Three New Species of Jaltomata (Solanaceae) from Ancash,

Thomas Mione Department of Biological Sciences, Central Connecticut State University, New Britain, Connecticut 06050-4010, U.S.A.

Segundo Leiva G. Museo de Historia Natural, Universidad Antenor Orrego, Avenida America Sur 3145, Trujillo, Peru

Leon Yacher

Department of Geography, Southern Connecticut State University, New Haven, Connecticut 06515-1355, U.S.A.

ABSTRACT. Three new Jaltomata species from the served flowers were used for drawing, while dried department of Ancash, Peru, are described and il- specimens were used for measurements. Seeds of lustrated. The three species are distinguished from J. lomana were not available for study. others in the genus by features of the flowers, hairs, Trichomes are not gland-tipped unless indicated and leaves. Fruits of Jaltomata cajacayensis S. Lei- as such. Finger hairs are uniseriate, unbranched, va & Mione are gathered for consumption. Jalto- and multicellular. Branchlet hairs (Fig. 3B) have mata lomana Mione & S. Leiva is known only from multiple termini (Seithe, 1979). A gland-tipped fin- a single fog-dependent plant community, a lomas ger hair has a bulbous terminal cell (Fig. 1C) that formation. Jaltomata yungayensis Mione & S. Leiva stains densely with neutral red, as does the mul- is widely distributed at high elevations. ticellular head of a stalked gland (illustrated in Mione & Serazo, 1999). Species of Jaltomata have simple, often gemi- nate leaves, 5-merous flowers with rotate to cam- Jaltomata cajacayensis S. Leiva & Mione, sp. panulate or tubular corollas, and a bicarpellate ova- nov. TYPE: Peru. Ancash: Bolognesi, highway ry housing numerous ovules. The inflorescence is from Pativilca to Recuay and , km 90, umbellate; anthers dehisce longitudinally and the town of Cajacay, 2540 m, open sun, roadside ovary is girdled by a disk at its base. The berry, with Agave andina and Carica candicans, 18 orange or red on most South American species and Jan. 1998, T Mione, S. Leiva G. & L. Yacher dark purple to black on most Central American 624 (holotype, NY; isotype, CONN). Figure 1. species, is not enclosed by the calyx and is often Planta fruticosa; axes juvenes, folia, faciesque abaxialis edible (Leiva, 1998; Mione, 1992). The genus in- calycis obtecta pilis indivisis apice glandiferentibus; folia cludes about 50 herbaceous and shrubby species ovata vel deltata, ad 9 cm longitudine, 10 cm latitudine, distributed from Arizona, U.S.A., to Bolivia, with petiolo ad 3.2 cm longitudine; inflorescentia floribus 14 one species each on the Galapagos Islands and ut maximum; calyx 13 mm diametro maturitate fructus; corolla brevissime, tubulosa, limbo reflexo 10.3-19.2 mm Greater Antilles (Mione et al., 1993). These de- diametro, olivacea; filamenta villosa secus proximales 4/5 scriptions of three new species are a contribution partes longitudinis; bacca aurantiaca, ad 9 mm diametro. to ongoing systematic studies of this genus (e.g., Mione, 1999; Mione & Leiva, 1997). Shrub to 1.1 m high. Young axes, peduncles, pedicels, and abaxial faces of calyx villous with MATERIALS AND METHODS erect, gland-tipped finger hairs to 3 mm long. Woody stems glabrous, terete, to at least 1.5 cm The compound light microscope was used for diam. Leaves alternate, often geminate, the blade measurement of stalked glands and stigma papillae; ovate or deltoid, to 9 x 10 cm, pubescence of flowers placed in 70% ethanol in Peru were used. gland-tipped finger hairs on both faces, the margin The sizes of structures as indicated by the scale repand, sinuate or dentate with up to 5 pairs of bars in the figures may differ somewhat from the broad, rounded teeth; petiole to 3.2 cm long. Inflo- sizes given in the descriptions because alcohol-pre- rescence axillary or less commonly arising from a

NovoN 10: 53-59. 2000.

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stem dichotomy, agricultural umbellate, fields. One 6- to woman 14-flowered. told us Pe- that a tea duncle tomade 15 mmfrom long, leaves pedicel and flowers to 20 ismm used long. to treat Ca- stom- lyx green ach at ache anthesis, and children's 3-4 mm diarrhea,from pedicel and toto tipbring reg- of lobe, ularity ciliate towith the bothmenstrual branchlet cycle. hairs The Quechuato 0.25 name mm long is and"musho." gland-tipped Flowering finger in January, hairs to and1 mm flowering long; adaxially and fruiting pubescent in November. with finger hairs to 0.4 mm long; Jaltomata calyx at fruitcajacayensis maturity is similar(from Leivato J. 1739)propinqua stellate and(Miers) 13 mm Mione diam. & Corolla M. Nee short-tubular in that both with bear gland- a reflexed tipped limb, hairs dull andolive-green, have a short-tubular the tube (obscured corolla with by pressing) a much 1.5-2 broader X 6.5-8 limb. mm, Jaltomata within cajacayensisthe tube has a two dark green green corolla maculae and cream-coloredflanking the radialanthers, vein the style extending is exserted to the tip a few of eachmillimeters corolla lobe,beyond narrow the stamens, purple ringand itsurrounding is known only mouth from of the tube, vicinity the limbof one mon- 10.3-19.2 tane mm town diam., (Cajacay) the 5 lobesin the alternating department with of Ancash; lobules, J. adaxially propinqua glabrous has blue within to purple tube corollasand the and an- limb pilosulose, thers, the the style hairs is 0.21-0.5twice the mm length long, of abax- the stamens, ially with and finger it is widelyhairs, somedistributed gland-tipped, in the 0.15-1.2department of mm long, , ciliate Peru with (Mione finger et hairs al., 1993).to 0.6 mm long. Stalked multicellular glands 75-90 Im long, abun- dant on Paratypes. both faces PERU. of calyx Ancash: and Bolognesi, abaxial corolla, Cajacay, 2600 m, 18 Nov. 1995, Leiva 1739 (AAU, CCVS, CORD, FK uncommon on adaxial corolla. Stamens 7 mm long; HAO(), HUT, MO, NY, USM), 18 Jan. 1998, Leiva et al. filaments 21.32 villous (F, HAO). on proximal V3 of the length, the finger hairs purple, to 1.5 mm long, decreasing in length distally;Jaltomata anthers lomana cream-colored, Mione & S. 1-2.1 Leiva, mm sp. nov. long. Style TYPE: 3.5-10 Peru. mm Ancash: long; stigma Casma, capitate, Lomas de dark Mon- green and gon, exserted 9037'S, to 78043'W, 5.5 mm beyond450 m, anthers17 Sep. 1938,on living plants, H. E. Stork,0.5-0.9 O.mm B. wide, Horton the & papillae C. Vargas 45 tmC. 9183 long. Nectar (holotype, disk broad, K; isotypes, girdling G, base GH). of Figure the ovary 2. (Fig. 1H). Gynoecium glabrous. Mature berries or- ange (not Planta seen, fruticosa; from informant axes juvenes, at folia, type calyx locality), fere glaber; nearly spherical,folia ovata ad5-9 6 cmmm longitudine, diam. Seeds 3.4 (Leivacm latitudine, 1739) petiolo ad 7 mm longitudine; inflorescentia floribus 7 ut maxi- numerous, subtriangular to reniform, alveolate, mum; calyx 8-11 mm diametro, fructu; corolla breviter 1.4-1.7 Xtubulosa, 1.1-1.3 limbo mm, patente0.5-0.55 11-14 mm mm thick. diametro, alba; fila- menta villosa secus proximale dimidium longitudinis; bac- Uses, habitat, ca ad 6 mm diametro. local name, phenology. When the type collection was made, two women told us that the fruits Shrub to 8 dm tall.are Young axes, peduncles, sweet ped- and commonly eaten by every- one, and icels, leaves, thatand calyx nearly glabrous, thethe finger plants are not deliberately culti- vated but hairs to 0.15 mm longare where present. Larger common in and along the edge of

Figure 1. Jaltomata cajacayensis S. Leiva & Mione. -A. Branch with inflorescences and leaves. -B. Flower, oblique view. -C. Calyx, lateral view. -D. Corolla, overhead view showing basal adnation of stamens. -E. Stamen, ventral view, with abundant finger trichomes, base flattened by adpression to ovary. -F, G. Anther and distal portion of filament, lateral and dorsal views, respectively. -H. Gynoecium including basal disk. -I. Berry with calyx, lateral view. -J. Seed. (From S. Leiva et al. 2132, drawn by S. Leiva.)

Figure 2. Jaltomata lomana Mione & S. Leiva. -A. Branch with inflorescences and leaves, with segment of stem 2 mm wide to left. -B. Flower, lateral view. -C. Calyx, lateral view. -D. Corolla, dissected, overhead view showing basal adnation of stamens, with marginal hair 0.12 mm long to left. -E. Stamen, ventral view, with abundant finger hairs, base flattened by adpression to ovary. -F, G. Anther and distal portion of filament, lateral and dorsal views, respectively. -H. Gynoecium including basal disk. (From S. Leiva et al. 2140, drawn by S. Leiva.)

Figure 3. Jaltomata yungayensis Mione & S. Leiva. -A. Branch with inflorescences and leaves. -B. Flower, lateral view, with branchlet hair 0.15 mm long at lower right. -C. Corolla, dissected, overhead view showing basal adnation of stamens. -D. Stamen, ventral view, with finger trichomes, base flattened by adpression to ovary. -E, F. Anther and distal portion of filament, lateral and dorsal views, respectively. -G. Gynoecium including basal disk. -H. Berry with calyx, lateral view. -I. Seed. -J. Embryo from seed. (From S. Leiva et al. 2138, drawn by S. Leiva.)

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branches Planta glabrous, fruticosa; terete. axes Leavesjuvenes, alternate, folia, faciesque often abaxialis calycis puberula, pilis ramosis vel simplicibus; folia lan- geminate, blade elliptical to ovate, to 6 x 3.4 cm, ceolata vel elliptica, ad 6 cm longitudine, 2.1 cm latitu- the margin dine, entire, petiolo ciliolate ad 2 cm with longitudine; finger hairsinflorescentia to 0.15 floribus mm long, 11 utthe maximum; apex acute, calyx the 10 mmbase diametro symmetrical maturitate to fructus; somewhat corolla oblique; campanulata petiole tovel 7 rotata, mm long. 11.5-15.5 Inflores- mm diametro, cence axillary purpurea, or lessfauce commonly viride; filamenta arising villosafrom secusa stem proximalem dichotomy, Y, partem umbellate, longitudinis; 5- to 7-flowered.bacca aurantiaca, Peduncle ad 8 mm dia- metro. 3-8 mm long; pedicel 5-14 mm long. Calyx at an- thesis green, 3-4.8 mm from pedicel to tip of lobe, Shrub to 2.3 m high. Young axes, peduncles, ciliolate with finger hairs to 0.1 mm; 8-11 mm pedicels, leaves, and abaxial faces of calyx puber- diam. on mature (?) fruit. Corolla short-tubular with ulent, with both branchlet and finger hairs to 0.15 a spreading limb, white, the tube 1.6 x 4.0-4.3 mm long. Woody stems brown, terete and hollow, mm at base (obscured by pressing), the limb 11- 14 mm glabrousdiam., with age. Leaves alternate, often 5-lobed, geminate, adaxially with finger hairs 0.12 mm the blade long,lanceolate to elliptical, to 4.9(-6)erect x and abundant on limb, absent in tube, 2.1(-4.8) abaxially cm, margins entire to repand, apex usu- with finger hairs 0.09 mm long, erect and ally acute, the baseconcentrated of some leaves somewhat on veins, margins ciliate with both oblique; petiole tofinger 2 cm long. Inflorescence axillary and branchlet hairs 0.12 mm long. Stalked or lessmulticellular commonly arising from a stem dichotomy, glands 55-75 Am long, abun- dant on umbellate, both sometimes branched, 4- tofaces 11-flowered. of the young leaves and calyx, and the Peduncle abaxialto 10 mm long; pedicel to 9 mm long. face of the corolla (Mione et al. 631). Stamens Calyx green at anthesis, 2.5-3 mm from pedicel5.5-6.5 to mm long; filaments villous on proximal tip of lobe, adaxially glabrous, %,margins ciliate withfinger hairs to 0.75 mm long; an- thers 0.6-1.5 finger hairs to 0.25 mm long; at fruit maturitymm to 10 long. Style 4.7-5.4 mm long, slender mmand diam. Corolla campanulate straight; to rotate, 11.5-15.5 stigma capitate, shallowly bi- lobed, 0.36-0.51 mm diam., 5 lobes alternating with inconspicuous mm wide, the papillae (Mione et al. 631) lobules 20-25 (obscured by pressing); dark purple Am with a long. Nectar disk at base of ovary. Berries pale green throat having 5 pairs of dark-green (mature?) mac- 4-6 mm diam. Distribution ulae in a ring, each pair of maculae straddlingand the habitat. Known only from a single small main vein that extends radiallymountain to the tip of the on the coast, the plant com- munity corolla of lobe, abaxial facewhich pubescent with finger is fog-dependent and therefore known hairs,as and a few abranchlet lomas hairs, to 0.9 mm but formation (Rundel et al., 1991). Flowering commonly 0.3 mmin long, adaxial faceJanuary glabrous, mar- and September, and fruiting in September. gins off-white, ciliate with finger hairs to 0.75 mm Jaltomata long. Stalked multicellular lomana glands 75-90 gm long is an erect, nearly glabrous shrub with abundant on abaxial andpetioles adaxial faces of calyx and to 7 mm long and a white, short- tubular abaxialcorolla face of corolla. Stamens 2.8-3.9 mm havinglong; a spreading limb 11-14 mm in diameter. The type specimen of J. lomana (Stork filaments villous on proximal Y5, finger hairs unpig- et al. 9183) mented, 0.3 mm long; anthers was cream-colored, 1.25- considered to represent Saracha dentata Ruiz & Pav6n (Macbride, 1962), but the 1.55 mm long. Style 6.4-7.6 mm long, straight. latter species, now J. dentata (Ruiz & Pav6n) Ben- Stigma capitate, shallowly bilobed (obscured by ftez, is procumbent, has rotate-campanulate purple pressing), 0.54-0.72 mm wide. Nectar disk at base corollas, and does not occur in lomas formations. of ovary, orange on living plants but red/orange Paratypes. PERU. Ancash: Prov. Casma, km 350 of nectar not observed. Gynoecium glabrous. Berries PanAmerican highway, walk ca. 4 km NW across desert to and up subspherical, Lomas to 8 mm across, orange at maturity.de Mongon, 630 m, growing in the pro- tection of Seeds numerous,boulders, subreniform, alveolate, 1.6-1.8 x not on the open hillside, 20 Jan. 1998, Leiva et 1.3-1.5al. mm, 0.45-0.54 2140 mm thick. Infructescence (F, HAO), Mione et al. 631 (MOL, NY). Jaltomata up to 8-fruited. yungayensis Mione & S. Leiva, sp. nov. TYPE: Peru. Ancash: Yungay, Cordillera Blan- ca, about Habitat, phenology, 200-500 and local name. Jaltomata m into Huascar~in National Park along yungayensis has been collectedroad from 3000 to 4150 from city of Yungay to lake Chinan m on steepCocha south-facing scree slopes, in shrub- and lake Llanganuco, sunny, rocky lands,roadside and on roadsides among shrubs. Flowering with shrubs, 3525 m, 19 Jan. 1998, T and fruiting Mione, in January, October, and December, S. Leiva G. & L. Yacher 628 (holotype, and flowering in AprilNY; and May. A local nameisotypes, is CONN, MOL). Figure 3.

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"lampuxso," and fruits are not eaten (Leiva et Davidal. Spooner for helpful correspondence, Neil R. 2139). Smith for translation to Latin, and the staff of the Jaltomata yungayensis is similar to J. diversa cited herbaria. A National Geographic Society grant (Macbride) Mione: both are shrubs of high altitude, (6008-97) to T. M. funded fieldwork and supplies. and have branchlet hairs on young axes and leaves, Literature Cited and rotate to campanulate corollas. Jaltomata yun- gayensis, from Ancash, has pedunculate inflores- Leiva G., S. 1998. Las Especies del Genero Jaltomata Schlech. (Solanaceae: Solaneae) del Norte del Perl. cences, and the corolla is purple with a green Master's Thesis, Universidad Nacional Mayor de San throat. In contrast, J. diversa from the departments Marcos, Lima, Peru. of Apurfmac, Arequipa, Ayacucho, and Cuzco, Macbride, J. F. 1962. Solanaceae. Field Mus. Nat. Hist., Peru, lacks peduncles (umbels are sessile) and has Bot. Ser. 13, part V-B, No. 1. a white corolla (Mione, 1999). Mione, T 1992. The Systematics and Evolution of Jalto- mata (Solanaceae). Ph.D. Dissertation, University of Connecticut, Storrs, Connecticut. Paratypes. PERU. Ancash: Prov. Yungay, Huascarain -~. 1999. Jaltomata II: New combinations for five National Park, regione Huaraz, 13 Oct. 1976, Bernardi et South American species (Solanaceae). Brittonia 51: 31- al. 16680 (G, NY); Huascarain National Park, Llanganuco 33. Sector, near park entrance, alt. 3330-3525 m, 30 Dec. - & S. Leiva G. 1997. A new Peruvian species of 1984, Smith & Goodwin 8875 (NY), 19 Jan. 1998, Leiva Jaltomata (Solanaceae) with blood-red floral nectar. et al. 2138 (CONN, HAO); along road from Yungay to lake Rhodora 99: 283-286. Llanganuco, pueblo de Huashau, 3000 m, 19 Jan. 1998, -~ & L. A. Serazo. 1999. Jaltomata lojae (Solana- Leiva et al. 2139 (HAO), Mione et al. 629, 630 (CONN); ceae): Description and floral biology of a new Andean Prov. Huaylas, Huascarain National Park, Quebrada Santa species. Rhodora 101: 136-142. Cruz between Lago Chico and Cashapampa - , G. J. Anderson & M. Nee. 1993. Jaltomata I: exit, 3870-3350 m, 17 Jan. 1985, Smith et al. 9333 (MO); Circumscription, description and new combinations for Huascarain National Park, Auquispuquio area of ruins, five South American species. Brittonia 45: 138-145. 3800-3900 m, 7 Apr. 1986, Smith et al 11992 (MO, NY); Rundel, P. W., M. O. Dillon, B. Palma, H. A. Mooney, S. Pucapampa (Lampanin-Huaylas), 4150 m, ladera, 3 May L. Gulmon & J. R. Ehleringer. 1991. The phytogeog- 1987, Mostacero & Leiva 1897 (HUT, NY); Prov. Corongo, raphy and ecology of the coastal Atacama and Peruvian Cerro Clarinirka (arriba de Corongo), 3500-3600 m, 7 deserts. Aliso 13: 1-49. Aug. 1993, Leiva & Lezama 865 (HAO), 28 June 1998, Seithe, A. 1979. Hair types as taxonomic characters in Leiva et al. 2200 (HAO). Solanum. Pp. 307-319 in J. G. Hawkes et al. (editors), The Biology and Taxonomy of the Solanaceae. Linnean Acknowledgments. We thank Michael Nee for Society Symposium Series No. 7. Academic Press, New review, Gabriel Bernardello, Michael Dillon, and York.

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