Two New Species in Cuphea (Lythraceae), and a Note on Alzateaceae Shirley A. Graham Department of Biological Sciences, Kent State University, Kent, Ohio 44242 U.S.A. Abstract. Cuphea (sect. Melvilla) alaniana from dish purple, bulbous-based, minute (< 0.25 mm) central Mexico and C. (sect. Diploptychia) armata or 0.5-1 mm long; intemodes up to half as long as from Central America are described and illustrated. the subtending leaves. Leaves with petioles 5—10 The monotypic family Alzateaceae, long known mm long, becoming subsessile or sessile distally in from Peru and Bolivia, and more recently discov¬ the inflorescence; blades 25-100 mm long, 5-35 ered in Costa Rica, Panama, and Colombia, is here mm wide, membranous, crowded on the upper reported from Ecuador for the first time. Of the two stem, narrowly lanceolate-linear, base acute, apex Ecuadorian collections, one belongs to the north¬ acute; blade surface minutely scabrous, or scabrous ern, broad-leaved Alzatea verticillata subsp. ampli- with sparse to abundant white, weak, slightly ap- folia; the other is most similar to the southern sub¬ pressed hairs to 1 mm long, the margins of young species verticillata. Apical leaf glands are a newly leaves densely ciliate, the hairs erect, glandular- recognized feature of the genus. They are inter¬ viscous, to 1 mm long; upper leaves very gradually preted as extra-floral nectaries. diminishing in size to become the lanceolate-linear bracts of the inflorescence. Inflorescences densely In revisionary studies of Cuphea, two new North flowered, terminal, leafy racemes; flowers solitary American species were discovered among collections at the nodes, interaxillary; pedicels 4—6 mm long; on loan from a number of North American herbaria. bracteoles 1.5 mm long, lanceolate-linear or re¬ Cuphea alaniana is a large-flowered perennial of duced to a few fleshy hairs, positioned 1.0-1.5 mm section Melvilla, subsection Polyspermum (Koehne, below the apex of the pedicel. Floral tubes 25-32 1903). It is one of four species of the subsection, mm long, including a rounded, horizontal base ex¬ which is endemic to the western and southern moun¬ tending 1 mm beyond the pedicel, 4—5 mm wide in tains of Mexico. Cuphea armata, discovered in lateral view at anthesis, dorsally slightly convex, neighboring areas of Guatemala, El Salvador, and the neck slightly or not at all narrowed, the mouth Honduras is a new species of section Diploptychia. flaring, oblique in lateral view, the ventral side ex¬ tended 1 inin or less beyond the dorsal side, the Cuphea alaniana S. A. Graham, sp. nov. TYPE: floral tube more distinctly convex and distended in Mexico. Guerrero: Distrito Adama, Temisco, fruit, the neck contracted; outer surface reddish Barranca del Consuelo, 500 m, 15 Nov. 1937, purple, straw-colored ventrally at the mouth, glan¬ Mexia 8810a (holotype, UC; isotype, GH). Fig¬ dular-puberulous and glandular-hispid with abun¬ ure la-d. dant, erect, thin, colorless or reddish purple hairs Suffmtices vel frutices, probabiliter ad 1.5 m, caulis to 1 mm long; inner surface pale, neither bialate simplex, erectus, viscosissimus. Petioli 5-10 mm longi, ad nor vesiculate, glabrous or lightly villous above the summum caulis sessiles; folia 25-100 mm longa, 5-35 stamens, glabrous below; calyx lobes subequal, the mm lata, anguste lanceolata vel linearia. Flores 25-32 mm dorsal lobe deltoid, 2X3 mm, the others 1X2 longi. rubro-purpurei, glanduloso-hispidi; lobus dorsalis calycis ceteris paullo major; appendices prominentes, se- mm, margins not ciliated; appendages at the sinus¬ tosae. Petala 6, roseo-purpurea, inaequalia, 2 dorsalia 7- es of the lobes prominent, 1 mm long, equaling the 9 mm longa, manifeste unguiculata, 4 ventralia 1.0-2.5 calyx lobes, thick, green or reddish purple, the mm longa. Ovula 40-50. Discus 2-3 mm longus, 2.5 mm apex bulbous, crowned by 5-8 setae 1—2 mm long. latus, 2 mm crassus, basi ovarii late affixus. Petals 6, rose-purple, the 2 dorsal 7—9 mm long Suffrutescent or shrubby perennials, probably including a slender claw 1-2 mm long, narrowly 0.5-1.5 m tall, stems simple, erect, from a fibrous- obovate, the 4 ventral 1.0-2.5 mm long, linear-spa- rooted base, branched distally in the inflorescence; thulate, caducous. Stamens 11, all inserted at % the stems and pedicels highly viscous, the vestiture length of the floral tube, the 2 dorsalmost shortest, densely, minutely glandular-puberulous and glan¬ included, the 5 antesepalous exserted, the 4 ante- dular-hispid, the hairs thin, erect, colorless or red- petalous included to subexserted; filaments gla- Novon 5: 272-277. 1995. Volume 5, Number 3 Graham 273 1995 Cuphea Figure 1. a-d, Cuphea alaniana S. A. Graham, —a. Habit. —b. Lateral view of flower. —c. Nectariferous disc at base of the ovary within the floral tube. —d. Floral tube opened dorsally, ovary removed to show position and number of stamens. —e. Cuphea armata S. A. Graham, lateral view of flower. Drawn from the holotypes. brous to lightly villous, reddish; anthers purple. ly, the gibbosity pronounced in flower; ovules 40- Pollen triangular in polar view; tricolporate, syn- 50. Seeds numerous, 2 mm long, 1.5 mm wide. Disc colpate, pores not protruding; exine uniformly fine¬ massive, 2-3 mm long, 2.5 mm wide, 2 mm thick, ly striate, the striae extending ca. Vi the distance horizontal, oblong-rectangular in dorsal view, to the poles, partially anastomosing; diameter 24- broadly attached at the base of the ovary. 28 gm in lactic acid. Style exserted at maturity, Phenology. Collected in flower and fruit in No¬ glabrous; stigma punctiform; ovary gibbous dorsal¬ vember. 274 Novon Figure 2. Distribution of Cuphea alaniana and C. armata in Mexico and Central America. Distribution and habitat (Fig. 2). In southern xaca by its slightly enlarged dorsal lobe and large Mexico state and adjacent Guerrero; low spiny, de¬ petals, but C. heteropetala lacks a gibbous ovary ciduous woods, dry slopes, and sandy streamsides; and has a small, erect disc. Attempts to relocate C. 500-700 m. alaniana in the states of Mexico and Guerrero were Etymology. The species is named after Alan unsuccessful. The type area, along the river near Graham, who has generously and ably assisted on Temisco, Guerrero, is now occupied by dwellings field excursions many times since our first trip to and cultivated fields. Exploration further upstream Mexico for Cuphea in 1961. in less disturbed sites also failed to locate the spe¬ The recognition of Cuphea alaniana adds a cies. The paratype locality, directly north of Tem¬ fourth species to subsection Polyspermum of section isco in Mexico, was not accessible by road in 1989, Melvilla. Species of the subsection are endemic to and was not visited by us. the western and southern mountains of Mexico. The subsection is characterized by spurless, deeply col¬ Paratypes. MEXICO. Mexico: Cerro de San Antonio, ored, thick-bodied floral tubes 20-32 mm long and Tlatlaya, Matuda 32110 (GH, MEXU). ovules numbering 40-120. In C. alaniana, the red¬ dish purple floral tube with two large erect, clawed Cuphea armata S. A. Graham, sp. nov. TYPE: El dorsal petals, four small ventral petals, and a mas¬ Salvador. Chalatenango: 4 km SSE of La Pal¬ sive horizontal nectary termed the “disc” serve best ma, pine forest, 950 m, 11 June 1970, G. Dav- to distinguish it from C. micropetala var. hirtella idse & R. Pohl 2073 (holotype, F; isotype, Koehne, to which it is surely most closely related. MO). Figure le. The latter differs in having six minute petals of Suffrutices, 0.5-1 m, caules erecti vel suberecti, pilis equal size and a small disc. Both species share an minutis albidis antrorsis adpressisque. Folia sessilia vel unusual, dorsally gibbous ovary in addition to sim¬ subsessilia, 20-50 mm longa, 5-12 mm lata ad medium ilar vegetative, trichome, and pollen morphology. caul is, ovata, oblonga, vel sublinearia, basi obtusa vel cor- Within the subsection, C. alaniana also resembles data, utrinque scabrella. Inflorescentia sat distincta, ra- cemosa, interdum subcomposita; pedicelli gracili, 2-5 mm the poorly known C. heteropetala Koehne from Oa¬ longi, persistentes; bracteolae 0.5 mm longae, lineari-lan- Volume 5, Number 3 Graham 275 1995 Cuphea ceolatae. Flores 12—15 mm longi, pallidi purpureo-rosei, angular to comiculate, thickened scale free at the basi puberuli saepe prominente bulboso-setosi, calcari 4 tip for 0.2 mm, the 4 ventral petals 3-5 mm, oblong mm longo, distincte ascendente etiam conico; intus infra stamina bialatus, alae glabrae. Petala 6, 2 dorsalia intense to obovate, pale purple, held horizontally. Stamens violacea, majora, 7—9 mm longa; 4 ventralia minora 3-5 11, the 2 dorsalmost stamens shortest, included, 9 mm longa, pallide violacea. Stamina 11, 9 longiora exser- inserted at ca. 2-3 mm below the rim of the floral ta. Stylus demum exsertus; ovula 8—10. Semina 2 X 1.5 tube, exserted; filaments probably pale purple, gla¬ mm, oblonga, margine obtusa. Discus crassus, deflexus. brous; anthers purple. Pollen oval-triangular to Semi-shrubby perennials, 0.5—1 m, erect or nearly spherical, pores very slightly protruding; tri- weakly erect and sprawling, irregularly branched colporate, syncolpate; exine distinctly and uniform¬ from above the base, branches shorter than to lon¬ ly striate, striae covering the surface of the grain, ger than the main stem, bearing minute white, op¬ and possibly extending to the poles without disag¬ pressed, antrorse, sharp-pointed hairs and a lon¬ gregation; diameter 18-20 gm in lactic acid. Style gitudinal row of minute, falcate, antrorse hairs; ultimately well-exserted; stigma punctiform; ovules intemodes mostly equal to or shorter than the sub¬ 8-10. Seeds 2 X 1.5 mm, oblong, brown, margin tending leaves. Leaves sessile or subsessile, petioles thick, rounded. Disc 1 mm long, 0.5 mm wide at 0-1 mm; blades at midstem 20-50 mm long, 5—12 base of ovary, thick, triangular in outline, deflexed mm wide, decreasing in size above and below the into the spur. midstem, membranous, ovate, oblong, or sublinear, Phenology. Collected in flower and fruit in base rounded to cordate, apex acute to acuminate; June, September, and December. surfaces finely scabrous, appearing glabrous, the Distribution and habitat (Fig. 2). Southeastern hairs minute, appressed, the margins of the upper Guatemala, Honduras, and El Salvador; pine for¬ leaves and bracts ciliolate, bearing sparse bulbous- ests; 900—1640 m. based setae, upper surface darker green than the Etymology. The specific epithet refers to the lower; upper leaves gradually diminishing in size red-purple, bulbous setae that arm the floral tube to become the bracts of the inflorescence. Inflores¬ and upper bracts of the inflorescence. cences terminal bracteolate racemes or loose, short- The species is placed in section Diploptychia by branched thyrses; one flower interaxillary at a node, the presence of the key character of the section, 1—3 additional flowers sometimes present on slen¬ the alae, two wings of tissue that extend from the der, alternate, axillary branchlets ca. 1 cm long; base of the dorsalmost stamens in the floral tube to pedicels 2—5 mm, slender, persistent; bracteoles the base of the tube (Koehne, 1903). Cuphea ar- 0.5 mm, linear-lanceolate, persistent at the apex of mata bears close resemblance to Cuphea pinetorum the pedicel. Floral tubes 12—15 mm long, including Bentham and C. avigera Robinson & Seaton but an ascending, conical spur 4 mm long, the apex differs from both by its unequal, bicolored petals. sometimes slightly descending, floral tubes dorsally The prominent reddish, bulbous-based setae and concave, 1-2 mm wide at anthesis, in lateral view pale purple floral tube further distinguish it from the base wider than the midportion, the neck not C. pinetorum, which has a white to rose, hirsute and strongly contracted at anthesis, the mouth oblique, pubescent floral tube. Cuphea armata resembles C. the ventral side extended 1 mm or less beyond the avigera especially by its conical, ascending spur, dorsal, the dorsal and adjacent lateral ribs diverg¬ but lacks the glandular-viscous patches on the ing approximately 1 mm; outer surface pale rose- stems that distinguish C. avigera. Neither C. pine¬ purple, darkest dorsally, puberulent with sparse but torum nor C. avigera has been collected within the prominent red-purple, bulbous-based setae present range of C. armata. Because agamospermy with ac¬ on the dorsal and lateral ribs, especially toward the companying infertile pollen is present in several base of the tube, rarely individual flowers lacking species of the section, pollen of C. armata was test¬ setae, setae 0.5 mm long; inner surface lightly vil¬ ed for pollen viability. In cotton blue-lactic acid lous to glabrous above the stamens in a shallow after 24 hours, 99% of all pollen stained deeply, ventral pouch, glabrous below, the alae glabrous; indicating viable protoplasm. The pollen exine calyx lobes subequal, broadly triangular, 1.0 X 1.5 seems to lack the meshwork of striae at the poles mm, margins of the dorsal lobe setulose-ciliolate, that is typical of the pollen group to which C. avi¬ rarely glabrous, the others glabrous; appendages gera and C. pinetorum pollen types belong (Gra¬ 0.5 mm long, terminating in a short bristle. Petals ham, in prep.), but this needs to be confirmed by 6, unequal, the 2 dorsal largest, 7—9 mm long, ob¬ SEM observations when more abundant material long to nearly orbicular, not clawed, deep purple, becomes available. held erect or reflexed slightly back over the dorsal side of the tube, subtended at the base by a tri¬ Paratypes. EL SALVADOR. Chalatenango: carretera 276 Novon Figure 3. Distribution and variation in leaf shape, size, and petiole length in Alzatea verticillata. Examples 1-5 represent subspecies amplifolia, characterized by large, broad, oblong, sessile or subsessile leaves. Examples 6-8 represent subspecies verticillata, characterized by small, elliptical, oblong or obovate, distinctly petiolate leaves. Petioles on plants represented by examples 5 and 6 are approximately intermediate between the two subspecies. The major distinction remaining between the subspecies is leaf size, especially leaf width. a La Palma, Montalvo 3682 (MO); Rfo Nunuapa near La 1986, was found in Colombia on the Cerro del Tor- Palma, R. Seiler 574 (F). GUATEMALA. Chiquimula: pine forest between Guatemala-Honduras border and Atu- rd, Departamento del Choco (Silverstone-Sopkin & lapa, Molina R. * Molina 25260 (F, MO). HONDURAS. Graham, 1986). The distribution of the species is Intibuca: 12.7 mi. N of Marcala along the road from Mar- now further enlarged by collections made by the eala to La Esperanza, Davidse et al. 35191 (MO). late Alwyn Gentry on his last field expedition to Ecuador. Two collections. Gentry 80008 and 80334 Note on Alzateaceae (MO), from the Cordillera del Condor, near the dis¬ The New World tropical family Alzateaceae is puted Ecuador-Peru border in Morona-Santiago represented by the single species Alzatea verticil¬ Province, represent the first record of the family in lata Ruiz & PavtSn. The species had been placed Ecuador, and tenuously establish its continuous in eight families in five orders, including the Lyth- distribution in mid- to low-montane forests along raceae, before its segregation to the monotypic fam¬ the eastern slopes of the mountains from Bolivia to ily Alzateaceae (Graham, 1984). Alzatea verticillata Costa Rica (Fig. 3). The species is one of an esti¬ was described first from the eastern side of the Pe¬ mated 26% of Ecuadorian species restricted to ruvian Andes in 1798 (Ruiz & Pavdn, Syst. Veg. mid-elevations (900-3000 m) with widespread dis¬ FI. Peruv. Chil. 1: 72). It was considered restricted tribution in the Neotropics (Balslev, 1988). to Bolivia and Peru until 1936, when it was col¬ The two Ecuadorian collections differ in leaf lected in the low montane forests of Costa Rica. In shape and size, and represent two subspecies. Gen¬ 1978, it was discovered in the elfin cloud forest of try 80334, collected from an 18-m-tall tree at 2100 Veraguas Province, Panama, and subsequently, in m, is typical of the elliptical to oblong or obovate, Volume 5, Number 3 Graham 277 1995 Cuphea petiolate, small-leaved subspecies verticillata from pamento Achupalla, 15 km E of Gualaquiza, 03;27'S. Peru and Bolivia (Fig. 3). Gentry 80008, collected 78°22'W, dense tangled scrubby forest, 22 July 1993, A. Gentry 80334 (MO). from a 5-m-tall tree at 1350 m, has the broad, ob¬ long, large leaves of subspecies amplifolia, until Acknowledgments. I thank Ishmael Calzada for now known from Colombia, Panama, and Costa his energetic assistance in the field during our un¬ Rica. Both collections have short-petiolate leaves successful attempts to relocate C. alaniana. I am and are intermediate for this character between the also grateful to Ronald Liesner, who first deter¬ distinctly petiolate-leaved subspecies verticillata mined the Alzatea collections, and whose intimate and the sessile-leaved subspecies amplifolia. The knowledge of plant families has resulted in spe¬ distinctions between the subspecies, which were cialists everywhere receiving many interesting and described before the species was known from Co¬ important specimens that otherwise might have lombia or Ecuador (Graham, 1984), are diminished gone undetermined or unnoticed among exsiccatae. by the partial intermediacy of the Gentry collec¬ Wendy Mahon-Hils prepared the illustrations in tions. If future collections of Alzatea from northern Figure 1. This research was supported by NSF Peru and Ecuador sample a fuller range of leaf grant DEB 8806523. shape and size, it should be easy to decide whether or not to continue recognition of these intraspecific Literature Cited taxa. Balslev, H. 1988. Distribution patterns of Ecuadorean plant species. Taxon 37: 567—577. Leaves of Gentry 80334, among all collections of Graham, S. A. 1984. Alzateaceae, a new family of Myr¬ Alzatea, are the first to display apical leaf glands. tales in the American tropics. Ann. Missouri Bot. Gard. The glands are similar to the hydathodes and extra¬ 71: 757-779. floral nectaries described for several other genera Koehne, E. 1903. Lythraceae. Pp. 1-326 in A. Engler (editor). Das Pflanzenreich, IV. 216. Heft 17. H. R. En- of the Myrtales (Ross & Suessenguth, 1926; Rao & gelmann, Leipzig. Chakraborti, 1982; Turner & Lersten, 1983; Gra¬ Rao, T. A. & S. Chakraborti. 1982. A little looked at ham, 1984). The shiny, hardened secretion at the attribute of the leaves of Sonneratia caseolaris (L.). Curr. Alzatea leaf apex suggests the secretory product is Sci. 51: 303-305. Ross, H. & K. Suessenguth. 1926. Das Apikalorgan der a sugar solution and the gland is probably an extra¬ Blatter von Lafoensia. Flora 120: 1—18. floral nectary. Silverstone-Sopkin, P. A. & S. A. Graham. 1986. Alza¬ teaceae, a plant family new to Colombia. Brittonia 38: Specimens examined. ECUADOR. Morona-Santia- 340-343. go: Gualaquiza Cantdn, Cordillera del Cdndor, ridge top Turner, G. W. & N. R. Lersten. 1983. Apical foliar nec¬ above Banderas, primary forest, 03°28'S, 78°22'W, pri¬ tary of pomegranate (Punica granatum: Punicaceae). mary forest, 17 July 1993, A. Gentry 80008 (MO); Cam- Amer. J. Bot. 70: 475-480.