(Cactaceae)

LIBRARY. JAN 1 2 1987

BOTANICAL GARDEW BEAT ERNST LEUENBERGER

Memoirs of the New York Botanical Garden Volume 41 Memoirs of The New York Botanical Garden

ADVISORY BOARD PATRICIA K. HOLMGREN, Director JAMES L. LUTEYN, Curator of the Herbarium The New York Botanical Garden The New York Botanical Garden GHILLEAN T. PRANCE, Senior SCOTT A. MORI, Chairman, Vice-President for Science Publications Committee, The New York Botanical Garden and Curator The New York Botanical Garden

EDITORIAL BOARD

WILLIAM R. BUCK, Associate Editor The New York Botanical Garden Bronx, New York 10458 H. DAVID HAMMOND, Associate Editor The New York Botanical Garden Bronx, New York 10458

WALTER S. JUDD (1984-1989), Herbarium, Department of , University of Florida, Gainesville, Florida 32611, U.S.A. AMY ROSSMAN (1984-1989), B011 A, Room 329, BARC-West, Beltsville, Mary­ land 20705, U.S.A. LESLIE R. LANDRUM (1985-1990), Herbarium, Department of Botany and Mi­ crobiology, University of Arizona, Tempe, Arizona 85281 U.S.A. MELINDA DENTON (1986-1991), Herbarium, Department of Botany, University of Washington, Seattle, Washington 98195 U.S.A.

The MEMOIRS are published at irregular intervals in issues and volumes of various sizes and are designed to include results of original botanical research by members of the Garden's staff, or by botanists who have collaborated in one or more of the Garden's research programs. Ordinarily only manuscripts of fifty or more typwritten pages will be considered for publication. Manuscripts should be submitted to the Editor. For further information re­ garding editorial policy and instructions for the preparation of manuscripts, ad­ dress the Editor.

Orders for past and forthcoming issues and volumes should be placed with: Scientific Publications Department The New York Botanical Garden Bronx, New York 10458 USA PERESKIA (CACTACEAE) Beat Ernst Leuenberger

MEMOIRS of THE NEW YORK BOTANICAL GARDEN Volume 41

The New York Botanical Garden Bronx, New York 10458

ISSUED 4 DECEMBER 1986 Copyright © 1986

The New York Botanical Garden

Published by The New York Botanical Garden Bronx, New York 10458

International Standard Serial Number 0071-5794

Library of Congress Cataloging in Publication Data

Leuenberger, Beat Ernst, 1946- Pereskia (Cactaceae)

(Memoirs of the New York Botanical Garden; v. 41) Bibliography: p. 129. Includes indexes. 1. Pereskia—Classification. 2. Pereskia—American - Classification. 3. Botany—America—Classification. I. Title. II. Series. QK1.N525 vol. 41 581 s 86-28483 [QK495.C11] [583'.47] ISBN 0-89327-307-4

Copyright © 1986 The New York Botanical Garden International Standard Book Number 0-89327-307-4 DECEMBER 1986 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN 41: 1-141 Pereskia (Cactaceae) Beat Ernst Leuenberger1

Contents

Summary 2 Introduction 3 Taxonomic History 4 Morphology 5 Habit 5 Roots 8 Development of the Stem and Its Appendages 11 Areoles 13 15 Spines 15 16 Development of Rower and 17 Receptacle, Receptacular Bracts, and Areoles 21 Sepaloid and Petaloid Perianth Segments 21 Androecium 23 Development 23 Stamens 23 Staminodes 24 Nectaries 24 Gynoecium 24 Ovules 25 Infructescence and Fruit 26 29 Embryo 29 Cotyledons 31 Perisperm 31 Seedlings 31 Anatomy 34 Root Succulence 34 Epidermis : 34 Stomata 35 Lenticels 35 Periderm 35 Cortex 38 Sclereids 38 Crystals and Druses 39 Mucilage Cells and Cavities 39 Wood and Pith 40 Pollen Morphology 40 Phytochemistry 42 Chromosome Numbers 42 Evolution 42 Distribution 45 Ecology 45 and Climate 45 Phenology 46

1 Botanischer Garten und Botanisches Museum, Konigin-Luise-Strasse 6-8, D-1000 Berlin 33, Federal Republic of Germany. 2 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Pollination 46 Dispersal 4' Conservation 4° Uses 48 Hybridization 50 Infrageneric Relationship 5 * Intergeneric Relationship 53 Systematic Treatment 53 Generic Description 53 Notes on Keys, Terms, and Abbreviations 55 Keys to of Pereskia 55 Key A for Flowering Specimens 55 Key B for Fruiting Specimens 57 Key C for Sterile Specimens 57 1. Miller 58 2. P. humboldtii Britton & Rose 66 a. var. humboldtii 67 b. var. rauhii (Backeberg) Leuenberger 69 3. P. diaz-romeroana Cardenas 71 4. P. weberiana K. Schumann 73 5. P. lychnidiflora De Candolle 76 6. P. aureiflora Ritter 83 7. P. guamacho F. A. C. Weber 85 8. P. zinniiflora De Candolle 89 9. P. portulacifolia (Linnaeus) De Candolle 93 10. P. quisqueyana Liogier 97 11. P. bleo (Kunth) De Candolle 99 12. P. nemorosa Rojas Acosta 103 13. P. sacharosa Grisebach 107 14. P. grandifolia Haworth Ill a. var. grandifolia 113 b. var. violacea Leuenberger 116 15. P. bahiensis Gurke 119 16. P. stenantha Ritter 122 Insufficiently Known Taxa 126 Doubtful Names and Excluded Taxa 127 Acknowledgments 129 Literature Cited 129 Numerical List of Taxa 134 List of Exsiccata 134 Index of Local Names 138 Index of Scientific Names 139

Summary

Leuenberger, B. E. (Botanischer Garten u. Botanisches Museum, D-1000 Berlin 33, Federal Republic of Germany). Pereskia (Cactaceae). Mem. New York Bot. Gard. 41: 1-141. 1986.—This monograph of Pereskia, neotropical of Cac­ taceae with 16 species, which range from southern and the Caribbean to northern and Uruguay, is based on the study of herbarium and liquid- preserved material as well as on the observation of seven species in the field and of all except one species in cultivation. The taxonomic history, morphology, anatomy, pollen morphology, phytochemistry, chromosome numbers, evolution, distribution, ecology, conservation, uses, hybridization, and infrageneric and in­ tergeneric relationships are discussed. Pereskia is a genus of , shrubs, and scramblers characterized by the presence of regular, slightly succulent leaves, bearing in their axils typically cactaceous areoles, i.e., brachyblast shoots 1986] PERESKIA (CACTACEAE)

which produce spines and, in about half of the species of the genus, periodically also brachyblast leaves. Several species have tuberous roots. The predominantly receptacular nature of the and and the transition from superior to inferior gynoecia are described and illustrated. Besides paniculate and racemose or single terminal or axillary flowers, proliferation from the recep­ tacles resulting in chain-like clusters of fruits is observed within the genus. Chro­ mosome numbers of In = 22 were found in the 13 species successfully investigated. The evolutionary origin of the genus and family may be sought in northwestern South America, as suggested by character correlations and distribution. Local uses of several species are known; the planting of living fences and hedges being the most widespread. Separate keys for flowering, fruiting, and sterile specimens are provided in the systematic treatment. All 16 species are illustrated by line drawings of relevant parts. The generic concept follows that of previous monographers (Britton & Rose, 1919; Vaupel, 1926) and includes (Berger) F. Knuth in the synonymy of Pereskia. No formal infrageneric classification is upheld. Sixteen of the previously described 59 species are recognized; 26 names are placed in synonymy and 17 are doubtful or excluded from the genus. The list of correct names as accepted here includes: Pereskia aculeata Miller, widespread with dis­ junct distribution, recurved geminate spines, and fragrant flowers; P. humboldtii Britton & Rose, P. diaz-romeroana Cardenas, and P. weberiana K. Schumann, tuberous-rooted Andean species with small leaves, flowers, and fruits; P. lychni­ diflora De Candolle, an orange-flowered with primitive floral characters; P. aureiflora Ritter, only recently described; P. guamacho F. A. C. Weber, a tree with small yellow flowers appearing during the leafless stage; P. zinniiflora De Candolle, P. portulacifolia (L.) De Candolle, and P. quisqueyana Liogier, endemics from the Greater Antilles and distinctive for their dioecious flowers; P. bleo (Kunth) De Candolle, a shrub with petiolate leaves and bright red flowers from mesic forest in and ; P. nemorosa Rojas Acosta, P. sacharosa Gri­ sebach, P. grandifolia Haworth, and P. bahiensis Giirke, pink-flowered species with medium to large leaves, all of South American origin; P. stenantha Ritter, closely related to the former but distinctive for its urceolate-campanulate flowers. One new combination, P. humboldtii var. rauhii (Backeberg) Leuenberger, is nec­ essary, and one new variety, P. grandifolia var. violacea Leuenberger, is described in this paper.

Introduction Pereskia is a genus of 16 species of neotropical trees and shrubs. It differs from typical succulent Cactaceae in its woody habit and the presence of normal leaves and is therefore not always easily recognized as a member of the family at first sight. However, it exhibits the basic characters of the family, namely the spine- bearing areoles and the peculiar nature of the floral cup which incorporates few to many -bearing nodes. Other floral characters distinctive of the genus and the family include: numerous perianth segments, spirally arranged centrifugal stamens, and a gynoecium of few to many connate carpels, adnate to the recep­ tacular cup. The superior to inferior ovary is more diverse in Pereskia than in other genera of the family, where it is always distinctly inferior. By reason of its general habit and the morphological and anatomical features of its stem, leaf, and , Pereskia is generally considered to be the most primitive genus within the Cactaceae. The genus has repeatedly received the attention of botanists and, to a lesser extent, that of amateur collectors. It is better represented in herbaria than are most other Cactaceae and has been less affected by nomenclatural fluctuations MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

than the more popular genera. Nevertheless a formal taxonomic treatment is needed because Vaupel's monograph (1925) is out of date. His study was based on a rather narrow selection of specimens, mainly consisting of the material at the Berlin-Dahlem herbarium, subsequently destroyed in 1943. Later publications (Backeberg & Knuth, 1935; Backeberg, 1958) were not based on standard taxo­ nomic methods. The separation into two separate genera, Pereskia and Rhodo­ , relied mostly on supposed, rather than observed, characters of the ovary. An impressive number of publications deals with the morphology and anatomy of individual species of Pereskia, namely the anatomy of leaves and stems (Bailey, 1960, 1961a, 1961b, 1962, 1963a, 1963b, 1963c, 1963d, 1964, 1965; Bailey & Srivastava, 1962; Gibson, 1975, 1976) and the developmental morphology of the shoot and flower (Boke, 1954, 1963a, 1963b, 1964, 1966, 1968). These thorough investigations demonstrated the significance of the genus for evolutionary con­ siderations and for the understanding of the phenomenon of succulence within the family. This monograph is based on herbarium material from over 50 herbaria as well as on the study of seven species in the field and observations of cultivated of all except one species. Sixteen species are recognized. One variety is described as new and a new combination at the varietal level is made. A possibly new taxon is discussed but treated under "insufficiently known taxa." Taxonomic History The first specimens of Pereskia were probably collected by Plumier between 1689 and 1695 in the , but are known only from manuscript descrip­ tions (Hunt, 1984). Other pre-Linnean collections were described with phrases such as "Portulaca americana latifolia adfoliorum ortum lanugine obducta" (Plu- kenet, 1696), "Grossulariae fructu majore arbor spinosa, fructu folioso e viridi albicante" (Sloane, 1696), and "Malus americana spinosa portulacaefolio, fructu folioso, semine reniformi splendente" (Commelin, 1697). It is noteworthy that both the specimens and the illustrations belonging to these phrases were later thought to be the same as "Pereskia aculeata, flore albo, fructu flavescente" of Plumier (1703), which became Cactus pereskia L. and Pereskia aculeata Miller. However, they can now be identified as P. guamacho, a species named nearly 200 years later. The pre-Linnean generic name Pereskia dates back to Plumier (1703), who gave a short description of flower and fruit characters and mentioned one species by the above mentioned phrase "Pereskia aculeata, flore albo, fructu flavescente." Rower, fruit, and were illustrated, and a lengthy laudatio commemorated the merits of the French scholar Nicolas Claude Fabri de Peiresc (1580-1637), latinized by Plumier as "Nicolaus Fabricius Peireskius, Senator Aquisextiensis." Plumier's spelling of the generic name was nevertheless Pereskia, probably re­ flecting the correct French pronunciation, as Weber (1898a) clear-sightedly ex­ plained. Both Linnaeus (1753) (Cactus pereskia) and Miller (1754) (Pereskia) retained this spelling, which is therefore nomenclaturally correct. Subsequent authors modified the spelling to "Peirescia" (Zuccarini, 1837; Pfeif­ fer, 1837a, 1837b; Salm-Dyck, 1850), "Perescia" (Lemaire, 1838), and "Peires- kia" (Schumann, 1898; Vaupel, 1925), assuming that "Pereskia'" was an error. Sometimes awkward polemics, speaking of a mutilation of the name of the person to be honored, have continued on the subject ever since (Dolz, 1937; Buxbaum, 1940; Backeberg, 1966; Ritter, 1979), disregarding the fact that Werdermann (1937) also had demonstrated not only Plumier's obvious intention of latinizing the name in this spelling (mentioned four times in the publication, hence not a 1986] PERESKIA (CACTACEAE) 5 typographical error), but also its correctness under the International Rules of Botanical Nomenclature. The same applies today under the current International Code. Only one genus of Cactaceae was accepted by Linnaeus (1753) in "Species plantarum." The two leafy cacti hitherto known from Plumier's collections from the West Indies were listed as Cactus pereskia L., and C portulacifolius L. Only the former species was in cultivation in Europe at that time. In the "Gardeners Dictionary," ed. 4, Miller (1754) reinstated Pereskia as a genus, but he commented on the species only in the 8th edition (Miller, 1768) where he formally described Pereskia aculeata, retaining Plumier's first adjective of the phrase as the epithet. Miller's authorship for the genus is to be accepted under , although some writers still attribute it to Plumier or add Plumier's name in parentheses (Backeberg, 1958; Bravo-Hollis, 1978). In 1812, Haworth named two species, of which P. grandifolia is well-known today, while the other, P. longispina, remains in the doubtful category. Two species discovered by Humboldt and Bonpland in Colombia and Peru, first published as Cactus bleo and C. horridus (Humboldt, Bonpland & Kunth, 1823), were trans­ ferred to Pereskia by de Candolle (1828a). Four new species were later described by him (de Candolle, 1828b), solely from the plates of Mocifio's "Flora Mexicana" ined., hence supposedly from Mexico. The dramatic fate of these plates is described by McVaugh (1980a, 1982). The elucidation of the four names, representing two species of Pereskia and one of , is discussed under Pereskia lychni­ diflora, P. zinniiflora, and in the list of excluded names under P. rotundifolia. Pfeiffer (1837a, 1837b), Forster (1846), Salm-Dyck (1850), and Riimpler (1886) included several species in Pereskia that were excluded from the genus by later monographers and attributed to Opuntia and Pereskiopsis. The circumscription of the genus by Schumann (1898) approaches the generic limits as upheld today. Britton and Rose (1919) added several new species and reduced Schumann's subgenera to the rank of series. Berger (1926, 1929), having a more phylogenetic approach to , emphasized differences in the position of the ovary and ovules and proposed a new subgenus Rhodocactus, though without knowledge of the morphology of the gynoecium of some of the species he included therein. Knuth (in Backeberg & Knuth, 1935) elevated the subgenus Rhodocactus to the rank of a genus, recombining some of the names, again based, however, on supposed rather than observed characters of the gynoecium. Backeberg (1956) classified the remaining species of Pereskia into two subgenera by creating a subgenus Neopeireskia for the Andean species with small leaves. Later authors questioned the generic status of Rhodocactus (Bravo-Hollis, 1978; Buxbaum, 1956; Hunt, 1967) and upheld only one genus, Pereskia, the position taken in this monograph. With a few supposed novelties, but also remarkable discoveries by Cardenas (1950), Liogier, (1980), and Ritter (1979) during the last few decades, the number of names had increased to about 50 by 1979. Of these, 16 are recognized as correct at the species level in this monograph.

Morphology

HABIT The species of Pereskia have been variously described by collectors and monog­ raphers as small trees, shrubs, or . The basic growth forms in Cactaceae have MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 1. Habit of Pereskia trees and shrubs. A-C, Pereskia lychnidiflora. A, juvenile on a pasture near El Rancho, Progreso, Guatemala {Leuenberger & Schiers 2515a). B, apex of juvenile plant with dormant shoot apex surrounded by a whorl of subverticillate branches {Leuenberger & Schiers 2512). C, mature tree over 10 m near Tehuantepec, Oaxaca, Mexico {Leuenberger & Schiers 2533a). D, Pereskia portulacifolia, two-stemmed tree, ca. 5 m tall, in dry forest near Lago Enriquillo, (same loc. as Leuenberger 3048). E, Pereskia grandifolia, canes of 5-10 cm diam. on a mature shrub in cultivation at Huntington Botanical Garden, San Marino, California. (All photographs of this and the following plates by the author unless otherwise stated.) been discussed by Buxbaum (1957) and Rauh (1979), who showed that Pereskia species exhibit the normal growth patterns found in other woody dicotyledonous plants. Typically arborescent species like P. lychnidiflora produce a primary stem with subverticillate branching. During seasonal phases of inactivity the shoot apex in this species is concealed among the densely spaced areoles of the obtuse shoot tip, which is surrounded by the uppermost whorl of branches (Fig. 1A, B). Even in seedlings the shoot apex sometimes aborts, and the primary stem continues from one of the uppermost areoles. Even though young plants exhibit a distinctly monopodial growth pattern, the same habit may result from sympodial growth, which at least rarely occurs on juvenile plants and is obviously the rule at a later stage of development in the branching pattern of the crown. The primary stem eventually terminates growth, and a broad, umbrella-shaped crown is formed by repeated dichotomous to trichotomous branching of the sprays (Figs. 1C, 5 A). A similar growth habit is found in P. zinniiflora and P. guamacho. The height of these trees may at least partially depend on the density of the 1986] PERESKIA (CACTACEAE)

B

FIG. 2. Development of shrubby growth in Andean small-leaved Pereskias. A-D, Pereskia we­ beriana—hybrid {cult. hort. Berol. 206-01-80-70). A, one-year-old seedling. B, the same seedling during the second year, with terminal inflorescence. C, the same plant with lateral branching during the third year. D, the same plant with flowering shoots arising from near the base of the primary stem (filled circles represent flowers or flower buds; stems of one year or older are drawn in black throughout). stand. Free standing trees of P. lychnidiflora develop a much shorter trunk and a more rounded crown, comparable to a lime-tree (Tilia) (Eichlam, 1909) or an apple-tree (Standley & Williams, 1962). Transitional growth forms, called trees by some, but shrubs by others, occur in species where the primary shoot terminates growth during the first few years with or without formation of an inflorescence. Lateral erect or nearly erect shoots of 8 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

often increasing strength develop from near the base, and branching of these shoots produces a short-trunked or many-stemmed tree called "Baumstrauch" (= tree- shrub) by Rauh (1979). Depending on the level of primary branching, the aspect is more tree-like or more shrub-like (Figs. ID, 32L, 38G, 47J, 49J). Individual shoots (canes) may become tree-like. The seemingly contradictory field notes on the habit of these species is therefore not surprising. A shrubby specimen of P. grandifolia is shown in Figure IE, but the best examples of shrubs within the genus are the small-leaved Andean Pereskias which, according to Rauh (1979), belong to the group of cane-forming shrubs ("Schosslingsstraucher"), like roses (Rosa) or black elder (Sambucus). Long shoots of increasing strength initiate from the very base of the epicotyl and soon bend over (Fig. 2D). Lateral branching occurs mainly on the distal portion and is oriented upward. Rapid growth is probably enhanced by the assimilates stored in the tuberous roots characteristic of this group of species. Pereskia aculeata, the only species normally described as a , climber, or straggling shrub, has basically the same branching pattern as the second, transi­ tional, group mentioned above, but the flexuous primary and secondary branches may elongate and form vines if adequate support is available. They may attain a length of 10 m. The tallest growing arborescent member of the genus is probably P. lychnidiflora which is reported by MacDougall (1947) to reach 20 m. This would make it the tallest arborescent cactus known.

ROOTS Only scattered observations on the root system of Pereskia species have been made up to now. Cardenas (1950) indicated woody roots with fusiform thickenings in Pereskia diaz-romeroana. Bailey (1963a) noted that the Andean small-leaved species have succulent tuberous roots and Rauh (1979) reported fusiform to tu­ berous thickenings of up to 60 cm long and 5 cm diam. in cultivated plants of P. weberiana grown from cuttings. Little is known about the non-succulent root systems. Several species tend to form a tap root in cultivation (P. lychnidiflora, P. sacharosa and others). Nothing, however, is known about the dimension of the deep root system of mature trees and shrubs in nature. In cultivation a ca. 20-year-old tree of P. sacharosa at the Berlin Botanical Garden showed both deep-reaching tap roots and weaker shallow roots. Roots spreading at the soil surface were observed by the author around the trunk of a large specimen of P. lychnidiflora (Fig. 3 A) near Tehuantepec in Mexico. As succulence of stem and leaves is not very pronounced in Pereskia and as the leaves are deciduous during long periods of drought, the presence of both shallow and deep roots seems adequate. The deep root system may be crucial for survival over long drought periods. This is also suggested by experiences with cultivated plants, where roots tend to escape from even large pots into the substrate below and allow the plants to keep their leaves for a prolonged period without watering. On the other hand, the shallow root system may facilitate quick water uptake after the first rains. This could be important for competition with other plants, as it appears to be for the more succulent Cactaceae in seasonally dry woodland. Further observations on root succulence in Pereskia were made by the author during a visit to the Dominican Republic in 1983. Fleshy root thickenings of even larger dimensions than the ones mentioned above were found in P. quisqueyana, partly exposed on the coral limestone habitat at the type locality of this species. Root tubers up to 20-30 cm long and 10 cm thick were observed at the base of one shrub (Fig. 3B). Cultivated specimens of P. portulacifolia in the collection of 1986] PERESKIA (CACTACEAE)

FIG. 3. Roots of Pereskia. A, P lychnidiflora, shallow root system of a large specimen near Tehuantepec. B-D, P. quisqueyana {Leuenberger 3044). B, base of shrubby plant with large fusiform root tubers. C, tuberous root and cross sections of the same. D, tuberous root, x.s., showing core of unlignified tissue formed by the vascular cambium, and the narrow ring of cortex with stone cells. E, F, P. weberiana {cult. hort. Berol. 037-01-37-80). E, root stock with large tuberous roots. F, tuberous root, x.s.

Padre Julio Cicero at San Cristobal, Dominican Republic, reveal the same char­ acter. However, the closely related P. zinniiflora does not seem to produce tu­ berous roots, at least not by the few plants I have seen in cultivation. On the other hand, I noticed root thickenings in cultivated P. guamacho, but only after several years of cultivation. It is noteworthy that within the genus succulent roots occur in two different groups which do not seem very closely related. Formation of large tuberous roots in P. weberiana occurs within one or two years, as a specimen grown from a cutting planted in the display glasshouse at the Berlin Botanical Garden in 1981 shows. When the sparingly branched shrub, about 3 m tall, had to be removed in 1983, a massive Dahlia-like tuberous rootstock with individual tubers up to 40 cm long and 15 cm diameter (Fig. 3E) was discovered. Individual tubers that were removed and kept dry at room temperature showed some shrink­ ing but stayed alive for more than one year with a green cortex below the shreddy periderm of the root. Formation of buds and stems was not observed. However, it is evident that the succulent root system enables these plants to survive for a long period, more than one year under experimental conditions, at the expense 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 4. A-E, foliage and development of brachyblast and lateral auxoblast shoots (schematic, not in scale). A-D, development of areole in species with brachyblast leaves. A, areole of new growth, with spines and subtending leaf (auxoblast leaf)- B, areole with spines, brachyblast leaf and leaf scar of auxoblast leaf. C, older, elongated areole (spur shoot in the strict sense used here) bearing spines and brachyblast leaves at the tip. D, spur shoot with subsequent growth as a long shoot (long internodes) bearing auxoblast leaves. E, stem of P. aculeata, a species without brachyblast leaves (schematic, from cultivated material); a, young areole with a pair of recurved spines (auxoblast leaf fallen); b, older 1986] PERESKIA (CACTACEAE) 11 of a nearly total loss of stems and leaves. The tendency for long shoots to die back seasonally was observed by Boke (1968), in P. diaz-romeroana.

DEVELOPMENT OF THE STEM AND ITS APPENDAGES Detailed studies on the development of the stem of three species of Pereskia were made by Boke (1954), and there is no doubt that these results are represen­ tative for the genus as a whole. Boke noted that early growth in width of the axis is a striking feature of Pereskia. The flank meristem initiates dispersed leaf pri­ mordia; axillary buds arise from the flank zone of the apical cone very early, and the axillary bud, which in Pereskia becomes a spine- and sometimes also leaf- bearing areole, attains an advanced stage of development at or near the level of the shoot apex in the region where growth in width predominates (Boke, 1954, p. 621). This proleptic development of axillary buds produces the characteristic spiny areole of the Cactaceae (Rauh, 1979), located on a persistent leaf base, which separates early in development by abaxial constriction from the leaf proper. Stems of long shoots are fleshy when juvenile, but become more or less woody, with varying thickness of cortex and pith. The auxoblast leaves subtend the axillary bud (areole) and are normally deciduous after the first season (Fig. 4A, B). The areoles continue to produce foliar appendages, i.e., spines, and sometimes brachy­ blast leaves, beyond the first season (Fig. 4C). Little or no elongation takes place, and therefore the areole may also be called a spur shoot. The areoles always produce trichomes, in all species usually spines, and in some species leaves also. Leaves, or leaves and spines, are produced seasonally for one to several years after the auxoblast leaves have fallen. Spine formation may continue for many years after leaf production has ceased. The areoles of distal twigs in the upper region of the crown are often spineless, in contrast with the fiercely spiny trunks and lower branches and twigs. A similar tendency is observed in other arborescent or columnar Cactaceae, e.g., Consolea, Neoabbottia, Pachycereus, and Browningia. Although most areoles remain active and may thus be termed spur shoots, I prefer to retain, for practical reasons in the descriptive part, the term areole, unless a conspicuous spur-like elongation has taken place (Fig. 4B, C). In herbarium material it is often impossible to distinguish between leaf-producing and only spine-producing or inactive buds, as the leaves easily fall during the drying process. The term "spur shoot" —suggesting a prominent structure—is hence applied here only to those conspicuous knob-like to twig-like structures with suppressed in­ ternodes observed in several species (Fig. 4C). Spur shoots may resume growth as normal long shoots (Fig. 4D). In arborescent species the arrangement of spur shoots and lateral shoots of varying length produces a distinctive branching pat­ tern, e.g., in P. portulacifolia (Fig. 5B, E). In cultivated juvenile plants of P. nemorosa and P. grandifolia the formation of brachyblast leaves occurs even before the abscission of the subtending leaf (Fig. 5G). The number of leaves on a single areole in one season varies. A maximum of five leaves was observed in cultivated specimens of P. grandifolia. The leaves are formed in or near the center of the areole, surrounded by older spines, in P. grandifolia or P. nemorosa (Fig. 5F, G), or with the adaxial side lacking spines, as in P. sacharosa (Fig. 43H). New

areole with additional, straight spines; c, areole with a suppressed lateral shoot bearing only one leaf and an aborted shoot tip on a very short but conspicuous internode; d, areole with developing lateral shoot. 12 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 5. Branching pattern and foliage of Pereskia. A, P. lychnidiflora {Leuenberger & Schiers 2718), dichotomous and trichotomous branching and brachyblast leaves. B, P. portulacifolia {Leuen­ berger 3048), branches with numerous spur shoot and short perpendicular twigs. C, P. lychnidiflora {Leuenberger & Schiers 2508, cult.), top whorl of seedling, exhibiting anisophylly of primary foliage. D, P. lychnidiflora {Leuenberger 2513), trunk, ca. 15 cm in diam. with some areoles producing brachyblast leaves. E, P portulacifolia {Leuenberger 3048), branch with epitonous brachyblast foliage and young lateral shoot (arrow). F, P. grandifolia var. grandifolia {cult. hort. Berol. 047-01-78-84), areole with three brachyblast leaves and new spines developing towards the end of the season. G, P. nemorosa {Gonjian s.n., cult.), seedling with brachyblast leaves formed before the auxoblast leaves have fallen. H, P. bleo {cult. hort. Berol. 082-18-82-70), smooth periderm with lenticels; areoles without brachyblast leaves, the lower right areole with central bud initiating a lateral shoot. I, P. aculeata {cult. hort. Berol. 169-02-79-80), areole with three primary spines, lateral shoots with scale- 19°°] PERESKIA (CACTACEAE) 13

brachyblast leaves and new spines are not formed simultaneously. The leaves appear at the beginning of the wet season, the new spines during and towards the end of the growing season, but new spines may be formed on areoles which still bear brachyblast leaves (Fig. 5F, J). According to Boke (1954, p. 625), "each brachyblast leaf produces its own axillary bud meristem, and this frequently becomes as large and conspicuous as that of the spur itself. Primordia produced by the large axillary bud seem to abort." It seems possible that in some cases bud meristems in the axils of brachyblast leaves may become active and produce spines or even flower buds. This could explain the formation of several flowers on a single spur shoot of P. guamacho. Developmental studies of older areoles are extremely difficult, however, because sectioning of the very tough spine bases lying side by side with meristematic tissues poses practical problems. Eight of the 16 species lack brachyblast leaves completely, but some areoles may produce suppressed lateral shoots with only one or few leaves. They are the first leaves of a lateral shoot which eventually elongates during the same season. If it aborts, it is still notably different from a true brachyblast due to the short but conspicuous internode and the scale-like prophylls which usually accompany it (Figs. 4E, 51). Leaves never appear on older areoles of these species (Fig. 5H). In contrast, brachyblast leaves sometimes occur even on old areoles of the trunk in species of the first group, e.g., in P. lychnidiflora (Fig. 5D).

AREOLES2 The areoles of Cactaceae are proleptic axillary buds which produce trichomes, usually spines, and in some Pereskia species also leaves. The trichomes charac­ teristic for all Cactaceae, according to Boke (1954), arise from protodermal cells at either side of the leaf primordium at the time the axillary bud is initiated. They ultimately surround the axillary bud and all structures it produces. Growth of the trichomes is mostly restricted to their base. In about half of the species of Pereskia all trichomes are uniseriate. Boke (1954) noted some multiseriate trichomes in P. aculeata. Intermediate appendages, ranging from soft multiseriate trichomes to coarse hairs or tortuous bristles, are visible to the naked eye in several species, particularly in P. humboldtii (Fig. 6C) and on main shoots and in seedling plants of P. lychnidiflora (Fig. 6B). They are mostly distinct from even the weakest spines and the tips of the thicker ones are trichome-like, distinguishing them from spines. They are particularly conspicuous in new growth and often caducous on older areoles. They are restricted to the lower end of the areole adjoining the base of the petiole of the subtending leaf. A detailed study of the developmental mor­ phology of these appendages seems worthwhile to evaluate their possible evolu­ tionary significance for areole structure within Cactaceae, and particularly in com-

2 Here, as well as in the descriptions in the systematic part, the areoles are treated before the leaves, although the logical sequence would be "auxoblast leaves—areoles—spines—brachyblast leaves." Since the distinction of auxoblast and brachyblast leaves is often difficult in herbarium material with fallen leaves, and because the areoles are a conspicuous element in all specimens and a basic character of all Cactaceae, this sequence has been adopted here.

like prophylls (withered), the upper one aborting. J, P. grandifolia var. violacea {Horst & Ubelmann HU 226, cult.), ramification; a, subtending areole with continuing spine formation; b, first areole of axillary shoot 14 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 6. Morphology of the stem and its appendages. A, P. nemorosa {Gonjian s.n., cult.), areoles with short tomentum of uniseriate hairs. B, P. lychnidiflora {Leuenberger & Schiers 2508, cult.), areoles with tomentum, longer multiseriate, coarse and flexuous hairs, and spines. C, P. humboldtii var. humboldtii {Hutchison 1452, cult.), areoles with short tomentum, coarse hairs, and spines. D-F, P. aculeata {cult. hort. Berol.), areoles showing spine dimorphism and increase of spine number with age. G, P grandifolia var. grandifolia {Leuenberger et al. 2656), spiny trunk of old specimen. H, P. grandifolia var. grandifolia {cult. hort. Berol. 047-01-78-80), spineless young stem with stomatal cavities shining through as light dots. I, P. nemorosa {Gonjian s.n., cult.), beginning periderm for­ mation. J, P aureiflora {Leuenberger & Martinelli 3054), stem with conspicuous lenticels after peri­ derm formation. parison with the glochids of the . Rutishauser (1981, pp. 26, 27, fig. 9e, f) described and illustrated the early stages of such appendages of P. aculeata in a paper discussing the formation of stipules, emergences, and hairs at the leaf bases of Centrospermae. Length and color of the trichomes seem to be characteristic for some species. White, gray, brown, and rusty brown tomentum is found in the genus. Boke (1954) noted that maturing cells of trichomes of P. zinniiflora (=P. cubensis) rapidly become filled with deeply staining ergastic materials. This species has conspicu­ ously rusty brown areoles. Seasonal growth of new leaves is announced by flocks of new trichomes which are partly pushed out of the densely pulvinate mass of older trichomes by the developing brachyblast leaves. 1986] PERESKIA (CACTACEAE) 15

LEAVES The leaves of Pereskia are not appreciably different from those of other woody dicotyledons. They are rather variable in size, shape, and thickness, partly de­ pending on environmental conditions, and soft fleshy to somewhat succulent. In shrubby species large leaf size is quickly attained in seedling plants while, e.g., in P. lychnidiflora, juvenile plants tend to have consistently smaller and narrower leaves. The petiole is long and distinct in P. bleo, but short to inconspicuous in other species. Shape and size of the leaf blade shows great plasticity and the observation of living plants over several years leads to the conclusion that seem­ ingly great differences in herbarium specimens have to be interpreted with great care, as they do not necessarily represent specific differences. Moreover, the high degree of shrinkage during the drying process calls for much care also in the comparison of living and dried material. Puzzling and apparently uncorrected variation of leaf shape, even a weakly defined heterophylly, sometimes occurs on stems of P. sacharosa and P. bahiensis, with acute or broadly rounded leaves both among brachyblast and auxoblast leaves. In P. portulacifolia, the difference seems to be positively correlated with the position of the leaf. Emarginate leaves in this species occur on the areoles only, the subtending leaves are acute. An- isophylly is conspicuous in young plants of P. lychnidiflora on horizontal twigs and branches (Fig. 5C). Venation is generally pinnate, approaching pseudopalmate (P. guamacho), or pseudopalmate (P. lychnidiflora). Hickey (1973) used the terms eucamptodromous to cladodromous for the pinnate venation observed in Pereskia, and imperfect acrodromous for the pseudopalmate type. The angle of diversion of the lateral veins is acute to moderate and proves to be a significant character by which to distinguish two frequently confused species, P. sacharosa and P. nemorosa.

SPINES Repeated investigations on the nature of the spines of Cactaceae, notably by Wetterwald (1889), Leinfellner (1937), Buxbaum (1951), and particularly the de­ tailed studies by Boke (1954, 1980), showed that spines arise from the axillary bud in acropetal order and in the same general manner as leaves arise from an apical meristem. They are homologous with leaves. In the initial stage, spine primordia resemble leaf primordia, but the apex of a spine primordium usually shows early signs of maturation (Boke, 1954, p. 623). A basal meristem is re­ sponsible for growth in length and increase in diameter. Gibson (1976) noted that no vascular tissue enters the spine. After periderm formation it is possible to peel off the whole areole including dead spines and trichomes. The axillary bud remains undamaged in the cortex. Small green protrusions of cortical tissue are located under each hollow spine base, where no periderm is formed in P. lychnidiflora. Spine arrangement on seedlings and on older branches and the trunk is usually more or less radially symmetrical (Figs. 5H, 6G) but in some species there is a tendency of spine formation towards the abaxial side of the areole, while the brachyblast leaves appear on the adaxial side. Deflexed lateral spines by contortion upon drying of the mature spines occur in P. sacharosa, and on strong canes of P. weberiana almost claw-like spines with broad base are observed, which recall the distinctly claw-like first pair of spines of P. aculeata, the only species with regularly heteromorphic spination. Although P. aculeata has two very different spine forms, of which the first claw-like pair was interpreted by Buxbaum (1951) as homologous with a pair of prophylls, the sequence of the first formed spine 16 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 primordia is, according to Boke (1954), no different from that of species with uniform spines as, e.g., P. grandifolia. The two primordia on either side of the axillary bud soon develop a broader base in P. aculeata and thus form a pair of spines which spread outward and backward. In cultivated material I have some­ times observed a third median spine developing in the same way (Fig. 51), but normally the subsequent spines are straight (Fig. 6D-F). In the Caribbean species the first spines formed are usually the lateral ones, followed by a third, median, and additional adaxial spines. In P. lychnidiflora usually only a median, very stout and long spine develops on some twigs. The number of spines is variable and a rather unreliable character. Up to 100 spines are formed on old areoles of the trunk in P. grandifolia (Fig. 6G), but this feature is insufficiently known in many species. Spine length varies greatly within an individual areole and also to some degree with the age of the plant as well as with environmental conditions. A maximum within the genus (16 cm) was measured in P. lychnidiflora. The living, growing spines are usually brightly colored, often reddish or yellowish, sometimes with both colors occurring within the same species. They are usually darker at the tip. The mature spines are variously brown to nearly black in many species, becoming gray with age.

INFLORESCENCE While the complex nature of the flowers of Pereskia has been studied by several authors, little information is available on the inflorescence as a whole. Buxbaum (1953) illustrated a racemose inflorescence of Pereskia aculeata and mentioned lateral flowers initiating from the areoles of the receptacle in this and other species. Rauh (1979) discussed two basic types of inflorescence in the genus. The paniculate inflorescence of P. aculeata (Figs. 7E, 8) has loosely arranged flowers, and that of P. weberiana (Fig. 7F) has basically the same structure but with extremely short internodes, whereby the flowers appear as axillary clusters. The other type is the "cymose" inflorescence, produced by a chain-like proliferation from the recep­ tacular areoles, as in P. grandifolia (Fig. 7D). Additional pedunculate cymes may initiate from areoles of the peduncle and from lower vegetative areoles producing a combined paniculate-cymose, often subcorymbose, inflorescence with up to 50 flowers, as in P. grandifolia. The inflorescence of P. bleo (Fig. 7C) differs in a remarkable way from the two types described by Rauh (1979). This species de­ velops a racemose inflorescence with a terminal flower on the main shoot or on lateral branches. Unlike P. grandifolia, however, the stem is not gradually thick­ ening into the receptacle but is constricted into a leafless pedicel in a terminal position. The lateral flowers initiate in an acropetal order from the uppermost areoles of the stem in the axils of regular leaves of leaf-like bracts. Their pedicels are also ebracteate, and it is noteworthy that their abscission zone is not at the level of the areole but slightly raised, leaving a green stump on the stem, when a nonfertilized, withering flower falls after two or three days (Fig. 39E). In Pereskia guamacho the flowers are axillary, subsessile, and solitary to fas­ ciculate (Fig. 7B). The areoles or spur shoots obviously have several meristems of which only one produces leaves while the others produce flowers. Unfortunately not enough material was obtained to study this properly, and a detailed investi­ gation should elucidate the morphology of the spur shoots in this species, which is also noteworthy because it flowers profusely before the leaves appear, a phe­ nomenon not observed in other species. Areoles with clustered flowers are known from several other genera of Cactaceae, e.g., Myrtillocactus and Weingartia, and specialized flowering spur shoots from Neoraimondia (Rauh, 1957, 1979). 1986] PERESKIA (CACTACEAE) 17

FIG. 7. A-F, interpretation of inflorescence structure of Pereskia. A, single flower in terminal, subterminal, or lateral position {P. lychnidiflora). B, flowers fasciculate or single on areoles and spur shoots {P. guamacho). C, closed racemose inflorescence with lateral flowers developing in acropetal sequence {P. bleo). D, cymose-paniculate inflorescence with proliferation from the receptacle {P. grandifolia). E, paniculate inflorescence {P. aculeata). F, condensed paniculate inflorescence with subfasciculate clusters of flowers in the axils of regular leaves {P weberiana). Receptacular, pedicellar, and peduncular zones loosely stippled; main axis densely stippled, transitional in D and E.

DEVELOPMENT OF FLOWER AND FRUIT According to Boke (1963b) the first stages of the development of the flower resemble those of the shoot. The observation that the primordia of bracts and perianth segments are hardly distinguishable from those of leaves is not surprising, but the basipetal initiation of stamen primordia simultaneous with or even after carpel initiation merits attention. Boke's detailed description of the ontogeny of the flowers of Pereskia lychnidiflora, P. aculeata, and P. diaz-romeroana gives important insight into the nature of the floral cup, the interpretation of the gy­ noecium, and the question of parietal versus axile placentation (Boke, 1963b, 1966, 1968), and Ross (1982) illustrated the ontogeny in P. bleo with SEM- micrographs. Longitudinal sections of the flowers of all species and of the fruit of all but two species could be studied and compared with Boke's results. These 18 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 1986] PERESKIA (CACTACEAE) 19

FIG. 9. Generalized flower structure and terminology, a, pedicel (peduncle); b, lower receptacular bracts; c, receptacular areole with short tomentum; d, upper receptacular bracts; e, upper receptacular areole with longer hairs; f, sepaloid perianth segments; g, petaloid perianth segments; h, staminodial hairs; i, stamens; j (in downward sequence), stigma lobes, style, locule with septal ridges at the ovary roof, ovules, slightly concave ovary floor. observations agree with the theoretical implications of Boke better than with those of Buxbaum, particularly in regard to the gynoecium. For the species of Pereskia with perigynous flowers the following characteriza­ tion of development probably can be generalized: "At anthesis the gynoecium appears to be slightly sunken into the floral cup, its sides adnate to the bases of the stamens. Following pollination, the floral cup deepens markedly. . . . The fruit appears to be distinctly inferior" (Boke, 1963b, p. 857). In P. lychnidiflora the whole flower axis elongates, becoming the columella. In others, as in, e.g., P. grandifolia, distortions mainly affect the peripheral parts of the receptacular cup around the inferior ovary, which thus becomes completely enclosed in fruit as in P. lychnidiflora, but without a columella. In P. bleo, post- floral growth of the ovary roof results in a very large naked umbilicus. It should be noted that the term flower cup, receptacular cup, or receptacle for short, is used here in the broader sense of Boke (1963b). Buxbaum (1953, 1957) used the

FIG. 8. Inflorescence and sequence of anthesis in Pereskia aculeata (cult. Jardin Exotique, Leuen­ berger 2817, B). Explanation of symbols: small circles = young flower buds; large circles = mature flower buds; double circles = open flowers; filled circles = withered flowers. For simplicity, bracts on lateral peduncles, pedicels, and receptacle of flower are omitted. 20 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

P. lychnidiflora

umbilicus: flower: narrow perigynous ovary roof: ovary: enclosed superior to half inferior columella: accrescent placentation: "basal-axile"

P. weberiana (similar: species no. 1, 2, 3, 6) flower: umbilicus: narrow perigynous ovary: ovary roof: enclosed superior to half inferior columella: placentation: indistinct to "basal" slightly accrescent

P. sacharosa (similar: species no. 7, 12, 14, 15, 16) umbilicus: flower: • * narrow perigynous ovary roof: ovary: enclosed half inferior columella: placentation: none parietal

umbilicus: large ovary roof: accrescent and flower: exposed epigynous columella: none ovary: inferior placentation: parietal

P. portulacifolia (similar: species no. 8, lO)

flower: *~ umbilicus: epigynous medium ovary: ovary roof: inferior exposed placentation: columella: parietal none

FIG. 10. Flower and fruit structure of Pereskia (schematic, not in scale; carpellary tissue stippled, species with similar to identical flower and fruit types are listed in parentheses). Left two columns: flowers, l.s.; right two columns: fruits, l.s.

term "receptacle" exclusively for the cylindric elongation of the flower cup above the level of the adnation of the carpels, that is, for the receptacular tube present in many Cactaceae, but less conspicuous in Pereskia. The lower part of the flower axis at the level of the ovary is called "pericarpellum" by Buxbaum, a descriptive term useful in the subfamily . In Pereskia, such a distinction is not feasible, and the term is not used here. A schematic drawing of a generalized flower of Pereskia is presented in Figure 9. As to the fruit (Fig. 10), it should be 1986] PERESKIA (CACTACEAE) 21

clear that the fruit wall is not formed by the carpels alone, hence it is not a true pericarp, but rather, it is formed mainly by the flower cup (receptacle), which surrounds or encloses the pericarp, leaving a narrow to broad depression (um­ bilicus). The relative position of carpellary and receptacular tissues in flower and fruit are outlined in Figure 10.

RECEPTACLE, RECEPTACULAR BRACTS, AND AREOLES The receptacle of Pereskia flowers is composed of a few to many nodes, and internodes of varying length. It is terete to angled-turbinate, cup-shaped, or sub­ globular, and gradually narrowed into a short stalk (peduncle, "pedicel"), or is subsessile to sessile (P. humboldtii, lateral flowers, Fig. 25A, B). The bracts are leaf-like to scaly and more or less fleshy. They are often not sharply delimited in shape and size from regular leaves, particularly in compound inflorescences. Towards the rim of the receptacle the bracts often diminish in size, and the uppermost are scale-like in many species, and appressed to the bud. These are called "upper bracts" in the descriptive part, and their number and shape is a distinctive character for some species. The receptacular areoles always have at least a trace of tomentum, and in some species produce longer hairs, rarely even spines. The presence of longer hairs seems to be a good diagnostic character. Spines are formed regularly in the receptacle of P. aculeata, but only occasionally in individual specimens of other species (P. sacharosa, P. stenantha). The delim­ itation of bracts and sepaloid perianth segments is illustrated in Figure 11A-I. The limit between bracts and sepaloids is apparently at the point where the foliar appendages increase in size again. The sepaloids also have a thinner blade, show transitional stages towards petaloid pigmentation, and lack an axillary areole. Receptacular bracts are numerous and more or less regularly spaced, giving the receptacle a cone-like appearance, as in P. lychnidiflora, or they may be inserted on raised podaria, as in the inner flowers of the inflorescence of P. grandifolia, or be absent from the lower part of the receptacle, as in P. bleo. The bracts are usually green, sometimes tinged with red at the margin, or entirely pigmented as in P. grandifolia var. violacea. The lower bracts are sometimes accrescent with the fruit but often deciduous at maturity. The uppermost bracts are often persistent at or in the umbilicus of the fruit.

SEPALOID AND PETALOID PERIANTH SEGMENTS Sepaloids and petaloids in Pereskia differ mainly in size, shape, and pigmen­ tation, and they are often difficult to delimit. Therefore I prefer not to use the term "petals" and "sepals," but "tepals" does not seem wholly adequate either. In Figure 11A-I, the second arrow of each sequence marks the transition between sepaloids and petaloids. The sepaloids are conspicuous in bud, imbricate, thicker, and with broader bases than the petaloids, often greenish but with petaloid pig­ mentation towards the margin. Their shape varies from broadly ovate to obovate, with acuminate, acute, or rounded apices. Petaloids of several species are illustrated in Figure 11A-J. The largest ones are found in P. nemorosa, but some species show great variability of shape as well as size, as exemplified by selected petaloids of P. grandifolia, illustrated in Figure 10J. Pereskia lychnidiflora is distinctive with its irregularly dentate-lacin- iate petaloids, and P. stenantha is characterized by its narrow, erect petaloids, which are more fleshy at the base than in most other species. The lamina is very delicate in some species, particularly in P. aculeata, especially in the forms with 22 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

ooO

FIG. 11. Receptacular bracts and perianth segments. A-I, sequence of receptacular bracts and perianth segments on individual flowers of different species. A, Pereskia bleo {cult. hort. Berol. 277- 01-80-80). B, P. nemorosa {cult. hort. Berol. 090-01-79-80). C, P. aculeata {cult. SSZ). D, P. lych­ nidiflora {Leuenberger & Schiers 2513). E, P portulacifolia {Leuenberger 3048). F, P. stenantha {Leuenberger 3073). G, P. quisqueyana {Leuenberger 3044). H, P weberiana {cult. hort. Berol. 037- 01-77-80). I, P. bahiensis {Leuenberger 3061). J, variability of perianth segments in different specimens of the same species; Pereskia grandifolia var. grandifolia (all cult. hort. Berol). All half size. (Note: dots represent bracts and perianth segments omitted in the drawing; the first arrow of each sequence marks the transition between receptacular bracts and sepaloids, the second one the transition between sepaloids and petaloids.) 1986] PERESKIA (CACTACEAE) 23

greenish-white flowers where it is almost translucent. The flowers wither on the day of anthesis, rarely on the second day. Flower color was observed to fade during anthesis in P. grandifolia and seems to be rather variable in this species and even on individual plants, possibly due to environmental conditions. An interesting color change was observed by Padre Julio Cicero (pers. comm.) in P. bleo, where the brightly scarlet petaloids turn pinkish at the margin at high temperatures during anthesis.

ANDROECIUM3 Development The initiation of the androecium in Cactaceae has been studied by Buxbaum (1953), Ross (1982), and in Pereskia particularly by Boke (1963b, 1966, 1968), who showed that an annular meristematic region extending from the carpel pri­ mordia to the inner perianth segments gives rise to numerous stamens in basipetal (=centrifugal) sequence. The term centrifugal seems preferable, because in flowers with an elongated receptacle, i.e., in those of most genera of the other subfamilies (Opuntioideae and Cactoideae, except Rhipsalis), the outer stamens are inserted at a much higher level due to growth of the receptacular tube. In Pereskia this situation is less pronounced although evident in P. lychnidiflora. According to Boke (1968, p. 1256), "there seems to be nothing unusual about stamen devel­ opment" except for the sequence, which is evident in Figure 13A. Stamens The number of stamens ranges between 50 and 100 in the small-flowered Andean species and 200 to 300 in P. guamacho and P. lychnidiflora. Stamens are more or less spreading, with the anthers dispersed in the open flowers in, e.g., P. guamacho and P. aculeata, and densely spreading in P. lychnidiflora and the Caribbean species. They are more or less clustered around the style in the other species, and erect and densely surrounding the style in P. bleo. Style and stigma are hidden among the stamens in P. lychnidiflora and the staminate flowers of the Caribbean species, while they surpass the level of the anthers in the other species, at least at a stage where the stigma is receptive. The filaments are terete, smooth, and narrowed just below the point of attach­ ment to the connective. They are mostly colorless at the base but in some species conspicuously pigmented farther up, either correlated with the color of the pet­ aloids, as in P. diaz-romeroana, P. bleo, and P. guamacho, or contrasting with the petaloids, as in forms of P. aculeata where yellow, orange, or red filaments occur. Anthers in Pereskia are of the normal two-locular, tetrasporangiate type, dor- sifixed, with the sacs joined for about half to most of their length, with complete longitudinal dehiscence and rectangular in outline when dehisced. The range of size and shape is moderate within the genus, but differences occur even between closely related species: P. portulacifolia has nearly square anthers, and P. quis­ queyana has long rectangular anthers. Anthers and pollen are yellow or cream in

3 A detailed study of the development of the androecium and gynoecium of Pereskia with SEM- methods is currently in preparation by Sylvia Schwitalla at the Institute for Systematic Botany of the University of Heidelberg. See also Leins and Schwitalla (1986), a preliminary paper which appeared in February 1986, after termination of this manuscript. 24 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 most species, but in P. portulacifolia, P. zinniiflora, and P. quisqueyana the an­ thers are pinkish. Staminodes The only species where structures probably best interpreted as staminodes occur regularly is P. nemorosa. Clusters of uniseriate and multiseriate, curled trichomes are found between the petaloids and stamens in flowers of this species (Fig. 14B). In fresh flowers, the trichomes are turgid and glassy white, in dried flower they are similar to trichomes of the areoles. However, they are more curly, and their position is unlike that of receptacular areoles. Fascicles of the curly hairs occur alternating with and sometimes opposite to the innermost petaloids. Individual larger appendages transitional between trichomes and filaments suggest the prob­ ably staminodial nature of these appendages. I have seen individual curly hairs only in one case in another species, in a single flower of a cultivated specimen of P. diaz-romeroana (Rauh 40627). Nectaries In the three species of Pereskia studied in detail by Boke, longitudinal and transverse sections of flowers clearly show nectarial tissues. In P. lychnidiflora (Boke, 1963b, p. 853) the nectary is small and "arises at the base of the innermost stamens and takes the form of a narrow ring of secretory tissue surrounding the base of the gynoecium." A similar small nectary is discernible in P. diaz-rome­ roana (Boke, 1968, p. 1259). In P. aculeata the inner side of the short but con­ spicuous floral tube forms a cylindrical nectary closely surrounding the style base and periphery of the ovary (Boke, 1966, p. 542). Buxbaum (1953) studied the floral nectaries of many other Cactaceae and found them also at the base of the primary stamens, or on what he considered to be decurrent filament bases. Al­ though no histological studies were made, the examination of sectioned fresh flowers revealed nectaries at basically the same position as in the species studied by Boke. While the nectarial ring-shaped furrow is very narrow in most species, a few exceptions are noteworthy. In P. stenantha the furrow is broader but constricted above the secretory tissue by bulging filament bases (Fig. 49E). A weakly defined nectar chamber is the result. A short elongation of the receptacular rim combined with a flat ovary roof forms a broad, moat-like furrow in the pistillate flowers of the Caribbean species, e.g., in P. portulacifolia (Fig. 14H) and P. sp. A (Fig. 14C). In P. bleo the broad and flat ovary roof is surrounded by knee-like projections of the filament bases, concealing a narrow ring of secretory tissue (Fig. 39C).

GYNOECIUM Long before the morphology of the ovary of Pereskia had been properly in­ vestigated, its characters nevertheless had been used to split the genus in two. Studies by Buxbaum (1940, 1944, 1948, 1953) on the gynoecium of P. aculeata and P. sacharosa and by Tiagi (1955) of P. grandifolia (misidentified as P. bleo) showed remarkable differences and the inherent difficulty of interpretation of carpel shape. Again it was the publications of Boke (1963b, 1964, 1966, 1968) which elucidated the morphology of the gynoecium, although according to this author there already seemed to be universal agreement that in Cactaceae the gynoecium is sunken in the end of a modified branch. The recurrent bundles 1^86] PERESKIA (CACTACEAE) 25

supplying the ovary and the ontogenetic distortions during bud and fruit devel­ opment present strong evidence for this. However, widely diverging opinions still exist on the interpretation of placentation. Prior to Buxbaum's and Boke's studies, placentation had been thought to be of the conventional parietal type and con­ tinued to be so by some authors (Hutchinson, 1967). Detailed investigations of several species by Boke not only showed diversity of characters of the gynoecium but also led to a new interpretation of the gynoecium of Cactaceae in general (Boke, 1964), which differs from that of Buxbaum (1944, 1948, 1953, 1957). Buxbaum based his conclusions on the peltate theory of carpel form and on the assumption that carpels with one or few ovules were primitive. Boke (1964, p. 609) concluded that parietal placentation in Cactaceae "appears to have been derived from axile placentation and is not strictly homologous with parietal pla­ centation in other families" and that "the inferior position has been derived in a very different manner from that in most flowering plants." Boke's interpretation of the carpel of Pereskia is illustrated in Figure 12, and his model has been amplified here to distinguish more transitional forms from superior to inferior ovaries and from "axile" to "basal" and parietal placentation. Possible evolu­ tionary tendencies are indicated by arrows. The terms "axile" and "basal" refer to the position within the locule only. It is understood that the placentae are initiated on carpellary tissue on or near the margin of the carpels. However, in species with a sunken gynoecium, the free tip of the receptacle forms part of the locule floor (Figs. 12C, D, 13B-E). Stages of the development of the gynoecium of the type illustrated in Figure 12C are shown in the SEM-micrographs of Figure 13 A-F, which clearly show that the ovules are initiated at the lower end of the septal ridges. The extremes of gynoecium structure within the genus are found in P. lychnidiflora, with an essentially superior ovary and individual carpel locules, and in P. sp. A, with one large locule, low septal ridges on the ovary roof, and extended, U-shaped parietal placentae (Fig. 14C). The morphology of the gynoe­ cium of P. lychnidiflora was discussed in detail and illustrated in a stylized drawing by Boke (1963b, p. 854, fig 32). The "peltate theory" of Buxbaum (1953) is not discussed further at this point, because comparison of gynoecium structure of all species and evaluation of other features and their possible evolutionary signifi­ cance suggest that the model designed by Boke (1964) is preferable to that of Buxbaum (1953), whose theory infers that parietal placentation was derived from an elongated and displaced cross zone of peltate and originally one-seeded carpels. No detailed study of the gynoecium of the Caribbean species has been made, and even for this monograph, pistillate flowers were not available for closer examination until after completion of relevant chapters and illustrations. The elongated U-shaped placentae in the large locule of P. portulacifolia (Fig. 14H) and P. sp. A (Fig. 14C) support Boke's model of the cactus gynoecium. Carpel shape approaches the interpretive diagram illustrated in Figure 12D. However, the placentae are more elongated and ovules are more numerous. Gynoecia of staminate flowers are rudimentary (Figs. 34B, 37C, 38D).

OVULES The morphology of the ovules of Cactaceae was described by Buxbaum (1958, 1966). However, only one species of Pereskia has been studied in detail (Neumann, 1935): the ovules of P. nemorosa (as P. amapola var. argentina) were described as bitegmic, crassinucellate, and more or less distinctly campylotropous. Engleman (1960) compared three other cacti with Neumann's illustrations of P. nemorosa and his own slides of P. sacharosa, and found similarities between some Cac- 26 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

r\ B

rec rec FIG. 12. A-D, interpretive diagrams of carpel structure and placentation. A, interpretation of a carpel of Pereskia lychnidiflora. B, interpretation of a carpel of P. aculeata. C, interpretation of a carpel of the type represented by P. sacharosa and several other species with semi-inferior gynoecium and conical ovary roof and style base. D, interpretation of a carpel of P. bleo, a species with distinctly inferior locule and horizontal, free ovary roof. The arrows indicate possible evolutionary lines as supported by other character correlations; d, dorsal carpel bundle; r, recurrent bundle system; rec, free tip of receptacle; v, ventral bundle system; broken lines indicate theoretical carpel limits (diagrams A-C modified from Boke, 1964, p. 608, 609). toideae and Pereskia, but noted the larger funicular hump in Pereskia. The ovules of 14 species seen so far are all similar, anatropous to slightly campylotropous at the time of anthesis and varying in length from ca. 0.4 mm in P. diaz-romeroana to ca. 0.9 mm in P. sp. A. The funicles are of variable length and single or in fascicles, but never branched as in certain Cactoideae (Buxbaum, 1957; Vaupel, 1925). They are partly covered by papillae pointing towards the micropyle, a character typical for many (Centrospermae) according to Neumann (1935), who interpreted this as a functional feature in ectotropous guidance of the pollen tube (Fig. 13D, F). Locules without ovules or abnormal ovules were found in staminate flowers of P. portulacifolia, P. quisqueyana, and P. zinniiflora. These flowers are apparently unisexually staminate, with short style, filiform stigma lobes without papillae, and a very small locule (Figs. 34D, 37C, 38D). In a cultivated specimen of P. zinniiflora the staminate flowers showed teratological features in the gynoecium. The locule was filled by another series of carpels or stamens with abnormal anthers rising from the parietal placentae.

INFRUCTESCENCE AND FRUIT The development of flower and fruit has been outlined above. In species with proliferating flowers, infructescences with chain-like aggregates of fruits develop. 1986] PERESKIA (CACTACEAE) 27

FIG. 13. A-F, development of the gynoecium of Pereskia grandifolia var. grandifolia (A-C from cult. hort. Berol. 047-01-78-80; D-F from cult. Palmengarten Frankfurt). SEM-micrographs A-F by courtesy of Sylvia Schwitalla, Institute for Systematic Botany, University of Heidelberg. A, stamen and carpel primordia (primordia of perianth segments removed except for the two innermost); B, ovary of younger bud, l.s., with broadening locule floor and ovule primordia around the base of the septal ridges of the ovary roof; C, the same at an advanced stage; D, young ovules and papillate septal ridges; E, locule of older bud; F, ovules with funicular papillae.

If only the distal flowers are pollinated, the floral parts of the non-pollinated flowers wither down to the ovary, but the receptacle remains intact as a link to the distal flowers and fruits. Therefore, the infructescence of these species can vary from dense to lax. There is some evidence that proliferation is prolonged if the proximal flowers are not pollinated. Long chains of receptacles and fruit are the result in, e.g., P. grandifolia var. violacea (Fig. 14D, E). The fruit of Pereskia is of a predominantly receptacular nature, with only minor parts formed by carpellary tissue. Fruit shape and size is diverse, ranging from the small, smooth, black, and juicy berries of P. diaz-romeroana to the pear-sized and -shaped, irregularly angled, greenish, and sometimes bracteate fruits with 28 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 14. A-H, flower and fruit characters of Pereskia. A, P. stenantha {Leuenberger et al. 3081), flower at full anthesis, with spreading bracts and erect sepaloids and petaloids. B, P. nemorosa {cult, hort. Berol. 164-01-80-70), flower, l.s., exposing long hairs in the axils of the receptacular bracts and tufts of staminodial hairs between the petaloids and filaments. C, Pereskia sp. A. {Marcano & Cicero s.n., cult. hort. Berol. 259-02-82-30), pistillate flower, l.s., with conspicuous stigma lobes (darkened by FAA-fixation) and large locule with elongate U-shaped placentae. D, E, P. grandifolia var. violacea {Horst HU 226, cult. hort. Berol.). D, branch of inflorescence with chain-like proliferation from the receptacles of unpollinated flowers; E, branch of infructescence with receptacles of unpollinated flowers bearing the distal fruits. F, P. grandifolia var. grandifolia {cult. hort. Berol. 168-02-79-80), old fruit 19 8 " J PERESKIA (CACTACEAE) 29 tough, fleshy, mucilaginous fruit walls seen in P. grandifolia and related species. Different relative growth of parts of the receptacle and gynoecium is responsible for the distinctive fruit characters. Growth of the ovary roof leads to the broadly turbinate fruit of P. bleo with its distinct and very large, smooth umbilicus (Figs. 10, 39E). Moderate growth of the ovary roof as well as of the flanks of the receptacle results in the globular to ficiform fruit of P. portulacifolia, with an equally smooth but smaller umbilicus (Figs. 10, 37F). Enhanced growth of the upper rim and flanks of the receptacle nearly encloses the ovary roof in species with obovate to pyriform fruit, leaving a narrow umbilicus with more or less persistent flower remnants (Fig. 10, P. sacharosa). In most species, the carpels play a minor quan­ titative role in fruit development, but they may contribute to the formation of a mucilaginous pulp, at least at the ovary roof. An outstanding feature is the presence of a columella developing from the accrescent free tip of the receptacle in fruits of P. lychnidiflora (Figs. 10, 29F, I). This species is also distinguished by the occurrence of stinging sclereids in the wall of the moat-like locule of the fruit. The color of mature fruits ranges from greenish to yellow (P. bleo) and chocolate brown (P. aureiflora) and vinaceous to black in the small-leaved Andean species. In most species the fruit wall is thick and mucilaginous or juicy, but in P. por­ tulacifolia and probably the other Caribbean species as well it is relatively thin, and the locule is not hollow but filled by pulp formed by the funicles, a character widespread in Cactaceae.

SEEDS The seeds of Pereskia are medium-sized to fairly large compared to those of other genera of Cactaceae. Seed size is not correlated with fruit size and varies between a minimum length of 1.8 mm in P. weberiana and a maximum of 7.5 mm in P. bleo. Seed shape ranges from obovate to subreniform or lenticular in lateral view (Fig. 15A-D) and the seeds are more or less flattened to elliptic in transsection. The seed coat is black and smooth or nearly so, usually glossy, and in some species slightly ribbed along the periphery with rows of rectangular testa cells (Fig. 15D). The micropyle is discernible at the basal umbo adjoining the hilum, which is separate and covered by a flat to pulvinate or protruding, whitish, spongy tissue with central funicle scar (Fig. 15C, E, F). Seed size and shape is characteristic for some species, but a considerable amount of variation has to be taken in account both within one sample and within the species as a whole. Seed number also varies greatly within the genus. Only two to five seeds are normally observed in fruits of P. aculeata, which are also distinctive because they are flattened to slightly concave on one side, apparently the side adjoining the col­ umella in fruit.

EMBRYO The internal structure of the seeds was studied by simply breaking or cutting the tough to rather brittle seed coat and removing the black testa of seeds mounted

seen from above, with sunken umbilicus, bract scars, and oily exudations after long storage. G, H, Pereskia portulacifolia {Zanoni 35204), G, from left to right, flower bud, pistillate flower, l.s., and young fruit after dehiscence of flower remnant, exposing umbilicus; H, locule with ovules, receptacular rim with dwarfed stamens and flat ovary roof, stigma lobes darkened by fixation in FAA. 30 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

h i \^ j k *^ I m

5mm

F

FIG. 15. A-F, seeds of Pereskia. A, macro-photographs of a series of seeds of different species; a, P. aculeata {Araujo 766); b, P. bahiensis {Leuenberger et al. 3063); c, P. stenantha {Leuenberger et al. 3072); d, P. nemorosa {Solomon 6918); e, P sacharosa {Ahlgrimm 41); f, P. guamacho {Araque 24); g, P. diaz-romeroana {Knize 943); h, P. aureiflora {Leuenberger & Martinelli 3055); i, P. bleo {Pittier 4704); j, P. grandifolia var. grandifolia (cult. hort. Berol. 038-04-77-80); k, P. grandifolia var. violacea {Leuenberger et al. 3060); 1, P. lychnidiflora {Leuenberger & Schiers 2531); m, P. portulacifolia {Czer­ wenka s.n.); n, P. zinniiflora {Wood & Atchison 74040). B-F, SEM-micrographs of seeds and hilar regions of seeds. B, C, P. lychnidiflora {Leuenberger & Schiers 2508); B, lateral; C, hilar view. D, P. weberiana {cult. hort. Berol. 258-08-80-50), lateral view with hilum at the lower end (center) and micropylar nose (left). E, P. sacharosa {Kiesling s.n.), micropylar nose, micropyle and large hilum with central funicle scar. F, P bleo {Fairchild Tropical Garden), hilum. on a glass slide with glue. All species examined have curved embryos and copious perisperm, both tightly enclosed by a thin membrane of dead cells. This mem­ brane, which is also found in seeds of other Cactaceae, was classified by Buxbaum (1955) as the "inner testa," while Barthlott and Voit (1979) more cautiously call it a layer of dead cells composed of elements of the degenerated inner integument, of the nucellus, and possibly of the endosperm. Its ontogeny in Pereskia has not yet been studied. In P. grandifolia no trace of endosperm is left in mature seeds, while in immature seeds with brownish testae the embryo is still embedded in endosperm. Fresh seeds of P. bleo did not germinate at all in repeated sowings, and examination of such seeds showed that the embryo was not mature but still embedded in endosperm, even though the testa was entirely black. This may suggest that at least in some species the embryo is not mature at the time of seed dispersal. 1986] PERESKIA (CACTACEAE) 31

Cotyledons These are folded with their laminae flat in those species with small to medium- sized seeds. In species with larger seeds, the embryo is superficially similar, but aestivation of the cotyledons is obvolute. This also becomes evident from the folded appearance of the cotyledons in the first stages of germination in, e.g., P. bleo, P. aculeata, and P. grandifolia. The embryo is cream-colored and turns green only during germination. Perisperm A hard, starchy perisperm occupies the space left on the ventral side of the seed by the strongly curved embryo. The perisperm is used up during germination. It may attain about one fourth of the total volume of the seed. Similar amounts of perisperm are found in other Cactaceae (Buxbaum, 1953, 1966), with a tendency towards reduction of perisperm along phylogenetic lines postulated by that author.

SEEDLINGS No young seedlings have been observed by the author in their natural habitat and all previous information on seedlings was also based on cultivated material (Bailey, 1965; Boke, 1963a; Buxbaum, 1953; Duke, 1969). I have seen one seedling collection of P. sacharosa (collected by K. Ahlgrimm near Camiri, Bolivia), which in leafless condition resembled a Quiabentia due to its very stout epicotyl of 7 cm length and over 1 cm diam. I observed older seedling plants of P. lychnidiflora in moderate number on a pasture near Teculutan, Guatemala, and the plants grown in the greenhouse from seeds collected at the same locality showed no differences. The first stages of germination, illustrated also by Bregmen (1982, p. 226, figs. 7, 8), seem to be similar in all species observed. The seed coat usually splits lengthwise or in a more or less irregular way at the base of the seed. Different stages of germination are shown for P. nemorosa in Figure 16, drawn from a series of photographs taken over a period of five weeks. Germination normally takes place after a few days. From my experience fresh seed does not germinate well. Best results were observed after several months to one or two years, but five-year- old seed of P. lychnidiflora still yielded more than 50 percent germination. Growth of the seedling depends much on nutrition, temperature, and light. Under experimental condition (greenhouse or sunny window) the hypocotyl at­ tains its full length within two or three weeks. Primary growth in thickness con­ tinues for weeks and the cotyledons also grow even after development of the primary shoot. Seedlings of all species except P. aureiflora, P. guamacho, P. humboldtii, P. quisqueyana, P. zinniiflora, and P. sp. A could be studied. They are all basically alike, although differences in size, pigmentation, and nyctinastic behavior occur. Three size classes more or less correlated to seed size can be distinguished. Seed­ lings of species with seed between 3.5 and 7.5 mm in length are about the same size. Under experimental conditions two- to three-week-old seedlings of this category have a hypocotyl of 3-5 cm long and ovate cotyledons 3-4 cm long, e.g., P. bahiensis, P. grandifolia, and P. sacharosa. Pereskia lychnidiflora has medium- sized, slow-growing seedlings. The smallest seedlings with a hypocotyl of barely 2 cm and cotyledons of ca. 1 cm length are found in the Andean species. These differ in several characters, such as red pigmentation of the lower side of the 32 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

1 cm I

B

FIG. 16. Germination of seed in Pereskia nemorosa {Gonjian s.n.). A, B, sequences showing the development of two seedlings from germination to the age of five weeks at intervals of increasing length, from 12 hours (far left) to several days (far right). Drawing based on a series of photographs by the author. cotyledons, lack of nyctinastic movement of the cotyledons, and finely papillate hypocotyl (observed in P. weberiana). All other species studied fold up their cotyledons at night for the first two to three weeks prior to formation of the primary shoot. Spiny areoles, but no brachyblast leaves develop in the axils of the cotyledons. In P. weberiana and P. diaz-romeroana the hypocotyl is sometimes thickened at the base, but the swelling does not continue into the primary root. Thickened roots seem to develop only at a later stage of ontogeny. A weak lignotuber was observed in young plants of P. nemorosa. Seedlings of few days to one year old are shown in Figure 17A-E. Nyctinastic movement of leaves occurs in P. gran­ difolia var. grandifolia, where the primary leaves of seedlings are drooping at night. 1986] PERESKIA (CACTACEAE) 33

FIG. 17. Seedlings of Pereskia. A, Pereskia lychnidiflora {Leuenberger & Schiers 2508), different seedlings between one day and one year old. B, P. aculeata {cult. hort. Berol. 128-01-82-70), one week and three months old. C, P. sacharosa {Ahlgrimm 68), one week and three months old. D, P. grandifolia {cult. hort. Berol. 047-01-78-80), different seedlings from three days to three months old. E, P. we­ beriana {cult. hort. Berol. 258-08-80-84), one week and three months old. 34 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Anatomy A number of anatomical features of Pereskia have been studied in detail by Bailey, namely the xylem (Bailey, 1962, 1963a, 1963b, 1963c, 1963d, 1966; Bailey & Srivastava, 1962), the vascular system (Gibson, 1976), the form and distribution of oxalate crystals and druses (Bailey, 1961b), the foliar vasculature (Bailey, 1960, 1965), and the structure and distribution of sclerenchyma in phloem and cortex (Bailey, 1961a). These studies provided basic data for the discussion of evolu­ tionary aspects, and some tentative taxonomic conclusions were outlined by Bailey and reviewed by Gibson (1975). However, anatomical data should not be con­ sidered in isolation. Anatomical similarities in the plants investigated can reflect close relationship of well delimited species, or they can be explained by the fact that the samples received under different names actually represent conspecific taxa or that they were misidentified. All three cases in fact occur in the material studied by Bailey. Examination of other characters during this revision clearly revealed that the taxa suggested to be conspecific by Gibson (1975), based on Bailey's studies (P. autumnalis, P. pititache, and P. nicoyana), are indeed the same as P. lychnidiflora, and that P. colombiana is a synonym of P. guamacho. On the other hand, three other taxa referred to as possibly conspecific by Gibson (1975), P. humboldtii, P. diaz-romeroana, and P. weberiana, as suggested by anatomical data, prove to be distinct even though closely related. In the case of P. bleo and P. grandifolia (Bailey, 1963c; Gibson, 1975) the material from Costa Rica {Rodriguez s.n.) identified as P. bleo was undoubtedly P. grandifolia, the only species cultivated and subspontaneous in that area, and likely to be confused with P. bleo, a species which has not been reported from Costa Rica. Hence the similarity in this case is due to a misidentification. The limited number of samples makes it difficult to evaluate the taxonomic significance of anatomical data, but conspicuous and consistent differences in structure and distribution of sclerenchyma, succulence of roots, and presence or absence of stomata on the stem seem to contribute useful complementary data with a taxonomic bearing. Only a few points of Bailey's detailed work are sum­ marized here and some complementary observations added.

ROOT SUCCULENCE Investigations by Bailey (1963a) on the root swellings of P. diaz-romeroana and other Andean species show that a core of unlignified cells is formed by the vascular cambium. Primary xylem is often discernible to the naked eye as a ring of bundles near the center of the axis. Secondary xylem with libriform fibers is formed only at the final stage of cambial activity, i.e., after formation of varying amounts of soft tissue. Radial seriation of these soft derivatives of the cambium is lost soon due to enlargement and displacement of the cells which have large vacuoles containing water and starch (Bailey, 1963a; Gibson, 1975). The structure of root swellings larger than those reported by Bailey (1963a) is shown in Figure 3C, F for P. quisqueyana and P. weberiana. The structure of the core is basically alike, but the sclerenchymatous cells of the cortex (discussed below) are completely different in the two species.

EPIDERMIS The difficulty of obtaining comparable data on epidermal features has been discussed by Bailey (1964). As cultivated material of nearly all the species was 1986] PERESKIA (CACTACEAE) 35

available during preparation of this revision, hand sectioned material of com­ parable parts of the stem grown under the same conditions was scanned for possibly useful characters. All taxa except P. humboldtii var. rauhii have a smooth epidermis with cell sizes between 10 and 30 /xm (height of cell in transverse section) and cuticle of 2-20 /mi thickness. In those species with precocious periderm formation (Table I), the cuticle is thin and inconspicuous. Epidermis which has not begun to make periderm by periclinal divisions can only be observed on very young stems where the internodes are still elongating. In species with retarded periderm formation the cuticle is conspicuous, measuring about 6-10 Aim in the Andean species and up to 20 /Jim or more in Pereskia nemorosa and P. stenantha, where it almost equals the height of the epidermal cell. In P. grandifolia and P. bahiensis wedge- shaped accumulations of cutin extend inward between the epidermal cells. The same is to a minor degree observed in other species. A remarkable feature unique in the genus, but not within the family, is the finely papillate epidermis of P. humboldtii var. rauhii. Stout papillae to ca. 50 jam long and 10-15 /mi thick are formed on about one fourth to one half of all epidermal cells. The cuticle of the papillae is of equal thickness as that of the rest of the cell surface (Figs. 18A-C, 25N).

STOMATA According to Eggli (1984), the stomata on leaves of Pereskia are of the paral- lelocytic type, while those on stems, if present, are not of the parallelocytic type common to Cactoideae, but termed "opuntioid" by Eggli (1984, p. 200). Stomata density on leaves is ca. 30-80 mm2 and nearly equal on both surfaces, or higher on the lower surface (in P. aculeata, P. bleo, and P. grandifolia, according to Eggli, 1984). Stomata are absent from the stems of a number of species, where periderm formation is precocious, and they are lacking in at least one species with retarded periderm formation, P. nemorosa. In P. bahiensis and P. sacharosa usually only few scattered stomata are found near the areoles. Large intercellular cavities below the numerous stomata are conspicuous as whitish dots on the stem surface of living plants of P. grandifolia (Fig. 6H). The highest density of stomata on stems is found in the Andean species with ca. 40 per mm2. The presence of stomata is not correlated with the presence or absence of brachyblast leaves in the species.

LENTICELS In two species, P. guamacho and P. aureiflora (Fig. 6J), lenticels are numerous and conspicuous on younger stems, and in P. aculeata their number apparently varies from specimen to specimen. Scattered, prominent lenticels occur on stems of P. bleo (Fig. 5H), P. quisqueyana, and P. sp. A, where they contrast with the greenish stem by their pale brown color. Mauseth (1984) noted and illustrated the lenticels of P. aculeata.

PERIDERM Precocious periderm formation is characteristic for about half of the species of Pereskia and is a feature shared by but probably no other genus of Cactaceae. In the other species periderm formation is retarded for about one year or more, and anticlinal divisions of epidermal cells compensate for primary growth in thickness of the stem. In highly succulent Cactaceae the epidermis stays intact 36 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

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V <•«*! F 100 urn J 10pm ,'; FIG. 18. A-J, anatomical features and chromosomes of Pereskia. A-C, P. humboldtii var. rauhii {Rauh 40184), papillate epidermis at increasing magnification. D, E, P. aureiflora {Leuenberger & Martinelli 3054), simple calcium oxalate crystals from the cortex of older branch, F, P. weberiana {cult. hort. Berol. 037-01-77-80), druses of calcium oxalate crystals from cortex of stem. G, H, P. lychnidiflora {Leuenberger & Schiers 2508), fusiform sclereids from fruit wall. I, P. weberiana {cult, hort. Berol. 037-01-77-80), fusiform sclereids in the secondary phloem of broken stem. J, P. stenantha {Leuenberger et al. 3072), somatic chromosomes, metaphase plate. 38 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 for many years, or a very thin, translucent periderm may be formed (Mauseth, 1984). The first cork cambium is initiated by periclinal divisions in the lower half of the epidermal cells, producing a dense layer of thin-walled, translucent to corky cells covered by the more or less intact outer portion of the epidermis and cuticle. In living stems, periderm formation is usually evident macroscopically by the color change from green to olive-green or pale brown. In species with precocious periderm formation it seems to begin simultaneously as soon as longitudinal growth of the shoot has stopped, but has no corky appearance. In species with retarded periderm formation, specific patterns of periderm formation occur. In P. sacharosa and P. bahiensis it begins in more or less irregular blotches around the areoles, while in P. nemorosa and to a lesser extent in P. grandifolia, stra­ mineous longitudinal striations form by rupture of the epidermis and rapid for­ mation of many layers of cork cells (Fig. 61). Older bark becomes longitudinally fissured or irregularly flaky in most species.

CORTEX All species of Pereskia have a green cortex with numerous oxalate crystals or druses, and large mucilage cells. When the brownish-gray bark is peeled off from old trunks of P. lychnidiflora, a dark green layer appears below the phellogen. The cortex is thin to more or less fleshy and together with the pith and unlignified rays causes the half-succulence of younger stems. Sclereids initiated in the sec­ ondary phloem are a remarkable feature of the inner cortex and periderm of Pereskia.

SCLEREIDS Morphology and taxonomic significance of sclereids of the secondary phloem and cortex in stems and roots of Pereskia have been discussed by Bailey (1961a) and Gibson (1975), based on the investigation of material belonging to ten species as circumscribed in this monograph. Three groups were distinguished by Bailey, and the three types of sclereid form and distribution are confirmed by my own observations on additional species. The first group has simple fusiform sclereids to 1500 /urn long or longer and ca. 18-100 nm in diam., which are diffusely distributed. They occur in seemingly unrelated species such as P. aculeata, P. lychnidiflora, and the Andean group (see Table I). They were also the reason for the fluctuation in taxonomic position of the names here treated as synonyms of P. lychnidiflora: P. conzattii and P. piti­ tache. The skin-irritating sclereids falling from dry and broken twigs (and also present in mature fruits) were confused with glochids, hence the combinations of both names under Pereskiopsis, a genus of Opuntioideae. The sclereids are visible to the naked eye or with a hand lens in broken stems of P. weberiana. Sclereids protruding from broken tissues are illustrated in Figure 181, and individual scler­ eids from dried fruit of P. lychnidiflora in Figure 18G, H. The sclereids are numerous also in the cortex of the root swellings of P. weberiana. The second type, consisting of variably shorter sclereids aggregated in longitudinally oriented clusters, occurs in several species from South America. The aggregates are easily visible to the naked eye in dried pieces of cortex, and the individual sclereids attain a length of 1000 ixvci and a diameter of 200 /xm. Transitional forms may exist between the second and the third type, more or less isodiametric stone cells aggregated in massive, irregular clusters. They are conspicuous but of somewhat transitional shape in the cortex of older branches 19 8 6] PERESKIA (CACTACEAE) 3 9

of P. aureiflora and occur in typical form in the secondary growth of root swellings of P. quisqueyana, besides the species mentioned by Bailey (1961a). They are difficult to section as they are embedded in soft cortical tissue. The structure of the sclereids has been described in detail by Bailey (1961a). Their size ranges from 40 to over 100 /mi in diam., and the larger aggregates attain ca. 600 /xm diam. The sclereid type is to some extent correlated with the geographical distribution of the species (Fig. 22).

CRYSTALS AND DRUSES Older literature referring to crystals and druses of calcium oxalate in Cactaceae has been reviewed by Bailey (1961b), who discussed not only the difficulty of securing strictly comparable material but also the wide variation in size, form, and number of druses in Pereskia. The diffuse distribution of rotund forms of druses in the outermost layers of the cortex was found to be a taxonomically significant character for distinguishing Pereskia anatomically from the superfi­ cially similar genera Pereskiopsis and Quiabentia (subfamily Opuntioideae) with densely packed druses in the outermost, collenchymatous layer. There seems to be slight differentiation within the genus itself, both in form and arrangement of the druses. Bailey (1961b, p. 338), based on observations after different chemical treatments, concluded that the druses are not composed solely of calcium oxalate but contain organic substances as well. This merits further attention and is sup­ ported by my own experience with FAA-preserved material mounted in Kaiser's glycerine gelatine. Some druses showed considerable size increase by swelling. Druses are a conspicuous character in the cortex and pith of all species of Pereskia. The largest druses were found in P. grandifolia, attaining 60-80 utm in diam., whereas in the other species variable sizes between 20 and 60 Aim diam. were observed in the cortex and pith of twigs. In some species the druses are scattered, in others (e.g., the Andean group and P. guamacho) they are arranged in longitudinal chains of 3-10 druses. From the small number of samples it is impossible, however, to draw conclusions on the usefulness of this character. Simple crystals were reported by Bailey (1961b, p. 335), for the pith and cortex of P. lychnidiflora (as P. conzattii) and for the older secondary phloem of P. guamacho. Simple crystals of about the same size as entire druses or even larger occur in the cortex of older branches of P. aureiflora, where they are numerous in at least one layer of cells. The tissue is difficult to section both because of the crystals and the stone cells. Stellate druses of P. weberiana and simple rhombic crystals of P. aureiflora are illustrated by SEM-micrographs in Figure 18D-F.

MUCILAGE CELLS AND CAVITIES Examination of sections of young stems of most species, and of the flower axis and fruit shows that mucilage cells and cavities of varying size occur in all taxa. Numerous and very large cavities of up to more than 300 ixm diam. are found in stems of P. grandifolia, while their number is apparently much smaller in P. sacharosa. The Andean species also have smaller mucilage cells or cavities (ca. 30-70 /mi diam.), and in P. guamacho and P. zinniiflora no mucilage cavities were found in stem samples, although mucilage cavities are conspicuous in the flower axis and fruit of the same species. They attain more than 1 mm in diam. in the fruit wall of several species, e.g., in the cortex and columella of the fruit of P. lychnidiflora, of which Boke (1963a, p. 852, 1963b, p. 8, fig. 9), illustrated mucilage cavities of to 2.5 mm diam. When fresh or FAA-preserved, sectioned 40 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 fruits of P. lychnidiflora and P. grandifolia are put in water, the mucilage swells and exudes in large amounts from the cut mucilage cavities.

WOOD AND PITH Only a few observations on macroscopic characters of Pereskia wood are re­ ported in the literature. Crespo (1973, p. 36) described the wood of Pereskia lychnidiflora as yellow, without difference between softwood and "heartwood," without characteristic odor or taste, with medium texture, soft grain, straight fiber, clearly discernible growth rings, larger pores of spring wood, radially arranged numerous pores in summer wood, and numerous rays visible to the naked eye. Wood samples of other species show basically the same characters, but growth rings are inconspicuous at least macroscopically in, e.g., P. aureiflora (sample from a branch of 6 cm diam., Leuenberger & Martinelli 3054) and in the stem of an old cultivated specimen of P. sacharosa. Gibson (1975, p. 80) noted the large unlignified pith of more than 5 cm diam. in P. lychnidiflora and described it as "stellate at the macroscopic level, particularly in the basal portion of a large trunk." Large pith is observed also in canes of P. grandifolia and P. bleo. In longitudinal sections it appears transversely laminate, a character observed also in other woody succulents, like Didiereaceae (Rauh, 1978; Rauh & Dittmar, 1970), and in Euphorbia (Leuenberger 1979, p. 82). In older wood samples of P. aureiflora, P. sacharosa, and P. portulacifolia, the pith is inconspicuous in the few samples seen.

Pollen Morphology Pollen of several species of Pereskia was studied by Tsukada (1964) and Leuen­ berger (1976a, 1976b), who showed the broad range of size and aperture number within the genus. Neumann (1935) described the development of anthers and pollen of P. nemorosa (P. amapola var. argentina), a species in which I observed indehiscent anthers or clumped pollen grains with abnormal exine in most spec­ imens. Aperture number ranges from three in the Andean species (tricolpate) to 15 in some grains of P. guamacho and a few other species (15-pantocolpate). Different aperture numbers occur within the same sample in most species, and configura­ tions aberrant from the series 3-6-9-12-15 are occasionally observed (Fig. 19C). Pollen size ranges from ca. 45 to 80 ^m and is within the range of the family as a whole. The exine is tectate in all species, with perforations of varying size and with medium- to medium-large-sized spinules in moderate number compared to other genera. Some of the species not studied in previous works are illustrated in Figure 19. Pereskia quisqueyana agrees with the other Caribbean species (Fig. 19F, G). Pereskia aureiflora approaches the southern group in exine characters but is linked, on the other hand, to the Caribbean species and P. guamacho in other features (Fig. 19H, I and Table I). The fine spinules and perforations of P. stenantha (Fig. 19J, K), although somewhat variable, are remarkable together with the other floral features mentioned above. Pollen characters pose a particular problem for phylogenetic interpretation with­ in the genus as has been noted earlier (Leuenberger, 1976a). If tricolpate pollen is considered to be more primitive than polypantocolpate, one has to accept widely differing evolutionary levels of characters within the species of the Andean group. While their habit, the tuberous roots and the small flowers and fruits were inter­ preted as advanced characters by Boke (1968) and Bailey (1963a), and Gibson 1986] PERESKIA (CACTACEAE) 41

FIG. 19. Pollen of Pereskia (all grains acetolysed). A, P. weberiana {cult. Jard. Exotique, Leuen­ berger 2718), tricolpate and one hexacolpate pollen grains. B, P. humboldtii {Hutchison 1452), exine with fine spinules and perforations. C, P. sacharosa {cult. hort. Berol. 001-15-74-80), 9-15-colpate pollen grains. D, P. lychnidiflora {Leuenberger & Schiers 2509), hexacolpate grain with coarse, irreg­ ular-sized perforations. E, P. lychnidiflora {Boke 25), tectate exine, x.s. F, G, P. quisqueyana {Leuen­ berger 3044); F, 12-colpate grain; G, exine with coarse spinules. H, I, P. aureiflora {Leuenberger & Martinelli 3054); H, 15-colpate grain; I, exine with fine spinules and numerous perforations of variable size. J, K, P. stenantha {Leuenberger et al. 3072), 15-colpate grain with fine spinules and perforations.

(1975) noted the presence of both primitive and advanced anatomical features, palynological data could be interpreted as primitive or, if one accepts the possi­ bility of a reduction series in aperture number, also as advanced. The phenomenon of differing evolutionary levels of characters was termed "heterobathmy" by Takh- tajan (1959, p. 13). It seems a useful term to characterize evolutionary features of Pereskia. 42 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

On the subfamily level, pollen characters of Pereskia are more similar to those of the Cactoideae than to those of the Opuntioideae, supporting the proposal to distinguish only two subfamilies, Cactoideae (containing the tribe Pereskieae) and Opuntioideae (Hunt & Taylor, 1986).

Phytochemistry No systematic phytochemical study of Pereskia has been made hitherto, but some scattered data are found in the compilatory publications of Gibbs (1974) and Hegnauer (1964), who consider the Cactaceae to be a phytochemically uniform and fairly well characterized family. Betalains, the pigments typical for the order Centrospermae (Caryophyllales) are found in Pereskia, namely betanin, isobetanin, and phyllocactin, according to Gibbs (1974). Betacyanin and flavonols are present in young leaves (Reznik, 1955) and citric acid in the stem of Pereskia aculeata (Hegnauer, 1964). Young leaves are often conspicuously purplish-red, purplish-green, or olive, but the quan­ tity of pigment depends much on environmental conditions. In other species only the leaf margin is finely lined with red (P. nemorosa). In P. weberiana cotyledons and seedling leaves are conspicuously reddish below while new leaves of mature plants lack this feature. Clones with and without red pigments in leaves and stem are known in P. aculeata. A character evident to the naked eye is the richness of mucilage as observed in cut stems and particularly in the fruit wall. Mucilage is apparently the active agent in the use of crushed Pereskia leaves and for clarification of water as reported by Humboldt, Bonpland, and Kunth (1823) for P. bleo.

Chromosome Number Chromosome counts for only four species of Pereskia have been mentioned in the literature, indicating 2n = 22 for P. aculeata (Arp, 1973) and P. grandifolia (Spencer, 1955) and a gametic chromosome number of n = 11 for P. diaz-ro­ meroana and P. bleo (as P. corrugata) (Ross, 1981). From 1978 to 1984 repeated attempts were made to count chromosomes of all species cultivated at Berlin. Although plenty of root tips were preserved, suitable mitotic stages were obtained only sporadically. Root tips from seedlings and cuttings in pots or in hydroculture were pretreated in oxyquinoline and preserved in ethanol/acetic acid 3:1. In several species only few metaphase plates could be studied, but in all cases a number of 2n = 22 is confirmed or reported for the first time (Table II). No significant differences in the shape of chromosomes were observed, but no detailed study of shape and size of chromosomes was made (Fig. 17 J). The chromosome number of Pereskia is thus the same as in other genera of Cactaceae, where only a few exceptions with polyploid numbers have been reported hitherto.

Evolution Pereskia is generally accepted as the genus of Cactaceae in which are united most of the primitive characters, and Schumann (1899, p. 112) was undoubtedly right in his statement that it is closest to the ancestors of the Cactaceae. In his biogeographic analysis of the family he also noted the widely scattered distribution of the species as evidence for the age of the genus. Backeberg (1942) adopted the Wegener theory of continental drift and vividly depicted the evolution of the family from ancestors to be sought in the tropical "West Indian-Middle American 1986] PERESKIA (CACTACEAE) 43

Table II Chromosome numbers of Pereskia species (counts by M. Schroder and the author).

Origin and collector Accession number Species (voucher specimens at B) at Hort. Berol. 2n = P aculeata cult. Berlin-Dahlem; coll. unknown 001-19-74-80 22 P. bahiensis cult. Berlin-Dahlem; coll. unknown 001-13-74-70 22 P bleo cult. Kew Gardens; coll. unknown 277-01-80-80 22 P. diaz-romeroana Bolivia: Samaipata; Rauh 40627 039-02-77-30 22 P. grandifolia cult. Wilhelma Stuttgart; coll. unknown 168-02-79-80 22 var. grandifolia P. grandifolia var. cult. Hamburg Bot. G. (); coll. un­ 047-01-78-80 22 grandifolia known P. grandifolia Brazil: Minas Gerais; Horst HU 226 036-01-77-30 22 var. violacea P. guamacho cult. Berlin-Dahlem; coll. unknown 001-16-74-70 22 P. humboldtii Peru: Amazonas; Hutchison 1452 256-01-82-30 22 var. humboldtii P. lychnidiflora Guatemala: Zacapa; Leuenberger & 003-04-78-10 22 Schiers 2508 P. nemorosa : San Pedro; Esser 14763 039-05-77-30 22 P nemorosa cult. Gent Bot. G. 090-01-79-80 22 P. sacharosa Bolivia; Ritter 640 038-02-77-30 22 P stenantha Brazil: Bahia; Leuenberger et al. 3072 166-72-83-10 22 P. weberiana cult. Munich Bot. G.; coll. unknown 037-01-77-80 22 P. zinniiflora ; Alain s. n. 046-02-78-30 22 region" and the subsequent development of the family in the northern and south­ ern arid zones following the dislocation of the climatic belts. The studies of Raven and Axelrod (1974) and a recent paper on biogeography and plate tectonics of the American continent by Coney (1982) provide data which leave questions as to the paleogeography of the Middle American-Caribbean region. However, they suggest to me the necessity to assume the origin of the family somewhere on the west or northwest Gondwanan continent, most likely in northwestern South Amer­ ica. This would explain the evidently early dispersal of Pereskia, Opuntioideae, and Cactoideae to the Caribbean and to North America, as well as the failure of all but one genus to reach the African continent, which was much closer to South America than North America was during the probable time of evolution of the family. Since Gentry (1982, p. 583) suggested that dry area-centered taxa of neo­ tropical origin reached Central America and southern Mexico by "late Cretaceous island hopping along the proto-Antilles," one has to assume that a species exhib­ iting many primitive characters, like P. lychnidiflora, is a relic on an outpost. Similarly, in the Southern Hemisphere, the small genus Maihuenia is a Patagonian element of pereskioid stock geographically distant from the possible center of origin. Schumann (1899) sought the nearest allies of the ancestors of Pereskia among Portulacaceae and Aizoaceae, while Buxbaum (1949) noted similarities in stem morphology with certain Phytolaccaceae (Seguieria), but his observations were influenced by the interpretation of Pereskia aculeata as the most primitive species of the genus. Although P. aculeata combines some ancestral characters, it was interpreted as rather advanced in anatomical features (Bailey, 1962; Boke, 1954, 1966) and with its specialized, scandent habit and spination it may represent a side line of evolution within the genus. 44 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Published by the State University of Utrecht, the Netherlands Department ot Systematic Botany FIG. 20. Distribution of Pereskia. Dots represent specimens of native, subspontaneous, or un­ certain status; circles indicate locally cultivated material.

As to the ancestors of Pereskia and Cactaceae as a whole, one may merely speculate about a protocactaceous plant in a tropical, seasonally dry climate as a tree with brachyblasts, leafy or spiny or both, and with flowers on leafy shoots, with an increasing number of nodes becoming incorporated in the receptacle. Nonspecific insect pollination, perhaps saurochory, and a tendency towards stem or leaf succulence seem possible in a temporarily drier habitat. A plant of the habit of P. lychnidiflora unites all these characters, combining them with a prim­ itive ovary structure (Boke, 1963b, p. 855). The tendency towards succulence of both leaves and stems is still present in some Opuntioideae, in which morpho­ logical progressions and reduction series from flat, succulent to terete, and short­ lived scaly succulent leaves, correlated with increasing stem succulence {Peres­ kiopsis, Quiabentia, Opuntia), are well known. Also, it does not seem necessary that present-day, large-leaved Pereskias of more mesic habitats (P. bleo, P. gran­ difolia) represent more-ancestral forms. On the whole they are no less succulent than the other species, and they have a number of rather derived features as well as presumably primitive ones. Since all Pereskias have areoles and spines, and many also brachyblast leaves, one may assume that areoles and spines represent a character possibly evolved before and independently of succulence and not necessarily correlated with particularly arid conditions. Pronounced heterobathmy 1986] PERESKIA (CACTACEAE) 45

is not surprising in a genus like Pereskia, and it renders the construction of a phylogenetic tree more difficult. The best way to demonstrate possible evolution­ ary lines is in connection with geographical distribution (Fig. 22). Only a few (but well delimited) species occur near the postulated center of origin, perhaps due to drastic climatic changes, whereas speciation in three major zones (Andean, Ca­ ribbean, and Caatinga-Chaco-zone) peripheral to the northwestern South Amer­ ican origin of the genus expresses itself in several groups of closely related species and subspecific taxa (Table I, Fig. 22). Distribution The genus Pereskia is distributed from Guerrero, Mexico, southward through Central America and the eastern side of the to northern Argentina, and eastward to the West Indies, , , eastern Brazil, and northern Uruguay (Fig. 20). It is doubtfully native in Florida and apparently lacking from the Pacific side of the Andes in South America, as well as in the lower part of the Amazon basin. There are no significant concentrations in any region, but areas with slightly increased species number are the Greater Antilles and eastern Brazil, where pronounced endemism occurs. The size of the distributional range varies greatly from species to species. Pereskia aculeata, a very polymorphic taxon, is widespread in two disjunct areas north and south of the equator (Fig. 24), while P. quisqueyana is as yet known only from the type locality and is very likely to be of limited range, as are two other closely allied species from Hispaniola, P. portulacifolia, and the as yet insufficiently known P. sp. A. A number of closely related taxa occur in dry valleys of the eastern Andes of Peru and Bolivia, and are possibly of very limited distribution, but they are also insufficiently known. Collections of P. bahiensis available prior to the author's field studies in Bahia gave the impression of very sporadic occurrence. However, even with the additional data available, the documented distribution may not properly reflect the real distribution pattern. Yet another drastic example is the well-known P. grandifolia, of which dozens of old collections exist from the vicinity of Rio de Janeiro and from more recent, cultivated sources. However, little is known about its possibly much larger natural range, partly because potential areas are poorly collected, and partly because cacti are underrepresented in existing collections. Only few species seem to have over­ lapping ranges of distribution as, e.g., P. sacharosa and P. diaz-romeroana in Bolivia, and P. aculeata and several other species throughout its range. P. gran­ difolia and P. bahiensis were observed side by side in southern Bahia, where both species are used for planting hedges, but they are probably not naturally sympatric. Two species not distinguished by evident vegetative features but strikingly dif­ ferent in flower characters, P. bahiensis and P. stenantha, may have overlapping ranges of distribution, as is suggested by observations of G. C. Pinto (pers. comm.).

Ecology

HABITAT AND CLIMATE Most species of Pereskia occur in dry forest and thorn-scrub habitats in a tropical climate with a more or less pronounced dry season of about two to five months. They are associated with other woody plants in vegetation types known as "selva baja espinosa" in Mexico, "caatinga" in Brazil, and "chaco" in Paraguay, or described as cactus forest in Bolivia (Cardenas, 1969) or thorn scrub in the West 46 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Indies (Howard, 1973). Some species, e.g., P. aureiflora and P. grandifolia, grow in less xeromorphic vegetation types, although perhaps on locally drier sites. The polymorphic P. aculeata seems to have also a very broad ecological range, as it is reported from rocky areas in mesophytic forest, from dry forest, restinga and from dunes near the sea. Pereskia bleo is a notable exception, if the information available is correct. This species is reported growing on sandy soil along streams in lowland evergreen forests in Panama and Colombia. Most Pereskia habitats are on alluvial soils on diverse geological formations, including igneous and calcareous rock. Pereskia aculeata is found on both, P. bahiensis and P. aureiflora grow in areas where granite, gneiss, and quartzites are predominant, while P. stenantha was observed by the author near calcareous formations in the Rio Sao Francisco valley in Bahia, but it is not known whether its range is correlated with this geological factor. The Caribbean species mostly grow in soils derived from coral limestone, and P. lychnidiflora occurs on alluvial soil of diverse origin. In cultivation, all species seem to do well in any substrate.

PHENOLOGY Pereskia species are deciduous under conditions with seasonal drought and, in cultivation, also at mean temperatures below ca. 15 C. Species without brachyblast leaves, like P. bleo, may be only partly deciduous, as is the case in cultivation under favorable conditions, but I have no data on the phenology of this plant in its native habitat. The Andean species with tuberous roots shed their leaves completely and, even though they do not have brachyblast leaves, seem very well adapted to xeric conditions. Species with brachyblast leaves may produce these independently of new growth of long shoots. If water is limited, they may set new brachyblast leaves but not produce new shoots. Thus they can nevertheless quickly restore full photosynthetic capacity. Shedding of only part of the leaves during shorter periods of drought is observed in cultivation. According to data compiled from herbarium labels most species flower on leafy stems during the rainy season. The flowers of P. aculeata are produced on long shoots at the end of the growing season. Pereskia guamacho is a notable exception, as it flowers in the leafless stage at the end of the dry period. The flowers of Pereskia open during the day and usually last one day or only a few hours. In cultivation, P. grandifolia is an exception, at least on cool days, with a prolonged anthesis of two days during which the petaloid perianth segments are somewhat accrescent. Anthers dehisce early during anthesis and usually well before the stigma lobes begin to spread. Weingart (1904, p. 85) noted sensitive stamens in P. nemorosa, in which he saw filaments curling around the style upon touch, an observation which I cannot confirm.

POLLINATION Rowley (1980) summarized the pollination syndromes in cactus flowers with reference to both earlier (Porsch, 1938-1939) and recent (Grant & Grant, 1979) works. Floral syndromes in Pereskia, with two exceptions, include diurnal opening time, various colors, scent faint or none, pollen abundant and freely exposed, nectar freely exposed or absent, size various, profile flat. These characters suggest unselective pollination, but unfortunately direct observations are scarce. Porsch (1938, pp. 36, 42) mentioned P. aculeata and P. grandifolia as unselective and P. lychnidiflora (as P. nicoyana) as having bee flowers. The latter observation is confirmed by the author, based on sightings in the field in Guatemala in 1978, 1986] PERESKIA (CACTACEAE) 47

and also documented by a photograph. A single undocumented observation (by the author) of a hummingbird visiting a staminate flower of P. portulacifolia suggests at least potential pollination by birds, although the species does not have flower characters particularly adapted for bird pollination. In P. zinniiflora, a. species with identical floral syndromes, Areces (1984, as Rhodocactus cubensis) observed Hymenoptera and Lepidoptera visiting the flowers. Pereskia aculeata likewise has unselective flower characters, but the flowers are fragrant, the scent being variously described as pleasing or offensive by several observers. The species is mentioned as a bee plant on a herbarium label (Mexia 4129), and I have recently seen bees, bumble-bees, and syrphids visiting flowers in a greenhouse-grown spec­ imen. The notable exception in floral characters is P. stenantha, the only species with a flower suggesting bird pollination. Unlike the case in other species of the genus, the perianth segments remain erect, causing the urceolate to campanulate shape of the flower. The rather fleshy sepaloids are orange-red to red, rarely with a purplish tinge, and thus contrast with the pink to purplish-pink tip of the petaloids, which are surrounded and enclosed by the sepaloids for more than half of their length. Stamens and style are tightly enclosed, and a conspicuous nectary and diffuse nectar chamber are present (Figs. 14A, 49E). The difference in floral characters between this species and the vegetatively indistinguishable P. bahiensis can only be understood as a case of evolution caused by pollination vectors. Ritter (1979, p. 22) commented on this without mentioning possible ornithogamy. Un­ fortunately it was not possible to observe any pollinators during a short stay in the field in 1983. Butterflies visiting nearby Lantana camara flowers were not attracted to P. stenantha. All Pereskia species are basically self-sterile although Ross (1981) reported that self pollination produces seed in P. diaz-romeroana, a species observed by the author to occasionally set fruit without cross pollination. In other species self pollination rarely yields fruit. However, in these cases the fruits are then often underdeveloped, seedless or few-seeded. Incompletely unisexual flowers in the genus were noted 200 years ago by La­ marck (1785) for Cactus portulacifolius, based on a manuscript of Plumier. This feature has been overlooked by subsequent monographers and collectors until recently. I have seen many staminate flowers with obviously non-functional gy- noecia, but only one collection of pistillate flowers, of P. portulacifolia (Zanoni 35204). The plants are apparently dioecious (Zanoni, pers. comm.). The same was reported for P. zinniiflora under its synonym Rhodocactus cubensis by Areces (1984), who described and illustrated the unisexual flowers. His observations are based on a large number of specimens and confirms the phenomenon which, however, is not distinctive of the species, but apparently a characteristic of the entire group of endemic species of the Greater Antilles, including also P. quis­ queyana and the insufficiently known P. sp. A. Unisexual flowers are an exception in the family, but known in, e.g., Mammillaria dioica and Selenicereus innesii (Kimnach, 1982).

DISPERSAL Little is known about dispersal agents of Pereskia, but, from the fruit type, endozoochory or synzoochory by generalist frugivores (Roosmalen, 1985) seem most probable. The only species listed by that author, P. aculeata, is described as endozoochorous (bats and monkeys), but the indication of the seed dispersal strategy did not necessarily mean that it has been directly observed (Roosmalen, 1985). Other species with -like fruits, juicy fruit walls (P. weberiana), or 48 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 sticky pulp (P. portulacifolia) seem likely to be dispersed by birds. Only speculation is possible for species with greenish-yellow fruits, tough fleshy fruit walls, and indetectable odors, as in P. grandifolia and others. Birds, bats, monkeys, or even should be considered as potential dispersal agents. The fruit of P. lych­ nidiflora is of particular interest because its seeds are surrounded by stinging sclereid fibers of the locule wall. Although this may not be a factor preventing dispersal of the intact fruit, the fibers might act as a repellent against consumption of the seeds. Nevertheless, the fruits collected by the author during a field trip to Guatemala in 1978 proved to be heavily infested with the larvae of a beetle, obviously consuming the seeds. Different modes of fruit maturation are observed within the genus and should perhaps be considered in a discussion on dispersal. In species with small, black, and soft berries the fruits mature within a few weeks and change color from green to blackish within a few days, whereas the large greenish fruits of P. grandifolia under the same conditions in cultivation take several months to mature without changing color significantly. A large fruit of this species harvested in March 1984, and kept at room temperature in the laboratory for ten months, gradually lost weight without external shrinking, and the outer surface remained smooth and exuded an oily substance for many months while slowly turning yellowish-green (Fig. 14F).

Conservation Although most species of Pereskia do not seem immediately endangered, many of them may eventually be threatened in large areas of their habitat due to clearing of land for dry-farming or replacement of hedges by barbed wire. In grazing areas, plants would probably survive and some would be kept as ornamentals. They could well be preserved in cultivation at least in tropical gardens as they are easily propagated by seed or cuttings. Pereskia quisqueyana probably represents the only immediately threatened taxon unless further localities are found in addition to the only known habitat in a thicket near a village and beach at Bayahibe, east of La Romana in the Dominican Republic.

Uses The genus is of little economic importance although many local uses, listed under the species, are known. The most widespread use is for hedges, by means of cuttings taken during the dry season. These root easily and a living fence of spiny canes about 1 m tall soon develops into an impenetrable hedge. Species reported to be used this way are P. bahiensis, P. stenantha, and P. grandifolia in Brazil, and P. guamacho in Venezuela and P. lychnidiflora in El Salvador. In Bahia, Brazil, such hedges were frequently observed even along main roads during a trip by the author in 1983 (Fig. 21A-E). The crushed leaves of P. bleo were reported by Humboldt, Bonpland, and Kunth (1823) to be used to clarify water, the abundant mucilage apparently being the active agent of this process. The leaves of P. aculeata are still widely used as a pot herb in rural areas of Minas Gerais and Bahia and even sold on the markets. The fruits of P. aculeata and P. guamacho are reported to be edible (Romero- Castafieda, 1971; Standley, 1924). Eichlam (1909) indicated that the spines of P. lychnidiflora (as P. autumnalis) were used as needles in Guatemala. Some species have beautiful flowers and are often found in cultivation in botanical collections and to a lesser extent in private gardens and public parks in tropical and subtropical 1986] PERESKIA (CACTACEAE) 49

FIG. 21. Fences and hedges of Pereskia bahiensis. A, a new fence planted near Caetite, Bahia. B, a fence developing into a hedge at Brejinho das Ametistas near Caetite. Bahia {Leuenberger et al. 3071). C-E, fences and hedges near Brumado, Bahia {Leuenberger et al. 3063). regions. Pereskia grandifolia is particularly widespread because it flowers profusely and even small cuttings taken from flowering shoots will continue to produce flowers every year as small pot plants. Pereskia aculeata is a decorative vine, but the intensely aromatic and somewhat offensive fragrance of the flowers is said to limit its use. The wood of Pereskia is apparently not used except for fence posts. Charcoal burners exploiting dry forests in the arid parts of the southwestern Dominican Republic apparently leave P. portulacifolia untouched, but this could at least partly be due to the spininess of the plants. No part of the plants is known to be poisonous, but Standley and Williams (1962) consider P. lychnidiflora plant­ ed as a hedge to be dangerous to man and . The stinging sclerenchymatic fibers falling from the withering cortex of dead branches and trunks and blown 50 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Table III Hybridization of Pereskia in cultivation.

Fruit develop- Seed Cross ment viability F, generation P. diaz-romeroana x P. weberiana yes yes vigorous, intermediate, flowers (pink flowers) like either of the parents, or white P. weberiana x P. diaz-romeroana yes yes vigorous, intermediate, flowers (pink flowers) like P. diaz-romeroana, or white P. grandifolia var. grandifolia x yes yes vigorous, intermediate, no flowers var. violacea seen yet P. grandifolia var. violacea x yes yes vigorous, intermediate, flowers var. grandifolia more like var. grandifolia P. bahiensis x P. stenantha yes no b P. sacharosa x P. grandifolia yes no — P. grandifolia x P. sacharosa no a — P. bleo x p. grandifolia no — — P. bleo x p. diaz-romeroana no — — P. nemorosa x P. diaz-romeroana no — — a None. bNoF,. away with the wind irritate the skin and are dangerous to the eyes. These fibers apparently make the wood unsuitable for fuel.

Hybridization No natural hybrids of Pereskia have been reported. Under experimental con­ ditions, however, hybridization is possible between some closely related taxa. Others seem to be incompatible insofar as evidence from the few cross-pollination experiments between flowering specimens available at the Berlin-Dahlem Botan­ ical Garden. The results are listed in Table III. The first flowers on hybrid specimens of P. weberiana x P. diaz-romeroana appeared on a one and one-half-year-old seedling with a main shoot only 30 cm long. This is the youngest and smallest flowering seedling plant of Pereskia ob­ served by the author. All hybrids between the two closely related and vegetatively similar species are more or less intermediate but closer to one or the other parent in individual characters. It is noteworthy that about half of the F, hybrids between pink-flowered plants of the two species produce white flowers. Interpretation of this result is difficult due to the small number of hybrid specimens available. White-flowered specimens are known in P. weberiana but have not been reported so far in P. diaz-romeroana, which may nevertheless possess recessive alleles for white flowers, a character known also in the third Andean species, P. humboldtii. Possibly a more complex genetic control of flower characters has to be assumed. Of four hybrids between P. diaz-romeroana and P. weberiana two produce white flowers, one has pink flowers resembling P. weberiana and one has the pigmented stamens typical for P. diaz-romeroana. In all four hybrid plants the receptacle and fruit resembles P. weberiana. The hybrids between the two varieties of P. grandifolia again yielded plants with mostly intermediate characters. The color of the bracts—a particularly striking feature of P. grandifolia var. violacea—is 1986] PERESKIA (CACTACEAE) 5 1

intermediate in the only hybrid specimen yet flowered, but the petaloids and stamens are exactly as in the pollen-supplying parent in this hybrid of P. gran­ difolia var. violacea x var. grandifolia.

Infrageneric Relationship The first author to circumscribe the genus as we do today was Schumann (1898), who excluded species of Opuntia, Pereskiopsis, and Maihuenia that had been included in Pereskia by earlier authors (Riimpler, 1886; Salm-Dyck, 1850). His infrageneric concept is therefore the first one to be considered. His two subgenera were based on the presence of spines on the receptacle (subgenus Eupeireskia K. Schumann) and their absence (subgenus Ahoplocarpus K. Schumann). The first consisted of one species only, P. aculeata. Britton and Rose (1919) equally main­ tained two entities, series Typicae and series Grandifoliae. Berger (1926, 1929) again recognized two subgenera, however, based on different diagnostic characters, namely the internal structure of the ovary. His interpretation of the position of the ovules in P. aculeata and P. sacharosa as being "at the base of the style" and in subgenus Rhodocactus as "enclosed in a true locule" seems strange, as was already noted by Boke (1964). They are, rather, enclosed in a locule of varying relative position within the receptacle in all species. Backeberg and Knuth (1935) elevated Berger's subgenera to the rank of genera, classifying the species without knowing the internal structure of the flower in many cases. Taxa now known to be conspecific were even attributed to different genera (P. conzattii, P. nicoyana). A more recent classification by Backeberg (1958) is based on the same principle, although his new subgenus Neopeireskia (Backeberg, 1956) is an acceptable entity comprising the Andean species with small leaves and densely clustered flowers; but I doubt that it should deserve more than the rank of a series. The groups outlined by Gibson (1975), based on anatomical data, clearly show the limits between Pereskia and genera of the Opuntioideae supported by ample character combinations, but his three infrageneric groups are based solely on the nature of the phloem sclereids. The three groups, impractical to distinguish in a key, appear to have some evolutionary significance, but are not entirely satisfactory as taxonomic units. The question is whether one single character should be ac­ cepted to delimit infrageneric entities. Equally, the position of the ovary and placenta is, in my opinion, not a tangible character to delimit two subgenera. The transitional nature of ovary and ovule position, subject to distortions during ontogeny of flower and fruit, makes it an unreliable character. Where to draw the line between two or three tentative units continues to be controversial. Two possible entities based on the position of the placenta in fruit do not coincide with the above mentioned subgenera and would also mean the over-emphasis of one character, which is not easily available as a key character in material to be identified. A closer look at character correlations suitable for infrageneric classification reveals that it is possible to outline only more or less distinctive species groups, but not clear-cut entities. Intricate and reticulate relations occur between these species groups and individual species, as expressed in Table I. Little is to be gained by naming seven possible groups of which three would be monotypic. When it is necessary to speak of some groups of closely related species, geo­ graphic and descriptive terms seem adequate in this relatively small genus. A diagram indicating evolutionary affinities, as expressed by the data shown in Table I and suggested by overall similarities, is given in Figure 22. Conspicuous specific characters like hairiness of the upper areoles of the re- 52 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 198^] PERESKIA (CACTACEAE) 5 3

ceptacle, or flower color, are apparently not correlated with the evolutionary groups and only would make a very artificial infrageneric classification.

Intergeneric Relationship Pereskia belongs to the subfamily Pereskioideae K. Schumann (Schumann, 1898). The only other genus of this subfamily is Maihuenia, a genus of low, cushion- or mat-forming succulent spiny shrubs. The stems of these lack stomata and have early periderm formation, and their leaves are long-lived as in Pereskia but much smaller, terete, and succulent and resemble those of Opuntioideae, although the arrangement of stomata is different (Eggli, 1984). Maihuenia resem­ bles Opuntia in habit but lacks glochids. Seed structure, pollen, and anatomical features also clearly negate inclusion in Opuntioideae (Gibson, 1977; Leuenberger, 1976a, 1976b). Pereskia is therefore without very close or even hard-to-distinguish relatives. Nevertheless there has been some confusion between species of Pereskia and Pereskiopsis, a genus of the subfamily Opuntioideae. Superficial similarities of Pereskiopsis, and the probably congeneric Quiabentia, with Pereskia are readily disproved by their barbed spines, opuntioid flowers, porate pollen, and seeds provided with a bony "aril," as well as by anatomical features discussed by Bailey (1960, 1961b, 1964) and Gibson (1975). A recent proposal of an ad hoc working party (including the author) under the auspices of the International Organization for Study is to divide the family into only two subfamilies (Hunt & Taylor, 1986). This reduces the Pereskioideae to a tribe (Pereskieae) within subfamily Cactoideae, based on char­ acters shared with tribe Cacteae, such as lack of glochids, seeds without bony aril, and anatomical and palynological similarities.

Systematic Treatment

GENERIC DESCRIPTION Pereskia Miller, Gard. diet. abr. ed. 4. 1754. Haworth, Syn. pi. succ. 198, 199. 1819; De Candolle, Prodr. 3: 474, 475. 1828; De Candolle, Mem. Mus. Hist. Nat. 17: 73-77, figs. 17-20. 1828; Pfeiffer, Enum. diagn. Cact. 175-179. 1837; Salm-Dyck, Cact. hort. Dyck. 75, 76. 1850; Hooker in Bentham & Hooker, Gen. pi. 1(3): 851. 1867; Schumann in Martius, Fl. brasil. 4, 2: 308-314, fig. 43. 1890; Schumann in Engler & Prantl, Nat. Pflanzenfam. 3 (6a): 203-205, fig. 71. 1894; Schumann, Gesamtb. Kakt. 757-766. 1898; Weber in Bois, Diet. hort. 937-939, fig. 678. 1898; Britton & Rose, Cactaceae 1: 8-24, pi. 2, 3, figs. 1-20. 1919; Vaupel in Engler & Prantl, Nat. Pflanzenfam. ed. 2, 21: 615-617, fig. 274. 1925; Vaupel, Die Kakteen 1: 3-18, figs. 1-4. 1925; Britton & Wilson, Sci. Surv. Porto Rico & Virgin Isl. 5: 608. 1924; Standley,

FIG. 22. Infrageneric groups, character correlations, and geographic distribution in Pereskia. Map shows approximate present center of distribution marked by a number (see numerical list of taxa and graph at right) for each species as well as their hypothetical links as geographical-evolutionary groups to an assumed center of origin in NW South America. The diagram at right, with explanation of pictograms at lower left, shows the seven indistinctly delimited groups and the distribution of seven selected characters demonstrating heterobathmy. 54 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Trees and shrubs of Mexico, Contr. U.S. Natl. Herb. 23: 858-863. 1924; Descole, Gen. et sp. pi. Argent. 1: 121, 122, fig. 46. 1943; Leon & Alain, Fl. Cuba 3: 358, fig. 159. 1953; Rauh, Beitr. peruan. Kak- teenveg., Sitzungsber. Heidelberger Akad. Wiss. math.-nat. Kl. 1958(1): 188-192, figs. 79, 80. 1958; Woodson & Schery, Bora of Panama VII, Ann. Missouri Bot. Gard. 45: 69-72, fig. 21. 1958; Backeberg, Die Cactaceae 1: 104-113. pi. 1, 2, figs. 41-46. 1958; Standley & Williams, Flora of Guatemala VII, Fieldiana Bot. 24: 222-226, fig. 35. 1962; Backeberg, Kakteenlexikon, 354-357. 1966; Hunt in Hutchinson, Gen. Fl. PL 433. 1967; Kiesling, Los generos de Cactaceae de Argentina, Bol. Soc. Argent. Bot. 16: 202, 204, figs. 1-3. 1975; Bravo, Las Cac­ taceas de Mexico 1: 143-154, figs. 64-72. 1978; Ritter, Kakteen in Siidamerika 1: 19-22, figs. 1-5, 217. 1979; Benson, The Cacti of the and Canada, 264-268, 911, 969, figs. 234-237. 1982. Type species: Pereskia aculeata Miller (=Cactus pereskia Linnaeus, cf. Britton & Rose, 1919). Named as a tribute to the French scholar Nicolas Claude Fabri de Peiresc (1580-1637). Rhodocactus (A. Berger) F. Knuth in Backeberg & Knuth, Kaktus-ABC 48, 96. 1935. Type. Pereskia grandifolia Haworth (cf. Backeberg, 1958, 113). Leon & Alain, Fl. Cuba 3: 359, 360. 1953; Backeberg, Cactaceae 1: 113-118, figs. 47-53. 1958; Backeberg, Kakteenlexikon, 395-397, figs. 370-372. 1966. Leafy and spiny trees, shrubs, or scramblers, sometimes with fleshy, tuberous root thickenings; twigs straight or slightly zigzag, with green cortex; epidermis with few to numerous stomata and retarded periderm formation, or stomata lacking, cuticle thin and periderm formation precocious; epidermis glabrous, or rarely with minute papillae; bark smooth or longitudinally fissured. Areoles in the axils of deciduous leaves, sparsely to densely tomentose with uniseriate and some­ times multiseriate trichomes, producing spines and sometimes brachyblast leaves. Leaves alternate, broad, flat, exstipulate, petiolate to subsessile, somewhat fleshy, deciduous; margin entire, sometimes undulate; venation pinnate or pseudopal­ mate. Spines solitary, paired, or numerous, uniform but unequal in length, rarely dimorphic and then the first spines paired, claw-like and recurved, straight or nearly so, acicular to subulate, sometimes with markedly broadened, conical base, sometimes lacking on flowering twigs but increasing in number on the trunk. Flowers solitary, terminal or axillary, or in a racemose, paniculate or cymose- paniculate inflorescence sometimes by proliferation from the flower receptacle; flowers hypogynous-perigynous to epigynous; receptacle with few to many areoles, smooth or with prominent podaria, rarely naked except for the receptacular rim; receptacular areoles with sparse to dense tomentum, rarely spiny or with coarse longer hairs; receptacular bracts leaf-like to scaly, somewhat fleshy, the lower ones spreading, the upper ones erect and appressed in bud, green or sometimes tinged like the sepaloids; perianth multiseriate, the segments few to numerous, free; sepaloids somewhat intermediate between bracts and petaloids, lacking axillary areoles and partly tinged like the petaloids; petaloids spreading, rotate, rarely erect and forming a campanulate-urceolate corolla; staminodial hairs rarely present between petaloids and stamens; stamens numerous; filaments inserted at the base of the perianth and shorter than it; ovary superior to inferior, unilocular, with septal ridges at the ovary roof only, or rarely septate with pocket-like locules around a central conical columella; placentae "basal-axile," "basal," or parietal; style shorter to longer than the stamens; stigma lobes 3-20, erect to spreading! Fruit solitary, or clustered-proliferous, or in axillary clusters, rarely in a racemose or paniculate infructescence; fruit pear-shaped, turbinate, globular or broadly 1 986] PERESKIA (CACTACEAE) 5 5 depressed-obovate, terete or angled, with narrow to broad umbilicus and persistent or deciduous flower remnants and persistent or deciduous leafy or scaly bracts; fruit wall formed mostly by the receptacle, fleshy to juicy, often rich in mucilage, rarely with fusiform sclereids; locule hollow or filled by a pulp formed by the accrescent funicles. Seeds few to numerous, obovate to lenticular reniform, 1.8- 7.5 mm long, with black, mostly smooth and glossy testae, the micropylar end rounded to nose-like; hilum flat to prominent, whitish, small but conspicuous. Embryo strongly curved around the starchy perisperm; cotyledons foliaceous, flat or obvolute. A neotropical genus of 16 species, ranging from southern Mexico and the West Indies to northern Argentina and Uruguay, primarily in the lowlands.

NOTES ON KEYS, TERMS, AND ABBREVIATIONS Three separate keys for flowering, fruiting, and sterile specimens are given for practical reasons. No safe and easily visible characters are available to key out in a single key the natural groups worked out in Table I and expressed in the sequence of species in the systematic treatment. Therefore, all keys are artificial. The quality of herbarium specimens depends on the drying technique, and shrinkage and color change are often considerable. Moreover, many specimens are incomplete and sometimes lack important parts, besides telling little about the variability of leaf shape and size and of spination. The number of characters both reliable and present in an average specimen is accordingly small, particularly in dried material. Measurements were basically taken from dried material unless specifically stated and with some degree of tolerance apply to dry, liquid-preserved, and fresh ma­ terial. However, measurements of internal flower parts (style, locule, ovules) and of thickness of leaves are taken from liquid-preserved and/or fresh specimens. Dried material with obvious shrinkage, e.g., withered petaloids, should be ex­ amined carefully. Leaf and bract measurements are given as length x width including petioles. Measurements of the receptacle are taken at its greatest di­ ameter. The size of leaves and thickness of stems vary greatly with environmental conditions. Very thin and crinkled brachyblast leaves are usually immature and should be disregarded for measurements. Colors refer to living material unless otherwise stated. Stem and leaves sometimes turn black upon drying, particularly in specimens where boiling or freezing has been used to facilitate drying. The pink to purplish color of the petaloids is difficult to describe properly by standard terms. Deepness of color seems to depend on environmental conditions, and colors sometimes fade during anthesis. Dried specimens often show darker tones than fresh material, if flower color is retained at all. Morphological terms used in the key and descriptions are discussed in the relevant chapters and illustrated in Figures 4-6 (stem and leaves), and Figure 9 (flower). Photographs are by the author unless otherwise stated. Herbarium acronyms follow Holmgren et al. (1981).

KEYS TO SPECIES OF PERESKIA Key A for Flowering Specimens* 1. Scandent shrub with racemose to paniculate inflorescences and whitish, fragrant flowers; spines on new growth geminate, recurved. 1. P. aculeata.

4 For fruiting and sterile specimens, see Keys B and C. 56 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

1. Shrub or tree, inflorescence and flowers various, not fragrant; spines straight or nearly so. 2. Axils of the upper receptacular bracts with short tomentum or conspicuous hairs less than 5 mm long, or naked. 3. Receptacle with more than 10 bracts or areoles; petaloids yellow to orange-red. 4. Receptacle more than 10 mm in diam.; flowers orange-red; petaloids laciniate- dentate to erose. . 5- P lychnidiflora. 4. Receptacle less than 10 mm in diam.; flowers yellow; petaloids entire. 5. Rowers distinctly pedicellate. 6- p- aureiflora. 5. Rowers sessile, solitary or clustered on the areoles and knob-like spur shoots. 7. P. guamacho. 3. Receptacle with less than 10 bracts or areoles; petaloids not yellow, if orange-red not laciniate-dentate. 6. Rowers with campanulate to urceolate perianth; perianth segments erect, recurv­ ing at apex only; stamens and style enclosed. 16. P. stenantha. 6. Rowers rotate; perianth segments spreading, not erect and enclosing stamens and style at anthesis. 7. Receptacle either less than 5 mm in diam., or larger and then lacking prominent podaria; flowers solitary, racemose, or in condensed axillary cymes (then ap­ pearing fasciculate). 8. Axils of upper receptacular bracts with silky hairs; flower buds often partly masked by the hairs. 9. Rowers reddish, orange, or white; tomentum of the areoles uniformly white to grayish-white. 2. P. humboldtii. 9. Rowers purplish-pink; tomentum of the areoles grayish-brown, some­ times masked by longer white hairs. 3. P. diaz-romeroana. 8. Axils of the upper receptacular bracts without longer hairs; flower buds appearing naked. 10. Receptacle 3-4 mm in diam.; flowers usually less than 25 mm in diam., pink or white. 4. P. weberiana. 10. Receptacle 5-16 mm in diam.; flowers usually more than 25 mm in diam., orange-red, red, or pink to purplish. 11. Rowers bright red, salmon, or orange-red; receptacle angled; an­ thers cream; leaves mostly larger than 9 x 2.5 cm; flowers perfect. 11. P. bleo. 11. Rowers pink to purplish; receptacle terete; anthers grayish-pink or yellow; leaves mostly smaller than 9 x 2.5 cm; flowers often imperfect, staminate or pistillate. 12. Tomentum of areoles rusty brown; leaves acute. 8. P. zinniiflora. 12. Tomentum of areoles grayish to brown; leaves various. 13. Brachyblast leaves rounded to emarginate; anthers 0.8- 1.2 mm long. 9. P. portulacifolia. 13. All leaves acuminate or acute. 14. Receptacle (staminate flowers) less than 10 mm in diam., turbinate-poculiform; anthers 1.6-2.5 mm long. 10. P. quisqueyana. 14. Receptacle (pistillate flowers) ca. 16 mm in diam., subspheroidal. p. sp. A. 7. Receptacle over 5 mm in diam.; receptacular bracts and areoles on prominent podaria; flowers usually proliferating, in a terminal cymose-paniculate (cor­ ymbose) inflorescence, or occasionally solitary. 15. Leaves normally more than twice as long as broad, elliptic-lanceolate; apex acute to acuminate. 14. p. grandifolia. 15. Leaves normally less than twice as long as broad, obovate to elliptic obovate; apex acute to rounded. 15. p. bahiensis. 2. Axils of the upper receptacular bracts with conspicuous hairs more than 8 mm long. 16. Rowers with staminodial hairs between the petaloids and stamens; leaves narrowly elliptic-obovate; lateral nerves diverging at angles of 40-60°. 12. P. nemorosa. 16. Rowers without staminodial hairs between the petaloids and stamens; leaves ob­ ovate; lateral nerves diverging at angles of 25-40°. ' 13. p. sacharosa. 1 9 8 6] PERESKIA (CACTACEAE) 5 7

Key B for Fruiting Specimens

Note: Fruits are yet unknown in species no. 10 and insufficiently known in "sp. A." Both are placed in the key based on fruit characters predictable from flower characters. 1. Fruit turbinate and with large flat naked umbilicus nearly as large as the truncate end of the fruit. ii. p. bleo. 1. Fruit shape various; umbilicus distinctly smaller, naked or with bracts, hairs, or flower remnants. 2. Umbilicus narrow, with conspicuous long hairs arising from the areoles of its constricted rim. 3. Fruit normally more than 15 mm in diam., greenish to yellow or reddish. 4. Average leaves more than twice as long as broad or narrower; seeds 4.5-5 mm long. 12. P. nemorosa. 4. Average leaves ca. twice as long as broad; seeds 3.2-4.2 mm long. 13. P. sacharosa. 3. Fruit less than 15 mm in diam., vinaceous to black. 5. Tomentum of the areoles white or nearly so. 2. P. humboldtii. 5. Tomentum of the areoles brown (sometimes masked by longer white hairs). 3. P. diaz-romeroana. 2. Umbilicus wide or narrow, without long hairs, rim naked or with short-tomentose areoles. 6. Mature fruit chocolate-brown to black, 5-15(-20) mm in diam. 7. Fruit chocolate-brown; seeds ca. 5 mm long. 6. P. aureiflora. 7. Fruit black; seeds ca. 2 mm long. 4. P. weberiana. 6. Mature fruit greenish, or yellow to reddish, normally more than 15 mm in diam. and often much larger. 8. Fruit not angled, subglobular to ficiform, mostly less than 5 cm long, never pro­ liferating. 9. Seeds few, more than 4 mm long; young fruit spiny. 1. P. aculeata. 9. Seeds numerous, less than 4 mm long; fruits not spiny. 10. Fruit with numerous bracts or bract scars and inconspicuous areoles; umbilicus narrow, concealed by the connivent, scaly upper bracts. 11. Fruit 3-5 cm in diam.; locule with a distinct columella; sclereids present in the fruit wall. 5. P lychnidiflora. 11. Fruit 1.5-3 cm in diam.; locule without columella; sclereids absent. 7. P. guamacho. 10. Fruit with 0-4 bracts or bract scars and minute areoles; umbilicus con­ spicuous, naked (fruit unknown in no. 10). 12. All leaves acute to acuminate. 13. Areoles with rusty brown tomentum. 8. P. zinniiflora. 13. Areoles with pale brown to gray tomentum. 14. Spines stramineous to brown; fruit unknown. 10. P. quisqueyana. 14. Spines dark reddish-brown to black; fruit depressed-globose. P. sp. A. 12. Brachyblast leaves obtuse to emarginate. 9. P. portulacifolia. 8. Fruit angled-pyriform or angled-turbinate, ca. 4-10 cm long, often proliferating. 15. Leaves narrowly elliptic to obovate-lanceolate; lateral veins (7-) 10-13; seeds (5_)6-7 mm long. 14. P. grandifolia. 15. Leaves obovate to broadly elliptic obovate; lateral veins 5-7; seeds 4-5.5 mm long. 16. Infructescence densely clustered; fruits sessile to subsessile. 16. P. stenantha. 16. Infructescence lax; fruits (except the outermost) distinctly pedunculate. 15. P. bahiensis. Key C for Sterile Specimens

Note: Well-developed distal twigs with leaves are necessary, in some cases both new growth and stem parts formed in the previous season (brachyblast leaves). It must also be kept in mind that plasticity of stem thickness, leaf shape, and leaf size may cause difficulties in individual specimens. 58 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

1. First spines recurved, claw-like, usually geminate, with very broad base; clambering shrub or liana. l- R aculeata. 1. All spines straight or nearly so, acicular to subulate; shrubs or trees. 2. Leaves distinctly petiolate; petiole equalling to longer than the spines; blade acuminate at apex; lateral veins 4-7, bifurcate. \\. P. bleo. 2. Leaves indistinctly to short-petiolate; petiole shorter than the spines; leaf apex and venation various. 3. Venation pseudopalmate. 4. Distal twigs less than 4 mm thick; secondary phloem and cortex with aggregated stone cells. 1. P. guamacho. 4. Distal twigs more than 4 mm thick; secondary phloem and cortex with dispersed fusiform sclereids. 5- p lychnidiflora. 3. Venation pinnate, or lateral veins inconspicuous. 5. Leaves of distal twigs obovate-spathulate to obcordate, 5-15(-30) mm long, with obtuse to emarginate apex. 9. P. portulacifolia. 5. Leaf shape and size various; leaves more than 20 mm long, with rounded, acute, or acuminate apex. 6. Distal twigs 4 mm thick or less; lateral veins mostly 0-5; roots often tuberous. 7. Areoles rusty brown. 8. P. zinniiflora. 7. Areoles whitish, gray, or pale brown. 8. Areoles, particularly in new growth, with coarse longer hairs or bristles besides the short tomentum. 9. Tomentum of the areoles white to pale gray; color of trichomes uniform. 2. P. humboldtii. 9. Tomentum of the areoles grayish-brown, sometimes masked by longer white hairs; trichomes with dark base. 3. P. diaz-romeroana. 8. Areoles without conspicuous longer hairs or bristles besides the tomen­ tum of the areoles. 10. Spines subulate, with very broad and angled or flattened base; sto­ mata on the stems numerous. 4. P. weberiana. 10. Spines acicular to subulate, slightly thickened at the base, stomata on the stems few or absent. 11. Lenticels few; Caribbean species. 12. Spines stramineous to brown. 10. P quisqueyana. 12. Spines reddish-brown to nearly black. P. sp. A. 11. Lenticels numerous; South American species. 13. Lateral veins diverging at angles of ca. 20-40°. 7. P. guamacho. 13. Lateral veins diverging at angles of ca. 40-60° 6. P. aureiflora. 6. Distal twigs more than 4 mm thick; lateral veins mostly more than 4; roots not tuberous. 14. Leaves mostly narrowly elliptic-obovate to elliptic-lanceolate, more than twice as long as broad; lateral nerves 6-13, diverging at angles of 40-60°. 15. Stomata present, the stomatal cavities visible on living stems as light dots; epidermis often reddish. 14. P. grandifolia. 15. Stomata absent on stem; epidermis smooth, green, never reddish. 12. P. nemorosa. 14. Leaves mostly obovate or elliptic-obovate and to twice as long as broad, but brachyblast leaves and leaves of strong shoots sometimes narrower; lateral nerves mostly 4-7, diverging at angles of 25-40°. 16. Tomentum of the areoles pale gray, the trichomes uniform in color; stomata lacking on stem, or very few around the areoles; species of the Chaco and eastern Andes. 13. P. sacharosa. 16. Tomentum of the areoles brown at least in youth, the trichomes with darker transverse cell walls; stomata usually present on stem, but density variable; species of the caatinga of eastern Brazil (species not distinguishable by evident vegetative characters). 15. P. bahiensis. 16. P. stenantha.

1. Pereskia aculeata Miller, Gard. diet. ed. 8. 1768. Figs. 4E, 51, 6D-F, 7E, 8, 11C, 15Aa, 17B, 23A-M, 24. 1986] PERESKIA (CACTACEAE) 59

Cactus pereskia Linnaeus, Sp. pi. 469. 1753. Type. Rate 227, fig. 294 of Dillenius, Hortus elthamensis (1732), herewith designated as lectotype, rejecting the lectotype selected by Ben­ son (1982), a specimen without direct reference in Species plantarum, ed. 1 (Linnaeus, 1753). Cactus lucidus Salisbury, Prodr. stirp. 349. 1796 (nom. illeg.). Pereskia pereskia (Linnaeus) Karsten, Deut. R. 888. 1882 (nom. illeg.). Pereskia aculeata Miller var. rubescens Pfeiffer, Enum. diagn. Cact. 176. 1837. Type. Not cited. Pereskia aculeata Miller var. rotundifolia Hort. Dyck. ex Pfeiffer, Enum. diagn. Cact. 176. 1837. Type. Not cited. Pereskia aculeata Miller var. lanceolata Pfeiffer, Enum. diagn. Cact. 176. 1837. Type. Not cited. Pereskia undulata Lemaire, Illustr. hortic. 5 (Misc.). 11. 1858. Type. Brazil. Santa Catarina, 1854, Fr. Devos s.n., cult, in coll. Verschaffelt (not preserved). Pereskia godseffiana hort., Gard. Chron. 43: 257. 1908. Type. Cult, in Hort. Sander (not pre­ served). Pereskia rubescens A. Houghton, J. Cact. Succ. Soc. Amer. 1: 3. 1929. Type. Not cited. Pereskia aculeata Miller f. rubescens (Pfeiffer) Krainz, Die Kakteen, A, 1 Mar 1969. Pereskia aculeata Miller var. godseffiana (Hort. ex Gard. Chron.) Backeberg et F. Knuth ex Krainz, Die Kakteen, A, 1 Mar 1969. Straggling to clambering shrub or liana, forming vines 3-10 m long; trunk 2- 3 cm in diam., possibly thicker; bark grayish-brown, longitudinally fissured; long shoots straggling-scandent to winding, distal twigs to ca. 4 mm thick, green, olive, or reddish when fresh. Areoles ca. 2 mm diam. on twigs; tomentum brown to gray, short; additional longer, white hairs often present in new growth; areoles accrescent, cushion-like on the trunk and to 15 mm in diam. and 5 mm high; producing dimorphic spines but no brachyblast leaves. Leaves lanceolate to oblong or ovate, rarely broadly ovate to nearly orbicular, variable in shape and size on the same plant, 4.5-7(-l 1) x 1.5-5 cm, in broad-leaved specimens often 6-7 x 3-4 cm, in narrow-leaved specimens often 8-11 x 2.5-4 cm; petiole distinct but short, 3-7 mm long; leaf base variable, cuneate, attenuate, or rounded in fresh leaves, greatly affected by shrinking and often attenuate in dried specimens; blade green, concolorous, or purplish below when fresh (conspicuously discolorous or variegated in cultivars); apex short-acuminate or acute; venation pinnate; midrib prominent below, often slightly impressed above when fresh; lateral veins 4-7, often inconspicuous in fresh material, diverging at angles of 35-55°. Spines di­ morphic; primary spines geminate, rarely three or one, sometimes anisomorphic, always recurved, claw-like, sometimes lacking on weak shoots and lateral twigs but usually vigorous on main shoots, 4-8 mm long, 2-5 mm thick at base and flattened to elliptic in transverse section, brown, dark brown, often greenish-white or green when young, becoming gray with age, persistent or caducous on older areoles; secondary spines straight, produced on older areoles of the main shoots, to 25 per areole on the trunk, spreading, subulate, 10-35 mm long and 0.5-1.5 mm thick, at base abruptly thickened to 2-3 mm diam.; brown to nearly black, with variously colored base in new growth, becoming gray with age. Inflorescences terminal and lateral on long shoots, racemose to profusely paniculate; lateral panicles with up to 70 flowers or more, the entire long shoots with hundreds of flowers; panicles drooping, bracteate. Flowers 2.5-5 cm in diam., whitish, fragrant; pedicels 5-15 mm long; receptacle cup-shaped to turbinate, 5-6(-8) mm in diam., green, with 6-15 areoles on inconspicuous to prominent podaria; areoles with sparse and short brown, yellowish-brown or grayish tomentum, the upper ones often densely hairy with trichomes l-2(-3) mm long; lower areoles often with 1- 3(-5) spreading, brown or stramineous, easily detachable spines 4-5(-9) mm long; lower bracts (6-)9-12(-16), spreading to recurved, lanceolate to linear-lanceolate, or narrowly elliptic, 8-25 x 2-6 mm, petiolate, green, often fallen in dried spec­ imens; upper bracts 3-8, erect or appressed in bud, narrowly to broadly triangular, 1-3 mm long and broad, fleshy, green; sepaloids 2-5, ovate to obovate, 12-21 x 60 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 23. Pereskia aculeata. A, flowering twig. B, flower. C, flower, l.s. D, flower bud. E, fruit without spines. F, fruit, l.s. G, spiny fruit. H, fruit, l.s. I, seeds. J, younger shoot (leaves fallen) with recurved spines. K, older stem with additional straight spines. L, areoles of older stem. M, average leaves of different specimens. A from Brade 17425 (B); B from cult. SSZ, photograph; C, D from cult. SSZ (B); E, F from Czerwenka OV744 (B); G, H from cult. hort. Held. (HEID); I from Widgren s.n. (UPS); J-L from cult. hort. Berol. 001-11-74-80 (B); Ma from Santoro 7675 (UC); Mb from 1986] PERESKIA (CACTACEAE) 61

4-11 mm, acute, to rounded with a point, pale green or with green center and whitish, very thin margin; petaloids 6-11, obovate to spathulate, 15-25 x 6-10 mm, the inner ones usually narrower, very thin and delicate, almost translucent, white to greenish-white or with yellowish or pink hue; base cuneate, apex rounded and apiculate to subacute or truncate; stamens ca. 100-260, 5-10 mm long, the outer twice as long as the inner and all shorter than the style; filaments with the connate bases adnate to the elongated receptacular cup, forming a narrow nectarial furrow around the cylindrical style base, bending outward and spreading to as­ cending, white at base, white, yellow to orange-red, or bright coppery-red towards the tip; anthers 1-1.2 x 0.4-0.5 mm, yellow; pollen mostly 6-9-colpate, variable, medium-sized, tectum perforate and with small spinules; ovary cylindric, not distinctly delimited from the style and of about the same diameter, adnate to the receptacular cup with its base only, thence the locule appearing to be a cavity of the style base of ca. 1 mm diam. at anthesis; ovary half superior to superior, but surrounded by the short-tubular rim of the receptacular cup and filament bases; locule very small, with "basal" placenta and ca. five ovules; style 10—12(—19) mm long and ca. 1-2 mm thick, gradually thickened towards the stigma, white; stigma lobes 4-7, erect or suberect, white, 3-5(-7) mm long. Infructescence racemose or paniculate, or fruit solitary, pedicellate, fleshy, 15-25 mm in diam., globular, sub­ globular to depressed-obovate, yellow to orange at maturity, bearing 6-15 areoles with sparse tomentum, usually bracteate and spiny when immature, but mostly naked at maturity, the bracts to 25 x 6 mm, spines 3-8 mm long, spreading; umbilicus narrow, filled by the connivent upper bracts and flower remnants (if persistent); fruit wall fleshy, ovary roof enclosed but large and dome-shaped, with gelatinous tissue enclosing the seeds, or locule eventually hollow if seed incom­ pletely developed. Seeds 2-5, lenticular, 4.5-5 mm in diam., laterally flattened and often slightly concave on one side (adaxial side in the locule); 1.4-1.6 mm thick, black, smooth on the sides, with fine low peripheral ribs; hilum basal. Seedlings (three weeks old) with hypocotyl ca. 25 mm long; cotyledons elliptic- ovate, ca. 20 x 6 mm, accrescent and broadly ovate to subcordate at base and with pinnate venation in seedlings with developing primary shoot. Chromosome number: In = 22; cult. hort. Berol. 001-19-74-80 (B). Distribution (Fig. 24) and phenology. West Indies, northern, eastern, and south­ eastern South America; in Florida, Central America, and southern Mexico prob­ ably only subspontaneous or spontaneous after cultivation; in and outside its natural range also often cultivated and subspontaneous, or spontaneous after cultivation; from sea level to ca. 1000 m, in rocky areas, both calcareous and igneous, in and at the fringe of mesophytic forest, in dry forest, restinga, secondary vegetation, capoeira, and on dunes near the sea; flowering mostly during the humid season, north of the equator mainly from August to October, south of the equator mainly from January to April; fruits maturing ca. 2-3 months after flowering. The data of origin given by Linnaeus (1753) are erroneous. Plants from Mar­ garita (Isla de Coche, Venezuela) were in fact P. guamacho, and Jamaica could not be confirmed as a locality for P. aculeata, except for cultivated specimens (Adams, 1972). In most cases the cultivated or native status is not indicated on herbarium labels and is often doubtful. Although the species is well represented in herbaria,

Broadway 4290 (E); Mc from Glaziou 18267 (R); Md from Czerwenka OV744 (B); Me from Reitz 4591 (NY). 62 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Prepared by Hendrik R Rypki

© 1979 by the University of Utrecht Published by the State University of Utrecht, the Netherlands Department of Systematic Botany FIG. 24. Distribution of Pereskia aculeata. its natural range and its distribution as a cultivated plant for local use remain rather insufficiently known. Since the species constitutes a conspicuous and pos­ sibly subspontaneous element of the local garden flora and in secondary vegetation, specimens from outside botanical gardens and major parks are included in the distribution map with a different symbol but not listed among the specimens cited. Some of the specimens listed may also represent cultivated rather than indigenous plants. Variability and problems of delimitation of possible subspecific taxa are discussed below after the taxonomic history. Introduced P. aculeata is becoming a threat to local vegetation in coastal regions of Natal, (H. G. Zim- mermann, pers. comm.).

Representative specimens. USA. FLORIDA: Highlands County, vie. Avon Park, 2 Aug 1931 (st), McFarlin 6346 (MICH); Manatee County, Oneca, 1910, Reasoner s.n. (US); Palm Beach County, W of Palm Beach, 2 Dec 1919 (st), Small et al. 9220 (NY); St. Lucie County, 6 mi S of Ft. Pierce, 20 Dec 1917 (st), Small 8458 (G, MICH, NY, US), 10 May 1918 (fr), Small 8751 (NY). MEXICO. YUCATAN: Without exact locality, Gaumer 24374 (F, NY, US). PANAMA. PANAMA: Cerro Campana, 800 m, 7 Jan 1947 (fr), Allen 3972 (G). CUBA. SANTIAGO DE CUBA (ORIENTE): Rd. to El Chote, Alto Songo, Dec 1947 (fr), Clemente 5713 (GH). PINAR DEL Rio: Pinar del Rio, km 12 of highway to La Paloma, 28 Oct 1923 (fl), Ekman 17801 (S). . Cul-de-Sac, near Port-au-Prince, 10 Oct 1927 (st), Ekman s.n. (S); vie. Port-au-Prince, 1 Mar 1926 (st), Leonard 10122 (GH, NY, US). DOMINICAN REPUBLIC. BARAHONA: 50 m, Sep 1910 (fl), Fuertes 589 (E, G, GH, LIL, NY P, 1986] PERESKIA (CACTACEAE) 6 3

U, US). LA ALTAGRACIA: Km 15 from Punta Cana on rd. to Otra Banda, 18°34'N, 68°29'W, 120-130 m, 14 May 1980 (st), Mejia & Zanoni 6343 (JBSD), 29 Mar 1981 (fr), Zanoni et al. 12010 (JBSD). PEDERNALES: 10 km SW Oviedo, Loma Toussaint, 18 Oct 1982 (fr), Czerwenka OV744 (B). . VIEQUES: Cerro Encanta, 230 m, 28 Jan 1914 (fr), Shafer 2563 (NY, US). . ST. CROIX: St. Croix, 6 Jul/8 Aug 1870 (fl), Eggers s.n. (C). ST. THOMAS: "Ad habitationes naturalis," Nov 1882 (fl), Eggers 875 (B, BR, G, GH, GOET, LE, M, P, W). LEEWARD ISLANDS. ANTIGUA: Sugar Loaf Mtn., 250 m, 24 Sep 1938 (st), Box 1539 (BM, MICH, US); 4-16 Feb 1913 (fr), Rose et al. 3434 (NY, US). SABA: From Windwardside to Bottom, Boldingh 1858a (U). ST. BARTHELEMY: Forsstrom s.n. (S). WINDWARD ISLANDS. GUADALOUPE: DUSS 977 (P); Baillif, Mar 1894 (fl, fr), Duss 3053 (NY); Baillif, Gombeyre, Basse Terre, Duss 3073 (F, GH, NY, US). : Case Pilote, Duss 958 (NY, US). VENEZUELA. BOLIVAR: E of Miamo, Hato de Nuria, 400-550 m, 25 Jan 1961 (st), Steyermark 88832 (NY, U). CARABOBO: Vic. Puerto Cabello, 28-29 Oct 1916 (fr), Rose 21868 (NY, US); Puerto Cabello-Valencia, nr. El Palito, Hacienda Taborda, 20-100 m, 30 Oct 1919 (fl), Pittier 8633 (GH, US). DISTRITO FEDERAL: S of La Guayra, ca. 300 m, Fendler 67 (MO); La Guayra, Boldingh 4131 (U); Cambari nr. La Guayra, Karsten s.n. (W). MIRANDA: Morros de la Guairita, 900-950 m, 16 Aug 1975 (fr), Berry 1009 (VEN). SUCRE: Vic. Cristobal Colon, Jan-Feb 1923 (fr), Broadway 372 (NY, US). ZULIA: Vic. Perija, Tejera 184 (US). TRINIDAD. Magueripe, 3 Mar 1920 (st), Britton et al. 189 (GH, NY, US). TOBAGO. Windward, 15 Oct 1912 (fl), Broadway 4290 (BM, E, F, G, L). GUYANA. Mabaruma, Aruka River, 15 Jul 1952 (fl), Fanshawe 3411 (K, NY, S, U), British Guiana Forest Dept. 6975 (NY, S). BRAZIL. BAHIA: Mun. de Aiquara, 2 km N of Aiquara, 13 May 1969 (fl), Jesus 375 (CEPEC); Mun. de Alcobaca, km 5 of rd. from Alcobaca to Caravelas, 4 May 1973 (fl), Pinheiro 2172 (CEPEC, RB); Barragem de Bananeiras, Rio Paraguacu and Jacuipe, 12°32'S, 39°05'W, Apr 1980 (fr), Cavalo 4 (ALCB); Cruz das Almas, 30 May 1981 (fl), Pinto 83 (HRB); Mun. de Ilheus, km 22, rd. from Ilheus to Itabuna, on grounds of CEPLAC, 25 Apr 1983 (fl), Leuenberger & Brito 3103 (B, CEPEC, K, SI, W); Ilheus, 11 Jun 1917 (fr), Zehntner s.n. (US); on rd. from Itajuipe to Ubaitaba, 12 km N of Banco Central, 24 Apr 1965 (fl), Belem & Magalhaes 877 (CEPEC, UB); Mun. de Itaberaba, Fazenda Monte Verde, Nov 1973 (st), Costa s.n. (ALCB); between Prado and Alcobaca, 16 Apr 1967 (fl), Castellanos 26413 (CEPEC, K, M); Mun. de Rui Barbosa, Serra de Orobo, 1 Jul 1973 (fl), Pinto s.n. (ALCB); vie. Salvador de Bahia, 26 May 1915 (st), Rose & Russell 19639 (NY, US); vie. Toca de Onca, 27-29 Jun 1915 (fl, fr), Rose & Russell 20069 (NY, US). ESPIRITO SANTO: Colatina, 30 Apr 1934 (fl), Kuhlmann 290 (MO, RB, UC); Itapemirim, Bello 472 (R); Vitoria, Gagzo s.n. (HBG); Mun. de Vila Velha, between Lagoa do Milho and Barra do Jucu, 20 Jul 1973 (fr), Araujo 369 (RB). MINAS GERAIS: Caldas, Regnell III-1776 (UPS); Itabira, Cane, 11 Feb 1934 (fl), Sampaio 7069 (R); Lagoa Santa, Warming s.n. (C); Mun. de Sao Joao del Rei, Apr 1896 (fl), Silveira 943 (R); Vicosa, 19 Dec 1929 (fl), Mexia 4129 (BM, F, G, GH, K, LIL, MICH, MO, NY, S, U, UC, US); Vicosa, km 4.5 on rd. to Sao Miguel, 750 m, 27 June 1930 (fr), Mexia 4804 (BM, F, G, GH, K, MO, NY, S, U, UC, US). GOIAS: Near Goias, 20 Jul 1928 (st), Burchell 7347-2 (K). PARANA: Campina do Sul, Rio Pardinho, 30 Jan 1969 (fl), Hatschbach 20973 (UC); Cascavel, 10 Jan 1953 (fl), Rambo PACA 53529 (B); Foz do Iguacu, 31 Mar 1941 (fl, fr), Kuhlmann s.n. (MO, RB); Mun. de Sta. Helena, Barra do Rio Sao Francisco, 26 Mar 1977 (fl), Hatschbach 39856 (CTES); Theresina, 29 Jan 1911 (fl), Dusen 11282 (P, S); vie. Umuarama, 30 Feb 1980 (fl), Furtado 160 (HRB); Mun. de Umuarama, Ivate, 18 Apr 1968 (fl), Hatschbach 19046 (C, CTES, F, K, NY, S, UC, US). PERNAMBUCO: Olinda, restinga do Rio Doce, 5 Jan 1948 (st), Leal 41 (RB). Rio GRANDE DO SUL: Cerro Largo, nr. S. Luiz, Jan 1943 (fl), Rambo 11159 (B, LIL); Santo Angelo, Jan 1893 (fl), Lindman s.n. (S); Schwabenschneis, Feitoria river, 25 May 1949 (fr), Rambo 41779 (F, LIL, P, W); Taquara, Amaral Ribeiro, 5 Jul 1949 (fr), Rambo 42387 (LIL). Rio DE JANEIRO: Campos, Apr 1939 (fl), Sampaio s.n. (R); Carmo, Neves-Armond 11 (R); Itaquiatara, 29 Mar 1964 (fl), Pabst HB 29478 (B); Itatiaia, 1000 m, 13 Feb 1945 (fl), Brade 17425 (B) Brade 17435 (RB); Itatiaia, Maromba, 11 Apr 1929 (fl), Campos Porto 1714 (RB); Mt. Itatiaia, vie. Ponte Maromba, 1200 m, 22°28'S, 44°38'W, 12 Apr 1929 (fl), Ferreira 2316 (BM, F, GH, K, S, US); Mt. Itatiaia, 1060 m, 26 Oct 1927 (fl), Ginzberger 61 (F); Marica, Lagoa de Padre, 16 Apr 1975 (fl), Araujo 680 (RB), 26 Jun 1979 (fr), Araujo 766 (RB); Rio de Janeiro, Barra da Tijuca, 26 Mar 1964 (fl), Trinta 535 (M, R); Rio de Janeiro, Corcovado, Luschnath s.n. (LE); Mun. de Rio de Janeiro, Restinga do Grumari, 24 Feb 1972 (fl), Jesus 1333 (B, RB); Praia de Grumari, 26 Mar 1979 (fl), Yautone 655 (RB); Ilha Furtada, Bahia de Sepetiba, 31 Mar 1968 (fr), Sucre 2623 (RB); Ilha de Paqueta, 20 Jun 1960 (fr) Scheinvar 112 (R); Jacarepagua, May 1898 (fl), Vie 4690 (HBG); Reserva Biol de'Jacarepagua, 3 May 1969 (fr), Colaris 1032 (U); Morro da Viracao, Praia Grande, 3 Apr 1891 (fl), Glaziou 18267 (C, G, K, LE, P, R); Praia Vermelha, Pao de Acucar, 13 Apr 1977 (fl), Rodrigues 75 (CTES, RB); Recreio dos Bandeirantes, 12 May 1959 (fl), Duarte 4806 (RB), 14 Apr 1931 (fl), Lutz 566 (R, US); Restinga de Tijuca, 24 Apr 1942 (fl), Machado 63 (MO, RB); Sepetiba, 2 Apr 1896 (fl), Ule 3816 (HBG); Mun. de Sao Jose do Rio Preto, Valverde, Feb 1963 (fl), Pabst 7310 64 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

(M); Mun. de Saquarema, Praia Jacone, 30 May 1978 (fl), Martinelli 4511 (RB); Serra dos Orgaos, Jan 1834 (fl), Luschnath s.n. (BR, MO); Socavao, Bananal, Mar 1883 (fl), Palma s.n. (R). 5>ANTA CATARINA: Ilha Santa Catarina, Rio Tavares, 11 Mar 1953 (fl), Reitz & Klein 310 (LIL, S); Itapiranga, 15 Feb 1934 (fl), Rambo 1522 (LIL); Joinville, 15 Feb 1910, Ehrhardt 15 (HBG); Mun de Mondai, Pinheiral, S of Sao Miguel d'Oeste, 600-700 m, 23 Feb 1957 (fl), Smith & Klein 11720 (K, NY, UC, US); Pirao Frio, Sombrio, 10 m, 17 Mar 1960 (fl), Reitz & Klein 9564 (L, UC, US); Rio Capinzal, 26 Feb 1916 (fl), Dusen 17842 (S). SAo PAULO: Campinas, 1 Apr 1906 (fl), Heiner 637 (S); Mun. de Campinas, mata da Fazenda Sta. Genebra, 10 Apr 1978 (fl), Neto 7377 (UEC); Fazenda Sao Quinno, 28 Jan 1970 (fl), Souza s.n. (CTES); Morro das Pedras, Brade 9023 (R); Serra do Caracol, 20 Feb 1874(A), Mosen 1330 (S). PARAGUAY. ALTO PARANA: Fiebrig 6485 (G, GH, LIL, SI, US). CENTRAL: Asuncion, cult., orig­ inally from Colonia Independencia, Guaira, 2 Apr 1973 (fl), Schinini 6165 (CTES, MO, SI). SAN PEDRO: Primavera, 13 Jul 1958 (fr), Woolston 1002 (K, NY, S, SI). GUAIRA: Villarnca, Feb 1876 (fl), Balansa 2502 (G, K, P). ARGENTINA. CORRIENTES: "Bresil, Corrientes," Jul 1821 (fr), Bonpland 641 (MO, P). MISIONES: Cainguas, Puerto Tabay, 6 Apr 1949 (fl), Schwarz 7670 (LIL), 16 Apr 1950 (fr), Schwarz 10463 (LIL, MO); Candelaria, 19 Feb 1948 (fl), Bertoni 3628 (LIL); Candelaria, San Juan, 7 Jul 1947 (fr), Schwindt 313 (LIL); Eldorado, 8 Mar 1944 (fl), Meyer 6747 (LIL); Iguazu, Reserva Forestal Parque Nacional de Iguazu, 25°44'S, 54°26'W, 1 Feb 1976 (fl), Romanczuk et al. 648 (SI); Monte Carlo, Ayo. Piray Guazu y ruta 12, 15 Feb 1980 (fl), Schinini 19896 (CTES, SI); Obera, Guarany, 420 m, 1 Mar 1950 (fl), Schwindt 3319 (LIL); Posadas, 12 Mar 1917 (fl), Shafer 129 (G, GH, K, MO, NY, US); Posadas, Garupa, 12 Mar 1946 (fl), Bertoni 2770 (LIL); San Pedro, Barranon, 560 m, 2 Mar 1950 (fl), Schwindt 4092 (LIL). Local names and uses. Cuba: grosellero, jasmin de uvas, ramo de novia. Do­ minican Republic: camelia blanca. Puerto Rico: gooseberry. St. Thomas: Surinam gooseberry. , Martinique: groseiller criole, groseille-pays, groseille-Barbade, groseiller-pays, groseiller de Barbades, groseiller de la Barbade. Brazil: Ora pro nobis, azedinha (Bahia); lobolobo (Minas Gerais); cipo estrela (Parana); padre nostro (Rio de Janeiro). Paraguay: curuzu ipochi. The leaves are used as a pot herb or vegetable, particularly in rural areas of Minas Gerais and Bahia. The fruits are edible. Because of the showy flowers it is also planted as an ornamental, but the strong, rather unpleasant fragrance limits its use. It is probably the most frequently cultivated Pereskia in botanical gardens. Taxonomic history and typification. Pereskia aculeata was first described in Plumier's unpublished manuscript "Botanica americanarum . . ." (Hunt, 1984). The much shorter description given by Plumier (1703) was followed by a lengthy dedication to Nicolas Fabricius de Peiresc, for whom the genus was named. Linnaeus (1753) accepted only one genus in the Cactaceae and consequently named the species Cactus pereskia. His citations, however, contain two different elements, one being Pereskia aculeata (four citations), the other representing the plant described much later as P. guamacho (two citations). No specimen or il­ lustration is directly linked to the citation "hort. Cliff. 122" (correctly 183). Lec- totypification following normal procedures for Linnean names fortunately guar­ antees nomenclatural stability despite the fact that Cactus pereskia L. contains different elements. Nevertheless, the choice of a lectotype has been the subject of discussion. The Linnean herbarium of the Swedish Museum of Natural History at Stockholm holds two specimens from the Uppsala Botanical Garden that are linked to Linnaeus' citations. The phrase name and citation "Hort. Ups. 122." can be linked to only one of these specimens. The older one, apparently a de­ pauperate sterile plant without spines and merely three elliptic leaves, bears the same phrase name (in the handwriting of D. Solander, before 1758) as Cactus pereskia L. in "Species plantarum," ed. 1 (Linnaeus, 1753), viz., "Cactus caule tereti arboreo spinoso, foliis lanceolato ovatis. Linn. spec, plant. 469. 21. Hort." It would be preferable as the lectotype to the one selected by Benson (1982, p. 969), which only has a direct reference to "Systema naturae," ed. 13 (Linnaeus 1986] PERESKIA (CACTACEAE) 65

1770), in L. J. Montin's handwriting "specimen ex horto Upsal. communicavit Hortulanus Nietzel." It is possible that both specimens were taken from the same cultivated plant. Depauperate specimens often lack the characteristic geminate recurved spines and vary greatly in leaf shape. The character "aculeis geminis recurvis," so typical for the species, was only added to the phrase in the second edition of "Species plantarum" (Linnaeus, 1762). The second specimen from the Hortus Upsaliensis exhibits this character, which is why Benson (1982) probably chose it, but unfortunately it is evidently more recent than both the first and second edition of "Species plantarum," hence not suitable as a lectotype. Although I am convinced that the older, but poor, specimen represents P. aculeata, its choice as lectotype for Cactus pereskia could easily be questioned, as it lacks the crucial characters to separate this species from P. guamacho, the other element represented by some citations under Linnaeus' Cactus pereskia. Among the other citations identified with P. aculeata, that of Plumier (1703; "Plum. cat. 37") lacks a specimen. The illustration belonging to it is not cited by Linnaeus (1753). The plate of Dillenius (1732), cited as "Dill. elt. 303, t. 227, f. 294," is more representative and seems to be the best choice for lectotype of Cactus pereskia, the correct name for both Plumier's plant and the specimens cultivated in the Hortus Elthamensis and Hortus Upsaliensis. In agreement with Wijnands (1983), I have therefore selected this plate as lectotype. A specimen at the Fielding-Druce Herbarium, Oxford, annotated by Sherard with plate and figure number referring to Dillenius's Hortus Elthamensis and possibly belonging to the plate, although sterile, is undoubtedly P. aculeata. The genus Pereskia was resurrected by Miller (1754), and the new name for Cactus pereskia published by Miller (1768) in the "Gardeners Dictionary," 8th edition, Pereskia aculeata, takes up Plumier's first epithet and is legitimate for nomenclatural reasons instead of the tautonym Pereskia pereskia. As already noted by Schumann (1898, 1903) and Britton and Rose (1919), P. aculeata is a polymorphic species, possibly a complex of varieties, forms, and clones. To these authors the varieties described earlier by Pfeiffer (1837a) seemed untenable, being based on variable or unstable characters such as size, shape, and coloration of leaves. Even today, with a much larger number of specimens avail­ able for study, it is impossible to delimit adequately the infraspecific entities which possibly exist. Herbarium material alone does not allow a thorough analysis of potentially useful character combinations and their correlation with geograph­ ical and ecological distribution. Paucity of characters, lack of notes on badly preserved characters, and artefacts caused by different drying methods are the main reasons besides the usually fragmentary or incomplete specimens. In the future, collectors should make: 1) complete specimens, including areoles from the trunk, representative vege­ tative twigs and inflorescence(s); 2) notes on habit, size of plant, trunk, color of leaves, sepaloids, petaloids, filaments and style, fragrance; 3) notes on shrinkage of leaves, method of drying (it is noteworthy that shrinking may depend on several yet insufficiently known factors; broad leaves may remain so in the dry condition or become narrow and indistinguishable from narrow leaves); 4) observations on variability in nature and under controlled conditions (leaf size and shape particularly); and 5) notes on ecological data (habitat, soil, geology, phenology). Possible diagnostic characters for subspecific entities are the spination of re­ ceptacle and fruit, color of filaments, number and shape of petaloids, and leaf 66 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL- 41

shape and color. However, at this time, a key based on such characters would not permit identification of most herbarium specimens available, which, however incomplete or fragmentary, can still be easily identified to species. Separation of subspecific taxa would therefore be impractical. Since certain distinguishable clones exist in cultivation, cultivar names seem appropriate for horticultural use, and are to be discussed in a separate paper. Varietal names proposed by Pfeiffer (1837a, 1837b) can be used as cultivar names. Two of them are documented by possibly authentic material from the Salm-Dyck collection dated 1857 (P. aculeata var. rotundifolia, sheet no. 2755742 and P. aculeata var. lanceolata, sheet no. 2755743 both at MO, photographs at B). For this monograph it has seemed preferable to treat this taxon as one polymorphic species. The analysis of character correlations, based on more than 200 herbarium specimens and a few living collections of known wild origin, shows a reticulate pattern showing some general trends. In the northern range of distribution, narrow to very broad leaves, weakly spiny receptacles, and spineless fruits (Fig. 23E, Mb, d) seem to be almost the rule, and the few specimens seen in flower all had white filaments. In the southern distributional area, different leaf shapes also occur (Fig. 23Ma, c, e), but the southern-most collections mostly have very narrow and long leaves with acuminate apices; and spiny receptacles and fruits are frequent, as are brightly colored stamens. However, Esser (1982) found plants with white filaments in Paraguay as well, and this character is known in less than 10 percent of all herbarium specimens. The difficulty of a safe interpretation and evaluation of different leaf shapes (a very conspicuous character in fresh material) has been outlined above. Variegated forms found in cultivation represent cultivars, e.g., P. aculeata 'Godseffiana,' with entirely apricot-colored upper leaf surface and reddish underside. Nomina nuda listed by Forster (1846), Pfeiffer (1837a, 1837b), and Backeberg (1958) as synonyms of P. aculeata are P. acardia, P. brasiliensis, P. foetens, P. fragrans, P. hortensis, P. lanceolata Hort. Hamb., and P. longissima. Pereskia discolor F. A. C. Weber is also a nomen nudum. Plants found in cultivation under this name are P. aculeata 'Godseffiana.'

2. Pereskia humboldtii Britton & Rose, Cactaceae 4: 251. 1923. Shrub, to 3 m high, or small tree, 2-6 m, with divaricate and epitonous branch­ ing. Roots with fusiform to subglobular thickenings (observed in cultivated spec­ imens); distal twigs 1.5-3(-4) mm thick, often markedly zigzag; internodes 1-3 cm long; epidermis glabrous or minutely papillate with stout papillae to 0.1 mm long. Areoles 2-4 mm in diam. on twigs, accrescent on branches and trunk, with grayish-white to white tomentum of trichomes to ca. 1 mm long, particularly in new growth with few to many additional longer, white, crisp to flexuous, coarse hairs 5-20 mm long; areoles producing spines but no brachyblast leaves. Leaves elliptic-ovate to narrowly or broadly elliptic ovate or obovate, 15-40(-60) x 10- 25(-30) mm; petiole inconspicuous or to 1-2 mm long and thick; blade thick, base cuneate to attenuate or rounded; apex shortly acute to acute; venation rather inconspicuous, in fresh leaves slightly impressed above; midrib prominent below; lateral veins 0-2(-4). Spines on twigs (0-)l-4 per areole, increasing in number on the branches and main shoots to 10 or more (areoles of trunk not seen), porrect to spreading, straight, acicular to subulate, very unequal in length, (0.5-)l-4(-5) cm long and 0.3-1.0(-l.2) mm thick, at base to 1.5(-3) mm thick, not conspic­ uously bulbous but often angled, flattened or contorted, stramineous or yellow or white when young, brown to gray in age. Inflorescence condensed cymose-pa- 1986] PERESKIA (CACTACEAE) 67 niculate. Flowers terminal and in axillary clusters of 2-6, perigynous to nearly hypogynous, 5-15(-20) mm in diam.; receptacle broadly cup-shaped, becoming depressed subglobular soon after anthesis, ca. 3-4 mm in diam.; receptacular areoles to ca. five, with scarce to abundant wool of trichomes 1-2 mm long, rarely with 1-3 spines, the upper ones with longer, woolly or silky hairs 2-3(-5) mm long; lower receptacular bracts 0-3, very variable in size, leafy, elliptic to obovate, or triangular, smaller, 2-10(-15) x l-5(-10) mm, on terminal flowers nearly attaining the size of regular leaves; upper bracts 3-5, triangular to narrowly tri­ angular, 1-2 mm long and broad; sepaloids ca. three, triangular-ovate, 4-10 x 2-3 mm, greenish-red or greenish-white; petaloids usually five, ovate to lanceolate, variable in shape, ca. 5-10 x 3 mm, acute to acuminate, dull red, orange-red, brownish-orange, or white; stamens ca. 40-60, 4-5 mm long; filaments reddish towards the tip, entirely white in white flowers; anthers 0.7-0.9 x 0.3-0.4 mm, yellow; pollen 3(-6)-colpate, small, tectum finely perforate and with small spinules; ovary superior to half inferior, the free ovary roof often bell-shaped, adnate to the receptacle with its base only; locule with flat floor, ca. 0.7-1 mm in diam., septal ridges extending to the ovary floor; placentae at the base of the locule; style 4-5 mm long, style base conical; stigma lobes 3-5, 1-2 mm long. Fruit a soft, very broadly obovate berry, 5-6 mm in diam., green to vinaceous when immature, black when mature, juicy; fruit wall with 0-3 white-hairy areoles, the apical, depressed, narrow umbilicus with longer hairs and persistent flower remnants; locule hollow, filled by the seeds. Seeds variable, lenticular to obovate, 1.5-1.8 mm long, 1.3-1.5 mm broad and 0.6-0.7 mm thick, with shallow peripheral ribs, smooth on the sides, black, glossy; hilum lunate to subquadrangular, white, with central funicle scar and flat to prominent rim.

Key to the Varieties of Pereskia humboldtii 1. Twigs (internodes) smooth, without minute papillae; flowers dull-red to orange-red, or brownish-orange, rarely white. 2a. var. humboldtii. 1. Twigs minutely papillate, with stout papillae 0.05-0.1 mm long (Figs. 18A-C, 25L-N; use hand lens, at least 10 x magnification); flowers white. 2b. var. rauhii.

2a. Pereskia humboldtii Britton & Rose var. humboldtii. Figs. 6C, 25A-J, 26. Pereskia humboldtii Britton & Rose, Cactaceae 4: 251. 1923. Cactus horridus Kunth in Humboldt, Bonpland & Kunth, Nov. gen. & sp. 6: 70. 1823 (non Salisbury, 1796). Type. Peru. Jaen de Bracamoros, in collibus siccis, ad flumen Maranon (Aug 1802), Humboldt & Bonpland 3594 (lectotype, P; isotypes, B-W, MO, P). Pereskia horrida (Kunth) De Candolle, Prodr. 3: 475. 1828. Rhodocactus horridus (Kunth) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. IPereskia vargasii H. Johnson [var. vargasii], Cact. Succ. J. (Los Angeles) 24: 115. 1952. Type. Peru. Jaen, east bank of Rio Maranon opposite Bella Vista, 500 m, discovered by C. Vargas and H. Johnson, but no specimen cited and no preserved or living material located. Shrub to small tree; twigs with smooth internodes; leaf shape varying from broadly ovate or obovate to narrowly elliptic; flowers dull-red to orange-red, brownish-orange, or rarely white. Chromosome number: In = 22; Hutchison 1452, hort. Berol. 256-01-82-30 (B). Distribution (Fig. 26) and phenology. Peru, dry valleys of the Rio Maranon drainage system, in dry forest and brushland (monte bajo subxerofilo) from ca. 500 to 760 m as far as known from the few collections available; distribution insufficiently known. Specimens examined. PERU. AMAZONAS: Bagua, near Bagua Chica, 500-600 m, 3 Mar 1961 (fl, fr), Ferreyra 14202, 14204 (A); Bagua, nr. Bagua Grande, Rio Utcubamba, Hacienda Marerilla, 500 [VOL. 41 68 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN

FIG. 25. Pereskia humboldtii and varieties. A-J, Pereskia humboldtii var. humboldtii (A-C, E-G, J from Hutchison 1452 (B); D, H, I from Rauh 24505 (B)). A, one-year-old twig with flower and flower buds. B, flower bud of the same twig. C, twig of new growth with terminal flower. D, terminal flower of white-flowered specimen. E, flower, l.s. F, twig with fruits. G, young and mature axillary fruits. H, terminal fruit. I, fruit, l.s. J, seeds. K-N, Pereskia humboldtii var. rauhii (K-N from Rauh 1986] PERESKIA (CACTACEAE) 69

m, 29 Sep 1957 (fl), Hutchison 1452 (G, GH, K, LIL, MICH, P, S, US); Bongara, Capallin, Cajaruro, 11 Dec 1952 (fl), Diaz s.n. (UC); between Monte Negro and Imacita, 9 Aug 1970, Rauh 24505, cult, hort. bot. Heid. et cult. hort. bot. Berol. (B). CAJAMARCA: 23.6 km E of Pucara, 760 m 19 Jan 1964 (fl), Hutchison & Wright 3621 (US). The new name used by Britton and Rose (1923) was repeatedly questioned by later authors (Ritter, 1981; Vaupel, 1925) but it is accepted here because Cactus horridus Salisbury (1796) was effectively published and may be considered as a specific name for a particularly spiny variety of Cactus tuna L. The principal diagnostic feature (smooth stem), which separates the typical variety from var. rauhii (with papillose stem), needs further study based on more material. Although the material is easily sorted into two groups, the occurrence of two slightly papillose twiglets among typically glabrous isotype material of P. humboldtii (at B-W and MO) is puzzling and could be used to deny any taxonomic status to the varieties accepted here. Flower color does not seem to be useful to distinguish species or varieties within the Andean group, as the example of P. weberiana (see also under Hybridization) shows. Therefore, a glabrous, white-flowered specimen {Rauh 24505) is included in this variety. Rauh (1958) already noted that Pereskia vargasii may just be a variety of P. humboldtii and this opinion is supported here with the modification that P. vargasii var. vargasii is placed, with a question mark, in the synonymy of P. humboldtii var. humboldtii. Pereskia vargasii lacks a type specimen and the description is ambiguous, because the Latin diagnosis states "flowers white" but "outer petals green, . . . the inner purple or pink" while the English description merely says "flowers white" (Johnson, 1952). Nothing is said about the stem surface, but from the illustration one could assume that P. vargasii was a smooth plant. In that case, however, it is best placed in the synonymy of P. humboldtii var. humboldtii. Rauh (1958) did not comment on this. The varietal names proposed by Backeberg, and Rauh and Backeberg (in Backeberg, 1956), on the other hand, represent a distinctive taxon with papillose stems, which is discussed below.

2b. Pereskia humboldtii Britton & Rose var. rauhii (Backeberg) Leuenberger comb. nov. Figs. 18A-C, 25K-N, 26. Pereskia vargasii H. Johnson var. rauhii Backeberg, Descr. Cact. Nov. 6. 1956. Type. Peru. Olmos-Jaen, Maranon valley, 1000 m, 7 Oct 1954 (fl), Rauh & Hirsch P2162 (HEID). Cited as "Jaen, near Bella Vista, 1000 m, type-no. Heidelberg" by Backeberg (1956), and illustrated in Backeberg (1958: 112, fig. 46). Pereskia vargasii H. Johnson var. longispina Rauh et Backeberg, Descr. Cact. Nov. 6. 1956; amplified description in Rauh, Beitrag Kenntnis Peruan. Kakteen veg. 190. 1958. Type. Peru. Jaen, Rauh K80 (specimen lost, according to Rauh, pers. comm.). Shrub with minutely papillate twigs; papillae 0.05-0.1 mm long; flowers white. Distribution (Fig. 26). Peru, in dry valleys of the Rio Maranon drainage system, from ca. 500 to 2100 m; occurring together with var. humboldtii at lower altitudes, but range of distribution insufficiently known.

Specimens examined. PERU. WITHOUT EXACT LOCALITY: "Hda. Valor," Ritter 283 (SGO). AMAZONAS: East of Balsas, Ritter 284 partly (SGO); between Imacita and Bagua Grande, 17 Jul 1976, Rauh 40184, cult. hort. Berol. (B). CAJAMARCA: Jaen, 5 km S of Jaen, 950 m, 28 Sep 1957 (fl, fr), Hutchison 1442 (US); Chamaya, Rauh 53666, cult. hort. Berol. (B). LA LIBERTAD: Huamachuco, Rio Maranon canyon,

40184, B). K, twig. L, M, papillate epidermis at increasing magnification. N, epidermis and hypodermis, t.s., showing epidermal papillae and a druse of calcium oxalate crystals. 70 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 26. Distribution of Pereskia humboldtii and its varieties, P. diaz-romeroana and P. weber­ iana.

18 km below Aricapampa, 2100 m, 8 Aug 1964 (fl), Hutchison 6219 (US); El Chagual, Ritter 1079 (SGO). The papillose stem surface of this variety is a remarkable feature, unique within the genus. Material of the species is easily sorted into two groups, and varietal status seems appropriate. The observation of several living plants of different origin shows that the papillosity of the stem is a constant character. Nevertheless, the distribution of this character remains to be studied both in its geographical and ecological aspects and in relation to flower color. Rauh (1958) did not mention P. vargasii var. rauhii but only described P. vargasii var. longispina in detail, using diagnostic characters ("receptacular bracts rarely foliaceous") which do not prove suitable to distinguish it from P. humboldtii as a species or variety, because of the great variability of this character in all Andean species of Pereskia. However, a type specimen is not available for var. longispina, and var. rauhii is the only name available, which is documented by a type specimen representing a white- flowered plant with papillose stem. 1986] PERESKIA (CACTACEAE) 71

3. Pereskia diaz-romeroana Cardenas, Lilloa 23: 15, fig. 1, t. 1. 1950. Type. Bolivia. Cochabamba: Prov. Mizque, Ttacko Laguna-Pulquina, 1300 m, Oct 1947 (fl), Cardenas 4388 (lectotype and isotype, US). Figs. 15Ag, 19B, 26, 27A-K. Shrub 0.5-3 m high; main shoots erect and arching, with basitonous and epi- tonous branching. Main shoots (canes) to ca. 2 cm in diam., with gray or brown exfoliating bark; branches straight to zigzag, the lateral ones plagiotropic; twigs 2-3 mm thick, internodes mostly 5-20 mm long, with retarded periderm for­ mation. Roots with fusiform to globular, fleshy thickenings to 30 cm long and 4 cm thick (measurements from cultivated plants less than 1 m high), probably attaining larger size. Areoles on twigs rounded, 2-3 mm in diam., with short grayish-brown tomentum and few to numerous longer, shaggy, white hairs ca. 3- 4 mm long; trichomes with dark base; areoles accrescent to ca. 6-7 mm diam. on older branches, producing spines but no brachyblast leaves. Leaves elliptic to obovate, in some specimens broadly obovate, 2-3.5(-4) x 1.2-2.2 cm, sometimes larger when fresh; petiole inconspicuous, 1-2 mm long and thick when fresh; leaf blade 0.7-0.8 mm thick, base cuneate to cuneate-attenuate or almost rounded in very broad-leaved specimens; apex acute; venation pinnate, midrib somewhat prominent below when fresh, slightly impressed above; lateral veins 3-5, diverging at angles of 25-30(-40)°, arching, often inconspicuous in fresh leaves. Spines on twigs 2-5(-7) per areole, spreading, (5-)15-22(-30) mm long and 0.4-0.6 mm thick, thicker towards the base and often flattened and bulbous or conical at the base, straw-colored to brown, yellow in new growth; spines on older branches to 12 (or more) per areole, of unequal length, 5-30 mm long and 0.8-1.0 mm thick, often flattened or slightly contorted and to 2 mm in diam. at base, straight to somewhat deflexed. Inflorescence condensed cymose-paniculate, or flowers some­ times solitary, terminal and axillary, but usually in dense clusters of 2-3(-5) per areole, partly by proliferation from the receptacle. Flowers hypogynous to perig­ ynous, ca. 1-2 cm in diam., pedicellate to sessile; receptacle 4-5 mm in diam., turbinate or cupuliform; lower areoles hairy and rarely with 1-2 spines, upper areoles with numerous silky to shaggy hairs ca. 5 mm long which often nearly mask the bud; lower receptacular bracts 0-4, often leaf-like, elliptic-obovate to lanceolate, 8-16 x 3-10 mm, usually smaller on lateral flowers of each cluster; upper bracts 5-7(-9), erect, oblong, linear to narrowly or broadly triangular, ca. 1-2x1 mm, green, fleshy; sepaloids 2-4, ovate, to 5-7 x 3-5 mm, with broad base, green with pink or purplish-red margin; petaloids 4-6, pink to bright pur­ plish-red, narrowly to broadly obovate, 8-12 x 4-6 mm, base cuneate, apex rounded to acute. Stamens ca. 50-70, erect; filaments 3-7 mm long, conspicuously pink to purplish-red, often paler at base; anthers 0.7-0.8 x 0.3-0.4 mm, yellow. Pollen tricolpate, small, tectum with mostly fine perforations and spinules ca. 0.5 urn long. Ovary variously superior to half inferior, adnate to the receptacle with the base only, its upper part looking like a swollen style base, ca. 1.5 mm in diam.; locule with flat to slightly convex floor and incompletely septate roof; placenta at the base of the septal ridges along the margin of the locule floor; styles 5-7 mm long, white, stigma lobes 4-6, suberect, 1-2 mm long, white. Ovules ca. 10-20, 0.5 mm long. Fruit very broadly obovate to depressed subglobular, fleshy soft and juicy, 4-6 mm in diam. when dried, to ca. 13 mm in diam. when fresh, black, glossy, smooth except for 2-3 leaf scars and transverse linear areoles with white hairs 2-3 mm long; umbilicus depressed, narrow, filled by the connivent upper bracts, silky to shaggy white hairs and the crumpled, usually persistent flower remnants; fruit wall juicy, thick, brown or purplish-black inside at maturity; locule 72 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 27. Pereskia diaz-romeroana. A, flowering twig. B, flower, l.s. C, flower bud. D, flower at anthesis. E, flowering branch at an early stage, the lateral twigs with terminal flowers. F, twig with terminal and axillary clusters of withered flowers, one flowerbu d and one fruit. G, axillary infructes­ cence. H, fruit, l.s. I, seeds. J, areole from older stem. K, habit. A-I from Rauh 40627 (B); J, K from Cardenas 4388 including a photograph (US). 1986] PERESKIA (CACTACEAE) 73

hollow, with flat or convex floor, more or less filled by the seeds. Seeds few (ca. 5), obovate, 1.8-2.0 mm long, 1.3-1.6 mm broad, and 0.7-1.0 mm thick, with shallow peripheral ribs, otherwise smooth, black, glossy; hilum lunate, brown or cream, with central funicle scar; micropylar nose prominent but blunt or rather inconspicuous. Chromosome number: In = 22; Rauh 40627, cult. hort. Berol. 039-02-77-30 (B). A gametic chromosome number of n = 11 was reported by Ross (1981). Distribution (Fig. 26) and phenology. Central Bolivia, in dry valleys from 1300 to ca. 2000 m, in xerophytic open "cactus forest," rare. Rowers observed in March, June, and October. Fruits mature a few weeks after flowering.

Specimens examined. BOLIVIA. COCHABAMBA: Between Huerta Molino and Perez Totora, 1800 m, Oct 1947 (fl), Cardenas 4069 (US). COCHABAMBA/SANTA CRUZ: Rio Mizque, Ritter 904 (SGO). SANTA CRUZ: Nr. Mataral, 1500 m, Mar 1911 (fl), Herzog 1869 (L, NY); Saipina, Knize 934, cult, hort. Berol. (B); Samaipata, 1700 m, Rauh 40627, cult. hort. Berol. (B); between Taco Laguna and Elele, 26 Jun 1948 (fl), Scolnik & Luti 778 (K, LIL, P, SI); Vallegrande, Chilon, 25 Mar 1920 (fl), Steinbach 3954 (LIL). Local name. Uturunku. Pereskia diaz-romeroana is still a little known species, and the variability of leaf characters and flower color needs further study. A broad-leaved form {Knize 934) is distinctive in cultivation but for lack of other distinctive characters hardly merits taxonomic status. Ritter (1980, p. 483) reported having observed a yellow- flowered local variety, but neither specimens nor further data or color photographs are available. The striking feature of the succulent ("woody") roots was first observed by Cardenas (1950) but it has since been found in the other Andean species and in some West Indian taxa as well. The epithet is a tribute to Dr. Belisario Diaz Romero, Bolivian naturalist at the beginning of this century.

4. Pereskia weberiana K. Schumann, Gesamtb. Kakt. 762. 1898. Type. Bolivia. Cochabamba: Tunari mountains, May 1892, Kuntze s.n. (lectotype, NY; holotype at B destroyed). Figs. 2A-D, 3E, F, 7F, 10, 11H, 15D, 17E, 18F, I, 19A, 26, 28A-K. Rhodocactus antonianus Backeberg, Descr. Cact. Nov. 3: 13. 1963. Type. "Regio orientalis cor- dillerarum" {cult, in coll. Backeberg). Not preserved, but illustrated in Backeberg, Kakteen­ lexikon 695, fig. 370; 696, fig. 371. 1966. Pereskia antoniana (Backeberg) Rauh, Kakteen an ihren Standorten 89. 1979 (comb, inval., with­ out basionym citation). Slender shrub with basitonous and epitonous branching, forming erect to arching canes, also described as spreading or clambering, 1-3 m high. Trunk to 12 cm diam., sometimes thickened at base (hypocotyl); canes ca. 1 cm thick, with light brown longitudinally fissured papery bark; branches green, with late periderm formation, becoming pale brown; twigs straight or slightly zigzag, 2 mm thick, glabrous, smooth when fresh, ribbed when dry; internodes 5-20 mm long. Roots observed in cultivated specimens only, with conspicuous globular to fusiform swellings of up to 40 cm length and 15 cm diam. Areoles 2-3 mm in diam. on twigs, to ca. 7 mm in diam. on branches and trunk, with rather sparse and short, brownish-gray tomentum between the thickened spine bases; trichomes 0.1-0.3 mm long; longer white hairs (frequent on seedlings) of up to 1.5 mm length sometimes present; areoles producing spines but no brachyblast leaves. Leaves elliptic to narrowly elliptic-lanceolate, 2.5-6 x 1-3 cm, varying greatly in size, and attaining 8 x 4 cm on strong shoots in cultivation; petiole short but distinct, 2 mm long and 1 mm thick (larger in fresh specimens); leafblade ca. 0.7-0.8 mm 74 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 28. Pereskia weberiana. A, floweringtwig . B, flower, l.s. C, flowerbud . D, stem with axillary clusters of flower buds and immature fruit. E, fruit. F, fruit, l.s. G, seed. H, stem with areoles. I, areoles of trunk. J, tuberous roots of cultivated plant grown from a cutting. K, habit. A-C, from cult, hort. Berol. 037-01-77-80 (B); D, from Cardenas 5008 (US); E, F, G from cult. hort. Berol. 258-08- 80-80; H, from Cardenas 5005 (US); I from Cardenas 5006 (US); J from cult. hort. Berol. 046-03- 78-80 (B); K from a photograph by Cardenas (Cardenas, 1967, p. 19). 1986] PERESKIA (CACTACEAE) 75

thick, fleshy; base cuneate to cuneate-attenuate, apex acute; venation pinnate, midrib prominent below; lateral veins 2-4, rather inconspicuous when fresh. Spines (0-)3-5(-7) per areole on twigs, spreading, stiff, subulate, some deflexed or reflexed, (3-)8-13(-17) mm long, 0.2-0.6 mm thick, often angled or flattened at base, with distinct basal thickening of 1—1.5(—2) mm diam., brown or yellowish- brown; spines to 20 per areole on branches and main shoots, 1-2 cm long and to 2.5 mm thick at base; spines often lacking on distal flowering twigs (particularly in cultivation). Flowers terminal and axillary, solitary or in cymose-paniculate inflorescences and in dense axillary clusters of 2-4(-6) partly formed by prolif­ eration from the receptacle. Flowers perigynous, ca. 17-25(-35) mm in diam., lasting 1-2 days; receptacle turbinate or obovate, 3-4 mm long and in diam., terete, smooth; receptacular areoles l-2(-4), with sparse tomentum of trichomes 0.4-1 mm long, white but with dark base, rarely 1-2 longer hairs present, but receptacle appearing essentially naked. Lower bracts 0-4, spreading, the lowermost (cult, specimens observed only) often leaf-like and to 15x7 mm, the others elliptic to ovate or lanceolate, 2-6 x 1-3 mm; upper bracts 4-6, erect, appressed in bud, lanceolate to narrowly or broadly triangular, 1-5 x 0.5-2 mm; sepaloids 2-3, ovate to obovate, 6-8(-l 0) x 2.5-5(-6) mm, greenish-white to greenish-pink; petaloids 4-8, spreading-recurved, obovate to spathulate, 6—10(—18) mm x 4-7 mm; base cuneate, apex rounded, truncate, or mucronate; blade accrescent during anthesis, thin, pink (pale pink to purplish-pink) or white. Stamens ca. 70-90, at first erect and clustering around the style, later somewhat spreading, 4-11 mm long, shorter to nearly as long as the style; filaments white or pale pink (in pink flowers); anthers 0.9-1.0 x 0.3-0.4 mm, yellow. Pollen tricolpate, small, tectum with fine perforations and small spinules. Ovary half inferior, with free ovary roof; locule 1-2 mm in diam., with flat to convex floor and conspicuous septal ridges protruding to ca. lk of the total diameter of the locule; placentae at the base of the ridges along the margin of the ovary floor, hence appearing basal; style 7- 10 mm long, white; stigma lobes 5-6, suberect to spreading, 1-2 mm long, white. Ovules ca. 30, 0.5 mm long. Fruit a juicy hollow berry, depressed subglobular to very broadly obovate, ca. 5-6 mm long and 7 mm in diam. (to 13-17 mm in diam. when fresh in cultivated specimens); bracts deciduous, mature fruit with 1-2 minute areoles and bract scars, otherwise the surface smooth, glossy black; umbilicus depressed, narrow, with or without the crumpled flower remnants and few hairs less than 1 mm long, or naked; locule in the upper half of the fruit, sometimes with a central flat cone formed by the ovary floor; fruit wall fleshy, dark purplish-brown to black. Seeds ca. 20, obovate, 1.8-2.1 mm long, 1.3-1.6 mm broad and 0.7-0.8(-1.0) mm thick, with shallow ribs along the antiraphe, otherwise smooth, black, glossy, micropylar end somewhat prominent, blunt; hilum basal, flat, lunate, with central funicle scar, cream-colored. Seedlings (three weeks old) with hypocotyl ca. 15-20 mm long and 0.7-0.9 mm thick, finely papillose, reddish; cotyledons ovate (narrowly ovate at earlier stages), ca. 8 x 5 mm, conspicuously red below, without nyctinastic movement, remaining ex­ panded at night. Older seedlings (one year) often with basally thickened hypocotyl of ca. 4 mm diam. Root thickenings only developing at later stages. Chromosome number: In = 22; cult. hort. Berol. 037-02-77-80 (B). Distribution (Fig. 26) and phenology. Bolivia, in the Andean valleys of the Rio Beni drainage system, in dry open forests, from 1100 to 1900 m. Cardenas (1967, p. 20) observed it on dry slopes and river banks. Flowering from July to October. In cultivation the fruits mature within a few weeks after flowering, and the nearly mature fruits which are green at first then quickly turn vinaceous and finally black within few days. The fleshy root swellings enable the plant to survive long periods 76 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 of drought during which some or even all distal stem parts may die off to near the rootstock. The roots alone, however, do not seem capable of regenerating meristems able to form new shoots, although detached root swellings survive for months. Specimens examined. BOLIVIA. LA PAZ: Sud Yungas, La Plazuela, 1500 m, Jul 1949 (flowers pink), Cardenas 5005 (US), (flowers white), Cardenas 5006 (US); La Plazuela, Ritter 328 (B, photo; SGO); Espia, 1060 m, 5 Aug 1921 (fl, fr), Rusby 116 (NY, US); Espia, head of Rio Boopi, 1060 m, 27 Aug 1921 (fl), Rusby & White 428 (NY, US). COCHABAMBA: Ayopaya, Rio Yakonko, 1800 m, Oct 1949 (fl white), Cardenas 5007 (US), (fl purple), Cardenas 5008 (US). Local names and uses. Uturunku or cervetano. The fruits are said to be edible. Pereskia weberiana was described by Schumann (1898) as white-flowered, but Kuntze's label reads "white, sometimes pink," as already noted by Vaupel (1925). Backeberg (1963) described Rhodocactus antonianus from a plant cultivated in his collection and, unknowing of its origin, he gave very vague data "Eastern Andes." One can only speculate that it was grown from seeds received from Cardenas. Backeberg (1962) had misidentified the same plant earlier as Pereskia sparsiflora Ritter, whereas he later (Backeberg, 1966) indicated northeastern Peru as the supposed origin of Rhodocactus antonianus. The plant illustrated by Backe­ berg and plants grown under this name and distributed by the Munich Botanical Garden are definitely conspecific with Pereskia weberiana. The plant illustrated in Krainz, Die Kakteen, 1.10. 1967, as P. diaz-romeroana, as well as plants grown under this name in the Jardin Exotique de Monaco are also P. weberiana. The two species are closely related, but P. weberiana lacks the long hairs on the areoles of the receptacle and fruit and the purplish-red stamens typical for P. diaz-rom­ eroana. The epithet is a tribute to Schumann's contemporary, Dr. Frederic Albert Constantin Weber (1830-1903), French physician and author of several publi­ cations on Cactaceae and Agave.

5. Pereskia lychnidiflora De Candolle, Mem. Mus. Hist. Nat. 17: 75, t. 18. 1828. Type. Plate of "Cactus fimbriatus" in Mocino, Fl. Mex. ined. no. 1689, preserved at the Hunt Institute for Botanical Documentation, Pitts­ burgh. A copy of the plate was used by De Candolle to illustrate the species on pi. 18 of the publication cited above. "Legionensis Cactus caule arborescente foliis ovatis acuminatis spinis alternis longissim- is. . . . Nicaraguae montibus . . ." in Mocifio's manuscript of the Flora of Guatemala (no. 13, 4a div. in the Instituto Botanico Cavanilles in Madrid) apparently refers to the same, but an equivalent specimen was not located. Figs. 1A-C, 3A, 5A, C, D, 6B, 7A, 10, 11D, 15A1-C, 17A, 18G, H, 19D, E, 29A-L, 30. Rhodocactus lychnidiflorus (De Candolle) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Pereskia opuntiiflora De Candolle, Mem. Mus. Hist. Nat. 17: 76, t. 19. 1828. Type. Plate of "Cactus opuntiaeflorus" in Mocino, Fl. Mex. ined. no. 1842, preserved at the Hunt Institute for Botanical Documentation, Pittsburgh. "Matiari Cactus caule arborescente, foliis ovatis retusis, spinis duplo longioribus. .. Nicaraguensis Provinciae sepibus" in Mocifio's manuscript of the Flora of Guatemala (no. 13, 4a div. in the library of the Instituto Botanico Cavanilles in Madrid) evidently refers to the same. Equivalent specimen. Nueva Espafia, Herb. Pavon (G). The specimen is labeled "Cactus macrochanthos" and is the same branch that was used for the Fl. Mex. ined. drawing (mirror image!). Pereskiopsis opuntiiflora (De Candolle) Britton & Rose, Smithsonian Misc. Collect. 50: 332. 1907. Opuntia golziana K. Schumann, Gesamtb. Kakt. 654. 1898 (nom. illeg. based on Pereskia opun­ tiiflora). 1986] PERESKIA (CACTACEAE) 77

Pereskia pititache Karwinski ex Pfeiffer, Enum. diagn. Cact. 176. 1837. Type. Mexico. (Not preserved; description based on a young plant grown from seed collected by Karwinski.) Opuntia pititache (Pfeiffer) F. A. C. Weber, Bull. Mus. Hist. Nat. (Paris) 4: 166. 1898. Pereskiopsis pititache (Pfeiffer) Britton & Rose, Smithsonian Misc. Collect. 50: 332. 1907. Pereskia calandriniifolia Hort. Berol. ex Salm-Dyck, Cact. hort. Dyck. ed. 2, 252. 1850 ("P. calandriniaefolia"). Type. Not preserved. Pereskia nicoyana F. A. C. Weber, Bull. Mus. Hist. Nat. (Paris) 8: 468. 1902. Type. Costa Rica. Guanacaste: Nr. Nicoya (no specimen cited); description based on living material collected by A. Tonduz and sent by Biolley in 1900. Neotype designation. Nicoya, May 1900 (fl), Tonduz 14001 (neotype, US) (possibly a duplicate of the type collection). Rhodocactus nicoyanus (F. A. C. Weber) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Pereskiopsis autumnalis Eichlam, Monatsschr. Kakteenk. 19: 22. 1909. Type. Guatemala. Pro­ greso: San Augustin, Eichlam s.n. (holotype, B destroyed). A drawing by Weingart of a fruit sent by Eichlam is at NY (copy at B). Pereskia autumnalis (Eichlam) Rose, Contr. U.S. Natl. Herb. 12: 399. 1909. Rhodocactus autumnalis (Eichlam) F. Knuth in Backeberg & Knuth, Kaktus-ABC 96. 1935. Pereskia conzattii Britton & Rose, Cactaceae 1: 24. 1919. Type. Mexico. Oaxaca: Salina Cruz, 23 Jan (Feb?) 1913 (fr), Conzatti s. n. (syntypes, US, NY); Tehuantepec, Apr 1913 (fr), Conzatti s.n. (syntype, US). Rhodocactus conzattii (Britton & Rose) Backeberg, Cactaceae 1: 118. 1958.

Tree up to 10 m or rarely to more than 15 m tall; trunk to 30-40 cm in diam.; young trees monopodial, with subverticillate primary, horizontal to diffuse, di- chotomous to trichotomous branching (Figs. 1 A, 5 A); older trees in open condition with short trunk of 1-2 m, and broad crown, resembling an apple tree, or, in dense stands, trunk 2-4 m tall, with crown narrower but broadened towards the top. Twigs stout, 4-5 mm thick, olive-green to reddish-brown when fresh, var­ iously brown when dry, lacking stomata, periderm formation early; main shoots thicker, twigs and branches with smooth periderm, trunk of younger trees with smooth leathery bark ca. 1 mm thick, brownish-gray, with green cortex below, containing masses of fusiform, stinging sclereids; bark of old trees rough, longi­ tudinally fissured mainly above and below the accrescent areoles. Roots imper­ fectly known; young plants with taproot, older trees with (additional?) thick roots spreading near the soil surface. Areoles 3-5 mm in diam. on young twigs, some­ times forming protuberances of ca. 5 mm length and diameter on slow growing distal twigs with seasonal brachyblast leaf formation, accrescent to 15-20 mm diam. and 10-30 mm length on the trunk; tomentum pale brownish-gray, tri­ chomes ca. 1.5 mm long, longer on juvenile plants; few or to ca. 20, tortuous, coarse, brittle and eventually deciduous hairs 5-20 mm long (mainly on new shoots and particularly on seedling plants) also present. Brachyblast leaves and/ or spines produced. Leaves of long shoots often smaller than brachyblast leaves and usually narrower, elliptic to obovate, acute; brachyblast leaves 1—2(—3) per areole on twigs and branches, rarely also on older branches and trunk, variable in shape and size; foliage of horizontal branches showing conspicuous epitony and anisophylly (Figs. 5C, 29A); leaves obovate to nearly orbicular, or elliptic to narrowly elliptic-obovate, or oblanceolate, (1—)2—8(—10) x l-4(-5) cm; petiole inconspicuous or l-3(-4) mm long and 1—2(—3) mm thick; blade fleshy, 0.7-1 mm thick when fresh, green, smooth, mature leaves drying leathery; base shortly attenuate, or cuneate, or rounded-subcordate in suborbicular fresh leaves; apex shortly acute, acute, rounded or slightly emarginate, or obtuse with a triangular point, usually acute in auxoblast leaves; venation pseudopalmate, conspicuous in dried leaves only, not prominent in fresh leaves, lateral veins 2-6, arising from the lower Vl0 to V5 of the midvein, diverging at angles of 5-20(-30)°. Spines often lacking on distal flowering twigs but present on lower branches, often one per areole on twigs of mature plants, or 1-3 on twigs, more numerous on juvenile [VOL. 41 78 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN

T 1 mm

VereskicL LychnldifLora

FIG. 29. Pereskia lychnidiflora. A, twig of juvenile plant. B, stem with brachyblast leaves. C, average leaves from mature plant. D, main shoot with small auxoblast leaves. E, flower. F, flower, l.s. G, flower bud with lower and upper bracts. H, fruit. I, fruit, l.s., with detail of sclerenchymatic fibers. J, seeds. K, areole from trunk. L, habit, (a) of a young plant in open habitat, (b) of a tall tree in dry forest, (c) of a tree in open pasture, (d) of a tree in dry forest. A from Leuenberger & Schiers 2508 cultivated seedling (B); C, H, I, J, Lc from Leuenberger & Schiers 2508 (B); B from Cook & 1986] PERESKIA (CACTACEAE) 79 plants, subulate when young, acicular to subulate when mature, (l-)2-7(-9) cm long and ca. 1 mm thick, shorter and thinner on young plants; older areoles on branches with 0-3(-4) spines 2-10 cm long; areoles of trunk with (5-)10-30(-40) spreading spines of very unequal length, 3-12(-16) cm long and 1-2.5 mm thick; spines yellowish or reddish in new growth, with reddish-brown tip, becoming gray with dark tip. Flowers single, terminal or on short lateral persistent twigs, or on short pedicels arising from subterminal and lateral areoles, to ca. 6 cm diam. when fully expanded; flower buds 15-20 mm in diam. with spreading bracts and numerous imbricate sepaloids; pedicels 5-10(-30) mm long, with one to several leafy, acute bracts, gradually broadened into the turbinate receptacle; receptacle 10-15 mm long and in diam. (ca. 20 mm in diam. when fresh), bearing ca. 10- 20 spirally arranged, sessile, fleshy bracts; receptacular areoles transverse-linear, ca. 3 mm long, with sparse tomentum of several rows of short, white to pale brown trichomes ca. 1 mm long, very rarely with a short spine; lower receptacular bracts elliptic, 4-20 mm long, with narrow base, acute; upper bracts spreading, elliptic to suborbicular or semi-orbicular, 4-8 mm long and broad, with broad base; sepaloids ca. 20, erect, imbricate, appressed to the bud, broadly ovate- triangular, the outer ones ca. 3 x 3 mm, the inner ones to ca. 6 x 6 mm, green to reddish-green, conspicuously parallel-veined, intergrading with the petaloids; petaloids 10-20, obovate, ca. 20 x 8-15 mm, fleshy at base but with delicate blade, usually emarginate to deeply cleft at apex, with laciniate-dentate to erose margin and palmate-flabellate venation, bright yellowish-orange at anthesis but carrot- to brick-red in bud and in closing flowers. Stamens numerous, ca. 250 or more, (5—)9—11 mm long, filaments cream, longer than the style; anthers 1.3- 1.6 x 0.5-0.6 mm, pale yellow; pollen 6-9(-12)-colpate, medium-sized, tectum with variable perforations and medium to large spinules. Ovary half superior, ca. 9 mm in diam., with broadly conical, free ovary roof ("style base"), carpels 10- 18, connate, with their involute portion adnate to the conical floral axis; style ca. 5 mm long and 2-3 mm thick, often longitudinally ribbed, stigma lobes 10-18, nearly erect, 4-5 mm long, equalling or a few millimeters below the level of the anthers, usually not visible in open flowers; placentation basal-"axile"; ovules numerous in each pocket-like locule, ca. 0.6 mm long. Fruit broadly pyriform to spherical, 25-40 mm in diam. (3-5 cm when fresh), greenish-yellow (yellow to reddish according to Bravo-Hollis, 1978), nearly smooth; lower bracts deciduous at maturity, upper bracts persistent or deciduous, areoles transverse-linear, ca. 5 mm long, naked or nearly so; uppermost bracts and outer sepaloids connivent over the narrow umbilicus; fruit wall somewhat leathery, fleshy inside; locule forming a moat-like cavity around a central, peg-like columella, fruit wall and columella with large mucilage cavities; locule filled by the seeds and numerous acicular sclereids formed in the ovary wall. Seeds numerous, to 100-150, obovate, 3-3.5 mm long, 2-2.5 mm broad and ca. 1.5 mm thick, smooth, black, glossy; hilum subbasal, semicircular or lunate, low or protruding, cream. Seedlings (three weeks old) with hypocotyl ca. 20-26 mm long and 1.2 mm thick, olive to reddish; cotyledons elliptic, ca. 20 x 7 mm, slow-growing. Chromosome number: In = 22; Leuenberger & Schiers 2508, cult. hort. Berol. 003-04-78-10 (B). Distribution (Fig. 30) and phenology. Southern Mexico to Costa Rica. In low-

Collins s.n., photograph (NY); D, from Leuenberger & Schiers 2718 (B); E, G from Leuenberger & Schiers 2513 (B); F, K from Leuenberger & Schiers 2533 (B); La from Leuenberger & Schiers 2515a, photograph (B); Lb from a photograph by McDougall; Ld, Leuenberger & Schiers 2533a, photograph (B). 80 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 30. Distribution of Pereskia lychnidiflora. land dry forest, "selva baja caducifolia," on dry plains and hillsides in the Pacific lowland and inland in dry valleys, from sea level to 1000 m; flowering mainly from June to October; fruits mature from September to January; leaves deciduous or partly deciduous ca. from January/February to April. The disjunct distribution of this species was commented upon by the author earlier (Leuenberger, 1980), after confirmation of a single record from Guerrero. The locality data for Costa Rica are not very precise, and during a trip by the author to the type locality of Pereskia nicoyana ("around houses at Nicoya") it was not possible to locate the plant. The localities "Puntarenas" and "Punta Arenas" may refer to place names or to the whole region, and unfortunately no recent collections are available to clarify this. Pereskia nicoyana is mentioned by Pittier (1908) as "mateares" in his list of useful plants of Costa Rica. The spiny trunk illustrated without identification in the caption of plate 2 in his work cer­ tainly belongs to this species growing in a coastal forest at Salinas, Guanacaste. Living twigs of P. lychnidiflora collected in NW Guanacaste (Gomez, pers. comm.) confirm this record.

Representative specimens. MEXICO. GUERRERO: 9 km SE of San Luis San Pedro, 2 km on side rd. to Los Tarros, 9 Mar 1980 (st), Leuenberger & Schiers 2718 (B, MEXU); 4 mi N of San Luis de La Loma, 30 Jul 1952 (fl), Rowell 3140A (MICH). OAXACA: 3-4 mi E of Tehuantepec, 3 Jul 1962 (fl), Boke 25 (GH, RSA, UC); isthmus of Tehuantepec, Jun 1965 (fl), Bravo 35 (MEXU);'Salina Cruz beach, 18 Oct 1919 (fr), Conzatti 3667 (NY, US); between Totolapan and Tehuantepec, 23 Dec 1966 (st), Delgadillo s.n. (MEXU); Tequisistlan, 15 Sep 1975 (st), Hernandez 89 (MEXU);'Tequisistlan, 1838, Karwinski s.n. (G); 7 km E of Salina Cruz, beach at La Ventosa, 50 m, 3 Jul 1959 (fl) King 1288 (MICH, NY, UC, US); 16 km NE of Tehuantepec, 50 m, 7 Jul 1959 (st), King 1400 (MICH, 1986] PERESKIA (CACTACEAE) 81

NY, UC, US); 3 km W of Ixtepec, 50 m, 9 Jul 1959 (fl), King 1476 (MICH, NY, UC, US); Salina Cruz, 30 Jan 1960 (fr), King 2461 (GH, POM); 4 km N of La Ventosa, 13 Oct 1978 (st), Leuenberger & Schiers 2504 (B, MEXU); 12 km S of Tehuantepec on side rd. to Pishishe, ca. 30 m, 19 Oct 1978 (fr), Leuenberger & Schiers 2521 (B, MEXU); 4 km W of Tehuantepec, ca. 50 m, 20 Oct 1978 (fr), Leuenberger & Schiers 2531 (B, MEXU); 43 km NW of Tehuantepec, ca. 300 m, 20 Oct 1978 (fl), Leuenberger & Schiers 2533 (B, MEXU); Tehuantepec, 1841-43, Liebmann 11157 (C); Tehuantepec, Cerro Calderona, 27 May 1972 (fl), MacDougall H 483 (NY); Tehuantepec, Jul 1936 (fl), Matuda 2296 (MEXU); Juchitan, 26 Jan 1956 (st), Matuda 38571 (MEXU); 0.5 mi W of Bahia Ventosa, 20 Nov 1959 (st), Moran 7740 (UC); San Geronimo (Ixtepec), May 1923 (fr), Purpus s.n. (NY, US); Tehuantepec, km 3 (20) on rd. to Oaxaca, 17 Sep 1967 (fl), Pennington & Sarukhan 9213 (A, K, MEXU, NY). GUATEMALA. BAJA VERAPAZ: Between Salama and Rabinal, ca. 1000 m, ca. 1905, Cook & Collins 7459 & 7853 (photos, US); between Salama and Purula, 4 Jun 1904 (fl), Cook & Doyle 291 (US). CHIQUIMULA: Chiquimula, 250 m, 19 Sep 1946 (fr), Clover 9134 (MICH); 2 mi W of Chiquimula, 260 m, 1 Oct 1946 (fr), Clover 9228 (MICH). EL PROGRESO: Between El Rancho and Tulemaje, 400 m, 10 Nov 1946 (fr), Clover 9493 (MICH); El Rancho, 9 Apr 1902 (st), Cook 806 (US); El Rancho, 300 m, 8 Jun 1909 (fl), Deam 6246 (E, MICH, MO, NY, US); nr. San Augustin Acasaguastlan, 6 Mar 1970 (fl), Harmon 2574 (MO); El Rancho, banks of Rio Motagua, 350 m, 28 Feb (Dec?), 1907 (fr), Kellermann 7011 & 7014 (K, MO, NY, US); between El Progreso and El Rancho, ca. 3 km above bridge over Rio Motagua, 15 Oct 1978 (st), Leuenberger & Schiers 2507 (B, MEXU); nr. San Cristobal Acasaguastlan, ca. 350 m, 16 Oct 1978 (fl), Leuenberger & Schiers 2513 (B, MEXU); below El Rancho and beyond rd. to El Augustin, 350 m, 7 Nov 1959 (fr), Moore & Cetto 8210 (GH). JUTIAPA: 3 km ENE of Jutiapa, NW of El Tablon, 930-960 m, 31 Dec 1975 (st), litis & Gillespie G 28 (F); vie. Jutiapa, 850 m, 24 Oct-5 Nov 1940 (st), Standley 75634 {¥). ZACAPA: Gualan, ca. 220 m, 15 Jun 1909 (fl), Deam 6269 (US); Gualan, 122 m, 28 Dec 1902/05 (st), Kellermann 5726 (US); Gualan, 2 Jan 1906 (fr), Kellermann 5020 (MEXU, RSA, US); 3 km W of Teculutan, ca. 350 m, 16 Oct 1978 (fr), Leuenberger & Schiers 2508 (B, MEXU); 3 km W of Teculutan, 16 Oct 1978 (fl), Leuenberger & Schiers 2509 (B); 2 mi E of Gualan, Los Limones, 26 Jan 1927 (fr), Record & Kuylen G. 122 (US); between Rio Hondo and Santa Cruz, 200 m, 11 Oct 1940 (fr), Standley 74123 (F); nr. Estanzuela, 200 m, 5 Oct 1939 (fl), Steyermark 29131 (F). HONDURAS. CHOLUTECA: 1 km N of Choluteca, 200 m, 2 Jun 1965 (fl), Lent 593 (MICH); 4 km N of Choluteca, 7 Dec 1971 (fl), Harmon & Fuentes 6356 (MO); 8 km SE of Choluteca, 200 m, 2 Jun 1965 (fl), Lent 592 (F, NY); 9 km N of Pespire, 170 m, 5 Aug 1962 (fl), Webster et al. 12778 (F, GH, MO). COMAYAGUA: Vic. Comayagua, 600 m, 12-23 Mar 1947 (st), Standley & Chacon 5451 {¥). MORAZAN: Miramonte, D.C., 2 Aug 1969 (fl), Barkley 39448 (GH); Pedregal, drainage of Rio Yeguare, 900 m, 23 Jun 1948 (fl), Glassman 1713 (F, NY, UC); Zamorano, 800 m, 6 Oct 1943 (fl), Rodriguez 1152 (F, LIL); vie. El Zamorano, 780-900 m, 26 Nov 1946-9 Jan 1947 (st), Standley 156 {¥). OCOTEPEQUE: Vic. Ocotepeque, along Marchala river, 800 m, 30 Aug 1968 (fl), Molina 22484 (F, NY). VALLE: 5 km N of Jicaro Galan, 120 m, 3 Oct 1946 (fl), Williams & Molina 12671 (GH, MICH, MO); 2 km S of San Lorenzo, 27 Sep 1973 (fl), Hazlett 908 (MO). EL SALVADOR. LA LIBERTAD: La Libertad, 4 m, 15 Sep 1935 (fr), West 3549 (UC). SAN MIGUEL: Hacienda San Antonio, Jan 1924 (fr), Calderon 2117 (US). SAN SALVADOR: San Salvador, Aug 1912 (fl), Calderon 1164 (GH, NY, US). SONSONATE: Acajutla, Jul 1923 (fl), Calderon 1757 (NY, US). NICARAGUA. WITHOUT LOCALITY: 7 Jul 1927 (fl), Chaves 281 (US). Vic. Managua, Jul 1932 (fl), Gamier 900 (MICH); Esteli, Cerro Quiabu, Atwood & Neill AN 292 (MSC, n.v.). COSTA RICA. WITHOUT LOCALITY: Alfaros.n. (US). "Puntarenas," Feb 1909 (fl), Brade 2553 (BR); "Punta Arenas," 1912, Werckle s.n. (US). Local names and uses. Mexico: guititache, guichitache, guitache, patilon, cruz del matrimonio, arbol del matrimonio (Oaxaca); cuncu, cuncu mar in (Guerrero). Guatemala: manzanote, matial. Honduras: mateado, amatilla. El Salvador: matial, matiare. Nicaragua: matiari, mateare. Costa Rica: mateares. According to Eichlam (1909) the spines were used as needles in Guatemala. Pittier (1908) noted that it is sometimes used for planting hedges in Costa Rica. The leaves are eaten by goats when other food is rare. In El Salvador it is frequently planted in hedges, but Standley and Williams (1962, p. 225) note that the sclereids (erroneously called glochids) are irritating and dangerous for man and domestic . The taxonomy of this species has received little attention compared with the detailed morphological and anatomical studies by Bailey (1963b), Boke (1963b), and Crespo (1973), although Boke (1963a) gave a brief historical account of Pereskia pititache and its synonyms. Bravo-Hollis (1978) recognized the older 82 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 name P. lychnidiflora as correct for the plant occurring in Mexico, but she main­ tained P. autumnalis from Guatemala as distinct, admitting that it may just be a geographical form of the former. Examination of flowering trees in the field in 1978 both in Guatemala and Mexico made it very clear to the author that there is only one species and that the characters used in the keys by Britton and Rose (1919) and Bravo-Hollis (1978), petaloids toothed vs. fimbriate, are too variable and unspecific. The petaloids of P. lychnidiflora were very adequately described by De Candolle (1828b) as "dentes ou franges a leur sommet," i.e., "dentate or fringed at the tip." The comparison of the two plates of the Flora Mexicana ined. of Mocino, on which P. lychnidiflora and P. opuntiiflora are based, would not suggest the inclu­ sion of P. opuntiiflora in the synonymy of P. lychnidiflora due to the different petaloids. While the plate of P. opuntiiflora very accurately depicts the habit of a flowering twig of P. lychnidiflora as interpreted here, the flowers are likely to have been drawn from a withered specimen and wrongly reconstructed, or the spreading bracts were later mistaken for petaloids and adapted and colored ac­ cordingly by the artist. This interpretation is supported by the comparison with a photograph published by Bravo-Hollis (1978, fig. 71), as compared with the plate of P. opuntiiflora. The strongest evidence, however, comes from a specimen located at the Geneva herbarium, labelled as ''Cactus macrochanthos" from Nueva Esparia, Herb. Pavon, and annotated by J. N. Rose in 1912 as "Pereskia lych- niflora." According to McVaugh (1977) and Bernardi (1977), specimens from the Sesse and Mocino collection were sold by Pavon with his own labels between 1814 and 1828. This specimen, easily identified as P. lychnidiflora by comparison with dried material of this species, is a nearly perfect mirror image of the plate of P. opuntiiflora and likely to be the same twig that was used in preparation of the painting by Cerda, the artist accompanying the expedition of Mocino to Central America. In any case it demonstrates the link between the plate of P. opuntiiflora and dried specimens of P. lychnidiflora. Schumann (1898) and Britton and Rose (1919) misidentified the somewhat blurred structures on the receptacle of P. opuntiiflora in the De Candolle copy of the painting as fascicles of glochids, an error possibly favored by inaccurate copying of the plate. The original plate, which is among the ones rediscovered recently (McVaugh, 1982), and of which a color slide was kindly made available to the author by the Hunt Institute for Botanical Documentation, clearly shows bracts rather than areoles with glochids on the flower receptacle. Although no specimen belonging to the plate of P. lychnidiflora (De Candolle, 1828b, pi. 18) could be located, this name poses no interpretation problem thanks to the direct comparison of fresh flowering specimens matching the plate wjth dried material prepared from the same collection {Leuenberger & Schiers 2513 & 2533). De Candolle (1828b) incorrectly assumed all the plants illustrated in the Flora Mexicana ined. of Mocino to have originated from Mexico, and this error has ever since been handed down by authors of local floras and monographs. However, as McVaugh (1977) showed and as is further discussed in this paper under P. zinniiflora, some of Mocifio's plates refer to plants of Central American or even West Indian origin. From the descriptions in the Flora of Guatemala manuscript of Mocino kept at the Instituto Botanico Cavanilles in Madrid the plants later described as P. lychnidiflora and P. opuntiiflora were in fact from Nicaragua, where the royal expedition stayed from May to December 1797 and collected the two plants in August and October respectively. The ex­ pedition party may have seen this cactus earlier around Tehuantepec and on the 1986] PERESKIA (CACTACEAE) 83 way to Nicaragua but possibly only in the non-flowering condition, depending on the season. No recent and living material from Costa Rica could be studied, but from the old herbarium material and the original descriptions it is beyond any doubt that P. nicoyana is a synonym of P. lychnidiflora.

6. Pereskia aureiflora Ritter, Kakteen in Siidamerika 1: 22, fig. 5. 1979. Type. Brazil: Minas Gerais, Itaobim, Ritter 1413 (U, apparently not deposited). Figs. 6J, 15Ah, 18D, E, 19H, I, 31A-I, 48. Small tree or shrub up to 6 m tall, trunk up to 20 cm in diam.; bark brownish- gray, rough, branches erect or arching; distal twigs ca. 2-3 mm thick. Areoles 2- 3 mm in diam. on twigs, with dense, short, grayish-brown tomentum, producing spines and 1-3 brachyblast leaves, accrescent on trunk to ca. 6 mm in diam. Leaves extremely variable in shape and size, obovate to elliptic-suborbicular on distal twigs under shady conditions, elliptic-lanceolate and thick-fleshy in exposed habitat, lanceolate on strong main shoots; auxoblast leaves up to 11 cm long and 2-3 cm broad, with distinct petiole and attenuate or cuneate base; brachyblast leaves 4-10 cm long and (1.5—)2.5—5 cm broad, petiole 2-4 mm long, blade plane or somewhat boat-shaped, 0.5-1 mm thick, base broadly cuneate, apex acute; venation pinnate, midrib prominent below, lateral veins 3-5(-7), ascending, di­ verging at angles of 40-60°. Spines 0-3 on twigs, 4-5 on young longshoots, 1-3 cm long, up to 35 on trunk, 1-3 cm long, 0.4-0.8 mm thick and up to 2 mm in diameter at the conically thickened base. Flowers solitary, terminal or on short lateral twigs, ca. 4 cm in diam.; pedicels 3-13 mm long, with 0-2 leafy bracts; receptacle 4-5 mm in diam., turbinate, bracteate; areoles with very sparse to­ mentum of hairs up to 1 mm long; lower bracts ca. 5-8, lanceolate or elliptic- lanceolate, short petiolate, spreading, 7-23 x 3-10 mm, thick and fleshy, green, sometimes with reddish margin and apex; base attenuate, apex acute; upper bracts 5-8, erect, triangular, 1-2 mm long and broad, fleshy, green, keeled; sepaloids ca. three, obovate, ca. 11 x 5 mm; petaloids ca. 10-12, spreading to reflexed 15- 20 x 8-12 mm, obovate, yellow, narrowed at base; apex obtuse; stamens nu­ merous (ca. 100 or more), 5-10 mm long; filaments pale yellow; anthers 1-1.6 x 0.5-0.7 mm, golden-yellow; pollen 12-15-colpate, large, tectum densely perforate and with small spinules; ovary ovate, half inferior, adnate to the receptacular cup with its base only, but its free part surrounded by the filament bases and the rim of the receptacular cup; locule with flat floor and "basal" placentae. Fruit a globular berry, 12-15(-20) mm in diam., reddish-green to chocolate-colored at maturity; pedicel distinct; areoles transverse linear, naked or with few short hairs; lower bracts leafy but usually deciduous at maturity, leaving a smooth, pale gray leaf scar which is more conspicuous than the areole; umbilicus narrow, filled by the connivent upper bracts and the crumpled flower remnants, or to ca. 4 mm diam.; fruit wall ca. 4 mm thick, fleshy; roof of the ovary to ca. 5-6 mm thick, with mucilaginous tissue; locule cup-shaped, ca. 6-7 mm in diam. Seeds few, often only 1-3, obovate, (4-)5-5.5 mm long, 4.2-5.1 mm broad and 2.5-3.0 mm thick, often slightly concave or flattened on one side, smooth, black, glossy; hilum subbasal, reniform to semicircular, whitish; micropylar end rounded. Distribution (Fig. 48) and phenology. Northeastern Minas Gerais and southern Bahia, Brazil, at ca. 300 to 700 m, in the transition zone of caatinga and cerrado, also in disturbed vegetation and planted for hedges, not frequent, flowering in October and November, fruiting in March and April. [VOL. 41 84 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN

3 mm

Pereskia aureiflora

FIG. 31. Pereskia aureiflora. A, floweringtwig . B, flower(closed) , from boiled herbarium material, l.s. C, twig with immature fruit. D, fruit. E, fruit, l.s. F, seeds. G, young main shoot with auxoblast leaves. H, brachyblast leaves. I, areole of trunk. A from photograph of Ritter (Ritter 1979, p. 278, fig. 5); B from Duarte 10534 (B); C from Leuenberger & Martinelli 3055, photograph (B); D-I from Leuenberger & Martinelli 3054 (B). 1986] PERESKIA (CACTACEAE) 85

Specimens examined. BRAZIL. BAHIA: Mun. de Rio de Contas, 13 km E of Vila do Rio de Contas on rd. to Marcolino Moura, ca. 700 m, 25 Mar 1977 (fr), Harley 19992 (B, K). MINAS GERAIS: Aracuai, 300 m, 12 Nov 1981 (fl), Pinto 398/81 (B, HRB); 4 km E of Itaobim on rd. to Jequitinhonha, 9 Mar 1977 (fr), Shepherd et al. 4422 (UEC); 8 km W of Itaobim, 2 km S of rd. Itaobim-Aracuai, ca. 500 m, 9 Apr 1983 (st), Leuenberger & Martinelli 3056 (B, CEPEC, RB); 9 km W of Itaobim, 9 Apr 1983 (fr), Leuenberger & Martinelli 3055 (B, CEPEC, RB); 15 km SW of Itinga, on rd. Aracuai-Itaobim, Taquaral, Lavra da Pinheira, ca. 400 m, 8 Apr 1983 (fr), Leuenberger & Martinelli 3054 (B, CEPEC, K, MO, NY, RB, SI, W); between Virgem da Lapa and Aracuai, 18 Oct 1967 (fl), Duarte 10534 (B, RB). Local names and uses. Facho, or ora pro nobis de mata (Minas Gerais). Some­ times used for fences. Pereskia aureiflora was discovered by Ritter, probably in 1964, and described and illustrated in 1979, but the type specimen, said to be deposited at U, is missing and no duplicate material was located at SGO, where some other fragmentary material of the Ritter collection of Cactaceae is extant. This species is still incom­ pletely known, and I have seen only fruiting material in the field. The variation in leaf shape under different conditions within the same area is remarkable. G. R. Pinto (pers. comm.) believes that another yellow-flowered species exists in Bahia. This species is remarkably similar to P. guamacho in some vegetative features and the yellow flower color but easily distinguished by the pedicellate flowers and much larger seeds.

7. Pereskia guamacho F. A. C. Weber in Bois, Diet. hort. 938. 1898. Type. "Orinoco basin." No specimen cited. Neotype designation. Venezuela. Bolivar: Ciudad Bolivar and vie. on the Orinoco, 70 m, Feb-Mar 1921 (fl), L. Bailey & E. Bailey 1351 (US). Figs. 7B, 15Af, 32A-L, 33. Rhodocactus guamacho (F. A. C. Weber) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Pereskia colombiana Britton & Rose, Cactaceae 1: 17, fig. 11. 1919. Type. Colombia. Magdalena: Santa Marta, Bonda, 50 m, 5 Apr 1898/99 (fl), H. H. Smith 1886 (lectotype, NY; isotypes, BM, BR, E, F, G, GH, L, LE, MICH, MO, P, S, U, UC, US, VT). Rhodocactus colombianus (Britton & Rose) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Tree, 4-8 m tall, or shrub; trunk usually short, l(-2) m high and to 25(-40) cm thick; young trees monopodial, with subverticillate, diffuse branching, sometimes many-stemmed by formation of lateral erect long shoots; bark brownish-gray on trunk, smooth (?), branches with pale brown to reddish-brown, smooth bark with few whitish lenticels; twigs 2-3(-4) mm thick, green to reddish when fresh, pale brown after periderm formation. Wood yellowish; pith conspicuous, 1.2-2 mm in diam. in branches of 1 cm thickness, transverse-laminate. Areoles 2-4 mm in diam. on twigs, 5-7 mm in diam. on branches, forming knob-like spur shoots 3- 5 mm long on older flowering branches and branchlets, accrescent on trunk to ca. 1 cm diam.; tomentum of the areoles pale gray to pale brown, dense and short, consisting of uniseriate, multicellular trichomes l(-2) mm long; additional longer hairs usually in new growth only; areoles producing spines and brachyblast leaves. Leaves of long shoot not differing from brachyblast leaves; brachyblast leaves 1- 3(_4) per areole, fascicled in the upper portion of the areole. Leaves variable in shape and size, narrowly to broadly obovate or elliptic, 2-7(-9) x 1-6 cm, average leaves ca. 4 x 2.5 cm; petiole 1—4(—5) mm long and 1 mm broad, appearing shorter and thicker and often indistinct when fresh; leafblade 1-1.5 mm thick when fresh, fleshy, base attenuate; apex shortly acute to acute-acuminate or obtuse with a short triangular point; venation pseudopalmate to pinnate; midrib thick­ ened but not markedly prominent below when fresh; lateral veins (2-)3-4(-5), diverging at angles of (15-)20-40(-45)°, often inconspicuous in fresh leaves. Spines on young twigs 0-2 per areole, porrect, subulate, 5-20 mm long and 0.5-1.0 mm 86 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 32. Pereskia guamacho. A, leafless branch with flower bud. B, flower and flower buds. C flower, basal view, exposing receptacular bracts. D, flower, l.s. E, fruit. F, fruit, l.s. G, seeds. H stem with brachyblast leaves in the lower part. I, J, leaf variation. K, areole of trunk. L, habit A from Manara 88883 (VEN); B from Pittier s.n. (photo, NY); C, D from Pittier 6179 (NY)- E F G from Breteler 4553 (U); H from cult. hort. Berol. 001-16-74-70 (B); I from Alston 8547a (BM)- J from Aristeguieta et al. 6744 (VEN); K, L from L. Bailey & E. Bailey 1351, including a photograph (US). 1986] PERESKIA (CACTACEAE) 87

FIG. 33. Distribution of Pereskia guamacho. thick, with slightly bulbous thickening at the base, reddish brown to black, bright dark red at base when young (fresh); spines often lacking on flowering twigs and spur shoots; spines 0-5 on branches, spreading, (5-)10-30(-40) mm long, brown to black, becoming gray with age; spines on trunk 50-60 or more, subulate, 3-5 cm long (said to be to 10 cm long, but no specimens seen), 1-1.7 mm thick. Flowers axillary on young and older areoles (spur shoots), solitary or fascicled (2- 3), sessile or nearly so, appearing before the leaves and rapidly falling unless pollinated, flower buds with subcylindrical receptacle ca. 5 mm in diam. including the dense tomentum and appressed receptacular bracts. Flowers perigynous, ro­ tate, 2-5(-6) cm in diam. when fully expanded; receptacle ca. 5 mm in diam., turbinate, receptacular areoles densely silky-tomentose with several rows of tri­ chomes 1-3 mm long, usually masking the receptacle in bud; hairs white to pale brown; receptacular bracts 15-28, linear-lanceolate, very variable in size, the lower and uppermost 1.5-3 x 1 mm, the middle ones 3-5 x 1-2 mm, rarely leaf-like and to 18 x 5 mm, with acute or rounded apex; sepaloids ca. 3-5, obovate, 10- 15 mm long, apex acute or obtuse with a point, blade yellowish or flushed with red below; petaloids 10-12, obovate to narrowly obovate, to 25 x 13 mm, with rounded to truncate-emarginate apex; blade yellow, thin and short-lived. Stamens 200-300, spreading, 9-15 mm long, filaments yellow; anthers 1.0-1.4 x 0.3-0.4 mm, yellow, with darker connective. Pollen (9-) 12-15-colpate, medium-sized, tectum with perforations and spinules of ca. 1 ixm length. Ovary half inferior, with free roof narrowed into the conical style base; locule ca. 2 mm in diam., with shallow floor and roof with conspicuous septal ridges; placentae parietal, in the middle or lower third of the locule at the base of the septal ridges; style ca. 10 mm long; stigma lobes 5-6, ca. 2-3 mm long. Ovules numerous, ca. 0.25 mm long. Fruit globular to obovate, ca. 15-20 mm in diam., green, fleshy, main body formed by the middle portion of the receptacle, hence bearing only to ca. 10 bracts; bracts lanceolate, to ca. 12 x 4 mm, spreading, persistent, or deciduous at maturity; fruit smooth except for the transverse elliptic leaf scars and the sparsely tomentose inconspicuous areoles; umbilicus narrow, filled by the ca. 10 connivent upper bracts surrounding the crumpled, usually persistent flower rem­ nant; fruit wall 2-4 mm thick; locule hollow. Seeds ca. 10-20 or more, reniform, 88 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

3.7-4.1 mm long, 2.9-3.4 mm broad and 1.5-1.6 mm thick, strongly flattened and with a peripheral ridge, smooth, black, glossy; hilum low to rather prominent, cream, with central, depressed funicle scar; micropylar end rounded. Seedlings not observed. Distribution (Fig. 33) and phenology. Colombia, Venezuela, uncertain in south­ ern Dutch Antilles (Bonaire). From sea level to 800 m, rarely to 1800 m, in dry coastal formations, deciduous seasonal woods, "espinares," "selvas claras," "bos- que caducifolio," flowering mainly from March to May in the leafless condition, at the end of the dry season; fruiting from July to September. The record for Bonaire seems doubtful, as Boldingh (1913) did not mention it and because it was not confirmed by Wagenaar Hummelinck (1938), who only mentioned the species as introduced in Curacao but not for Bonaire. The species is apparently common in the flat land and in the coastal ranges of northern Colombia and Venezuela and very conspicuous during the flowering season, where the golden yellow color of flowering Pereskia guamacho trees dominates the dry forest land­ scape (Hoyos, 1978). Pittier (1978) reported large stands of this species from the plains at Monagas and Barcelona, and from around Caracas.

Specimens examined. SOUTHERN DUTCH ANTILLES. BONAIRE: 1909-10, Boldingh 5442 (U). COLOMBIA. ATLANTICO: Puerto Colombia, 50 m, 15 Mar 1930 (fl), Cufodontis 49 (W); nr. Bar­ ranquilla, Limon trail, 25 Sep 1932 (st), Dugand 112 (F); nr. Galapa, 16 Apr 1933 (f), Dugand 406 (F); between Palmar de Varela and Ponedera, 2-4 Aug 1943 (fl), Dugand & Jaramillo 3453 (US, COL); Barranquilla, 1927, Elias 151 (US); Barranquilla, 5-10 m, 29 Apr 1960 (fl), Mora 1391 (COL); S of Barranquilla, vie. Aeropuerto de Soledad, 6 m, 29 Apr 1960 (fl), Mora 1467 (AAU, COL); Puerto Colombia, Jun-Jul 1906, Pittier s.n. (US); nr. Salgar, 30 m, 12 Apr 1974 (fl), Plowman 3518 (GH). BOLIVAR: Galerazamba, 13 Oct 1962 (st), Saravia 1324 (COL); Cartagena, Cerro Pelado, Mar 1858 (fl), Schott 16 (BM, US). GUAJIRA: Rancheria Valley, Fonseca, 175 m, 10 Apr 1944 (fl), Haught 4065 (NY, US); Serrania La Macuira, Jassai region, nr. the dunes of Arehuara, 4 Mar 1963 (st), Saravia 2324 (COL, US); Torinche, between Uribia and Malcao (Maicao), 18 Oct 1963 (st), Saravia 2868 (COL, US); 8 km from Riohacha on rd. to Malcao, 20 Apr 1964 (fl), C. Saravia & M. Saravia 3684 (US, COL). MAGDALENA: Sierra Nevada de Santa Marta, ca. 300 m, 19 Jul 1948 (fr), Barkley & Araque, 24 (COL, US); Santa Marta, hills above Santa Marta Bay, 30-90 m, Jun 1916 (fl), Curran 357 (K, NY, US); Santa Marta, 1844, Goudot s.n. (P); Valledupar, 35 m, 10 Apr 1939 (fl), Hanbury-Tracy 304 (K); Santa Marta, 50-100 m, 1 Apr 1918 (fl), Pennell 4765 (NY); Santa Marta, 20 Jan 1930 (st), Record & Jaca 76 (F, GH, NY); Mamatoco, 4 Mar 1948 (fl), Romero-Castaheda 1122 (COL); Isla de Salamanca, nr. Rincon de Najagualito, 13 May 1966 (fl), Schnetter 4 (COL). VENEZUELA. ANZOATEGUI: Cantaura, 15 Feb 1949, F. D. Smith 17 (US), 7 Apr 1950 (fl), 122 (US). ARAGUA: Guamitas, 760 m, 10 Dec 1938 (st), Alston 5857a (BM); San Mateo, May 1957 (st), Curso Dendrol. 300 (U); nr. Colonia Tovar, 1856/57, Fendler 2326 (K, MO); Girardot, U. C. V. Maracay, 23 May 1974 (st), Guevara 1565 (VEN); Ocumare de la Costa, 2 Apr 1926 (fl), Pittienl2157 (VEN); Ocumare, 23 Apr 1937 (fl), Pittier 13967 (US), 14967 (VEN); Ocumare de la Costa, 0-15 m, 9 Dec 1943 (st), Steyermark 54932 (F, NY); Maracay, Vogl 76 (M). BOLIVAR: Ciudad Bolivar and vie, Feb/Mar 1921 (st), L. Bailey & E. Bailey 1921 (US); Ciudad Bolivar (Guyana Angostura), Grosourdy 13 (P). CARABOBO: Guacara, 400 m, 27 Jul 1938 (st), Williams 10276 (F, VEN), Pittier 10276 (G); vie. Puerto Cabello and Valencia, 27 Oct 1916 (st), Rose 21837 (NY), 21834 (US). DISTRITO FEDERAL: Caracas and vie, 1000-1200 m, 19 Dec 1920 (st), L. Bailey & E. Bailey 304 (US), 3 Jan 1921 (st), 658 (US); La Guaira-Rio Grande (Cabo Blanco), 12 Jun 1917 (fr), Curran & Haman 968 (GH, NY, US); El Cardonal, Cordillera de la Costa, S of La Guaira, 26 Mar 1972 (fl), Manara s.n. (VEN); Caracas, Bosque de Catuche, 1200-1800 m, 15 May 1913 (fl), Pittier 6179 (NY, US); vie. Caracas, 1913 (st), Pittier s.n. (US); La Guaira, 12 Jul 1900 (fl), Robinson & Lynn s.n. (US); between Caracas and La Guaira, 19 Oct 1916 (st), Rose 21704 (NY, US). FALCON: Between Coro and Alta Gracia (Zulia), 25 Apr 1917/1 May 1917 (fl), Curran & Haman 741 (GH, NY, US). GUARICO: Calabozo, 10 May 1962 (fl), Blydenstein 383 (VEN); S of Camatagiiito, 18 Apr 1925 (fl), Christ 10 (VEN); vie. El Sombrero, 17 Apr 1927 (fl), Pittier 12365 (G, M, US, VEN). LARA: Between los Yavos and El Chirio, 610 m, 10 Mar 1944 (fl), Steyermark 55651 (F); 30 km N of Barquisimeto along rd. to Coro, 3 Nov 1967 (fl), Tillet 6711-554 (VEN). MERIDA: Chama valley, near Estanques, 1 Sep 1965 (fr), Breteler 4553 (U, US); Lagunillas, 700 m, 11 Jan 1922 (fl), Jahn 965 (VEN, US); Estanques, 450 m, 31 Oct 1953 (fl), Little 15795 (VEN); above Los Gonzalez, canyon bordering Rio Chama, 1220- 1820 m, 2 May 1944 (fl), Steyermark 56225 (F, NY). MIRANDA: Las Barrancas de Guatire, May 1958 (fl), Aristeguieta 3052 (VEN); Santa Lucia, 150-200 m, 6-8 Mar 1943 (fl), Killip & Tamayo 37038 1986] PERESKIA (CACTACEAE) 89

(A, NY, US). NUEVA ESPARTA: El Cercado, Aug 1953 (fr), Gines 3386 (US); Isla de Margarita, Pt. Moreno, Aug 1903 (fl, fr), Johnston 216 (C, F, G, GH, K, NY, US, W); Isla de Coche, 5 Aug 1903 (fr), Johnston s.n. (GH); Isla de Margarita, Jul 1959 (fr), Ramia 1929 (VEN); Isla Margarita near Pampatar, 7 Aug 1936, Wagenaar Hummelinck 101 (U). SUCRE: Isla Arapo al Oeste, Norte de Bahia de Arapo, 10°16'N, 64°29'W, 1 Sep 1973 (st), Steyermark & Manara 107935 (VEN); Peninsula de Manare, Quebrada de Manare, Playa Garrapata south of Manare, between Punta Garrapata and Punta Aguirre, 0-15 m, 11 Sep 1973 (st), Steyermark et al. 108023 (VEN); SE of Cumana, Playa Manzanillo, 0-1 m, 8 Sep 1973 (st), Steyermark 108178 (VEN, U); Isla Caracas del Este, southwestern part, Playa El Corral, 0 m, 9 Sep 1973 (st), Steyermark et al. 108359 (VEN). ZULIA: Km 25 Maracaibo-Perija, 10 Nov 1968 (st), Aristeguieta et al. 6744 (VEN); Pararu (Goajira), 5 Jul 1975 (st), Perrin 1 (VEN); Maracaibo, 1824, Plee I (F, P); Maracaibo, 1826, Plee s.n. (MO, P); 18-22 km WSW of Maracaibo, rd. to Villa de Rosaria, 100 m, 21 Aug 1967 (st), Steyermark & Fernandez 99539 (US, VEN). Local names and uses. Colombia: guamacho. Venezuela: guamacho (Guarico, Merida, Miranda), supi (Falcon), suspiro (Merida), suspire, siichi (Zulia). The fruits are edible and the leaves are eaten by cattle. Trunks and larger branches are used for fence posts, which root and grow to considerable size. It is also grown as a hedge. Although Pereskia guamacho was described as late as 1898, specimens of this taxon were the first ones to be collected in pre-Linnean times besides those of P. aculeata. Dillenius (1732) recognized the plant illustrated by Plukenet (1692) as probably different from Plumier's "Pereskia aculeata flore albo fructu flaves- cente," but Linnaeus (1753) cited both Dillenius's plate and that of Commelin (1697, Comm. hort. 1, p. 135, t. 70) under Cactus pereskia. Later monographers evidently failed to pay closer attention to these plates and overlooked specimens in the Sloane Herbarium (BM), which reveal that more than one species was involved in Linne's Cactus pereskia. Commelin's plate shows an erect tree-like plant originating from Margarita Island (Nueva Esparta, Venezuela), and Plu­ kenet's straight-spined plant is also very distinct from the scandent, geminately hooked P. aculeata. Both are in fact P. guamacho. In the 18th and 19th century only a few specimens, unidentified or misidentified, were collected of this species, namely by Mutis in Colombia, between 1760 and 1808, and later by Schott, Plee, and Fendler, antedating Weber's description. None of these seem to have been known of or seen by Weber, who gives the Orinoco basin as the origin but cites no collector or herbarium specimen with his fairly accurate description of a flow­ ering plant. Leaf shape and size of P. guamacho are variable even on the same plant and the characters on which Britton and Rose based P. colombiana are not reliable, as noted already by Wagenaar Hummelinck (1938).

8. Pereskia zinniiflora De Candolle, Mem. Mus. Hist. Nat. 17: 75, t. 17. 1828 (as "zinniaeflora"). Type. Plate of Cactus zinniaeflorus in Mocino, Fl. Mex. ined. (no. 1012, preserved at the Hunt Institute for Botanical Documentation, Pittsburgh). Equivalent specimen. "Nova Hispania," 1787-1804, Sesse et al. 2137 (F, MA); "Nueva Espaha," Herb. Pavon (G). The specimen distributed by Pavon was apparently taken from the Sesse collection and represents the major part of the material. Figs. 15An, 34A-K, 35. Rhodocactus zinniiflorus (De Candolle) F. Knuth in Backeberg & Knuth, Kaktus-ABC 96. 1935. 1935. Pereskia cubensis Britton & Rose, Torreya 12: 13. 1912. Type. Cuba. Oriente, 1856-57, Wright 205 (lectotype, US; isotypes, G, GH, NY). Rhodocactus cubensis (Britton & Rose) F. Knuth in Backeberg & Knuth, Kaktus-ABC 96. 1935. Small tree, 4-8(-10) m tall, with diffuse branching and flat, much branched top, or shrub-like; trunk to 25(-30) cm in diam.; bark smooth, brown, or longi- [VOL. 41 90 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN

FIG. 34. Pereskia zinniiflora. A, twig with auxoblast leaves. B, flower. C, flower buds. D, flower bud, l.s. E, twig with fruits. F, G, seeds from different specimens. H, branch with young brachyblast leaves. I, spiny erect stem. J, areole from trunk. K, habit. A, B, from Alain s.n., cult. hort. Berol. 046- 02-78-30 (B); C from Jervis 1504 (MICH); D from Ekman 18911 (S); E, G from Wood & Atchison 7404A (A); F from Jervis 1905 (MICH); H, K from Britton 2013 with photograph (US); I from Britton et al. 13034 (NY); J from Wright 205, the type (GH). 1986] PERESKIA (CACTACEAE) 91

FIG. 35. Distribution of Pereskia zinniiflora.

tudinally fissured, with spiny areoles, or marked by transverse black bands formed by withered areoles; twigs 2-3 mm thick; internodes 4-20 mm long; twigs green when young, reddish-brown or pale brown after periderm formation. Roots (ex­ amined in cultivated specimens only) without thickenings (?). Areoles 2-3 mm in diam. on twigs, often prominent, with conspicuous rusty brown tomentum of trichomes of up to 0.5 mm length, producing spines and 1-3 brachyblast leaves, often elongating into spur shoots on distal flowering twigs; areoles accrescent on branches and trunk, broadly elliptic, to 10 mm long, 50 mm broad and 8 mm high on trunk. Leaves narrowly elliptic or ovate to oblanceolate, broadest near or above the middle, 10-40(-65) x (7-) 10-15(-24) mm; petiole 2-3 mm long, base cuneate to attenuate; blade 0.5-1 mm thick, fleshy; apex acute to shortly acu­ minate, rarely obtuse; venation pinnate; midrib prominent below; lateral veins 2-3, inconspicuous. Spines 0-5 per areole on twigs, acicular, often two or three at first, the lateral spreading, the median porrect, 7-30(-40) mm long and 0.3- 0.7 mm thick, with conspicuous conical thickening of up to 2-3 mm diam. at base; young spines subulate, yellow to brown at base, becoming brown to black; spines increasing in number on branches and trunk to 30-80, 2-5 cm long and 0.8-1.0 mm thick, brown or black, becoming gray with age. Flowers solitary, terminal on main branches and twigs and from lateral areoles and spur shoots, epigynous, 30-40 mm in diam.; pedicels to ca. 3 mm long. Staminate flowers with receptacle turbinate, 5-10 mm long, 5-8 mm in diam., bearing 2-6 areoles with sparse rusty brown tomentum; lower bracts 1-3, obovate to lanceolate, leaf­ like, 8-14 x 4-6 mm, with cuneate base and acute apex; upper bracts (0-)l-3, lanceolate to broadly obovate, ca. 5-7 x 5 mm, with rounded apex; sepaloids 2- 3, broadly ovate to suborbicular, 6-15 x 6-9 mm, with broad base, green when fresh, venation nearly parallel; petaloids 7-9, obovate, 15-25 x 8-12 mm; base cuneate to attenuate, apex rounded to emarginate; blade purplish-pink, variously described as pink to crimson-purple; stamens ca. 150, 4-8 mm long; filaments white at base, apically pink; anthers 1.2-1.5 x 0.5-0.7 mm, pale pink (pollen 92 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 yellow); pollen 12-colpate, medium-sized, tectum perforate and with large spi­ nules; ovary inferior but with free ovary roof; locule small, ca. 1 mm in diam.; placentae parietal; style 3-5 mm long and ca. 1 mm thick, abruptly rising from the ovary roof, narrowing into the 3-9 filiform-subulate stigma lobes 1-2 mm long; shorter than the stamens; ovules aberrant or absent. Pistillate flowers (data from Areces, 1984, p. 161, as Rhodocactus cubensis) with receptacle broadly ovate or subglobose, 9-14 mm long, 9-13 mm in diam.; receptacular bracts 1-3, elliptic to oblanceolate, 8-17 x 4-8 mm; sepaloids ca. five, orbicular or ovate, 4-8 x 4-7 mm, pale green with pink margins, transitional to the petaloids; petaloids 8- 11, subspathulate to obovate, 11-18 x 5-12 mm, emarginate or rarely rounded or truncate, bright reddish-pink; stamens abnormal, dwarfed, ca. 2 mm long; anthers empty; style stout, terete, finely striate, 3-5 mm long, 1-2 mm thick; stigma lobes 7-10, spreading, 3-8 mm long, 1 mm thick; locule subglobose, 4-7 mm in diam.; ovules numerous, parietal. Fruit a globular to depressed ficiform berry, ca. 14-20 mm in diam. (18-30 mm in diam. according to Areces, 1984); pedicel ca. 2 mm long and 1 mm thick; areoles few, nearly naked, ca. 2 below the receptacular rim, 2-3 at the rim around the accrescent, naked, flat, smooth umbilicus of 6-9 mm diam. formed by the accrescent ovary roof; flower remnants deciduous. Seeds to 50 or more, obovate to reniform, (2.5-)2.8-3.0 mm long, 2.2-3.0 mm broad and 1.0-1.5 mm thick, smooth on the sides with shallow peripheral ribs, black, glossy; hilum subbasal, transversely elliptic; micropylar end rounded or nose-like. Chromosome number: In = 22; Alain s.n., cult. hort. Berol. 046-02-78-30 (B). Distribution (Fig. 35) and phenology. Cuba, endemic to the southern and south­ western part of the island, in the lowlands, in dry coastal thickets, xerophytic scrub thickets, "poor forests," and salty pastures, according to collectors' data; flowering from ca. March to August, fruits reported from May to August. Plants of this species are dioecious according to Areces (1984), and this feature is apparently common in other Caribbean species of the genus. I have seen many staminate flowers in herbarium specimens and on one cultivated plant, but no perfect flowers. One single pistillate flower seen with a fruiting specimen {Alain 1080) has papillate stigma lobes ca. 5 mm long, and dwarfed sterile stamens. In staminate flowers the locule is very small, and the ovules, if present, are aberrant. In one case, teratological stamens arising from the placenta were found {cult. hort. Berol. 046-02-78-30). The reproductive biology of this species remains to be studied. Areces (1984) observed Lepidoptera and Hymenoptera visiting the flow­ ers.

Specimens examined. WITHOUT LOCALITY (probably Cuba): "Iter Mexicanum," 1841-42, Karwinski s.n. (LE). CUBA. WITHOUT EXACT LOCALITY: Sagra s.n. (P); "Abrojo de la Florida," Sagra 6 (P); Jacqua- Oran, Sagra s.n. (P). CAMAGUEY: Santa Cruz del Sur, 2 May 1917 (fl), Ekman 8609 (G, K, S). HABANA (doubtfully native): "La Havane," 1821, Ferrero s.n. (G-DC). LAS VILLAS: Lomas San Juan, 16 Jul 1947 (fr), Wood & Atchison 7404A (A). SANTIAGO DE CUBA (ORIENTE): Pilon, S of Manzanillo, Jul 1949 (fl), Alain 1080 (GH); vie. Guantanamo, Los Canos, 19-31 Mar 1909 (st), Britton 2013 (NY, US); Ensenada de Mora, 28 Mar 1912 (fl), Britton et al. 13034 (K, NY, US); Caimanera, May 1889 (st), Eggers 5441 (NY); Mir, Rio Rioja, 4 Mar 1915 (st), Ekman 4871 (S); Santiago, western shore of Santiago Bay, 22 Jun 1914 (fl), Ekman 1421 (S); Guantanamo, Novaliches, 16 May 1918 (fr) Hioram 1835 (GH, NY), Jervis 1061 & 1504 (MICH), Leon 16731 (GH); Guantanamo valley, Glorieta, 8 Aug 1951 (fr), Jervis 1879 (MICH); E of Santiago de Cuba, between Sigua and Verraco, 11 Aug 1951 (fl, fr), Jervis 1904 & 1905 (MICH). SANTA CLARA: Trinidad, Loma de la Vigia 29 Mar 1924 (fl), Ekman 18911 (S), Britton & Wilson 5513 (NY, US). Local names. Erizo, abrojo (Leon & Alain, 1953). When describing this species from the plate of "Cactus zinniaeflorus" of Mo- 1986] PERESKIA (CACTACEAE) 93

cirio's unpublished Flora Mexicana paintings, de Candolle assumed it to be from Mexico. However, according to McVaugh (1977), some of the plants illustrated there were in fact from the West Indies, collected by Sesse in 1795. Two specimens apparently belonging to the same collection and referable to the plate were located, one at Madrid (MA) in the Sesse and Mocino collection, and one at Geneva (G) labeled as from "Herb. Pavon." As noted by McVaugh (1977), specimens from the Sesse and Mocino collection were sold by Pavon. Comparison of the specimens and the plate with old and recent collections of all taxa from the West Indies and from Mexico and Central America shows that Pereskia zinniiflora is the same as the Cuban plant later described as P. cubensis and that it is entirely different from any species from the mainland. It is noteworthy that the specimen at Geneva was annotated by J. N. Rose as "Pereskia zinniiflora ?," but this was not mentioned in the monograph of Britton and Rose (1919). Bravo-Hollis (1959) saw the spec­ imen {no. 2137) in the Sesse and Mocino collection at Madrid and also correctly identified it, but she later included a different taxon, Pereskia tampicana, in the synonymy of P. zinniiflora, assuming that both were from Mexico and apparently influenced by the superficial similarity of two illustrations which, however, differ widely in scale (Bravo-Hollis, 1978, p. 149, figs. 67, 68). Pereskia tampicana is discussed under P. grandifolia. The original plate of" Cactus zinniaeflorus" which was only recently rediscovered (McVaugh, 1982) is of excellent quality, and while the spines and areoles of the older stem do not seem entirely typical the flower and leaves as well as floral details illustrated separately are very accurate. The exact origin of Sesse's plant is unknown, and it is possible that it was not collected within the natural range of distribution of the species but perhaps from a plant cultivated at or near Havana, where Sesse stayed for nine months in 1795 (McVaugh, 1977). Several cases of plants in the Sesse collection originating from eastern Cuba but not collected there by Sesse himself are reported by McVaugh (1977). There is a specimen dating back to 1821 {Ferrero s.n.) provided with locality data "La Havane" but there is no evidence that the species is native there. Another collection {Sagra s.n.) cannot be placed. Ramon de la Sagra was the director of a botanical garden and professor at Havana from 1822 to 1835 (Stafleu & Cowan 19.83; Urban, 1902-1903). Furthermore, Karwinski's collection of 1840- 41, labeled as "Iter Mexicanum 1841-42," may well be from the same source, for he also visited Havana but not the southern and southeastern part of Cuba (McVaugh, 1980b). More recent collectors and authors report the species only from the provinces of Santiago de Cuba (Oriente) and Las Villas (Leon & Alain, 1953) and from Santiago de Cuba, Camagiiey, and Santa Clara (Werdermann, 1931).

9. Pereskia portulacifolia (Linnaeus) De Candolle, Prodromus 3: 475. 1828. Figs. ID, 5B, E, HE, 14G, H, 15Am, 36, 37A-K. Cactus portulacifolius Linnaeus, Sp. pi. 469. 1753. Type. "Opuntia arbor, spinosissima, foliis Portulacae cordatis" Plum. cat. p. 6. Specimen apparently lost. Linnaeus (Spec. pi. ed. 2, 671. 1762) refers to the illustration in Plumier, PI. amer. fasc. (ed. Burmann, 1758) pi. 197, fig. 1, drawn from Plumier's collection, herewith designated as lectotype. According to Plu­ mier's manuscript "Botanicum americanum .. ." (cf. Hunt, 1984), the plant was collected by Plumier himself in St. Domingue (Haiti), Fond Parisien, towards the Grand Cul-de-Sac. Rhodocactus portulacifolius (Linnaeus) F. Knuth in Backeberg & Knuth, Kaktus-ABC 96. 1935. Small tree 3-5(-10) m tall, sometimes branching from near the base, shrubby; main branches suberect to arching; trunk to 15-20 cm in diam.; bark gray, nearly smooth but interrupted by the often prominent areoles; twigs 2-3(-4) mm thick, 94 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL- 41 brown, reddish or gray, dark green before periderm formation, older twigs and branches often with knob-like spur shoots to 5-10(-15) mm length and 2-2.5 mm in diam.; internodes 5-15 mm long; pith of main shoots ca. 4 mm in diam., septate. Roots with fusiform thickenings. Areoles rounded to transverse-elliptic, 1-2 mm in diam. on twigs, accrescent on branches, to 5 mm long, 7-9 mm broad and 5 mm high on trunk; tomentum of areoles short, gray, trichomes ca. 0.5 mm long, brown at base; areoles producing spines and l-2(-3) brachyblast leaves, often elongating into spur shoots but then normally spineless or older spines deciduous. Leaves variable in shape and size; auxoblast and brachyblast leaves not differing on distal twigs, obovate-spathulate to cordiform, 5-15(-30) x 4- 10(-12) mm; auxoblast leaves of main shoots larger, spathulate, 20-35 x 8-15 mm; petiole obscure, 0-1 mm long, blade fleshy, ca. 0.6 mm thick, base attenuate to cuneate, apex broadly acute to rounded, truncate or emarginate, usually acute on long shoots; venation pinnate; midrib prominent below, lateral veins obscure, 0-2(-3), ascending. Spines 0-3 on twigs, (5-) 10-16 mm long and 0.3-0.4 mm thick, stiff, spreading, reddish-brown to black, with conspicuous conical thickening to 1 mm in diam. at base; spines increasing in number to about 50 on old areoles of trunk, 10-22 mm long and 0.5-0.6 mm thick, black, base to 1.5 mm in diam. Flowers imperfectly unisexual (plants functionally dioecious), terminal or on spur shoots, or in subterminal groups, one flower from each of the approximated areoles; 30-55 mm in diam., corolla in bud spherical; pedicels 2-3 mm long. Staminate flowers with turbinate receptacle of 6-8 mm diam., smooth, green; lower bracts 2-3, leaf-like, obovate-spathulate, attenuate at base, 4-9 x 2-7 mm; upper bracts 0-2, spreading, obovate to spathulate, 4-7 x 2-5 mm; sepaloids 3- 5, appressed in bud, obovate to suborbicular with broad base and rounded apex, 8-12 x 4-9 mm; petaloids 8-9, spreading, obovate 15-25 x 7-15 mm, bright pink to purplish-rose; base cuneate to attenuate, apex rounded to slightly emar­ ginate or nearly erose; stamens ca. 200, 5-8 mm long, distinctly longer than the style and completely masking it; filaments pink; anthers 0.8-1.2 mm x 0.6-1.0 mm, pink; pollen 12-colpate, medium-sized, tectum perforate and with large spinules; ovary inferior, but with horizontal, free, ovary roof abruptly narrowed into the style; locule with concave floor, ca. 1 mm in diam., with parietal placentae but no ovules; style reduced, 2-3.5 mm long and 0.5-1 mm thick, narrowed towards the tip and ending in ca. five erect, subulate stigma lobes ca. 0.5 mm long, without papillae. Pistillate flowers similar to the staminate except in the shape of the receptacle and characters of the androecium and gynoecium; recep­ tacle ca. 10 mm in diam., cup-shaped, subglobular; stamens dwarfed, ca. 2-3 mm long, sterile, arising from the low rim of the receptacle surrounding the flat ovary roof; locule ca. 5 mm in diam., spherical, with conspicuous septal ridges at the roof and numerous ovules on long U-shaped placentae in parietal position; ovules ca. 0.8 mm long, with funicles to 1 mm long; style ca. 1 mm long and in diam., grooved; stigma lobes ca. 10, to 7 mm long and 1 mm broad, suberect to spreading. Floral parts above the ovary roof deciduous in early postfloral stage, exposing the umbilicus in young fruits. Fruit subglobular to broadly ficiform, 18-30 mm in diam., pedicel ca. 2 mm long and thick (when fresh); fruit smooth to rough leathery when dry; areoles inconspicuous, bracts 0-2, persistent or deciduous, broadly obovate, ca. 4-5 x 5 mm; umbilicus depressed, flat, circular (often rounded tri­ angular when dried), ca. 6-11 mm in diam., formed by the accrescent ovary roof, fruit wall ca. 2 mm thick; locule filled by the seeds and pulp formed by juicy gelatinous funicles. Seeds numerous, obovate to reniform, ca. 3-3.5 mm long, 2.5-1.2 mm broad and 1.2 mm thick, with low peripheral ribs, smooth on the sides, black, glossy. 1986] PERESKIA (CACTACEAE) 95

2Q0

• Pereskia portulacifolia

A Pereskia quisqueyana

A Pereskia spec. A.

FIG. 36. Distribution of Pereskia portulacifolia, P. quisqueyana, and P. sp. A.

Distribution (Fig. 36) and phenology. Endemic to Hispaniola; arid regions of Haiti and adjacent areas in the southwestern Dominican Republic, in the lowland, in dry forests and thorn scrub with Acacia, Stenocereus, Neoabbottia, and Opuntia; common in Haiti; main flowering period from March to August, fruits collected in May and August. This species has imperfectly unisexual flowers occurring on separate plants, as only recently documented by the collections Zanoni et al. 35204 ($) and 35205 {6), following earlier unconfirmed observations by the author. Since the plate for P. portulacifolia (Fig. 37) was already finished, a separate illustration of the pis­ tillate flowers is given in Figure 14G, H. Lamarck (1785) noted sterile and fertile flowers, based on a manuscript of Plumier's, and Plumier's plate 197, figure 1 (Plumier, 1758), reproduced in Britton and Rose (1919, p. 23, fig. 20), shows one staminate flower between two pistillate flowers with long, spreading stigma lobes. This feature has been overlooked by subsequent authors and collectors. In one specimen with flower and fruit {Clover 1130), I found the only flower to have sterile anthers of normal size and inconspicuous stigma lobes without papillae. The numerous flower buds which I collected at the beginning of the flowering season at two different sites in the Lago Enriquillo valley in the Dominican Republic {Leuenberger 3046 & 3048) are all staminate, but fruits were later col­ lected from trees in the same region {Czerwenka s.n.). Specimens examined. HAITI. Morne Cahos, nr. Gonaives, 80 m, May 1900 (fl), Buch 341 (NY); lie de la Gonave, Grande Lagune, 12 Feb 1928 (st), Ekman H9593 (S); Dept. de l'Ouest, Cul-de-Sac plain, 8 m, 23 Aug 1948 (fl, fr), Clover 11030 (MICH, US); Cul-de-Sac, foot of Morne a Cabrits, 18 Jul 1924 (fl), Ekman H953 (K, NY, S); vie. Etang Saumatre, mid regions of Cul-de-Sac, 4-12 Apr 1920 (fl), Leonard 3503, 3606 (NY, US); vie. Maneville, Etang Saumatre, 13-17 May 1920 (fl), Leonard 4251 (NY, UPS, US); vie. Port-au-Prince, 5-9 Jul 1920 (st), Leonard 5325 (NY, US); nr. Fond Parisienne, 25 m, 15 Oct 1941 (fl), Holdridge 845 (MICH, NY, US); Riviere Blanche district, 18 Sep 1917 (st), Cook s.n. (NY, US); Savane Desole, 12 Jun 1941 (fl), Bartlett 17385 (LIL, MICH, US); Thomazeau, 2-4 Apr 1917 (fl), Bautsch 137 (NY, US); S of GonaiVes, Jun 1985 (fl), Zanoni 35204, 35205 (B, JBSD). DOMINICAN REPUBLIC. INDEPENDENCE: Enriquillo Valley, 1.5 mi from Lago Limon, 24 Aug 1950 (fl), Howard 12598 (A); km 23 from Jimani at junction of rds. to Duverge and Puerto Escondido, 96 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 37. Pereskia portulacifolia. A, branch with staminate flower. B, leaf dimorphism and vari­ ation; a from long shoot; b, brachyblast leaves. C, flower bud, l.s. (staminate flower). D, twig with flower buds. E, fruit (dried specimen). F, fruit (preserved in alcohol). G, fruit l.s. H, seeds. I, erect stem with auxoblast leaves. J, areole of trunk. K, habit. A, D, K from Leuenberger 3048, including photographs (B); B, C, I, J from Czerwenka 28 (B); E, H from Clover 1130 (MICH); F, G from Czerwenka s. n. (B). 1986] PERESKIA (CACTACEAE) 97

18°21'N, 71°42'W, ca. 10 m, 27 Apr 1982 (fl), Czerwenka 28 (B); 23 Mar 1983 (fl), Leuenberger 3046 (B, JBSD); 3 Sep 1960 (st), Marcano & Jimenez 4305 (US); 23 km SE of Jimani, 10-20 m, 3 Jul 1980 (fl), Zanoni & Mejia 7152 (JBSD); on rd. from Jimani to El Limon, N of Loma Las Trincheres, 18°28'N, 71°49'W, 120 m, 23 Mar 1983 (fl), Leuenberger 3048 (B, JBSD). Local name. Camelia roja (Liogier, 1974, p. 226). This species was discovered by Plumier between 1689 and 1695 and listed as Opuntia arborescens spinosissima foliis portulacae cordatis in his manuscript "Bo- tanicum americanum. . ." (Hunt, 1984) but not included in his new genus Pereskia (Plumier, 1703). The original material seems to be lost, but a plate drawn from it apparently still exists at Paris (Hunt, 1984). There are also copies (tracings) of the original plate at the University Library at Groningen and at Kew besides the printed version by Burmann (Plumier, 1758), which is chosen as lectotype for the direct reference given to it by Linnaeus (1762). The copies at Groningen were annotated by W. T. Stearn (T. P. Hidma, pers. comm.), who identified one on thin paper as a tracing of the original drawing by Claude Aubriet made under the supervision of S. Vaillant at Paris from Plumier's drawing done in the West Indies. The engraving on thicker paper with several plants on one plate is identified as a "proof before lettering and numbering of J. Burmann's Plant. Amer. . . ." The first author to recognize this plant as a species of Pereskia was Haworth (1812, p. 199). However, he did not make the formal combination but mentioned it only as "OBS. Cactus portulaccaefolius is another species of this Genus," noting that it had not yet been introduced into English gardens. The species was not collected again until 1900 and to my knowledge it had not been introduced into cultivation outside Hispaniola until 1985. Pereskia portulacifolia is closely related to P. zin­ niiflora, and together with P. quisqueyana, it belongs to a group of endemic Caribbean species of common ancestry. Some specimens of P. zinniiflora seem to approach this species in leaf shape, and a rather narrow species concept must be applied in this case to avoid bulky trinomials.

10. Pereskia quisqueyana Liogier, Phytologia 43: 183, fig. 6. 1980. Type. Do­ minican Republic. La Altagracia ("La Romana"): Bayahibe, 9 Jul 1977 (fl), Alain H. Liogier 27032 (holotype, UPR; isotypes, NY, US). Figs. 3B-D, 11G, 19F, G, 36, 38A-H. Shrub 2-4(-5) m tall, main shoots to ca. 10 cm thick; trunk to 25 cm in diam. at base, bark of main shoots smooth, brown; branches with pale brown to greenish- gray, smooth bark; twigs 2-3(-4) mm thick, olive-green to brown, drying pale brown. Roots with fusiform thickenings 20-30 cm long and 5-10 cm thick, fleshy. Areoles rounded, 1-3 mm diam. on twigs, to 15 mm diam. on trunk, with white to light gray or pale brown tomentum of trichomes ca. 1 mm long, producing spines only. Leaves elliptic to obovate-lanceolate, petiolate; base attenuate; apex acuminate or acute-acuminate, variable in size, those of the distal twigs elliptic- obovate, (2-)3-5(-7) x (1.4-)2.5-3.0 cm; petiole ca. 1-5 mm long; blade 0.8 mm thick; midrib prominent below, lateral veins ca. 2-3, inconspicuous; leaves of strong shoots 5-12 x 1.5-2.5 cm; petiole 5-15 mm long, midrib prominent below, lateral veins 3-5(-6), rather inconspicuous. Spines 0-4 on twigs, often lacking on flowering twigs, 10-15 mm long and 0.2-0.4 mm thick, spreading, stiff, pungent, straw-colored to brown with light tip, base not or slightly thickened, increasing in number on branches, to 120 per areole on trunk, 2-6 cm long and 0.5-0.8 mm thick, spreading, pungent. Staminate flowers terminal and on short spur shoots, (4_)5_7 cm in diam.; pistillate flowers not seen; flower buds with subspherical- ovoid corolla; pedicels (0-)2-3 mm long; receptacle 7-8 mm in diam., turbinate- 98 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 38. Pereskia quisqueyana. A, branch with young leaves, flower and flower buds. B, leafy shoot with terminal flower. C, flower bud. D, flower, l.s. E, leaf variation. F, areoles of older stem. G, habit. H, root-stock with partly exposed fusiform root thickenings. A, C-H from Leuenberger 3044 including photographs (B); B from a photograph by A. Liogier of cultivated material of the type collection. 1986] PERESKIA (CACTACEAE) 99

poculiform, green; areoles 3-5, inconspicuous, almost naked or with sparse white wool of trichomes 1 mm long, the lower ones rarely with 1-2 weak spines up to 5 mm long; lower bracts 0-3, leaf-like at the base of the pedicel and receptacle; upper bracts on the rim of the receptacle 0-3, obovate-lanceolate, to 14 mm long and 6 mm broad, thick, green, with midrib prominent below; sepaloids three, obovate, 15-17 mm long and 7-9 mm broad, apex rounded, blade greenish-pink; petaloids ca. 8-9, spreading, obovate-spathulate, (25-)30-35 mm long and 18- 22 mm broad, pink; base narrow cuneate; apex emarginate; stamens ca. 150, longer than the style, filaments 7-11 mm long, the outer ones longer than the inner ones, pink; anthers 1.6-2.5 mm long and 0.5-0.9 mm broad, dull pink; pollen 12-colpate, medium-sized, tectum perforate and with large spinules; ovary of staminate flowers deeply sunk in the turbinate receptacle, locule ca. 1 mm in diam., placentae in the upper part of the locule, parietal; ovules aberrant; style 4 mm long, 0.5 mm thick, narrower towards the stigma, stigma lobes ca. five, 0.5- 1 mm long, erect, concealed by the stamens, without stigmatic papillae. Fruits unknown. Distribution (Fig. 36) and phenology. Endemic to the southeastern part of the Dominican Republic, at sea level in coastal thickets on coral limestone and sand, flowering observed in March and July; only known from the type locality.

Specimens examined. DOMINICAN REPUBLIC. LA ALTAGRACIA: Bayahibe, 2 Oct 1976 (st), A. Liogier et al. 25424 (JBSD); 19 Mar 1983 (fl), Leuenberger 3044 (B, JBSD). The diagnostic characters mentioned in the original description of this species, namely flower and leaf size, hardly seemed to justify specific status as distinct from P. zinniiflora, but examination of the type specimen and further material as well as my own observations at the type locality revealed that the taxon is well delimited by leaf shape, spine color and dimensions, tomentum of the areoles, and size of the anthers. The fusiform thickenings of the roots hitherto only known in Andean species were observed during the same visit at the type locality and in cultivated specimens in the collection of Padre Cicero at San Cristobal, Do­ minican Republic. Unless it is found at other localities this plant could become highly endangered due to its extremely small and exposed habitat on the peninsular shoreline near a village with sand beaches. Further observations on the repro­ ductive biology are necessary, because normal ovules and fruit have not been reported yet.

11. Pereskia bleo (Kunth) De Candolle, Prodromus 3: 475. 1828. Figs. 5H, 7C, 10, 11A, 15Ai, F, 39A-J, 40. Cactus bleo Kunth in Humboldt, Bonpland & Kunth, Nov. gen. & sp. pi. 6: 69. 1823. Type. Colombia. Bolivar: nr. Badillas on the Magdalena river, May 1805 (fl), Humboldt & Bonpland 1546 (lectotype, P; isotypes, B-W, P). Rhodocactus bleo (Kunth) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1939. Pereskia panamensis F. A. C. Weber in Bois, Diet. Hort. 939. 1898. Type. "Isthme de Panama." No specimen cited, but a specimen, probably labeled in Weber's handwriting, may serve as lectotype: "isthme de Panama 1888-89, Geay 27, cult, in caldar. mus. Paris (P)." Pereskia corrugata Cutak, Cact. Succ. J. (Los Angeles) 23: 173, figs.95 , 96. 1951. Type. Cultivated at Missouri Bot. Gard., origin unknown, 10 Sep 1951 (fl), Cutak s.n. (MO). Rhodocactus corrugatus (Cutak) Backeberg, Cactaceae 1: 118. 1958. Shrub or small tree, 2-8 m; trunk to 15 cm diam., armed or unarmed; bark of trunk and branches smooth, olive-green to brownish-gray; twigs 2-5 mm thick, main shoots and fresh twigs often thicker, dark green to olive when fresh; stomata lacking, periderm formation early. Areoles transversely oblong, 2-5 mm wide on 100 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 39. Pereskia bleo. A, twig with flower buds. B, flower bud. C, flower, l.s. D, flower. E, fruit. F, fruit, l.s. G, seeds. H, leaf. I, one-year-old stem with areoles. J, areole from older main shoot. A- F, I from cult. hort. Berol. 277-01-80-80; G from Pittier 4704 (US); H from Steam et al. 128 (GH); J from cult. hort. Berol. 082-18-82-70. 1986] PERESKIA (CACTACEAE) 101 twigs, circular and 8-10 mm diam. on trunk; tomentum whitish to light brown; hairs uniseriate, ca. 0.5-1 mm long; areoles producing spines but no brachyblast leaves. Leaves narrowly elliptic to oblong or lanceolate, (4-)6-20(-25) x (l-)2- 7(-8) cm; average leaves ca. 16 x 6 cm, smaller and narrower at the base of new shoots and on distal twigs; petiole distinct, 0.5-3 cm long, ca. 3 mm wide and to 4 mm thick when fresh, rounded below and slightly grooved above; leafblade fleshy, 0.5-1 mm thick, base gradually narrowed into the petiole, attenuate; margin entire, often undulate when fresh; apex acuminate; venation pinnate; lateral veins 4-6(-7), arching, often bifurcate in the distal third of their length, impressed above in fresh leaves. Spines usually 1-5 on twigs, sometimes lacking, to 40 per areole on trunk, subulate, 5-10(-l 5) mm long and 0.4-0.6 mm thick on twigs, ca. 20(-50) mm long on trunk, thickened at base, fasciculate to spreading, black or nearly so, with red base in youth, becoming gray with age. Inflorescence condensed racemose, terminal on main shoots and lateral twigs; main shoot narrowed into the indistinct pedicel of the terminal flower; lateral flowers arising in acropetal sequence from the axils of the uppermost leaves and leafy bracts; receptacle not proliferating. Flowers epigynous, 4-6 cm in diam., pedicels 5-10 mm long and ca. 3 mm thick at base, naked; receptacle turbinate, 5—10(—12) mm in diam. at the rim, with ca. five irregular blunt angles, usually ebracteate below its greatest diameter, rarely with 1-2 linear-lanceolate, lower receptacular bracts; upper bracts above the rim of the receptacle, 4-5, erect, lanceolate, up to 18 x 2 mm, the inner ones shorter, triangular, ca. 8 x 5 mm; sepaloids 2-3, ovate, ca. 12-21 x 12-17 mm, with broad base, green or reddish towards the margin; petaloids 10-15, obovate to broadly obovate, 2-3(-4) x 2 cm, the outer with broad base, the inner with narrow base, cuneate, apex broadly rounded and softly mucronate to emarginate, blade thin and delicate, bright red, scarlet, orange-red in bud, also described as salmon, flesh-colored, or (according to Weber, 1898b, p. 93, P. panamensis) yellowish- white with pink margins (unconfirmed); margins sometimes turning pink at high temperatures; venation palmate-flabellate. Stamens ca. 120, 5-13 mm long, fil­ aments erect, clustering around the style, pale at base, conspicuously red in the upper two thirds; anthers 0.9-1.4 x 0.3-0.4 mm, cream; pollen 9-12-colpate, medium-sized, tectum perforate and with spines barely 1 ^rn long. Ovary inferior but with free, flat ovary roof, locule cup-shaped, unilocular with septal ridges at roof; placentae parietal in the upper half of the locule at the base of the septal ridges; ovules numerous, ca. 1 mm in diam.; style abruptly rising from the flat ovary roof, 10-14 mm long, ca. 1 mm thick, thinner at base; stigma lobes 5-6, 4 mm long, cream. Fruit 2L turbinate, truncate berry 4-5(-7) cm long and in diam., naked or with 1-2 persistent bracts on the rim; distal end with a broad, flat, and smooth umbilicus formed by the accrescent ovary roof and surrounded by a circular groove along the raised, rounded rim of the receptacle; yellow at maturity, sour; wall ca. 1 cm thick, locule 1.5-2 cm in diameter. Seeds few or to 10-20, mostly obovate, but variable in outline, 6-7.5 mm long, 5-6.5 mm broad and 3- 4 mm thick, smooth, black; hilum subbasal, reniform to subquadrangular, cream- colored, with a low and broad rim formed by spongy tissue and central funicle scar. Seedlings (three weeks old) with hypocotyl ca. 3 cm long and 3 mm thick; cotyledons ovate, ca. 30 x 20 mm, venation pinnate, intercostal areas convex; cotyledons accrescent on older seedlings to ca. 38 x 27 mm; primary leaves like regular leaves but smaller, reddish-olive, areoles with numerous spines 1-5 mm long; juvenile plants with basitonous branching from the lowermost areoles above the cotyledonar areoles. Chromosome number: In = 22; cult. hort. Berol. 277- 01-80-80 (B). A gametic chromosome number of n = 11 was reported by Ross (1981). 102 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

0 50 100 150 200 km

FIG. 40. Distribution of Pereskia bleo.

Distribution (Fig. 40) and phenology. Panama and Colombia, from sea level to about 500 m, in Colombia to 1300 m, on sandy soil along streams and on river banks, also in secondary forest and in hedges; flowering mainly from March to June; fruiting from October to March (data insufficient). The species is notable for its rather mesic habitat, but the stems and leaves, although fairly soft, show a degree of succulence similar to large-leaved species of more xeric habitats. Three specimens are omitted from the distribution map because I could not place them precisely (all from Colombia, on the Rio Mulatos, Rio Sinu, and "lands of Loba")-

Specimens examined. PANAMA. WITHOUT LOCALITY: 1850, Duchassaings.n. (GOET); Nov 1909 Gaillard s.n., NY 09.510 (NY); Goldman 1956 (US). CANAL ZONE: NOV 1909 (fr), Gaillard s.n. (NY, US); Pittier 3507 (US); around Culebra, 50-150 m, 27 Nov 1911 (fl), Pittier 4821 (US); lake shore along Gatun River valley, 25 Mar 1925 (fl), Pittier 6510 (US). DARIEN: Between Pinogana and Yavisa, ca. 15 m, 17 Mar 1937 (fl), Allen 281 (MO); "forest of Darien," Dawe 869 (K, NY); Rio Pirre, 14 Jul 1971 (fl), Croat & Porter 15519 (F, MO); vie. Cana, 540 m, 10 Mar 1912 (1915), Goldman 1884 (NY, US), Pittier 1884 (NY); forests around Pinogana, 16-21 Apr 1914 (fl), Pittier 6545 (US); Pinogana- Yaviza, 1 km from Tuira, 6 Jun 1959 (fl), Stern et al. 128 (G, GH, LE, MO, UC); Chonta, banks of Rio Cupe, 27 June 1959 (fl), Stern et al. 732 (GH, LE, US); Marraganti, 3-65 m, 4 Apr 1908 (fl), Williams 661 (NY); Quebrada Camachimuricate, nr. Casa de Bartolo, 15 Mar 1973 (fl), Kennedy 2868 (MO). PANAMA: Chepo, Apr 1933 (fl), Paul 339 (US); Panama City, 1 Mar 1905 (fr), Pittier 1 (US); nr. Chepo, 30 Oct 1911 (fr), Pittier 4704 (P, NY, US); Taboga Island, Perlas Islands, 22 Jun 1969 (fl), Tyson 5581 (MO). SAN BLAS: Caledonia Harbour, Mount Vernon, 26 Apr 1939 (fl) Elmore L34 (MICH). COLOMBIA. ANTIOQUIA: 10 km E of Turbo, 50 m, 7 Apr 1945 (fl), Haught 4547 (NY, US); Turbo, forest on Rio Mulatos, 21 Apr 1946 (fl), Haught 4798 (COL, US); Chigorodo, 100 m, 26 Apr 1972 1986] PERESKIA (CACTACEAE) 10 3

(fl), Plowman & Saldarriaga 3177 (GH, K, US). BOLIVAR: 25 km E of San Onofre, arroyo Jojan, 300 m, 9 Mar 1962 (fl), Beuther 91 (COL, MO); lands of Loba, Amargamiento Rico and vie, Apr-May 1916 (fl), Curran 355 (K, NY, US); Boca Verde on Rio Sinu, 100-300 m, 4 Mar 1918 (fr), Pennell 4579 (GH, MO, NY, US); vie. Since, 6 Sep 1963 (st), Romero-Castaheda 9794 (COL). BOYACA: MUZO, Goudot s.n. (P). CHOCO: Mun. de Riosucio, Tilupo, 16 Jun 1957 (fl), Romero-Castaheda 6269 (AAU, NY). HUILA: W slope of the Cordillera Oriental, above Guadalupe, 1000-1300 m, 20 Mar 1940, Perez-Arbeldez & Cuatrecasas 8405 (COL). SANTANDER: Vic. Puerto Berrio, between Carare and Magdalena River, 100-700 m, 16 Apr 1935 (fl), Haught 1643 (COL, US). TOLIMA: Mariquita, 547 m, Apr 1952 (fl), Perez-Arbeldez A-1030 (COL). VALLE DEL CAUCA: Pradera, 1000 m, Dec 1937 (fl), Dryander 1962 (US). Local names and uses. Panama. Naju de culebra, najii or naju de espinas. Colombia. Bleo, chupa, bleo de chupa (Bolivar), chupa melon (Choco). In Co­ lombia it is used for hedges. The leaves were used for clarification of water, and the fruits are edible, but sour. Pereskia bleo is probably the most remarkable member of the genus and a most unusual cactus in general because of its large petiolate leaves, the showy bright red flowers, the distinctive turbinate fruit, and the terrestrial rain forest habitat. It is cultivated occasionally outside its natural range, particularly in the West Indies, but until recently was unknown in European collections. Pfeiffer (1837a, 1837b) mentioned P. cruenta Hort. as a synonym, which indicates that the species has been in cultivation earlier, but later reports of P. bleo from cultivation refer to misidentified plants of P. grandifolia. Ever since Schumann (1890, 1898) mis­ applied the name to plants of P. grandifolia, following earlier illustrations under this name (Edwards, Bot. Mag. 17: pi. 1473; Bot. Mag. 63: pi. 3478, and Pfeiffer & Otto, 1843: pi. 30), the confusion has persisted in botanical collections, en­ cyclopedic and pictorial works, e.g., in Tropica, p. 238 upper right (Graf, 1978), and Exotica (ed. 4), p. 622, upper left (Graf, 1982). Pereskia panamensis was described by Weber (1898a) as having yellowish-white petals with pink margin, but no further material with such flower colors has been found. Even if the type specimen had whitish flowers, all other characters perfectly agree with P. bleo. Cutak (1951) described P. corrugata from a plant originally received from the Bureau of Plant Industries, U.S. Department of Agriculture at Glenn Dale, Mary­ land, saying that "the meager data which accompanied the cutting proved erro­ neous" but unfortunately without stating what these were. Apparently Cutak did not know the true P. bleo and hence believed himself to have found a new species. The undulate, crinkled leaves are typical for living specimens of P. bleo, but the character is not as evident in older leaves and certainly not in herbarium speci­ mens. Britton and Rose (1919) did not note this leaf character but were the first to resolve the confusion between P. bleo and P. grandifolia.

12. Pereskia nemorosa Rojas Acosta, Cat. hist. nat. Corrientes 64. 1897 {"Pa- reskia nemerosa"); Flora del Chaco Austral 31. 1907 (orthography corrected). Type. Argentina. Corrientes, no specimen cited. Neotype designation. Corrientes: Paso de los Libres, 10 Mar 1917 (fl, fr), Shafer 127 (neotype, NY; isotypes, K, MO, US). Figs. 5G, 6A, I, 11B, 14B, 15Ad, 16, 41A-I, 42. Pereskia amapola F. A. C. Weber in Bois, Diet. hort. 938. 1898. Type. Paraguay. Asuncion, Mar 1877 (fl), Balansa 2671 (holotype, P; photo, GH). Specimen not cited with the description but mentioned in a letter from Weber to Weingart dated 6 Aug 1902 (Weingart, 1904: 85). Pereskia argentina F. A. C. Weber in Bois, Diet. hort. 938. 1898. Type. Not cited. Description based on young, sterile plant grown from seed from Argentina. Pereskia amapola F. A. C. Weber var. argentina F. A. C. Weber ex Weingart, Monatsschr. Kakteenk. 14: 87. 1904. Type. Not preserved. Description based on living plant grown from seed collected by Grosse in Paraguay, ca. 1899. 104 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 41. Pereskia nemorosa. A, flowering twig. B, flower, l.s. C, fruit. D, fruit, l.s. E, seed. F, G, leaf variation. H, areoles of trunk. I, habit in open condition. A, B from cult. hort. Berol. 260-02-80- 80 (B); C from Krapovickas 22229 (CTES); D from Gonjian s.n. (B); E from Herter 82375 (B); G, H from Shafer 137 (MO); I from Shafer 117, photograph (NY). 1 ^ 8 6] PERESKIA (CACTACEAE) 10 5

Small tree or shrub, 3-5(-8) m high; trunk to 10 cm in diam. or more; bark smooth, brownish-gray; branches and twigs green, olive, or yellowish-green, drying yellowish-brown; periderm formation beginning as pale brown longitudinal stria- tions contrasting with the dark green epidermis; twigs ca. 5 mm thick, straight. Areoles on twigs rounded, 3-8 mm in diam., with dense, gray to light brown wool; trichomes ca. 3 mm long; areoles producing spines, and 1-3 brachyblast leaves attached in the center of the areole. Leaves obovate to narrowly elliptic-obovate, more than twice as long as broad, 4-14(-16) x 2-6(-7) cm; petiole 0-5 mm long and to 4 mm thick when fresh, gradually merging into the blade; leafblade fleshy, ca. 0.8 mm thick; base narrowly cuneate to attenuate; apex acute; venation pinnate; midrib prominent below, slightly impressed above; lateral veins (5-)6-10(-14), diverging at angles of (35-)40-60°. Spines 0-5 per areole on twigs, fasciculate to spreading, 0.5-5 cm long and 0.5-1 mm thick, acicular to subulate, basally flat­ tened just above the thickened point of attachment; spines on branches 1-12 per areole, 1-6 cm long, on trunk 15-20 or more, (2-)4-6 cm long and to 2 mm thick. Inflorescence terminal, cymose-paniculate by proliferation from the receptacle, 2-5(-8)-flowered, or flowers solitary; lateral peduncles and pedicels 1-4 cm long, with 0-2 small bracts; bracts subtending the pedicels and peduncles resembling normal leaves but smaller, ca. 3-6 cm long. Flowers perigynous, 4-7(-9) cm in diam. when fully expanded. Receptacle turbinate, 12-15 mm in diam., podaria somewhat prominent, areoles 4-6 mm in diam., densely woolly, rarely with 1-2 short spines, the upper ones transverse-linear, with numerous coarse hairs of (6-) 10-15 mm length, the hairs brown or white with brown base, often nearly covering the bud in dried specimens; lower bracts leaf-like, 3-9, spreading, elliptic to lanceolate, with broad cuneate base, ca. 25-40 x 7-12 mm or smaller; upper bracts 3-6, erect, triangular acute or acuminate with a dark point, 4-8 x 3-4 mm, the outer ones narrowly triangular, the inner broadly triangular and shorter; sepaloids 4-5 (-7), lanceolate to ovate or obovate, softly mucronate, pale green or green with pink margins, 20-35 x 10-15 mm, somewhat cucullate when fresh; petaloids 6-12, obovate, softly mucronate or retuse to emarginate, 25-45 x 15- 28 mm, pink, base narrow, cuneate, pale pink to white; venation pseudopalmate. Staminodial hairs present at and between the bases of the inner petaloids, forming tufts of glassy white (when fresh), curly hairs, the tufts less than half as long as the filaments (Fig. 14B). Stamens more than 100, erect, grouped around but shorter than the style, 8-15 mm long; filaments white; anthers dorsifix, 1.2-1.5 x 0.6- 0.7 mm, often not or incompletely dehiscent and with clumped pollen. Ovary superior to half inferior, locule with flattish floor and ogival ovary roof with conspicuous septal ridges extending to near the bottom of the locule; placentation parietal along the base of the septal ridges; style 15-23 mm long, 1-2 mm thick, broadened at base into the free part of the ovary of ca. 4 mm diam.; stigma lobes ca. six, 3-4 mm long and 1 mm thick, white, often brown in dried or liquid- preserved specimens. Ovules numerous, ca. 0.8 mm long. Fruit solitary or clus­ tered, pyriform to broadly pyriform but variable in shape and size, often slightly angled, (3-)4-6 cm long and (2.5-)3.5-5 cm thick, green to yellowish-green when mature, areoles with short gray tomentum, rarely with 1-2 short spines of 5-10 mm length; bracts accrescent on young fruit but usually deciduous in mature fruits; umbilicus ca. 5-10 mm in diam., filled with the crumpled flower remnants and the coarse hairs of the upper receptacular areoles; fruit wall ca. 1 cm thick; locule concave to convex at the bottom, funicles thick, green, arising from the parietal placentae in the lower half of the locule. Seeds to ca. 50, obovate, 4.5-5 mm long, 3.0-3.6 mm broad and 2.0-2.5 mm thick, smooth, black, glossy, with 106 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 42. Distribution of Pereskia nemorosa and P. sacharosa. prominent micropylar nose; hilum lunate to semicircular, with distinct rim, ca. 1 mm in diam., cream. Seedlings (three weeks old) with hypocotyl ca. 5 cm long and 2 mm thick, green to olive, cotyledons ca. 30 x 15 mm, ovate-elliptic, with rounded apex, pinnate venation, lateral nerves 2-4. Juvenile plants with obovate, apically rounded leaves and areoles with 5-13 spines, 5-15 mm long. Chromosome number: In = 22; Esser 14763, cult. hort. Berol. 039-05-77-30 (B), and cult. hort. Berol. 090-01-79-80 (B). Distribution (Fig. 42) and phenology. Southern Brazil, Paraguay, northeastern Argentina and northwestern Uruguay. In drier parts of the forest and shrubland, in "high woodland" (humid semideciduous forest), in secondary vegetation and cultivated in hedges. Main flowering period is from October to March; fruits mature from November to April. According to Esser (1982, p. 69), the leaves are partly deciduous from May to August.

Specimens examined. BRAZIL. Rio GRANDE DO SUL: Uruguayana, 6 May 1927 (fl), Langeron s.n. (P). PARAGUAY. AMAMBAY: Parque Nacional Cerro Cora, Solomon et al. 6918, cult. hort. Berol. (B). CENTRAL: Barranca de Asuncion, 30 Jan 1908 (fl), Hicken 317 (SI); Asuncion, Recoleta, 6 Oct 1893 (fl), Lindman s.n. (S); Asuncion, 2 Nov 1888 (fl), Morong 188 (BM, E, GH, K, MO, NY, UC, US). CONCEPCION: Concepcion, Dec 1909 (st), T. Rojas s.n. (BAF). CORDILLERA: Between San Bernardino and Altos, 17 Oct 1981 (fl), Schinini & Bordas 21326 (SI). ITAPUA: Trinidad, 17 Mar 1917 (fr), Shafer 137 (G, GH, K, MO, NY, US). SAN PEDRO: 15 km SW Lima, Estancia Carumbe, 1966, Esser 14763, 1986] PERESKIA (CACTACEAE) 107 cult. hort. Berol. 090-01-79-30 (B); Alto Paraguay, Primavera, 17 Nov 1957 (fl), Woolston 910 (B, C, K, NY, S, U, UC). WITHOUT LOCALITY: Hassler 1326 (G). ARGENTINA. CORRIENTES: Dept. Bella Vista, Bella Vista, barranca del Rio Parana, 13 Oct 1974 (fl), Schinini & Cristobal 9899 (CTES); Dept. Mercedes, vie. Mercedes, Nov 1936 (fl), Rodrigo 802 (NY); Dept. Santo Tome, Santo Tome, 1 Feb 1976 (fl), Krapovickas & Cristobal 29165 (AAU). ENTRE Rios: Concordia, 6 Mar 1917 (fl, fr), Shafer 117 (K, MO, NY, US). MISIONES: Posadas, 30 Apr 1944 (fl), Bertoni 14 (LIL). URUGUAY. ARTIGAS: Sta. Rosa de Cuareim, 50-100 m, Nov 1927 (fl, fr), Herter 82375 (B, F, G, GH, GOET, HBG, LE, LIL, M, MO, NY, S, SI, U, UC, US). Local names and uses. Paraguay. Amapola, mori, suruby-i. Argentina. Ama­ pola, quisca del bosque, tuna quisca (Corrientes). Locally cultivated as a hedge and reported to form impenetrable stands due to the prohibitive armament of the trunk and branches. Several collectors note that the species is grown as an ornamental, apparently within its natural range of distribution, e.g., in Corrientes {Curran s.n.) and in the Dept. Mburucuya, on estancia Santa Teresa {Krapovickas & Mroginski 22229). According to a note on a herbarium label with a specimen {Arenas 1776) collected in Paraguay, Dept. Central, Ita, the leaves are sold in local markets for medical purposes. This species, better known hitherto as Pereskia amapola, has repeatedly been confused with P. sacharosa and P. grandifolia, but Weingart (1904) rightly rec­ ognized it as distinct, noting important key characters like "hairs between the petals and stamens," large flower size and leaf shape, while Schumann (1898) had also recognized P. amapola, but used key characters which it actually shares with P. sacharosa. The appropriate name P. amapola is unfortunately antedated by the poorly documented but still valid older name given by Rojas Acosta. Despite the poor and general description there can be no doubt on the identity of P. nemorosa, as there is no other pink-flowered species of Pereskia in Corrientes. Pereskia nemorosa is very closely related to P. sacharosa, but their ranges are distinct (Fig. 42). Further data on the distribution of the two species in Paraguay and Brazil seem desirable. Superficial similarities with P. grandifolia exist also, particularly in habit, habitat, leaf size and leaf shape, but flower characters, smaller seed size and lack of stomata on the stem distinguish it. A good illustration of P. nemorosa, unfortunately mislabeled as P. grandifolia, is given by Benson (1982, p. 268, fig. 237). Pereskia nemorosa is not only remarkable for having the largest flowers within the genus but also for the staminodial hairs and the frequently aberrant anthers and clumped pollen observed in nearly all cultivated plants as well as in most field collected specimens. Seasonal differences in leaf shape already noted by Weingart (1904) speak against justification of a distinct variety with narrow, acute leaves {P. amapola var. argentina), although different clones may exist. However, remarkable variability of leaf shape is observed in cultivation in this species and P. sacharosa. A fragmentary and sterile specimen labeled by Ritter as "FR 640 cf." from "Cerro Membuy" (Cerro Memby, Paraguay) most likely belongs here, notwithstanding that "FR 640" is the type collection of P. sparsiflora, a synonym of P. sacharosa.

13. Pereskia sacharosa Grisebach, Abh. Konigl. Ges. Wiss. Gottingen 24: 141. 1879. Type. Argentina. Salta: Oran, frequens in sepibus, 1873, Lorentz & Hieronymus 410 (holotype, GOET). Figs. 10, 15Ae, E, 17C, 19C, 42, 43A-J. Rhodocactus sacharosa (Grisebach) Backeberg, Kakteenlex. 397. 1966. Pereskia moorei Britton & Rose, Cactaceae 1: 15, fig. 8. 1919. Type. Brazil. Matto Grosso: Corumba, Tannay, 1891-92, Moore 955 (holotype, BM; drawing, NY). Pereskia saipinensis Cardenas, Cactus 19 (80-81): 17, with figs. 1964. Type. Bolivia. Santa Cruz: 108 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 43. Pereskia sacharosa. A, B, flowering twig. B, exposing receptacle, bracts and sepaloids. C, flower, l.s. D, fruit. E, fruit, l.s. F, seed. G, young stem with auxoblast leaves. H, stem with brachyblast leaves. I, J, leaf variation. A-C from cult. hort. Berol. 001-15-74-80 (B); D-F from Ahlgrimm 41 (B)- G, H, J from Ritter 640, cult. hort. Berol. (B); I from West 8359 (UC). 1986] PERESKIA (CACTACEAE) 109

Prov. Caballero, nr. Saipina, 1500 m, Nov 1962, Cardenas 6122 (Herb. Cardenasianum, specimen not located). Rhodocactus saipinensis (Cardenas) Backeberg, Kakteenlex. 397. 1966. Pereskia sparsiflora Ritter, Kakteen in Siidamerika 2: 482, fig. 330. 1980. Type. Bolivia. Tarija: Villa Montes, 1958, Ritter 640 p.p. (holotype, U). Small tree to 5(-7) m high, or shrub 2-4 m high; twigs 5-7 mm thick, often slightly zigzag, green or olive to pale brown when fresh, brown, stramineous, reddish or gray when dried; periderm formation initiating in rounded spots below the areoles. Areoles 4-7 mm in diam. on twigs, densely and very short-tomentose with trichomes 1.5-2 mm long, light gray to pale brown, cell walls uniform in color; areoles accrescent on branches and trunk to ca. 12 mm diam. and 5-8 mm height, producing spines and brachyblast leaves during the first few years, later producing additional spines only; brachyblast leaves mainly formed in the adaxial side of the areole. Leaves very variable in shape and size, mostly obovate, flat or folded along the midrib and hence appearing boat-shaped, 3-12 x (1—)2—7 cm, usually about twice as long as broad or less, but brachyblast leaves on strong shoots and in cultivation often longer and narrower; petiole 2-8 mm long and 3(-5) mm wide, or inconspicuous in fresh leaves, leafblade ca. 1 mm thick, fleshy; base attenuate, apex rounded and often with a short triangular point, or broadly acute, rarely acuminate, venation pinnate, rather inconspicuous in fresh leaves; midrib thick and prominent below in fresh leaves; lateral veins 4-6(-9), thin, rather inconspicuous in fresh leaves, diverging at angles of 25-40°. Spines 0-5 per areole on twigs, fasciculate to spreading, the lateral ones often deflexed on main shoots, very stout, subulate, 1—4(—8) cm long and 0.5-1.5 mm thick, on branches 1-8 per areole and 1-6 cm long and to 2 mm thick, on trunk to 25 per areole and 3-5 cm long and ca. 2 mm thick, often flattened at base and conically thickened at the point of attachment; brown to black, red at base in new growth, becoming gray with age. Flowers solitary, terminal, or in a terminal 2-4-flowered cymose-paniculate inflorescence by proliferation from the receptacular areoles. Flowers perigynous, 3-7 cm in diam. when fully expanded; receptacle turbinate with slightly prominent podaria, 10-15(-20) mm in diam., areoles 3-4 mm in diam., with abundant short gray tomentum, the upper ones with wool and nu­ merous silky to coarse, white or brown hairs 10-15 mm long (particularly con­ spicuous in bud); lower receptacular bracts 7-14, variable in size, obovate to lanceolate or linear-lanceolate with broad base, 15-40 x 5-15 mm, green, often finely lined with red at the margin; upper bracts erect, narrowly to broadly tri­ angular, to ca. 1 cm long and broad, acute; sepaloids 3-5, ovate-lanceolate to obovate, ca. 15 x 6 mm, acute, pale green to cream or pink towards the margin; petaloids 10-13, obovate to narrowly obovate, 25-35 x 15-20 mm, base cuneate, apex broadly acute to rounded with a point or emarginate, pink (but variously described by collectors as pink, rose-colored, mauve-pink, rose-purple, lilac, or deep pink), pale pink to white at base; stamens ca. 300, erect, clustering around the style, 6-15 mm long, shorter than the style; filaments white, pale pink towards the tip; anthers 1.3-1.5 x 0.5-0.6 mm, yellow; pollen mostly 9-12-colpate, me­ dium-sized, tectum rather densely and coarsely perforate and with medium to large spinules; ovary half inferior, with conical free roof and style base, unilocular, with shallow locule floor, roof with septal ridges; placentae basal-parietal at the lower end of the septal ridges; style 13-18 mm long, 1-1.5 mm thick, white; stigma lobes 7-11; ovules numerous, ca. 0.5 mm long. Fruit pyriform to subglobular or somewhat angled, 4-5 cm long and in diam.; areoles ca. five, semicircular to transverse-linear, with short gray tomentum; umbilicus narrow, filled by the flower remnants (unless fallen) and the persistent coarse hairs of the upper receptacular 1 10 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN LVOL- 41 areoles; bracts often accrescent on young fruit but usually deciduous at maturity; wall fleshy, green or yellowish; locule hollow or nearly filled by the seeds and thick funicles at least in the upper part. Seeds numerous, to ca. 50 or more, obovate to broadly obovate, 3.2-4.2 mm long, 2.4-3.1 mm broad and (1.5-)1.8- 2.2 mm thick, smooth, black, glossy, with prominent micropylar nose and subbasal semicircular to reniform hilum; hilum cream-colored, with low rim and central funicle scar. Seedlings (three weeks old) with hypocotyl ca. 25-45 mm long and ca. 1.6 mm thick, reddish; cotyledons ca. 30 x 15 mm, ovate-elliptic; primary leaves obovate. Chromosome number: In = 22; Ritter 640, cult. hort. Berol. 038- 02-77-30 (B). Distribution (Fig. 42) and phenology. Southeastern Brazil, Bolivia, Paraguay, and northwestern Argentina; mainly in lowlands along the eastern slope of the Andes and in the Chaco and Matto Grosso region, in low, open dry forest, to ca. 1000 m in Argentina and ca. 2500 m in Bolivia, frequent in northwestern Ar­ gentina, but range and frequency insufficiently known in the northern and north­ eastern distribution areas; main flowering period from October to February, fruits maturing from January to May; leaves deciduous in May and June (according to Hosseus, 1939).

Specimens examined. BRAZIL. MATTO GROSSO: Corumba, 1908, Hoehne 4863 (NY, US); Co- rumba, Feb 1911 (fl), Hoehne 3859 & 3860 (R); S. Anna da Chapada, 2 Nov 1902 (fl), Robert 714 (BM); on rd. Corumba-Baia Jacadingo, 19°12'S, 57°51'W, 11 Oct 1980 (fl), Paschoal 17 (HRB). BOLIVIA. WITHOUT LOCALITY: "Perou" (between 1747 and 1755), /. de Jussieu s.n. {Herb. A. L. Jussieu 13.547) (P-JUSS). BENI: Rurrenabaque, 1500 m, 10 Jul 1921 (fl), Rusby 1557 (NY, US); Rurrenabaque, 10 Dec 1930 (fl), Fleischmann 512 (S); Baures, Itinez, 29 Nov-25 Dec 1947 (fl), Scolnik & Luti 667 (SI). CHUQUISACA: Between Sucre and La Palma, 2380 m, Feb 1979 (fl), Ceballos et al. Bo 356 (SI). COCHABAMBA: Nr. Mizque on Rodeo rd., 2600 m, 6 Sep 1966 (fl), Badcock831 (K); Tako Tako, farm nr. Miske [Mizque?], 2100 m, 23 Dec 1949 (st), Brooke 5929 (BM); Rio Mizque, Ritter 640 p.p. (SGO). SANTA CRUZ: 14 km E of Comarapa, May 1981 (fr), Ahlgrimm 41 (B); 14 km S of Camiri, May 1981 (fr), Ahlgrimm 68 (B); 20 km NW of Boyuibe, May 1981 (st), Ahlgrimm 74a, cult, hort. Berol. (B); "Santa Cruz," 400 m, 12 Nov 1949 (fl), Brooke 5790 (BM); "Santa Cruz," 422 m, 1 Oct 1937 (fl), Gilson 1517 (BM). TARIJA: Between Canada and Palos Blancos, 25 Oct 1980, Zuloaga et al. 1366 (SI); Entre Rios, Ritter 640 p.p. (SGO). PARAGUAY. BOQUERON: Mision Santa Rosa, 21°45'S, 61°35'W, Nov 1981 (fr), Arenas 1776 (B, SI). CHACO: Cerro Leon, 3 Oct 1979 (fl), Schinini & Bordas 17817 (CTES), Schinini 17813 (B, SI); Mayor Pedro Lagerenza, 10 Apr 1978 (fr), Schinini & Bordas 14964 (CTES, SI); Agua Dulce, 5 Oct 1979 (fl), Schinini & Bordas 18067 (CTES). ARGENTINA. CATAMARCA: Dept. Capital, Las Rejas, 28 Oct 1973 (fl), Ulibarri 496 (B, SI). CORDO­ BA: Altos Sud, 8 Jan 1899 (fl), Stuckert 6068 (G). FORMOSA: Without locality, cult, at Bot. Gard. Agricult., Buenos Aires, 19 Nov 1938 (fl), Eyerdam & Beetle 23035 (G, UC). JUJUY: Without locality, Humbert 21157 (P, US), Schuel 163 (W); Dept. Capital, between Jujuy and La Mendieta, 800-1000 m, 24 Oct 1948 (fl), Killip 39578 (US), Killip 39580 (US); Dept. El Carmen, Pampa Blanca, Kiesling s.n. (B); Dept. El Carmen, Perico, 6 Jan 1971 (fl, fr), Krapovickas & Cristobal 17505 (CTES); Dept. El Carmen, Maquinista Veron, 800 m, 28 Nov 1949 (fl), Araque & Barkley 19Ar517 (K, LIL); Dept. El Carmen, on rd. no. 34, 6 km from Pampa Blanca, 2 Feb 1975, Zuloaga & Deginani 406 (SI); Dept. Ledesma, 23 Feb 1940 (fr), Burkart & Troncoso 11424 (SI); Dept. Ledesma, way to Higueritas, 23 Nov 1980, Cabrera et al. 32259 (SI); Dept. Ledesma, nr. Ledesma, 24 Oct 1948 (fl), Fries s.n. (S); Dept. Ledesma, Fraile Viejo, 400 m, 10 Oct 1932 (fl), Pastrana 9135 (LIL); Dept. Ledema, Calilegua, 22 Jan 1917 (fr), Shafer 59 (G, K, MO, NY, US); Dept. San Pedro, San Pedro, 7 Jan 1957 (fr), Burkart 20135 (SI); Dept. San Pedro, nr. San Pedro, 5 Jan 1978 (fr), Kiesling 1653 (B); Dept. San Pedro, along Rio San Pedro, W of La Mendieta, 800 m, 16 Feb 1937, West 8359 (UC); Dept. Santa Barbara, way from San Lucas to Santa Clara, 12 Nov 1978, Cabrera et al. 29817 (SI). SALTA: Without exact locality, Dec 1878, Grisebach s.n. (K); Dept. Anta, Joaquin V. Gonzalez, 9 Nov 1974 (fl), Krapovickas et al. 26709 (CTES); Dept. Jose de San Martin, ca. 15 km E of Hickmann, Pozo El Milagro, 10 Dec 1972 (fl), Maruhak et al. 575 (CTES); Urundel, 750 m, Nov 1930 (fl), Kanter 111 (HBG); Campo Santo, 18 Apr 1873 (fr), Lorentz & Hieronymus 524 (GOET, US); Cobos nr. Salta, 1 Mar 1873 (fr), Lorentz & Hieronymus 562 (LE); Campo Santo, 20 Nov 1933 (fl), Peirano s.n. (LIL); Oran, way to Mision Zenta, 380 m, 17 Mar 1974 (fr), Tiirpe et al. 4899L (LIL); Oran, Abra Grande, 750 m 17 Nov 1927 (fl), Venturi 5619 (GH, NY, US); Campo Santo, 800 m, 20 Apr 1941 (fl), Zabala 137 (LIL) TUCUMAW 1986] PERESKIA (CACTACEAE) 111

Dept. Capital, 450 m, Nov 1897 (fl), Lillo 2106 (LIL); Dept. Capital, Alto de la Polvora, 480 m, 4 Dec 1924 (fl), Venturi 3818 (LIL, NY, SI, US). Local names and uses. Argentina: sacharosa. Bolivia: cuguchi and guyapa (Santa Cruz), according to Pflanz (Vaupel, 1925, p. 13). In Bolivia and Argentina the plant is used for hedges (Bailey, 1963a, p. 130; Boelcke, 1981, p. 340). The oldest specimen of P. sacharosa, collected by J. de Jussieu in "Perou" more than a century before the type specimen and description of the species, apparently remained unnoticed—labeled as Cactus pereskia—hitherto. Jussieu stayed in Bolivia (at that time part of the viceroyalty of Peru) from 1749 to 1755, according to Lacroix (1936), and travelled to places like Chuquisaca and Santa Cruz, areas in which P. sacharosa occurs. Three taxa described later prove to be conspecific with this species. Weak spines occasionally occur on the receptacular areoles and this character, observed on a single specimen, was used by Britton and Rose (1919) to segregate a new species, P. moorei from Corumba, Brazil, a locality then thought to be far away from the distributional range of P. sacharosa. The type specimen of P. saipinensis could not be located, but from the description, illustration, and origin of the plant there is no doubt that it is conspecific with P. sacharosa. Plants distributed by Knize under the name P. saipinensis, however, are P. diaz-romeroana, a plant also growing in the area of Saipina, Bolivia. For P. sparsiflora, Ritter (1980, p. 482) mentioned "flowers solitary" as a main di­ agnostic character delimiting this species from P. sacharosa, but he erred in his statement that, according to the original description, P. sacharosa has branched inflorescences. Grisebach (1879) does not mention the nature of the inflorescence but merely states "floribus subsessilibus." Contrary to Ritter's observation, a plant raised from seed collected by Ritter (FR 640) and seen by the author at the Jardin Exotique in Monaco in 1980 produces few-flowered inflorescences as well. Vari­ ability of this character is observed in cultivated plants of P. sacharosa. The type collection of P. sparsiflora ("FR 640 from Villa Montes"), a fragment without flowers and leaves, and cultivated plants raised from "FR 640" seed (not nec­ essarily from the type locality due to Ritter's species-based numbering system) are indistinguishable from P. sacharosa. Ritter (1980, p. 482) mentioned the species also for Mairana and Mataral, Bolivia, and further specimens labeled as FR 640 were collected in Entre Rios, and Rio Mizque, Bolivia. Another example of Ritter's awkward numbering system is a fragmentary specimen labeled "FR 640 cf." at SGO from "Cerro Membuy." The locality is not cited in Ritter (1979, 1980) but it is probably Cerro Memby in Paraguay, Dept. Concepcion, and the specimen most likely is P. nemorosa. The epithet is derived from the local name "sacha rosa," an apparent Quechua and Spanish compound meaning "false rose" (C. Beer, pers. comm.), literally translated as "wild rose" (Vaupel, 1925, p. 13) or "woods rose," "jungle rose," or "tree rose" (M. Nee, pers. comm.). It is still often erroneously spelled "saccharosa" in horticultural literature.

14. Pereskia grandifolia Haworth, Rev. pi. succ. 85. 1819. Shrub or small tree, 2-5 (-10) m tall, with erect to arching main shoots; trunk to 20 cm in diam., bark grayish-brown, slightly rough; twigs 5-8 mm thick, after periderm formation yellowish-brown when dry, young twigs green or reddish when fresh and with conspicuous fine white spots (stomatal cavities) not visible in dried material. Areoles rounded, cushion-like, 3-7(-8) mm in diam. on twigs, to 12 mm in diam. on trunk, 2-4 mm high, with dense and short brown, grayish-brown or 1 12 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 gray tomentum of uniseriate multicellular hairs 1-2 mm long, producing spines and l-3(-4) brachyblast leaves, growth zone in the center or in the upper part of the areole. Leaves variable in size, elliptic or narrowly elliptic-oblong, narrowly ovate to obovate-lanceolate, (6-)9-23(-30) x (3-)4-6(-9) cm; petiole (3-)5-12 (-15) mm long and 1-2 mm broad (to 3-4 mm thick when fresh); blade fleshy, 0.4-0.7 mm thick, base attenuate, apex acute to acuminate, plane or recurved; venation pinnate, prominent below; lateral veins (6-)7-l 3(-l 6), ca. 10 in medium- sized leaves, diverging at angles of 40-60°, mostly straight but somewhat arching and reticulate towards the margin. Spines 0-8(-ll) on twigs, fasciculate (when young) to spreading, 1-4 cm long and 0.5-1.0 mm thick, acicular to subulate, black, brownish-black or brown with darker tip, red or yellowish at base in new growth, increasing in number on older branches to 10-15 or more, 2.5-6 cm long, on main shoot and trunk to 25-90 per areole, 2-6.5 cm long and 0.8-1.5 mm thick, brownish-black to black, becoming gray with age. Inflorescence terminal, densely cymose-paniculate by proliferation from the receptacles; additional lateral inflorescences developing from the upper areoles of the main shoots and twigs, usually ca. 10-15-flowered, but sometimes to 30(-50) flowers on strong shoots; lateral peduncles and pedicels 3-9, the upper ones 1-3 cm long, leafless or with 1-2 leafy bracts; terminal flower without a distinct pedicel, the axis broadened into the receptacle. Flowers 3-5(-7) cm in diam., showy, rose-like, lasting 1-2 days; receptacle 5-11 mm in diam., turbinate, usually with conspicuous angles or furrows and raised podaria, green, with bracts and areoles in the upper half only; areoles woolly, with hairs ca. 1 mm long; lower bracts ca. 4-7, spreading to ascending, variable in shape and size, small or leaf-like, elliptic-ovate to narrowly obovate-lanceolate, 4-30 x 4-15 mm, green or purplish, shortly and broadly petiolate, blade plane or keeled below; apex acute to acuminate, flat or recurved; upper bracts 2-5, erect, appressed to the bud, ovate to obovate-lanceolate or broadly triangular, to ca. 5 x 5 mm, with broad base and acute apex; sepaloids 2-5, ovate to obovate-lanceolate, often asymmetric, 8-18 x (3-)5-8(-12) mm, greenish to pink or purplish, with broad, cuneate to attenuate base; petaloids 5- 12, obovate, to spathulate, 10-32 x 5—13(—17) mm, accrescent during anthesis, pink, rose, pale pink, or purplish-pink; base cuneate, broad to narrow, paler; apex rounded with a point to deeply emarginate; stamens ca. 110-130, 5-10 mm long, clustering around the style below the level of the stigma lobes; filaments white; anthers 1.0-1.8 x 0.5-1.0 mm, pale to golden-yellow; pollen mostly 6-9(-12)- colpate, medium-sized, tectum densely and coarsely perforate; spinules ca. 1 /*m; ovary distinctly inferior but with free conical ovary roof narrowing into the style; locule cup-shaped, ca. 3-4 mm in diam., locule floor concave, ovary roof with septal ridges; placentae parietal, in the upper part of the locule around the base of the septal ridges; style 8-12 mm long, white; stigma lobes 5-8 suberect, 1-2 mm long, white to pale pink. Infructescence a cymose-paniculate cluster, usually pendent, dense to lax. Fruits 1—12(—30), pyriform or turbinate, usually with con­ spicuous angles formed by the podaria of the receptacular bracts, very variable in shape and size, 5-10 cm long and 3-7 cm in diam., green or reddish-green to yellowish at maturity; pedicel 0-20 mm long, areoles 4-5(-8), with grayish to­ mentum; bracts small to leafy, persistent or mostly deciduous at maturity; um­ bilicus constricted, narrow, filled by the upper bracts and the crumpled flower remnants, 4-9 mm in diam.; locule in the upper half of the fruit, cup-shaped to obtriangular; fruit wall 7-12 mm thick, ovary roof to 12 mm in diam. and ca. 5- 7 mm thick. Seeds ca. 20-60, obovate-elliptic, (5.0-)6.0-6.7(-7.0) mm long, (3.6-) 4.5-5.0(-5.4) mm broad and 2.5-3.2 mm thick; smooth, black, glossy; hilum subbasal, creamy white, reniform, with low rim, micropylar end rounded to nose­ like. Seedlings (three days old) with hypocotyl 15-40 mm long, cotyledons ca. 10 1986] PERESKIA (CACTACEAE) 113

x 8 mm, ovate, acute, accrescent on seedlings with developing primary shoot to 30-40 x 22 mm.

Key to the Varieties of Pereskia grandifolia 1. Receptacular bracts green; sepaloids greenish to pink; lower receptacular bracts rarely with recurved apices; petaloids 15-33 mm long; anthers golden-yellow. 14a. var. grandifolia. 1. Receptacular bracts and sepaloids bright purplish-pink to dark purplish; lower receptacular bracts with recurved apices; petaloids 10-18 mm long; anthers pale yellow. 14b. var. violacea.

14a. Pereskia grandifolia Haworth var. grandifolia. Figs. IE, 5F, G, 6G, H, 11J, 13A-F, 14F, 15Aj, 17D, 44A-H, 45. Pereskia grandifolia Haworth, Rev. pi. succ. 85. 1819. Type. "Vigebat in regio horto Kewense A. D. 1818, sub nomine Cacti Portulaccaefolir (description based on a cultivated sterile plant). Neotype designation. Cult. hort. Kew. "raised from seed collected by Bowie & Cun­ ningham in 1816 in the neighbourhood of Rio de Janeiro," drawing by T. Duncanson, June 11, 1824 (K,ic). Cactus grandifolius (Haworth) Link, Enum. pi. hort. Berol. 2: 25. 1822. Rhodocactus grandifolius (Haworth) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Pereskia ochnocarpa Miquel, Bull. Sci. Phys. Nat. Neerl. 1: 48. 1838. Type. Brazil. Illustration of Cactus rosa Vellozo (nom. nud.), Fl. flum. 5: pi. 27. 1827. Pereskia rosea Hort. ex A. Dietrich, Allg. Gartenzeitung 19: 152. 1851. Type. Cult, in Hort. Berol., received from F. A. Haage, Erfurt, a specimen of unknown origin. Not preserved. Pereskia tampicana F. A. C. Weber, Bull. Mus. Hist. Nat. (Paris) 4: 167. 1898. Type. Mexico. Tamaulipas: Nr. Tampico, at El Paso de Dona Cecilia on the Rio Panuco, ca. 1895, Heese s.n. (not preserved). Rhodocactus tampicanus (F. A. C. Weber) Backeberg, Cactaceae 1: 115. 1958. Shrub, 2-5 m high, or individual shoots becoming tree-like and to 10 m high. Leaves variable in shape and size, elliptic-oblong, elliptic or narrowly elliptic- lanceolate, shortly acuminate or acute, rather thin, ca. 0.4-0.5 mm thick when fresh, green, concolorous. Bracts on the inflorescence and receptacular bracts green, usually plane, rarely recurved; receptacular bracts shorter to longer than the peta­ loids; sepaloids greenish-pink to pink; petaloids 15-32 mm long, pink to deep pink; anthers golden-yellow. Fruit variable, pyriform to obtriangular, angled; often nearly as broad as long; locule cup-shaped to broadly obovate-obtriangular. Chro­ mosome number: In = 22; cult. hort. Berol. 047-01-78-80 (B), and cult. hort. Berol. 168-02-79-80 (B). Distribution (Fig. 45) and phenology. Eastern Brazil (even there insufficiently known), elsewhere probably subspontaneous after cultivation; from the lowland to ca. 1000 m, in forest, secondary forest, restinga, on waste ground and cultivated in hedges, also cultivated as an ornamental; flowering almost throughout the year (not observed in June, July, and November), fruiting observed in March and April, probably extending over a longer period. The natural range of this variety is insufficiently known, but it is widespread in cultivation as an ornamental both in and outside of Brazil and appears to be established in several countries of tropical America. Numerous collections are without indication of its native, spontaneous, or cultivated status. Where I regard them as spontaneous after cultivation or doubtful I have omitted them from the list of specimens. Such collections from Amazonian Peru, Bequia (island S of St. Vincent), Cuba, the Dominican Republic, Honduras, Puerto Rico, St. Croix, Trinidad, and Venezuela nevertheless represent conspicuous elements of the flora of these countries and they are included in the distribution map, marked accord­ ingly. Abundant collections of plants cultivated in gardens and parks exist from Costa Rica, Cuba, Florida, Guadeloupe, Guatemala, Haiti, Mexico (Chiapas, 114 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 44. Pereskia grandifolia var. grandifolia. A, flowering shoot. B, flower, l.s. with base of lateral bud. C, flower bud. D, shoot with auxoblast leaf (upper left) and three brachyblast leaves. E, infruc­ tescence. F, fruit. G, fruit, l.s. H, seed. A, F, G from cult. hort. Berol. 047-01-78-80 (B); B, C from Horst & Ubelmann HU492 (B); D, H from cult. hort. Berol. 038-04-77-80 (B); E from photograph of specimen cult, at Huntington Botanical Gardens. 1986] PERESKIA (CACTACEAE) 115

© 1979 by the U Department of Systematic Botany FIG. 45. Distribution of Pereskia grandifolia and its varieties.

Morelos, Oaxaca, Veracruz), Puerto Rico, St. Croix, St. Eustatius, St. Kitts, Trin­ idad, and Venezuela.

Representative specimens. BRAZIL. BAHIA: Mun. de Andarai, 12 Jan 1983 (fl), L. A. M. Silva et al. 1592 (B, CEPEC); Cruz das Almas, May 1973 (fl), Pinto s.n. (ALCB), Pinto 29/81 (HRB); Porto Castro Alves, Vale dos Rios Paraguacu e Jacuipe, 10 Dec 1980 (fl), Cavalo 1011 (ALCB, BAH, HRB); Mun. Porto Seguro, Monte Pascoal, on rd. to Caraiba, 14 May 1971 (fl), T. S. Santos 1641 (B, CEPEC); vie. Salvador de Bahia, 27 May 1915 (fl), Rose & Russell 19656 (NY, US); Mun. de Vitoria da Conquista, 20 km E Anage, 13 Apr 1983 (fr), Leuenberger et al. 3062 (B, CEPEC, K, SI, W). CEARA: Chapada do Araripe, 30 Aug 1971 (fl), Gifford & Fonseca G 335 (E). ESPIRITO SANTO: Cachoeiro, Bello 653 (R); Campo Novo, Neuwied s.n. (BR, MO). MATTO GROSSO: Cuiaba, Aug 1892 (fl), Kuntze s.n. (F, NY). Rio DE JANEIRO: Sertao de Cacimbas, on right embankment of Rio Itabapoana, 10 Oct 1909 (fl), Sampaio 985 (R); between Campos and Morro do Coco, 8 Aug 1964 (fl), Trinta 1051 (R); Cantagalo, Peckolt 121 (W); Monte Alegre, 595 m, Feb 1927 (fl), Vidal s.n. (R); Nova Friburgo, 8 Dec 1918 (fl), Curran 621 (F, GH, US); Mun. de Rio Claro, Manguaratiba, 400 m, 16 Mar 1978 (fr), Ferreira et al. 348 (RB); "Rio de Janeiro," Burchell 901, 1364, 1665 (K), Gaudichaud 912 (G, MO, P), Gay s.n. (P), Glaziou 18268 (K), Miers 3833 (K), Mosen 2865 (S), Riedel 89 (P), Weddel 350 (P); on the way from Leme to Botafogo, 8 Aug/Oct 1890 (fl), Glaziou 12868 (P, R); Boa Vista, Jan 1923 (fl), Freire & Vidal s.n. (R); Jacarepagua, Aug 1925 (fl), Lutz 15701 (R); Terezopolis, Boa Fe, 8 Apr 1943 (fl), Velloso 375 (R). SANTA CATARINA: Laguna, 19 Sep 1951 (fl), Reitz & Klein 3 (NY); SAo PAULO: Paraguacu Paulista, 1 km N on rd. to Bora, 500 m, 22°24'S, 50°35'W, 10 Feb 1965 (fl), Eiten et al. 6011 (K, NY, US); Saliceos-Novo Horizonte (Sales?), 800 m, 26 Sep 1982 (fl), Rauh 53389a (HEID). 1 16 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL- 41

Local names and uses. Brazil: sabonete (Espirito Santo), ora pro nobis (Rio de Janeiro), quiabento (Bahia). It is sometimes planted as a hedge and often cultivated as an ornamental. In Venezuela, where it is not native, it is called "guamacho morado" according to Hoyos (1978). There is no doubt that Duncanson's 1824 drawing of Pereskia grandifolia rep­ resents this species and variety, and that it probably shows the same plant used as a yet juvenile, sterile specimen by Haworth for the publication of the species in 1819, even though N. E. Brown annotated the drawing as "scarcely the plant described by Haworth." The seed collected by Bowie and Cunningham in 1816 represent the oldest known collection of the species, but apparently no herbarium specimen exists. The next oldest collections are those of Gay, Gaudichaud, and Riedel from between 1828 and 1833. These, as well as many later collections, are all from Rio de Janeiro. Schumann (1890, 1898) misinterpreted P. grandifolia as a synonym of P. bleo, and Weber (1898b), who was aware of the differences between the two species, nevertheless illustrated P. grandifolia with the wrong caption in Bois (1893-1899, fig. 678). Britton and Rose (1919) recognized the difference between the two species and noted that all previous illustrations with the caption "P. bleo" show in fact P. grandifolia. This species and variety is still found mislabeled as P. bleo in many collections and pictorial works. The spelling "P. grandiflora" is an equally widespread error. This name was mentioned by Pfeiffer (1837a, 1837b), is a nomen nudum for the same species, and is still found and erroneously attributed to Haworth even in recent publications (Hoyos, 1978, p. 208). For P. tampicana no type specimen is available; however, from Weber's description it seems evident that this name is a synonym of P. grandifolia var. grandifolia and not of P. zinniiflora as Bravo-Hollis (1978) suggested. All spec­ imens identified by previous workers as P. tampicana are apparently from cul­ tivated sources in Mexico and undoubtedly the same as P. grandifolia. Pereskia gigantea hort., illustrated by Hertrich (1956, p. 108, 109, figs. 68, 69) is a nomen nudum belonging here also.

14b. Pereskia grandifolia Haworth var. violacea Leuenberger var. no v. Type. Brazil. Minas Gerais: Mun. de Santana do Riacho, area do futuro Parque Estadual da Serra do Cipo, 650 m, 23 Sep 1981 (fl), F. C. F. da Silva 89 (holotype, HRB; isotype, B). Figs. 5J, 14D, E, 15Ak, 45, 46A-J. A var. grandifolia differt bracteis et foliis sepaloideis atropurpureis ad laete violaceo- roseis, recurvatis; foliis petaloideis minoribus, 10—15(—18) mm longis, variabile co- loratis, pallide roseis ad laete violaceo-roseis; antheris pallide flavis; fructu et loculo plerumque obtriangulari, umbilico angustiore, 4-5 mm lato. Shrub, ca. 2-4 m high; leaves narrowly elliptic-lanceolate to oblong; blade rather thick and fleshy, 0.6-0.7 mm thick when fresh, dark green, often slightly dis- colorous, matt below, nearly always distinctly acuminate and with recurved apex; spines variable in size and number, red at base when young; bracts all dark purplish to bright purplish-pink even in bud, the normally ca. five lower receptacular bracts very showy and resembling open flowers from a distance; lower bracts nearly as long as the petaloids, with acuminate, recurved apex, thick and keeled below; petaloids 10—15(—18) mm long, fleshy and thick at base, color of blade variable, pale pink to purplish-pink, pale at base; anthers pale yellow; fruit and locule usually narrowly obtriangular; bracts purplish to tinged with green or becoming 1986] PERESKIA (CACTACEAE) 117

Vereskla grandifolia var. violacea

FIG. 46. Pereskia grandifolia var. violacea (A-D, F-J from Horst & Ubelmann HU 226, cult. hort. Berol. 036-01-77-30, (B); E from a photograph published by Uil (1978, p. 262, fig. 5) of material from Buining, presumably of the same collection). A, inflorescence, lateral view. B, flowering shoot with terminal flower and flower buds. C, flower. D, flower, l.s. E, infructescence. F, fruit. G, fruit, l.s. H, seeds. I, stem. J, stem with brachyblast leaves. 1 18 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 nearly green; umbilicus narrow, ca. 4-5 mm in diam. Chromosome number: In = 22. Horst & Ubelmann HU 226, cult. hort. Berol. 036-01-77-30 (B). Distribution (Fig. 45) and phenology. Espirito Santo and Minas Gerais, Brazil; from ca. 600 to 1400 m, in the transition zone of forest and savanna (cerradao), also in secondary forest and locally cultivated; planted near houses in northeastern Minas Gerais and in southern Bahia, but apparently not native there; flowering from September to April, fruits observed in April; range of distribution and natural habitat insufficiently known.

Specimens examined. BRAZIL. ESPIRITO SANTO: SW of Colatina, Braun 476 (B, photo). MINAS GERAIS: Mun. Itabira, Caue, 11 Feb 1934 (fl), Sampaio 7068 (R); Itabira de Matto Dentro, 1386 m, 24 Jan 1922 (fl), G. Santos s.n. (R); 8 km W of Itaobim, ca. 6 km S of rd. Itaobim-Aracuai, near rural house, 9 Apr 1983 (fl, fr), Leuenberger & Martinelli 3057 (B, CEPEC, RB); Mun. de Marlieria, Parque Estadual do Rio Doce, 19°46'S, 42°36'W, 27 Sep 1975 (fl), Heringer & Eiten 15252 (UB, US). Local names and uses. Ora pro nobis (Minas Gerais). Planted as an ornamental in northeastern Minas Gerais and southern Bahia. This new variety was discovered by the commercial collectors L. Horst and W. Ubelmann before 1972 and introduced into European collections as Pereskia bahiensis HU 226. Uil (1978) received material of probably the same origin from Buining in 1974 and described and illustrated it as Rhodocactus grandifolius, while Leuenberger (1976a), Barthlott(1977,1979), and Rauh (1979), misidentified it as P. bahiensis again. It is definitely closer to the former species in all major characters, and dried material is often impossible to distinguish from P. gran­ difolia, while living plants are strikingly different due to the coloration of the bracts and flower buds. Smaller flower size than in well developed specimens of P. grandifolia seems to support delimitation of a separate taxon, but most other characters seem too weakly defined to give this plant specific status. It is best treated as a variety of P. grandifolia. The material distributed by Horst and Ubelmann as HU 226 seems to consist of two slightly different clones. One plant cultivated at Berlin, received from Horst with locality data "Minas Gerais, Ma- lacacheta, or Capelinha, Jaguarina" has a deeper tone of color of the petaloids and rather broad petaloids; the other received from Ubelmann under the same number as from "W of Teofilo Otoni, on the road to Itamarandiba" has narrower and pale petaloids and even darker purplish bracts. Since an original herbarium specimen with exact locality data is not available I have selected as type a more recent field-collected specimen accompanied by a note on the diagnostic color character. It is likely to be from the natural habitat of the variety and preferable to a cultivated specimen. My journey to Minas Gerais in 1983 in search of the plants seen by Horst and Ubelmann around Capelinha was unsuccessful, although the local curandero recognized the plant from photographs as existing in the area. Later I found an identical plant, thanks to enquiries of G. Martinelli, planted near a rural house in the hills southwest of Itaobim. It is also planted near houses on the main highway in northern Minas Gerais and in Vitoria da Conquista, Bahia {Leuenberger et al. 3060). Under experimental conditions this variety hybridizes with the typical variety. The first and only Vx hybrid specimen flowered so far is somewhat intermediate in the color of the bracts but identical with var. grandifolia in flower size and color of the petaloids. This demonstrates the close relationship but may also support its status as a variety rather than just a mere form. The above cited specimens collected by Sampaio, Santos, and Heringer and Eiten unfortunately lack notes on the color of the bracts and are included here with some reservations. 1986] PERESKIA (CACTACEAE) 119

15. Pereskia bahiensis Gurke, Monatsschr. Kakteenk. 18: 86. 1908. Type. Brazil. Bahia: Calderao, Oct 1906 (fl), Ule 7050 (lectotype, HBG; isotype, L). Figs. Ill, 15Ab, 21A-E, 47A-K, 48. Small tree or shrub, 1-6 m tall, with erect to arching main shoots; trunk to 30 cm in diam.; distal twigs stout, (3-)5-7 mm thick; green to reddish when fresh, with few stomata mainly around the areoles only; stramineous to grayish-brown after periderm formation; bark of trunk longitudinally fissured, grayish-brown. Areoles rounded, 3-6(-8) mm in diam. on twigs and main shoots, accrescent to ca. 10 mm diam. and rounded, or transverse-elliptic and to 20 mm diam. on trunk; tomentum short, dense, pale gray or brown when young; trichomes ca. 1.5- 2 mm long, with dark transverse cell walls, all uniseriate; areoles producing spines and l-3(-4) brachyblast leaves. Leaves variable in shape and size, elliptic or obovate, often narrowly obovate on main shoots, flat or folded upward along the midrib and recurving, (4-)5-12 x 2-7 cm, often ca. 7 x 5 cm; petiole indistinct or 1—2(—5) mm long thick and with a dorsal spur-like rim at the base when fresh; blade ca. 1 mm thick, fleshy, base attenuate; apex rounded to broadly acute, usually acute in narrow leaves; venation pinnate; midrib prominent and thick at the base below; lateral veins (3-)4-7(-8), rather inconspicuous in fresh leaves. Spines 0-6 per areole on distal twigs, fasciculate to spreading, (l-)2-5 cm long and 0.5-1.0 mm thick, often flattened towards the base, brown or brownish-gray, reddish at base when young, with darker, nearly black tip, becoming gray in age, increasing in number on branches and trunk to ca. 45, 2-6.5(-9) cm long and to 1.8 mm thick, gray to nearly black. Inflorescence cymose-paniculate by prolif­ eration from the receptacle, 2-12-flowered, to 10 cm in diam., dense to lax, or flowers solitary. Flowers perigynous, 4-7 cm in diam. when fully expanded, lasting one day only; receptacle 7-10 mm in diam., turbinate, with prominent podaria at least on the central flowers; pedicels ca. 1 cm long; areoles 3-8, mainly in the upper half of the receptacle, 3(-5) mm in diam., with short dense tomentum, the upper ones transverse-linear with a dense row of pale gray to grayish-brown hairs ca. 2-4 mm long and surpassing the margin of the erect upper bracts; lower bracts 4-6(-9), often calyx-like, spreading, obovate to elliptic, with broad base and acute to rounded apex, sometimes resembling regular leaves and to 5 x 3 cm on solitary flowers, but mostly smaller, 5-10(-20) x 5-7 mm, thick, fleshy, green, sometimes with reddish margin; upper bracts 4-6, erect, narrowly to broadly triangular or triangular-ovate, ca. 2.5-3 mm long and broad, thick, fleshy, green with purplish to nearly black margin, apex acute to mucronate, keeled; sepaloids 2-5, ovate to obovate, broadly acute, 6-10 x 6-12 mm, fleshy, greenish-red to reddish-purple; petaloids ca. 7-9, spreading to recurved, obovate to spathulate, 20-30 x 8-13(- 15) mm, pink to reddish-purple, pale at base; blade finely spotted with more deeply stained cells; base cuneate; apex rounded-truncate to emarginate, entire. Stamens ca. 160-200, 5-10 mm long, clustering around the style; filaments in 4- 5 rows, white; anthers yellow, ca. 1 x 0.3-0.5 mm. Pollen (9-)12(-15)-colpate, medium-sized, tectum with perforations of variable size without ring-like collar, spinules rather small. Ovary half inferior to inferior; locule ca. 3 mm in diam., with variously flat to concave floor and distinct septal ridges at the roof; placentae parietal around the base of the septal ridges; style (7—)10—12(—15) mm long and ca. 1 mm thick, white, broadened into the conical, free ovary roof of ca. 3 mm diam.; stigma lobes 5-10, 1-3 mm long, suberect, white. Ovules numerous, ca. 0.5 mm long. Infructescence a rather lax cymose-paniculate cluster of up to 12 fruits and to 15(-20) cm in diam., often pendent, or fruits solitary. Fruit pyriform 120 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 47. Pereskia bahiensis. A, twig with flower and solitary fruit. B, flower bud. C, flower, l.s. D, leaf shapes. E, fruit. F, fruit, l.s. G, infructescence. H, seeds. I, areole of trunk. J, habit. K, planted as a fence and hedge. A from Leuenberger et al. 3069 (photo, B). B, G from Leuenberger et al. 3063 (B); C from cult. hort. Berol. 001-13-74-70 (B). D from Leuenberger et al. 3086 (B). E, F, H from Leuenberger et al. 3071 (B). I, J from Rose & Russell 19786 (photo, US). K from photographs by the author at and near locality of Leuenberger et al. 3063. 1986] PERESKIA (CACTACEAE) 121

to turbinate and more or less angled, 3-6 cm long and 3-5 cm in diam.; peduncles and pedicels 6-15 mm long, usually shorter on distal fruits; bracts often present on immature fruits, variable in size, often leaf-like on solitary fruits, smaller and rounded to obovate, thick and fleshy on peripheral fruits of the cluster, usually deciduous at maturity; areoles 4-7, ca. 4 mm in diam., with conspicuous, dense short, gray tomentum; umbilicus narrow, 5-8 mm in diam., filled by the upper, connivent bracts and the crumpled flower remnant; fruit wall ca. 10-15 mm thick, with numerous mucilage cells and cavities, very sticky; locule obovate to obtrian- gular, 10-20 mm in diam., roof of the ovary flat below the umbilicus, to ca. 10 mm diam., ca. 7 mm thick; locule hollow, more or less filled by the seeds and the sticky remnants of the funicles, and in the upper part by mucilaginous tissue formed by the ovary roof. Seed numerous, to more than 30, obovate, 3.8-5.5 mm long, 2.8-3.5 mm broad and 1.9-2.8 mm thick, black, smooth, glossy; mi­ cropylar end rounded to nose-like; hilum reniform, 0.7-1.2 mm broad. Seedlings (three weeks old) with hypocotyl 30-55 mm long and 1.5-2.5 mm thick when fresh, reddish; cotyledons ovate-elliptic, 30-45 x 15-20 mm, base and apex rounded, blade green, concolorous, midrib faint, lateral veins 0-2. Chromosome number: In = 22; cult. hort. Berol. 001-13-74-70 (B). Distribution (Fig. 48) and phenology. Bahia, Brazil, in caatinga, from 400 to 900 m, also on disturbed ground and often planted for hedges, flowering from October to April, fruiting from March to June, leafless during the dry season from about June to October. Northern and western limit of distribution insufficiently known.

Specimens examined. BRAZIL. BAHIA: Mun. Boa Nova, 20 km S of Manuel Vitorino, 19 Apr 1983 (fr), Leuenberger et al. 3083 (B, CEPEC, K); Mun. Caetite, km 17 on rd. from Caetite to Brumado, 750 m, 14 Apr 1983 (fl, fr), Leuenberger et al. 3069 (B, CEPEC, K); Mun. Caetite, Brejinho das Ametistas, hedges at the village, 900 m, 15 Apr 1983 (fl, fr), Leuenberger et al. 3071 (B, CEPEC, K, SI); Mun. Irece, 26 Oct 1970 (fl), Costa s.n. (ALCB); Mun. Irece, on rd. to Presidente Dutra, 7 Oct 1980 (fl), Ferreira 118 (ALCB, BAH); Mun. Itaete, 19 Oct 1978 (fl), Almeida 28 (HRB, RB); Mun. Itaete, between Machado Portella and Bandeira de Melo, 19 Oct 1978 (fl), Araujo 103 (HRB, RB); Mun. Jaguarari, 64 km N of Senhor do Bonfim on highway to Juazeiro, ca. 400 m, 25 Feb 1974 (fl), Harley 16352 (CEPEC, K, RB); Barrinha, 7 Jun 1915 (fr), Rose et al. 19786 (NY, US); Mun. Jequie, 11-17 km W of Jequie on rd. to Lafayete Coutinho, 19 Nov 1978 (fl), Mori et al. s.n. (CEPEC); Mun. Joazeiro, Nov 1913 (fl), Zehntner 743 (M, US); Mun. Lencois (?), Nova America, 17 May 1975 (st), Costa & Barroso s.n. (ALCB); Mun. Malhada de Pedras, km 11 on rd. from Brumado to Caetite, 480 m, 14 Apr 1983 (fl, fr), Leuenberger et al. 3063 (B, CEPEC, W); Mun. Maracas, 26 Jan 1980 (fl), Nunes 207 (RB); Mun. Marconilio Sousa (?), vie. Machado Portella, 19-23 Jun 1915 (fr), Rose & Russell 19901 (NY, US); Mun. Marconilio Sousa (?), Machado Portella, 18 Dec 1912 (fl), Zehntner 677 (RB, US), Inspectoria de Obras contra as Seccas 5049 (M); Mun. Palmeiras(?), km 16 from Rio Preto to Riacho de Mel, 19 Mar 1980 (fr), Brazao 165 (HRB); Mun. Planaltino, km 26 on rd. from Maracas to Tamburi, 500-550 m, 20 Apr 1983 (fr), Leuenberger et al. 3086 (B, CEPEC, K, SI); Mun. Planaltino, km 26 on rd. from Maracas to Tamburi, 24 Jan 1965 (fl), Pereira 9719 (R); Mun. Planaltino, on rd. from Maracas to Planaltino, 29 May 1979 (fr), Brazao 109 (HRB, RB); Mun. Rio de Contas, 19 Jun 1978 (st), Araujo 47 (RB); Mun. Vitoria da Conquista, 20 km E of Anage, 13 Apr 1983 (fr), Leuenberger et al. 3061 (B, CEPEC, K, W); between Vitoria da Conquista and Brumado, 31 Mar 1966 (fr), Castellanos 25946 (M). Local names and uses. Quiabento ("inhabento," "iniabento"), espinha de Santo Antonio and flor de cera. Frequently used for planting hedges, very frequent near houses and in hedges along roads between Anage, Brumado, and Caetite. Cuttings of about 1 m length are planted in a dense row or in a double row with the sticks arranged in an X perpendicular to the row, thus forming a broad and impenetrable hedge with a broad crown, which may be cut repeatedly. Pereskia bahiensis is closely related to P. grandifolia and P. stenantha. In general habit, P. bahiensis is stouter than P. grandifolia and has shorter, more fleshy leaves and often fewer flowers per inflorescence, and smaller seeds. Like P. 122 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 48. Distribution of Pereskia aureiflora, P. bahiensis, and P. stenantha.

stenantha, it is a plant of the caatinga. Even in the field I could not find any reliable vegetative characters distinguishing P. bahiensis from P. stenantha, al­ though the latter species is quite distinct when in flower, with its orange-reddish flower buds and urceolate flowers.

16. Pereskia stenantha Ritter, Kakteen in Siidamerika 1: 21, figs. 3, 4. 1979. Type. Brazil. Bahia: Caetite, Ritter 1251 (holotype, U, fide Ritter, but

rr~ 1986] PERESKIA (CACTACEAE) 12 3

not located there, probably not deposited; figure 3 of Ritter, 1979: 277, shows the type specimen and may serve as lectotype). Figs. 11F, 14A, 15Ac, 18K, 19J, K, 48, 49A-J. Shrub or small tree, 2-4(-6) m, branching at or from near the base; trunk to ca. 15 cm diam. in the few large specimens seen; main shoots erect and arching, with epitonous branching; bark gray, smooth or longitudinally fissured; distal twigs ca. 4-6 mm thick, with few to many stomata, grayish-brown after periderm formation. Areoles rounded, 2-5 mm in diam. on twigs, larger on main shoots, accrescent to 15 mm diam. on trunk, with short and dense, pale grayish-brown tomentum of uniseriate trichomes 2-3 mm long, producing spines and 1-3 brachy­ blast leaves. Leaves variable in shape and size, obovate to elliptic, often folded upward along the midrib, (5—)7—11(—15) x (2-)4-6(-9) cm; petiole 2-10 mm long, thick when fresh and with basal spur-like rim below; blade 0.6-1.0 mm thick, fleshy; base cuneate to attenuate; apex broadly acute to rounded, often recurved; midrib prominent below; lateral veins 5-7, patent, often inconspicuous in fresh leaves. Spines 0-7 on twigs, fasciculate to spreading, stiff, 1-5 cm long and to 1 mm thick, increasing in number to ca. 20 on main shoots and up to 40 on trunk, to 5 cm long and 1.5 mm thick, brown to grayish-brown, base dark red when fresh on young spines. Inflorescence terminal, densely cymose-paniculate by pro­ liferation, the receptacle of the terminal flower, 5-15 cm in diam., to ca. 15- flowered, with leafy bracts; flowers sometimes solitary, terminal or axillary. Flow­ ers perigynous to epigynous, opening very little, campanulate-urceolate, 20-35 mm long and 10-20 mm in diam., odorless; flower buds orange-red; flowers orange-red without, pink within; pedicels 1-3 mm long, hardly distinguishable from the turbinate receptacle; receptacle 7-10 mm in diam., bracteate in the upper half; bracts varying in number and size depending on the position of the flower within the inflorescence; areoles ca. 5-8, with short grayish-brown tomentum, the upper ones only visible as tufts of tomentum on both sides of the erect upper bracts but without longer hairs visible along the margin of the bracts; lower bracts 3-5(-7), usually five on the distal flowers and calyx-like, spreading, 5-20(-40) x (2-)4-10(-30) mm, thick, fleshy, green, with broad base and obtuse apex; upper bracts (l-)3-5(-6), erect, broadly triangular, 4-5 x 5-7 mm, keeled, fleshy, green to reddish; sepaloids 4-5, erect, ovate to narrowly ovate, to ca. 18 x 9 mm, with broad base, to 1.5 mm thick and fleshy, concave, apex acute or rounded with a point, red to orange-red, with a purplish-pink tinge usually at the margin only and particularly in bud, more reddish than in P. bahiensis; petaloids 6-8, erect, imbricate, spreading at the tip only, linear-lanceolate, 20-25 x 4-12 mm, the inner ones narrower than the outer; apex acute to broadly acute, blade pink to purplish-pink, base ca. 1 mm thick when fresh, pale pink; stamens ca. 140, 12- 15 mm long, erect, surrounding the slightly longer style and stigma lobes, not attaining the throat of the corolla; filaments in ca. 2 rows, white, their base slightly prominent at the level of insertion above the conspicuous 1.5-2 mm high nectarial wall; anthers ca. 1 x 0.3 mm, pale yellow; pollen mostly 12-colpate, medium- sized, tectum with fine perforations and rather small spinules; ovary half inferior to inferior, locule ca. 3 mm in diam., floor concave, roof with septal ridges, placentae parietal; style 13-20 mm long and 0.8-1 mm thick, white, base widened into the conical ovary roof of 3-5 mm diam.; stigma lobes 4-7, 4 mm long, slender, suberect to spreading. Infructescence a densely cymose-paniculate cluster of proliferating fruits, erect or arching at maturity, to 20 cm in diam., with few to 15 fruits, or fruits solitary. Fruit variable, 3-7 x 2-6 cm, pyriform or turbinate and more or less angled, green to yellowish-green when mature; pedicel to 3 mm 124 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

FIG. 49. Pereskia stenantha. A, B, flowering shoots. C, D, flower buds. C, several days before, and D, just before anthesis. E, flower, l.s. at full anthesis. F, infructescence. G, fruit, l.s. H, seeds. I, areoles from main shoot. J, habit, natural, and in fence. A from Leuenberger et al. 3076, with photo (B); B-I from Leuenberger et al. 3072 with photos (B); J from photographs by the author (left from Leuenberger 3079). 1986] PERESKIA (CACTACEAE) 12 5

long; areoles conspicuous, with dense grayish-brown tomentum, rarely spiny with 1-5 spines of 1-3 cm length (only one such specimen seen); bracts obovate, ca. 5-10 mm long or larger leaf-like, usually deciduous at maturity; umbilicus con­ stricted to ca. 5 mm diam., usually filled by the flower remnant and the fleshy upper bracts; fruit wall 5-10 mm thick, containing many mucilage cavities, ovary roof to 15 mm in diam., smooth, green, 5-10 mm thick, with mucilaginous tissue inside; locule broadly obovate, to 25 mm. Seeds ca. 30(-50), obovate, 4.5-5.5 mm long, 3.2-3.6 mm broad and 2.0-2.5 mm thick, smooth, black, glossy; hilum transversely lunate to semicircular, ca. 1 mm in diam., whitish; micropylar nose mostly conspicuous. Seedling (three weeks old) with hypocotyl ca. 20-30 mm long and 2-2.5 mm thick, reddish; cotyledons ovate-elliptic, ca. 30-40 x 15-20 mm, midrib faint, lateral veins 0-2, inconspicuous. Chromosome number: 2n = 22; Leuenberger et al. 3072, cult. hort. Berol. 166-72-83-10 (B). Distribution (Fig. 48) and phenology. Southern central Bahia, Brazil, in the caatinga, also planted in hedges, ca. 400-600 m, flowering from November to April, mature fruits collected in April, not frequent; distribution insufficiently known.

Specimens examined. BRAZIL. BAHIA: Bom Jesus da Lapa, on rd. to Riacho de Santana, 28 Oct 1965 (fl), Duarte 9531 (RB); Bom Jesus da Lapa, ca. 8 km down rd. to Calderao, Fazenda Imbuzeiro da Onca, ca. 500 m, 19 Apr 1980 (st), Harley 21543 (B, K, CEPEC); Bom Jesus da Lapa, 2 km E of town, 450 m, 16 Apr 1983 (fl, fr), Leuenberger et al. 3076 (B, CEPEC, K); Bom Jesus da Lapa, 8 km on rd. to Ibotirama, 3 km E of rd., 450 m, 17 Apr 1983 (fl, fr), Leuenberger et al. 3079 (B, CEPEC, K, SI); Bom Jesus da Lapa, nr. S. Antonio, Nov 1912 (fl, fr), Zehntner 631 (US), Inspectoria de Obras contra as Seccas 5047 (M); Mun. de Igapora, 6 km W of Igapora, 16 Apr 1983 (fl, fr), Leuenberger et al. 3072 (B, CEPEC, K, SI, W); Porto Novo do Rio Corrento, 12 Nov 1912 (fl), Zehntner 563 (US), Inspectoria de Obras contra as Seccas 5048 (M); Mun. Riacho de Santana, 10 km W of Igapora, 18 Apr 1983 (fl, fr), Leuenberger et al. 3081 (B, CEPEC, K). Local names and uses. Quiabento, espinha de Santo Antonio, flor de cera. Planted in hedges near Bom Jesus da Lapa, Igapora and Riacho de Santana but according to local people increasingly replaced by barbed wire because the spines are feared. Specimens of this species were collected in Bom Jesus da Lapa by Zehntner in 1912, and a specimen of P. bahiensis, collected in the same year {Zehntner 639), is annotated with the remark that the plant from Lapa (Zehntner 631) differs in its longer and red-orange flower buds and the flowers which open only a little. However, the specimens and the notes were not sufficient to make the diagnostic characters evident to later workers. They are most striking in fresh flowers, but in specimens dried without special care the flowers of P. stenantha look much like withered flowers or flower buds of P. bahiensis to the observer who has not seen both as living specimens. Ritter recognized the species as new during field observations, probably in 1964, when he discovered several new Cactaceae in southern Bahia (Ritter, 1979, pp. 66-69). He cites Caetite as the type locality, but during a visit to that area in 1983 I found only P. bahiensis in flower and fruit. Local people at Brejinho das Ametistas near Caetite say that two different plants occur in the area. Pereskia stenantha is a remarkable species because it does not seem to differ from P. bahiensis in vegetative characters. Those given by Ritter (1979) (time of spine and leaf formation, color of leaf margin), do not prove reliable from the material I have seen. All the more surprising are the flowers, which are unique in the genus with their urceolate corolla. The flowers are not structurally different, but differ greatly from P. bahiensis in orientation, shape, and color of the sepaloids and petaloids and the larger nectary, all of which suggests a hummingbird pollination syndrome. Ritter (1979) inferred that this is 126 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41 a case of rapid evolution. Unfortunately no observations on pollinators are avail­ able yet. Near Bom Jesus da Lapa a shrub of similar habit and shape and color of inflorescence, Jatropha mutabilis (Pohl) Baillon, is easily confused with Pereskia stenantha at a distance.

INSUFFICIENTLY KNOWN TAXA Pereskia sp. A. Fig 14C. Small tree, resembling P. zinniiflora, but leaves larger, tomentum of areoles not rusty brown; only pistillate flowers known (a single flower from a cultivated specimen). Twigs 2-4(-5) mm thick, grayish-brown after periderm formation. Leaves lanceolate to oblanceolate (2-)3-6(-12) x 0.8-3 cm, petiole ca. 1 mm long, leaf base cuneate to attenuate, blade fleshy, green, rather thin; apex acu­ minate; midrib prominent below, slightly impressed above when fresh; lateral veins 0-3(-4), often inconspicuous, the longer ones sometimes forked. Spines 0- 3, often two on areoles of young twigs, 5-15 on areoles of older branches, 10-20 mm long and 0.2-0.4 mm thick, acicular, grayish or reddish-brown to nearly black, conically thickened at the base. Flowers (only one pistillate flower seen) terminal on leafy twigs, ca. 3 cm in diam., bright purplish-pink with green recep­ tacle and bracts; receptacle spheroidal to depressed spheroidal, 16 mm in diam., smooth to slightly grooved when fresh, upper rim constricted to ca. 7 mm diam.; receptacular areoles inconspicuous; receptacular bracts ca. four, foliaceous, lan­ ceolate, 20-35 x 6-12 mm, apex acuminate; receptacular rim naked, without areoles and upper bracts; sepaloids 3-5, appressed, 7-11 x 5-10 mm, oblong to obovate with broad base and obtuse apex, greenish to pink, fleshy at base; sepaloids ca. nine, obovate, ca. 15 x 12 mm, thin, purplish-pink, apex obtuse; stamens dwarfed, 1 mm long, sterile, inserted on the short-cylindric receptacular rim which forms an open nectarial pit delimited by the flat ovary roof at the bottom. Ovary inferior; locule large, with nearly flat bottom and ovary roof, ca. 10 mm in diam., placentae parietal, forming a narrow and long U below the septal ridges of the ovary roof; style abruptly rising from the flat ovary roof, 3 mm long and 1.5 mm thick when fresh; stigma lobes eight, spreading, 4-5 mm long; ovules 0.9 mm long, numerous. Fruit not seen, but according to Padre Julio Cicero resembling that of Hura crepitans in shape (depressed globose), but smaller, bracteate, green, containing ca. 150 seeds. Distribution (Fig. 36) and phenology. Haiti and Dominican Republic. From sea level to ca. 400 m; range and ecology insufficiently known. Specimens examined. Without locality: Cult. hort. reg. Paris, Herb. A. L. Jussieu 13546A (P, US). HAITI. Massif de la Selle, Morne des Commissaires, Anses-a-Pitre, limestone cliffs nr. La. Hatte, 21 Aug 1926 (st), Ekman H6709 (G, K, NY, S, US). DOMINICAN REPUBLIC. LA ESTRELLETA: Banica, Cerro San Francisco, 400 m, 18 Sep 1977, Marcano & Cicero s.n., cult, in coll. Cicero, Nov. 1982 (st), Czerwenka s.n. (B); cult., 25 Mar 1984 (fl), hort. Berol. 259-02-82-30 (B). During the preparation of this monograph the first three of the above cited specimens posed a puzzling identification problem, all being sterile and not exactly matching any known species. During a brief visit to the Dominican Republic in March 1983, I saw the specimen from Banica in Padre Julio Cicero's garden at San Cristobal. When I had seen Pereskia portulacifolia and P. quisqueyana in their natural habitat, it became fairly clear that the plant would eventually prove to be a new taxon, intermediate between P. quisqueyana and P. zinniiflora. Un­ fortunately, it was not possible to study the plant in its natural habitat, and no 1986] PERESKIA (CACTACEAE) 12 7 flowers or fruit could be obtained. Only in March 1984, a small cutting cultivated at Berlin produced a flower, strikingly different from all seen before by the sub­ globular receptacle, large spreading stigma lobes, and dwarfed sterile stamens. It seemed doubtful that this was a normal feature of the plant. However, it was found that all collections of P. portulacifolia and P. quisqueyana have equally imperfect but staminate flowers. The same was observed in a cultivated flowering specimen of P. zinniiflora at Berlin in fall, 1984, and only in 1985 did I see material of both pistillate and staminate flowers of P. portulacifolia (q.v.). The shape of the receptacle of the pistillate flower of P. sp. A is not different from that of P. portulacifolia and P. zinniiflora. In P. quisqueyana it is still unknown. A larger number of complete specimens is needed. Vegetative characters like leaf shape and spine color, first thought to be distinctive, prove to be rather variable in cultivation and need further study as well. It is noteworthy that the first collection cited above, a specimen with leaves identical to sp. A in the Jussieu Herbarium (18th or early 19th century), labeled as "Pereskia spinosissima arborescens fructu fici formi, in h. reg." is annotated with "sp. nov." in different handwriting, hence considered as something new long ago. The specimen from the southern coast of Haiti collected by Ekman was annotated by Werdermann as "Pereskia aff. cubensis," thus also recognized as possibly distinctive. Without sufficient evidence to include it in P. quisqueyana or to treat it as separate, I prefer to leave it among the insufficiently known taxa.

DOUBTFUL NAMES AND EXCLUDED TAXA Cactus longispinosus (Haworth) Link, Enum. pi. hort. reg. Berol. 2: 25. 1822. Orthographic error based on Pereskia longispina Haworth (q.v.). Cactus neglectus Dehnhardt, Rivista Napol. 1(3): 166. 1839. Type. Not cited. Described from a plant cultivated at Naples as "Cactus pseudo-tuna." Erroneously referred to "Pereskiae sp." by Index Kewensis, vol. 1. The diagnosis "C. articulato-prolifer, articulis latis ovatis turgidis ..." def­ initely excludes it from this genus. It can probably be identified with some Opuntia species described earlier. Pereskia ajfinis Meinshausen, Wochenschr. Gartnerei Pflanzenk. 2: 118. 1859. Type. Not cited. Described from a cultivated plant similar to Opuntia subulata (Miihlenpfordt) Engelmann; to be excluded from Pereskia and doubtful under Opuntia. Pereskia crassicaulis Zuccarini ex Pfeiffer, Enum. diagn. Cact. 176. 1837. Nom. nud., listed as synonym under P. spathulata (q.v.). Pereskia glomerata Pfeiffer, Enum. diagn. Cact. 179. 1837. Type. Not cited. De­ scribed from a plant observed by Meyen (1834: 452) in the Andes of Tacna, Peru. Excluded from the genus and tentatively mentioned in the synonymy of Opuntia atacamensis Philippi by Britton and Rose (1919: 94). Pereskia grandispina Hort. ex Forbes, Hort. Tour Germ. 159 (cited by Vaupel 1925: 17 as "Journ. Hort. Tour Germ. 159. 1837"). Doubtful. Pub­ lication not seen. Pereskia higuerana Cardenas, Cactus 19(80-81): 18. 1964. Type. Bolivia. Valle- grande: Near Higuera, 1600 m, Apr 1954, Candia s.n. in Herb. Car- denasianum no. 6123 (holotype not located, isotype indicated for US, but not located). From the description and the poor illustration this 128 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

taxon is a Pereskiopsis rather than a Pereskia. A plant cultivated at the Reservegarten Schonbrunn, Vienna, and which, according to E. Zecher, was received as Pereskia higuerana from R. Vasquez, Cochabamba, and presumably originated from Cardenas, is apparently the same as the plant widespread in amateur collections as stock and is probably correctly identified as Pereskiopsis spathulata (Otto) Britton & Rose, a plant of Mexican origin. As long as the original material is not available for study and the origin of the specimen is not confirmed, I prefer to leave this name in the doubtful category, to be excluded from Pereskia. Pereskia lanceolata Hort. Berol. ex Salm-Dyck, Cact. hort. Dyck. 252. 1850. Described as "hardly different from P. spathulata . . . ." A doubtful name probably belonging to Pereskiopsis spathulata (Otto) Britton & Rose. Pereskia longispina Haworth, Syn. pi. Succ. 178. 1812. Type. "cult, ante 1808, communicavit Dom. Gul. Anderson," apparently not preserved. The description does not allow identification of this plant, and although it has been considered as a synonym of P. aculeata by previous authors, it seems best placed in the doubtful category. Pereskia pflanzii Vaupel, Z. Sukkulentenk. 1: 56. 1923. Type. Bolivia. Tarija: In the region of Santa Isabel, 50 km downriver from Villa Montes, Pflanz 6 (B, destroyed) = Quiabentia pflanzii (Vaupel) Backeberg & F. Knuth. Pereskia philippii F. A. C. Weber in Bois, Diet. hort. 939. 1891. Type not cited. =Maihuenia philippii (F. A. C. Weber) K. Schumann. Pereskia plantaginea Hort. Gotting. ex Pfeiffer, Enum. diagn. Cact. 179. 1837. Nom. nud. Doubtful. Pereskia poeppigii (Pfeiffer) Salm-Dyck, Cact. hort. Dyck. ed. 2: 76, 252. 1850 [basionym: Opuntia poeppigii Otto ex Pfeiffer, Enum. diagn. Cact. 174. 1837] = (Pfeiffer) K. Schumann. Pereskia rotundifolia De Candolle, Mem. Mus. Hist. Nat. 17: 77. Type. Plate of "Cactus frut escens" in Mocino, Fl. Mex. ined., no. 287 (cf. McVaugh, 1980a: 126), preserved under no. 265 at the Hunt Institute for Botanical Documentation. No specimen located. = Pereskiopsis rotundifolia (De Candolle) Britton & Rose. Pereskia scandens (Britton & Rose) Standley, Publ. Field Mus. Nat. Hist. Chicago, Bot. Ser. 3: 369. 1930. =Pereskiopsis scandens Britton & Rose. Pereskia spathulata hort. Berol. ex Pfeiffer, Enum. diagn. Cact. 176. 1837. = Pereskia spathulata Link et Otto ex Pfeiffer, Beschr. Syn. Cact. 201. 1837. Described from a living plant at the Berlin Botanical Garden. =Pereskiopsis spathulata (Pfeiffer) Britton & Rose. Pereskia subulata Miihlenpfordt, Allg. Gartenzeitung 13: 347. 1845. Type. Not cited. Described from a living plant. = Opuntia subulata (Miihlen­ pfordt) Engelmann. Pereskia verticillata Vaupel, Z. Sukkulentenk. 1: 55. 1923. Type. Bolivia. Tarija: Laguna Santa Isabel, 50 km downriver from Villa Montes, Pflanz 5 (holotype, B, destroyed). = Quiabentia verticillata (Vaupel) Backeberg & F. Knuth. Pereskia zehntneri Britton & Rose, Cactaceae 1: 14. 1919. Type. Brazil. Bahia: Bom Jesus da Lapa, 16 Nov 1912, Zehntner 630 (US). = Quiabentia zehntneri (Britton & Rose) Britton & Rose. Rhodocactus higueranus (Cardenas) Backeberg, Kakteenlex. 396. 1966. See under Pereskia higuerana Cardenas. 1986] PERESKIA (CACTACEAE) 129

Acknowledgments The following institutions, to which I am most grateful, have kindly provided facilities during and after field work: Instituto de Biologia and Jardin Botanico, Universidad Nacional Autonoma de Mexico, Mexico, Jardin Botanico Rafael M. Moscoso, Santo Domingo, Dominican Republic, and Centro de Pesquisas do Cacau, Itabuna, Bahia, Brazil. My special thanks for loan of material, in many cases for prolonged periods, for help in locating specimens, or for advice during my visits to the herbaria are due to the curators of the following herbaria: A, ALCB, BAB, BAH, BM, BOLV, BR, C, CEPEC, COL, CR, CTES, CUZ, E, F, Fl, FR, FTG, G, GH, GOET, H, HBG, HEID, HNT, HRB, JBSD, K, L, LD, LE, LIL, LP, M, MA, MEXU, MICH, MO, NY, OXF, P, POM, R, RB, RSA, S, SGO, SI, U, UB, UC, UEC, UPR, UPS, US, VEN, VT, W, Z, ZSS. I am most grateful to Christof Schiers, Gustavo Martinelli, Andre M. de Carvalho, Luiz A. Mattos Silva, Jiirgen Czerwenka, and Eberhard Bolay, who have helped me in field work, to Monika Schroder for the chromosome counts and photographic processing, to Christine Hillman-Huber for the ink drawings, to H. Loose, H. Ehrke, and M. Swiatkowiak for taking care of the living collection, to Susanne Schwerdtfeger for preparation of garden herbarium specimens, to Brigitte Schilling for handling the loans, to Sylvia Schwitalla for the SEM-micrographs of the gy­ noecium, and to the many persons who made other useful contributions, namely K. Ahlgrimm, T. Antonio, Julio Cicero, P. Fantz, L. D. Gomez, M. Hakki, L. Horst, D. R. Hunt, R. Kiesling, M. Kimnach, A. Krapovickas, A. Liogier, A. Lourteig, E. Medina, M. Muhoz Schick, W. Rauh, Th. Raus, H. Sanchez-Mejo- rada, M. Sousa Sanchez, L. Scheinvar, J. Solomon, J. D. Supthut, N. P. Taylor, and T. Zanoni.

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. 1982. Initiation of stamens, carpels, and receptacle in the Cactaceae. Amer. J. Bot. 69: 369- 379. Rowley, G. 1980. Pollination syndromes and cactus taxonomy. Cact. Succ. J. Gr. Brit. 42: 95-98. Riimpler, Th. 1886. Carl Friedrich Foerster's Handbuch der Cacteenkunde. Leipzig. Rutishauser, R. 1981. Basale Blattauswuechse bei Centrospermen. Pages 21-27 in W. Van Cotthem (ed.), Morphologie-Anatomie und Systematik der Pflanzen, 5. Symposium, Gent 1979. Waege- man, Ninove (Belgium). Salisbury, R. A. 1796. Prodromus stirpium in horto ad Chapel Allerton vigentium. Londini. Salm-Dyck, J. 1850. Cacteae in Horto Dyckensi cultae anno 1849. Henry & Cohen, Bonnae. Schumann, K. 1890. Cactaceae. In F. von Martius (ed.), Flora brasiliensis 4(2): 185-334. . 1898. Gesamtbeschreibung der Kakteen. Neumann, Neudamm. . 1899. Die Verbreitung der Cactaceae im Verhaltniss zu ihrer systematischen Gliederung. Konigl. Akad. Wiss., Berlin. 1903. Gesamtbeschreibung der Kakteen. 2., um die Nachtrage von 1898 bis 1902 vermehrte Auflage. Neumann, Neudamm. Sloane, H. 1696. Catalogus plantarum, quae in insula Jamaica sponte proveniunt vel vulgo coluntur. Londini. Spencer, J. L. 1955. A cytological study of the Cactaceae of Puerto Rico. Bot. Gaz. 117: 33-37. Stafleu, F. A. & R. S. Cowan. 1983. Taxonomic literature, ed. 2, vol. 4. Regnum Veg. 110. Utrecht. Standley, P. C. 1924. Trees and shrubs of Mexico. Part 4. Contr. U.S. Natl. Herb. 23: 849-1312. & L. O. Williams. 1962. Flora of Guatemala. Part VII, no. 2. Fieldiana Bot. 24: 187-281. Takhtajan, A. 1959. Die Evolution der Angiospermen. G. Fischer, Jena. Tiagi, Y. D. 1955. Studies in floral morphology. II. Vascular anatomy of the flower of certain species of the Cactaceae. J. Indian Bot. Soc. 34: 408-428. Tsukada, M. 1964. Pollen morphology and identification. II. Cactaceae. Pollen et Spores 6: 45-84. Uil, Ir. G. 1978. Rhodocactus grandifolius (Haw.) Knuth een interessante cactus. Succulenta 57: 258-265. Urban, I. 1902-1903. Symbolae antillanae seu fundamenta florae Indiae Occidentalis. Vol. 3. Lipsiae. Vaupel, F. 1925. Die Kakteen. Monographic der Cactaceae. Vol. 1. Berlin. Wagenaar Hummelinck, P. 1938. Notes on the Cactaceae of Curacao, Aruba, Bonaire and North Venezuela. Recueil Trav. Bot. Neerl. 35: 29-54. Weber, F. 1898a. Les Pereskia et Opuntia pereskioides du Mexique. Bull. Mus. Hist. Nat. (Paris) 4: 162-167. . 1898b. Pereskia. Pages 937-939 in D. Bois (ed.), Dictionnaire d'Horticulture. Klincksieck, Paris, 1893-1899. Weingart, W. 1904. Peireskia amapola Web. Monatsschr. Kakteenk. 14: 83-87. Werderman, E. 1931. Die von E. L. Ekman in Westindien, besonders auf Cuba and Hispaniola gesammelten Cactaceae. Repert. Spec. Nov. Regni Veg. 29: 222-243. . 1937. Beitrage zur Nomenklatur. Kakteenk. & Kakteenfr. 1937: 17-24. Wetterwald, X. 1889. Blatt- und Sprossbildung bei Euphorbien und Cacteen. Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 53: 377-440. Wijnands, D. O. 1983. The botany of the Commelins. Balkema, Rotterdam. Zuccarini, J. G. 1837. Plantarum no varum vel minus cognitarum, quae in horto botanico herba- rioque regio monacensi servantur, fasc. 3, Cactaceae. Abh. Math.-Phys. CI. Konigl. Bayer. Akad. Wiss. 2: 597-742. 134 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Numerical List of Taxa

1. Pereskia aculeata Miller 9. P. portulacifolia (Linnaeus) De Candolle 2. P. humboldtii Britton & Rose 10. P. quisqueyana Liogier a. var. humboldtii 11. P. bleo (Kunth) De Candolle b. var. rauhii (Backeberg) Leuenberger 12. P. nemorosa Rojas Acosta 3. P. diaz-romeroana Cardenas 13. P. sacharosa Grisebach 4. P. weberiana K. Schumann 14. P grandifolia Haworth 5. P lychnidiflora De Candolle a. var. grandifolia 6. P. aureiflora Ritter b. var. violacea Leuenberger 7. P guamacho F. A. C. Weber 15. P. bahiensis Gurke 8. P. zinniiflora De Candolle 16. P. stenantha Ritter

List of Exsiccata Ahlgrimm, K., 41, 68, 74a (13). British Guiana Forest Dept., 6975 (1). Alain, Bro., 1080 (8). Britton, N. L., 2013 (8); 19270 (1). Alfaro, A., s.n. (5). Britton, N. L., E. G. Britton & J. F. Cowell, 9702 Allen, P. H., 281 (11); 3972(1). (14a). Almeida, Edson de Faria, 28 (15). Britton, N. L., E. G. Britton & T. E. Hazen, 189 Alston, A. H. G., 5857a (7). (1). Araque J. & F. A. Barkley, 19Ar517 (13). Britton, N. L. & J. F. Cowell, 347 (14a). Araujo, Adonia P. de, 47, 103 (15). Britton, N. L., J. F. Cowell & J. A. Shafer, 13034 Araujo, D., 369, 680, 766 (1). (8). Arenas, P., 1776 (13); 1917 (12). Britton, N. L. & P. Wilson, 5513 (8). Arganaraz, J., 468 (13). Broadway, W. E., 372, 4290 (1); s.n. (14a). Aristeguieta, L., 3052 (7). Brooke, W. M. A., 5790, 5929 (13). Aristeguieta, L. et al., 6744 (7). Brooks, Th., s.n. (8). Arnoldo, Fr. M., 1652(11). Buch, W., 341 (9). Asplund, E., 14586 (14a). Buck, P., s.n. (1). Atwood & Neill, AN292 (5). Burchell,W. J.,901, 1364, 1665 (14a); 2858,4675, Badcock, 831 (13). 7347-2 (1). Bailey, L. H. & E. Z. Bailey, 304, 658 (7); 798 Burkart, A., 20135 (13). (14a); 1351, 1921 (7). Burkart, A. & N. S. Troncoso, 11424 (13). Balansa, B., 2502 (1); 2671 (12). Cabrera, A. L., 16070(13). Balbis (see Bertero). Cabrera, A. L. & Fabris, 21003 (13). Barkley, F. A., 39448 (5). Cabrera, A. L. et al., 24045, 29817, 32259 (13). Barkley, F. A. & J. Araque M., 24 (7). Calderon, S., 1164, 1757, 2117 (5). Bartlett, H. H., 17385 (9). Campos Porto, P., 1714 (1). Bautsch, P., 137(9). Cardenas, M., 4069, 4388 (3); 5005, 5006, 5007, Belem, R. P. & M. Magalhaes, 877 (1). 5008, s.n. (4). Bello, W., 472 (1); 653 (14a). Carillo, Gil, s.n. (14a). Bernardi, L., 435 (14a). Carlson, M. C, 3222 (14a). Berry, 1009 (1). Case, P. etal., 169 (5). Bertero, C. G., s.n. (7). Castellanos, A., 25946 (15); 26413 (1). Bertoni, 14 (12); 2770, 3628 (1). Castillon, L., 3073 (13). Beuther, A., 91 (11). Cavalo, G. Pedra do, 4 (1); 1011 (14a). Blanchet, M., 1654, 1746 (1). Ceballos, A. et al., Bo356 (13). Blydenstein, J., 383 (7). Chavelas, J. & C. Zamora, ES4815 (5). Boke, N. H., 25 (5). Chaves, D., 281 (5). Boldingh, I., 1858a, 4131 (1); 5442 (7); s.n. (14a). Christ, P., 10 (7). Bonpland, A., 641 (1). Cicero, Julio, s.n. (San Jose de Ocoa) (1); s.n. Box, H. E. 1539(1). (Bayahibe)(10). Brade, A., 2553 (5); 9023, 17425, 17435 (1). Clemente, Bro., 5713 (1). Braun, P., 454 (1); 476 (14b); 481 (1). Clover, E. U., 9134, 9228, 9493 (5); 11030 (9). Bravo H., H., 35, s.n., s.n. (5). Colaris, W. J. J., 1032 (1). Brazao, Jose E. M., 109, 165 (15). Conzatti, C, 3 (14a); 3667 (5); s.n. (Oaxaca) (1); Breedlove, D. E. & E. McClintock, 23488 (1). s.n. (Orizaba) (14a); s.n. (Salina Cruz and Te­ Brenes, A. M. & J. Valerio, 77, 137 (14a). huantepec) (5). Breteler, F. I., 4553 (7). Cook, O. F., 806 (5); s.n. (9). 1986] PERESKIA (CACTACEAE) 135

Cook, O. F. & Collins, 7459, 7853 (5). Gagzo, s.n. (Bahia) (1), (Vitoria) (1). Cook, O. F. & C. B. Doyle 291 (5). Gaillard, D. D., s.n. (11). Costa, A. L., s.n. (ALCB 02910) (1); s.n. (ALCB Gamier, Bro. A., 900 (5). 02911) (15). Gaudichaud, C, 911 (1); 912 (14a); 913 (1); s.n. Costa, A. L. & G. M. Barroso, s.n. (15). (Rio de Janeiro) (1, 14a). Croat, T. B., 294 (14a). Gaumer, G. F., 24374 (1). Croat, T. B. & D. M. Porter, 15519 (11). Gay, s.n. (14a). Cufodonti, G., 49 (7). Geay, 27(11). Curran, H. M., 355 (11); 357 (7); 621 (14a); s.n. Gentry, A., 951 (14a). (12). GirTord & Fonseca, G335 (14a). Curran, H. M. & M. Haman, 741, 968 (7). Gilson, H. C, 1517(13). Curso Dendrol., 300 (7). Gines, H., 3386 (7). Cutak, L., s.n. (11). Ginzberger, A., 61 (1). Czerwenka, J., 28 (9); OV744 (1). Glassman, S. F., 1713 (5). Daniel, Bro., 1181 (11). Glaziou, A., 12868 (14a); 18267 (1); 18268 (14a). Davidse, G. & A. C. Gonzalez, 13690 (1). Goes, Campos & D. Constantino, s.n. (1). Davidson, 3839 (3). Goldman, E. A., 1884, 1956 (11). Dawe, M. T., 869(11). Gongrijp, J. W., BW 5208 (7). Deam, C. C, 6246, 6269 (5). Gonjian, B., s.n. (12). Delgadillo, D., s.n. (5). Goudot, J., s.n. (St. Martha) (7); s.n. (Muzo) (11). Diaz, A., s.n. (2a). Graham, s.n. (1). Dryander, E., 1962(11). Gregory, L. E., 209 (1). Duarte, A. P., 4806 (1); 9531 (16); 10534 (6). Grisebach, A., s.n. (13). Duchassaing, s.n. (11). Grosourdy, de, 13 (7). Dugand, A., 112, 406(7). Groth, s.n. (14a). Dugand, A. & R. Jaramillo, 3453 (7). Guevara, L. Cardenas de, 1525, 1565 (7). Duque, J. M., 1854a (11). Hage, J. L., 1640(1). Dusen, P., 11282, 17842(1). Hanbury-Tracy, J., 304 (7). Duss, P., 958, 977, 3053, 3073 (1); 3895 (14a, 1 Harley,R.M., 16352(15); 19992 (6); 21543 (16). partly). Harmon, W. E., 2574 (5). Echeverria, J. A., 426 (14a). Harmon, W. E. & J. A. Fuentes, 6356 (5). Eggers, H., 275 (14a); 875 (1); 5441 (8); s.n. (1). Harris, W., 38 (14a). Ehrhardt, W., 15(1). Hassler, E., 1326(12). Eichlam, F., s.n. (5). Hatschbach, G., 7896 (14a); 19046,20973,39856 Eiten, G., L. T. Eiten & I. Mimura, 6011 (14a). (!)• Ekman, E. L„ H953 (9); 1421, 4871 (8); H6709 Haught, O., 1643 (11); 4065 (7); 4547, 4798 (11). (spec. A); 8609 (8); H9593 (9); 17801 (1); 18911 Hazlett, D., 908 (5). (8); s.n. (1). Heiner, A., 637 (1). Elias, Bro., 151 (7). Herb. A. L. de Jussieu, 13546A (sp. A); 13547 Elmore, F. H., L34(ll). (13). Esser, G., 14763 (12). Herb. Pavon, s.n. (Nueva Espaiia) (8); s.n. (N. E., Eyerdam, W. J. 8c A. A. Beetle, 23035 (13). Cactus macrochanthos) (5). Fabris, 5078 (13). Herb. Sloane, 97 f. 118 (7); VI.98.1 (7). Fabrisetal., 3100,3167(13). Herb. Vahlii, s.n. (1). Fanning, K. A. & M. B. Meagher, KF122 (11). Heringer, E. P., 652 (14a); 12921 (14). Fanshawe, D. B., 3411 (1). Heringer, E. P. & G. Eiten, 15252 (14b). Fantz, P. R., 4185(11). Herminier, L., s.n. (1). Faris, J. A., 374 (14a). Hernandez M., R., 89 (5). Fendler, A., 67 (1); 2326 (7). Herter, W. G., 82375 (12). Fernandez Casas, J. & J. Molero, 4392 (13). Herzog, T., 109 (13); 1869 (3). Ferreira, J., 2316 (1). Hicken, C. M., 317 (12). Ferreira, M. do Socorro G., 118 (15). Hioram, Bro., 1835 (8). Ferreira, V. F., 348 (14a). Hoehne, F. C, 3859, 3860, 4863 (13). Ferrero, M., s.n. (8). Holdridge, L. R., 209 (1); 845 (9). Ferreyra, R., 14200, 14204 (2a). Holton, I. F., s.n. (11). Fiebrig, K., 6485 (1). Holway, E. W. D. & M. M. Holway, 1123 (14a). Fleischmann, E., 512 (13) Horst, L. & W. Ubelmann, HU226 (14b); HU492 Forest Dept. British Guiana, 6975 (1). (14a). Forsstrom, s.n. (1) Howard, R. A., 6636 (14a); 12598 (9). Freire, C. V., 611 (1). Humbert, H., 21157(13). Freire, C. V. & J. Vidal, R91095 (14a). Humboldt, F. & A. Bonpland, 1546 (11); 3594 Fries, R. E., s.n. (13). (2a). Fuertes, M., 589, s.n. (1). Hutchison, P. C, 1442 (2b); 1452 (2a). Furtado, P. P., 160 (1). Hutchison, P. C. & J. K. Wright, 3621 (2a). 136 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Hutchison, P. C. et al., 6219 (2b). Little, E. L., 15795 (7). litis, H. H. & L. Gillespie, G28 (5). Lorentz, P. G. & G. Hieronymus, 410, 524, 562 Inspectoria de Obras contra as Seccas, 5047, 5048 (13). (16); 5049(15). Luna, A., 511 (14a). Isert, 337 (1). Luschnath, B., s.n. (Rio de Janeiro) (1); s.n. (Serra Jack, J. G., 4300 (1); 4556 (14a); 8445 (7); 8478 dos Orgaos) (1). (1); 8479 (14a). Lutz, B., 566(1); 15701 (14a). Jahn, A., 965 (7). MacDougall, T., H483, s.n. (5). Jervis, R. N., 1061, 1504, 1879, 1904, 1905 (8). Machado, O., 63, s.n. (1). Jesus, J. Almeida de, 375, 1333 (1). Maidana, J. I., 119(13). Jimenez, J., 2578, 4325 (9). Malvarez, 379(12). Johnston, J. R., 216, s.n. (7). Manara, B. J., s.n. (7). Jussieu, J. de, s.n. (13). Marcano & Cicero, s.n. (spec. A). Kanter, 111 (13). Marcano & J. Js. Jimenez, 4305 (9). Karsten, s.n. (1). Marquez, 552 (14a). Karwinski, W., s.n. (Tequisistlan) (5); s.n. (Iter Martinelli, G., 4511 (1). Mexicanum 1841/42) (8). Marunak, V., C. Quarin & A. Schinini, 575 (13). Kellermann, W. A., 4318, 5020, 5726, 7011, 7014, Matuda, E., 2296 (5); 17050 (1); 17608 (14a); s.n. (5). 38571 (5). Kennedy, H., 2868(11). McFarlin, J. B., 6344 (14a); 6346 (1). Kiesling, R., 33, 1653, s.n. (13). Mejia, M. & T. Zanoni, 6343 (1). Killip, E. P., 39578, 39580 (13). Meyer, T., 6747 (1). Killip, E. P. & A. C. Smith, 26912 (14a). Meyer, T. & Vaca, 22473 (13). Killip, E. P. & F. Tamayo, 37038 (7). Mexia, Y., 4129, 4804(1). Kimnach, M., 172(5). Miers, J., 3833 (14a). King, R. M., 1288, 1400, 1476, 1552, 2461 (5). Molina R., A., 22484 (5). Klug, G., 40 (14a). Molina R., A. 8c A. Molina, 24283 (14a). Knize, K., 934 (3). Montes, J. E., 16621 (1). Krapovickas, A. & C. L. Cristobal, 17505 (13); Montes, R. A., 355 (14a). 29165(12). Moore, H. E. & M. Cetto, 8210 (5). Krapovickas, A. & L. Mroginski, 22229 (12). Moore, S., 955 (13). Krapovickas, A. & A. Schinini, 28535 (1). Mora, L. E., 1391, 1467(7). Krapovickas, A., A. Schinini & C. Quarin, 26709 Moran, R., 7740 (5). (13). Mori, S. A., T. S. dos Santos & C. B. Thompson, Kuhlmann, J. G., 290 (1); 6188 (14a); s.n. (Foz 11211 (15). do Iguacu) (1); s.n. (Rio de Janeiro) (1). Morong, T., 188 (12). Kuntze, O., s.n. (Tunari mountains) (4); s.n. (Cu- Mosen, H., 1330 (1); 2865 (14a). yaba) (14a). Mutis, J. C, 2269 (7). Langeron, M., s.n. (12). Neto, J. V., 7377(1). Leal, C, 41 (1). Neuwied, Princ, s.n. (14a). Lent, R. W., 592, 593 (5); 842 (14a). Neves-Armond, A., 11 (1). Leon, Bro., 71, 4631 (14a); 16731 (8). Nunes, S., 207(15). Leonard, E. C, 3503, 3606, 4251, 5325 (9); 9841 Omawale & R. Persaud, 152 (11). (14a); 10122 (1). Pabst, G., 7310, HB29478 (1). Leuenberger, B., 3044 (10); 3046, 3048 (9). Pabst, G., A. Castellanos, R. Santos & L. Duarte, Leuenberger, B. & H. S. Brito, 3103 (1). 7168 (14a). Leuenberger, B., A. de Carvalho & L. A. M. Silva, Palma, M., s.n. (1). 3060 (14b); 3061 (15); 3062 (14a); 3063, 3069, Paschoal, 17 (13). 3071 (15); 3072, 3076, 3079, 3081 (16); 3083, Pastrana, 9135 (13). 3086(15). Patin, C, s.n. (11). Leuenberger, B. & G. Martinelli, 3054, 3055, 3056 Paul, Bro., 339(11). (6); 3057 (14b). Peckolt, 121 (14a); 252, s.n. (1). Leuenberger, B. 8c Ch. Schiers, 2503, 2504, 2507, Pedersen, T. M., 13372 (1). 2508, 2509, 2510, 2511, 2512, 2513, 2515a, Peirano, s.n. (13). 2521,2525,2528,2531, 2533,2533a, 2718 (5). Pennell, F. W., 4579 (11); 4765 (7). Leuenberger, B. & C. Todzia, 2656 (14a). Pennington, T. D. & J. Sarukhan, 9213 (5). Liebmann, F. M., 11157 (5). Pereira, E., 9719(15). Lillo, M., 2106(13). Perez Arbelaez, E., A-1030 (11). Lindman, C. A. M., s.n. (Sto. Angelo) (1); s.n. Perez Arbelaez, E. & J. Cuatrecasas, 8405 (11). (Asuncion) (12). Perrin, M., 1 (7). Liogier, A. & P. Liogier, 27082 (9). Petersen & Hjerting, 529, s.n. (13). Liogier, A., P. Liogier & J. J. Jimenez, 25424 (10). Pinheiro, R. S., 2172(1). Liogier, A. H., 18612 (14a); 27032 (10). Pinto, G. C. P., 83 (1); 28/81 (15); 29/81 (14a); 1986] PERESKIA (CACTACEAE) 137

398/81 (6); s.n., ALCB2907 (1); s.n., ALCB 2908 Rusby, H. H. & White, 428 (4). (15); s.n., ALCB 2909 (14a). Rutten-Pekelharing, C, 147 (14a). Pittier, H., 1, 1884, 3507, 4704, 4821 (11); 6129 Sagra, Ramon de la, 6, s.n. (8). (14a); 6179, 6453 (7); 6480 (14a); 6510, 6545 Salzman, M., s.n. (1). (11); 7151, 7433 (14a); 8633 (1); 12157, 12365, Sampaio, A., 985 (14a); 7068 (14b); 7069, 8065 13967, 14967 (7); s.n. (photo) (11); s.n. (Puerto (1); 8215 (14a); s.n. (1). Colombia) (7); s.n. (Caracas) (7). Santoro, J., 7675 (1). Pittier, H., 10276 see Williams, L., 10276. Santos, E. B. dos, 10(1). Plee, 1, s.n. (7). Santos, G., s.n. (R 91091 & 91094) (14b). Plowman, T., 3518 (7); 3552 (11). Santos, T. S. dos, 1641 (14a). Plowman, T. & J. G. Saldarriaga, 3177 (11). Santos, T. S. et al., 3827 (14a). Poveda, L. J., 792 (14a). Saravia T., C, 1324, 2324, 2868 (7), s.n. (13). Purpus, C. A., s.n. (Mirador) (14a); s.n. (San Ge- Saravia T., C. & M. E. de Saravia, 3684 (7). ronimo) (5) Scheinvar, L., 112 (1). Quentin, R. P., 793 (14a). Schinini, A., 4915,6165,19896,21055(1); 23219 Questel, A., 1772 (1); 2214 (14a). (13). Rambo, B., 1302, 1522, 11159, 41779, 42387, Schinini, A. & Arbo, 24308 (1). 53529(1). Schinini, A. & E. Bordas, 14964, 17813, 17817, Ramia, M., 1929 (7). 18067 (12); 21326 (12). Rauh, W., 24505, 24601 (2a); 38889 (5); 40184 Schinini, A. & C. L. Cristobal, 9899, 10860 (12). (2b); 40627 (3); 53389a (14a); 53666 (2b); Schnetter, M. L., 4 (7). 58595a (9). Schott, A., 16(7); 5810(1). Rauh, W. & G. Hirsch, P2162 (2b). Schreiter, 3477(13). Raunkiaer, C, s.n. (1). Schuel, 163(13). Read, R. W., 1471 (11). Schulz, 18759(1). Reasoner, Bros., s.n. (1). Schwarz, G. J., 7670, 10463 (1). Record, S. J. & Don Jaca, 76 (7). Schwindt, E., 313, 3319, 4092 (1). Record, S. J. & H. Kuylen, G122 (5). Scolnik, R. & R. Luti, 667 (13); 778 (3). Regnell, A. F„ III, 1776(1). Scott, H. & M. Biraben, 1457 (13). Reiche, K., 189(1). Servico Florestal do Brasil, 209 (1). Reitz, P. R., 4591,6873(1). Sesse, Mocino, Castillo & Maldonaldo, 2137 (8). Reitz, P. R. & R. M. Klein, 3 (14a); 310, 9564 Shafer, J. A., 59(13); 117, 127(12); 129(1); 137 (1). (12); 2563 (1); 10352, 13861 (14a). Reviralta, M. del Carmen, 1 (14a). Shamel, Popenoe & Dorsett, 138b (14a). Ricksecker, A., 385 (14a). Shepherd, G. J. et al., 4422 (6). Ricksecker, J. J., 294 (14a). Silva, F. C. F. da, 89 (14b). Riedel, L., 89, s.n. (Rio de Janeiro 1832) (14a); Silva, L. A. Mattos et al., 1592 (14a). s.n. (Rio de Janeiro) (1). Silveira, A., 943 (6). Ritter, F., 283 (2b); 284 p.p. (Maranon) (2a); 284 Sintenis, P., 4896, s.n. (14a). p.p. (Hda. Valor) (2b); 284 p.p. (E. Balsas) (2b); Small, J. K., 8458, 8751, s.n. (1). Ritter 328 (4); 640 (13); 640cf. (12); 904 (3); Small, J. K., N. L. Britton & J. B. De Winkeler, 1079 (2b); 1250(15). 9220 (1). Robert, A., 714(13). Smith, F. D., 17, 122(7). Robinson, W. & M. W. Lynn, s.n. (7). Smith, H., 1886(7). Rodrigo, A. P., 802(12). Smith, H. H. & G. W. Smith, s.n. (14a). Rodrigues, I. A., 75 (1). Smith, L. B. & R. Klein, 11720 (1). Rodriguez, J. V., 235, 1152 (5); 2817 (14a). Solomon, J. et al., 6918 (12). Rojas, C. E. B. de, 258 (14a). Souza, H. M. de, s.n. (1). Rojas, T., s.n. (12). Standley, P. C, 156 (5); 30343, 30904 (11); 74123, Romanczuk, C. et al., 648 (1). 75634 (5); 78054 (1). Romero-Castaneda, R., 1122 (7); 6269, 9794 (11). Standley, P. C. & J. Chacon, 5451 (5); 6538, 6661 Rose, J. N., 21704, 21834, 21837 (7); 21850 (14a); (14a). 21868(1); s.n. (8). Stehle, M. & H. Stehle, 2542, 3542 (1). Rose, J. N., W. R. Fitch & P. G. Russell, 3434 Steinbach, I., 3954 (3). (1); 3611 (14a); 19786(1). Stern, W. L. et al. 128, 732 (11). Rose, J. N. & Rose, 22030 (7). Stevenson, J. A., 3026 (14a). Rose, J. N. & P. G. Russell, 19656 (14a); 19786 Steyermark, J. A., 29131,46550 (5); 54932, 55651, (15); 19639 (1); 19901 (15); 20069 (1); 20102 56225 (7); 88832 (1). (15). Steyermark, J. A. & J. E. Fernandez, 99539 (7). Rosengurtt, B-3675 (12). Steyermark, J. A. & B. J. Manara, 107935 (7). Rowell, Ch. M., 3140A(5). Steyermark, J. A. et al., 108023, 108178, 108359 Runyon, R., s.n. (1). (7). Rusby, H. H., 116(4); 1557(13). Stienstra, B. de, s.n. (12). 138 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

Stoffers, A. L., 3894 (14a). Weir, J., s.n. (1). Stuckert, T., 2456 (1); 6068 (13); 6754 (1). Werckle, C, s.n. (5). Sucre, D., 2623, 3611, 4940 (1). West, J., 3549 (5); 8359 (13). Swartz, O., s.n. (1). Widgren, J. F., 995 (1); 1010 (14a); s.n. (1). Tejera, E., 184(1). Williams, L., 8131 (14a); 9761 (5); 10276 (7). Tillet, S. S., 6711-554(7). Williams, L. & A. H. G. Alston, 72 (14a). Tonduz, A., 14001 (5). Williams, L. O. & A. Molina R., 12671 (5). Trelease, W., 2/26/02, 72/02/11 (5). Williams, R. S., 661 (11). Trinta, Z. A., 535 (1); 1051 (14a). Wood, C. E. & E. Atchison, 7404A (8). Tiirpe, Legname & Romero, 4899L (13). Woolston, A. L., 910 (12); 1002 (1). Tyson, E. L., 5581 (11). Wright, C, 205 (8); 2626 (1). Ule, E., 3816, 4690 (1); 7050 (15). Wullschlaegel, s.n. (1). Ulibarri, E. A., 496(13). Yautone, L., 655 (1). Velloso, H., 375 (14a). Zabala, S., 137(13). Venturi, S., 3818, 5619(13). Zanoni, Th. & M. Mejia, 7152 (9). Vidal, J., s.n. (14a). Zanoni, Th. et al., 12010 (1); 35204, 35205 (9). Vogl, C, 76 (7); A655, s.n. (14a). Zehntner, L„ 563, 631 (16); 639, 677, 743, s.n. Wagenaar Hummelinck, P., 101 (7). (Juazeiro), s.n. (Maracas) (15); s.n. (Ilheus) (1). Warming, E., s.n. (1). Zuloaga, F. O. & N. B. Deginani, 406 (13). Webster, G. L. et al., 12778 (5). Zuloaga, F. O. et al., 1366 (13); 2036 (1). Weddel, 350 (14a).

Index of Local Names abrojo (8), 92 guitache (5), 81 amapola (12), 107 guititache (5), 81 amatilla (5), 81 guyapa (13), 111 azedinha (1), 64 inhabento (15), 121 barbados gooseberry (1), 64 iniabanto (15), 121 bleo (11), 103 jasmin de uvas (1), 64 bleo de chupa (11), 103 lobolobo (1), 64 camelia blanca (1), 64 manzanote (5), 81 camelia roja (9), 97 mateado (5), 81 cervetano (4), 76 mateare (5), 81 chupa (11), 103 mateares (5), 81 chupa melon (11), 103 matial (5), 81 cipo estrela (1), 64 matiari (5), 81 cruz del matrimonio (5), 81 mori(12), 107 cuguchi (13), 111 najii de espinas (11), 103 cuncu (5), 81 naju de culebra (11), 103 cuncu marin (5), 81 ora pro nobis (1, 14a, 14b), 64, 116, 118 curuzu ipochi (1), 64 ora pro nobis de mata (6), 85 erizo (8), 92 Padre nostro (1), 64 Espinha de San Antonio (16), 125 patilon (5), 81 Espinha de Santo Antonio (15), 121 quiabento (14a, 15, 16), 116, 121, 125 facho (6), 85 quisca del bosque (12), 107 flordecera(15, 16), 121, 125 ramo de novia (1), 64 groseille-barbade (1), 64 sabonete (14a), 116 groseille-pays (1), 64 sacharosa (13), 111 groseiller-pays (1), 64 siichi (7), 89 groseiller criole (1), 64 supi (7), 89 Groseiller de Barbades (1), 64 Surinam gooseberry (1), 64 Groseiller de la Barbade (1), 64 suruby-i (12), 107 grosellero (1), 64 suspire (7), 89 guamacho (7), 89 suspiro (7), 89 guamacho morado (14a), 116 tuna quisca (12), 107 guichitache (5), 81 uturunku (3, 4), 73, 76 1986] PERESKIA (CACTACEAE) 139

Index of Scientific Names New names and combinations are in bold face and synonyms are in italics. Page numbers in bold face indicate primary page references. Page numbers with an asterisk (*) indicate pages with illustrations or maps.

Acacia 95 Pereskia 3, 4, 5, 11, 13, 15, 16, 19, 20, 21, 23, Agave 76 24, 25, 26, 27, 29, 34, 35, 38, 39, 40, 41, 42, Aizoaceae 43 43, 44*, 45, 46, 47, 48, 50, 51, 52*, 53, 97 Browningia 11 subgen. Ahoplocarpus 51 Cactaceae 3, 15, 16, 23, 24, 25, 29, 35, 39, 42, subgen. Eupeireskia 51 44, 64, 76 subgen. Neopeireskia 5, 51 Cacteae 53 subgen. Rhodocactus 5, 51 Cactoideae 23, 25, 26, 35, 42, 43, 53 ser. Grandifoliae 51 Cactus ser. Typicae 51 bleo 5, 99 acardia 66 fimbriatus 76 aculeata 4, 5, 8, 10*, 13, 14*, 15, 16, 17*, 18*, frutescens 128 21, 22*, 23, 24, 26*, 29, 30*, 31, 33*, 35, grandifolius 113 36, 38, 42, 43, 45, 46, 47, 48, 49, 51, 55, horridus 5, 67, 69 57, 58, 60*, 61, 62*, 64, 65, 89, 128 longispinosus 127 var. godseffiana 59 lucidus 59 var. lanceolata 59, 66 macrochanthos 76, 82 var. rotundifolia 59, 66 neglectus 127 var. rubescens 59 opuntiaeflorus 76 fa. rubescens 59 pereskia 4, 5, 59, 64, 65, 89, 111 cv. Godseffiana 66 portulaccaefolius 97 affinis 127 portulacifolius 5, 47, 93 amapola 103, 107 pseudo-tuna 127 var. argentina 25, 40, 103, 107 rosa 113 antoniana 73 tuna 69 argentina 103 zinniaeflorus 93 aureiflora 14*, 29, 30*, 31, 35, 36, 37*, 39, 40, Caryophyllales 26, 42 41*, 46, 56, 57, 58, 83, 84*, 85, 122* Centrospermae 26, 42 autumnalis 34, 48, 77, 82 Consolea 11 bahiensis 15, 22*, 30*, 31, 35, 36, 38, 43, 45, Dahlia 9 46, 47, 48, 49*, 50, 56, 57, 58, 118, 119, Didiereaceae 40 120*, 121, 122*, 125 Euphorbia 40 bleo 15, 16, 17*, 19, 20*, 21, 22*, 23, 24, 26*, Hura crepitans 126 29, 30*, 31, 34, 35, 36, 40, 42, 43, 44, 46, Hymenoptera 47, 92 48, 50, 56, 57, 58,99, 100*, 102*, 103, 116 Jatropha mutabilis 126 brasiliensis 66 Lantana camara 47 calandriniaefolia 11 Lepidoptera 47, 92 colombiana 34, 85, 89 Maihuenia 35, 43, 51, 53 conzattii 38, 39, 51, 77 philippii 128 corrugata 42, 99, 103 poeppigii 128 crassicaulis 127 Mammillaria dioica 47 cruenta 103 Myrtillocactus 16 cubensis 14, 89, 93, 127 Neoabbottia 11, 95 diaz-romeroana 8, 17, 23, 24, 27, 30*, 32, 34, Neoraimondia 16 36, 42, 43, 45, 47, 50, 56, 57, 58, 71, 72*, Opuntia 5, 44, 51, 53, 95 73, 76, 111 atacamensis 127 discolor 66 golziana 76 foetens 66 pititache 11 fragrans 66 poeppigii 128 gigantea 116 subulata 127, 128 glomerata 127 Opuntioideae 23, 38, 39, 42, 43, 51, 53 godseffiana 59 Pachycereus 11 grandiflora 116 Pareskia nemerosa 103 grandifolia 5, 6*, 8, 11, 16, 17*, 19, 21, 23, 24, Peirescia 4 29, 31, 33*, 34, 35, 36, 38, 39, 40, 42, 43, Peireskia 4 45, 46, 47, 48, 49, 50, 54, 56, 57, 58, 93, Perescia 4 103, 107, 111, 113, 115*, 116, 118, 121 140 MEMOIRS OF THE NEW YORK BOTANICAL GARDEN [VOL. 41

var. grandifolia 12*, 14*, 22*, 27*, 28*, 30*, tampicana^, 113, 116 32,43, 50, 51,113, 114*, 115*, 116, 118 undulata 59 var. violacea 12*, 21, 27, 28*, 30*, 43, 50, vargasii 67, 69 51, 115*, 116, 117* var. longispina 69, 70 grandispina 127 var. rauhii 69, 70 guamacho 4, 6, 9, 13, 15, 16, 17*, 23, 30*, 31, var. vargasii 67, 69 34, 35, 36, 39, 40, 43, 46, 48, 56, 57, 58, verticillata 128 61,64, 65,85, 86*, 87*, 88, 89 weberiana 7*, 9*, 16, 17*, 20*, 22*, 29, 32, 33*, higuerana 127, 128 34, 36, 37*, 38, 39, 41*, 42, 43, 47, 50, 56, horrida 67 57, 58, 69, 73, 74*, 76 hortensis 66 zehntneri 128 humboldtii 13, 21, 31, 34, 36, 41*, 50, 56, 57, zinniaeflora 89 58, 66, 67, 68*, 69, 70* zinniiflora 5, 6, 9, 14, 24, 26, 31, 36, 39, 43, var. humboldtii 14*, 43, 67, 68*, 69, 70* 47, 56, 57, 58, 82, 89, 90*, 91*, 93, 97, 99, var. rauhii 35, 37*, 68*, 69, 70* 116, 126, 127 lanceolata 66, 128 sp. A 25, 28*, 31, 35, 45, 56, 57, 58, 95*, 126 longispina 5, 128 Pereskieae 42, 53 longissima 66 Pereskioideae 53 lychnidiflora 5, 6*, 7, 8, 9, 12*, 13, 14*, 15, 16, Pereskiopsis 5, 38, 39, 44, 51, 53, 128 17*, 19, 20*, 21, 22*, 23, 24, 25, 26*, 29, autumnalis 11 30*, 31, 33*, 34, 36, 37*, 38, 39, 40, 41*, opuntiiflora 76 43, 44, 46, 47, 48, 49, 56, 57, 58, 76, 78*, pititache 11 80*, 82, 83 rotundifolia 128 lychniflora 82 scandens 128 moorei 107, 111 spathulata 128 nemorosa 11, 14*, 15, 21, 22*, 24, 25, 28*, 30*, Phytolaccaceae 43 32*, 35, 36, 38, 40, 42, 43, 46, 50, 56, 57, Portulacaceae 43 58, 103, 104*, 106*, 107, 111 Quiabentia 31, 39, 44, 53 nicoyana 34, 46, 51, 77, 80, 83 pflanzii 128 ochnocarpa 113 verticillata 128 opuntiiflora 76, 82 zehntneri 128 panamensis 99, 103 Rhipsalis 23 pereskia 59, 65 Rhodocactus 4, 5, 54 pflanzii 128 antonianus 73, 76 philippii 128 autumnalis 11 pititache 34, 38, 77, 81 bleo 99 plantaginea 128 colombianus 85 poeppigii 128 conzattii 11 portulacifolia 6*, 8, 11, 12*, 15, 20*, 22*, 23, corrugatus 99 24, 25, 26, 28*, 29, 30*, 36, 40, 45, 47, 48, cubensis 47, 89, 92 49, 56, 57, 58, 93, 95*, 96*, 97, 126, 127 grandifolius 113, 118 quisqueyana 8, 9*, 22, 23, 24, 26, 31, 34, 35, guamacho 85 36, 39, 40, 41*, 45, 47, 48, 56, 57, 58, 95*, higueranus 128 97, 98*, 126, 127 horridus 67 rosea 113 lychnidiflorus 76 rotundifolia 5, 128 nicoyanus 11 rubescens 59 portulacifolius 93 saccharosa 111 sacharosa 107 sacharosa 8, 11, 15, 20*, 21, 24, 25, 26*, 29, saipinensis 109 30*, 31, 33, 35, 36, 38, 39, 40, 41*, 43, 45, tampicanus 113 50, 56, 57, 58, 106*, 107, 108*, 111 zinniiflorus 89 saipinensis 107, 111 Rosa 8 scandens 128 Sambucus 8 sparsiflora 76, 109, 111 Seguieria 43 spathulata 127, 128 Selenicereus innesii 47 stenantha 21, 22*, 24, 28*, 30*, 35, 36, 37*, Stenocereus 95 40, 41*, 43, 45, 46, 47, 48, 50, 56, 57, 58, Tilia 7 121, 122*, 124*, 125 Weingartia 16 subulata 128 1986] PERESKIA (CACTACEAE) 141

BEAT ERNST LEUENBERGER Dr. Beat Ernst Leuenberger was born 27 August 1946 in Burgdorf, Switzerland, where he also went to school. In 1963/64 he was an American Field Service high school exchange student to Portales, New Mexico. From 1966 on he studied botany, zoology, and chemistry at the universities of Berne, Switzerland (diploma 1972) and Heidelberg, Germany (Dr. rer. nat. 1975). The subject of his thesis was the pollen morphology of the Cactaceae. He was an assistant at the Systematic- Geobotanical Institute of the University of Berne for one year and in 1976 he became a staff member of the Botanical Garden and Museum of Berlin-Dahlem, where he has worked in the phanerogam herbarium and since 1979 mainly with the living collection of tropical and subtropical plants. His field studies have taken him to Togo, Mexico, Central America, the Dominican Republic, Brazil, Namibia, and Argentina. He contributed to the "Flore analytique du Togo" and is partic­ ipating in the Flora of the Guianas project. In 1985 he was appointed honorary research associate of the New York Botanical Garden. His main research interests are the taxonomy of Cactaceae, the biology of succulents, and neotropical flora and vegetation.