A taxonomic revision of Lycianthes series Meizonodontae (Solanaceae)

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1 Blackwell Science, LtdOxford, UKBOJBotanical Journal of the Linnean Society The Linnean Society of London, 2004? ? Original Article LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) E. A. DEAN Botanical Journal of the Linnean Society, 2004, 145, With 16 figures A taxonomic revision of Lycianthes series Meizonodontae (Solanaceae) ELLEN A. DEAN* UC Davis Herbarium, Section of Plant Biology, One Shields Ave., University of California, Davis, CA (current address) and University Herbarium, University of California, Berkeley, CA 94720, USA Received January 2003; accepted for publication February 2004 Species descriptions and identification keys are provided for the eight species and three varieties of Lycianthes series Meizonodontae (Solanaceae), a group of perennial herbs that ranges from Mexico to Costa Rica. An image of a herbarium specimen of each species is provided. A new species, L. hintonii from Nuevo León, Mexico, is described, as are two new varieties: L. moziniana var. oaxacana, from the Sierra Juárez of Oaxaca, Mexico, and L. moziniana var. margaretiana, from Nuevo León and San Luis Potosí, Mexico. Images of the pollen, seeds and fruits of the species are included The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145, ADDITIONAL KEYWORDS: Capsicum taxonomy. * eadean@ucdavis.edu INTRODUCTION Lycianthes Hassler is the closest relative of the pepper genus Capsicum L. (Bohs & Olmstead, 1997; Walsh & Hoot, 2001). The two genera have a similar calyx morphology, in which the five lobes of the calyx are truncated into a sleeve, below which may protrude five to ten appendages (calyx teeth). However, whereas Capsicum species have anthers that dehisce by longitudinal slits, the species of Lycianthes typically have poricidal anther dehiscence. The majority of the approximately 200 species of Lycianthes are distributed in the New World from Mexico to Argentina; approximately 30 species are found in the Old World, distributed between Asia and Australia. The entire genus was last monographed in 1919 by the German botanist Georg Bitter, who divided the species into sections and series but provided no identification key (Bitter, 1919). The eight species of Lycianthes series Meizonodontae are a monophyletic group of perennial herbs from Mexico and Central America (Dean, 1995; Dean & Potter, 1998). All members possess enlarged tuberous roots, single-flowered inflorescences, relatively large green to purple fruits, and relatively large, thick, dark-coloured seeds. These characteristics stand out within the genus; most other members of the genus are woody perennials (small trees, shrubs, vines, epiphytes) with relatively small brightly coloured fruits and planar seeds. The species of series Meizonodontae are found in regions of temperate vegetation, usually in conifer or oak forest or xeric scrub community types. In most cases this is indicative of higher elevations (above 700 m); the exception to this is in south-western Mexico and Central America, where one species (Lycianthes acapulcensis) is found below 500 m in lowelevation oak forest. The soils on which the species grow are derived from a range of parent material such as relatively old limestone deposits, granite and various volcanics. Many of the species occur in either naturally occurring or anthropogenically disturbed situations, preferring eroding slopes, drainages, cultivated fields or roadsides. The species are often widespread geographically, but rare where found. However, several of the species are common in hedgerows or traditionally managed agricultural fields in Mexico (Dean, 1995). In some of these regions, the fruits of the species have been used as a food source. One of the species, Lycianthes moziniana, was routinely gathered for sale at market until about 20 years ago; it has been postulated that this 385

2 386 E. A. DEAN species was a crop of prehispanic Mexico that has reverted to a volunteer and protected plant of agricultural fields (Williams, 1993). From field interviews with farmers and other local people, it appears that some of the species are becoming rarer in agricultural fields, due to the introduction of agrochemicals, including herbicides, and the use of heavy machinery for ploughing. Grazing pressure in some regions is also having an adverse impact on some of the species (Dean, 1995). The taxonomy of series Meizonodontae as a whole has not been looked at for over 80 years, and since Georg Bitter s treatment, many new collections of the species have been made. As identification of the species has been problematic, a systematic revision of the group and identification key were needed. This paper reports the findings of a thorough investigation of all parts of the life-cycle and morphology of the species, and a new taxonomic treatment is proposed. METHODS AND RESULTS PLANT MATERIAL Specimens from the following herbaria were examined and the localities noted: ASU, BH, BM, BR, C, CAS, CR, ENCB, F, G, GH, GOET, HAL, HBG, IEB, INBIO, JE, K, MEXU, LL, M, MO, NY, P, S, TEX, UC, US, W, WIS, WU and XAL. Localities for all species except Lycianthes hintonii were visited and specimens collected in Mexico and Costa Rica during five field trips totalling 18 months between June 1990 and October Whenever possible, flowers and fruits were measured in the field, before the plants were pressed and dried. Roots and seeds collected in the field were propagated at the University of California at Berkeley between 1990 and 1995 and at the University of California at Davis between 1995 and Limited crossing experiments were performed in an insect-free greenhouse and screenhouse in order to produce fruits and seeds for study. Live plants were measured in the greenhouse for all species except L. hintonii, which is known to me only from herbarium specimens. All species were included in the following growth, pollen and seed studies except L. hintonii. SEEDLINGS The seeds of seven of the eight species germinate easily in 2 4 four weeks in Petri dishes or sand without any special treatment; germination time depends on the size of the seed. Germination is phanerocotylar in most species, although it is cryptocotylar in Lycianthes starbuckii and L. rzedowskii. Early in development, the seedlings produce a taproot. The hypocotyl root transition zone then begins to thicken into a storage organ. Although the hypocotyl root transition zone is above ground when thickening starts, it is subsequently pulled below the ground by contractile regions that appear as horizontal lines in the thickened area. Above-ground growth ceases, after the production of several leaves, and plants then die back to the root crown. Second-year growth is initiated from buds that form in one of the axils of the cotyledons; thereafter, root-borne shoots can be initiated from the root crown or other parts of the root. ROOTS The roots of seven of the eight species were observed both in the field and in the greenhouse. I have recognized two main variants in mature root form among the species: fusiform segments and moniliform segments (thickened segments separated by very narrow, string-like segments). Only one species, Lycianthes acapulcensis, has the latter type. Both arrangements promote root fragmentation when the roots are disturbed. LIFE CYCLE The above-ground growth of all species arises from the roots during the rainy season (May December); and the plants die back to their roots during the dry season. From herbarium label data, field observation and greenhouse studies, I estimated the timing of growth and flowering for the species. Very few herbarium specimens with fruits were available, and so fruiting times were estimated based on field observations and the timing of fruit development in the greenhouse. Under greenhouse conditions, most first-year plants do not flower. With few exceptions, flowers appear on second-year plants and thereafter. Under natural conditions, flowering begins as early as June in Lycianthes dejecta and L. peduncularis, with most species flowering between July and September. Fruit development takes 3 4 months in most species, the exception to this being L. moziniana, with a mean fruit development time of 2 months. The non-human seed dispersal agents of these species are not known. ABOVE-GROUND ARCHITECTURE The terminology used to describe the architecture of the above-ground plant body follows that of Child & Lester (1991; Fig. 1). All branching is sympodial, with the axis terminated by a flower or sterile bud after one to many nodes. The sympodial branching point can be dichasial or monochasial. The first sympodial unit (initiated from the root) bears many nodes and has spirally arranged leaves. Thereafter, all branches are short sympodial units bearing one to two leaves.

3 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 387 Lycianthes dejecta. These first scale-like leaves are generally orbicular to spathulate and sessile, whereas the first leaves produced in L. dejecta are longerpetiolate with a reniform blade. Subsequent leaves produced on the first sympodium and subsequent sympodia are expanded, often each leaf larger than the subsequent one, reaching their largest size either in the middle or upper parts of the first sympodium or in the second to fourth sympodia. Figure 1. Branching, leaf arrangement and flower location pattern in Lycianthes series Meizonodontae. Figure shows: A first sympodial unit with multiple, spirally arranged leaves ending in a mature flower and dichasial branching point; B second sympodial units with the one on the right ending in a nearly mature flower and dichasial branching point; C third bifoliate sympodial units, each ending in a bud and monochasial branching point (with leaves geminate). If the branching is monochasial, there are two geminate leaves placed below the flower or sterile bud; if the branching is dichasial, there is one leaf. If the plant becomes large enough for branching to be initiated from the axils of lower leaves, the entire sequence is repeated again. In the species descriptions that follow, a detailed description is given for the first sympodial unit, which is usually erect. A size range is given for the first sympodial branching point which terminates the first sympodial unit. Then, a short description is given for subsequent sympodial growth. LEAVES Several different types of leaves are produced by mature plants. Leaves produced on underground stems are reduced and colourless. The first leaves produced by the above-ground section of the first sympodial unit are reduced and scale-like, except in TRICHOME TYPES There are two classes of trichomes in series Meizonodontae: macroscopic cover trichomes and microscopic multicellular glandular trichomes. These two classes of trichomes are present in other Solanaceae (Seithe & Sullivan, 1990; Bohs, 1994). The cover trichomes can be seen with the naked eye and are easily observed with a dissecting microscope; these are the trichomes that are characterized in the species descriptions that follow using the terminology of Roe (1971). The microscopic glandular trichomes occur throughout the plant body of all species, but are most numerous in the inflorescence, especially on the inside of the calyx; these trichomes are uniform in all of the species and are not discussed in the species descriptions. INFLORESCENCE STRUCTURE AND PEDUNCLE DEVELOPMENT Observations of inflorescence development were made in the greenhouse and measurements were made over time on the same flower. All species of series Meizonodontae have one-flowered inflorescences, a rarity in the genus. The peduncle begins as an erect stalk bearing a small bud; it then elongates rapidly, becoming recurved, so that the bud faces the ground. In the final stage before anthesis, the peduncle straightens, remaining recurved only at the tip, with the flower orientated plagiotropically (sideways). Peduncle elongation can be quick and dramatic in species with elongate peduncles such as Lycianthes peduncularis and L. starbuckii. In these species, the peduncle may elongate five- to ten-fold in 2 or 3 days to a final length of up to 11.5 cm. Peduncle elongation continues at a slower pace after anthesis and during fruiting. CALYX The calyx of Lycianthes has been described in detail elsewhere (D Arcy, 1986). The species of series Meizonodontae have a calyx that is broadly cupulate, conic or urceolate with ten ribs and ten well-developed calyx teeth (Fig. 2C, D). Calyx measurements were made over the lifetime of individual flowers in the

4 388 E. A. DEAN A B C D Figure 2. Corolla, stamen and calyx morphology in Lycianthes series Meizonodontae. A. Adaxial side of flower of L. ciliolata (Dean & Starbuck 295) showing corolla lobes and membrane; B. Sideview of flower of L. ciliolata (Dean & Starbuck 213) showing unequal stamens; C. Abaxial side of flower of Dean & Starbuck 212 (placed in L. rzedowskii) showing calyx with spreading teeth; D. Abaxial side of flower of L. starbuckii (Dean & Starbuck 315) showing calyx with lax teeth.

5 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 389 greenhouse and were found to remain constant. Calyx size at the time of anthesis can be useful in distinguishing the species, that of L. moziniana being the largest and that of L. peduncularis the smallest. The degree to which the teeth are reflexed at anthesis is also useful in species recognition. Most species within the series have teeth that are widely spreading or reflexed backward at anthesis (Fig. 2C). Only in L. moziniana and L. starbuckii are the teeth either just slightly spreading or lax and somewhat undulate, lying against the corolla (Fig. 2D). During fruit development, the calyx and calyx teeth undergo size and positional changes, with the calyx enlarging and the teeth usually becoming more reflexed or even recurved. In L. moziniana var. moziniana the teeth can remain lax, lying against the fruit, or become stiff and straight; in L. starbuckii (and rarely in L. acapulcensis) the teeth become stiff, but remain near the fruit surface. Unfortunately, these helpful calyx characteristics can be difficult to see in pressed and dried specimens. COROLLA All species of the series have rotate, sympetalous corollas. The constricted base of the corolla tube is very short and well included within the calyx. The limb is divided into two main parts: the five thickened corolla lobes, which are ovate, obovate or oblanceolate in outline, and the corolla membrane, which is composed of thinner tissue that connects the corolla lobes to nearly their tips (Fig. 2A). Flowers were observed and measured over the several days that they opened in the greenhouse. The corollas of all species grown exhibit diurnal floral movements, opening early in the morning and closing 2 6 h later. Flowers may open for two to seven successive days, depending on the species, genet and weather conditions. Over the several days during which a flower opens and closes, its corolla grows in size. This change can be very dramatic in flowers that open for five or more days in a row (Dean, 2001). ANDROECIUM The five stamens are adnate at their bases to the corolla tube and are unequal in length (Fig. 2B). The lowest stamen is generally the longest, followed by the two lateral stamens and then the two upper stamens. As anther size is usually more or less equal among the stamens (the anther of the lowest stamen is sometimes longer than the other four), the stamen size differences are mostly due to differences in filament length. Filaments vary in their cross-sectional appearance, ranging from nearly round (Lycianthes dejecta and L. peduncularis) to laterally compressed (all other species). In most species the filaments hold the anthers close to the style, whereas in L. dejecta and L. peduncularis the filaments are curved downward. As was done with the corolla, the androecium was observed and measured over the several days that an individual flower was open. It was found that over the days that the corolla opens, the filaments elongate, so that it is difficult to use absolute filament lengths for taxonomic purposes (Dean, 2001). However, the ratio of the lower to lateral filament length can be helpful for identifying species. The shape of the lowest anther and the shape and orientation of the pores of that anther are useful in distinguishing some of the species of the series. The lowest anthers of most species range from lanceolate to widely elliptic, and the lateral to terminal pores are round to ovate, dehiscing toward the style or stigma. However, the lowest anthers of Lycianthes acapulcensis and many populations of L. ciliolata are oblong to lanceolate with lateral pores that are vertical slits that dehisce toward one another rather than toward the style. In addition, in some coastal and Central American populations of L. acapulcensis the pores of the lateral and sometimes the upper anthers are located on the abaxial side, so that the pores dehisce away from the style (extrorse dehiscence). Most of the species exhibit asynchronous anther dehiscence with the lateral, or lateral and upper, anthers dehiscing first and the lower anther dehiscing last. Only in L. peduncularis is dehiscence synchronous, with all anthers dehiscing on the first day of anthesis (Dean, 2001). POLLEN I examined the pollen using pollen acetolysis (Erdtman, 1969) and light microscopy. I examined grains taken from two to seven accessions per species, generally measuring 15 grains per accession. Pollen grain images were examined using a Zeiss microscope attached to a Panasonic PANA video camera and a video imagery program (IMAGE), which allowed measurement of the polar and equatorial axes of individual pollen grains that I examined. Photographs were taken using a Zeiss microscope with a Nikon camera attachment. Three types of pollen grains occur within the species of the series: tricolporate, with nearly equal pore to pore distance in polar view (the most common type found in the Solanaceae) is found in all but three species (Fig. 3A); dicolporate (two pores) is found in Lycianthes acapulcensis and L. rzedowskii (Fig. 3C); and dicolporate with a much reduced remnant of a third pore is found in L. ciliolata (Fig. 3B). Lycianthes starbuckii has very irregular pollen, sometimes appearing tricolporate, sometimes with the third pore

6 390 E. A. DEAN reduced to a remnant, often asymmetrical in polar view, but with none of the sides equal. The pollen grains of Lycianthes dejecta and L. peduncularis are the smallest of the series and similar to one another in their dimensions. Although the pollen grains of L. moziniana are tricolporate, they are slightly larger, on average, than those of L. dejecta and L. peduncularis. POLLINATORS Bees were observed and collected from seven populations representing four species of series Meizonodontae, and they were identified as solitary bees in the Anthophoridae, Colletidae and Halictidae (Dean, 2001); I had observed these insect visitors travelling between flowers within a population, buzzing the anthers of each flower visited. In two of the species (Lycianthes peduncularis and L. ciliolata), bees belonging to two different genera (and families) were observed visiting a single population. From these data, one can infer that the relationship between pollinator and plant species may not be a tight one. This type of loose relationship was found in the only other pollination study that has been conducted with Lycianthes (de Nevers, 1986). In that study, the pollinators were observed visiting three different species of plants, only one of which was a Lycianthes. FRUIT Fruit type is very helpful in distinguishing some of the species from one another and is emphasized in the identification key. Before this study, the fruits of most of the species were unknown. Now, only that of Lycianthes hintonii remains uncharacterized. In the species descriptions that follow, the fruits are described in terms of their exocarp, mesocarp and placental area. A B C Figure 3. Pollen types in Lycianthes series Meizonodontae. A. Tricolporate grain from L. moziniana (Dean & Starbuck 270); B. Dicolporate grain with remnant third pore from L. ciliolata (Dean & Starbuck 295); C. Dicolporate grain from L. acapulcensis (Dean & Starbuck 332). Scale bars = 9 mm. Three of the species have greenish exocarps and a mostly smooth placental area (L. moziniana, L. dejecta and L. peduncularis), whereas the other four that are known (L. acapulcensis, L. ciliolata, L. rzedowskii and L. starbuckii) all have dark blue to purple exocarps with a powdery texture to the placental area (Fig. 4). The occurrence of sclerotic granules (enlarged stone cells) in the mesocarp of L. peduncularis is an anomaly in the series, and these structures are uncommon in Lycianthes as a whole (Fig. 4C). The variation in fruit shape and exocarp coloration is rather remarkable in L. acapulcensis and is discussed under that species treatment (see Fig. 7). SEEDS The seeds of series Meizonodontae are unusual in Lycianthes for their dark brown to black colour and thickness. The typical seeds of Lycianthes are tan to light brown and very flattened. I have seen the seeds of all of the species of the series except L. hintonii. The outline of the seeds of the species of the series is usually compressed obovate or reniform, but can be angular trullate or depressed rhombic (outline descriptors are taken from Radford et al. (1974: 131). The seeds range from somewhat laterally compressed to subglobose. In addition, many of the species have seeds that possess a ridge that protrudes along one side. The seed epidermis of all species but Lycianthes hintonii was examined using scanning electron microscopy. Seeds were taken from field collections and herbarium specimens, sonicated with water to remove residual pulp, then treated with cellulase as described in Lester & Durrands (1984) for h, depending on seed size. This treatment removes the anticlinal walls of the epidermal cells, exposing the structure of the periclinal walls. Treated seeds were dried in a vacuum chamber for 48 h, mounted on stubs with silver

7 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 391 paste and sputter coated with gold (30 50 nm). An ISI-130 scanning electron microscope set to 10 kv was then used to examine the seeds. In all seven species, I was able to see the pattern of the anticlinal walls of the epidermal cells (Fig. 5). The number of cell wall convolutions per cell is very similar from species to species, but in Lycianthes ciliolata, L. acapulcensis, L. rzedowskii and L. starbuckii, the anticlinal cell walls are well separated from one another, so that the lumen is much larger and more easily seen than in the other three species. In Lycianthes peduncularis and some populations of L. moziniana, I discovered the presence of surface A F Figure 4. Fruit types in Lycianthes series Meizonodontae (note: grid squares are 2.0 cm on a side): A. L. moziniana (Dean & Starbuck 328) whole fruit; B. L. moziniana (Dean & Starbuck 326) long section; C. L. peduncularis (Dean & Solano 303) seeds and sclerotic granules; D. L. peduncularis (Dean & Starbuck 298) whole fruits; E. L. dejecta (Dean & Starbuck 304) exocarp (left) and long-section showing inside of fruit (right); F. L. ciliolata (Dean & Starbuck 295) whole fruit; G. L. acapulcensis (Dean & Starbuck 280) long section; H. L. starbuckii (Dean et al. 315) whole fruit. D B G papillae and in L. dejecta and L. peduncularis, I found fibrils protruding from the anticlinal walls. The exact nature and function of the fibrils are not known, but their presence or absence is constant within a species. The fibrils appear to be protruding from the region between adjacent cells. In L. moziniana, that region is occupied by a smooth membrane-like structure, rather than fibrils. Although I did not study seed development, I did examine one immature seed of L. rzedowskii. That seed possessed fibrils between adjacent cells, but those were much longer than those of L. dejecta and L. peduncularis. It may be that fibrils are present C E H

8 392 E. A. DEAN 315 µm 24.8 µm A B 313 µm 22.0 µm 437 µm during seed development in some species but become poorly defined in the mature seeds. C E G Figure 5. Seeds in Lycianthes series Meizonodontae: A. L. moziniana (Dean & Starbuck 256) whole seed; B. Same, closeup showing smooth membrane between cells; C. L. dejecta (Dean & Starbuck 276) whole seed; D. Same, close-up showing seed fibrils; E. L. peduncularis (Dean & Starbuck 308) whole seed; F. Same, close-up showing surface papillae and seed fibrils; G. L. acapulcensis (Dean & Starbuck 310) whole seed; H. Same, close-up µm 345 µm 23.1 µm 2n = 24 (voucher information given at the end of each species description). This chromosome number is very common in the subfamily Solanoideae. D F H CHROMOSOME NUMBERS I examined the chromosomes of all of the species except Lycianthes hintonii using a standard method for acetocarmine staining of root tip preparation (Tanaka & Kamemoto, 1984). All seven species have TAXONOMY SPECIES CONCEPT The species circumscriptions in this taxonomic treatment are based on a combination of phylogenetic and

9 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 393 morphological species concepts, and all species except Lycianthes hintonii were grown in a common garden to ensure that morphological and micromorphological characteristics were not caused by environmental differences. In separating L. dejecta, L. peduncularis and L. moziniana, I emphasized fruit and seed characters. However, in the other five species, character evolution followed a different path, and characters such as pollen type, flower colour, androecium type and habitat preference are more important. Limited crossing experiments were also performed among the species, which indicated that many of these species are good biological species; however, several species can hybridize with one another, and this is discussed under each species treatment. With the exception of L. hintonii, which is known from only a few collections and which I have never seen in the field, all of the species that I have circumscribed are monophyletic in phylogenetic analyses based on morphological characters (Dean, 1995). In most cases, my species circumscriptions are broad, and only well-defined entities were described. The exception to this rule is the semidomesticated species Lycianthes moziniana, which due to its long history with humans in the agricultural environment has a more complicated taxonomy. I have broken this species into three varieties found in Mexico: a northern variety that grows in relatively undisturbed oak forest and other habitats on limestone soils of the Sierra Madre Oriental from San Luis Potosí north; a southern variety found in agricultural fields and old fields of the Sierra de Juárez of Oaxaca; and the typical variety that grows in agricultural fields and disturbed old fields on volcanic soils of the transvolcanic belt. Sequence data from the ITS region indicate that the Oaxacan and northern varieties are very closely related, whereas the widespread typical variety, which has been gathered, perhaps cultivated, by farmers has undergone extensive molecular evolution and perhaps hybridization with other species of the series (my unpubl. data). When crossed in the greenhouse, the typical variety and the Oaxacan variety do have limited crossability; these data have not been obtained for the northern variety. Because these varieties are so close morphologically, they are kept together as one species in this treatment. GENERIC SYNONYMY Lycianthes (Dunal) Hassler, Ann. Conserv. Jard. Bot. Genève 20: Name conserved over Otilix Raf. (1830) and Parascapolia Baill. (1888) (McVaugh, 1973). Solanum subsection Lycianthes Dunal, in DeCandolle, Prodr. 13(1): Solanum section Lycianthes (Dunal) Wettst., in Engler and Prantl (Eds.), Die Natürl. Pflanzenf. 4(3b): Solanum subgenus Lycianthes (Dunal) Bitter, Bot. Jahrb. Syst. 54: TYPE: Based originally on Solanum subsection Lycianthes Dunal. LECTOTYPE: Solanum lycioides L. (D Arcy, 1973: 631). Otilix Raf., Medical Fl. 2: TYPE: Solanum lycioides L. Parascopolia Baill., Hist. Pl. 9: TYPE: P. acapulcensis Baill. SPECIES TREATMENT Lycianthes acapulcensis (Baill.) D Arcy (Fig. 6) Parascapolia acapulcensis Baill., Hist. Pl. 9: Lycianthes acapulcensis (Baill.) D Arcy, Solanaceae Newsl. 2(4): TYPE: MEXICO. State of Guerrero, Acapulco, Punto Griffon, 1888, M. Thiebaud 1002 (HOLOTYPE: P!; photos NY!, WIS!). Lycianthes grandifrons Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: COSTA RICA. Province San José, Llanos de Turrucares, 600 m, 18.ix.1888, H. Pittier & T. Durand 478 (HOLOTYPE: Figure 6. Herbarium specimen of Lycianthes acapulcensis (Dean & Starbuck 280 (DAV)).

10 394 E. A. DEAN KEY TO SPECIES OF SERIES MEIZONODONTAE 1a. Indument of entire plant of multi-angulate-dendritic trichomes (all rays originating from one point) mixed with dendritic and simple trichomes, the individual multiangulate-dendritic trichomes usually less than 0.5 mm long; species of arid habitats...l. dejecta 1b. Indument of simple or dendritic trichomes, not multi-angulate-dendritic, if dendritic trichomes present, these often more than 0.5 mm long; species of both arid and non-arid habitats...2 2a. Plant prostrate (rarely decumbent or ascending); fruits with yellowish sclereids in the mesocarp; styles often strongly curved downward; ovaries short, rounded to ovoid, less than 2 mm long; indument of simple, antrorsely appressed trichomes... L. peduncularis 2b. Plants prostrate or not; fruits without sclereids; styles straight to slightly curved, never strongly curved downward; ovaries conic, usually longer than 2 mm (sometimes shorter in L. starbuckii); indument of simple or dendritic trichomes, these spreading or retrorsely appressed (rarely, antrorsely appressed in Chiapan or Guatemalan populations of L. ciliolata)...3 3a. Corolla white, with or without maroon to purple nectar guides; corolla lobes usually glabrous abaxially; plant body usually erect, with first sympodial unit well developed above the ground; first two branching points on the plant body usually dichasial...4 4a. Filament of each lower stamen usually more than twice as long as those of the lateral stamens; pores of the anther of the lower stamen dehiscing laterally, usually slit-like; stigmas generally deeply bilobed; widespread species of the transvolcanic belt, southern Mexico and Central America... L. acapulcensis 4b. Filament of each lower stamen usually less than twice as long as those of the lateral stamens; pores of the anther of the lower stamen oval, nearly terminal, not lateral and slit-like; stigmas not deeply bilobed...5 5a. States of Morelos, Michoacan and Mexico, on volcanic soils; first sympodial unit usually very well developed (to 90 cm long) with numerous internodes (usually 10 21) and leaves; lateral branching from the nodes of the first sympodial unit usually not present at the time of flowering; subsequent sympodial growth poorly developed; pollen dicolporate...l. rzedowskii 5b. State of Nuevo León, limestone soils; first sympodial unit c. 25 cm long, the internodes c. 13 in number; lateral branching from the nodes of the first sympodial unit usually present at the time of flowering; subsequent sympodial growth about equal to the length of the first sympodial unit; pollen tricolporate... L. hintonii 3b. Corolla lilac, violet or light purple, with maroon to purple nectar guides; corolla lobes glabrous or pubescent abaxially; plant body erect, decumbent or prostrate with first sympodial unit sometimes not well developed above ground; second sympodial branching point sometimes monochasial...6 6a. Fruit green to tan at maturity, sometimes with purple blotches; seeds less than 3 mm long, smooth and shiny to the naked eye; second sympodial branching point usually monochasial...l. moziniana 6b. Fruit purple at maturity; seeds greater than 3 mm long, rough textured and dull to the naked eye; second sympodial branching point usually dichasial...7 7a. Calyx teeth at anthesis lax or slightly spreading, not widely spreading or reflexed; corolla lobes usually noticeably pubescent abaxially; first sympodial unit poorly developed above ground, the plant body often prostrate; rare species of the Sierra de Nanchititla (State of Mexico)...L. starbuckii 7b. Calyx teeth at anthesis widely spreading to reflexed; corolla lobes generally glabrous abaxially (rarely slightly pubescent in Oaxaca); first sympodial unit well-developed above ground, the plant body usually erect; widespread in SE Mexico, Guatemala...L. ciliolata BR!; ISOTYPES: CR! (mixed collection with Witheringia solanacea L Hér.), US!). Lycianthes guatemalensis Bitter, Abh. Naturwiss. Vereine Bremen 24(2): Type: GUATE- MALA. Department Retalhuleu, Retalulëu, v.1877, Bernoulli & Cario 2384 (LECTOTYPE here designated: GOET!). Chosen from syntypes: Bernoulli & Cario 2384, Bernoulli & Cario 2404 (B, destroyed; photos G!, GH!, NY!).

11 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 395 Lycianthes villosula Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: COSTA RICA. Province Alajuela, El Brazil, gorges of Virilla River, 800 m, 14.vii.1911, H. Pittier 3676 (HOLOTYPE: US!). Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. cladotricha Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO. State of Morelos, Cuernavaca, in moist copses, 5000 ft, vi vii.1896, C. Pringle 6399 (LECTOTYPE here designated: MEXU!; ISOLECTOTYPES: BM!, BR!, CAS!, G!, GH!, GOET!, HBG!, JE!, K!, M!, MEXU!, MO!, NY!, S!, UC!, W!, WU!, Z!). Chosen from syntypes: Pringle 6399, Pringle (BH!, C!, CAS!, GH!, K!, L!, MO!, S!), Schmitz 1091 (W!). Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. ramosipila Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO. State unknown, plaines de terre froide, JDP (Jardin des Plantes) 82 (HOLOTYPE: P!). Perennial HERBS, the habit variable, decumbent to erect, often recumbent with age (4) (100) cm tall. INDUMENT of simple trichomes in coastal Mexico and Central America, of simple and dendritic trichomes in more inland Mexican populations. ROOTS moniliform, usually present within 20 cm of the soil surface. STEMS arising from the root one to several; below-ground growth not extensive, white, aboveground growth green or greenish-purple, mostly herbaceous, terete in cross-section, becoming somewhat woody with age, much compressed when dried, the new growth often villous to tomentose, sometimes just pubescent, the older growth pubescent to tomentose, rarely nearly glabrous, the trichomes (2) mm long, spreading to appressed-retrorse. FIRST SYMPO- DIAL UNIT decumbent to erect above-ground (1.5) 5 30 (70) cm long (1.3) 2 5 (13) mm diameter at widest point; internodes 2 10 (14) in number; lateral branching well developed or not. FIRST TWO SYMPODIAL BRANCHING POINTS usually dichasial (the first with a branching angle of (30) (195) ), followed by monochasial branching, sometimes lateral branching from monochasial or diachasial branching points creating false dichasia or trichasia. SUBSEQUENT SYMPO- DIAL GROWTH quite variable in extent of development, ascending to decumbent, often exceeding the first sympodium in length (in some Costa Rica and Nicaragua populations spreading widely on the ground, sometimes rooting at the nodes), extending laterally a total of (2) 5 30 (50) cm, the second sympodial units cm long, mm diameter, these generally followed by one to several longer sympodia, with subsequent sympodia becoming more reduced distally. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT chartaceous (thick-chartaceous in some coastal populations) (2) 4 10 (14) in number; leaves closest to the soil surface reduced, scale-like, obovate to spathulate, sessile, cm long, 2 10 mm wide, early deciduous; subsequent leaves usually expanded (rarely only scale leaves present), often deciduous, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for (1/8) 1/4 1/2 of total leaf length, the lamina obovate, elliptic or ovate, the margin smooth to undulate, the tip rounded to short-acuminate, the base usually obtuse to cuneate (rarely truncate), short-attenuate (rarely long-attenuate) into the petiole (or sessile), the largest expanded leaves (usually at the distal end) 3 15 (18) cm long, 1 5 (8) cm wide, the petiole (if present) (2.5) cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate, similar in shape and texture to the distal-most leaf of first sympodial unit, but the lamina usually ovate to elliptic, rarely obovate, the tip acute to shortacuminate (rarely long-acuminate), and the base short-attenuate into the petiole, or sessile; largest leaves (15.5) cm long, 2 6 (8) cm wide, with petiole to cm long, the leaves becoming reduced and more sessile on more distal sympodial units; geminate leaf pairs unequal in size (the smaller 1/4 3/4 of the larger), the smaller of the pair with shorter and more oblique leaf base. LEAF PUBESCENCE similar throughout, the leaves nearly glabrous to villous or tomentose, the trichomes (1.5) mm long, erect to appressed on both surfaces, the trichomes of the main veins of the abaxial sides usually spreading, rarely somewhat antrorsely appressed, the trichomes of the margin mm long, spreading to appressed-antrorse. PEDICELS AT ANTHESIS green or greenish-purple (1) 3 6 (8.5) mm long, usually (0.1) times as long as subtending leaf (rarely to 2 times if subtending leaf is the distalmost leaf of the first sympodial unit and it is scale-like), glabrous (in some coastal populations) to villous, the trichomes (0.1) (1.5) mm long, spreading (in most inland Mexican populations) to appressed-retrorse (in most coastal and Central American populations). CALYCES AT ANTHESIS green or greenish purple, obconic to campanulate to urceolate, (6.25) mm long, mm diameter when fresh (often longer than wide) (2) (6) when pressed and dried, the teeth usually reflexed (sometimes just widely spreading in coastal populations), emerging from the ribs at approximately the same level or at two distinct levels (these levels about mm apart), of approximately equal length or of two alternating lengths (rarely every other tooth nearly suppressed in some coastal populations) (0) (9) mm long, the ribs and teeth glabrous, pubescent, villous, or tomentose, the trichomes (2) mm long, spreading to slightly retrorse. COROLLAS orientated laterally, not nodding, the

12 396 E. A. DEAN tube mm long, the limb 2 5 cm diameter, the membrane white, the adaxial sides of the lobes of similar colour but with darker violet or maroon veins (in inland Mexico) or greenish coloration at the base (in coastal and Central American populations), the abaxial sides of the lobes green, usually glabrous (very rarely with a few trichomes along the midvein in Oaxaca). ANDROECIUM with green, erect, glabrous to pubescent, laterally compressed filaments, the lowest 4 9 mm long, the two lateral (4.5) mm, the two upper (3.5) mm, the length of the lowest nearly always 2 4 times that of the lateral (rarely times in some atypical populations in the state of Mexico); anthers exuding a citrus scent, mm long, the lowest anther usually lanceolate to oblong (rarely ovate), the pores slit-like, lateral, generally dehiscing toward one another, the pores of the lateral and upper anthers dehiscing toward the style in most Mexican populations but dehiscing away from the style in coastal and Central American populations (in some of these forms, the two pores of these anthers are at different levels, one higher, the other lower); pollen grains dicolporate. GYNOECIUM with conic ovary (sometimes attenuate into style) mm long, the ovules (34) (94); style straight to slightly curved, 9 14 mm long; stigma green, usually strongly bilobed (rarely weakly bilobed or rounded in some atypical populations in the states of México and Morelos). FRUITS remaining attached to calyces at maturity, variable in shape, short-ovoid to elongate-fusiform (10) (70) mm long (4.5) 9 20 mm diameter at widest point, the tip apiculate to long-attenuate; exocarp purple to bluish, ranging from glossy blue-black, to grey-blue, to bright blue, or dull purple, glabrous; mesocarp variable in texture, dark purple and juicy to light purple and powdery, lacking sclerotic granules; placental area light purple, powdery; fruiting calyx enlarged (1.5) 2 4 (6) mm long, (14) mm diameter, the teeth usually reflexed, sometimes appressed to fruit, often broken, 0 10 mm long; fruiting pedicels 2 7 (9) cm long, deflexed, the fruit pendant; seeds (11) (90) per fruit, black, irregularly depressedobovate to depressed rhombic, rough in texture, ridged along one side, usually with blisters along the ridge, mm long, mm wide. Chromosome number: 2n = 24 (Dean & Starbuck 313, 314, 329). Indigenous names: Maravilla, huevo de cuervo, chimpin. Distribution: Central America (Guatemala, El Salvador, Nicaragua, and Costa Rica) and Mexico (states of Chiapas, Colima, Guerrero, Jalisco, México, Michoacán, Morelos, Oaxaca and Puebla). In inland Mexico and Central America, this species generally grows on volcanic soils (rarely on limestone near Chilpancingo, Guerrero); coastal populations near Puerto Vallarta, Mexico, grow on granitic soils, and this may be true of other coastal populations. Vegetation types range from oak pine woodland to more tropical vegetation types. Habitats include drainages, clearings, paths and agricultural fields, m. Diagnostic features: Lycianthes acapulcensis is the most variable species of the series, and it may be that some of the local forms deserve varietal status. It is variable in habit, pubescence (both trichome type and degree of pubescence), leaf shape, presence or absence of nectar guides, fruit shape (Fig. 7) and fruit coloration. However, my attempts to use these features to separate the species into infraspecific taxa have not been successful, because the variation extremes are connected by a series of intermediate populations. Therefore, I have elected to treat all of the populations as one variable species. Lycianthes acapulcensis may be confused with L. ciliolata and L. rzedowskii. It is separated from those species by both vegetative and floral characteristics. It is best recognized by its combination of white corollas that may or may not have maroon to purple nectar guides and its pattern of filament lengths (the lowermost filament nearly always more than twice as long as the lateral filaments). The anthers have a lemony odour, which is unlike that of any other anther (pollen) odour in the series. The root shape (moniliform rather than fusiform segments) is helpful if underground parts are available for examination. On dried specimens, the length of the pedicels of the youngest mature flowers relative to their subtending leaves is often a useful character for separating L. acapulcensis from L. ciliolata. In the former, the length of those pedicels is usually less than that of the subtending leaves, whereas in L. ciliolata the length of the pedicels generally exceeds that of the leaves. Lycianthes acapulcensis appears to hybridize with L. moziniana where the two species co-occur. In the regions of the Sierra del Tigre, the Sierra del Halo and the Nevado de Colima, in the state of Jalisco, rather atypical forms of L. moziniana can be found with dendritic trichomes, short peduncles and, at times, white corollas. In particular the collections of McVaugh from the Sierra del Halo (McVaugh 16154, near a lumber road leaving the Colima hwy 7 miles SSW of Tecalitlán) indicate that intermediates between the two species occur. One of my greenhouse accessions from that region (Dean & Starbuck 328) has lower than normal pollen stainability (56% of grains stained). In the state of México, Lycianthes acapulcensis and L. rzedowskii may hybridize. Intermediates between the two species have been collected in the area sur-

13 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 397 A B C Figure 7. Fruit types in Lycianthes acapulcensis (note: grid squares are 2.0 cm on a side): A. Dean & Starbuck 310 from state of México; B. Dean & Starbuck 314 from state of México; C. Dean & Starbuck 332 from state of Jalisco. rounding the towns of Temascaltepec and Ixtapan de la Sal. Both species are known to occur in the moist valleys near Temascaltepec. I have collected only L. acapulcensis near Ixtapan de la Sal; some of my greenhouse accessions from that region have sterile pollen. Specimens examined: COSTA RICA: PROVINCE PUNTARENAS: Monteverde, 1 2 km downstream from the village of San Luis along the Río Guacimal, N, W, m, 30.v.1992, Haber & Joyce (INBIO). SAN JOSÉ: El Brazil, rd between El Brazil and the electric substation of the River Virilla, along tributary of the Virilla River, 9 56 N, W, 800 m, 29.ix.1993, Dean et al. 361 (CR, DAV, INBIO, NY); along rd between Guacima and Ciruelas, at the crossing of the Ciruelas River, near railroad station, 9 59 N, W, 700 m, 29.ix.1993, Dean et al. 362 (CR, DAV, INBIO, UC); Villa Colon, Finca San Luis, 800 m, 26.v.1965, Jiménez-M (CR, F, MO, NY). EL SALVADOR: DEPARTMENT UNKNOWN: Finca San Nicolás, v.1923, Calderón 1643 (US). SON- SONATE: vicinity of Izalco, iii.1922, Standley (GH, US). GUATEMALA: DEPARTMENT UNKNOWN: S. Sebastian, ix.1874, Bernoulli & Cario 2404 (B destroyed, photos NY, GH, G). MEXICO: STATE OF CHIAPAS: Mpio. Cacahoatán, Monte Bello, 450 m, 19.iv.1985, Ventura & López 1541(ENCB, G, UC). COLIMA: Mpio. Comala, Rancho El Jabalí, c. 22 km NNW of city of Colima, near Jalisco state line, S and E sides of Lago Calabozo (near N, W), 1475 m, 29.viii.1988, Sanders et al (NY); Mpio. Comala, Rancho El Jabalí, 22 air km NNW of Colima in the SW foothills of the Volcán de Colima, on the Colima/Jalisco border, N, W, 1350 m, 12.vii.1991, Vázquez- V. & Phillips 837 (NY); Mpio. Comala, Rancho El Jabalí, 22 air km NNW of Colima in the SW foothills of the Volcán de Colima, on the Colima/Jalisco border, along road from headquarters to Lago Epazote, N, W, 1300 m, 14.vii.1991, Vázquez- V. & Phillips 876 (CAS, NY); Mpio. Comala, Rancho El Jabalí, 22 air km NNW of Colima in the SW foothills of the Volcán de Colima, on the Colima/Jalisco border, N, W, 1300 m, 15.vii.1991, Vázquez- V. & Phillips 887 (NY, UC). GUERRERO: S of Chilpancingo, 24.vi.1935, Clark 7196 (MO); N of Chilpancingo, 24.vi.1935, Clark 7196 (NY); 5 mi NE of Ocotito, 2900 ft, 13.vi.1954, Crisman & Willis 194 (TEX, UC); Mpio. Taxco, Just west of main part of town of Landa, along road to Ixcateopan, 1890 m, 12.vii.1990, Dean & Starbuck 209 (DAV, MEXU); Mpio. Tetipac?. 2.4 rd mi from the town of Tepitac along the Tepitac-Taxco rd, 4.x.1991, Dean & Starbuck 278c (XAL); Mpio. Mochitlan, along hwy 95 between Chilpancingo and Tierra Colorada, c. 5.0 rd mi N of El Ocotito, W side of rd, 3000 m, 7.x.1991, Dean & Starbuck 279 (DAV, MEXU, XAL); Mpio. Chochihualco, along rd between hwy 95 and Filo de Caballo, N of Chilpancingo, c. 9 rd mi W of Xochipala, N side of rd,

14 398 E. A. DEAN 6500 m, 7.x.1991, Dean & Starbuck 280 (DAV, MEXU, NY, XAL); Mpio. Chilpancingo, Botanical Garden of the Universidad Autónoma de México at Chilpancingo, 4400 m, 8.x.1991, Dean & Starbuck 281 (DAV, ENCB, MEXU, MO, NY, UC, XAL); Mpio. Chilpancingo, Jardín Botánico of the Universidad Autónoma de México at Chilpancingo, 4400 m, 5.xi.1991, Dean & Starbuck 312 (DAV, MEXU, NY, UC, XAL); Mpio. Taxco, Town of Landa, just W of main part of town, along rd to Ixcateopan, 1890 m, 6.xi.1991, Dean & Starbuck 313 (DAV, ENCB, MEXU, NY, UC, XAL); Dist. Montes de Oca, Capiral, 780 m, 13.vi.1937, Hinton et al (G, GH, K, LL, NY, S, UC); Dist. Mina, Manchón, 6.vii.1937, Hinton et al (BM, DS, GH, K, LL, MEXU, NY, UC); Mpio. Zirandaro, en La Saiba Amarilla a 7 km al E de Guayameo, camino Guayameo-Los Placeres del Oro, 1150 m, 14.vii.1982, Martínez-S. & Soto-N (MEXU, NY); Limón Mt., 4000 ft, 2.vii.1910, Rusby s.n. (NY); Landa, 5 km al SW de Taxco, camino a Ixcateopan, 1790 m, 7.vii.1982, Soto-N. & Martínez 4000 (CAS, ENCB, MEXU, WIS); road from Milpillas to Atoyac de Alvarez, 9 mi WSW of Xochipala, 5400 ft, 30.vi.1982, Thomas et al (NY); Chilpancingo, Jardín Botánico de la Universidad Autónoma de Guerrero, 1330 m, 29.vi.1978, Toledo-C. 386 (XAL); Mpio. Acapulco, La Venta, 10.vii.1966, Villanueva-O. 10 (ENCB, F). JALISCO: Mpio. La Huerta, camino antiguo Chachalaca-Arroyo Zarco, 28.vii.1982, Bullock 1217 (BH); Mpio. Casimiro Castillo, 7 8 km NNE of Casimiro Castillo, Arroyo La Calera, m, 14.vii.1988, Cuevas & Nuñez 3066 (WIS); Mpio. Tecalitlán, rd km along dirt rd to San Isidro that leaves old Colima-Tecalitlán road about 7 rd mi S of city of Tecalitlán, 1200 m, 15.viii.1991, Dean & Starbuck 249 (DAV, XAL); Mpio. Tecalitlán, Sierra del Halo, c. 2 rd mi along rd to San Isidro (or Jilotlan) that leaves old Colima-Tecalitlán rd c. 7 rd mi S of Tecalitlán, 1340 m, 23.xi.1991, Dean & Starbuck 329 (DAV, MEXU, NY, UC, XAL); Mpio. El Tuito, About 3 rd mi N of El Tuito along hwy 200, footpath enters woods above a running river on W side of hwy, 2400 m, 27.xi.1991, Dean & Starbuck 332 (DAV, MEXU, NY, UC, XAL); Sierra de Manantlán, 3 km al SE de Lagunillas, La Mochota, Cuautitlán, N, W, 850 m, 1.vii.1990, De Niz et al. 173 (WIS, ZEA); entre el Tuito y Puerto Vallarta, a 20 km de Puerto Vallarta y a 20 km de El Tuito, 19.vii.1976, Delgado-S. & Hernández 358 (Hernández & Delgado-S. 2609) (MEXU); no exact locality, Diquet s.n. (NY); Mpio. Tecalitlán, alrededores de Plan de Lego, 51 km directamente al SSE de Ciudad Guzmán, al E de P. de Lego, brecha a Las Animas, 8.viii.1988, Fuentes-O. 496 (NY); 4 km S of Mazamitla, 1600 m, 7.vii.1967, Hernández-M. 407 (GH, MEXU); steep N-facing hills, 0.5 km S of Puente San Pedro, on road from Colima to Ciudad Guzmán, N, W, m, 31.vii.1960, w 609 (ENCB, WIS); Mpio. La Huerta, Rancho Cuixmala and environs, S side of Río Cuixmala, dirt road heading upriver (E), c. 3 km inland from Puerto Vallara-Barra de Navidad (Mex. 200) hwy, N, W, 9.vii.1991, Lott et al (CAS, NY); Sierra del Halo, near a lumber road leaving the Colima hwy 7 mi SSW of Tecalitlán and extending SE toward San Isidro, about 2 mi from the hwy, 1400 m, 13.viii.1957, McVaugh (TEX; duplicate specimens at MEXU, NY, and TEX are L. moziniana); Sierra de Manantlán, 40.5 km al SE de Autlán, 1 2 km al NE de Telcruz, Cuautitlán, N, W, 1100 m, 18.vii.1987, Vázquez & López-V (UC, ZEA). MÉXICO: Mpio. Ixtapan de la Sal, 2 rd km NE of the NE gates of the town of Ixtapan de la Sal, along hwy 55 (to Toluca), 1920 m, 12.vii.1990, Dean & Starbuck 210 (DAV, MEXU); Mpio. Ixtapan de la Sal, 2 rd km NE of the NE gates of the town of Ixtapan de la Sal, along hwy 55 (to Toluca), 1920 m, 12.vii.1990, Dean & Starbuck 211 (DAV, MEXU, UC, XAL); Mpio. Ixtapan de la Sal, 2 rd km NE of the NE gates of the town of Ixtapan de la Sal, along hwy 55 (to Toluca), 1830 m, 2.xi.1991, Dean & Starbuck 310 (BM, DAV, ENCB, MEXU, MO, NY, UC, XAL); Mpio. Temascaltepec, c rd mi S of Temascaltepec along hwy 134, E and W sides of hwy, 2030 m, 7.xi.1991, Dean & Starbuck 314 (DAV, ENCB, MEXU, NY, UC, XAL); District Temascaltepec, Vigas, 1080 m, 30.vii.1932, Hinton et al (BM, F, G); District Temascaltepec, Rincón, 1960 m, 10.ix.1932, Hinton et al (GH, K, NY); Plaza del Gallos, 1200 m, 27.viii.1933, Hinton et al (BM, F, G, MO); Los Bejucos, Tejupilco, 700 m, 27.viii.1954, Matuda (MEXU); 2 km N of Ixtapan de la Sal along hwy 55, 1650 m, 28.vii.1964, Mick & Roe 348 (WIS); 6 km S of Temascaltepec on Hwy 130, N, W, 2000 m, 3.ix.1965, Roe et al (ENCB, UC, WIS); 4 km S of Temascaltepec, N, W, 1800 m, 4.ix.1965, Roe et al (ENCB, UC, WIS); 2 km N of Ixtapan de la Sal, at km 145 on hwy 55, N, W, 1930 m, 9 10.ix.1965, Roe & Roe 1902 (ENCB, UC, WIS); Temascaltepec, cañada al oeste de la población, 1800 m, 4.ix.1965, Rzedowski (DS-CAS). MICHOACÁN: District Coalcomán, Pto. Zarzamora, 3.viii.1939, Hinton et al (F, GH, LL, MO, NY); Mpio. Tancítaro, rd from Tancítaro to Apatzingán, 5000 ft, 30.vii.1940, Leavenworth 386 (F); just above Tacámbaro on highway to Pátzcuaro, 1700 m, 20.vii.1948, Moore & Wood 4020 (A). MORELOS: Huitzilac, 1.vii.1930, Lyonnet 713 (MEXU mixed collection with L. rzedowskii, NY; duplicate specimens at BM, GH, MO AND MEXU are L. rzedowskii); barranca of Cuernavaca, 5000 ft, 17.vi.1904, Pringle (BH, C, CAS, K, GH, L, MO, S); Cuernavaca, Schmitz s.n. (GH, W); Cuernavaca, Schmitz 1091 (W); OAXACA: 13 mi E of Pinotepa Nacional on Mex Hwy 200, NW of Puerto Escondido,

15 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) v.1973, Hansen et al (WIS, MEXU); Dist. Putla, 13 km al S de Copala o 2 km al S de la desviación a Tlaxiaco, hacia Putla, 2600 m, 19.vi.1982, Torres-C. & Cedillo-T. 598 (MEXU, MO, NY); Dist. Pochutla, 6 km al NE de Chacalapa en la brecha a la finca Ojo de Agua, 14.vi.1985, Torres-C. & García-M (MEXU, MO, NY). PUEBLA: Sierra de Chalchi, vi.1945, Miranda 3551 (MEXU). NICARAGUA: DEPARTMENT MANAGUA: Managua, 29.vi.1927, Chaves 278 (F, US); vicinity of Managua, viii.1932, Garnier 1179 (US); km 48 carretera al Ingenio Julio Guitrago (Montelimar), c N, W, 100 m, 25.ix.1982, Grijalva 1250 (MO); Esquipulas, 10 km SE of Managua, 13.vi.1977, Neill 2122 (MO); campus of the University Centroamericana, Managua, 125 m, 7.ix.1981, Stevens (MO). Lycianthes ciliolata (M. Martens & Galeotti) Bitter (Fig. 8) Figure 8. Herbarium specimen of Lycianthes ciliolata (Dean & Starbuck 225 (DAV)). Solanum ciliolatum M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12: Lycianthes ciliolata (M. Martens & Galeotti) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO. State of Oaxaca, Cordillère orientale de Oaxaca, dans les bois de la Sierra de Capulalpan et de Llano Verde, Rocheu de Castrasana, de 6000 à 7000 pieds, Septembre 1840, Galeotti 1230 (LECTOTYPE here designated: BR!, photos BH!, WIS!; ISOLECTOTYPE BR!). Solanum somniculentum Kunze ex Schltdl., Linnaea 19: Lycianthes somniculenta (Kunze ex Schltdl.) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: GERMANY, grown by G. Kunze in the Leipzig Botanical Garden c from seed brought from Mexico by C.A. Ehrenberg, G. Kunze s.n., 1845 (LECTOTYPE here designated: W!). Chosen from syntypes: G. Kunze s.n., Schiede s.n. (HAL!), Leibold 143 (B, probably destroyed). See discussion below after specimens examined section. Solanum andrieuxi Dunal, in de Candolle, Prodr. 13(1): Lycianthes mociniana Dunal var. andrieuxi (Dunal) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO, no exact locality (probably from southern Puebla or Oaxaca), no date, Andrieux 195 (HOLOTYPE: M!). Lycianthes ciliolata (M. Martens and Galeotti) Bitter var. pratorum Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: GUATEMALA. Department Baja Verapaz, Patal, 1600 m, Von Turckheim 2317 (LECTOTYPE designated here: W!; ISOLECTO- TYPES: BR!, F!, G!, MO!, NY!, US!). Chosen from syntypes Von Turckheim 2317, Von Turckheim 1434 (F!, GH!, K!, US!). Lycianthes somniculenta (Kunze ex Schltdl.) Bitter var. lanceolata Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO. Oaxaca, no exact locality, 1842, Ghiesbrecht 81 (HOLOTYPE: P!). See discussion below after specimens examined section. Perennial HERBS, erect, tall plants often recumbent with age (10) (100) cm tall. INDUMENT of simple trichomes in Puebla, Chiapas and Guatemala, these sometimes dendritic in Oaxaca. Roots with fusiform segments, usually present cm below the soil surface. STEMS arising from the root one to several; below-ground growth not extensive, white; aboveground growth greenish-purple, terete in cross-section, woody with age, usually not much compressed when dried, the new growth glabrous to pubescent, the stems often glabrate with age, the trichomes (1.5) mm long, varying from appressed-antrorse (Puebla) to appressed-retrorse (Chiapas and Guatemala) or spreading (Chiapas and Oaxaca). FIRST SYM-

16 400 E. A. DEAN PODIAL UNIT erect above ground, 5 60 cm long, 2 8 mm diameter at widest point; internodes 4 10 (19) in number; lateral branching usually not extensive. FIRST TWO SYMPODIAL BRANCHING POINTS nearly always dichasial, followed by monochasial branching (rarely followed by further dichasia or the second dichasial branching points lacking), sometimes lateral branching from monochasial or dichasial branching points creating false dichasia or trichasia; first dichasial branching angle (20) (120). SUBSEQUENT SYMPODIAL GROWTH often extensive and with age equalling or surpassing the first sympodium in length, extending laterally 3 40 cm, the second sympodial units (1) 2 12 (20) cm long, mm diameter, subsequent sympodial units becoming reduced distally. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT chartaceous; leaves closest to the soil surface reduced, scale-like, oblanceolate to round, sessile or with attenuate base, cm long, cm wide, early deciduous; subsequent leaves usually expanded (rarely only scale leaves present), often early deciduous, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for 1/8 1/2 of total leaf length, the lamina usually ovate (rarely obovate or elliptic), the margin undulate, the tip acute to acuminate, the base truncate, obtuse, or cuneate, attenuate into the petiole, the largest expanded leaves (generally at the top or in the upper one third of stem) cm long, cm wide, the petiole cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate, similar in shape and texture to the distal-most leaf of the first sympodial unit, but the attenuate portion plus the petiole accounting for 1/8 1/3 of total leaf length, the lamina shape more lanceolate, and the petiole shorter or lacking; largest leaves (often produced by second sympodial units except in Chiapas where they are produced by the third or fourth sympodial units), 2 14 cm long, 1 7 cm wide, with petiole to 1.5 cm long, the leaves becoming reduced and more sessile on more distal sympodial units; geminate leaf pairs unequal in size (the smaller 1/4 3/4 of the larger), the smaller of the pair with shorter and more oblique leaf base. LEAF PUBESCENCE similar throughout, the leaves glabrous to pubescent, the trichomes mm long, adaxially appressed, abaxially erect to appressed, the trichomes of the main veins spreading to slightly antrorse in Puebla and Oaxaca, appressedantrorse in Chiapas and Guatemala, the trichomes of the margin mm long, appressed-antrorse in most populations, spreading in Oaxaca. PEDICELS AT ANTHESIS green or purple, 3 9 cm long, (0.5) 1 2 (3) times as long as subtending leaf, glabrous to pubescent, the trichomes mm long, spreading to slightly retrorse in Puebla and Oaxaca, becoming appressed-antrorse in Chiapas and Guatemala, sometimes glabrate with age. CALYCES AT ANTHESIS green or purple, obconic, mm long, (2.5) (6.5) mm diameter when fresh, (4) when pressed and dried, the teeth reflexed, emerging from the ribs at approximately the same level, of equal length or of two alternating lengths, (0) 4 9 (11) mm long, the ribs and teeth glabrous or pubescent with trichomes mm long, these usually erect or slightly retrorse, becoming appressed-antrorse in Guatemala. COROLLAS orientated laterally, not nodding, the tube 2 mm long, the limb (2.1) (5.3) cm in diameter, the membrane lilac, the adaxial sides of the lobes of similar colour but with darker violet veins, the abaxial sides of the lobes green, usually glabrous (puberulent with very sparse trichomes, mm long in some populations in Oaxaca). ANDROECIUM with green, erect, glabrous, laterally compressed filaments, the lowest (2) 3 7 (8) mm long, the two lateral (5.5) mm, the two upper (0.5) (4.25) mm, the length of the lowest usually times that of the lateral (rarely times); anthers exuding a sweet odour, (3) (8) mm long, the lowest anther usually lanceolate to oblong (rarely narrowly elliptical), the pores slit-like to narrowly obovate, lateral, generally dehiscing toward one another; pollen grains dicolporate with a remnant third pore. GYNOECIUM with conic ovary, mm long, the ovules ; style straight to slightly curved, 9 12 (13) mm long; stigma green, usually round, rarely shallowly bilobed (in some Oaxaca populations) to very bilobed (in Guatemala). FRUITS remaining attached to calyces at maturity, ovoid, mm long, (7) 9 22 mm in diameter, the tip apiculate; exocarp purple to nearly black, rarely with lighter purple dots, glabrous; mesocarp dark purple, soft and juicy, lacking sclerotic granules; placental area light purple, powdery; fruiting calyx enlarged, 2 4 mm long, 5 11 mm in diameter, the teeth reflexed, often broken, (0) 3 10 (11) mm long; fruiting pedicels (3) (11) cm long, deflexed, the fruit pendant; seeds (8) (102) per fruit, black, depressed-obovate, rough in texture, ridged along one side, often blistered along the ridge, mm long, mm wide. Chromosome number: 2n = 24 (Dean & Starbuck 271, 295). Indigenous names: hoh, yich hoh, yichjoj, tintolon, ma u dsea nuu jgiaa, u dsea niquia, yich balam, campanilla, chichi de vaca, chichi de perro, chichi de venado, binduchi, binduchito, rshtisti katya, tronchichi, tonchicho, kuan xille bekue, manzanillo del campo, shashasto, la pera. Distribution: Guatemala and Mexico (states of Puebla, Oaxaca, Chiapas and possibly San Luis Potosí). Lycianthes ciliolata grows on limestone soils of the

17 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 401 Sierra Madre Oriental of Mexico and Guatemala. The populations from San Luis Potosí may be hybrids with L. moziniana var. margaretiana, and are not included in the general description provided for the species here. These collections resemble what I am calling L. ciliolata when in flower, and as with that species, they grow on limestone. Field collection of fruits was unsuccessful, but I was able to produce several fruits on one accession of Dean & Starbuck 251 in the greenhouse. Although the accession is self-incompatible, it set fruit when crossed with Dean & Starbuck 206 (L. ciliolata from Tehuacán, Puebla). Seeds within the fruit did not have embryos, a very unusual result in my crossing experiments with L. ciliolata, especially those involving Dean & Starbuck 206; generally, this accession is highly crossable (producing fruits with mature seeds when crossed with accessions from as far away as Chiapas). In addition, the fruit colour was not the usual opaque purple-black that one always sees with L. ciliolata. Rather, it was a translucent rose colour, with an exocarp texture more similar to that of L. moziniana. Until I am able to collect in this area further, and confirm that this fruit-type occurs in the field as well as the greenhouse, I hesitate to name these collections as a separate species or variety. Vegetation types occupied by the species range from cactus scrub (matorral of Rzedowski, 1988) in the states of Puebla and Oaxaca, to moist upland oak or oak pine woodland in Oaxaca and Chiapas. Habitats include drainages, clearings, paths and agricultural fields, m. Diagnostic features: Lycianthes ciliolata is variable in height, leaf size and pubescence (both trichome type and degree of pubescence). In dry habitats, plants are shorter and have smaller leaves than in moister habitats. Whereas L. ciliolata has simple trichomes in most areas, Oaxacan plants often have dendritic trichomes. In addition, in the Valley of Oaxaca and the surrounding upland areas, plants appear to fall into two groups, those that are nearly glabrous and those that are villous. The two forms often grow side by side without intermediates. Lycianthes ciliolata may be confused with L. rzedowskii and L. acapulcensis. It differs from those two species in having lilac rather than white corollas (or the very pale lilac sometimes found in L. rzedowskii). The pattern of filament lengths can be useful in separating it from L. rzedowskii. In L. ciliolata the lowermost filament is often more than twice as long as the lateral filaments, which is not the case in L. rzedowskii. As discussed above, the lengths of the pedicels of the youngest mature flowers relative to their subtending leaves is often a useful character for separating L. ciliolata from L. acapulcensis (see discussion under L. acapulcensis). There are a few older collections collected on the Sessé and Moçiño expedition and by Ghiesbrecht (presumably from Oaxaca) that are intermediate between L. ciliolata and L. acapulcensis. I have placed these specimens in L. ciliolata for the present (see discussion below). Intermediates between this species and Lycianthes moziniana occur in the Sierra de Juárez of Oaxaca and San Luis Potosí, and in those regions distinguishing the two species can be difficult. The best distinction is in the pollen grains (tricolporate in L. moziniana and dicolporate with a remnant third pore in L. ciliolata). Specimens examined: FIELD ORIGIN UNKNOWN: Bernhardi herbarium s.n. (MO ); Leipzig Botanic Garden, Germany, no date or coll. number, tag on specimen says 1939 (M); probably a European Botanic Garden specimen, C.H. Beer.? 853. Sep (W); probably from a German botanical garden, ex herbario Bitter s.n. (GOET); H. B. Leipzig (Leipzig Botanic Garden), Germany, 1857, Kunze? s.n. (BR); Leipzig Botanic Garden, Germany, Kunze s.n. (W); Horto. Bot. Turionor (Zurich Botanic Garden), Regel 478? or 1847 no. 8? (ZT); cultivated in the garden of Van Houtte, France, 17.ix.1849, Van Houtte 390 (K); H. B. Berol (Berlin Botanic Garden), Germany, 1854?, Herb. Vocke. s.n. (GOET); GUATEMALA: DEPARTMENT BAJA VERAPAZ: Agua Caliente, 755 m, 25.v.1906, Cook 38 (US); Patal, zwischen Santa Rosa und Tastic, 5000 ft, ix.1888, Von Turckheim 1434 (F, GH, K, US); HUEHUE- TENANGO: Vicinity of Chinacho, 10 km W of Zacaleu Ruins, 1900 m, 14.ix.1971, Molina-R (F); Trail between Ixteapoc and Soloma, Sierra de los Cuchumatanes (near Soloma), m, 8.vii.1942, Steyermark (F); Cerro Pixpix, above San Ildefonso, Ixtahuacan, m, 15.viii.1942, Steyermark (F, NY); Cumbre Papal, on S-facing slopes between Cuilco and Ixmoqui, m, 19.viii.1942, Steyermark (F, G); Slopes above La Libertad on Cerro Pueblo Viejo, 1900 m, 20.viii.1942, Steyermark (F). MEXICO: State unknown: 1833, Andrieux, 163 (G); Liebmann 1451 (C); Vermultl. Provinz Veracruz oder Oaxaca, vii.71961, Schwabe s.n. (B); , Sessé, Moçiño, Castillo et Maldonado 1529 (photo NY of Madrid specimen, F negative 48221; another photo at NY of a duplicate specimen is L. moziniana); , Sessé, Moçiño, Castillo et Maldonado 5389 (photo NY of Madrid specimen F negative 4822); , Sessé, Moçiño, Castillo et Maldonado 5389 (F- newer label says 5389, small old label says 239 and S. uniflorum); Toluca? (probably in error), Wawra 1214 (W). CHIAPAS: Mpio. Tenejapa, near the school house of Pokolum in the Paraje of Sibanil Ha, 5100 ft, 10.vii.1964, Breedlove 6100 (DS); Mpio. Teopisca, NW edge of Teopisca along Mexican

18 402 E. A. DEAN hwy 190, 5900 ft, 25.vi.1965, Breedlove (DS); Mpio. Tuxtla Gutiérrez, 15 km N of Tuxtla Gutiérrez along rd to El Sumidero, 3800 ft, 2.vii.1965, Breedlove (DS, F); Mpio. Tenejapa, near the schoolhouse of Pokolum, paraje of Sibanil Ha, 5200 ft, 15.vii.1965, Breedlove (DS, ENCB, F); Mpio. Venustiano Carranza, 3 mi S of Aguacatenango along rd to Pínola Las Rosas, 5600 ft, 22.vii.1965, Breedlove (DS); Mpio. Tenejapa, Paraje of Kulak tik, 5500 ft, 18.vii.1966, Breedlove (DS, F, LL); Mpio. La Trinitaria, Lagos de Montebello, 42 km NE of La Trinitaria, along the Comitán River at its sumidero, 1300 m, 23.x.1971, Breedlove & Thorne (DS); Mpio. Totolapa, Rancho Ch a ha, 5 6 km W of Teopisca, 1500 m, 26.xi.1971, Breedlove (DS); Mpio. San Andrés Larrainzar, near the summit of Chuchil Ton, NE of Bochil, 2700 m, 17.x.1972, Breedlove (DS, MO); Mpio. San Cristóbal de Las Casas, on road to San Lucas Zapotal, 2 4 km from Mex hwy 190, 2400 m, 8.ix.1974, Breedlove (DS, MO); a 5 km al N de la carretera San Cristóbal de las Casas- Comitán, con rumbo a Ococingo, 19.vi.1982, Cabrera & Cabrera 2813 (NY); Mpio. San Cristóbal de las Casas, just outside city of San Cristóbal de las Casas, along rd to hwy 190 which diverges from rd to Tenejapa, nearly opposite trailer park, near Almolonga River, m, 16.vii.1990, Dean & Starbuck 213 (DAV, MEXU); Mpio. Zinacantán, hills SE of the town of Zinacantán, along footpath to Paraje Sok on, several km uphill from Zinacantán, 2463 m, 17.vii.1990, Dean & Starbuck 214 (DAV, MEXU, NY, UC, XAL); Mpio. Teopisca, c. 8 rd km NW of Teopisca along hwy 190, NE side of road, 2103 m, 18.vii.1990, Dean & Starbuck 215 (DAV, MEXU); no exact locality, Ghiesbrecht 58 (GH -mixed collection with L. pilifera); terre froide, vii , Ghiesbrecht 827 (BM, GH, MO); Yochib Ja, 5 km al N de Amatenango del Valle, 5500 ft, 15.iv.1988, Gómez-L. 421 (MO); Ixtapa, carretera 190 en el km 75 entre Tuxtla Gutiérrez y San Cristóbal de las Casas, 2250 m, 2.vii.1991, González-V et al (WIS); 32 mi SE of San Juan Cristóbal de las Casas, 7100 ft, 23.vi.1950, Johnston (TEX); Mpio. Zinacantán, valley floor in Zinacantán Center, 6700 ft, 7.vi.1966, Laughlin 1012 (DS); Mpio. Zinacantán, Paraje Vo bits, 4300 ft, 7.vi.1966, Laughlin 1016 (DS); Paraje Ya al Ichin, 18 km al sur de Chamula, 6800 ft, 21.vii.1988, López-M. 955 (MO); Mpio. Tenejapa, Pocolum, 1600 m, 15.ix.1982, Méndez-T. & Shilom-T (NY); Tak i, 3 km al sur del poblado, 4050 ft, 12.vii.1988, Santíz-C. 686 (MO, TEX); Mpio. Tenejapa, paraje of Kulak tik, 5800 ft, i iii.1964, Shilom-T. 250 (DS); Mpio. Teopisca, 5 mi NW of Teopisca, N, W, 6100 ft, 20.viii.1972, Webster & Lynch (DS); Mpio. Pueblo Nuevo Solistahuacan, 3 km NW of Pueblo Nuevo Solistahuacan, W, N, 5400 ft, 19.vii.1970, Zuill 99 (DS); Mpio. Pueblo Nuevo Solistahuacan, 3 km NW of Pueblo Nuevo Solistahuacan, 5400 ft, 18.ix.1970, Zuill 362 (DS). OAXACA: Mt. San Felipe, vii.1834, Andrieux 182 (G-DC n.v., photos WIS, NY, GH); near Oaxaca, at San Felipe del Agua, 1650 m, 1.ix.1895, Conzatti 132 (GH); vi.1901, Conzatti & González 1217 (GH); vi.1901, Conzatti 1218 (GH); Mpio. Huajuapan de León, c. 5.5 rd mi NW of Huajuapan de León along hwy 190, just S of tourist turnout, SW side of the road, 1920 m, 16.vii.1991, Dean & Starbuck 229 (DAV, XAL); Mpio. Tlalixtac, along hwy 175, NE of the city of Oaxaca, c. 4.3 rd mi E of the Juárez monument, where electrical wires cross rd, SE side of road, across river, 1740 m, 19.vii.1991, Dean & Starbuck 231 (DAV, UC, XAL); Las Animas, along hwy 175, NE of the city of Oaxaca, 2380 m, 19.vii.1991, Dean & Starbuck 232 (DAV, NY, XAL); Mpio. Calpulalpan, Calpulalpan, 2040 m, 20.vii.1991, Dean & Starbuck 233 (DAV, MEXU, MO, NY, XAL); Mpio. Tlacochahuaya, a very short distance N of turnoff to Dainzu along hwy 190, S of the city of Oaxaca, in the Oaxaca Valley, W side of road, 1585 m, 22.vii.1991, Dean & Starbuck 234 (DAV, UC, XAL); Mpio. Tamazulapan, along hwy 175 between Miahuatlán and Puerto Angel in the Sierra Madre del Sur, 0.7 rd mi NW of the town of San Andrés, c rd mi SE of Miahuatlán, 2286 m, 25.vii.1991, Dean & Starbuck 237 (DAV, UC, XAL); Mpio. Matatlán, along cobblestone road to radio tower, c. 3.6 rd mi S of town of Matatlán, S of city of Oaxaca along hwy 190, c. 1 mi S of km 595, 1006 m, 29.vii.1991, Dean & Starbuck 241 (DAV, XAL); Mpio. Huajuapan de León, 5.5 rd mi NW of Huajuapan de León along hwy 190, just S of tourist turnoff, SW side of road, 1920 m, 9.x.1991, Dean & Starbuck 282 (XAL); Mpio. Nochixlan, NW side of town of Nochixlan, near large drainage that crosses hwy 190, 2073 m, 9.x.1991, Dean & Starbuck 284 (DAV, XAL); Mpio. Santa María Jaltianguis, upper areas of Sta. María Jaltianguis, along dirt road, in front of residence, 2287 m, 11.x.1991, Dean & Starbuck 286 (DAV, XAL); Mpio. Calpulalpan, Calpulalpan, 2042 m, 12.x.1991, Dean & Starbuck 287 (BM, DAV, F, MEXU, MO, NY, UC, XAL); Mpio. Matatlán, along cobblestone road to radio tower, about 3.6 rd mi S of Matatlán, along hwy 190 between Oaxaca City and Tehuantepec, 1006 m, 13.x.1991, Dean & Starbuck 290 (DAV, NY, UC, XAL); Mpio. Tlacochahuaya, a very short distance N of turnoff to Dainzu, along hwy 190, between Mitla and Oaxaca City, W side of the rd, 1585 m, 14.x.1991, Dean & Starbuck 291 (DAV, XAL); Mpio. Tamazulapan, along hwy 175 between Miahuatlán and Puerto Angel, c rd mi SE of Miahuatlán, near turnoff to town of San Andrés Paxtla, 2287 m, 18.x.1991, Dean & Starbuck 294 (DAV, MEXU, MO, NY, UC, XAL); Mpio. Sta. María Jaltianguis, Sta. María Jaltianguis, 2195 m, 20.x.1991, Dean & Starbuck 295 (DAV, MEXU, MO,

19 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 403 NY, UC, XAL); Oaxaca Valley, on the western foothills of Sierra Oaxacana, along hwy 175, 7.2 rd km E of hwy 190 (at Juárez Monument), N, W, 1799 m, 29.vii.1978, Diggs & Corcoran 2148 (WIS); along hwy 131 between Oaxaca and Puerto Escondido, 15 km by road S of Sola de Vega, N, W, 2012 m, 25.vi.1986, Diggs et al (NY, TEX); cordillera, 6500 ft, xi iv.1840, Galeotti 1166 (BR); San Pedro Nolasco (note by M. Nee, c. 60 km NE of city of Oaxaca), vii.1844, Galeotti 1225 (1225 L) (BR, NY); Mpio. Santiago Comáltepec, Santiago Comáltepec, N, W, 2000 m, 21.vi.1987, Hernández-G. 31 (UC); San Pedro Nolasco, Talea, & c., , Jurgensen 695 (BM, G); Mpio. Santiago Comáltepec, Santiago Comáltepec, N, W, 2000 m, 18.vi.1988, López-L. 132 (UC); Mpio. Santiago Laxopa, Santiago Laxopa, N, W, 2000 m, 26.v.1986, Maldonado-V. 19 (UC); 15 km (by road) SE of Miahuatlán on road to Puerto Angel in high mountains of Sierra Madre de Sur, N, W, 2400 m, 6.vii.1969, Marcks & Marcks 1031 (WIS); E slope of El Cumbre Mtns, N, W, 21.vii.1966, Schoenwetter JSOX25 (US); Tenango, 5.vii.1895, Smith 400 (GH); along road to microwave tower, c. 3.6 mi S of Matatlán on hwy 190, about 1 mi S of km 595, 22.vii.1971, Stevens 1297 (ENCB, MO, WIS). PUEBLA: Mpio. Tehuacán, Barranca Cruz de Quiote, in the hill to the NE of city of Tehuacán, NE of the army rifle range and cemetery, m, 9.vii.1990, Dean & Starbuck 206 (DAV, MEXU); Mpio. Tehuacán, El Riego Mesa on the W side of the city of Tehuacán, 1750 m, 9.vii.1990, Dean & Starbuck 207 (DAV, MEXU); Mpio. Caltepec, 0.85 road km E of junction of Atecoxco-San Pedro Atzumba road with Acatepec-Caltepec road, along road to Caltepec, fields on S side of road, 2165 m, 10.vii.1990, Dean & Starbuck 208 (DAV, MEXU); Mpio. Tehuacán, E side of city of Tehuacán, just W and SW of El Riego mesa, below El Riego cave and by road leading up to hills, 1677 m, 13.vii.1991, Dean & Starbuck 225 (DAV, MEXU, NY, UC, XAL); Mpio. Caltepec, between Cerro Pochote and Cerro Gavilán Chico in hills SE of town of Caltepec, along road to Atolotitlan, m, 15.vii.1991, Dean & Starbuck 227 (DAV, XAL); Mpio. Tehuacán, old field just to the E of the mesa at El Riego, and drainage that crosses dirt road to the SE of El Riego, 1677 m, 27.ix.1991, Dean & Starbuck 271 (DAV, MEXU, NY, UC, XAL); Mpio. Tehuacán, hills to the SW of the city of Tehuacán, up dirt road near El Riego, m, 27.ix.1991, Dean et al. 272 (DAV, XAL); Mpio. Caltepec, 0.85 km E of junction of Acatepec-Caltepec and Atecoxco-San Pedro Atzumba roads, along road to Caltepec, 2164 m, 30.ix.1991, Dean et al. 275 (DAV, NY, XAL); Mpio. Caltepec, hills SE of the town of Caltepec, between Cerro Gavilán Chico and Cerro Pochote, as well as an area called Rancho Gavilán, near rd to Atolotitlan, 2225 m, 30.ix.1991, Dean & Starbuck 277 (DAV, UC, XAL); Mpio. Zapotitlán de las Salinas, San Antonio Texcala, along hwy 125 S of Tehuacán, canyon with onyx mine just N of town, 1677 m, 22.x.1991, Dean & Starbuck 299 (DAV, XAL); 3 mi N of the city limits of Tehuacán, 5500 ft, 7.vi.1973, Hansen et al (MEXU, WIS); Chila Zapotitlán (perhaps between Chila and Zapotitlán), 15.vii.1943, Miranda 2845 (MEXU); Tehuacán, 5000 ft, 6.viii.1897, Pringle 6776 (BM, BR, G, GH, GOET, HBG, JE, LL, M, MEXU, NY, S, UC, W, Z); El Riego, vi.1905, Purpus 1279 (MO, UC); in the vicinity of San Luis Tultitlanapa, near Oaxaca, vii.1908, Purpus 3561 (MO, UC); vicinity of San Luis Tultitlanapa, near Oaxaca, vii.1908, Purpus 3562 (BM, UC); W of Tehuacán on the Mesa above El Riego, 1500 m, vii.1961, Smith et al (F, G, MEXU, NY); Mpio. Atzingo, Tlacuilosto al S de Atzingo, 1900 m, 22.vii.1985, Tenorio-L et al (TEX); 9.6 km al NW de Huajuapan de León, carr. a Acatlán, antes de la torre de microondas, 1770 m, 29.vii.1983, Torres-C (MEXU). SAN LUIS POTOSÍ: Mpio. Zaragoza, c rd mi W of outskirts of town of Santa Catarina along hwy 70 (Río Verde-San Luis Potosí hwy), , 3.vii.1991, Dean & Starbuck 222 (DAV, XAL); Mpio. Zaragoza, c. 1.6 rd mi E of little town of San Francisco, along hwy 70 (Río Verde-San Luis Potosí hwy), W of Santa Catarina, large turnout at site of small shrine, , 3.vii.1991, Dean & Starbuck 223 (DAV, NY, XAL); Mpio. Zaragoza, 6.1 rd mi E of turn off to Sierra de Alvarez at town of San Francisco, along hwy 70 between city of San Luis Potosí and Río Verde, 1680 m, 15.ix.1991, Dean & Starbuck 251 (DAV, MEXU, NY, XAL); 56 km E of San Luis Potosí or 18 km W of Santa Catarina on Hwy 86, N; W, 1100 m, 10.vii.1965, Roe & Mori 75 (ENCB, WIS). Schlechtendal cited three syntypes for the name Lycianthes somniculenta, one based on cultivated material sent to Schlechtendal by Kunze from the Leipzig Botanical Garden and two herbarium specimens: MEXICO. Mexici ad Acalingo, Leibold 143; pr. Zimapán et pr. Reglam, Schiede s.n. Schlechtendal mentioned at the end of his protologue that the three specimens are quite distinct in some of their characters, thus admitting that they do not all match his description. I have not seen authentic material of the Leibold specimen, which was at B and may have been destroyed in the World War II. It was determined by G. Bitter to be L. moziniana, and until I see authentic material, I will accept his judgement. I have seen the Schiede specimen, which I have determined is L. dejecta, and it is different from the material from the Leipzig Botanical Garden. Kunze herbarium specimens collected from the Leipzig Botanical Garden do exist, from which I have chosen a lectotype. According

20 404 E. A. DEAN to Planchon (1849), the material in the Leipzig Botanical garden was grown by Kunze from seed brought to him by M. C. Ehrenberg. The type collection of Lycianthes somniculenta var. lanceolata, Ghiesbrecht 81 (from Oaxaca), which I have included in the circumscription above, is somewhat atypical of the species I am calling L. ciliolata. It has rather large lance-ovate leaves with the attenuate portion very short, the one nearly mature pedicel is shorter than its subtending leaf, and the pollen, when stained with aniline blue, appeared to be dicolporate. However, it has an unlobed stigma, and its lateral anthers dehisce toward the style. Given that the two collections of L. acapulcensis from Oaxaca that I have examined both have very large bilobed stigmas and lateral anthers that dehisce away from the style, it is likely that this collection is conspecific with what I am calling L. ciliolata. I have encountered similar specimens of L. ciliolata (with lance-ovate leaves and short pedicels) from the mountains south of Miahuatlan in Oaxaca, although generally the pollen is 2.5-pored. It may be that the older pollen on this specimen when treated with aniline blue does not retain its original shape. I have had similar problems with the Sessé, Moçiño, Castillo et Maldonado specimens from MA and F, which I have included in the circumscription of L. ciliolata (listed under Mexico, no exact state). Again, the pollen when stained with aniline blue is not obviously 2.5-pored, yet the specimens do not fall comfortably within the circumscription of L. acapulcensis. I assume that these specimens are from Oaxaca, as is the Ghiesbrecht specimen discussed above. Lycianthes dejecta (Fernald) Bitter (Fig. 9). Solanum dejectum Fernald, Proc. Amer. Acad. of Arts: 35: Lycianthes dejecta (Fernald) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: MEXICO. State of Durango, near city of Durango, Iron Mountain and vicinity, rare in crevices of rocks, vii.1896, E. Palmer 347 (LECTOTYPE here designated: GH!; ISOLECTOTYPES: BM!, C!, F!, CAS!, G!, NY!, S!, UC!, US!). Chosen from syntypes: Palmer 347, Pringle 7207 (GH!), Dugès s.n. (GH!), Hahn (Bourgeau) 543 (BR!, F!, G!, GH!, M!). Lycianthes dejecta (Fernald) Bitter var. palmeri Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: as above, E. Palmer 347 (HOLOTYPE B, probably destroyed; ISOTYPES as listed above). Perennial HERBS, decumbent to erect, decumbent forms to 50 cm in diameter, tall forms often scrambling, supported by nearby shrubs, to 50 cm tall. INDU- MENT of multi-angulate-dendritic trichomes mixed with a few dendritic or simple trichomes. ROOTS with widely fusiform segments, often warty with age, usually present cm below the soil surface. STEMS arising from the root one to several; below-ground growth not extensive, white; above-ground growth with green and purple striations, terete to angulate in cross-section, without much woody tissue, generally much compressed when dried, pubescent to tomentose, the trichomes spreading, (0.75) mm long. FIRST SYMPODIAL UNIT decumbent to erect above ground, 1 35 cm long, 2 11 mm diameter at widest point, narrowest at the soil level; internodes 2 7 (13) in number; lateral branching occurring early from the leaf axils. FIRST SYMPODIAL BRANCHING POINT dichasial, followed by a mixture of monochasial and dichasial branching, with lateral branching common; first dichasial branching angle SUBSEQUENT SYM- PODIAL GROWTH extensive, often surpassing the monopodium in length, extending 6 45 cm, the second sympodial units 3 15 cm long, mm diameter, these generally followed by one to several longer sympodial units, with subsequent sympodial units becoming reduced distally. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT chartaceous, often early deciduous; leaves closest to the soil surface reduced, spathulate, 1 5 cm long, 1 4 cm wide; subsequent leaves larger, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for 1/3 2/3 of total leaf length, the lamina ovate to reniform, margin entire to slightly undulate, the tip rounded to acute, the base obtuse to truncate then long-attenuate into petiole, the largest expanded leaves (usually occurring in the middle to most distal portion of sympodium) cm long, cm wide, the petiole poorly defined, 2 4 cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate or not, due to the mixture of monochasial and dichasial branching, the leaves similar in shape and texture to the distalmost leaves of the first sympodial unit, but the lamina ovate to deltoid, the attenuate portion plus petiole accounting for 1/8 3/4 of total leaf length, the margin slightly undulate, the tip acute and the base slightly oblique; largest leaves (often produced by the second, third or fourth sympodial units) cm long, cm wide, with petiole to 5.5 cm long, the leaves becoming reduced and more sessile distally; geminate leaf pairs unequal in size (the smaller 1/4 3/4 of the larger). LEAF PUBESCENCE similar throughout, the leaves pubescent to tomentose adaxially, tomentose abaxially, the trichomes spreading, mm long. PEDICELS AT ANTHESIS green or purple, (11.5) cm long, 1 2 times as long as subtending leaf, pubescent to tomentose, the trichomes spreading, mm long. CALYCES AT ANTHESIS green or purple, obconic mm long, mm diameter when

21 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 405 Figure 9. Herbarium specimen of Lycianthes dejecta (Dean & Starbuck 228 (DAV)). fresh, when pressed and dried, the teeth somewhat reflexed, emerging from the ribs at approximately the same level, of approximately equal length, (1) 2 7 (8) mm long, the entire calyx tomentose, the trichomes spreading, (0.5) mm long. COROL- LAS orientated laterally, not nodding, the tube mm long, the limb cm diameter, the membrane white to purple, the adaxial sides of the lobes matching the membrane in coloration but with darker maroon to purple veins, the abaxial sides of the lobes green to purple, pubescent to tomentose, the trichomes spreading, (0.5) mm long. ANDRO- ECIUM with green, glabrous filaments, these only slightly laterally compressed, straight to curved downward, the lowest (2) mm long, the two lateral (1.5) 2 5 mm, the two upper (1.25) mm, the length of the lowest (2) times that of the lateral; anthers exuding a sweet odour, 3 6 mm long, the lowest anther ovate to lanceolate (rarely narrowly elliptic), the pores round to widely ovate, generally terminal; pollen grains tricolporate. GYNOECIUM with conic to ovoid ovary, mm long, the ovules ; style straight to curved downward, (6) mm long; stigma purple, bilobed, 0.5 mm diameter FRUITS remaining attached to calyces at maturity, ovoid to conic, mm long, mm diameter at widest point, the tip apiculate to long-attenuate; exocarp light to dark green with purple or black lines (becoming more yellowish, then brown, upon drying), puberulent; mesocarp whitish to light green, soft and juicy, lacking sclerotic granules; placental area resembling the mesocarp; fruiting calyx enlarged, (5) 6 10 (11.5) mm long, (9) (23) mm diameter, the teeth reflexed to recurved, often broken, (3.5) 4 15 (19) mm

22 406 E. A. DEAN long; fruiting pedicel (3.5) 5 11 (12.5) mm long, deflexed, the fruit pendant, sometimes near the ground; seeds (40) (185) per fruit, dark brown to black, reniform to depressed-obovate, slightly appressed, without ridges but with microscopic fibrils, mm long, mm wide. Chromosome number: 2n = 24 (Dean & Starbuck 276, 309). Indigenous names: Trompeta, chichi de perra. Distribution: Mexico (Distrito Federal and states of Guanajuato, Hidalgo, Jalisco, Michoacán, Nuevo León, Puebla, Querétaro and San Luis Potosí). The current distribution of Lycianthes dejecta suggests that it once had a more continuous distribution on limestone soils that was broken up by the uplift of the transvolcanic belt. It now grows on limestone on either side of the transvolcanic belt, as well as in some very eroded, ancient agricultural areas within the transvolcanic belt. On both soil types, it is confined to cactus-scrub (matorral of Rzedowski, 1988) vegetation. It has been suggested that eroded volcanic areas within the Valley of Mexico are often home to calciphiles, because erosion has exposed a lower soil layer that is calcium rich (Rzedowski, 1986). Habitats include pastures, paths and the sides of agricultural fields, m. Diagnostic features: Lycianthes dejecta is easily recognized by its multi-angulate dendritic trichomes, which cover all parts of the plant. Its leaf shape is distinctive as well, with a relatively long attenuate portion and truncate lamina bases. Its fruits and seed type are similar to those of L. moziniana. It differs from that species in having maroon to black lines on its fruit, reflexed to curled calyx teeth, and microscopic fibrils on its seeds. This is the only species in the series with a bitter secondary compound present throughout the plant, even in the fruits. Specimens examined: MEXICO: Distrito Federal: Sierra de Guadalupe, 7000 ft, 21.vii.1938, Balls 5073 (BM, K, UC); Valley of Mexico, fields, near Guadalupe, Mt. Zacualco, 10.vii.1865, Bourgeau 543 (BR, F, G, M) (Syntype of Solanum dejectum, cited by Fernald but as Hahn 543); Actopan, on high plateau by city of Mexico, vi.1827, Karwinski s.n. (M); Valley of Mexico, c. San Cristóbal, 2800 m, 25.v.1951, Matuda (MEXU); no exact locality, Schmitz s.n. (W), Schmitz 482 (BM, W); Lomas de Jaral, 18.vii.1886, Schumann 998 (JE). STATE OF GUANAJUATO: Mpio. San Luis de la Paz, Mesas de Pueblo, along road NW of town, 2348 m, 4.vii.1991, Dean & Starbuck 224 (XAL); Mpio. San José Iturbide, near Rancho El Guajolote, SW of San José Iturbide, one hwy exit S of exit to San José, on W side of hwy 57, take dirt rd E and cross river, 1829 m, 16.ix.1991, Dean & Starbuck 253 (DAV, MEXU, NY, UC, XAL); Mpio. San José Iturbide, near Rancho El Guajolote, SW of San José Iturbide, one hwy exit S of exit to San José, dirt rd that goes W to large drainage, 1829 m, 31.x.1991, Dean & Starbuck 309 (DAV, XAL); hwy 51 near San Miguel de Allende, 2200 m, 7.vii.1971, Genelle & Fleming 922 (MO, WIS); cerca de San Miguel Allende, Río Laja, Atotonilco, 1850 m, 8.viii.1980, Kishler 923 (MEXU); San José Iturbide, cerca de el Guajolote, 2200 m, 22.viii.1988, Rzedowski (IEB); Rancho La Misión, 8 km al NE de San Luis de la Paz, 2900 m, 11.x.1988, Ventura & López 6129 (IEB); Mesas del Pueblo, 8 km al N de San Luis de la Paz, 17.vii.1989, Ventura & López 6871 (IEB). HIDALGO: Mpio. Tepealpulco, outskirts of Tepealpulco, base of Cerro Tres Peñas, near ruin of Jihuingo and rock quarry, 2530 m, 21.ix.1991, Dean & Starbuck 261 (DAV, MEXU, UC, XAL); 11 mi up mine rd W of Mex 85, 2 mi N of Posada del Rey, Zimapán, on ridge-like plateau, 5.vii.1966, Mears 254a (TEX); trail from Zimapán to mines of El Monte, N of Zimapán, ft, 11.viii.1948, Moore & Woods 4460 (A); prope Zimapán, vi , Schiede s.n. (HAL); Mpio. Tepeapulco, Cerro Tres Peñas, 2500 m, 9.ix.1975, Ventura-A. 248 (CAS, F, MEXU); Mpio. Tepealpulco, terrenos de Tepeapulco, 2600 m, 14.x.1975, Ventura-A. 402 (F, MEXU); Mpio. Zempoala, Tlaquilpan, 2500 m, 22.vi.1976, Ventura-A (F, MEXU). JALISCO: Matanzas, along road from Ojuelos de Jalisco, Rodríguez-C. s.n. (IBUG). MÉXICO: Mpio. Huehuetoca, N of Huehuetoca along the road to Apaxco, c. 4.2 road mi from building Los Arcos in downtown Huehuetoca, W side of rd, 2200 m, 1.vii.1990, Dean & Solano 202 (BM, DAV, MEXU, MO, NY, UC, XAL); Mpio. Huehuetoca, N of town of Huehuetoca along the rd to Apaxco, c. 4.2 rd mi from building Los Arcos (in downtown Huehuetoca), W side of rd, 2165 m, 3.viii.1991, Dean & Starbuck 243 (DAV, MEXU, UC, XAL); Mpio. Huehuetoca, N of Huehuetoca, along the rd to Apaxco, c. 4.2 rd mi from building Los Arcos in downtown Huehuetoca, W side of rd, 2165 m, 29.x.1991, Dean & Starbuck 304 (DAV, MEXU, XAL). MICHOACÁN: Morelia, Punguato, viii.1910, Arsène s.n. (F); Morelia, Punguato, 16.viii.1909, Arsène s.n. (F); Morelia, Punguato, 1950 m, 16.v.1909, Arsène s.n. (G); Punguato, near Morelia, 2000 m, 26.vi.1909, Arsène 2714 (GH, L, MEXU, MO); Punguato, near Morelia, 2200 m, 11.viii.1910, Arsène 5233 (CAS, GH, MO); vicinity of Morelia, Punguato, 2100 m, 20.vi.1912, Arsène 8300 (F, GH, MO, NY). NUEVO LEÓN: Mpio. Galeana, Cerro El Gallo, 2085 m, 15.vi.1991, Hinton et al (GH, TEX). PUEBLA: Mpio. Caltepec, between Cerro Pochote and Cerro Gavilán Chico in hills SE of town of Caltepec, along road to Atolotitlan, near small

23 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 407 valley called La Laguna, m, 15.vii.1991, Dean & Starbuck 228 (DAV, MEXU, XAL); Mpio. Caltepec, 0.85 km E of junction of Acatepec-Caltepec and Atecoxco-San Pedro Atzumba roads, along road to Caltepec, 2164 m, 29.ix.1991, Dean & Starbuck 274 (DAV, MEXU, UC, XAL); Mpio. Caltepec, hills SE of the town of Caltepec, between Cerro Gavilán Chico and Cerro Pochote, as well as an area called La Laguna, near rd to Atolotitlan, m, 30.ix.1991, Dean & Starbuck 276 (DAV, MEXU, XAL); Mpio. Caltepec, Barranca El Tocotín al E de Caltepec, 2020 m, 5.vii.1983, Tenorio-L (MEXU). QUERÉTARO: Cadereyta, 2000 m, 15.vi.1975, Arguelles 73 (MEXU); camino a Huimilpán, km 11, cerca de barranca, 2297 m, 27.vii.1984, Arguelles 2147 (MEXU); camino entre Huimilpán-Querétero y carr. Mexico-Los Cues, km 6, 2020 m, 29.vi.1986, Arguelles 2538 (MEXU); km 12 camino a Huimilpán, lado del camino, entrando un poco, lado izquierdo, 2100 m, 12.vii.1987, Arguelles 2985 (MEXU); Mpio. Huimilpán, 0.5 road km S of marker for km 11 along the road between city of Querétaro and Huimilpán, N side of road leading down to upper reaches of barranca, 2297 m, 2.vii.1990, Dean & Solano 203 (BM, DAV, F, MEXU, MO, NY, UC, XAL); Mpio. Huimilpán, along rd to Los Cues, 6 rd km E of intersection with Querétaro-Huimilpán road, just NE of 3 km marker, 2297 m, 2.vii.1990, Dean & Solano 204 (DAV, MEXU, NY, XAL); Mpio. Huimilpán, rd to Los Cues, 6 rd km NE of intersection with Querétaro-Huimilpán rd, just NE of 3 km marker, 2020 m, 16.ix.1991, Dean & Starbuck 255 (DAV, UC, XAL); Mpio. Huimilpán, rd to Los Cues, 6.0 rd km NE of intersection with Querétaro- Huimilpán rd., near 3 km marker, 2020 m, 29.x.1991, Dean & Starbuck 305 (DAV, MEXU, NY, UC, XAL); Mpio. Cadereyta, along rd between Cadereyta and Lizarro, by town of Charco Frío, 2226 m, 30.x.1991, Dean & Starbuck 307 (DAV, MEXU, MO, NY, UC, XAL). San Luis Potosí: Santa María del Río, 23.vi.1959, Rzedowski 3251 (ENCB, WIS). Lycianthes hintonii E. Dean sp. nov. (Fig. 10). Herba erecta ad 55 cm alta, inflorescentia floribus solitariis, serie Meizonodontae affinis. Habitus L. rzedowskii E. Dean similis, sed floribus albis antheris poris subterminalibus, pollinis granulis poris 3 autem differt. Habitat in montibus ad terram calcariam in Nova Hispania septentrionali (Nuevo León). Figure 10. Holotype of Lycianthes hintonii (Hinton et al (DAV)). Type: MEXICO: State of Nuevo Leon, Mpio. Aramberri, Cerro El Viejo, 1200 m, 28.vii.1993, Hinton et al (HOLOTYPE: DAV; isotype TEX). HERBS, usually erect, to c. 55 cm tall. INDUMENT of simple, acute, several-celled trichomes present throughout, these intermixed with a few branched trichomes, especially on upper stems. ROOTS not seen. Below-ground STEMS white; above-ground growth green with darker green vertical slashes, probably terete in cross-section, becoming somewhat woody with age, especially at base of plant, compressed when dry, especially upper branches, the stems pubescent, the trichomes (1) mm long, spreading to extrorse. FIRST SYMPODIAL UNIT erect above the soil surface, c. 25 cm long, c mm in diameter at widest point; internodes c. 13 in number; lateral branches developing at time of flowering. FIRST TWO SYMPODIAL BRANCHING POINTS dichasial followed by monochasial branching; first dichasial branching angle SUBSEQUENT SYMPODIAL GROWTH c. 30 cm in total length, the second sympodial units 6 13 cm long, 3 5 mm in diameter, subsequent sympodial units becoming reduced distally. LEAVES OF FIRST SYMPODIAL UNIT thin-chartaceous; leaves closest to the soil surface not seen; subsequent c. 7 leaves expanded, divided into expanded lamina and attenuate base, the attenuate portion plus petiole accounting for 25 50% of total leaf length, the lamina deltoid to ovate-elliptic, the margin

24 408 E. A. DEAN irregularly sinuate, the tip rounded to acute or shortacuminate, the base truncate to cuneate, attenuate into the petiole, the largest expanded leaves (at the top of the stem) c cm long, cm wide, the petiole poorly defined, 2 3 cm long. LEAVES OF SUBSE- QUENT SYMPODIAL UNITS geminate above the first two dichasial branching points, similar in shape and texture to the distal-most leaf of the first sympodial unit, but with shorter attenuate portion (1/3 of total leaf length); largest leaves (generally on the second sympodial units) c cm long, cm wide, the petiole poorly defined, to cm long, the leaves becoming reduced on more distal sympodial units; geminate leaf pairs unequal in size (the smaller 1/2 2/3 the size of the larger). LEAF PUBESCENCE similar throughout, the leaves sparsely pubescent with trichomes mm long, adaxially appressed, abaxially erect, those of the margin antrorsely orientated toward leaf tip. PEDICELS AT ANTHESIS green, 4 7 cm long, times as long as subtending leaf, pubescent with trichomes (1) mm long, spreading, antrorse or extrorse. CALYCES AT ANTHESIS green, campanulate 3 4 mm long, 4 5 mm in diameter when dry, the teeth spreading slightly, emerging from the ribs at approximately the same level, usually of approximately equal length, 4 11 mm long, the ribs and teeth pubescent with spreading hairs mm long. COROLLAS orientated laterally, the tube 3 mm long, the limb 2 4 cm in diameter, the membrane white (notched?), the adaxial sides of the lobes of similar colour, the abaxial sides of the lobes green, glabrous. ANDROECIUM with green, erect, glabrous, laterally compressed filaments, the lowest 3 6 mm long, the two lateral mm long, the two upper mm long, the length of the lowest times that of the laterals; the lowest anther ovate, KEY TO THE VARIETIES OF LYCIANTHES MOZINIANA mm long, the pores ovate, nearly terminal. GYNOECIUM with conic ovary 2.5 mm long, the ovules not seen; style slightly curved, mm long; stigma (green?), capitate, round to slightly bilobed. FRUITS AND SEEDS not seen. Distribution: Mexico (state of Nuevo León). Lycianthes hintonii has only been collected in oak forests on limestone soils in the mountains of Nuevo León in the vicinity of Cerro El Viejo, m, and may be endemic to the state. Diagnostic features: Although I have not seen this species in the field, it is obviously related to the species of series Meizonodontae and I assume it has the characteristic tuberous roots. The fruit type is unknown and could be either the green Lycianthes moziniana type or the dark purple L. ciliolata type. This species is similar to L. rzedowskii in its white flowers and anthers with terminal pores, but differs from that species in having fewer, larger leaves on the first sympodium, triangular, tricolporate pollen, and in growing in basic limestone soils. It is quite disjunct from the populations of L. rzedowskii, and I have no doubt that it is a separate species. Etymology: L. hintonii is named for the Hinton family, who have been collecting plants in Mexico for decades and discovered this species. Paratype: MEXICO: STATE OF NUEVO LEÓN: Mpio. Aramberri, Cerro El Viejo, 1495 m, 3.viii.1993, Hinton et al (DAV, TEX). Lycianthes moziniana (Dunal) Bitter ( mociniana ) 1a. Calyx teeth in flower lax, lying against the corolla; calyx teeth in fruit appressed to the fruit, not spreading; abaxial side of corolla lobes densely hairy; leaves of first sympodial unit cuneate at base, not attenuate; leaves sessile or petiole less than 5 mm long; plants mostly of agricultural areas of the transvolcanic belt...var. moziniana 1b. Calyx teeth in flower slightly spreading; calyx teeth in fruit spreading, not appressed to the fruit; abaxial side of the corolla lobes slightly hairy to nearly glabrous (glabrous in Nuevo Leon); leaves of first sympodial unit attenuate at base; petiole to 1.5 cm long; plants not of the region of the transvolcanic belt (San Luis Potosí, Nuevo Leon, Oaxaca) often growing on limestone soil...2 2a. Filaments glabrous; exocarp of fruit green; placental area green; plants found at high elevations in disturbed clearings and agricultural fields of mountains of eastern Oaxaca...var. oaxacana 2b. Filaments glabrous or pubescent; exocarp of fruit green or tan, often with purple blotches; placental area often purplish and powdery; plants of forested areas or more disturbed situations on limestone in San Luis Potosí and Nuevo León... var. margaretiana

25 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 409 Solanum uniflorum Sessé ex Lag., Gen. Sp. Pl Solanum monanthum Roemer & Schultes, Syst. Veg. 4: Solanum mocinianum Dunal forma luteiflorum Dunal, in De Candolle, Prodr. 13(1): Solanum uniflorum Moçiño & Sessé, Pl. Nov Hisp TYPE: Painting made during the Royal Botanical Expedition to New Spain ( ) under the direction of Martín de Sessé y Lacasta (LECTOTYPE: Hunt Institute for Botanical Documentation, catalogue number (Dean, 1997)). Figure 11. Herbarium specimen of Lycianthes moziniana var. moziniana (Dean & Starbuck 219 (DAV)). Lycianthes moziniana (Dunal) Bitter var. moziniana (Fig. 11). Solanum mozinianum Dunal, in Poiret, Encycl., Suppl. 3: Solanum mozinianum Dunal, Solan. Syn Solanum mocinianum Dunal, in De Candolle, Prodr. 13(1): Lycianthes mociniana (Dunal) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): 408, TYPE: Painting made during the Royal Botanical Expedition to New Spain ( ) under the direction of Martín de Sessé y Lacasta (LECTOTYPE: Hunt Institute for Botanical Documentation, catalogue number (Dean, 1997)). Perennial HERBS, decumbent, ascending or erect, usually recumbent with age, 5 50 cm tall. INDUMENT generally of simple trichomes (dendritic in some areas of Michoacán and Jalisco). ROOTS with narrowly fusiform segments, usually deeply buried (>30 cm below the soil surface). STEMS arising from the root one to several; below-ground growth extensive, often elongate, brittle, branched, whitish; above-ground growth green to purple-green, with very little woody tissue, often weak-stemmed (woody with age in Oaxaca), round in cross-section, compressed and ribbed when dried, usually noticeably pilose, the trichomes (0.1) mm long, these sometimes of two distinct lengths, the shorter mm and retrorsely appressed, the longer mm and slightly retrorse, rarely becoming glabrate with age. FIRST SYMPODIAL UNIT decumbent to erect above-ground, (2) 5 35 (40) cm long, mm in diameter at widest point; internodes (3) 6 14 (21) in number; lateral branching usually well developed early. FIRST SYMPODIAL BRANCHING POINT usually dichasial (with a branching angle of (120), followed by monochasial branching (rarely with a second dichasial branching point). SUBSEQUENT SYMPODIAL GROWTH extensive, often surpassing the first sympodial unit in length, extending laterally 3 30 (50) cm, the second sympodial units 1 7 cm long, mm in diameter, subsequent sympodia becoming reduced distally. ABOVE- GROUND LEAVES OF FIRST SYMPODIAL UNIT herbaceous, usually soft and pliable; leaves closest to the soil surface reduced, obovate, cm long, cm wide, often early deciduous; subsequent leaves expanded, obovate, elliptical, ovate or lanceolate, the margin sinuate to irregularly angled, the tip rounded to acute, the base usually obtuse to cuneate, the largest expanded leaves (in the upper 1/3 of the stem) cm long, cm wide, sessile or with petiole to 5 mm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate or not, similar in shape and texture to the distal-most leaf of the first sympodial unit, but tip usually acute and the lamina becoming more lanceolate distally, the largest leaves (often produced by the third or fourth sympodial unit) cm long, cm wide, sessile or with petiole to 5 mm long, becoming reduced and more sessile on more distal sympodial units; the geminate leaf pairs equal in size or the smaller (1/3) 1/2 3/4 the size of the larger. LEAF PUBESCENCE similar throughout, the leaves usually soft-pubescent to pilose, the trichomes (0.3) mm long, adaxially appressed, abaxially erect, the trichomes of the main abaxial veins spreading, the trichomes of the margin (0.75) mm long, usually appressed-antrorse (spreading in some Jalisco popula-

26 410 E. A. DEAN tions). PEDICELS AT ANTHESIS green, sometimes with purple, (3) 5 15 (18) cm long, times as long as subtending leaf, the trichomes usually of two distinct lengths, the smaller mm and appressedretrorse, the larger mm and mostly spreading (slightly retrorse), rarely only the longer trichomes present. CALYCES AT ANTHESIS green, sometimes flecked with purple, widely to narrowly campanulate, mm long, 4 7 mm in diameter when fresh, mm when pressed and dried, the teeth lax, not spreading or reflexed, emerging from the ribs at approximately the same level, usually of approximately equal length (rarely of two distinct lengths), 2 10 mm long, the ribs and teeth pilose, the trichomes spreading or slightly retrorse, mm long. COROLLAS orientated laterally, not nodding, the tube 2 mm long, the limb cm diameter, the membrane lilac to dark purple, the adaxial sides of lobes of similar colour but with darker purple veins, the abaxial sides of the lobes uniformly green to light purple, puberulent to pubescent, the trichomes appressed, mm long. ANDROECIUM with green, erect, glabrous, laterally compressed filaments, the lowest mm long, the two lateral mm, the two upper mm, the length of the lowest times that of the lateral; anthers exuding a sweet odour, mm long, the lowest anther elliptic to ovate, the pores obovate, terminal or lateral; pollen grains tricolporate. GYNOECIUM with conic ovary mm long, the ovules , the style straight to curved, mm long, the stigma yellow-green, round to slightly bilobed. FRUITS not separating from the calyx at maturity, round to ovoid, mm long, mm diameter, the tip rounded to apiculate; exocarp green, glabrous; mesocarp and placental area green, soft, juicy, lacking sclerotic granules; fruiting calyx much enlarged, 4 11 mm long, mm diameter, the teeth appressed to exocarp, often broken, mm long; fruiting pedicel cm long, deflexed, the fruits pendant or lying on the ground; seeds (10) per fruit, brownish-black, reniform to depressed-obovate, slightly compressed, smooth, without ridges or fibrils, mm long, mm wide. Chromosome number: 2n = 24 (Dean & Starbuck 300, 306). Indigenous names: Berenjena, tlanoxtle, tlanochtle, tlalnonochtle, shimpe, tintolón, tilindon, tilapó, huevo de gato, chumpin, tochin, la chichi, chochocuero, coyotomate, purga de las animas. Distribution: Mexico (Distrito Federal and states of: Hidalgo, Jalisco, México, Michoacán, Nayarit, Puebla, Querétaro, San Luis Potosí, Tlaxcala and Veracruz). Restricted to the volcanic soils of the transvolcanic belt with the possible exception of the occurrence in San Luis Potosí. Vegetation types range from disturbed oak, oak pine, pine oak and fir forest. Habitats include agricultural fields, roadsides, paths and pastures, rarely forest clearings, m. Diagnostic features: Lycianthes moziniana s.l. is separated from other species of series Meizonodontae by the following character combination: fruits with green to tan exocarp, small smooth seeds (without fibrils on cell walls), a tendency toward having only one dichasial or no dichasial branching points, very elongate pedicels, and usually abaxially pubescent corolla lobes. Variety moziniana is separated from the other varieties of the species in having fruiting calyx teeth that stay appressed to the fruit, cuneate (rather than attenuate) leaf bases, relatively dense and long stem and leaf pubescence, consistently pubescent abaxial corolla lobes, and an affinity for the soils of the transvolcanic belt. This weedy variety has had an intimate relationship with the people of Mexico and may owe its current distribution to humans, who are probably its primary dispersal agents. Limited crossing experiments indicate that var. moziniana is capable of crossing with var. oaxacana, although with reduced fertility; crosses between this variety and mature accessions of var. margaretiana have not yet been performed. Possible hybridization between var. moziniana and L. acapulcensis (in Jalisco) is discussed under the latter species. Specimens examined: MEXICO: State Unknown: 1833, Andrieux 60 (G); H. Buek s.n. (HBG); Lerma, 3.vii.1959, Davis s.n. (TEX); Herb. Jan (W); viii.1840, Galeotti 1185 (BR); Hab. in Mexico ad Guayconalpo, Hage s.n., 1906 (HBG); Karwinski s.n. (M); Cumbre de Estepe, 1842, Liebmann 1450 (C); Pavon s.n., actually Sessé and Mociño specimens (G); Schmitz s.n. (W); Schmitz s.n. (GH); Canada, Schmitz 484 (BM, F, NY, W), , Sessé & Moçiño s.n. (BM, photo WIS; according to M. Nee, specimen is annotated by Dunal); Sessé & Moçiño s.n. (BM) , Sessé, Moçiño, Castillo et Maldonado 1529 (photo NY of Madrid Specimen F negative 48228; another photo at NY of a duplicate specimen is L. ciliolata); , Sessé, Moçiño, Castillo et Maldonado 1530 (F); , Sessé, Moçiño, Castillo et Maldonado 5388 (F). Distrito Federal: Pedregal, 7000 ft, 30.vi.1938, Balls B5207 (BM, K, UC); cultivated fields in the hills of the Pedregal by Zapan, 26.vi.1865, Bourgeau 351 (BR, G, GH, L, S); foothills, SE Mexico City, 28.vi.1935, Clark 7352 (MO, NY); Cañada de Contreras, ix.1927, Lyonnet 177 (BM, GH, K, MEXU, MO, NY); Mpio. Huixquilucan, Fraccionamiento La Herradura, 8.vii.1968, Perusquía-O. s.n. (F); Santa Fe,

27 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 411 vii.1905, Rose et al (NY); lava beds near Eslava, 7500 ft, 19.vii.1910, Rusby 131 (NY); 2 km al N de la Cabrera, delegación de Cuajimalpa, 2500 m, 4.viii.1971, Rzedowski (F, IEB, MEXU); Valle de México, 1875, Schaffner 551 (GOET); Loma de la Era, delegación de Alvaro Obregón, 2650 m, 11.vi.1977, Ventura-A (ENCB, F, MEXU); Limbo, delegación de Alvaro Obregón, 2700 m, 4.viii.1979, Ventura-A (ENCB, F, MEXU); Mpio. Naucalpan, 6 km al E de San Francisco Chimalpa, 2550 m, 20.viii.1966, Rzedowski (CAS, ENCB, F, MEXU); STATE OF HIDALGO: Mpio. Acatlan, Los Reyes, c. 9 rd mi from Acatlan along rd to Huasca, NW of Tulancingo, 2195 m, 21.ix.1991, Dean & Starbuck 259 (DAV, NY, XAL); Agua Blanca de Iturbide, 2100 m, 9.vii.1972, Gimate-L. 690 (F, IEB); Mpio. Acatla, 9 km al N de Acatlán, hacia Huasca, 2300 m, 11.viii.1981, Hernández-M (MEXU); Zimapán, 25.viii.1938, Kenoyer 305 (F); Samariel to Las Lajas, viii.1905, Rose et al (GH, NY). JALISCO: Rancho La Joya Grande, 10 km al N de Zapotlanejo, 1600 m, 7.vii.1977, Carvajal & de Puga 641 (WIS); El huerto el Zarzamoro, Las Joyas, 1900 m, 15.vii.1986, Cuevas-R (WIS); Huerto el Zarzamoro, Las Joyas, Autlán, 1900 m, 16.vii.1986, Cuevas-R (WIS); Mpio. Tapalpa, Juanacatlan, 2561 m, 7.viii.1991, Dean et al. 246 (DAV, XAL); Mpio. Jocotepec, near town of Zapotitan de Hidalgo, cultivated field in valley at the base of Cerro Viejo, m, 11.viii.1991, Dean et al. 247 (DAV, XAL); Mpio. Mazamitla, Sierra del Tigre, c. 4.6 rd mi along rd to El Corral de la Mula, S of Mazamitla, rd leaves hwy 110 at Puerto Zapotería and winds into the mountains, m, 22.xi.1991, Dean & Starbuck 328 (DAV, MEXU, NY, XAL); Mpio. Jocotepec, Cerro Viejo, area opposite the Presa de Chayote, 2200 m, 24.xi.1991, Dean et al. 330 (UC, XAL); Mpio. Venustiano Carranza, along rd between Ciudad Guzmán and El Grullo, rd to microondas Las Víboras, near Puerto el Floripondo, 2317 m, 1.xii.1991, Dean et al. 333 (DAV, MEXU, NY, UC, XAL); Rancho el Isote, just W of Sayulapan, along rd between El Grullo and Ciudad Guzmán, along dirt rd that enters on S side of road, 2012 m, 3.xii.1991, Dean et al. 335 (DAV, MEXU, NY, UC, XAL); Mpio. El Mirador, Cocula (Colula), 1800 m, 16.vii.1976, Delgado-S. 262 (MEXU); Mpio. Volcanes, La Campana, 1800 m, 29.viii.1968, Díaz-L (F); steep S-facing slope above Salto of Río Tapalpa, 2 km NW of Tapalpa (c N; W), m, 7.viii.1960, Iltis et al. 797 (WIS); Tepotitlán, 5 mi E. of Roadside, 18.vii.1960, Knoblock 1652 (MEXU); Mpio. Jocotepec, Cerro Viejo, 2380 m, 19.vii.1986, Machuca-N. 2781a (WIS); Sierra del Tigre, 3 mi S of Mazamitla, m, 19.ix.1952, McVaugh & Sooby (BM, MEXU, NY, TEX); N slopes of the Nevada de Colima, W of summit of the N ridge, near junction of the old pack road to Zapotlán with Atenqique-Jazmín rd, 2100 m, 20.vii.1957, McVaugh et al (MEXU); Sierra del Halo, near a lumber road leaving the Colima hwy 7 mi SSW of Tecalitlán and extending SE toward San Isidro, 2 mi from hwy., 1400 m, 13.viii.1957, McVaugh et al (MEXU, NY, TEX; a duplicate specimen at TEX is L. acapulcensis); Río Blanco, vii.1886, Palmer 186 (BM, G, GH, MEXU, MO, NY, WU); banks of ravines near Guadalajara, 5000 ft, 14.vii.1902, Pringle (GH, K, L, MEXU); Mpio. Venustiano Carranza, Sayulapa, faldas del Nevado de Colima, 2100 m, 22.viii.1987, Rodríguez-C. & Suárez-J. 941 (WIS); Carr. Mazamitla-Tamazula de Gordiano, cerca del Rancho El Ternero, 2100 m, 21.vii.1988, Rodríguez-C. & Reynoso-D (IEB, WIS); trail between Chante and Manantlán about 15 mi SSE of Autlán, 4500 ft, 1.viii.1949, Wilber & Wilber 2081 (DS, MEXU, WIS). MÉXICO: Near Toluca, iv.1834, Andrieux 196 (G-DC n.v., photos: F, GH, NY, WIS); San Cayetano, c. 1.5 rd mi from old hwy 15 along a gravel rd that begins c. 3.4 rd mi E of turnoff to Villa de Allende, NW of Valle de Bravo, 2515 m, 11.xi.1991, Dean & Starbuck 319 (DAV); Amecameca, 26.vii.1924, Fisher, s.n. (F); Mpio. Texcoco, lado sur de la Cañada de Aguas, 13.5 km al SE de Texcoco (11 km al ESE de Coatlinchán), 2740 m, 29.vii.1977, García-P. 343 (CAS, MEXU); Mpio. Tepetlaoxtoc, 18 km por carr. Texcoco-Calpulalpan, limite entre Edo. México y Tlaxcala, 2870 m, 3.ix.1982, García-P (MEXU); District Temascaltepec, Comunidad, 2610 m, 7.vii.1932, Hinton et al. 972 (BM, G, K); 30 mi E of Mexico City on the road to Puebla, 8500 ft, 9.vii.1940, Hitchcock & Stanford 7013 (DS, F, GH, UC); Mpio. Jilotepec, Jilotepec, 2300 m, 21.ix.1953, Matuda (MEXU); Cumbre de Acambay, 3000 m, 9.viii.1953, E. Matuda (MEXU); entre El Oro y Villa Victoria, 2000 m, vi.1954, E. Matuda (MEXU); San Cayetano, Villa Allende, 2500 m, 3.vii.1954, Matuda (MEXU); Mpio. Villa de Allende, San Cayetano (Estación Experimental) al N de Agua Escondida y a 26 km al NE de Valle de Bravo, 2490 m, 7.vii.1974, Maury-H et al. s.n. (MEXU); along old hwy 190 between turnoff to Chalco (hwy 115) and Santa Barbara, c. 30 m above Azotla, 21.vii.1964, Mick & Roe 300 (WIS); near Toluca, 23.vi.1887, Pringle 2926 (GH); near Tultenango, 13.vii.1901, Rose & Hay 5413 (NY); 22.viii-19.ix.1930, Rusby & Souviron 114 (CAS); vertiente N del Cerro Sacromonte, cerca de Amecameca, 2500 m, 24.vii.1966, Rzedowski (ENCB, F); Sierra de Alcaparrosa, 5 km al NW de Tepozotlán, 2600 m, 29.vii.1971, Rzedowski (ENCB, F, MEXU); 22 km al NE de Texcoco, sobre la carretera a Calpulalpan, 2900 m, 17.viii.1971, Rzedowski (ENCB, F); Cerro del Tigre, al NW de Atizapán, 2500 m, 4.viii.1974, Rzedowski (ENCB, F); Mpio. Tepotzotlán, alrededores de la Hacienda Lan-

28 412 E. A. DEAN zarote, 2350 m, 31.vii.1977, Rzedowski (ENCB, F, MEXU); parte alta de la Sierra de Alcaparrosa, 10 km al NW de Tepotzotlán, 2800 m, 7.viii.1979, Rzedowski (ENCB, F); Ixtapaluca, km 42 antigua carretera México-Puebla, 2650 m, 6.ix.1970, Segura- R. 117 (CAS, F, IEB); Mpio. Texcoco, San Pablo Ixayoc, 2600 m, 10.ix.1983, Ventura-V.1375 (IEB, MEXU). MICHOACÁN: near Capácuaro, 2150 m, 7.viii.1965, Correll et al (LL); Mpio. Villa Escalante/Pátzcuaro, along Pátzcuaro-Villa Escalante rd, W side of rd, across from the microondas, west of Cerro El Tecolote, 2360 m, 22.vii.1990, Dean & Starbuck 216 (DAV, MEXU); Mpio. Villa Escalante, meadows and roadside SW Lagunita de San Gregorio, south of Cerro El Barro, 2360 m, 22.vii.1990, Dean & Starbuck 217 (DAV, MEXU, XAL); Mpio. Villa Escalante, along rd to San Gregorio, c. 4.7 rd km E of intersection with Pátzcuaro-Villa Escalante rd., N side of road, 2470 m, 22.vii.1990, Dean & Starbuck 218 (DAV, MEXU); Mpio. Pátzcuaro, E of Pátzcuaro, along hwy 190, upslope (S and SSE) of Las Trojes, 2134 m, 23.vii.1990, Dean & Starbuck 219 (DAV, MEXU); Ejido Cruz de Caminos, along old hwy 15, W of Ciudad Hidalgo, 2195 m, 12.xi.1991, Dean & Starbuck 321 (DAV, MEXU, UC, XAL); Mpio. Villa Escalante/Pátzcuaro, Las Joyas de las Navas, along Pátzcuaro-Villa Escalante rd, about 3.2 rd mi N of turnoff to Tucambaro, 2348 m, 18.xi.1991, Dean et al. 324 (DAV, XAL); Mpio. Villa Escalante, SW of Laguna de San Gregorio, along rd leading to radio tower, 2439 m, 18.xi.1991, Dean et al. 325 DAV, NY (XAL); Mpio. Pátzcuaro, Las Trojes, E of Pátzcuaro along hwy 120, slopes SSE of town, 2134 m, 18.xi.1991, Dean & Starbuck 326 (DAV, MEXU, XAL); Mpio. Zinapécuaro, km de la carretera Toluca-Morelia, 2300 m, 21.vi.1988, González- M. & Díaz (MEXU); Cerro de la Campana, unos 5 6 km al norte de Puruandiro, 19.ix.1959, Hernández-M. 32 (MEXU); Sierra Torrecillas, District Coalcoman, 2350 m, 25.vii.1939, Hinton et al (F, GH, LL, MO, NY, TEX; specimen at W is L. acapulcensis); along hwy 15, 3 km ESE of Comanja, 16 km ESE of Zacapu, c N, W, 2300 m, 26.vii.1960, Iltis et al. 442 (WIS); Mpio. Uruapan, Tancítaro region, west of Uruapan on trail to Tancítaro, 6000 ft, 17.vii.1941, Leavenworth & Hoogstraal 1008 (F); 23 km E of Morelia, km 290 from México City, just E of Ciudad Hidalgo, along Mex. hwy 15, 2600 m, 28.vi.1964, Mick & Roe 162 (WIS); salida poniente de San Isidro, 2650 m, 4.vi.1981, Motte 289 (MEXU); Santa Clara del Cobre, 2200 m, 16.vii.1988, Pérez-C. 109 (IEB, TEX); Mpio. Santa Clara del Cobre, laguna cerca de San Gregorio, 2700 m, 7.vii.1985, Rzedowski (MEXU, TEX); 5 mi W of Cd. Hidalgo, 7.vii.1947, Sauer 1109 (UC); SSW of Morelia on the road to Villa Madero (just NE of Tiripetio), km 21, 2000 m, 19.vii.1963, Ugent & Flores-C (WIS); Mt. Punguato, Morelia, 2120 m, 4.viii.1965, Ugent et al (WIS); at km 19, S. of Carapan on road to Uruapan (c N, W), 2300 m, 27.vii.1960, Iltis & Koeppen 483 (WIS). NAYARIT: Sierra Madre, territory of Tepic, between Sta. Gertrudis and Sta. Teresa, 8.viii.1897, Rose 2068 (GH). PUEBLA: Mpio. Chignahuapan, N of town of Chignahuapan, E of small town of San Joaquín, both E and W sides of Barranca El Salto, 2225 m, 22.ix.1991, Dean & Starbuck 264 (DAV, NY, XAL); Mpio. Chignahuapan, N of Chignahuapan, near town of San Joaquín, both sides of Barranca El Salto, 2225 m, 27.x.1991, Dean & Starbuck 300 (DAV, MEXU, NY, UC, XAL); Barranca de Ocoxicuaya, Chignahuapan, 2300 m, 14.vii.1975, Hernández-M (CAS, MEXU, US); Amazoc, viii.1946, Martínez (MO). QUERÉTARO: camino entre Amealco y Chiteje de la Cruz, lado del camino a unos 4 km de caminata, 2850 m, 9.vii.1978, Arguelles 1127 (MEXU); Mpio. Amealco, W of Amealco, along rd to Chiteje de la Cruz which turns off hwy 120, c. 0.3 rd mi from junction with hwy 120, E side of road, 2530 m, 5.viii.1991, Dean & Starbuck 245 (DAV, XAL); Mpio. Amealco, W of Amealco, along rd to Chiteje de la Cruz which turns off hwy 120, c. 0.3 rd mi from junction with hwy 120, E side of road, 2530 m, 18.ix.1991, Dean & Starbuck 256 (DAV, XAL); Mpio. Amealco, W of Amealco, along rd to Chiteje de la Cruz which turns off hwy 120, c. 0.3 rd mi from junction with hwy 120, E side of road, 2530 m, 30.x.1991, Dean & Starbuck 306 (DAV, XAL). SAN LUIS POTOSÍ: It is doubtful that the following specimens were collected in San Luis Potosí. They are probably from a trip that Palmer took between San Luis Potosí and Mexico City: near San Luis Potosí, ft, iv.1879, Parry and Palmer 662 (GH, K, MO; K collection mixed with Lycianthes moziniana var. margaretiana); chiefly in the region of San Luis Potosí, ft, 1878, Parry and Palmer 662 (NY). TLAXCALA: Mpio. Españita, along rd to Españita, c rd mi from intersection with hwy 136, W side of rd near large turnout, 2720, 22.ix.1991, Dean & Starbuck 262 (DAV, NY, XAL); Mpio. Hueyotlipan, E side of town of Hueyotlipan, down dirt rd and across a stream, 2591 m, 22.ix.1991, Dean & Starbuck 263 (DAV, XAL); Mpio. Tlaxco, N side of town of Acopinalco del Peñon, along old rd to Chignahuapan, 2652 m, 27.x.1991, Dean & Starbuck 301 (UC, XAL); Mpio. Españita, 0.95 rd mi from intersection with hwy 136, along road to Españita, 2720 m, 28.x.1991, Dean & Starbuck 302 (DAV); Tlaxcala, ft, 19.vi.1938, Balls B4838 (K); Mpio. Ixtacuixtla, San Juan Nepopoalco, 2500 m, 10.ix.1982, Williams 1 (CAS); Mpio. Calpulalpan, 4 km al S de Calpulalpan, sobre el camino rural a Nanacamilpa, 2650 m, 12.viii.1983, Williams 87 (F, TEX); Mpio. Nanacamilpa, 1 km al S de Nanacamilpa, sobre el camino rural a San Martín Texmelucan,

29 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) m, 12.viii.1983, Williams 90 (CAS); Mpio. Hueyotlipan, terrenos de la hacienda Sta. Cruz Tenancingo, 7 km al S de Hueyotlipan, 2600 m, 12.viii.1983, Williams 96 (WIS); Mpio. Panotla, San Tadeo Huiloapan, 5 km por vereda al N de San Tadeo, por la Barranca del Río Temeyaya (cerca de Barranca Honda), 2480 m, 7.ix.1983, Williams 151 (CAS); Mpio. Tlaxco, Acopinalco del Peñon, 1 km al N del pueblo, camino a Chignahuapan, 2660 m, 21.ix.1983, Williams 201 (CAS, MEXU); Mpio. Panotla, San Tadeo Huilapan, 5 km al N de Huiloapan por vereda, Barranca del Río Temeyeya, 2480 m, 4.x.1983, Williams 449 (CAS, F, MEXU, TEX, WIS). VERACRUZ: Mpio. Acultzingo, high mountains near border of Veracruz and Puebla, El Sumidero, S of town of Puerto del Aire which is along hwy 150 between Orizaba and Tehuacán, 2439, 27.ix.1991, Dean & Starbuck 270 (DAV, NY, XAL); Orizaba, viii.1853, Müller 1117 (BR, NY); Orizaba, 1855, Müller (NY); Orizaba, 1853, Müller 1617 (L); Mt. Orizaba, 1891, Seaton s.n. (GH); Mt. Orizaba, Esperanza, 8000 ft, 15.viii.1891, Seaton 373 (GH, NY). L. moziniana var. margaretiana E. Dean var. nov. (Fig. 12). Habitus var. oaxacana similis corollarum lobis glabris vel puberulis autem, filamentis antherarum glabris vel puberula, et fructibus ex purpureo viridibus. Habitat ad terram calcariam in montibus Sierra Madre Oriental septentrionali (San Luis Potosí, Nuevo León). Type: MEXICO: State of Nuevo León: Mpio. Zaragoza, Cerro El Viejo, 2085 m, 17.vi.1993, Hinton et al (HOLOTYPE: DAV; ISOTYPES: TEX, XAL). Perennial HERBS, usually erect, (60) cm tall. INDUMENT of simple trichomes. ROOTS with fusiform segments, usually present within 20 cm of the soil surface. STEMS generally arising from the roots singly; below-ground growth not extensive, white; aboveground growth green, sometimes with purple (especially near the soil level), with very little woody tissue except near the base, round in cross-section, compressed and ribbed when dried, pubescent, the trichomes of two lengths, the smaller more common, mm long, the longer interspersed, mm long. FIRST SYMPODIAL UNIT erect above-ground, 9 30 cm long, 2 4 mm diameter; internodes 6 14 in number; lateral branches from leaf axils few to many. FIRST SYMPODIAL BRANCHING POINT dichasial (with a branching angle of (70) ), followed by monochasial branching. SUBSEQUENT SYMPODIAL GROWTH 3 20 cm in total length, the second sympodial unit 1 12 cm long, 1 3 mm in diameter, subsequent sympodia becoming reduced distally. ABOVE-GROUND Figure 12. Holotype of Lycianthes moziniana var. margaretiana (Hinton et al (DAV)). LEAVES OF FIRST SYMPODIAL UNIT chartaceous; leaves closest to the soil surface reduced, ovate to obovate, cm long, cm wide, often early deciduous; subsequent leaves expanded, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for up 1/3 2/3 of the total leaf length, the lamina ovate, elliptical or obovate, the margin irregularly sinuate, the tip rounded to acute, the base attenuate into the petiole, the largest expanded leaves (usually the distal-most leaf) (9) cm long, cm wide, the petiole to 1.5 cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate, similar in shape and texture to the distalmost leaf of the first sympodial unit, but the lamina ovate to elliptical, the tip acute to short-acuminate, and the base cuneate, shortly attenuate into the petiole; largest leaves (generally produced by the second to fifth sympodial unit) cm long, cm wide, the petiole cm long; geminate leaf pairs equal in shape and size or the smaller leaf 1/3 3/4 the size of the larger. LEAF PUBESCENCE similar throughout, the leaves sparsely pubescent with trichomes mm long, adaxially appressed to erect (often orientated

30 414 E. A. DEAN toward the leaf margin), abaxially erect, those of the margin spreading. PEDICELS AT ANTHESIS green to purple, 3 12 (19) cm long, 1 3 (5) times as long as subtending leaf, pubescent, the smaller trichomes retrorsely appressed, (0.05) mm long, the longer spreading, mm long. CALYCES AT ANTHESIS green to purple, campanulate, 3 5 mm long, 4 6 mm in diameter when dry, the teeth spreading slightly, emerging from the ribs at approximately the same level, usually of approximately equal length, 3 7 (12.5) mm long, the ribs and teeth pubescent, the trichomes spreading, mm long. COROLLAS orientated laterally, not nodding, the tube 2 mm long, the limb (2) cm in diameter, the membrane lilac, the adaxial sides of the lobes of similar colour but with darker maroon-purple veins, the abaxial sides of the lobes uniformly green, glabrous to sparsely pubescent, the trichomes erect, (0.5) mm long. ANDRO- ECIUM with green, erect, laterally compressed filaments, these glabrous or puberulent, the lowest mm long, the two lateral mm long, the two upper 2 3 mm long, the length of the lowest times that of the laterals; anthers ovate, 4 5 (6) mm long, the pores ovate, terminal; pollen grains tricolporate, of somewhat larger size than the grains found in the other two varieties. GYNOECIUM with conic ovary mm long, the ovules c in number, the style straight to slightly curved, c mm long, the stigma capitate, round to slightly bilobed. FRUITS not separating from the calyx at maturity, ovoid, c mm long, mm diameter, the tip apiculate; exocarp green, often with white or purple blotches, or becoming tan with maturity, glabrous; mesocarp green, soft and juicy, lacking sclerotic granules; placental area light green, whitish or purple, variable in texture, sometimes juicy, sometimes slightly powdery as in L. ciliolata; fruiting calyx enlarged, c. 8 9 mm long, mm diameter, the teeth spreading slightly, not lax, often broken, mm long; fruiting pedicel c cm long, often looped or curved, deflexed; seeds c per fruit, similar in appearance and size to those of var. moziniana. Distribution: Mexico (states of Nuevo León and San Luis Potosí). Restricted to the limestone soils of the northern Sierra Madre Oriental. Vegetation types include oak, oak pine, and pine forest. Unlike var. moziniana and var. oaxacana, this variety is not typically found in anthropogenically disturbed habitats, although it may be found in naturally disturbed situations such as drainages, m. Diagnostic features: Lycianthes moziniana var. margaretiana is closely related to var. oaxacana based on DNA sequence data, and it shares the attenuate leaf bases and spreading calyx teeth (in fruit) of that variety. However, its distribution is disjunct from var. oaxacana and presumably the two varieties have been separated for quite some time. Variety margaretiana has several characteristics that differ from the other two varieties: (1) upon maturity, the fruit exocarp can have tan or purple patches of colour; (2) the placental area of the fruits may be purple, sometimes even of a powdery texture similar to the fruits of L. ciliolata; (3) the pollen is somewhat larger than that of the other two varieties; (4) the stamen filaments can sometimes be pubescent; (5) the abaxial sides of the corolla lobes can sometimes be glabrous; and (6) this variety does not appear to be a weed of agricultural situations. Etymology: Lycianthes moziniana var. margaretiana is named for my daughter Margaret Starbuck, who helped delay this publication and the naming of this variety for nearly a decade. Paratypes: MEXICO: NUEVO LEÓN: Summit of Mt. Infernillo, 9.5 miles south of Pablillo and 9.2 miles south of Mex. hwy 60, 9000 ft, 17.vi.1963, Bell and Rice (UC); Mpio. Galeana, E of the town of Pablillo, San Francisco Canyon, 4.ix.1993, Dean et al. 360 (DAV, XAL, ANSM); Mpio. Galeana, above Agua Blanca, 2305 m, 4.vii.1991, Hinton et al (TEX); Mpio. Zaragoza, Cerro El Viejo, 2485 m, 29.vii.1992, Hinton et al (TEX); Mpio. Aramberri, W of La Escondida, 2200 m, 3.viii.1993, Hinton et al (TEX); San Francisco Canyon, about 15 mi SW of Pueblo Galeana, ft, 16.v.1934, Mueller & Mueller 404 (GH); Lower San Francisco Canyon, about 15 mi SW of Galeana, ft, 12.vi.1934, Mueller & Mueller 778 (GH); Sierra Madre Oriental, descent to Banco de Santa Ana, about 15 mi SW of Galeana, in woods below Puerta de Santa Ana, 17.vi.1934, Mueller & Mueller 855 (F, GH, TEX). SAN LUIS POTOSÍ: Hwy 86, 25 mi from Juárez Circle, beyond Xoconostle, 9000 ft, 5.vii.1971, Andreasen et al. 544 (MO); 96.2 mi SE of Huizache along hwy mi NW of El Salto ft 25.vii.1967, Clarke et al. s.n. (ASU); mi E of Ciudad del Maiz, ft, 23.vii.1953, Manning & Manning (GH), (GH, TEX); Alvarez, 5 10.ix.1902, Palmer 138 (NY); chiefly in the region of San Luis Potosi, ft, 1878, Parry & Palmer 662 (K, mixed collection with Lycianthes moziniana var. moziniana); Santa María del Río, 23.vi.1959, Rzedowski 3251 (ENCB, WIS); km 40 carretera San Luis Potosí-Rioverde, 2500 m, 20.viii.1955, Rzedowski 6228 (WIS); Sierra de Alvarez, cerca de Puerto Huerta, 2300 m, 10.vii.1956, Rzedowski 7730 (WIS); Cerro de San Pedro, Cerros al SE de Jesús María 2300 m, 28.vii.1956, Rzedowski 7876 (WIS); Mpio. Tierra Nueva, entre Joyita y Paso de Ordena, 1700 m, 9.vi.1966, Rzedowski 10727

31 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 415 (ENCB); Mpio. Zaragoza, Sierra de Alvarez, cerca de Puerto de la Huerta, 2300 m, Rzedowski (ENCB). Lycianthes moziniana var. oaxacana E. Dean var. nov. (Fig. 13). Differt a var. moziniana pubescentia brevi, foliis basi attenuatis, dentibus calyci fructificantibus patentibus. Habitat in montibus Sierra Madre Oriental Oaxacanis. Type: MEXICO: State of Oaxaca: Mpio. Santa María Jaltianguis, along hwy 175, c rd mi N of Ixtlán de Juárez, N of turnoff to Sta. María Jaltianguis, W side of road, downslope along footpaths, 2439 m, 11.x.1991, Dean & Starbuck 285 (HOLOTYPE: DAV; ISOTYPES: NY, XAL). Perennial HERBS, usually erect, 7 40 cm tall. INDU- MENT of simple trichomes. ROOTS with fusiform segments, the uppermost within 20 cm of the soil surface. STEMS generally arising from the roots singly; below Figure 13. Holotype of Lycianthes moziniana var. oaxacana (Dean & Starbuck 285 (DAV)). ground growth not extensive, white; above-ground growth yellow-green, purple near the soil surface, woody with age (only the younger stems compressed when dried), pubescent, the trichomes of two lengths, the smaller retrorsely appressed, mm long, the longer spreading, mm long, some populations lacking the longer hairs. FIRST SYMPODIAL UNIT erect above-ground, cm long, mm diameter; internodes 3 8 in number; lateral branches from leaf axils few to many. FIRST SYMPODIAL BRANCHING POINT dichasial (with a branching angle of ) or monochasial, followed by monochasial branching. SUBSEQUENT SYMPODIAL GROWTH 5 15 cm in total length, the second sympodial unit cm long, mm diameter, subsequent sympodial units becoming reduced distally. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT chartaceous, the leaves closest to the soil surface reduced, obovate, cm long, cm wide, often early deciduous; subsequent leaves expanded, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for up to 1/6 1/3 of the total leaf length, the lamina ovate to obovate, the margin irregularly sinuate, the tip widely acute to somewhat acuminate, the base cuneate to shortly attenuate into the petiole, the largest expanded leaves (in the upper 1/3 of the stem) cm long, cm wide, the petiole to 1.5 cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate or not, similar in shape and texture to the distalmost leaf of the first sympodial unit, but the lamina ovate, the tip acute to short-acuminate, and the base obtuse to cuneate, shortly attenuate into the petiole; largest leaves (generally produced on the third or fourth sympodial unit) 3 7 cm long, cm wide, the leaves sessile or with petiole to 0.5 cm long; geminate leaf pairs nearly equal in shape and size or the smaller reduced to 3/4 the size of the larger. LEAF PUBESCENCE similar throughout, the leaves sparsely pubescent, the trichomes mm long, adaxially appressed to erect (often orientated toward the leaf margin), abaxially erect, those of the margin antrorse to spreading. PEDICELS AT ANTHESIS green to purple, 5 14 cm long, 2 6 times as long as subtending leaf, pubescent, the smaller trichomes retrorsely appressed, mm long, these interspersed with longer spreading trichomes, mm long. CALYCES AT ANTHESIS green to purple, campanulate 4 5 mm long, mm diameter when fresh, similar when dry, the teeth spreading slightly, emerging from the ribs at approximately the same level, usually of approximately equal length, 2 10 mm long, the ribs and teeth pubescent, the trichomes spreading, mm long. COROLLAS orientated laterally, not nodding, the tube 2 mm long, the limb cm diameter, the membrane lilac, the adaxial sides of the lobes of similar colour but with darker purple veins, the

32 416 E. A. DEAN abaxial sides of the lobes uniformly green, pubescent, the trichomes appressed to erect, mm long. ANDROECIUM with green, erect, glabrous, laterally compressed filaments, the lowest mm long, the two lateral mm long, the two upper mm long, the length of the lowest times that of the laterals; anthers ovate, mm long, the pores ovate, terminal; pollen grains tricolporate and of similar size to var. moziniana. GYNOECIUM with conic ovary c. 3 mm long, the ovules c in number, the style straight to slightly curved, 8 12 mm long, the stigma pale green to whitish, round. Fruits not separating from the calyx at maturity, ovoid, c mm long, mm diameter, the tip apiculate; exocarp green, glabrous; mesocarp and placental area green, soft, juicy, lacking sclerotic granules; fruiting calyx enlarged, c mm long, mm diameter, the teeth spreading slightly, not lax, often broken, 7 10 mm long; fruiting pedicel c. 5 9 mm long, deflexed; seeds c per fruit, similar in appearance and size to those of var. moziniana. Indigenous name: Chichi de venado. Distribution: Mexico (state of Oaxaca). Restricted to the southern Sierra Madre Oriental in the high mountains of Sierra de Juárez of Oaxaca. Vegetation types include oak, oak pine, and pine forest. This variety is typically found in anthropogenically disturbed habitats such as roadsides, pastures, old fields and corn fields, m. Diagnostic features: Lycianthes moziniana var. oaxacana is closely related to var. margaretiana based on DNA sequence data, and it shares the attenuate leaf bases and spreading calyx teeth (in fruit) of that variety. However, unlike var. margaretiana, this variety is more similar to var. moziniana; this variety does not exhibit any of the more specialized fruit and androecium characters that are sometimes found in var. margaretiana. Paratypes: MEXICO: STATE OF OAXACA: Llano de las flores, on the Oaxaca-Valle Nacional Highway, 20 km E of Ixtlán, 2870 m, 22.vii.1960, Beaman 3703 (GH, LL); Distrito de Ixtlán, las Animas, carretera Oaxaca a Ixtlán de Juárez, 2340 m, 29.vii.1981, Cedillo-T et al. 872 (F, MEXU); vi.1901, Conzatti & González 1220 (GH); km from route 190 jct. on rd to Díaz Ordaz, 31.vii.1977, Davis 801 (MEXU); Field site same as 285 (Holotype), grown from field collected seed in the greenhouse at UC Berkeley, harvested 31.x.1993, Dean & Starbuck 285a (DAV, MEXU, UC, XAL); Mpio. Sta. María Jaltianguis, Sta. María Jaltianguis, 2195 m, 20.x.1991, Dean & Starbuck 296 (DAV, NY, UC, XAL); E-facing mountains along Route 175, 12 km north of Ixtlán de Juárez on the road to Valle Nacional, 2500 m, 26.vii.1959, King 2011 (MEXU, TEX, WIS); Cerro Verde, vii.1908, Purpus 3611 (UC); Salle s.n. (BM); District Etla, S. Juan del Estado, 18.vi.1888, Seler 812 (GH); Mts. San Juan del Estado, 25.vi.1895, Smith 401 (GH); Mpio. Coixtlahuaca, Cerro Verde al NE de Marcos Pérez, N, W, 2700 m, 7.vii.1986, Tenorio-L et al (MEXU). Lycianthes peduncularis (Schltdl.) Bitter (Fig. 14). Solanum pedunculare Schltdl., Linnaea 19: Lycianthes peduncularis (Schltdl.) Bitter, Abh. Naturwiss. Vereine Bremen 24(2): TYPE: GER- MANY. Leipzig Botanical Garden, 1842, Kunze s.n. (NEOTYPE, W!, here designated). Chosen from syntypes Schlechtendal s.n. (grown at the Halle Botanic Garden, presumably destroyed in World War II), C. Ehrenberg 81 (n.v., B, presumably destroyed), Schiede s.n. (n.v., B, presumably destroyed) (see discussion below after specimens examined section). Figure 14. Herbarium specimen of Lycianthes peduncularis (Dean & Starbuck 226 (DAV)).

33 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 417 Perennial HERBS, prostrate to decumbent, to 10 cm tall and 50 cm in diameter. INDUMENT generally of simple, curved trichomes, rarely with a few dendritic trichomes mixed with the simple ones. ROOTS with fusiform segments, usually deeply buried >20 cm below the soil surface. STEMS arising from the root one to several; below-ground growth often extensive, elongated, brittle, sometimes highly branched, white; above-ground growth with green and purple striations, slightly angled in cross-section, without much woody tissue, much compressed and ribbed when dry, pilose to villous, the trichomes antrorsely appressed, (1.25) mm long. FIRST SYMPODIAL UNIT mostly below ground, extending above ground 0 5 (7) cm, 2 4 mm diameter; above-ground internodes 4 6 (10) in number. FIRST SYMPODIAL BRANCHING POINT dichasial, followed by a mixture of monochasial and dichasial branching points; first dichasial branching angle SUBSEQUENT SYMPODIAL GROWTH extensive, resting on the soil surface, extending 2 35 cm, the second sympodial units cm long, mm diameter, often followed by units of greater length, with subsequent sympodial units becoming reduced distally. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT thick-chartaceous, often early deciduous; leaves closest to the soil surface very reduced, to 0.5 cm long, 0.2 cm wide; subsequent leaves larger, the lamina spathulate, the margin entire to slightly undulate, the tip rounded, the base cuneate, then short-attenuate, the largest expanded leaves (usually at most distal point) cm long, cm wide, leaves sessile or with petiole to 1 cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate or not, due to the mixture of monochasial and dichasial branching, the leaves similar in shape and texture to distalmost leaves of the first sympodial unit but with more expanded lamina and more attenuate base, the attenuate portion plus petiole accounting for 1/8 2/3 of total leaf length, the lamina spathulate, obovate, elliptic or ovate, the margin undulate, the tip rounded to short-acuminate, and the base obtuse to cuneate, short to long-attenuate; largest leaves (generally borne on the third or fourth sympodia) (1) (14) cm long, (0.5) (7) wide, the leaves sessile or with a petiole to 2 cm long, the leaves becoming reduced and more sessile distally; geminate leaf pairs unequal in size (the smaller 1/8 3/4 of the larger). LEAF PUBESCENCE similar throughout, the leaves glabrous to sparsely pubescent with curved, appressed trichomes, mm long, the trichomes of the main veins spreading to appressed-antrorse, the trichomes of the margin sparsely to densely ciliate, mm long. PEDICELS AT ANTHESIS green to purple, (2) cm long, 2 8 times as long as subtending leaf, pubescent, the trichomes sparse, antrorsely appressed, mm long. CALYCES AT ANTHESIS green or purple, broadly cupulate, (1.5) (4.5) mm long, mm diameter when fresh, mm when pressed and dried, the teeth widely spreading to reflexed, emerging from the ribs at approximately the same level, of approximately equal length, (0.5) (6) mm long, the ribs and teeth pubescent to villous, the trichomes antrorsely appressed to spreading, mm long. COROLLAS orientated laterally, not nodding, the tube mm long, the limb cm diameter, the membrane white to lilac, the adaxial sides of the lobes matching the membrane in coloration but with darker violet veins, the abaxial sides of the lobes green to lilac, pubescent, the trichomes antrorsely appressed, mm long. ANDROECIUM with green, glabrous filaments, these nearly round in cross-section and somewhat curved downward, the lowest (8.5) mm long, the two lateral (1.5) (5) mm, the two upper (1.25) (4) mm, the length of the lowest (2.5) times that of the lateral; anthers exuding a sweet odour, (2.5) (6.5) mm long, the lowest anther elliptic to ovate, rarely lanceolate or oblong, the pores round to ovate, usually terminal (rarely slitlike and lateral in a few populations in southern Puebla and southern Oaxaca). GYNOECIUM with widely conic ovary, mm long, the ovules ; style usually curved downward (rarely curved upwards at the tip in a few populations in southern Puebla and southern Oaxaca), (12) mm long; stigma usually green, rarely purple, round to shallowly bilobed, mm diameter. FRUITS remaining attached to calyces at maturity, round to ovoid, 9 24 mm long, 9 26 mm diameter at widest point, the tip rounded; exocarp green (yellowish with age) with black or purple lines, glabrous; mesocarp thin, green and juicy, with profuse (32 83 per fruit), round to angular, yellow sclerotic granules; placental area narrow, greenish white, juicy; fruiting calyx enlarged, mm long, (9.5) mm diameter, the teeth recurved, often making a complete loop, sometimes broken, mm long; fruiting pedicels mm long, deflexed, the fruit lying on the ground; seeds (12) (143) per fruit, dark brown to black, oblong to depressed-obovate, smooth, ridged, (2.5) 3 4 mm long, (3.7) mm wide. Chromosome number: 2n = 24 (Dean and Solano 303). Indigenous names: Ojo de venado, tonchichi, chichi de perra. Distribution: Mexico (states of Guanajuato, Hidalgo, México, Oaxaca, Puebla and Querétaro). Like Lycianthes dejecta, and L. ciliolata, L. peduncularis may once have had a broader and more continuous distribution on limestone soils. It is currently restricted to

34 418 E. A. DEAN limestone soils on either side of the transvolcanic belt and to some eroded volcanic areas within the transvolcanic belt. The explanation for its presence on eroded volcanics is similar to that discussed for L. dejecta. In addition, this species may tolerate other, more unusual, substrates. I collected L. peduncularis in a canyon where onyx is mined and near a marble quarry. Several of the localities where others have collected this species are in or near mining areas. Generally, it is found in dry cactus-scrub (matorral of Rzedowski, 1988) vegetation. Habitats include eroded floodplains, the sides of washes and canyons, paths, and within or at the sides of agricultural fields, m. Usually, there is drainage nearby. The German collector C. A. Purpus wrote an interesting article on the habitat preferences of this species, including information on how best to cultivate it in the gardens of Germany (Purpus, 1923). Diagnostic features: Lycianthes peduncularis is recognized by its combination of prostrate to decumbent habit, simple, antrorsely appressed trichomes, small calyces, and round green fruit with maroon to black striations. This species is the only one within series Meizonodontae with sclerotic granules in the fruit. Specimens examined: FIELD ORIGIN UNKNOWN: No locality data or collector data (W); Leipzig Botanic Garden, no collector, label handwriting matches that of neotype, but not signed by Kunze (NY); Leipzig Botanic Garden, 1844, Kunze s.n. (W). MEXICO: STATE OF GUANAJUATO: Mpio. San José Iturbide, near Rancho El Guajolote, SW of San José Iturbide, one hwy exit S of exit to San José, on W side of hwy 57, 1829 m, 16.ix.1991, Dean & Starbuck 254 (DAV, XAL); Mpio. San José Iturbide, near Rancho El Guajolote, SW of San José Iturbide, one hwy exit S of exit to San José, dirt rd that goes W to large drainage, farm of Margarita Vargaz Fuentes de Acosta, 1829 m, 31.x.1991, Dean & Starbuck 308 (DAV, XAL); Mpio. San José Iturbide, alrededores de El Guajolote, 2250 m, 10.vii.1988, Rzedowski (IEB); Mpio. San José Iturbide, cerca de El Guajolote, 2200 m, 22.viii.1988, Rzedowski (IEB). HIDALGO: Mpio. Atotonilco El Grande, N of Atononilco El Grande and E of hwy 105, in canyon just upstream of the baths of Santa María de Amajac, 2050 m, 30.vi.1990, Dean & Solano 200 (MEXU); Mpio. Huichapán, 0.6 road km E of town of Comodoje, which is 18 rd km E of first exit to Huichapán travelling E on Hwy 45, 2200 m, 2.vii.1990, Dean & Solano 205 (DAV, MEXU); cerca de Atotonilco, en los Baños de Amajac, 2050 m, 3.vi.1976, Delgado-S. & Hernández 232 (CAS, MEXU, US); autopista Querétaro, km 79, 6.vi.1964, Gold 135 (MEXU); Mpio. Huichapán, Comodoje, 13 km al este de Huichapán, 2200 m, 25.vi.1980, Hernández-M (CAS, MEXU); Mpio. Zimapán, camino a Minas de San Miguel, 10 km al N de Zimapán, 2100 m, 6.x.1980, Hernández-M. & Rodríquez-B (MEXU, US); Mpio. Zimapán, 10 km al N de Zimapán, hacia a la mina San Miguel, 2200 m, 28.vi.1981, Hernández-M (MEXU); Mpio. Ixmiquilpan, ft, vii. Aug, Purpus s.n. (UC); Tula, vii.1905, Rose et al (NY); Mpio. Ixmiquilpan, barranca de Tolantongo, E- facing slope, along main trail to bottom N, 99 4 W, 7.vii.1974, Sohmer 9329 (MEXU). MÉXICO: Mpio. Huehuetoca, N of Huehuetoca along the road to Apaxco, c. 4.2 road mi from building Los Arcos in downtown Huehuetoca, W side of road, 2200 m, 1.vii.1990, Dean & Solano 201 (DAV, MEXU, MO, NY, UC, WIS, XAL); Mpio. Huehuetoca, N of town of Huehuetoca along the rd to Apaxco, c. 4.9 rd mi from building Los Arcos (in downtown Huehuetoca), E side of rd, near where RR tracks come close to rd, 2134 m, 3.viii.1991, Dean & Starbuck 244 (DAV, UC, XAL); Mpio. Huehuetoca, N of Huehuetoca, along the rd to Apaxco, c. 4.9 rd mi from building Los Arcos in downtown Huehuetoca, near turn in rd, where RR tracks can be seen on E side of rd, 2134 m, 18.ix.1991, Dean & Starbuck 257 (UC, XAL); Mpio. Huehuetoca, N of Huehuetoca, along the road to Apaxco, c. 4.9 rd mi from building Los Arcos in downtown Huehuetoca, near curve in rd where RR tracks come close to road, both sides of rd, 2134 m, 29.x.1991, Dean & Starbuck 303 (DAV, MEXU, UC, XAL); Mpio. Apaxco, Cerro la Manga, ladera NE, 2350 m, 8.vii.1981, Romero-R (MEXU); Mpio. Huehuetoca, ladera oeste del Cerro Mesa la Ahumada, 2300 m, 15.vi.1981, Romero- R.1354 (ENCB); parte alta del Cerro de la Cruz, 7 km al N de Tepotzotlán, 2500 m, 12.vii.1975, Rzedowski (F, MEXU); 6 km al N de Huehuetoca, sobre la carretera a Apaxco, 2350 m, 11.ix.1977, Rzedowski (F); 6 km al N de Huehuetoca, sobre la carretera a Apaxco, 2350 m, 11.ix.1977, Rzedowski (IEB); Cerro Ahumada, al N de Huehuetoca, 2400 m, 5.vii.1981, Rzedowski (MEXU). OAXACA: Comotlán y cercanías, 1700 m, 29.iv.1953, Bravo s.n. (MEXU, very depauperate form); 55 mi SE of Oaxaca along the road to Tehuantepec, in the mountains 9 mi NW of La Junta, 13.ix.1971, Clarke (TEX); Valle de Oaxaca, 1550 m, 8.vi.1897, Conzatti & González 285 (BH, GH, GOET); no exact locality, 1750 m, vii viii.1900, Conzatti & González 1072 (GH); de Matatlán a Tlacolula, 1600 m, vi.1906, Conzatti & Vásquez 1470 (GH); Nochixtlán, 2000 m, 20.vi.1907, Conzatti 1844 (F, MEXU); Mitla, 27.vi.1900, C. Dean 10 (F, GH); Mpio. San P. Villa de Mitla, c. 2.7 rd mi E of main road into the town of Mitla along the Mitla-Zacatepec rd, where rd meets foothills and crosses over river, W side of rd and N side of river, 1799 m, 18.vii.1991, Dean & Starbuck 230

35 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 419 (DAV, MEXU, XAL); Mpio. Nochixtlán, NW edge of town of Nochixtlán, along hwy 190, NW of city of Oaxaca, 2073 m, 22.vii.1991, Dean & Starbuck 235 (DAV, MEXU, XAL); Mpio. Tamazulapan, E side of hwy 175, S of city of Oaxaca, c. 15 rd km S of town of Miahuatlán in the Sierra Madre del Sur, 2012 m, 25.vii.1991, Dean & Starbuck 236 (DAV, XAL); Mpio. San Carlos Yautepec, along hwy 190 between city of Oaxaca and Tehuantepec, c. 3 rd mi S of town of Las Minas, c. 7 rd mi S of turnoff to San Juan la Jarcia, dirt rd, S side of hwy 190, 770 m, 29.vii.1991, Dean & Starbuck 239 (DAV, MEXU, NY, UC, XAL); Mpio. San Juan La Jarcia, slopes above hwy 190 between city of Oaxaca and Tehuantepec, nearly opposite turnoff to San Juan la Jarcia, E side of the road, 915 m, 29.vii.1991, Dean & Starbuck 240 (DAV, XAL); Mpio. Teposcolula, slopes above town of Teposcolula, along rd to San Andrés Laguna, in the Mixteca region of Oaxaca, 2043 m, 31.vii.1991, Dean & Starbuck 242 (DAV, MEXU, XAL); Mpio. Nochixtlán, NW side of town of Nochixltán, near large drainage that crosses hwy 190, 2073 m, 9.x.1991, Dean & Starbuck 283 (DAV, MEXU, UC, XAL); Mpio. San P. Villa de Mitla, ruins of Mitla, 13.x.1991, Dean & Starbuck 288 (DAV, XAL); Mpio. San P. Villa de Mitla, c. 2.7 rd mi N of entrance to town of Mitla, along Mitla-Zacatepec rd, where road meets foothills, crosses drainage, and turns north, 1799 m, 13.x.1991, Dean & Starbuck 289 (DAV, MEXU, UC, XAL); Mpio. San Carlos Yautepec, along hwy 190 between Oaxaca City and Tehuantepec, c. 3 rd mi S of town of Las Minas, c. 7 mi S of turnoff to San Juan la Jarcia, W side of rd, 770 m, 17.x.1991, Dean & Starbuck 292 (DAV, MEXU, NY, UC, XAL); Mpio. San Juan la Jarcia, slopes above hwy 190 between city of Oaxaca and Tehuantepec, nearly opposite turnoff to San Juan la Jarcia, NE side of the rd, 915 m, 17.x.1991, Dean & Starbuck 293 (DAV, XAL); Dist. Teposcolula, sobre el camino Teposcolula-San Andrés Lagunas, Mixteca Alta, 2160 m, 16.iv.1981, García-M. 258 (MEXU); in recently burned clearing, 7 mi by road SE of San Juan la Jarcia along Mex. hwy 190, 1000 m, 31.v.1973, Hansen et al (MEXU, WIS); 15 rd km SE of Miahuatlán on rd to Puerto Angel, in high mountains of Sierra Madre del Sur, 2400 m, 6.vii.1969, Marcks & Marcks 1082 (WIS); Las Naranjas, in the vicinity of San Luis Tultitlanapa, Puebla, v.1908, Purpus 3565 (F, NY, UC); Las Naranjas, v.1908, Purpus 3566 (GH, MO, UC); Santa Catarina, 14.vii.1910, Rusby 73 (NY); Dist. de Teposcolula, 5 km al NE de Chilapa de Díaz, 1750 m, 2.vii.1977, Rzedowski (CAS, F); Mitla, vi.1888, Seler 49 (GH); along Mitla-San Lorenzo rd on mountainside, 28.vi.1968, Smith & Kitchen 4787 (MEXU); 3 mi N of Mitla, 5800 ft, 16.vii.1969, Weaver 2149 (MEXU). PUEBLA: Mpio. Tehuacán, 4.4 km al E de San Pablo Tepetzingo, 24.vii.1979, Chiang et al. 60 (MEXU); Mpio. Zapotitlán de las Salinas, canyon of the onyx mines of San Antonio Texcala, S of Tehuacán along hwy 125 (Tehuacán-Huajuapan de León rd), 1677 m, 13.vii.1991, Dean & Starbuck 226 (DAV, MEXU, UC, XAL); Mpio. Zapotitlán de las Salinas, San Antonio Texcala, canyon where there are onyx mines, N of town, S of Tehuacán along hwy 125, 1677 m, 28.ix.1991, Dean & Starbuck 273 (DAV, UC, XAL); Mpio. Zapotitlán de las Salinas, San Antonio Texcala, along hwy 125, just S of Tehuacán, canyon with onyx mine N of town, 1677 m, 22.x.1991, Dean & Starbuck 298 (DAV, MEXU, NY, UC, XAL); Mpio. Zapotitlán de las Salinas, minas de las canteras de San Antonio Texcala, 1700 m, 9.vi.1978, Ventura-A (CAS, ENCB, F, MEXU). QUERÉTARO: Mpio. Cadereyta, 5 km al S de Vizarrón, sobre el camino a Cadereyta, 2300 m, 16.iii.1989, Rzedowski (XAL). Schlechtendal cited three syntypes, one based on cultivated material from the Halle Botanical Garden (which he cited after the species epithet) and two herbarium specimens from B, cited near the end of the protologue: (1) MEXICO. Hidalgo: Mineral del Monte, Real del Monte, Omitlan, Velasco, Regla, C. Ehrenberg 81; (2) MEXICO. Michoacán, Angangueo, Schiede s.n. Schlechtendal mentioned at the end of the protologue that the plants in their natural habitat... (presumably represented by the specimens from B)... have denser pubescence, thus acknowledging that the herbarium specimens did not match the horticultural specimen. I have not been able to obtain authentic material of those two specimens; they may have been destroyed in Berlin in World War II. Georg Bitter determined them to be Lycianthes moziniana, and until I see authentic material, I am accepting his judgement. Because Schlechtendal placed primary importance on the horticultural syntype, it is assumed that he took his description from that specimen. I have not seen authentic herbarium material of the specimen from the Halle Botanical Garden, but according to G. Bitter, material existed at the time he completed his monograph (perhaps destroyed in Berlin during World War II). The only herbarium specimens that survived were made by Kunze at the Leipzig Botanical Garden. Those specimens match Schlechtendal s description very well. Until authentic material from the Halle Botanical Garden can be found, I am neotypifying Solanum pedunculare on a specimen collected from the Leipzig Botanical Garden by Kunze and annotated by G. Bitter. This specimen matches the description in the protologue. Lycianthes rzedowskii E. Dean (Fig. 15). Lycianthes rzedowskii E. Dean, Novon 4(4): TYPE: MEXICO. State of Michoacán: Mpio.

36 420 E. A. DEAN Figure 15. Herbarium specimen of Lycianthes rzedowskii (Dean & Starbuck 221 (DAV)). Charo, along hwy 15, 20 rd km E of Morelia, just E of Pontezuelas, 2165 m, 13.xi.1991, E. Dean & T. Starbuck 322a (HOLOTYPE: UC; ISOTYPES: DAV, NY, XAL). Perennial HERBS, erect, often recumbent with age, cm tall. INDUMENT of simple or dendritic trichomes. ROOTS with fusiform segments, usually present cm below the soil surface. STEMS arising from the root one to several; below-ground growth not extensive, white or purplish, often with stemborne roots just below the soil level; above-ground growth green to reddish purple, terete in cross-section, becoming somewhat woody with age, compressed and ribbed when dried, the new growth pilose to villous (sometimes glabrous), often glabrate with age, the trichomes mm long, rarely of two distinct lengths, the shorter less than 0.25 mm long, regularly interspersed with others mm long. FIRST SYM- PODIAL UNIT erect above ground, 7 90 cm long, 2 8 mm diameter at widest point; internodes (6) in number, usually numerous; lateral branching usually lacking. FIRST TWO SYMPODIAL BRANCHING POINTS usually dichasial (rarely only the first), followed by monochasial branching; first dichasial branching angle (100). SUBSEQUENT SYMPODIAL GROWTH poorly developed, much shorter than first sympodial unit, extending laterally 6 40 cm, the second sympodial units 1 11 cm long, 2 5 mm wide, subsequent sympodial units becoming reduced distally. ABOVE- GROUND LEAVES OF FIRST SYMPODIAL UNIT membraneaceous to chartaceous; leaves closest to the soil surface reduced, scale-like, early deciduous; subsequent leaves expanded, divided into expanded lamina and attenuate base, the attenuate portion plus petiole accounting for 1/8 1/3 (1/2) of total leaf length, the lamina narrowly to broadly ovate, elliptical or obovate, the margin undulate, the tip acute to acuminate, the base obtuse to cuneate, attenuate into the petiole, the largest expanded leaves (in the middle or upper one third of the stem) (4) cm long, 1 6 cm wide, the petiole poorly defined, 1 28 mm long. LEAVES OF SUB- SEQUENT SYMPODIAL UNITS usually geminate, similar in shape and texture to the distal-most leaf of the first sympodial unit, but the lamina ovate to elliptic, the base cuneate, short-attenuate into the petiole; largest leaves (generally on the second sympodial unit) cm long, cm wide, the petiole poorly defined, mm long, the leaves becoming reduced, narrower (lanceolate) and subsessile on more distal sympodial units; geminate leaf pairs unequal in size (the smaller 1/4 3/4 the size of the larger), the smaller of the pair with more oblique leaf base. LEAF PUBESCENCE similar throughout, the leaves puberulent, the trichomes mm, adaxially appressed, uniformly pointing at the same angle toward leaf margin, abaxially spreading, concentrated on leaf veins, the margins ciliate. PEDICELS AT ANTHESIS green, sometimes with a few purple spots, cm long, (2.2) times as long as subtending leaf, glabrous to nearly villous, the trichomes mm long, spreading to slightly retrorse, straight to slightly undulate. CALYCES AT ANTHESIS green, campanulate, 3 6 mm long, mm diameter when fresh, mm when pressed and dried, the teeth widely spreading to slightly reflexed, emerging from the ribs at two distinct levels c mm below margin, variable in shape and length, from small knobs to linear-lanceolate and 7.25 mm long, the ribs and teeth glabrous to villous, the trichomes spreading to somewhat retrorse. Corollas slightly nodding, the tube mm long, the limb cm in diameter, the membrane white, translucent, notched at the margin, pulled smooth and taut when lobes open, the lobes lanceolate to oblanceolate, the upper narrower, mm wide, the lower 3 6 mm wide, the adaxial sides of the lobes white with 3 5 violet veins near throat, the abaxial sides of the lobes green, usually glabrous.

37 LYCIANTHES SERIES MEIZONODONTAE (SOLANACEAE) 421 ANDROECIUM with pale green, erect, laterally compressed filaments, sometimes slightly pubescent, the lowest mm long, the two lateral mm, the two upper mm, the length of the lowest always less than twice that of the lateral; anthers exuding a sweet powdery fragrance, mm long, the lowest anther broadly lanceolate to elliptical, the pores round to oval, terminal. GYNOECIUM with conic ovary (often attenuate into the style), mm long, the ovules 30 70; style straight to gently curved, 7 11 mm long; stigma green, captitate, rarely somewhat lobed, slightly oblique, 0.5 mm in diameter. FRUIT remaining attached to calyces at maturity, turbinate, elongate, mm long, 8 19 mm in diameter, the tip attenuate; exocarp light green with lighter or darker green spots or lines when immature, dull light purple to black at maturity; mesocarp dark purple, soft and juicy, lacking sclerotic granules; placental area light purple, powdery; fruiting calyx enlarged, 2 10 mm long, mm in diameter, the teeth stout, stiff, remaining appressed to fruit or somewhat spreading, often broken, mm long; fruiting pedicels cm long, deflexed, the fruit pendant; seeds 3 38 per fruit, rusty brown to black, depressedobovate to angular trullate, rough in texture, ridged along one side, blistered along the ridge, mm long, mm wide. Chromosome number: 2n = 24 (Dean & Starbuck 212, 317, 336). Indigenous names: Chilillo. Distribution: Mexico (states of México, Morelos and Michoacán). Lycianthes rzedowskii grows on volcanic soils in oak, oak pine, pine oak and fir forests. Its preferred habitat is forested slopes near drainages, m. Dean & Starbuck 212 from the state of México is placed in this species but differs from typical Lycianthes rzedowskii in having tricolporate pollen and phanerocotylar germination. It is not interfertile with other populations of the species. This population obviously represents a cryptic species that needs further work. Diagnostic features: Lycianthes rzedowskii may be confused with L. acapulcensis and L. ciliolata. It can be distinguished from those species by its welldeveloped erect to reclinate first sympodial unit with numerous internodes (many of these with expanded leaves). At the time of flowering, this species has very limited sympodial growth (the first sympodial unit is much longer than subsequent total growth), although in some populations the sympodia elongate during fruit maturation. This species is also distinguished by its combination of smooth, white (rarely pale lilac) corollas, broad sweetly scented anthers, and terminal, round anther pores. One of the best ways to distinguish L. rzedowskii from the other two species is to look at the relative lengths of the stamen filaments. In L. rzedowskii the length of the lowest (and longest) filament is never more than twice that of the lateral filaments, whereas in the other two species, the length of the longest filament is almost always more than twice that of the lateral filaments. Lycianthes rzedowskii may hybridize with L. starbuckii and L. acapulcensis (see discussions under those species). Specimens examined: MEXICO: STATE OF MÉXICO: Mpio. Sultepec, NE of Capula along the road to Sultepec, c. 2.5 rd km from the outskirts of Capula, 1.0 rd km NW of the turnoff to Tejupilco, 2317 m, 13.vii.1990, Dean & Starbuck 212 (DAV, ENCB, MEXU, MO, NY, UC, XAL); Nanchititla, Barranca de la Cueva de Santos, downstream from dam, 1794 m, 9.xi.1991, Dean et al. 317 (DAV, XAL); Mex. hwy 130, 10.1 miles N of Temascaltepec, 1940 m, 27.vii.1972, Denton 1900 (UC); District Temascaltepec, Comunidad, 2610 m, 7.vii.1932, Hinton et al. 971 (BM, F, G, MEXU); District Temascaltepec, Nanchititla, 26.vii.1935, Hinton et al (F, GH, MO, NY); Nanchititla, 29.viii.1935, Hinton et al (GH, K); La Ciénega, 5 km al S de Sultepec, sobre el camino a Amatepec, 2400 m, 15.vii.1973, J. Rzedowski (CAS, ENCB, F). MICHOACÁN: 13 miles E of Morelia,13.viii.1947, Barkley et al (F, TEX); along route 15 between K268 and 269, c. 45 km E of Morelia, 2830 m, 10.viii.1966, Cruden 1171 (MEXU, UC); Mpio. Charo, along hwy 15, 20 rd km E of Morelia, just E of Pontezuelas, 2165 m, 24.vii.1990, Dean & Starbuck 220 (DAV, MEXU, XAL); Mpio. Queréndaro, along old hwy 15 c. 0.4 rd km W of San José de la Cumbre, near the km 195 marker, 2500 m, 25.vii.1990, Dean & Starbuck 221 (DAV, MEXU); Mpio. Zitácuaro, near Macho de Agua, E of Zitácuaro, along old hwy 15, c. 6 8 rd mi E of RR crossing, m, 11.xi.1991, Dean & Starbuck 320 (DAV, ENCB, UC, XAL); Mpio. Charo, waterfall along old hwy 15, E of Morelia, just E of intersection with rd to Tzitzio, 2250 m, 13.xi.1991, Dean & Starbuck 322b (NY, UC, XAL) and 7.xii.1991, Dean & Starbuck 336 (DAV, XAL); Jesús del Monte, near Morelia, Galeotti 1182 (BR); Zitácuaro, Cerro Pelón, 17.vi.1938, Hinton et al (GH, K); 23 km E of Morelia, km 290 from Mexico City (on old Highway 15), 2600 m, 28.vi.1964, Mick & Roe 163, 163a (WIS); Pontezuelas, 20 km al E de Morelia, sobre la carretera a Zitácuaro (km 289), 2100 m, 21.vii.1964, Rzedowski (ENCB, WIS); Las Peras, 38 km E of Morelia (c. 33 km air dist.), km 272 on hwy 15 (Ciudad Hidalgo to Morelia), 2515 m, 13.ix.1962, Ugent & Flores 2012 (WIS). MORELOS: Sierra de Morelos,

38 422 E. A. DEAN Figure 16. Holotype of Lycianthes starbuckii (Dean & Starbuck 315 (UC)). Cuernavaca, 2050 m, 26.vii.1969, Hinton et al (NY); Huitzilac, 1.vii.1930, Lyonnet 713 (BM, GH, MEXU (one duplicate is a mixed collection with L. acapulcensis), MO; duplicate at NY is L. acapulcensis); hacia el Valle del Tepeite, Canal Zempoala, viii.1932, Lyonnet 1002 (CAS, MEXU). Lycianthes starbuckii E. Dean (Fig. 16) Lycianthes starbuckii E. Dean, Novon 4(4): TYPE: MEXICO. State of México: Sierra de Nanchititla, oak forest across the reservoir from the town of Nanchititla, 1945 m, 8.xi.1991, E. Dean et al. 315 (HOLOTYPE: UC; ISOTYPES: BM, DAV, ENCB, F, MEXU, MO, NY, XAL). Perennial HERBS, usually prostrate to ascending, to 15 cm tall. INDUMENT of simple trichomes. ROOTS with fusiform segments, often buried more than 10 cm below the soil surface. STEMS emerging from the root, one to several; below-ground growth sometimes extensive and branching, whitish; above-ground growth green-purple, highly branched from near the soil level, terete in cross-section, without much woody tissue, collapsed when dry, pilose to velutinous, the trichomes mm long near the soil level, these shorter distally, mm long. FIRST SYMPODIAL UNIT mostly below-ground, extending above the ground (18.5) cm, 2 4 mm diameter; above-ground internodes 2 5 in number. FIRST SYMPODIAL BRANCHING POINT usually dichasial, followed by a mixture of monchasial and dichasial branching points; first dichasial branching angle SUBSEQUENT SYMPODIAL GROWTH extensive, sometimes below ground, usually resting on the soil surface, often forming mats, extending to 20 cm diameter, the second sympodial unit cm long, mm diameter, subsequent sympodial units showing similar variation in size, with those at the tips becoming reduced. ABOVE-GROUND LEAVES OF FIRST SYMPODIAL UNIT thick-chartaceous, fleshier than those of other species in the series, 2 5 in number; leaves closest to the soil surface reduced, often scalelike, early deciduous, the subsequent ones expanded, each larger than the one below it, divided into expanded lamina and attenuate base, the attenuate portion plus the petiole accounting for 1/3 1/2 of total leaf length, the lamina obovate to oblanceolate, the margin entire, the tip broadly acute to rounded, the base cuneate, attenuate into the petiole, the largest expanded leaves to 6 cm long, 3 cm wide, the petiole to 1.8 cm long. LEAVES OF SUBSEQUENT SYMPODIAL UNITS geminate or not (due to the mixture of monochasial and dichasial branching), the leaves similar in shape and texture to distal-most leaves of the first sympodial unit, but the lamina ovate to broadly elliptical to rhombic, the margin entire to undulate, the largest leaves to 3 6 cm long, cm wide, the petiole to 1.4 cm long (generally shorter), the leaves becoming reduced, narrowed, more elliptical, subsessile distally; geminate leaves similar in shape, unequal in size, the smaller 1/4 1/2 the size of the larger, with rounder tip, more oblique leaf base, and shorter petiole. LEAF PUBESCENCE similar throughout, the leaves puberulent with acute appressed trichomes, adaxially pointing uniformly toward the leaf margin, mm long, abaxially appressed-ascending along veins, to 0.25 mm long, the margins short-ciliate with ascending trichomes, mm long. PEDICELS AT ANTHE- SIS green with some purple, cm long, times as long as subtending leaf, pilose, the spreading trichomes of two distinct lengths, the shorter (to 0.25 mm long) regularly interspersed with the longer (0.5 mm). CALYX AT ANTHESIS green and purple, narrowly to broadly conical, 3 4 mm long, mm wide when fresh, when pressed and dried, always broader than long, the teeth lax and flexuous, not reflexed, emerging from the ribs at approximately the same level, of approximately equal length, mm