Pereskia lychnidiflora DC.

Authority

Leuenberger, Beat E. 1986. Pereskia (Cactaceae). Mem. New York Bot. Gard. 41: 1-140.

Family

Cactaceae

Scientific Name

Pereskia lychnidiflora DC.

Description

Species Description - 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, dichotomous to trichotomous branching (Figs. 1A, 5A); 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, variously 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, longitudinally fissured mainly above and below the accrescent areoles. Roots imperfectly 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, sometimes 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, trichomes 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 1-4(-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 1/10 to 1/5 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 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. 1020 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 1018, 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: 2n = 22; Leuenberger & Schiers 2508, cult. hort. Berol. 003-04-78-10 (B).

Discussion

Type. Plate of “Cactus fimbriatus” in Mociño, Fl. Mex. ined. no. 1689, preserved at the Hunt Institute for Botanical Documentation, Pittsburgh. A copy of the plate was used by De Candolle to illustrate the species on pl. 18 of the publication cited above. “Legionensis Cactus caule arborescente foliis ovatis acuminatis spinis alternis longissim-is. . . . Nicaraguae montibus . . .” in Mociño’s manuscript of the Flora of Guatemala (no. 13, 4a div. in the Instituto Botánico Cavanilles in Madrid) apparently refers to the same, but an equivalent specimen was not located. Rhodocactus lychnidiflorus (De Candolle) F. Knuth in Backeberg & Knuth, Kaktus-ABC 97. 1935. Pereskia opuntiiflora De Candolle, Mém. Mus. Hist. Nat. 17: 76, t. 19. 1828. Type. Plate of Cactus opuntiaeflorus in Mociño, 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 Mociño’s manuscript of the Flora of Guatemala (no. 13, 4a div. in the library of the Instituto Botánico Cavanilles in Madrid) evidently refers to the same. Equivalent specimen. Nueva España, 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 opuntiiflora). 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. Progreso: 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. 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 certainly belongs to this species growing in a coastal forest at Salinas, Guanacaste. Living twigs of P. lychnidiflora collected in NW Guanacaste (Gómez, pers. comm.) confirm this record. Local names and uses. Mexico: guititache, guichitache, guitache, patilón, cruz del matrimonio, arbol del matrimonio (Oaxaca); cuncú, cuncu marin (Guerrero). Guatemala: manzanote, matial. Honduras: mateado, amarilla. 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 animals. 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 name P. lychnidiflora as correct for the plant occurring in Mexico, but she maintained 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 “dentés ou frangés à leur sommet,” i.e., “dentate or fringed at the tip.” The comparison of the two plates of the Flora Mexicana ined. of Mociño, on which P. lychnidiflora and P. opuntiiflora are based, would not suggest the inclusion 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 accordingly 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 España, Herb. Pavon, and annotated by J. N. Rose in 1912 as “Pereskia lychniflora According to McVaugh (1977) and Bemardi (1977), specimens from the Sessé and Mociño collection were sold by Pavón 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 Mociño 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 with 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 Mociño 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 Mociño’s plates refer to plants of Central American or even West Indian origin. From the descriptions in the Flora of Guatemala manuscript of Mociño kept at the Instituto Botánico 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 expedition party may have seen this cactus earlier around Tehuantepec and on the 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.

Distribution

Distribution (Fig. 30) and phenology. Southern Mexico to Costa Rica. In lowland 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.

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