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Selbyana 19(2): 227-235 TAXONOMY OF THE GENERA ANANAS AND PSEUDANANAS-AN HISTORICAL REVIEW FREDDY LEAL Universidad Central de Venezuela, Facultad de Agronomia, Apartado 4736, Maracay, Aragua, Venezuela GEORGE COPPENS D'EECKENBRUGGE CIRAD-FLHOR/IPGRI, % CIAT, AA 6713, Cali, Colombial BRUCE K. HOLST Marie Selby Botanical Gardens, 811 South Palm Avenue, Sarasota, Florida 34236, USA ABSTRACT. From the first observations of the pineapple by European explorers to the classification into Pseudananas sagenarius and the seven Ananas species prevailing at present, the evolution of pineapple taxonomy has shown considerable variation. Most early botanists named or renamed species from previous dubious descriptions or from particular horticultural types. More recently, the genus Pseudananas was created, while horticultural types disappeared from the Ananas species. Subsequently, the total number of species increased again as botanical varieties and forms with minor variation were elevated to species rank. The resulting classification is questionable, as neither discontinuous morphological variation nor reproduc tive barriers exist in the genus Ananas. Ananas and Pseudananas are the only genera peduncle. The dry fibers constitute 6% of plant in the Bromeliaceae whose fused flowers devel weight (Camargo 1943). The absence of spines op into a sorose-type fruit formed by the coa facilitates manual fiber extraction, although lescence of up to 200 berries. The most recent some spiny or partially spiny mutants have been classification (Smith & Downs 1979) established observed. Ananas lucidus has never been found Pseudananas as a monotypic genus, Pseudan in the wild. anas sagenarius (Arruda) Camargo, and divided Ananas bracteatus (Lindl.) Schult. f. comes Ananas divided into eight species, one of which from southern South America (southern Brazil, has been invalidated (Leal 1990). The species Paraguay, and northern Argentina), where it is considered valid now are presented in TABLE 1. always found cultivated as a living hedge or for The best known species of Ananas is the cul fruit juice, or abandoned in ancient settlements. tivated A. comosus (L.) Merr., thanks to its im Its variegated form has been widely diffused as pressive and delicious fruit, which is considered a garden ornamental. The plant is vigorous with an important taxonomic character (more than 15 wide and long leaves, large spines, and suckers cm long according to the key in Smith & Downs abundantly. The inflorescence is characterized 1979). This large fruit is borne on a wide, short by its bright pink to red color and long bracts. to-long peduncle. In the spiny genotypes, spines The fruit and peduncle are medium-sized (ac are antrorse and generally smaller and denser cording to the key presented in Smith and than in other species. Partially spiny genotypes Downs (1979), the syncarp is more than 15 cm also exist as well as completely smooth geno long; however it is often less). Ananas bractea types, characterized by a folding of the lower tus, well adapted to cool conditions and altitude, epidermis over the leaf edge ("piping" character has been observed at 1,000 m in the subtropics. as named by Collins & Kerns 1946). The variability of this species is very limited. Ananas lucidus Mill., the curagua, is cultivat Ananas Jritzmuelleri Camargo is almost identical ed by the peoples of the Orinoco basin and to to A. bracteatus with some retrorse spines (Ca the North of the Amazon River. They use its margo 1943). Formerly included in A. bractea strong and long fibers to make breechcloths, tus (Smith 1939), A. jritzmuelleri has floral hammocks, fishing nets and rods (Leal & Amaya bracts that turn a pale green color at fruit ma 1991). This species is mainly characterized by turity. long, smooth, and erect leaves and a small, fi Ananas ananassoides (Baker) L.B. Sm., A. brous, inedible fruit borne on a medium-to-long parguazensis Camargo and L.B. Sm., and A. nanus (L.B. Sm.) L.B. Sm. are wild species. An anas ananassoides is the most widespread spe 1 Address for correspondence. cies, from southern Brazil to Venezuela and Co- 227 228 SELBYANA [Volume 19(2) TABLE 1. Current composition of the genera Pseu As commonly found in Bromeliaceae, the ge dananas and Ananas. Based on Smith and Downs nus Ananas is diploid, characterized by having (1979) and Leal (1990). 50 minute and almost spherical chromosomes in both root tips and pollen mother cells (Collins Scientific name Common name & Kerns 1931, Canpinpin & Rotor 1937, Mar Pseudananas sagenarius Gravata de cere a, gravata chant 1967, Sharma & Ghosh 1971, Lin et al. (Arruda) Camargo de rede, yvira, nana ca 1987, Brown & Gilmartin 1989, Dujardin 1991). ",aba, nana brava Giant unreduced gametes may appear and pro Ananas ananassoides Ananas de ramosa, curi duce natural triploids and tetraploids (Collins (Baker) L.B. Sm. bijul, maya pifion, na 1933, 1960). Most genotypes display reduced nlli, pifiuela fertility and a self-incompatibility system with Ananas nanus (L.B. Sm.) Ananai L.B. Sm. considerable variation in its expression (Cop Ananas parguazensis Ca Gravata, pilla montafiera pens d'Eeckenbrugge et al. 1993). Vegetative re margo and L.B. Sm. production is largely dominant over sexuality in Ananas lucidus Mill. Curagua, curana, curaua, Ananas. This is particularly true in domesticated kulaiwat types. Pineapples multiply by suckers (terrestrial Ananas bracteatus Ananas bravo, ananas do and aerial), slips (suckers from the peduncle), (Lindt) Schult. f. mato and crown. Ananas fritzmuelleri Ca Ananas silvestre, gravata Pseudananas sagenarius has been cultivated margo de cerca as a source of fibers, which explains the origin Ananas comosus (L.) Abacaxi, ananas, pilla, Merr. matzatli of its common name, yvira, meaning fiber, as well as of its Latin epithet, which refers to a fishing net (sagena). The fibers constitute 7.6% of the leaf dry weight and reach a length of 1.60 lombia. Although a few genotypes thrive in m (Correa 1952). In contrast to Ananas, P. sa dense rainforest (in the Guianas). it is generally genarius is a tetraploid (211 = 100), character observed in savannas or in low-shade forest, ized by a complete lack of crown and asexual growing well on soils with limited water-holding reproduction by stolons. Its leaf margins bear capacity (such as sand or rocks), and forming strong spines, which are retrorse at the leaf base. populations of variable densities. Most of these Floral bracts are longer than in Ananas como sus. populations are monoclonal, but some are poly The fruits are low in acid. The habitat of P. sa clonal, with variation of recent sexual origin genarius, limited to forest areas, is under semi (Duval et al. 1997). The plant has long and gen dense shade and subjected to a rainy season dur erally narrow spiny leaves and bears a small, ing most of the year, or even to periods of flood globular-to-cylindrical syncarp on a long and ing. This species may be found in southern thin peduncle. The fruit is often seedy, and its South America (southern Brazil, Paraguay, Bo pulp is white, firm and fibrous, with a high sugar livia and northern Argentina), and along the and acidity content, good flavor and aroma, and eastern Brazilian coast, up to Pernambuco. Pop a narrow heart. According to the key, A. anan ulations of P. sagenarius are rare now because assoides is distinguished from A. comosus by the of severely reduced habitat. The few populations size of the fruit (shorter than 15 cm). recently observed, however, showed significant Ananas nanus is characterized by an even variation (Ferreira & Cabral 1993, Ferreira 1996, smaller fruit (shorter than 4 em), In fact, Smith Duval et ai. 1997, Coppens d'Eeckenbrugge et (1939) fonnerly classified it as a dwarf variety al. 1997). of A. ananassoides. Ananas nanus has some times been used as an ornamental. HISTORY OF PINEAPPLE TAXONOMY Ananas parguazensis is also very similar to A. ananassoides but differs in the retrorse ori The main steps in the evolution of pineapple entation of some spines and a wider leaf slightly taxonomy are presented in FIGURE 1. constricted at its base. Ananas parguazensis The Amerindians had domesticated pineap seems less adapted to drought, but an anatomical ples and the curagua for fruit and fiber and had and physiological comparison showed no other dispersed the fonner all over Latin America and differences (Leal & Medina 1995). Its distribu the Caribbean and the latter from the North bank tion is also limited to the northern Amazon (Rio of the Amazon to the Caribbean well before the Negro basin) and Rio Orinoco, with a wider var anival of Columbus (Leal & Coppens d'Eeck iability in the Orinoco region (Duval et ai. enbrugge 1996). They also had domesticated the 1996). It grows in the lowland forests, under yvira, as observed by Thevet (1557) around Rio canopies of variable densities, from clearings or de Janeiro. Native Americans were the first pine riverbanks to dense forest. apple taxonomists. On the island of Hispaniola, ..... Plumier (1755) \0 Pseudananas Hassler ex Harms 1930 Ananas Miller 1754 \0 r< < ----- ----I <I Rumphius (1747) 23 XVIII Anassa domestica Anassa silvestris t t Linnaeus (1753) Bromelia ananas Bromelia comosa l' tIl I I Arruda da Ciimara (I 810) Bromelia Iuricata Bromelia sagenaria ? -----= >- Bromelik sylvestris«'<AKanassla bracteata Anana~sa debilis Ananassal!ucida Ananassal sativa Lindley (1827) l' tIl vell;zo 1825 t t ~ .., Ananas muricatus V Ananas sagenaria Ananas semiserratus Ananas sativus Schultes & Sch< (1830) >- XIX , Ananas macrodontes ~ Morren (1878) r-' Ananas sativus Mez(l892) :r: k sativus vaL br~cteatus ..... I ..,CIJ A. comosus Merril(l917) 0 (Ananas Section Pseudoananas) (Ananas Section Euanan~s) Hassler (1919) ::0 >-< ~ 0 '"Ij Ananas microcephalUS A bracteatus? A. g~."iri,c,~ti'W Bertoni (1919) Pseudananas macrodontes t Harms (1930) ~ (Ananas murica'tus) Ananas macrodontes A. sagenaria ~ Ac:mosus Mez(l934) kc< vaL microstachys Smith (1934) ~ ~ ~ V V:l Pseudananas sagenarius A.

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