the variants, comparatively: Form 1 . The medium or more usual: Leaf acu- minate, narrow, mostly linear to slightly spatulate; teeth along the mid-blade 10-16 mm long, slender, curved or hooked, usu- ally 1-2 cm apart; spine slender, straight, often short. Plants mostly large. Form 2. Leaf broad, large, spatulate, cor- neous edge wide, teeth heavy, 10-15 mm long; spine thick; leaf short-acuminate. Few. Form 3. Leaf medium in size, between Forms 1 & 2, somewhat spatulate; corneous mar- gin and teeth heavy, 10-15 mm long; ro- settes medium size. Abundant. Form 4. Leaf short; rosettes small to medi- um; spine contorted; distal end of leaf wavy; teeth long and slender 20-30 mm long. Few. Form 5. Teeth very broad, often forked at tip, 15-25 mm long; rosettes small, blade spat- ulate. Few. Panicle is mostly 3-5 m tall with 20-30 lat- erals, well spread to form a long oval outline, in contrast to the subcapitate panicle of A. shawii. Bracts broad with buds bulging them, but small- er than those of A. shawii. Flowers numerous, large, yellow at anthesis, greenish in bud, and more yellow in age. Capsules variable, large, short oblong, or long obovoid being thickest just below apex. Flower form fairly constant. Summer rains falling through the southern part of the goldmaniana range in 1951 following 5-7 years of drought caused abundant flowering shoots in the late summer and fall. All of the earlier September flowers failed to set fruit, while October flowers in some cases appeared to be setting fruits. Lack of fruit set in early fall was obvious that year. However, in these same populations in previous years, fruit was ob- served to set well during the spring months. Is % OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 1.^0 lack of fruit set in fall due to: lack of pollinators, such as migrant hummingbirds; decreasing length of daylight; decreasing temperatures? Young flowering shoots in A. jJoUhtuiniana and A. shawii in October of 1951 were the most abundant ever observed. A cryptic member of the Deserticolae. A. avelkinidens replaces A. shawii goldmaniana south and east of Rancho Mesquital. The former can scarcely be distinguished from goldman- iana, so similar are the green rosettes, unless A. avellanidcns is in bloom. At that time its small- er, nearly tubeless flowers and narrow, deep panicles are evident. The small peduncular bracts and lack of axil branching in neighboring goldmaniana populations further complicate field identification. It is as though the goldman- iana population is infused with genes of single- rosette avellanidens that inhibit bract growth and axil branching — a speculative inference, of course. Food for Animals. — Leaf bases and hearts are eaten extensively by pack rats (Neotoma), who gnaw tunnels through the leaves around the head. Agave shawii was frequently noted with the terminal bud undermined and dying. Ravens apparently eat the young flowers. Claw marks, like those of fox or kitfox, on the peduncle showed how a climber had secured two or three young, tender, lateral branches by breaking them from the stalk. Cattle frequently chew the tops off young flowering shoots. Agave sebastiana (Figures 59. 60: Table II) Agave sebastiana Greene. Bull. Cal. Acad. Sci. 1:214. 1885. Agave shawii var. sebastiana (Greene) Gentry. Allan Han- cock Pac. Exped. 13:49. 1949. 'lAgave disjuncta Trel. Missouri Bot. Gard. Rep. 22:51. 1912. Rosettes medium to rather large, elongate, 6- 12 dm tall, closely imbricate, single or cespitose, glaucous gray or green; leaves generally 25^5 X 8-24 cm, broadly linear to ovate, short-acu- minate, thick rigid, plane to slightly hollowed above, rounded below, sometimes slightly nar- rowed towards base, light yellowish to grayish green, bud-printed, the margin usually cor- neous, dark brown; teeth slender, reddish brown, the larger through mid-blade 5-10 mm long, frequently down-flexed, 1-2 cm apart, or smaller and more numerous; spine stout, black to somewhat gray, 2-3 cm long, rarely shorter. variously grooved above; inflorescence 2-3 m tall, peduncle stout, with deltoid, scarious ap- pressed bracts; panicle short, wide-spreading, rounded to nearly flat, congested with 8 to 12 large umbels of yellow flowers in upper 14 of shaft; flowers 70-90 mm long, opening yellow, thick-fleshy; buds green; ovary 35-55 mm long, cylindric; tube broadly funnelform, 14-20 mm deep, 18-22 mm broad; tepals 16-25 x 5-7 mm ['"17-33 X 11-15"],* lanceolate, markedly cu- cullate and glandular floccose at tip, the inner shorter and strongly keeled; filaments 50-60 mm ["70""]* long, stout, inserted in mid-tube at 8-14 mm above bottom of tube; anthers 20-21 mm ["24-30""]* long, yellow (measurements from pressed dry specimens, see Moran numbers in Exsiccatae); capsules large, ca. 30 x 60-80 mm, beaked but scarcely stipitate; seeds large, glossy black, 7x11 mm. Type. — Greene s.n.. Cedros Island, I May 1885, CAS. Agave sebastiana is closely related to A. shawii as shown by the particulars of habit and flower morphology. However, sebastiana dif- fers significantly in the broader flatter panicle, in the smaller, more remote, scarious peduncu- lar bracts, and in the pale green, somewhat glau- cous leaves with more slender teeth. Bud print- ing and the black spines are more conspicuous on sebastiana due to the thin glaucous covering on the leaves. Reid Moran has provided an ex- cellent series of specimens from San Benito Is- lands (Fig. 59 & 60) together with a series of color slides. They show that sebastiana forms large clones with new rosettes starting from the bases of old stems. Moran 15142 from Natividad Island, southeast of Cedros Island, is a small form without corneous margins, the leaf only 15-20 cm long and pale green, the floral stem only 1 m tall. The flowers, however, are quite normal for sebastiana except for the long sta- mens. Moran noted it as "occasional, near mid- dle of Natividad Island."/!, sebastiana is a very handsome plant which should respond well in California gardens and, perhaps, elsewhere. Agave disjuncta Trel. was based on small live plants collected by J. N. Rose, reportedly on San Benito Island. Specimens of these plants are no longer in existence. Reid Moran informs me that he has searched the island for plants of this Moran measurements of fresh flowers. GENTRY: AGAVES OF BAJA CALIFORNIA 97 Figure 59. Agave sebastiana on Cedros Island with light-glaucous leaves. Photograph by Reid Moran. description without success. The name has no present application. Group Datyliones Trelease, Missouri Bot. Gard. Rep. 22:45. 1912. Acaulescent; leaves fibrous-fleshy, stiff, straight, dagger-shaped; spine strong, grooved above, scarcely decurrent; prickles heavily tri- angular; scape slender; panicles narrowly ob- long, with greenish tubular flowers; filaments deep-seated. Baja California Sur. Agave datylio (Figure 61) Agave datylio Simon ex Weber, Bull. Mus. Hist. Nat. Paris 8:224. 1902. Rosettes small to medium size, 6-10 dm tall, 10-15 dm wide, suckering freely, the rhizomes frequently elongate; leaves lance-linear 50-80 cm long. 3-4 cm wide, green to yellowish green, somewhat glaucous in youth, canaliculate above, rounded below, radiately spreading, rather rigid, nearly straight; margin nearly straight; teeth mostly 3-5 mm long, deltoid, flat- tened, rather blunt, dark brown, usually remote or 3-6 cm apart, more closely spaced below; terminal spine large, conical to subulate, 2.5^ cm long, dark brown to grayish, shortly decur- rent, scarcely or flatly grooved above; panicles 3-5 m tall with 8 to 15 branches of small umbels in upper half of shaft; flower greenish yellow, 40-55 mm long; ovary 20-30 mm long; tube fun- nelform, 5-10 mm deep; tepals 15-20 mm long, erect to ascending; filaments 35-45 mm long, in- serted 4-6 mm above base of tube; anthers 15- 20 mm long, yellow; capsules oblong to pyri- form, 1.5-2 cm in diameter, 3.5-4 cm long, short-stipitate; seeds 7x6 mm. Type. — Not designated; described from La Paz. Cape Dis- trict. A. datylio is widely scattered in the Cape region at lower elevations on granitic sandy soils (Fig. 61). Agave datylio has no close relatives on the peninsula. It belongs with the widespread sword-leafed group, Rigidae. whose nearest geographic member is A. aktites Gentry of the Sonoran-Sinaloan coast. The latter, however, is amply distinct with its narrower bluish leaves, sharply cuspid teeth, larger rotund stem, and 98 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. L^O Figure 60. Af^uve sebasiiana on East San Benito Island in the Ced^o^ group. An old clone with reclining trunks. Photograph by Raid Moran. larger flowers with deep bulging flower tube. Trelease set off A. datylio in a monotypic group, Datyliones, but I would presently favor its in- clusion in Berger's (1915) series Rigidae along with the A. angustifoUa alliance. However, no fresh flower material of A. datylio is presently available, and the whole sword-leafed group has yet to be carefully revised. A. datylio is locally called "datiliyo" or "da- tilillo," while its variety vexans at Comondii is called "mescaliyo." Agave datylio var. vexans (Trel.) 1. M. Johnston. Proc. Calif. Acad. Sci. ser. 4. 12:1003. 1924. Agave vc.xcins Trel. Missouri Bot. Card. Rep. 22:62. 1912. This variety differs from the typical form of the species in its smaller rosettes and leaves, ranging from 30-50 cm long, and in its more glaucous or yellowish color. Teeth are usually remote, but specimens from Arroyo Purisima have close-set, small teeth, the larger being only 2-3 mm long. Johnston separated the variety as having shorter filaments (20-30 mm), but this does not hold, as I have specimens from near Comondu with filaments 35-40 mm long, the same as for the typical form of the species. The variety occurs on sandy soils in lower elevations on both sides of the Sierra de la Giganta (Fig. 61). Variety vexans appears to be a xerophytic ecotype of the species. If grown together, the varieties may show themselves to be morpho- logically inseparable. I had them growing for many years in Murrieta. California, but they did not flower there, perhaps because of their shady situation. GENTRY: AGAVES OF BAJA CALIFORNIA 99 Figure 61. Agave datylio. (Upper) Near La Paz. (Lower) A. datylio ve.xan.s on sandy plain 23 miles (i of Comondu, showing the widely spaced off-sets. .i" km) soulhucst 100 OCCASIONAL PAPERS OF 1 HE CALIFORNIA ACADEMY OF SCIENCES. No. L^O 116» 114» 112» 110° 108" • @ ( ■ 1 1 ^® \ \ 34'- — ® i V" \ 45 / e,e^®\ - 34* t--^^© \ ) / " *' ' r^ \ I J 46-10.47 (64) @ __ApHOrNIX (SAN DIEGO I ^ v^ ^—^^^PHOENIX,, 1 l\ ""^"^sii^* ^ i* .-^-^^-^iy ^K I "yt-V"" '•■■■^ », *^^o-Aa, y~^~^~^^ -**^ ^/''^--'-v.^ — ^^^~x ^^ ^^ 32« \ensenada\M '•S§69) >--. ' N i j\ ', ©\ \S. ^^\. ,' -s WucsoN 1 - 32» v\-*-^^-''>^\-\isan Felipe ^S \ '''^\ .' .. n \'' 6 \> \ ' r ^ )? \i V /-''>-i 30< - y _ ' ^i^-'-'L ^^jy ROSARIoTv-ili) ;V V\ --^-Z ^V\ 1" 1 l°)Vvl4 \1^ )'^''''' ' J"" N K. 15\'' Nl6 V" ~^ ' ] A 30" W^'^ V "\ '-'V'""-"'' \¥n V^ 'k \ U-/ ) PUNTAV*V->--xtjoSj M) \ \ \ ,4 1 prietavX r^ C}\^^-~>"r -'"'' ^^^ l\V----'22 \ •• ,.>J iiS / >--' \ > "^ A 5\ 23\>V" \ /---. -; 28« S N. ^Calmalli s _„ \ X. ■ J ^^"v^^-^.g^,^../ V o^^s^i;^ ) ) , ;, ,^,^-^ y ^ - 28° "^''^"-' -.\— ^,_JN ,-- '' ^-^S-J.''^ ; ''"]' ,'"- ^Vwsv ^''"^ 11^ ^ * ^*~^'''\_,''^ V ' ' 1 ' ' ^^►©j^' rxA^'' K-'Y':'"! /-;' >P^ @ vv^ ^^ V^^^^ ^-^' ^^- ^\ ^^^^ 26" - ^S> '. i \ ^\\ j/ •' COMONDuV-A^--36|'^;7 ^( j' oV/ 37/H^O /,W^' - 26° ti \ \^ ''^^v. V -^ ,1 /Iz V ' -'' V# ^"^ \ ^^~ 7'' ''-'" ''Ji V \ \i;> %^^ / "^ 1 ^\ ^^ z' • ^J ,' \ 'y-' 24" - ""^^iX \WYa PAZ main roads ^V\ ~^^?a^\^ ^:; side roads and mule trails \ '"-^ 1 ^v^-. 24" \ v®l } CABO SAN LU CASW>^ km 0 100 200 22" 1 1 1 1 114'" 112* 110' 108» GENTRY: AGAVES OF BAJA CALIFORNIA 101 APPENDIX— Specimens that document the taxonomy, the distribution maps, and illustrations are enumer- ated in this index. Taxa, state names, and col- lectors" names are listed in alphabetical order. To facilitate these data for ready reference, the collectors" last names with their collection num- bers are listed first in italics, followed by the containing herbaria in capital letters (see Abbre- viations below). The localities with dates, and notes in brackets follow. Type collections and the herbaria containing holotypes are given in boldface. Curators having duplicates of these listed specimens can, therefore, easily identify their materials by using this reference. I have been unable to annotate all specimens in all her- baria, and some of my earlier identifications have been changed. Hence, this list of speci- mens will bring identifications up to date, pro- vided curators use these listings. You all would EXSICCATAE do agaveology well to assign an assistant to this chore, as soon as this monograph is received. It is not usual to include dates of collection with exsiccatae citations, but I have done this to document flowering times. The plant parts composing the collections are listed as abbre- viations (see table below) in parentheses, as well as my comments on some collections. Locality citations are edited copies from collectors" la- bels; their appended notes are frequently given in quotes. Altogether, the data represent a rec- ord of the living agave populations and should be useful to future students. The map of Figure 62 locates many of the little known localities listed in the Exsiccatae. Some ranches are not on maps. Other Mexican place names are frequently duplicated in a single area, such as Agua Caliente and San Felipe. Names of mountains are circled. I have found no map Figure 62. Outline of the principal travels of Howard Scott Gentry in the Gulf of California region ( 1938-1975) and some reference localities of agave collections. Baja California 1. La Mision 2. Sierra Juarez 3. San Matias Pass 4. Valle de la Trinidad 5. Sierra San Pedro Martir 6. Rancho Agua Caliente 7. Meling Ranch 8. San Telmo 9. Socorro 10. San Fernando 11. Sierra San Miguel 12. San Agustin 13. Catavina 14. El Marmol 15. Jaraguay 45. Pinyon Flats 46. San Felipe Ranch 47. San Felipe Creek 48. Yaqui Wells 58. Burro Creek 59. Hillside 60. Harcuvar Mountains 61. Cunningham Pass 70. Yavaros 71. Quitovac 72. Puerto Libertad 73. Punta Tepopa 16. Bahi'a San Luis Gonzaga 17. Laguna Seca Chapala 18. Sierra San Luis 19. Tinaja Yubay 20. Bahia de Los Angeles 21. Isla Angel de la Guarda 2~> San Borja 23. Paraiso 24. Rancho Mesquital 25. San Andres 26. Islas San Benitos 27. Isla Cedros 28. Bahia Tortuga 29. Cerro Tordillo 30. Sierra Vizcaino California 49. Borrego Springs 50. Carriso Creek 51. Jacumba 52. Mountain Springs Arizona 62. Indian Springs 63. Harquahala Mountains 64. Kofa (S. H.) Mountains 65. Little Horn Mountains SONORA 74. Isla Tiburon 75. Turner's Island 76. Seal Island 77 Caborca 31. Picachos de Santa Clara 32. Isla Magdalena 33. Isla Margarita 34. Sierra de las Palmas 35. Rancho San Sebastian 36. Loreto 37. Mision San Javier 38. Sierra de la Giganta 39. Isla San Marcos 40. Isla Carmen 41. Bahia Concepcion 42. Isla Espiritu Santo 43. Sierra Laguna 44. Rancho Burrera 53. Providence Mountains 54. Whipple Mountains 55. Dad Granite Mountains 56. Ivanpah Mountains 66. Sand Tank Mountains 67. Tinajas Altas 68. Cabeza Prieta Mountains 69. Tule Mountains 78. Santa Ana 79. Hermosillo 80. Ciudad Obregon 81. Guaymas 102 OCCASIONAL PAPERS OF THK CALIFORNIA ACADEMY OF SCIENCES. No. L^O that locates Sierra de las Palmas (34), yet it is a prominent extension of the Sierra de la Giganta with a local montane vegetation of o'dk-Nolina grassland. Notable additions to the peninsular flora have been found there, e.g., A^ave ^i^an- tensis and Sediim alamosanum from the main- land. The main road through the peninsula rep- resents the old unpaved auto trail. However, on a scale of this ratio, it was necessary to forego close accuracy in detail. Abbreviations for Herbaria A = Arnold Arboretum, Harvard University, Cambridge, Massachusetts. ASU = Arizona State University, Tempe, Ari- zona. ARIZ = University of Arizona, Tucson, Ari- zona. BH = Bailey Hortorium. Cornell University, Ithaca, New York. CAS = California Academy of Sciences, Gold- en Gate Park, San Francisco, California. DES = Desert Botanical Garden, Phoenix. Ar- izona. DS = Dudley Herbarium, now in CAS. GH = Gray Herbarium, Harvard University, Cambridge, Massachusetts. MEXU = Instituto de Biologia. Universidad Nacional Autonoma de Mexico. MICH = University of Michigan, Ann Arbor, Michigan. MO = Missouri Botanical Garden, St. Louis, Missouri. NA = National Arboretum Herbarium, Wash- ington, D.C. POM = Pomona College, Claremont, Califor- nia. SBBG = Santa Barbara Botanic Garden, Santa Barbara, California. SD = San Diego Museum of Natural History, San Diego, California. UC = University of California, Berkeley, Cali- fornia. US = National Herbarium, Natural History Museum, Washington, D.C. Gentry Herbarium specimens are presently on deposit in the Desert Botanical Garden Herbar- ium and are included in that listing, DES. Parenthetical Abbreviations br = bract cap = capsule f = flower 1 = leaf photo = photograph Agave aurea Baja California Barclay & Arguelles 1988, DES, MEXU, US. 5 miles [8 km] N of Todos Santos along road to La Paz, 20 Apr. 1966 [1, f]- Brandegee s.n., UC, DS. Type. Purisima, 13 Feb. 1889 [margin of large 1, small 1, 5 f, pieces of 2 cap. Has typically close-set, regular teeth on regular teats forming undulate margin, as growing on lava mesas about Comondii and E of Purisima]. Brandegee s.n., MO. Cape Region mountains, 20 Sept. 1899 [1, f, mixed with f of A. salvia; type of A. brandegeei Trel.]. Carter 5132, UC. Mesa de San Geronimo, N from Rancho Viejo [on road from Loreto to San Javier], 8 May 1966; alt. ca. 1,110 m. Carter 5484 , UC. Arroyo de Puerta Vieja on road from Loretto to Comondii, Sierra de la Giganta, 4 July 1970; alt. ca. 1,400-1,500 ft [ca. 430-460 mj. Carter 5779 , UC. Mesa de Humi, a mesa on crest of Sierra de la Giganta opposite N end of Isla San Jose, 20 Mar. 1973; alt. ca. 750 m. Gentry 11299, DES. Polymorphic population on mesa NW of Comondii [photo]. Gentry 12341, DES, MEXU, US. 5 miles [8 km] N of Todos Santos, Cape District, 22 Nov. 1952 [1, f]. Gentry 12375, DES. MEXU, US. 3 miles [5 km] N of Comondii, 30 Nov. 1952 [t]. Gentry 10321, DES, MEXU, US. Comondii, 2 Apr. 1951 [1, f, cap]. Gentry 11253, DES, MEXU, US. Ca. 10 miles [ca. 16 km] W of San Jose del Cabo, 5 Oct. 1951 [1, photo]. Gentry 11198, DES, US. Ca. 12-15 miles [ca. 20-25 km] E of La Paz, Cape District, 29 Sept. 1951 [1, photo]. Gentry 4272, ARIZ, DES, DS, US. Cerro de la Giganta, 1 Mar. 1939 [f]. Volcanic slopes & crags up to 4,000 ft [ca. 1,200 m]. Gentry 12383, DES, MEXU, US. Ca. 3 miles [ca. 5 km] N of Mision San Javier. Sierra Gi- ganta, 2 Dec. 1952. Gently & Cech 11255, DES. MEXU, US. Las Cuevas, Cape District, 7 Oct. 1951 [1, photo]. Gentry & Cech 11283, DES, MEXU, US. 35 GENTRY: AGAVES OF BAJA CALIFORNIA 103 miles [ca. 56 km] NW of La Paz. 11 Oct. 1951 [1. photo]. Gentry & Cech 11295, DES, MEXU. US. 3 miles [ca. 5 km] NW of Comondii, 18 Oct. 1951 [1, photo]. Gentry & Cech 11301, DES, MEXU, US. 10 miles [ca. 16 km] W of Canipole along road to Purisima [1, photo]. Gentry & Gentry 23182, DES. MEXU. US. 3 miles [ca. 5 km] N of Comondii on lava fields, 10 Apr. 1973 [1, f, photo]. Harbison s.n., SD. Mesa 2 miles [ca. 3 km] N of Comondii; elev. 400 m. 7 Oct. 1967. Harbison s.n., SD. 6 miles [ca. 10 km] N of Todos Santos. Moran 7144, CAS. DS. MEXU. SD. Cape District. 4 km N of La Huerta. 400 m. 25 Jan. 1959 [\,f]. Piirpus s.n., MO. UC. San Jose del Cabo. Jan. -Mar. 1901 [1. f]. Wiggins 14470, CAS. DS. MEXU. 100 miles [ca. 160 km] W of Los Planes, alt. 490 m. south- facing slopes. 21 Dec. 1958 [1, f]. Wiggins 14531, CAS. DS. MEXU. UC. 6.4 miles [10.3 km] N of Todos Santos along road to La Paz, 25 Dec. 1958 [1, f]. Agave aveilanidens Baja California Brandegee s.n., UC. Type. Parafso, 1 May 1889 [upper '/2-% of leaf, flowers, and old caps. All similar to what I have seen and col- lected between Calmalli and Mesquital Rancho]. Gentry & Fo.x 11944, DES. MEXU. US. Ca. 9 miles [ca. 30 km] E of Punta Prieta. 19 May 1952. Gentry & Fox 11933, 11932, DES, MEXU. US. 6 miles [ca. 10 km] W of Calmallf. 17 May 1952. Gentry & Fox 11929, DES. MEXU. US. 6 miles [ca. 10 km] W of Calmalli. 17 May 1952 [ 1. f. photo]. Gently & Gentry 23184, DES. MEXU. US. 51/2 miles [ca. 9 km] W of Calmalli. 12 Apr. 1973 [1. f, photo]. Gentry & Gentiy 23186, DES, MEXU, US. 15-17 miles [ca. 24-27 km] SE of Mesquital, 12 Apr. 1973 [1, f]. Gentry & Gentry 23187, DES, MEXU, US. 7 miles [ca. 1 1 km] SE of Mesquital along road to El Arco, 12 Apr. 1973 [1, f]. Hammerly 69, CAS. 29 miles [ca. 47 km] N of Mesquital, 27 Sept. 1941 [doubtfully referred here]. Wiggins 5726, DS. Between Calmalli and Mesquital Rancho on mesas, 31 May 1931 [1,11. Agave capensis Baja California Barclay & Arguelles 1987, MEXU, US. Vi- cinity of Cabo San Lucas, 19 Apr. 1966. Brandegee s.n., UC. Cabo San Lucas, 18 Mar. 1892 [1, cap]. Gentry 10080, DES, MEXU, US. Huntington Botanical Gardens, San Marino, California, 9- 15 Jan. 1951 [1, cap, photo]. Gentry 19676, DES, MEXU, US. Huntington Botanical Gardens, 17 Apr. 1962 [1, f]. Gentry & Fox 11247, 11250, DES, MEXU, US. Type. Cabo San Lucas & vicinity, 5 Oct. 1951 [l,f, photo]. Gentry & Fox 11823, DES, MEXU, US. 3 miles [ca. 5 km] N of Todos Santos. Cape Dist., 4 May 1952. Agave cerulata cerulata Baja California Brandegee s.n., DS. CardcSn Grande (between San Ignacio & CalmallO, 22 Apr. 1889 [1, f. cap]. Carter & Kellogg 2953, UC. Isolated red hill in sandy plain. 18.2 km W of Mision Santa Ger- trudis. 18 Dec. 1950. Ferris 8576A, DS. 1 mile [ca. 1.6 km] S of Laguna Seca Chapala, 6 Mar. 1934 [1, cap]. Gentn- 10346, DES, MEXU, US. Calmalli. between mine and houses of town. 6 Apr. 1951, type locality. Gentry 10359, DES. MEXU. US. Ca. 7 miles [ca. 11 km] S of Tinaja Yubay and 15 miles [ca. 24 km] NEofPunta Prieta; 8 Apr. 1951 [Leap]. Gentry 10369, MEXU, US. Rancho Jaraguay, 9 Apr. 1961 [1, early f, eaten by cattle]. Gentry 11188, DES, MEXU, US. 4 miles [ca. 6 km] NW of Laguna Seca Chapala. 21 Sept. 1951 [1, cap]. Gentry 19973, DES. MEXU, US. 10 miles [ ca. 16 km] S of Laguna Seca Chapala, 29 Apr. 1963 [1, f, photo]. Gentry & Cech 11322, DES, MEXU, US. 4 miles [ca. 6 km] NW of Laguna Seca Chapala, 24 Oct. 1951 [1, photo]. Gentry & Fox 11919, 11921, 11924. DES, MEXU, US. 6 miles [ca. 10 km] W of Calmalli, 17 May 1952. Gentry & Fox 11953, DES, MEXU, US. 21 miles [ca. 34 km] E of Punta Prieta on road to Bahfa de Los Angeles, 20 May 1952 [1, photo]. 104 OCCASIONAL PAPERS OF THK CALIFORNIA ACADEMY OF SCIENCES. No. 130 Gentry & Fo.x 11961, 11962, DES, MEXU, US. 4 miles [ca. 6 km] NW of Laguna Seca Cha- pala. May 1952 [1. photo]. Gentry & Gentry 23159, DBS, MEXU. US. 20 miles [ca. 32 km) SE of San Agustin, elev. 2.100 ft [ca. 640 m], 5 Apr. 1973 [1. early t]. Gentty & Gentry 23185, DES, MEXU. US. 6V2 miles [ca. 10 km] W of Calmalli, 12 Apr. 1973. Gentry & McGill 23298, DES. MEXU. US. 10 miles [ca. 16 km] S of San Luis Gonzaga Bay along road to Laguna Chapala, 17 June 1973 [1, f]. Gentry & McGill 23302, DES. MEXU, US. W. of Sierra Calamajue. ca. 30 miles [ca. 48 km] N of Punta Prieta, 17 June 1973 [1. f. photo]. Gentry & McGill 23306, DES. MEXU, US. 16-20 miles [ca. 26-32 km] NE of Punta Prieta along road to Bahia de Los Angeles, 17 June 1973 [1, f]- Gentry & McGill 23307, DES, MEXU. US. Ca. 40 miles [ca. 65 km] NE of Punta Prieta along road to Bahia de Los Angeles, 17 June 1973 [1, f]- Gentry & McGill 233 14, DES. MEXU. US. 6- 8 miles [ca. 10-13 km] S of road fork from Los Angeles Bay along road to San Borja. 18 June 1973 [1. f]. Harbison s.n.. SD. 12 miles [ca. 19 km] E of Calmalli. 8 Apr. 1947 [l.tl Harbison s.n., SD. 20 miles [ca. 32 km] S of Punta Prieta, 9 Apr. 1947 [1, inflo]. Harbison s.n. , SD. Agua Amarga, ca. 15 miles [ca. 24 km] W of Los Angeles Bay, 15 Apr. 1947 [Kf]. Johnston 3487. 3489, CAS, GH, SD, UC, US. Los Angeles Bay, 6 May 1921, "in small groups on rocky mountainside," [yellow leaves with brown-ringed teeth, 1, f, cap]. Johnston 3405 a-g, CAS, US. Angel de la Guarda Island, Palm Canyon, 3 May 1921 [a se- ries of yellow leaves, mostly with reduced teeth, brown-ringed, and some toothless], "gregarious on hillside. " Moran 4106, BH. DS. Motherless Island. Los Angeles Bay. 10 May 1952 [1, cap]. Moran 2007, DS, UC. 16 miles [ca. 26 km] N of Pimta Prieta, elev. ca. 1,700 ft [ca. 520 m], 22 Apr. 1946 [1, f]. Moran 8167, DS, SD, UC. 3 miles [ca. 5 km] E of El Arco, elev. ca. 250 m, 5 Apr. 1960 [1, f]. Moran 8185, SD. Arroyo Estaton, Isla Angel de la Guarda. 15 Apr. 1960; elev. ca. 25 m [f, cap, doubtfully assigned here]. Nelson & Goldman 7180, US. Type. Calmalli. elev. 800 ft [ca. 240 ml. 29 Sept. 1905. [Leaves and flowers characteristic of what I have seen through middle Baja California; no problem here.] Raven et at. 12631 , UC. 1.6 km S of Rancho Santo Ignacito. elev. 560 m. 21 Apr. 1958. Stover & Harbison s.n., SD. 35 miles [ca. 56 km] N of Punta Prieta, 5 May 1939 [1, f]. Thomas 7972, SD, US. 1 mile [ca. 1.6 km] NW of Pozo Aleman, elev. ca. 800 ft [ca. 240 m], 26 May 1959 [1, bud]. Wiggins 5721 , DS. 15 miles [ca. 24 km] NW of San Ignacio. 30 May 1931 [1, f]. Wiggins 5724, DS, UC, US. Calmalli. 31 May 193 ui. tl Wiggins 5734, DS. 5 miles [ca. 8 km] N of Punta Prieta. 1 June 1931 [1, f]. Wiggins & Wiggins 14882, CAS. DS. S end of Isla Ventana. Bahia de los Angeles, near beach, 18 May 1959 [1. cap]. Agave cerulata dentiens Baja California Bostic s.n., SD. San Esteban Island, sandy arroyo near SE corner. 21 June 1965. Johnston 3194, CAS. UC. San Esteban Is- land. 20 Apr. 1921. "common in small colonies on hillsides," [has brown-ringed teeth like cer- ulata, 1, cap]. Moran 4079, SD. San Esteban Island. 6 May 1952. Moran 21748, SD. San Esteban Island, arroyo near E side. Apr. 1975, "large colonies on hill- sides and in arroyo" [in bud]. Rose 16819, US. Type. San Esteban Island, 12 Apr. 1911 [1, cap]. Agave cerulata nelsonii Baja California Gentry et al. 10370, 11155, 11162. 11164, 11165. 11665, 11666, DES, MEXU, MICH, US. 2-3 miles [ca. 3-5 km] N of San Fernando, Sier- ra San Miguel, type locality, 9 Apr. 1951, 10 Sept. 1951, 10 Apr. 1952 [1, f, cap]. Gentry et al. 10376, DES, MEXU, US. 18 miles [ca. 30 km] E of Rosario, 10 Apr. 1951. Gentry et al. 11178, DES, MEXU, US. 28 miles [ca. 45 km] E of Rosario on Sierra San Miguel, 13 Sept. 1951. Gentry et al. 11179, DES, MEXU, US. 17 miles [ca. 27 km] E of Rosario. 14 Sept. 1951 [ 1, n. GENTRY: AGAVES OF BAJA CALIFORNIA 105 Gentry et al. 11185, DES. MEXU, US. Ca. 4 miles [ca. 6 km] SE of San Agusti'n. elev. ca. 2,000 ft [ca. 3,200 m], 21 Sept. 1951 [1, f]- Gentry & McGill 23311, DES, MEXU, US. 4 miles [ca. 6 km] S of Los Angeles Bay road fork on road to San Borja. elev. 900 ft [ca. 275 m], 18 June 1973 [1, f]. Gentry & McGill 23315, DES. MEXU. US. 10 miles [ca. 16 km] N of San Borja. elev. ca. 1 .700 ft [ca. 500 m]. 18 June 1973 [1, f]. Gentry & McGill 23322, DES. MEXU. US. Ca. 5 miles [ca. 8 km] N of San Fernando, Sierra San Miguel, elev. ca. 1.750 ft [ca. 530 m], 22 June 1973 [1, f]. Gentry & McGill 23324, DES, MEXU. US. 11 miles [ca. 18 km] E of Rosario. Rancho Por- venir. elev. 450 ft [ca. 140 m]. 24 June 1973 [1. f]. Harbison s.n., SD. 1 mile [ca. 1.6 km] W of Rancho Arenoso; elev. ca. 500 m (?). Moran 22643, SD. S of Rancho San Miguel [Sierra San Miguel Range], elev. ca. 900 m. 9 Aug. 1975 [f]. Moran & Reveal 22064, SD. Ridge 3 miles [ca. 5 km] SW of San Isidro. 30°44'N, 115°34' W. elev. ca. 1.120 m. 20 July 1975. "occasional in chaparral" [1. f]. Nelson & Goldman 7111, US. Type. San Fer- nando [Sierra San Miguel], alt. 1,400 ft [ca. 425 m], 4 Sept. 1905 [1, old f]. Agave cerulata subcerulata Baja California Barclay & Arguelles 1991, DES, MEXU, US. Ca. 5 miles [ca. 8 km] W of San Ignacio. 29 Apr. 1966 [1. f]. Gentry 10330, DES. MEXU. US. Type. San Ignacio. 3 Apr. 1951 [1. f. cap. inflo]. Gentry 11892, DES, MEXU, US. Arroyo de la Teneria, Isla San Marcos. 13 May 1952 [1, f, cap. photo]. Gentry & Fo.x 11926, DES. MEXU. US. 6 miles [ca. 10 km] W of Calmalli. 17 May 1952. Gentry & Gentry 23170, DES. MEXU. US. 10 miles [ca. 16 km] W of San Ignacio. 8 Apr. 1973 [1, f, photo]. Gentry & Gentry 23175, DES, MEXU. US. 24 miles [ca. 40 km] E of San Ignacio along road to Santa Rosalia. 9 Apr. 1973 [1. f]. Harbison No. P, DES. Cuesta de las Vir- genes, 1972. Johnson 3649, 3650, CAS. GH, SD, UC. US. San Marcos Island, on gypsum. 12 May 1921 [1, f. cap]. Pinkava & McGill P12287 , ASU. DES. Ca. 70 miles [ca. 110 km] W of Santa Rosalia. Route 1. 30 May 1974. Butte base [1. fj- Agave datylio Baja California Brandegee s. n. CAS. La Paz. Cape District. 4 Nov. 1891 (l.f). Brandegee 581, UC. San Pedro. 29 Oct. 1891 [1, f, cap]. Brandegee s. n. UC. Paseo de los Dolores to Lake Ramon, 4 Apr. 1889 (1, cap). Gentry 11200, DES. MEXU. US. About 4 miles [ca. 6 km] E of La Paz, Cape District. 29 Sep. 1951 [l.f, photo]. Moran 3553, SD. Ensenada de los Muertos, Cape District, 1 Apr. 1952 [1, cap]. Peters 124, UC. Los Planes, Cape District, arroyo bottom, elev. 300 ft [ca. 90 m]. 27 Mar. 1948. "mescal." Rose 1302, US. La Paz. 14 June 1897. Wiggins 11501 , DS. 9 miles [ca. 15 km] W of La Paz. Wiggins 15475, CAS, DS. MEXU. Rancho del Obispo [Magdalena Plain], alt. ca. 150 m, 15 Nov. 1959 [1 with long blade of A. datylio and remote teeth of vexans]. Agave datylio var. vexans Baja California Brandegee s. n. UC. Purisima?, 1899 [infl]. Gently 10302, DES. Rancho Panales, Arroyo Purisima, 31 Mar. 1951 (1. f. cap). Rocky sedi- mentary slope; cardon-pitaya-Bursera-Larrea- Opuntia-etc. Gentry 4322, DES. Comondu. 10 Mar. 1939 (f). Hill slope, common over coastal plain and foothills. Gently & Cech 11292, DES. 23 miles [ca. 37 km] SW of Comondu. 16 Oct. 1951 (1, photo). Sandy valley bottomland. Hastings & Turner 64-377, ARIZ, CAS. 6 miles [ca. 10 km] by road W of San Luis Gonzaga (NW of La Paz), elev. 400 ft [ca. 120 m], 21 Oct. 1964, flowers yellow with green tinge [1, f\. M. E. Jones 23751, MO. La Paz, 15 Nov. 1926 [f]. Rose 1302, US. La Paz, 14 June 1897. Rose 16540, US. La Paz, 29 Mar. 1911. Wiggins 15205. CAS. Mesa 2 miles [ca. 3 km] N of Arroyo San Gregorio. 27 Oct. 1959 [1. f]. 106 OCCASIONAL PAPERS OF IHl- CALIFORNIA ACADEMY OF SCIENCES. No. 130 Agave deserti deserti Baja California Gentry s.n., DES. Granitic sandy highland on Mex. Rt. 2, S of Jacumba. Apr. 1963 [photoj. Gentry & Arguelles 22990, DES, MEXU, US. Valle de Trinidad, elev. ca. 2.600 ft [ca. 800 m]. 3 May 1972 [1. f]- Gentry & McGill 23285, DES. MEXU. US. 17-18 miles [ca. 27-29 km] W of San Felipe along road to Valle Trinidad, elev. ca. 1,950 ft [ca. 600 m], 13 June 1973 [I, f]- Gentry & McGill 23286, ASU. DES. MEXU. US. 2.6 miles [ca. 4 km] SE of Rancho Agua Caliente on E bajada of Sierra San Pedro Martir, elev. ca. 1,450 ft [ca. 440 m], 13 June 1973 [1, f]. Hastings & Turner 66-6. ARIZ. 12.6 miles [ca. 20 km] W of turnoff toward San Matias Pass, elev. 1,300 ft [ca. 400 m], 4 Oct. 1966 [1, cap]. Hutchison 710, UC. Between Alaska and Mexicali, km 140, E slopes of the sierra, 31 Dec. 1952 [flowered UC Botanical Garden, Aug. 1963; karyotype n = 59, Cave]. Pinkava & McGill 8648, ASU, DES. Along Rte. 2, 38.5 miles [ca. 62 km] W of junction with main route to Mexicali, 7 June 1971 [1, f, br]. Wiggins & Wiggins 16044, DS. Granitic sandy bajada 16 miles [ca. 26 km] W of San Felipe Hwy. along road to San Matias Pass, 2 Apr. 1960 [1, f]. California Abrams 3976, DS. Between San Felipe & Ca- risso Creek, San Diego Co., 4 July 1903 [1, fj. Ball & Everett 22777, UC. Ca. 3 miles [ca. 5 km] NE of Banner Trading Post, State Hwy. 78, 6 Nov. 1957 [1, cap]. Ball & Everett 22671. UC. 6.6 miles [ca. 11 km] SW of Palm Village Hwy. junction 1 11 along Hwy. 74, elev. 1,650 ft [ca. 500 m], 20 Aug. 1957 [toothless 1 and cap]. Barr 67-211, 67-210, ARIZ. 40 miles [ca. 65 km] W of El Centro near Ocotillo, 13 May 1967; elev. 1,700 ft [ca. 520 m] [I, f|- Cleveland 7218, SD. Jacumba. 25 June 1885 [f]. Crovello 295, UC. 2.6 miles [ca. 4 km] E of Vallecito Stage Station, San Diego Co., elev. 2,300 ft [ca. 700 m], 15 Dec. 1963 [f]- Eastwood 18639, CAS. Road to Mountain Springs, San Diego Co.. 25 Apr. 1932. Ferris & Rossbach 9685, DS. Borrego Springs, P. O. road and State Hwy. 78, San Di- ego Co., May 1938 [1, f]. Ferris 7060, DS. Between Jacumba & Mt. Springs, San Diego Co., 18 Apr. 1928 [1, cap]. Flemming 753, SD. Foot of Banner Grade, Sentenac Canyon, [San Diego Co.] Apr. 1926. Gander 1325, SD. Carriso Creek. [San Diego Co.] 3 Jan. 1936 [1. fj- Gander 4810, SD. Bull Willow Canyon. San Diego Co., 15 Dec. 1937 [1, f]- Gentry 10034, 10034a, DES, MEXU, MICH, US. 3-4 miles [ca. 5-6 km] S of Palm Desert, Santa Rosa Mt.. 20 Dec. 1950. Gentry 10041. 10044, DES. MEXU, MICH, US. Yaqui Wells near San Felipe Creek, San Diego Co., 29 Dec. 1950 [1, cap]. Gentry 10051, DES. MEXU. US. Pinyon Flats, Hwy. 74, Riverside Co., elev. ca. 4,000 ft [ca. 1,200 m], 5 Jan. 1951 [1, f, cap]. Gentry 17759, DES, MEXU, US. Pinyon Flats, [Riverside Co.], 7 July 1959. Gentry 19741 , DES, MEXU. US. San Felipe Ranch near Rt. 78 and road to Warner's Ranch, San Diego Co., 25 May 1962 [1, f[. Gentry et al . 11650. 11652, DES, MEXU, MICH, US. Pinyon Flats, [Riverside Co.], 3 Apr. 1952. Gentry & McGill 23326, DES, MEXU, US. Pinyon Flats, [Riverside Co.], 26 June 1973. Hall 21 17, DS, UC, US. E base of San Jacinto Mts., Colorado Desert. June 1901 [1. f]. Hitchcock & Palmer in 1875. Emory in 1846, MO. "'The types." E of San Felipe Ranch, San Diego Co. [Probably along San Felipe Creek, above or below The Narrows.] Howell 3243, CAS. San Felipe Wash, halfway from Borrego Valley to Yaqui Wells, San Diego Co., 26 Nov. 1927 [1, f, cap]. Huey s.n.. SD. Borrego Narrows, [San Diego Co.], Apr. 1931 [f]. Lester s.n.. SD. San Isidro Mt., N of Borrego, May 1926. Mearns 2972, US. E base of Coast Range, edge of Colorado Desert, 7 May 1894. Mearns 3109. 3025. 3147. DS, US. Mountain Springs, San Diego Co., May 1894 [1, inflo]. Palmer 462, 88, GH. Type collection in part. San Felipe Canyon. San Diego Co., 1875 [inflo]. Vasey 626, US. Mountain Springs grade. [San Diego Co.], June 1880 [1. f]- Woglum 156, SD. Mountain Springs grade [San Diego Co.], 7 May 1936. Wolf 9455, UC. Palms to Pines Hwy. 74, 2.7 miles [ca. 4 km] below Dos Palmos Spring, Riv- erside Co., 29 Sept. 1939 [1, fl- GENTRY: AGAVES OF BAJA CALIFORNIA 107 Woodcock s.n., SD. Foot of Banner Grade. Sentenac Canyon [San Diego Co.], 10 Apr. 1929 m. Yates 5459, UC. 5 miles [ca. 8 km] W of San Felipe Canyon. San Diego Co., elev. 1.500 ft [ca. 460 m], 8 Apr. 1936 [1, f, cap]. Agave desert! pringlei Baja California Broder547, 473, DS. 3 miles [ca. 5 km] WNW of Santa Catarina, 25 May 1961. elev. ca. 4,000 ft [ca. 1,200 m] [1, f. In Sierra Juarez]. Gentry 10287, DES, MEXU, US. Northwest end of Sierra San Pedro Martir, 23 Mar. 1951. Gently 16723, DES. MEXU, US. Near San Matias Pass, 22 June 1957 [1, f, photo]. Gentry 19959, DES, MEXU, US. San Matias Pass, 23 Apr. 1963 [1, f, photo]. Harbison s.n., SD. Near E! Progreso, Sierra Juarez, elev. ca. 1,500 m: 32°18' N, 115°54' W, 1 Aug. 1965 [1, f]. Hastings & Turner 66-17, ARIZ. San Matias Pass, 19.1 miles [ca. 31 km] E of Valle Trinidad, 5 Oct. 1966, elev. 1,950 ft [ca. 600 m] [1, cap]. Moran 9838, SD, UC. 5 miles [ca. 8 km] SE of Las Filipenas, Sierra Juarez, elev. ca. 1,620 m, 30 June 1962 [1, f]. Moran 9849, SD. Just E of San Matias Pass, elev. ca. 1,020 m, 30 June 1962 [1, f, caps]. "Rosettes clustered, to 1.2 m. of ca. 50 leaves: leaves green, channeled, curved, to 7 dm long, floral stem 3-5 m tall ...."" Moran 15256, SD. Rancho San Pedro Martir, Sierra San Pedro Martir, elev. ca. 1.700 m, 5 July 1968 [1. f|. •• Rosettes l-V/i m. of ca. 50 green or slightly glaucous leaves 4-8 dm long. Occasional in chaparral." Moran 18639, SD. 2 miles [ca. 3 km] NE of Alamito, Sierra Juarez, elev. ca. 1 . 150 m. 3 Oct. 1971 [1. f]. Moran 21983, SD. Sierra San Pedro Martir, 30°57' N, 115°36' W, elev. ca. 1,600 m, "on met- amorphic rock in small arroyo. mile NW of oak pasture" [1. cap]. Moulis & McGill 555, ASU, DES. Sierra San Pedro Martir, ca. 18.5 road miles [ca. 30 km] E of Meling Ranch. 21 Aug. 1972 [1, f, cap]. Orcutt s.n., UC. Hanson's Ranch, 29 July 1883. Orcutt s.n., DS. Lower California, 1892, ex. herb. Pringle. [Sierra Juarez.] Orcutt s.n. K, MEXU. Type. Sierras Cen- trales, elev. 6,000 ft [ca. 1,830 m], 7 Oct. 1882 [Leap]. Agave deserti simplex Arizona Engard s.n. , DES. Near Indian Springs. Har- cuvar Mt., Yuma Co., June 1974 [f only]. Gentry 23404, DES, SD, US. Type. N slope of Harquahala Mt.. 12 miles [ca. 19 km] W of Aguila, Yuma Co., 12 June 1974 [1 of 5 & f of 4 pis.]. Gentry 20590, DES. MEXU, US. Near a sheep tank in Cabeza Prieta Mts. [Yuma Co.], 3 May 1964. Gently & Engard 23562, ASU, DES, US. S end of Silver Bell Mts., Pima Co.. 17 June 1975. Limestone, elev. 3,000-3,500 ft [ca. 900-1,070 m], [1, n. Gentry & Ogden 9947, MEXU, US. N. slope Harquahala Mt., elev. 3,500-5,000 ft [ca. 1,000- 1,500 m], 10 Nov. 1950 [1, f]. Gentry & Ogden Photo, DES. Cunningham Pass, Harcuvar Mts.. Yuma Co.. 14 Nov. 1950. Gently & Weber 23410, DES. Sand Tank Mt., Yuma Co.. 27 July 1974; rocky volcanic slope, elev. 2,800-3.000 ft [ca. 850-900 m], [1, cap, 2 live]. Goldman 2310, US. Tinaja Altas, Yuma Co., 20 Nov. 1913. Harbison 4312 , ARIZ. SD. Canyon leading to Del Oso Pass, Kofa Mts.. Yuma Co., 8 June 1943 [The largest leaf seen of this species, with large teeth] [I, f]. Harrison et al. 7303, ARIZ. Table Top Mt., 16 Aug. 1930 [1, cap]. [Doubtfully referred here.] M earns 305, US. Tule Mts., Mexican bound- ary line, 11 Feb. 1894 [1 looks like A. zebra]. Peebles & Smith 14415, ARIZ. GH, US. Sier- ra Estrella, Maricopa Co., elev. 2,500 ft [ca. 760 m], 12 July 1939 [1, cap]. Peebles & Smith 13873, 13878, 13881, ARIZ. GH, US. Cunningham Pass. Harcuvar Mts.. Yuma Co.. 17 May 1938 [1, L cap]. Van Devender s.n., ARIZ. Along Hwy. I 8 in Telegraph Pass, Gila Mts.. N slope, elev. ca. 800 ft [ca. 250 m], 31 Dec. 1972 [1, cap]. Weber Photo, DES. Summit of Little Horn Mts., May 1970. Weber & McGill 2549, ASU. DES. Vi miles [ca. 0.8 km] S of Sheep Tank Mine. Yuma Co., Ariz., 1 June 1970 [1, f]. West s.n., ARIZ. 18-Mile Drive. Organ Pipe 108 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Cactus National Monument, Pima Co., 27 May 1962 II, n. Wigi^ins 8643, ARIZ. DS, GH, UC, US. Rocky canyon near small tanks ca. 10 miles [ca. 16 kml S of Hwy. 80 on E foot of Mohawk Mts., Yuma Co.. 1 Mar. 1937 [1, cap — Atypical elon- gate leaf; flowers should be collected]. California Blukley 3231A, DES. Above Mitchell's Cav- erns, Providence Mts. [flowered Santa Barbara Botanic Garden, 22 June 1972]. Bnindegee s.n., UC. Providence Mts., 31 May 1902, ''plants solitary." Ferris & Bacigalupi 8161, DS. Canyon above Bonanza King Mine, Providence Mts., San Ber- nardino Co., 27 Apr. 1932. Jaeger s.n., SD. 20 miles [ca. 32 km] NE of Amboy, E end of Granite Mts.. [San Bernardino Co.], 26 Mar. 1929. Munz et al. 4302, US. Vicinity of Bonanza King Mine, E slope of Providence Mts.. Mohave Desert, ah. 3.500-5.000 ft [ca. 1.070-1,500 m], 21-24 May 1920 [cap]. Shaw s.n., DES. Ivanpah Mt. (Riley's claim). San Bernardino Co., Spring, 1974 [1. cap]. Van Devender et al. 74-29, ARIZ. S slopes of Whipple Mts.. San Bernardino Co.. elev. 1,600 ft [ca. 500 m], 17 Feb. 1974 [1. f]. Wolf 10183, DS, US. N side of Old Dad-Gran- ite Mt. Range, Snake Spring, Mohave Desert, elev. 4.500 ft [ca. 1.400 m]. 30 Apr. 1941. Wolf 3165, ChS,, DS. UC. From Copper Basin on road to Parker Bridge, Whipple Mts., E Col- orado Desert, San Bernardino Co., elev. 1,400 ft [ca. 400 m], 30 Apr. 1932 [1, cap, buds]. Wolf & Everett 9023, CAS. DS. UC. US. Same locality as Wolf 3165, 29 July 1937 [1, cap]. SONORA Gentry 21203, DES, MEXU, US. Km 2509 in mountain pass SE of Qui'tovac, 4 Sept. 1965, elev. 1,800-2,000 ft [ca. 550-600 m], [1, cap]. Agave gigantensis Baja California Barclay & Arguelles 1990, MEXU. US. Mountains above Rancho San Sebastian. 28 Apr. 1966. Carter & Reese 4552, UC. Cuesta de las Par- ras just above Rancho de las Parras. road be- tween Loreto & San Javier. 5 June 1963. alt. ca. 350 m. Gentry 7693, 7713 [pars.] ARIZ. DES. MEXU. SD. UC, UM. Picachos de Santa Clara, 5-10 Nov. 1947 [1, n. Gentry 10339, 10342, DES, MEXU, US. Pi- cachos de Santa Clara, Vizcaino Desert, 4-5 Apr. 1951 [doubtfully assigned here]. Gentry & Arguelles 10327. 10324, MEXU, US. Between Llano San Julio & Sierra de la Giganta, 2 Apr. 1951 [1, f. photo]. Gentry & Fo.\ 11778. DES, MEXU. US. Sier- ra de las Palmas above Rancho San Sebastian. 27-29 Apr. 1952 [1. cap]. Gentry & McGill 23320, DES. MEXU, US. Type. Sierra de las Palmas, above Rancho San Sebastian, 20 June 1973, elev. ca. 4.000 ft. [ca. 1.200 m]. Johnston 3843, CAS. US. Puerto Escondido, 29 May 1921 [1. f]. "single plant in wash." Agave margaritae Baja California Beauchanip 2149, SD. Magdalena Island, on slopes along arroyos N of Punta Magdalena, elev. ca. 5 m. Apr. 1971 [1, cap]. Beauchamp 2109, SD. West end of N side of Margarita Island, elev. ca. 50 m, 6 Apr. 1971. Brandegee s.n., UC. Type. Magdalena Island, 14 Jan. 1889 [1, f, cap]. Gentry, Fo.x, Arguelles 11903, 11905, DES, US. Cienegita. Isla Santa Margarita, on sandy bajada or plain, 15 May 1952. Moran 3540, BH. Santa Maria Bay. 31 Mar. 1952 [1. f]. Moran 4187, BH. Man-of-War Cove. Mag- dalena Bay, 21 May 1952 [1, cap]. Rose 16261 , US. Santa Maria Bay. Isla Santa Magdalena, 18 Mar. 1911 [type of Agave con- nochaetodon Trel.]. Agave mckelveyana Arizona Bezy 286, ARIZ. 8.3 miles [ca. 13 km] W of Hillside. [Yavapai Co.]. elev. 3.900 ft [ca. 1.200 m]. 8 June 1964 [tl. Braem s.n., DS. Dean Mountain Rd.. Huala- pai Mts., 6 Oct. 1935 [1, caps]. Breitung 18156, DS. McClout Mts., ca. 18 miles [ca. 13 km] N of Congress, [Yavapai Co.], 23 Aug. 1959. Eastwood 18387, CAS. Aquarius Mts. [Mo- have Co.], 13 May 1931 [1, bud]. Eastwood s.n., CAS. Coyote Pass between Oatman and Kingman, May 1931. Gentry 21979, ARIZ, DES, US. Type. Sit- GENTRY: AGAVES OF BAJA CALIFORNIA 109 greave Pass in Black Mts., ca. 4 miles [ca. 6 km] NE of Oatman. 26 June 1966. Gentry 23000, DES. MEXU, US. 26 miles [ca. 42 km] SE of Wikieup along Route 93, Mo- have Co.. elev. 3,400 ft [ca. 1,040 m], 10 June 1972 [I, f]. Gentry 23002, DES, MEXU, US. 6 miles [ca. 10 km] E of Route 93 along road to Burro Creek, Mohave Co., elev. 3.600 ft [ca. 1,000 m], 10 June 1972. Gentry & Ogden 9981, DES, MEXU, US. Near Oatman, elev. 3,000-3,200 ft [ca. 900-975 m], low bush cover with scattered juniper and yucca in igneous mountains, [Mohave Co.], 15 Nov. 1950 [I, cap]. Gentry & Whitehead 22312 . ARIZ, DES, US. Hualapai Mts., along road to Recreation Park from Kingman, elev. ca. 5,000 ft [ca. 1,500 m], 1 July 1967 [I, first f]. Harrison & Kearney 7303, ARIZ. Table Top Mt., Mohave Co., 16 Aug. 1930 [I, cap]. Haskell, s.n., ARIZ. Near Gold Road, [Mo- have Co.], 6 Dec. 1941 [1, cap]. Jones 25167, POM. Near Oatman, [Mohave Co.], 1930 [in part, cap, mixed with leaf of A. utahensis]. Kearney & Peebles 12570, ARIZ. Rocky hills S of Hillside, Yavapai Co., elev. 3,600 ft [ca. 1,100 m], 17 Sept. 1935, "scape 8 ft. high (other plants with scapes twice as tall, but leaves al- ways small)" [1, cap]. McKelvey 2235, 1515, 1653. A. West of Burro Creek, Aquarius Mts., [Mohave Co.], 14 May 1931 [1, f, cap]. McKelvey 2250, A. Black Mts., between Kingman and Oatman, [Mohave Co.], 16 May 1931 [fin fluid]. McKelvey 4056A, A. Camp Verde region, 23 Apr. 1934 [1, cap]. McKelvey 4078, 4080, A. Juniper Mts. [Ya- vapai Co.], Apr. 1934. McKelvey 1648, 1649, 4071 , A. Between Kirk- land and Hillside, [Yavapai Co.], 29 Mar. 1930 [whole plant with cap]. Stephens s.n. , UC. Hualapai Mts., 5 July 1903 [Kf]. Agave moranii Baja California Chambers 629, DS. Canyon del Diablo to N and W of Picacho del Diablo, E flank of Sierra San Pedro Martir, 6 miles [ca. 10 km] from Can- yon mouth, elev. 4,500 ft [ca. 1,400 m], 17 June 1954 [f, cap, bract]. Gentry & McGill 23287, DES, MEXU, US. Type. 2-3 miles [ca. 3-5 km] SE of Agua Cal- iente, on E plain of Sierra San Pedro Martir, 13 June 1973 [1, f]- Moran 18295, DES, MEXU, US. SE side of San Felipe Valley [Sierra Santa Rosa?] elev. ca. 530 m. 9 Mar. 1971. Moran Photo, SD. Cerro Chato, S side of Sierra San Pedro Martir, elev. ca. 1,800 m, 3 June 1963. Moran 21562, SD. Sierra San Pedro Martir, Arroyo del Cajon ca. 2 miles [ca. 3 km] from the mouth, near 30°5r N, 115°16' W, elev. ca. 810 m, 28 Dec. 1974. Agave promontorii Baja California Barclay & Arguelles 1986, DES, MEXU, US. Western summit of Sierra Laguna, 15 Apr. 1966 [l.f]. Brandegee s.n., UC. Sierra de la Laguna, 24 Apr. 1892 [I, n. Brandegee s.n., UC. San Jose del Cabo, cul- tivated in San Diego, 1903. Gentty 10164, DES, MEXU, US. Huntington Botanical Gardens, San Marino, California, 9- 15 Jan. 1951. Gentry 11218, DES, MEXU, US. Rancho La- guna and vicinity. Sierra Laguna, Cape District, western summit, 3 Oct. 1951. Gentry 11229, DES, MEXU, US. Rancho Burrera at west base of Sierra Laguna, 1-4 Oct. 1951. Gentry 19671 , DES. Sierra Laguna, Cape Dis- trict, 1952 [photo]. Moran 7451, CAS, SD. La Aguja, elev. 1,900 m. Cape District, 18 May 1959 [I, f, cap]. Nelson & Goldman 7437, US. Type. From San Bernardo to El Sauz, Sierra La Laguna, 21 Jan. 1906, alt. 2,400-5,000 ft [ca. 700-1,500 m], [I. f. photo]. Agave sebastiana Baja California Anthony 264, DS. K. US. San Benito Island. Mar. -June 1897 [I, f, fine-toothed form]. Anthony s.n., CAS. San Benito Island, Mar.- June 1897 [t^. Beauchamp 2095, SD. West San Benito Is- land, 4 Apr. 1971, elev. ca. 130 m. Beauchamp 3193, SD. Isia San Benito Oeste, 28 Feb. 1972. Common on slopes, floral stem 4 feet high [fj. 110 OCTASIONAI. PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 BeUinii s.n.. CAS. May 1881 [fj. Benedict s.n.. SBBG. Cedros Island, upper Campo Punta. Norte Canyon, easterly from Cerrodel Norte, 15 Mar. 1971. Benedict s.n. , SBBG. West San Benito Island. 200 yards [ca. 180 m] S of fishing village. SE portion of island, 9 Mar. 1971 [side flowering shoot]. Brandei>ee s.n., UC. San Benito Island. 27 Mar. 1897 [I. f, br]. Brandegee s.n., UC. San Benito Island, 1 Apr. 1897 [1. f. br]. Greene s.n., CAS, DS, UC. Type. Cedros Is- land, 1 May 1885. [leaves 25-30 cm, lanceolate, broadest in middle with regular teeth mostly curved upwards, 2 cm apart, 5-7 mm long, spine slender 20-30 mm long, openly grooved above, flowers 65-67 mm long, 3 1, f]. Howell 10691 , CAS. SE side, Cedros Island, 16 Aug. 1932. Mason 1986. CAS, K, US. Cedros Island, 3 June 1925 [f, cap]. Moran 2954, CAS. Cedros Island. Momn 4198, BH, DS. West San Benito Is- land, 24 May 1952 [1, f — leaf linear-ovate with close-set, slender teeth, reduced below mid- blade, on continuous margin]. Moran 15142 , SD. Natividad Island, near mid- dle, elev. ca. 100 m, 24 June 1968 [1, f], "Floral stem 1 m high, plant smaller than usual. Fls. bright yellow, erect, visited by flies." Moran 17430. 17431, DES, SD. West San Benito Island, elev. ca. 100 m, 19 Apr. 1970 [1, f]. Moran 17449, SD. East San Benito Island, elev. ca. 25 m, 20 Apr. 1970 [I, f]. Leaves gray to gray-green. Moran 21206, DES, SD. Cedros Island, 2 miles [ca. 3 km] from N end, elev. ca. 600 m, 28 Mar. 1974 [I, f]. "Fl. stem 3 m. with 10 branches in upper 3 dm." Moran 19924, SD. East shore of Bahia Tor- tugas, elev. ca. 10 m. 8 Feb. 1973 [I, f]. Philhrick B75-31, SBBG. West San Benito Island, 20 Jan. 1975. W facing slope, halfway between terrace & summit [1, br, bud]. Philhrick & Benedict B72-77 , SBBG. Arroyo Madrid, above Playa Madrid, N of Colorada, SE portion of Cedros Island, 20 Apr. 1972 [I, f]. Soils s.n., MEXU. Isia Cedros, 3 June 1925 [l,f|. Agave shawii goidmaniana Baja California Barclay & Argnelles 1985, MEXU, US. Ca. 25 miles [ca. 40 km] N of Punta Prieta. 6 Apr. 1966 [1, f]- Ferris 8581 , DS. W of San Borjas Range (Sier- ra Calamajue), 12 miles [ca. 19 km] from Laguna Chapala along road to Punta Prieta, 6 Mar. 1934 [1, f, caps]. Gentry 10349, MEXU, US. Near Marmolito, 7 Apr. 1951. Gentty 10361. 10365. 10366. 10355, MEXU, US. Ca. 7 miles [ca. 11 km] S of Tinaja Yubay and 15 miles [ca. 24 km] NE of Punta Prieta, 8 Apr. 1951 [I, f, cap]. Gentry 10379. 10381. 10382, DES. MEXU, US. Six miles [ca. 10 km] N of Socorro, 1 1 Apr. 1951 [1. f]. Gentry 11318, 11319. 11320, MEXU, US. Ca. 10 miles [ca. 16 km] N of Punta Prieta, 24 Oct. 1951 [anal., photo]. Gentry 11935, MEXU. US. Ca. 5 miles [ca. 8 km] NW of Mezquital, Vizcaino Desert, 18 May 1952 [1, f, inflo, photo]. Gentr}.' 19974. MEXU. US. Valley W of Sierra Calamajue, 25 miles [ca. 40 km] N of Punta Prie- ta, 30 Apr. 1963 [I, f, photo]. Gentry & Fox 11948, DES, MEXU, MICH, US. Ca. 15 miles [ca. 24 km] NE of Punta Prieta. 20 May 1952 [1, f, cap]. Gentry & Gentry 23161 , DES, MEXU, US. 16 miles [ca. 26 km] NE of Punta Prieta, 5 Apr. 1973 [f]. Gentry & Gentry 23166, DES, MEXU, US. 8 miles [ca. 13 km] W of Mission San Borja. 7 Apr. 1973 [I, f, photo]. Harbison s.n., CAS. Near Tinaja Yubay, 30 Apr. 1964 [1, f]. Harhison s.n. , SD. Four miles [ca. 6 km] S of San Andres, 25 July 1941 [I, f]. Harbison s.n., SD. Punta Prieta, 29 Apr. 1940 [i.n. Hastings & Turner 63-231 , DS. Arroyo Agua- jito. 14.9 miles [ca. 24 km] E of Rosario, elev. 700 ft [ca. 200 m], 19 Oct. 1963 [I. f, cap]. Lindsay DES 42, DES. Near San Andres, 1930"s [I, f, cap]. Moran 17027, DES. SD. 7.6 miles [ca. 12 km] N of Puerto Santa Catarina, 28 Mar. 1970; "Only this one plant seen in the Santa Catarina drainage" [I, f, cap]. GENTRY: AGAVES OF BAJA CALIFORNIA in Moran 17053, SD. 6 miles [ca. 10 km] E of Punta Canoas, elev. ca. 50 m. 29 Mar. 1970. "Common but only one still flowering" " [1. f. cap]. Moran 17121 , SD. Puerto San Jose. elev. ca. 25 m. 30 Mar. 1970 [1. f]. Moran 17191, SD. 8 miles [ca. 13 km] S of Las Palomas. elev. ca. 140 m. 1 Apr. 1970. "Rosettes 13 dm wide, of 125 leaves. 45 x 12 cm'- [1. f]. Moran 17204, SD. Boca de Marron, elev. ca. 5 m. 2 Apr. 1970. "Common on rocky hillside. Rosettes 12-14 dm wide of 200 leaves'" [1. f]- Nelson & Goldman 7151, VS. Type. Yubay, 30 miles [ca. 48 km] SE of Calamajue, B.C., alt. 2,000 ft [ca. 600 m], 18 Sept. 1905 [1, cap, photo]. Wiggins 4475, DS. Between EI Marmol and Rosario, 40 miles [ca. 64 km] E of Rosario, 12 Mar. 1930 [1, f]. Wiggins & Thomas 171, DS, US. About 14 miles [ca. 23 km] toward the coast from Cerro Blanco (SE of Rosario), ah. ca. 1,200 ft. [ca. 350 m], 8 Feb. 1962 [1, f]. Agave shawii shawii Baja California Arnott 12, 21, UC. 4 miles [ca. 6 km] N of Socorro, 21 Apr. 1955 [f]. Brandegee s.n., UC. Colnett, May 1893 [1, f]- Cox s.n. , UC. San Telmo de Abajo. May 193 1 . Farmer s.n., SD. Santo Tomas Valley, 24 Dec. 1955 [series of leaves]. Ferris 8524, DS. 16 miles [ca. 26 km] from Colnett Wash on Santo Domingo Road. 2 Mar. 1934 [1. n. Ferris 8528, 8529, DS, US. 30 miles [ca. 48 km] N of Rosario on road to Santo Domingo. 3 Mar. 1934. Gentry 4001 , UC. Between San Vicente and Hamilton Ranch, 10 Nov. 1938 [1, f]. Gentry 10079, MEXU, US. Huntington Bo- tanical Gardens, 9-15 Jan. 1951 [1, f]. Gentry 10281, MEXU, US. Km 57 S of Ti- juana, 21 Mar. 1951 [1, f]. Gentry 10285, MEXU. US. Ca. 10 miles [ca. 16 km] NE of San Telmo, 23 Mar. 1951 [1. f]. Gentry & Arguelles 19968, MEXU. US. 15 miles [ca. 24 km] E of Rosario. 28 Apr. 1963 [t]. Gentry & Gentry 23154, DES. MEXU. US. 12-13 miles [ca. 19-21 km] E of Rosario. 4 Apr. 1973 [1, f, photo]. Gentn- & Gentry 23190, DES. MEXU. US. Near La Mision. 27 miles [ca. 43 km] N of En- senada by power station, old road. 15 Apr. 1973 [1, f, photo]. Harbison 45522, SD. Santa Maria Valley, 31 Aug. 1953. Harbison s.n., SD. 7 miles [ca. 1 1 km] SE San Quintin, 21 Apr. 1927 [1. f]. Harbison s.n., SD. San Simon (ca. 10 miles [ca. 16 km] E of San Quintin). 10 Sept. 1955. Harbison s.n., SD. Camalu Point, 29 Dec. 1949 [1, f. cap June 1939]. Harbison s.n., SD. Arroyo ESE of Rosario, 29 Dec. 1949 [leaves narrow with very fine or small teeth. 1-2 m long & very narrow corneous margin]. Harbison s.n., SD. Canyon above Hamilton Ranch. 15 Dec. 1953 [1, tl. Harbison s.n., SD. La Mision Point. 30 Dec. 1949 [1. f]. Harbison & Howe s.n., SD. Arroyo 8.4 miles [ca. 14 km] E of Rosario. elev. ca. 200 m, 23 Sept. 1965. Harding s.n., SD. Halfway between Ensena- da and Tijuana. 28 Dec. 1937 [br. f]. Jones s.n., DS, MEXU. West of Tijuana near the sea, 26 Dec. 1924 [f]. Moran 16711, DES, SD, UC. Jatay, (S of La Mision), elev. ca. 100 m [f]. "Abundant on coastal terrace 1 mile S of Jatay."" Wiggins 21. 442, DS. Valle de San Telmo. 15 miles [ca. 24 km] E of Hwy. No. 1. 17 June 1971 [1. f. insects & hummingbirds]. Wiggins & Gillespie 3911. 4006, CAS. DS. MEXU. US. 37 miles [ca. 60 km] S of Tijuana. 2 miles [ca. 3 km] S of "Halfway House."" 8 Sept. 1929. California Eastwood 2940, CAS. Boundary Monument, San Diego Co., 24 Apr. 1913 [1, f, topotype]. Gander 494, SD. Near Boundary Monument 258, on shore of Pacific, San Diego Co.. 28 Jan. 1936 [1. f]. Hall 3957, UC. SW corner of San Diego Co.. Calif., 30 Apr. 1903 [cap]. Harbison s.n., DES. International Monu- ment. San Diego Co., Calif., [topotype]. Hitchcock in 1875, MO. Type. Initial Bound- ary Monument, S of San Diego. Moran 16707, DES, SD. UC. West side of Point Loma [tl 100 m S of Cal- Western Cam- 112 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 pus, 18 Dec. 1969. "Perhaps 200 rosettes, brink of sea bluffs." Moran 21699, SD. Torrey Pines, San Diego Co., 29 Mar. 1975; "'uniform spination — may be one clone — it can be considered native." Srovcr s.n., SD. Point Loma, San Diego Co., 21 Apr. 1937 [1, f]- Wolf 5412, CAS. 8 ft. N of Mexican Boundary and second small hill from ocean, San Diego Co., elev. 30-40 ft [ca. 9-12 m], 31 Aug. 1933, topotype. Wolf 2660. DS. Rancho Santa Ana Botanical Garden from plant brought from SW San Diego Co., 15 Feb. 1932 [1. f, br]. Agave sobria frailensis Baja California Gentry & Cech 11264, MEXU, US. Type. 4 miles [ca. 6 km] northward of Punta Frailes, 7 Oct. 1951 [l.f]. Gentry & Cech 11257. MEXU, US. 5-8 miles [ca. 8-13 km] N of Punta Frailes, 7 Oct. 1951, [1, photo]. Gentry & Fox 11858. MEXU, US. 5-8 miles [ca. 8-13 km] N of Punta Frailes, 6 May 1952 [1, f, cap]. Agave sobria roseana Baja California Brandegee s.n.. UC. La Paz, 14 Apr. 1892. Gentry 11274, MEXU, US. Islote Gallo, 10 Oct. 1951 [1, f, cap]. Gentry & Cech 11277. MEXU, US. A W ridge of Isla Espi'ritu Santo, Golfo de California, 10 Oct. 1951 [1, photo]. Gentry & Fox 11869. MEXU, US. Ca. 3 miles [ca. 5 km] E of La Paz, 7 May 1952 [1, f, photo]. Hastings & Turner 64-1 54b. SD. 6 miles [ca. 10 km] N of La Paz. Johnston 3989, 4001. UC, US. The Isthmus, Espiritu Santo Island, 31 May 1921. Johnston 4002, 4003. UC, US. San Gabriel Bay, Espiritu Santo Island, 1 June 1921 [1, f]. Rose 16854, US. Type. Espiritu Santo Island, 18 Apr. 1911. Wiggins 17828. DS. Isla Partida, just N of Isla Espiritu Santo, 20 Apr. 1962. Agave sobria sobria Baja California Barclay & Arguelles 1989. MEXU, US. Vi- cinity of San Miguel de Comondii, 26 Apr. 1966, topotype [1, f]. Brandegee s.n., DS, UC. Type. Comondii Mesas, 23 Mar. 1889 [1, margins, br, cap, f]. Brandegee s.n., MO, UC. Cape region moun- tains, 20 Sept. 1899 [f, type of A. brandegee! , mixed with A. aurea]. Carter 5486, UC. Cumbre de la Cuesta de Las Parras, north of road, alt. ca. 1 ,750 m, road from Loreto to San Javier, 5 July 1970. Carter 5487, UC. Cuesta S of Arroyo Ligui, south of Loreto, 8 July 1970. Carter & Sharsmith 4940, UC. Along trail from San Jose de Agua Verde to Bahia Agua Verde, on Gulf drainage, alt. ca. 360 m, 4 June 1965. Gentry 10304, MEXU, US. Arroyo Puiisima several miles above Purisima, 31 Mar. 1951. Gentry 10308. MEXU, US. Arroyo Purisima above Purisima, 1 Apr. 1951. Gentry 11303. DES. Ca. 10 miles [ca. 16 km] W of Canipole [photo]. Gentry 12382. MEXU, US. Ca. 3 miles [ca. 5 km] N of San Javier, Sierra Giganta, 2 Dec. 1952. Gentry 12387. MEXU, US. E side of Bahia Concepcion, Rancho Salto, 3 Dec. 1952. Gentry & Cech 11291, DES, MEXU, US. Comondii, N slope of volcanic rim, 15 Oct. 1951 [1. photo]. Gentry & Fox 11811. MEXU, US. Rancho San Andreas, Sierra de las Palmas, S of Santa Rosalia, 27-29 Apr. 1952. Gentry et al. 11876. MEXU. US. 2-3 miles [ca. 3-5 km] NE of La Paz, 7 May 1952. Gentry et al. 11882. DES, MEXU, US. Com- ondu, 10 May 1952 [cap]. Harbison s.n., SD. Danzante Island. 7 Apr. 1962 [1, f, cap, very long ovaries]. Harbison s.n., SD. Top of grade on road from Comondii to Loreto, 6 Oct. 1967. Hutchison 7399, 7473, SD. 13 miles [ca. 21 km] S of EI Coyote, Concepcion Bay {Vi mile N of km 85) May & Jan. 1975. Johnston 3857, CAS, US. Ballenas Bay, Dan- zante Island, 24 May 1921 [1, f, cap — leaf and its armor similar to A. roseana]. Johnston 3887, CAS, UC, US. Agua Verde Bay, 26 May 1921. Moran 3936, BH, DS. Canyon S of Balandra Bay, Carmen Island, 18 Apr. 1952 [1, f]. Purpus s.n., MO, UC. San Jose del Cabo, Jan. — Mar. 1901 [f only, mixed with A. aurea]. Wiggins 11446, UC. 6 miles [ca. 10 km] W of Canipole, 17 Nov. 1946 [1, different form]. GENTRY: AGAVES OF BAJA CALIFORNIA 113 Agave subsimplex SONORA Felger2721 , ARIZ. Isla Cholludo (Roca Fuca. Seal Island), 30 Oct. 1958, "common and wide- spread, suckering, leaves bluish; inflo. branched, only one plant in fl.: anthers yellow, filaments and tepals dusty rose-colored; base of tube pink- ish white" [I, fj. Felger & Bezy 14171. ARIZ. Foothills at N side of Cerro Tepopa, 4 miles [ca. 6 km] by road W of 9 miles [ca. 14 km] by road S of Desem- boque, 15 May 1966 [f]. Felger & Bezy 14182, ARIZ. 6.3 miles [ca. 10 km] S of Desemboque San Ignacio, middle ba- jada with sandy soil, 14 May 1966, "few scat- tered colonies, leaves faintly cross-banded, te- pals white to pinkish" [f]. Gently 4486, DES, ARIZ. Puerto Libertad, 22 May 1939 [f]. Gentry 10217, DES, MEXU, US. N end of Sierra Coloral, 28 Feb. 1951 [1, cap, pi]. Gentry 10221. 10222, 10223, DES, MEXU, US. Cerro Punta Tepopa, 1 Mar. 1939 [I, cap, pl]. Hastings et al. 63-8, ARIZ. Punto Cirio, about 7 miles [ca. 11 km] S of Puerto Libertad, 3 Jan. 1963 [I, cap — small leaf with big teats]. MacDoitgiil & Shreve s.n. , ARIZ. Kino Point. 17 Nov. 1923. Moran 13022, SD. UC. Turner's Island, elev. ca. 25 m, "common on rocky slopes." Tepals light yellow or reddish, filaments pink to dark red. 25 Apr. 1966 [I. f]- Rose 16811, US. Type. Seal Island, just off Tiburon Island. 13 Apr. 1911. Whiting 9047, ARIZ. Tiburon Island, vicinity of Tecomate. 14-22 June 1951. "abundant on rocky mountain slopes on W side of island. This one from NW corner." Agave vizcainoensis Baja California Gentry 7469, ARIZ, DES, DS, MEXU, MICH, UC. Type. Cerro Tordillo. Sierra Viz- caino, 12-13 Mar. 1947, elev. 400-800 ft [ca. 120-240 m], [I, f]. Gentry 7713, 7693 ARIZ, DES, DS, MICH, UC, UM. Picachos de Santa Clara, Vizcaino Desert. 5-10 Nov. 1947 [I. cap. Doubtfully as- signed here, in part.]. Howell 10660, CAS. San Bartolome Bay ( = Bahi'a Tortuga). 14 Aug. 1932 [I, f, s]. 114 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 GLOSSARY OF ahii.xial. the side of a lateral organ away from the central a.xis. acaulescent , stemless or without visible stem below the leaves. iickular. needle-shaped. adaxial. the side of a lateral organ next to the central axis. adventitious buds , those produced in areas with- out visible bud initials, as from the stem in- stead of the axils of the leaves. allopatric. applied to allied species or popula- tions inhabiting separate geographic areas. Compare sympatric. antrorse , directed forward or towards the apex, as the prickles on a leaf margin. arcuate , moderately arched or curving. ascending , of leaves pointed upward and out- ward at about 20° or more from the horizontal. attenuate, gradually narrowed or prolonged. bud printing , of leaves when the margin of one leaf is impressed upon the surface of the next leaf. bulbiferous, producing bulbils. bulbil, small plant reproduced vegetatively in the axils of the inflorescence. A form of asex- ual reproduction. cabeza, Spanish, head, applied to the thick, short stem of agaves. campanulate , bell-shaped. castaneous , chestnut-colored. caulescent , having a stem or trunk below the leaves. cespitose, as applied to succulents, growing in clusters by the production of basal branches, suckers, or offsets. chartaceous , papery, dry, and thin. circadian (L. circa, about, and dies, day), relat- ing to biologic variations or rhythms with a cycle of about 24 hours. clone, a group of individual plants reproduced asexually from a single original parent. conduplicate , folded together lengthwise. conic, cone-shaped. contingent perennial, a plant living more than 2 years and whose flowering is contingent upon the proper climatic conditions, i.e., rainfall and higher temperature. crenate , applied to leaf margin strongly and ab- ruptly undulate with large teats. cucullate, hooded at the apex. cultigen, a plant known to exist only in culti- vation. Compare cultivate. SPECIAL TERMS cultivate, as a noun, a cultivated plant with known wild ancestors. deflexed, bent downward. descending , of leaves, directed below the hori- zontal. ensiform, sword-shaped. explanate, flattened, spread out flat. exserted, exceeding the corolla, as the filaments extending beyond the tepals. filiferae, threadlike structures along the leaf margins. filiferous , having threadlike structures. friable, said of teeth or leaf margins that are eas- ily brushed or rubbed off. funnelform , funnel-shaped. glaucous, whitened with a waxy coating over the epidermis. guttered, having the sides of the leaves raised to form a trough-shaped leaf, partly condupli- cate. half, handle, applied to the narrowed lower part of the leaf, which is usually the least prickly part for grasping with the hand. keel, the fleshy midrib on the tepal or the leaf. laterals, the main branches of the paniculate in- florescence. maguey, an American Indian name for agave. It was picked up by the earliest Spaniards and appears to have been in general use in the Caribbean region and Mexico. Cortes in "His- toria de Mexico" wrote, "miel de unas plan- tas, que llaman en las otras, y estas maguey, que es muy mejor que arrope; y de estas plan- tas hacen azucar. y vino, que es asi mismo venden."" mescal, an American Indian name applied to agave plants, to the cooked parts of the same, and to the distilled liquor made from the meristem. Used more in northern Mexico. Also applied loosely to Manfreda, Tillandsia, and other monocots. metl, a Nahuatl name for agave, still in use among native tribal people in central Mexico; '■papalo metl" (butterfly agave, A. potato- rum), "tlaca metl" (A. salmiana). monocarpic , a plant or rosette that flowers once and dies. Compare polycarpic. multiannual , a plant that flowers once and dies, but requires several to many years to mature. neck, apical portion ot ovary between the ovar- ian cells and the base of the tube. panicle, the branched inflorescence of the sub- GENTRY: AGAVES OF BAJA CALIFORNIA 115 genus Agave with flowers borne in umbellate clusters on lateral branches. paniculate, like a panicle. patulous, standing open, spreading. plane, applied to leaves having the upper surface flattened as compared with guttered or ex- planate. polycarpic, a plant or rosette that flowers re- peatedly, but not necessarily every year. Compare monocarpic. proterandrous , the condition of a perfect flower when the anthers dehisce before the pistil is receptive. Hence, the flower cannot normally be self-fertilizing. pruinose, having a waxy exudate on the surface of the leaf. pulque, the fermented juice of the larger agaves. Word derived from Nahuatl "poliuhqui"' or "ocli poliuhqui,'" but which was applied to soured or spoiled "ocli," fide Nufiez Ortega, the chronicler of Hernando Cortes. raceme, an inflorescence in which the flowers are borne on pedicels along a central axis. reclinate , reclining, applied to leaves pressing downward upon lower leaves or on the ground. recurved, recurvate, curved backward or down- ward. reflexed, bent sharply downward. retrorse, directed towards the base. rhizome , underground stem or shoot. rosette, a closely spaced group of radiating leaves limited to a portion of the stem, usually at the base of the inflorescence. sapogenin, a compound derived by hydrolysis from saponin. A large group of such com- pounds have been found in plant tissues and named according to their specific molecular configuration, e.g., diosgenin, smilogenin, he- cogenin. shaft , in Agave the central axis of the inflores- cence including the peduncle and the central rachis of the flowering portion or branches. spike, an inflorescence with the flowers more or less sessile along a common or single peduncle or shaft. spine, terminal spine, in Agave the pungent in- durated tip of the leaf. spreading, of leaves extending outward less than 20° from horizontal. subulate, awl-shaped, long tapering. surculose , producing suckers or offsets. sympatric, applied to related species or popu- lations inhabiting the same geographic area. teat , fleshy prominences under the teeth on the leaf margins. teeth, the prickles along the leaf edge. tepal, a combination of sepal and petal, applied when the segments of the perianth are not dif- ferentiated into two dissimilar ranks, (com- mon in monocotyledons). trigonous, three-angled with plane faces. tubular, tube-shaped. umbel , a flat-topped or low-rounded flower clus- ter with the pedicels of unequal length from a common point, like an inverted umbrella. umbellate, having the inflorescence in umbels or in similar form. undulate , applied to leaf margins with low teats, wavy, as compared to straight margins. urceolate, urn-shaped. vallecullate, little valley-shaped, as applied to agaves having folds in the leaf towards apex, plicate. 116 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES. No. 130 Literature Cited Anderson, Edgar. 1949. Intiogressive hybridization. John Wiley & Sons. New York. 109 pp. Aschmann. Homer. 1959. The central desert of California: demography and ecology. Univ. Calif. Press. Berkeley & Los Angeles. }\5 pp. Baegeri. Johann Jakob. 1972. Nachrichten von der Amer- ikanischen halbinsei. Californien. Mannheim. [Also. Span- ish translation by Pedro R. Hendricks Perez, Noticias dc In Fciiinsula Americana de California par el Rev. Padre Juan Jacoho Baegeri. Con una introduccion por Paul Kinhoff. Mexico. Antigua libraria robredo de J. Perriia e hijos. 1942. A resume of original published in Ann. Geogr. 5:237-239. 1866. An English translation by M. M. Brandenburg & C. L. Baumann entitled Observations in Lower California. Berkeley, 1952.] Barco, Miguel del. 1973. Historia natural y cronica de la Antigua California. [Adiciones y correcciones a la noticia de Miguel Venegas] Edicion estudio preliminar. notas y ap- pendices. By Miguel Leon-Portilla. Universidad Nacional Autcinomade Mexico. Inst. Invest. Hist. Mex. Ixxxv + 464 pp. Barrows. David Prescott. 1%7. The ethno-botany of the Coahuilla Indians of southern California. Including a Coa- huilla bibliography and introductory essay by Harry W. Lavvton. Lowell John Beand [and] William Bright. Malki Museum Press. Banning, Calif. 82 pp. [Reprint of Ph.D. Thesis. Univ. Chicago. 1897] Berger, Alwin. 1915. Die Agaven. 285 pp., illus. Jena, Verlag von Gustav Fischer. Bolton, Herbert Eugene (editor). 1919. Kino's historical memoir of Pimeria Alta. Cleveland, 2 vols., illus., maps. Burrus, Ernest J. {editor and translator). 1%6. Wences- laus Linck's diary of his 1766 expedition to northern Baja California. Dawson's Book Shop, Los Angeles. 115 pp. . 1967. Wenceslaus Lincks reports and letters, 1762- 1778. Dawson's Book Shop, Los Angeles. 94 pp. Castetter, E. F., W. H. Bell, and A. R. Grove. 1938. The early utilization and distribution of Agave in the Amer- ican Southwest. Univ. Mexico, Bull. 6. 92 pp. Cave, Marion S.. 1964. Cytological observations on some genera of the Agavaceae. Madrorio 17(5): 163-170. Clavigero, Francisco Xaviero. 1789. Storia della Cali- fornia. Venice. 2 vols., map. [Spanish translation by Nic- olas Garcia de San Vicente, entitled Historia de la Antiqua 6 Baja California. Mexico, 1933. 267 pp. English translation by Sara E. Lake & A. A. Gray. The history of Lower Cal- ifornia. Stanford Univ. Press. Stanford. 1937, 413 pp., maps.] Darton, N. H. 1921. Geological reconnaissance in Baja Cal- ifornia. J. Geol. 29:720-748. Felger, Richard, and Mary B. Moser. 1970. Seri use of Agave (century plant). The Kiva 35:159-167. Gentry, Howard Scott. 1972. The agave family in Sonora. U.S. Dep. Agric. Agric. Handbk. 399. Pp. 1-195, maps, illus. , and Jane R. Sauck. 1978. A study of the stomatal complex in Agave: groups Deserticolae, Campaniflorae, Umbelliflorae. Proc. Calif. Acad. Sci., .Ser. 4, 41(I6):371- 387, figs. 1-28, tables 1-2. Goldman, Edward A. 1951. Biological investigations in Mexico. Smithson. Miscell. Coll. 115:1^76, map. illus. Granick, Elsa B. 1944. A karyosystematic study of the ge- nus A^wve. Am. J. Bot. 31:283-289. Hastings, James R. 1964. Climatological data for Baja Cal- ifornia. Univ. Ariz. Inst. Atoms. Physics Tech. Rep. No. 14. , AND Robert R. Humphrey. 1969. Climatological data and statistics for Sonora and northern Sinaloa. Univ. Ariz. Inst. Atmos. Physics Tech. Rep. No. 19. Johnston, Ivan M. 1924. Expedition of the California Acad- emy of Sciences to the Gulf of California in 1921. XXX. The botany (the vascular plants). Proc. Calif. Acad. Sci., Ser. 4, 12:951-1218. Langman, Ida K. 1964. A selected guide to the literature on the flowering plants of Mexico. Univ. Penn. Press. Phila- delphia. 1015 pp. Larson. R. L.. H. W. Menard, and S. M. Smhh. 1968. Gulf of California: a result of ocean-floor spreading and transform faulting. Science 161:781-783. MoRAN, Reid. 1964. Floracion de A^(/v£' ^'()/(//«(v/(/a/k/ a los 31 anos. Cact. Succulentas Mex. 9:87-88. . 1967. Agave Suhsimpie.x Trelease. Cact. Succulentas Mex. 12:59-61. Nelson, Edward W. 1921. Lower California and its natural resources. Mem. Natl. Acad. Sci. 16( 1st Mem.): 1-194, illus. Rau, Charles. 1864. An account of the aboriginal inhabit- ants of the California Peninsula as given by Jacob Baegert .... Annu. Rep. Smithson. Inst. ( 1863):352-369. Schuchert, Charles. 1935. Historical geology of the An- tillean-Caribbean region. John Wiley & Sons, New York. 811 pp. Shreve. Forrest. 1951. Vegetation of the Sonoran Desert. Carnegie Inst. Washington Publ. 591. xii + 192 pp.. 37 pis. . and Ira R. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. Vols. 1 & 2. Stanford Univ. Press, Stanford, Calif. 1740 pp. Trelease, William. 1912. The agaves of Lower California. Missouri Bot. Gard. Annu. Rep. (191 1)22:37-65, pis. 18-72. Wall, M. E. 1954. Steroidal sapogenins XV. U.S. Dep. Agric. Agric. Res. Serv. AIC-367. 32 pp. (processed) ETAL. 1954a. Steroidal sapogenins XII. J. Am. Pharm. Assoc. 43:503-505. ET AL. 1954b. Steroidal sapogenins VII. Survey of plants for steroidal sapogenins and other constituents. J. Am. Pharm. Assoc. (Sci. Ed.) 43:1-7. ET AL. 1955. Steroidal sapogenins XXVI. U.S. Dep. Agric. Agric. Res. Serv. ARS-73-4. 30 pp. (processed) ET AL. 1957. Steroidal sapogenins XLIII. J. Am. Pharm. Assoc. (Sci. Ed.) 46:653-684. Wilson, J. Tuzo, et al. 1972. Continents adrift: readings from Scientific American, with introduction by J. Tuzo Wilson. W. H. Freeman & Co., San Francisco. 172 pp. Wolf. Ivan, et al. 1%0-65. U.S. Dep. Agric. Unpubl. Rep. Inter-departmental from North. Utilization Research & Development Division, Peoria, Illinois. GENTRY: AGAVES OF BAJA CALIFORNIA 117 INDEX Page numbers in bold face indicate taxon description and essay; synonyms marked with an asterisk^* Abbreviations 102 Abert's tree squirrel 27 adda 7 Aesciiliis parryi 90 Agave offinis* 48 aktites 97 ainericana 10 angiistifolia 98 aquariensis* 27 aiirea 2, 52, 72, 75, 78, 82 distribution 70 avellanidens 14, 15, 58, 59, 61, 63, 67, distribution 58 bmndegeei* 11, 78 cape fi sis 2, 72, 82 distribution 70 canninis* 48 cendata 2, 3, 14, 15, 18, 31, 35, 53 distribution 42 key to subspecies 43 ssp. dentiens 2, 43 ssp. nelsonii 43, 44 ssp. subcerulata 43, 44 connochaetodon* 48 consociata* 15, 16 datylio 97 var. vexans 98 dentiens* 16, 43 deserti 2, 14, 15, 27, 34, 35, 42, 60 distribution 16 ssp. pringlei 2, 16, 20, 59 ssp. simplex 2, 16, 22, 27 disjuncta* 96 gigantensis 3, 14, 15, 52, 58, 59, 63 distribution 58 goldmaniana* 93, 96 gracilipes 14 lechuguilla 42 niacroculinis 60 margaritae 2, 15, 48, 58 distribution 58 inckelveyana 14, 15, 24, 25 distribution 16 moranii 3, 15, 22, 58, 67 distribution 58 murpheyi 60 nelsonii* 16, 42, 44 neomexicana 14 orciittiana* 86 pachyacantha* 86 parrasana 60 parryi var. couesii 27 pringlei* 14, 16 promontorii 2, 75, 78, 80, 81 distribution 70 rosea na* 53, 54 sebastiana 86, 96 shawii 2, 3, 86 distribution 85 var. sebastiana* 96 96 ssp. goldmaniana 3, 63, 86, 93 slevinii* 48 5(^/7/7V/ 2, 14, 15, 18, 48, 78 distribution 52 key to subspecies 53 ssp. frailensis 53, 54 ssp. roseana 2, 44, 54 subsimplex 15, 27, 48 distribution 16 utahensis 14, 27 var. discreta 27 vrAY///.v* 98 vizcainoensis 2, 15, 48, 58, 67 distribution 58 agave age of, 83, 84 as animal food 20, 43, 96 chemical composition of, 18, 20, 42, 52, 53, 72. 80, 81, 82, 85 clones 3, 42, 75, 95 connective gland 75 distribution of, 16, 42, 52, 58, 70. 85 edibility of, 19, 35, 43, 52, 96 evolution of, 1,2 exploitation of, 3, 93 fairy ring 41 fibers of, 72, 78-80, 82, 93 flowers 7, 9 flower measurements of, 7, 8, 9, 73, 88 flowering seasons of, 4, 5, 72, 84 genus of, 10 habit of, 3, 85 habitats of, 2, 90, 93, 95 hybrids 22, 78, 82 ideographs of, 7, 9, 10, 12, 13, 71, 89 key to groups of, 10, 13 localities of, 100, 101 118 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, No. 130 measurements of. 7 nectar 7 populations 42, 95 stem branching 3. 83, 85 stomates of. 14, 70, 84 subgenus 10 uses of, 3-7, 18-20, 35. 43, 52, 60, 67, 78-80, 82, 93 variability of, 16, 22, 42, 95 Agricultural Research Service vii, 1, 20, 85 a-mul 19 aparejo 5 Arguelles, Juan vii, 52 Aschmann, Homer 3 A triplex 90 B Barclay, Arthur S. vii Barco, Miguel del 3, 5, 43, 52 Berger, Alwin 14, 98 Beri>erocaclns 90 bighorn sheep 20 Brandegee, T. S. 1, 52 California Gulf region 1 Campaniflorae 13, 70 distribution 70 key to species 72 Cech, Frank L. vii Clavigero, Francisco X. continental drift 1 datilillo 98 Datyliones 10, 97 datiliyo 98 Deserticolae 13 key to species 14 Earle, W. Hubert vii Engard, Rodney 27 Erianson, C. O. vii Exsiccatae 101 D Gentry, Bruce vii Gentry, Linnea 26 gitogenin 18, 36, 53, 80, 81, 82, 86 Goldman, E. A. 1 H Hamilton, Lucretia vii hecogenin 18, 36, 42, 52, 53, 80, 81, 82, 86 herbarium abbreviations 102 Hermann, Frederick J. vii Hodgson, Wendy vii Howell. Thomas vii Hughes, Regina vii Huntington Botanical Gardens 75, 78 Hyland, Howard L. vii I Idria 95 Imle, Ernest P. vii Indian tribes Apache 20 Chemehueve 20 Coahuilan 19 Cochimi 5, 20, 43, 52 Cocopah 20, 60 Digueiio 20 Kamia 20 Pap ago 20 Pima 20 Seri 35, 43. 44 Ute 20 Yavapai 20 Yuman 20 Indian names of agaves aamXW 35 ahmmo 35 den;kl 35 ?emme 44 Indian use of agave 5, 18-20 Indian women 5 Jones, Marcus 27 Jones, Quentin vii Johnston, Ivan M. 16 Felger, Richard 44 Fouqiiierla 63 Franseria 90 land movements 1 lechuguilla 52, 67 GENTRY: AGAVES OF BAJA CALIFORNIA 119 LeRoy, E. R. vii Lindsay, George E. 84 Link, Wenceslaus 43 Littaea 14, 84 M manogenin 18, 36, 42, 52, 53, 80, 81, 82, 86 Marginatae 14 McClure, John vii Meyer, Frederick G. vii mescal 5, 43, 52 mescal cutting tool 5 mescaliyo 98 mescal pits 18, 20 mezcal, see mescal Miocene 1 Moran, Reid vii, 31, 96 N National Science Foundation vii Nelson, E. W. 1, 79 Neotomo 20, 96 Nolina heldingii 67 O Opuntia 90 Pachycereus 95 Pachyconniis 63, 95 Parry anae 14 Pinkava, Donald vii Pleistocene 1, 78, 85 Rosa minutifolia 90 Rose, J. N. 1, 96 San Leandro fault 1 sapogenin 18, 35, 36, 42, 52, 53, 80, 82, 85, 86 Schubert, Bemice vii Sciurus abertii 27 Sedum alamosannm 102 Sierra de la Giganta 3, 67, 101 Sierra San Pedro Martir 3, 101 Simmondsia chinensis 38, 90 Smith, Earl C, Jr. vii subgenus Agave 10 tatema 7 tatemar 7 Tertiary 1, 2 tigogenin 18, 36, 42, 52, 53, 80, 81, 82, 86 Tillandsia 95 tree squirrel 27 Trelease, William 1 U uani 5 u-a-sil 19 Umbelliflorae 13, 82 distribution 85 key to species 86 Venable, Grady 80 R rainfall silhouettes 4, 5, 72, Rigidae 98 84 ya-mil 19 Yucca 63, 95 MCV 1979 MBl, WIIOI 1 IHKARY UH nCE Y