Corresponding author: Sandra Knapp (
Academic editor: L. Giacomin
The
El clado Elaeagnifolium contiene cinco especies arbustivas, usualmente rizomatosas, distribuidas en desiertos y bosques secos de Norte y Sudamérica. Previamente, los miembros del clado estuvieron agrupados en las secciones
Knapp S, Sagona E, Carbonell AKZ, Chiarini F (2017) A revision of the
The large size of
Plants collected by William Houstoun on the Caribbean coast of Mexico were cultivated by Philip Miller of the Chelsea Physic Garden in London and were described as
Exploration of the western United States along with the trade in seeds between European botanic gardens led to the description of many synonyms of the extremely variable species
In his treatment of the
In floristic works
Description of the Argentine endemics
All of the members of this small species group were only recognised as being related as the result of molecular phylogenetics (
Trichome morphology representative of that found in members of the Elaeagnifolium clade.
Habit and stems. Members of the
The stems of all members of the group are variably prickly; this variability is most pronounced in
Sympodial growth is characteristic of
Leaves. Leaf morphology in subgenus
Leaf size also varies with season and environmental conditions; plants collected in the wet season or from wetter areas always have larger leaves than those from drier microhabitats or that were collected in the dry season.
Pubescence. In common with the rest of the
In herbarium specimens, the trichomes of all these species are pale grey, giving the plants a silvery cast. On live plants, the trichomes of
Flowers of members of the Elaeagnifolium clade.
Prickles. Prickles in
Prickles can occur on all above-ground parts of a plant except the corolla and the fruit. Density and distribution of prickles vary with the age of the plant and environmental conditions and, thus, are not particularly useful characters; all species of the group can have branches (see Figs
Fruits of members of the Elaeagnifolium clade.
Inflorescences. As with all species of
Calyces. Members of the Elaeagnifolium clade have 5-merous flowers like most other species of
Corollas. Like the calyces, corollas of members of the Elaeagnifolium clade are most often 5-merous. Corollas are stellate, and usually divided about halfway to the base, the lobes are deltate to triangular with copious to sparse interpetalar tissue, and are usually spreading at anthesis (see Fig.
The abaxial surfaces of the corolla lobes are densely pubescent with stellate trichomes where they are exposed in the bud; the interpetalar tissue in the sinuses is glabrous on both surfaces in all species. In exceptionally prickly individuals of
Androecium. The anthers of all members of the Eleagnifolium clade, like those of most of the spiny solanums, are long-tapering, with distally directed pores that do not lengthen to slits with age. In all five species the anthers are heteromorphic, with three of the five slightly longer than the rest, and are usually slightly curved (see Fig.
Petanatti and Del Vitto (1991) record structures they called “bridas” (flanges) on the anthers of
The filaments in all members of the group are composed of a very short filament tube, and a glabrous free portion that is usually ca. 1/5 the length of the anthers themselves (see Fig.
Gynoecium. All of the species of the Elaeagnifolium clade except
Fruit. As with all species of
Protein extracts from ripe berries of
Seeds. Seed morphology has been suggested to be a useful character for species-level taxonomy in
Chromosomes. Of the five species in the Elaeagnifolium clade, only
Chromosome numbers in the Elaeagnifolium clade.
Species | Haploid chromosome number (n) | Reference or voucher |
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n=12, 24, 36 (Argentina); n=12 (North America) | |
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n=12 |
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n= ca. 24 | |
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n=12 | |
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not known | – |
In the southern part of its range
Habitats and distribution. Members of the
Habitats of members of the Elaeagnifolium clade.
The members of the group (excluding the invasive populations of
Unravelling dispersal history of members of the Elaeagnifolium clade will require more detailed phylogenetic sampling in the spiny solanums; the amphitropical pattern is found in other small groups in the larger
Pollination and breeding systems. Like all
Bee visitors recorded on members of the Elaeagnifolium clade. Scientific names of bees are as recorded in the original publications, with more recent taxonomic placements (taken from
Species | Country | Bee Species | Reference |
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Argentina (Prov. Mendoza, environs of Mendoza) |
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Argentina (Prov. Mendoza, environs of Mendoza) |
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Argentina (Prov. San Luis ?) |
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Australia (greenhouse study, Adelaide) |
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Greece |
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United States (Arizona, Cochise Co., near Portal, Chiricahua Mountains) |
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United States (Arizona, Cochise Co., near Douglas) |
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United States (Arizona, Cochise Co., near Portal, Chiricahua Mountains) |
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United States (Arizona, Cochise Co., near Portal, Chiricahua Mountains) |
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United States (Arizona, cultivated plants) | “great attractors of carpenter [ |
Arizona Sonora Desert Museum Fact Sheet ( |
The anthers of members of the Elaeagnifolium clade are not tightly connivent (pepper pot configuration of
Andromonoecy, possession of hermaphroditic and functionally staminate flowers in a single inflorescence, has evolved multiple times in
Conservation status. With the exception of
Preliminary conservation status for members of the Elaeagnifolium clade. See Methods for details of threat status calculation, and species discussions for further discussion.
Species | IUCN threat status | Criteria ( |
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Widespread invasive | |
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Widespread, but relatively local | |
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B1 ( |
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Widespread | |
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B1 ( |
Our goal for the treatment of the species of this small clade has been to provide circumscriptions for the members of this morphologically variable group of species, while clearly highlighting those taxa and populations where further in-depth research would be useful. Delimitation of species here basically follows what is known as the “morphological cluster” species concept (
This monograph is based on examination of herbarium specimens supplemented with field observations in North and South America. We examined approximately 2,100 collections (ca. 2,700 specimens) from the following herbaria (herbarium acronyms follow
Measurements were made from dried herbarium material supplemented by measurements from living material. Colours of corollas, fruits, etc., are described from living material or from herbarium label data. Specimens with latitude and longitude data on the labels were mapped directly. Some species had few or no georeferenced collections; in these cases, we retrospectively georeferenced the collections using available locality data. The extremely widespread
Where specific herbaria have not been cited in protologues we have followed
Type specimens with sheet numbers are cited with the herbarium acronym followed by a dash and the sheet number (i.e., MO–1781232); barcodes are written as a continuous string (i.e., G00104280). We have cited geographically representative specimens for taxa where more than 100 collections are known. Full specimen details are available on the
Citation of literature follows BPH-2 (Bridson 2004) with alterations implemented in IPNI (International Plant Names Index,
Shrubs, sometimes rhizomatous, armed or unarmed. Stems terete, pubescent with multangulate, porrect-stellate or lepidote trichomes, sometimes glabrescent. Sympodial units difoliate or occasionally trifoliate or plurifoliate, not geminate. Leaves simple to shallowly lobed to occasionally somewhat pinnatifid, concolorous or discolorous, densely pubescent with multangulate, stellate or lepidote trichomes; petioles well developed, sometimes channelled above. Inflorescences terminal to lateral, usually unbranched, occasionally furcate, not bracteate, with up to 10 flowers (exceptionally to 26 flowers in
An exclusively New World group occurring in North America (western United States and Mexico) and southern South America (Argentina, Paraguay, Brazil, Uruguay, Chile). One species,
As discussed above under Phylogeny and as can be seen by the synonymy of the clade, members of this group were previously not thought to be closely related. Their resolution as sister to the diverse and diversifying Old World clade of spiny solanums makes them of particular interest in terms of character evolution.
The clade, as is common with the groups of spiny solanums, has few unambiguous and unique synapomorphies. The andromonoecious habit (very weak in
1 | Pubescence of leaves lepidote, the trichomes with the rays fused at the base; prickles, if present, often orange; flowers mostly all perfect; stamens equal or nearly so; fruits more than 1 per infructescence |
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– | Pubescence of leaves stellate or multangulate, the trichomes with the rays not fused or if very slightly fused near the base; prickles, if present, brown or yellow; flowers heteromorphic, not all perfect; stamens unequal; fruits usually 1 per infructescence. |
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2 | Trichomes of stems and leaves multangulate, the rays in multiple planes; Argentina |
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– | Trichomes of stems and leaves stellate, the rays in a single spreading plane (porrect-stellate) |
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3 | Pedicels strongly deflexed in fruit; trichome rays always fewer than 10; midpoint equal in length to the lateral rays; flowers white; Argentina |
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– | Pedicels erect in fruit; trichome rays usually more than 10; midpoint shorter than the lateral rays; flowers purple or lavender (occasionally white); Mexico |
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4 | Flowers weakly heteromorphic, the anthers in hermaphroditic and staminate flowers not different, straight or only slightly curved; fruiting calyx not or only sparsely prickly, usually not covering the berry by more than 2/3 of its length; Sonoran Desert |
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– | Flowers strongly heteromorphic, the hermaphroditic flower with the anthers more or less equal in size, the staminate flowers with 3 anthers much longer than the rest and strongly curved; fruiting calyx prickly, accrescent and almost completely enclosing the berry; widespread in Mexico |
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Cultivated in Madrid from “America calidiore” [“del viaje de los españoles alrededor del mundo, Cult. en el R. J. Bot. 1793”],
Herbaceous, woody at base, 20 to 50 cm tall. Stems erect, with running underground stems, sparsely or densely armed, sometimes unarmed; young stems densely pubescent with stellate, sessile to subsessile, porrect lepidote trichomes, the stalk 0.1–0.15 mm long, the rays 10–14(16), 0.1–0.3 mm long, fused at the base, the midpoints 0.1–0.15 mm long, usually absent, unarmed or prickly with the prickles, if present, usually uniform throughout the plant, 2–6 mm long if dense, 0.3–2 mm long if sparse, 0.5–2 mm wide at base, usually straight, orange to brick-red; bark of older stems smooth, brown to grey, the leaf scars small brown stumps. Sympodial units difoliate, not geminate. Leaves simple or shallowly lobed, (1.5)3–6(12) cm long, (0.3)1–2(3) cm wide, (2)3–4(6) times longer than wide, elliptic, sometimes lanceolate or ovate, mostly concolorous, drying yellow-green or silvery green, densely pubescent on both surfaces with sessile to subsessile lepidote trichomes, the rays (9)12–14, 0.1–0.3 mm long, porrect, fused at the base, the midpoint up to 0.15 mm long, sometimes reduced; principal veins 4–7 pairs, raised adaxially, spreading at ca. 30–45° from the midvein, the finer venation mostly not visible to the naked eye; base obtuse to rounded or cuneate, often oblique; margins entire to shallowly lobed, the lobes 4 to 6 on each side, the sinus extending up to ½ of the distance to the midvein, to 2(4) cm long, rounded to deltate; apex obtuse to rounded, rarely acute; petiole 0.2–2(4) cm, 1/2–1/6 of the leaf length, densely stellate-pubescent like the young stems. Inflorescences terminal or lateral, (2)3–5(6) cm long, with (3)4–6(7) flowers, unbranched (simple), densely pubescent with lepidote trichomes like the stems; peduncle 0.8–2.5 cm long; rachis 0.5–3 cm long; pedicels 1–3 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely stellate-lepidote-pubescent like the young stems, usually armed, articulated very near the base; pedicel scars brown, spaced 0.5–1 mm apart, sometimes obscured by the dense pubescence. Buds ellipsoid, the calyx ca. 1/3 of the corolla length prior to anthesis. Flowers 5-merous, usually all perfect, but some distal ones short-styled, functionally staminate and the plants weakly andromonoecious. Calyx tube 2–3 mm long, conical or cup-shaped, the lobes 2- 4 mm long, 1–2 mm wide at base, cuspidate, venation not visible, densely lepidote-pubescent like the leaf blades, sparsely prickly or unarmed, if present the prickles to 2 mm long. Corolla 2.5–3(4) cm in diameter, purple to pale violet or white, stellate, lobed ca. 1/2 of the way to the base, the lobes 0.6–1(1.2) cm, 0.6–1 cm wide, deltate, reflexed to spreading at anthesis, often with a yellow midvein, stellate-pubescent abaxially, mostly glabrous adaxially, the trichomes variously irregular and reduced-multangulate. Stamens equal or slightly unequal in size, with the 2 adaxial anthers very slightly shorter than the 3 abaxial anthers; filament tube 0.8–1.5 mm; free portion of the filaments 1–2 mm; anthers 6–10(12) mm long, 1.5–2 mm wide, tapering, spreading or coherent, the surfaces often papillate, poricidal at the tips, the pores not lengthening with age. Ovary globose to ovoid, usually densely stellate-pubescent like the young stems, sometimes glabrous; style 7–15 mm long, glabrous, slightly dilated toward the apex, in rare short-styled flowers 2–4 mm long; stigma clavate, papillose. Fruit a globose berry, 1–5 per infructescence, 0.6–1.2 cm in diameter, the pericarp thin, smooth, glabrous, yellow to orange at maturity, drying orange to brown and cracking while still on the plant and the seeds released as a mass; fruiting calyx not markedly accrescent, the lobes reflexed or very occasionally covering up to 2/3 of the mature fruit, with prickles ca. 2 mm long; fruiting pedicels 1–2.5 cm long, 0.5–1(2) mm in diameter at the base, 1–2(3)mm in diameter at the apex, somewhat woody, strongly deflexed, usually armed. Seeds 20–60 per berry, 2–3 mm long, 2–3 mm wide, flattened reniform, orange-brown, the surfaces shiny, usually borne in a mass in the berry, the testal cells sinuate (not differing in size between ploidy levels, Nilda Dottori, pers. comm.). Chromosome number: n=12, 24, 36 (
(Figure
Distribution of
Argentina: capiquí, meloncillo de campo, meloncillo de olor, quillo, suriñado, tutiá chico, granadillo, tomatillo, pocotillo, revienta caballos (
In areas where
The berries of
(
Populations of
As discussed above in the section on Habitats and distribution it was long assumed that
Introduction of
Measures for the control of
Lectotypes for most of the synonyms of
Mexico. Baja California: Bay of Magdalena [Bahía Magdalena], [1841],
Erect shrub, 0.5-3 m tall. Stems armed or more often unarmed; young stems pale yellowish green, densely stellate-pubescent with sessile to short-stalked porrect-stellate trichomes with 8–12 rays (0.03-)0.1–0.2 mm long, the midpoints up to 0.07(0.1) mm long, often reduced; prickles if present 0.2–1.5 cm long, 0.2–2 mm wide at base, usually straight, reddish brown, bark of older stems smooth, brown or greyish or reddish brown, glabrescent. Sympodial units difoliate, not markedly geminate. Leaves simple, (1-)2–3 cm long, (0.5-)0.8–2.3(-6) cm wide, ovate to elliptic (unusually narrowly elliptic), usually concolorous, drying yellowish green to pale green, densely pubescent on both sides and the blade surface not visible, the pubescence usually more dense beneath, the trichomes porrect-stellate, sessile or short-stalked, translucent, the rays 8–14, 0.05–0.2(0.3) mm long, straight, slightly fused near the midpoint base, the midpoint to 0.1 mm, often absent; primary veins 4–7 pairs, raised adaxially, flat abaxially, spreading at ca. 45° to the midvein, the tertiary venation mostly obscured by the dense stellate pubescence; base rounded to truncate, usually oblique; margins entire, rarely shallowly lobed, if so the lobes 2–3 on each side, the sinus extending only to 1/8 of the distance to the midvein, up to 3 mm long, rounded; apex obtuse or rounded; petiole 0.5–1.5 cm long, 1/3–1/4 of the leaf length, densely stellate-pubescent like the young stem. Inflorescences terminal or lateral, 3–4 cm long, unbranched, with 3–4 flowers; peduncle 1–2 cm long, rachis 0.5–1 cm long; pedicels 0.4–0.5 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely stellate-pubescent like the leaf blade, unarmed, even in plants with prickly stems, articulated near the base; pedicel scars spaced 1–2 mm apart, forming prominent brown stumps. Buds ellipsoid, the calyx ca. ½ of the corolla length prior to anthesis. Flowers 5-merous, heteromorphic and the plants andromonoecious, the lower ones (1–3) hermaphroditic and long-styled and the more distal staminate and short-styled. Calyx tube 6–7 mm long, conical or cup-shaped, strongly keeled at the midvein, the lobes 2–3 mm long, 2–3 mm wide at base, long-triangular from a deltate base, densely stellate-pubescent abaxially, unarmed. Corolla 2.5–4(-5) cm in diameter, apparently expanding with age (slightly smaller in short-styled flowers), violet, drying yellowish-light brown, lobed for 2/3–1/2 of its length, pentagonal with abundant interpetalar tissue, the lobes 0.8–1.2 cm long, 1–1.2 cm wide, broadly deltate with an abruptly acuminate tip, spreading or slightly reflexed, densely stellate-pubescent abaxially where exposed in bud, the interpetalar tissue glabrous abaxially. Stamens equal or very slightly unequal, with the 2 adaxial anthers shorter than the 3 abaxial anthers; filament tube 1–1.5 mm long; free portion of the filaments 2–3 mm long, glabrous; anthers 6–10 mm long, free, slightly unequal, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, the anther surface smooth to finely papillose. Ovary globose, 2–3 mm in diameter, glabrous; style in long-styled flowers 13–18 mm long, glabrous, strongly curved, in short-styled flowers 5–6 mm long, straight or only slightly curved; stigma capitate in long-styled flowers, the surface minutely papillose. Fruit a globose berry, 1(-3) per infructescence, 1–1.5 cm in diameter when dry, the pericarp thin, smooth, glabrous, light green, sometimes with dark stripes or a marbled pattern when young, drying dark brown or brown-reddish, drying and cracking open to release the seeds when mature; fruiting pedicels 1–2 cm long, 1–2 mm in diameter at the base, 2–3 mm at the apex, woody, erect, usually unarmed; fruiting calyx not accrescent, up to 1.5 cm long, usually unarmed. Seeds ca. 10–30 per berry. 2–3 mm long, 2–3 mm wide, flattened reniform, golden to dark reddish brown, the surface minutely pitted, the margins thickened, the testal cells sinuate, more rectangular in shape near the margins. Chromosome number: n=12 (
(Figure
Distribution of
In the Sonoran Desert biome (
Mexico. Baja California: mariola (many collections); Baja California Sur: mariola (many collections), trompillo (MEXU-1203067), malva (
(
Argentina. Córdoba: Dpto. Sobremonte, a 6 km de San Francisco de Chañar, a orilla de ruta rumbo a Lucio V. Mansilla,
Erect rhizomatous shrub, 0.3–0.5 m tall. Stems sparsely armed or more commonly unarmed; young stems densely pubescent, the trichomes stellate, porrect, yellowish golden, sessile and short-stalked, the rays 6–8, ca. 1 mm long, the midpoints up to 1 mm long, equal in length to the rays; prickles if present 5–6 mm long, straight, slightly wider at the base, pale yellowish brown; bark of older stems smooth, brown or yellowish brown from persistent pubescence. Sympodial units difoliate or trifoliate, not markedly geminate. Leaves simple, 3–8 cm long, 1–3.1 cm wide, elliptic to narrowly elliptic, 4–5 times longer than wide, mostly concolorous, drying yellowish green to green; adaxial surfaces densely pubescent with translucent, mostly sessile porrect, stellate trichomes, the rays 8–14, 0.05–0.2(0.3) mm long, straight, the midpoint up to 0.1 mm, often reduced; abaxial surfaces densely pubescent with stalked porrect, stellate trichomes, the stalks to 0.5 mm long, the rays 8–10, 0.4–0.6 mm long, the midpoint shorter; principal veins 4–8 pairs, impressed adaxially, flat abaxially, spreading at ca. 45° to the midvein, the tertiary venation mostly not visible due to the dense pubescence; base acute; margins entire, rarely very shallowly lobed, the lobes 2–3 on each side, the sinus extending only 1/8 or less of the distance to the midvein, rounded; apex obtuse, rarely rounded; petiole 0.5–1.5 cm long, densely stellate-pubescent like the young stem, prickly, the prickles ca. 2 mm long, acicular. Inflorescences terminal or lateral, to 5 cm long, with up to 10 flowers, unbranched; peduncle less than 0.2 cm long, the lowest flower arising from very near the main axis; pedicels ca. 2 cm long, ca. 2 mm in diameter, robust, articulated near the base, densely stellate-pubescent like the leaf blades, the basal flower with the pedicel prickly, the prickles 2–3 mm long, orange-yellow; pedicel scars prominent, spaced 0.5–1.5 cm apart, space between the basal flower and the next much longer than that between the more distal staminate flowers. Buds ellipsoid, the calyx ca. 1/2 of the corolla length prior to anthesis. Flowers strongly heteromorphic and the plants andromonoecious, only the basal flower perfect and long-styled, the rest functionally staminate and short-styled. Calyx conical or cup-shaped, the tube 3–5 mm long, the lobes (3)6–8 mm long, 3–4 mm wide at base, lanceolate or long-deltate with acuminate apices, densely stellate-pubescent abaxially like the leaf blades, prickly in basal hermaphroditic flowers. Corolla 3–4 cm in diameter, white, drying yellowish-light brown, stellate, lobed for ca. 1/2 of the length, the lobes 0.8–1.5 cm long, ca. 1.2 cm wide, broadly deltate, densely stellate-pubescent abaxially. Stamens unequal, with the 2 adaxial anthers shorter than the 3 abaxial anthers; filament tube 1–1.5 mm, free portion of the filaments 3–5 mm; anthers 8–12 mm long, free, slightly unequal, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, the anther surface smooth to finely papillose. Ovary globose, minutely puberulent with simple glandular trichomes and some stellate trichomes; style of hermaphroditic flowers ca. 10 mm long, glabrous or minutely puberulent in the lower half; stigma capitate, papillose. Fruit a globose berry, 1 per infructescence, 1.2–3.5 cm in diameter when dry, the pericarp thin, smooth, glabrous, greyish brown and chartaceous when mature, when immature light green with dark stripes or marbled pattern, drying dark brown; fruiting pedicels 3–4 cm long, ca. 2 mm in diameter at base, 3–4 mm at apex, woody, strongly deflexed, sparsely armed with straight yellowish red prickles or unarmed, channelled in dry specimens; fruiting calyx accrescent, to 1.5 cm long, the lobes covering 2/3 or the entirety of the mature fruit, sparsely prickly. Seeds ca 10–30 per berry, 5–6 mm long, ca. 2.5 mm wide, flattened, reniform-rounded, brown, the surface minutely pitted and somewhat warty. Chromosome number: n=ca. 24 (
(Figure
None known.
(
Based on
Lectotype of
Shrub, 0.2–1.5 m tall. Stems erect, sparsely to densely armed, or unarmed; young stems densely stellate-pubescent; trichomes porrect, translucent and often reddish gold, sessile to subsessile, the rays 6 to 9, 0.1–0.3 mm long, the midpoints ca. 0.1 mm long, the prickles irregularly distributed throughout the plant, sometimes more dense on the calyx and pedicels, 1–10 mm long, 0.5–1.5 mm wide at base, straight, brown or reddish, sometimes yellow on the young stems, spaced 0.1–1 cm apart if dense, 1–10 cm apart if sparse; bark of older stems brown, glabrescent. Sympodial units usually difoliate, but not markedly geminate, sometimes plurifoliate. Leaves simple, variously lobed, 3–7(10) cm long, 1.5–3(5) cm wide, variable in shape, ovate to elliptic to broadly elliptic, 1.5–3 times longer than wide, mostly concolorous, drying yellowish green to green, densely stellate-pubescent adaxially and abaxially, the trichomes porrect, translucent, sessile to subsessile, the stalk up to 0.1 mm long, the rays 7 to 9, (0.1) 0.2–0.4 mm long, the midpoints 0.1–0.2 mm long; principal veins 3–6 pairs, raised abaxially, flat adaxially, spreading at ca. 45° to the midvein, the tertiary venation mostly not visible to the naked eye; base obtuse, rounded or truncate, sometimes oblique; margins variously lobed, rarely entire, the lobes 2–5 on each side, 0.1–1.5 cm long, usually rounded, rarely obtuse, the sinuses extending up to 1/3–1/2(2/3) of the distance to the midvein; apex rounded to acute; petiole 1–4 cm long, 1/2–1/10 of the leaf length, densely stellate-pubescent like the young stems. Inflorescence terminal or lateral, 4–7(15) cm long, usually unbranched but occasionally forked, with 4–9(26) flowers, usually with one hermaphrodite flower at the base, all distal flowers staminate; peduncle 0.5–2 cm long; rachis 1–6(9) cm long; pedicels 0.4–1 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely or sparsely armed only in hermaphroditic flowers, unarmed in staminate flowers, articulated near the base; pedicel scars irregularly spaced 2–15 mm apart, prominent and brown. Buds strongly curved and zygomorphic, more so in staminate flowers; the corolla exserted ca. halfway from the calyx tube before anthesis. Flowers 5-merous, heterostylous, heterandrous and markedly dimorphic, the plants strongly andromonoecious, the basal flower long-styled and hermaphroditic, the distal flowers short-styled and staminate. Calyx tube 0.8–2 cm long, cup-shaped, the lobes 0.7–2.5 cm long, 1–2 mm wide, long-triangular with an elongate acumen, unarmed in staminate flowers, densely armed in hermaphroditic flowers, the prickles 0.5–1 cm long, 0.5–1 mm wide at base, yellow or reddish, straight, spaced 0.5–2 mm apart. Corolla 2–4 cm in diameter (slightly larger in staminate flowers), lilac or purple or occasionally white, the midvein paler and greenish yellow, drying yellow or brownish tan in herbarium specimens, stellate to broadly stellate, lobed ca. halfway to the base, the lobes 1–1.5 cm long, 0.8–1 cm wide, spreading or slightly reflexed at anthesis with abundant interpetalar tissue, the abaxial surfaces densely pubescent along the middle portions where exposed in bud, the adaxial surfaces glabrous or with a few trichomes along the margins, the tips somewhat cucullate. Stamens strongly unequal in both hermaphroditic and staminate flowers, with the 2 adaxial anthers shorter than the 3 abaxial anthers; filament tube minute; free portion of the filaments 1.5–2 mm long, glabrous; anthers tapering, yellow or occasionally purple (Sinaloa and Sonora), the surfaces smooth, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, directed distally, in hermaphroditic flowers three longer abaxial anthers ca. 9 mm long, two shorter adaxial ca. 6 mm long, in staminate flowers three curved abaxial anthers 15–20 mm long, two straight adaxial anthers ca. 10 mm long. Ovary in hermaphroditic flowers ca. 2 mm in diameter, conical. porrect-stellate pubescent in the distal third, the trichomes sessile to subsessile, with 7–10 rays, the rays 0.1–0.2 mm long, the midpoints 0.1–0.15 mm long; ovary in staminate flowers vestigal; style 20–30 mm long in long-styled flowers, in staminate flowers ca. 1.5 mm long; stigma capitate and slightly bilobed in hermaphroditic flowers, indistinctly bilobed and much smaller in staminate flowers. Fruit a globose dehiscent berry, (1-)2–2.5 cm in diameter, pale green with darker green mottled areas when young, dark brown or orange-brownish when dry, dehiscing irregularly in several parts, usually breaking open irregularly in the upper half of the fruit, the pericarp thin, smooth, glabrous; fruiting pedicels 0.5–1.5 long, 2–3 mm in diameter at base, 2–5 mm in diameter at apex, erect, herbaceous to woody; fruiting calyx (hermaphroditic flowers only) strongly accrescent, the lobes covering up to the total length of the fruit, sparsely or densely armed with prickles up to 8 mm long. Seeds ca. 20–40 per berry, 3–4 mm long, 3–4 mm wide, flattened reniform, dark brown to black, the surface minutely pitted, the testal cells pentagonal. Chromosome number: n=12 (
(Figure
Distribution of
Mexico. Colima: mala mujer; Puebla: diente de burro; Sinaloa: melón de coyote, coyotillo; Sonora: sacamanteca; Veracruz: berenjena espinosa, necachane; Yucatán: berenjena de monte, k’omya’ axnik (Maya).
The leaves of
(
This species was long known as either
The protologue of
The lectotype we have selected for
Of the two varieties described in
Of the two sheets of
Argentina. Catamarca: Dpto. Capayán, San Pablo, between Concepción and Huillapima, 700 m,
Erect rhizomatous shrub, to 1 m tall. Stems erect, woody, armed or unarmed; young stem densely stellate-pubescent, the trichomes multangulate, translucent, short-stalked, the rays 10–12, ca. 0.5 mm long; prickles if present 5–6 mm long, needle-like and straight, pale yellowish brown; bark smooth, brown or yellowish brown from persistent pubescence. Sympodial units difoliate, not markedly geminate. Leaves simple, (2-)4–9 cm long, (1-)2–4 cm wide, elliptic, ca. 3 times longer than wide, discolorous, drying yellowish green to greyish green; adaxial surfaces densely stellate-pubescent but the leaf blade tissue visible, the trichomes multangulate, translucent, sessile, the rays 10–12, ca. 0.5 mm long; abaxial surfaces more densely pubescent with similar multangulate trichomes; principal veins 5–7 pairs, impressed adaxially, flat abaxially, spreading at ca. 45° to the midvein, the tertiary venation usually visible in dry material; base truncate to slightly cordate, often somewhat oblique; margins shallowly lobed, the lobes 4–7 on each side, of varying sizes, becoming smaller towards the leaf apex, the sinuses extending only 1/4 or less of the distance to the midvein, triangular; apex acute to somewhat rounded, rarely obtuse; petiole 1–2 cm long, densely stellate-pubescent like the young stem, unarmed. Inflorescences terminal or lateral, to 6.5 cm long, with up to 10 flowers, unbranched; peduncle 1.5–3 cm long, densely pubescent with multangulate trichomes like those of the stems; pedicels 0.4–1 cm long, ca. 1.5 mm in diameter, robust, articulated less than 0.5 mm from the base, densely pubescent like the leaf blade; pedicel scars prominent, spaced ca. 0.5 cm apart. Buds turbinate, the corolla strongly exserted from the calyx tube prior the anthesis. Flowers 5-merous, strongly heteromorphic and the plants andromonoecious, only the basal flower perfect (hermaphroditic), the distal flowers functionally staminate and short-styled. Calyx conical or cup-shaped, the tube 5–6 mm long, strongly keeled, the lobes 7–10 mm long, ca. 1.5 mm wide at base, subulate, densely pubescent abaxially with multangulate trichomes. Corolla 3–3.5 cm in diameter, pentagonal, pale lavender, drying pale brown, barely lobed, interpetalar tissue abundant, the lobes ca. 0.2 cm long, ca. 0.1 cm wide, mere acumens, densely pubescent with multangulate trichomes abaxially along the midveins and surfaces exposed in bud. Stamens equal or very slightly unequal and the adaxial anthers slightly shorter; filament tube ca. 0.5 mm, free portion of the filaments ca. 2 mm; anthers 7–12 mm long, free, occasionally very slightly unequal, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, the anther surface smooth to finely papillose. Ovary globose, glabrous or minutely glandular puberulent; style of hermaphroditic flowers ca. 1 cm long, glabrous; stigma capitate, papillose. Fruit a globose berry, 1 per infructescence, ca. 2 cm in diameter when dry, the pericarp thin, smooth, glabrous, light green, when immature with dark stripes or marbled pattern, drying dark brown or brown-reddish and shattering; fruiting pedicels 1–1.7 cm long, ca. 1.5 mm in diameter at base, ca. 2.5 mm at apex, woody, erect, sparsely armed with straight yellowish red prickles or unarmed; fruiting calyx somewhat accrescent, to 2 cm long, the lobes covering up to 1/3 of the mature fruit, usually unarmed, if prickly the prickles needle-like and straight. Seeds ca. 40 per berry, 4–5 mm long, 3–4 mm, flattened, reniform-rounded, black, the surfaces minutely pitted. Chromosome number: not known.
(Figure
Like some other members of the group,
None recorded.
(
Type. “Habitat in Ahualulci montibus” [México. Tabasco: Ocuapan, 17°51'N, 93°29'W, or San Luis Potosí: Ahualulco, 22°24'N, 101'10"W],
We thank the curators of herbaria cited in the text for permission to examine or borrow the specimens in their care; Sarah Ficinski for assistance in data recording and for preparing the maps; Andrea Cocucci for sharing information about bee pollinators; Bobbi Angell, Nidia Flury, Laura Ribulgo and Leonor Sánchez for line drawings of the taxa treated here, and the Flora of Argentina editorial committee for allowing us to use those that had been previously published; the Université de Montpellier for permission to reproduce the drawing held in their care (MPU310713); Eugene Sturla for allowing to use his photographs of
Only first collectors in collections made by two or more collectors are listed here; pairs of collectors are listed in the Specimens examined section, while groups of three or more are listed as
Abd El Ghani, M. 6908 (
Abrams, L. 6607 (
Abulaila, K. JOR3-1 (
Acevedo, R. 32 (
Ackermann, M. 218 (
Acosta, L.E. 21 (
Acosta, M. 663 (
Aguayo C, J.A. 2 (
Aguiar H, H. 59 (
Aguilar C, A. 164 (
Aguilar Zepeda, J.A. 344 (
Aguilar, L.M. 912 (
Aguilar, R.M. 425, 555 (
Aguirre T., S. 45, 116, 116, 151, 259 (
Agúndez, J. 132, 297 (
Ahles, H.E. 56834 (
Alba, A. de 45 (
Alcorn, J.B. 2028 (
Altamirano, F. 1678 (
Álvarez, R. 401, 499, 573, 631, 989 (
Ambrosetti, J.A. 1418, 1472, 1480, 1481, 1483 (
Anderson, W.R. 3504 (
Araque M., J. 19Ar341 (
Arenas, P. 2360 (
Argañaras, J.L. 51 (
Argüelles, E. 23, 2468 (
Ariza Espinar, L. 1575, 2393, 3587, 3588b (
Armenta, P.M.A. 132 (
Arnoldo-Broeders, M. 3621 (
Arsène, G. 10632a (
Arteaga Saucedo, M.C. 840 (
Articó, L. 241, 244, 250 (
Atwood, N.D. 11023, 16071, 28852 (
Austin, S.B. 277 (
Bacigalupi, R. 2663 (
Bailey, L.H. 21 (
Baines, R. 255 (
Baird, G.I. 1818 (
Baker, M.A. 7568, 7581 (
Baker, M.S. 11244, 11574 (
Balegno, B. 249 (
Balls, E.K. 10857 (
Barbour, M. 119 (
Barboza, G.E. 76, 220 (
Barclay, G.W. 3079 (
Barkley, F.A. 14A539, 783 (
Baro, D. 495 (
Barr, R.J. 61-121 (
Bartholomew, B. 2430 (
Bartlett, H.H. 10991, 19760, 19871, 19886, 19905 (
Beatley, J. 9028 (
Beetle, A.A. 26132 (
Bell, W.M. 48 (
Benedeto, ? 235 (
Benítez-Paredes, A. 1814 (
Bennett, H.R. 8486, 8659 (
Berg, C.C. 153 (
Bernardello, G. 449, 503, 522, 534, 746, 754, 788 (
Berro, M.B. 4411, 7942 (
Berry, E.O. 19, 34 (
Biloni, J.S. 175M (
Biltmore Herbarium 3438 (
Bisschoff, J. 1 (
Biurrun, F. 2103, 2729, 7225 (
Blakley, E.R. 3-1652 (
Blanchet, J.S. 6410 (
Blumer, J.C. 1732 (
Bodenbender, W. 55 (
Boege, W. 689 (
Boelcke, O. 1412, 11691 (
Bojórquez B., G. 721 (
Borsini, O.H. 1013, 1201, 1333 (
Botello, L. 14 (
Boulos, L. 18198 (
Bowen, J. 27 (
Boyd, S. 5335, 5539, 5778, 5956 (
Bradburn, A.S. 1134 (
Braunton, E. 697 (
Breedlove, D.E. 1150 (
Brigada Zacatecas 86 (
Brizuela, A. 1181 (
Brizuela, J. 847, 1159 (
Brother Anect 115 (
Broome, C.R. 313 (
Bruno, R. 4 (
Burge, A. 25 (
Burkill, I.H. 432 (
Búrquez, A. 96-15, 94-017 (
Bush, B.F. 266, 392, 1588 (
Butterwick, M. 7782, 8217 (
Bye, R. 28334 (
Cabrera, A.L. 7007, 30038, 30771, 31065 (
Cabrera, E. 4658, 8272, 8658, 8834, 8895, 8962, 9130, 9225, 9428, 9631, 10064, 10207, 10286, 10461, 10755, 11000, 11289, 11465, 11530, 11566, 11625, 11686, 11770, 12835, 13037, 13210, 13635, 13734, 13920, 14204, 14318, 14535, 15400, 15787, 17207 (
Cadena, B. 23 (
Calhoun, D. 129 (
Calzada, J.I. 1837, 3182, 6459, 7725 (
Campos, G. 2800, 2836 (
Cancino, J. 20, 55, 123 (
Canedo, F. 9075 (
Cano R, P. 21 (
Cantero, J.J. 6183 (
Cantino, P.D. 505 (
Carette, E. 22 (
Carleton, M.A. 171 (
Carnegie, F.G. 40 (
Carranza, E. 3292 (
Carrasco, E.L. 151 (
Carrillo-Reyes, P. 1974 (
Carter, A. 1892 (
Castañeda V, ? 64, 68 (
Castellanos, A. 90 (
Castello, L.V. 232 (
Castillo Campos, G. 189, 700, 841, 6971, 18152 (
Castillón, L. 1030, 1628 (
Cervantes M., H. 19 (
Chambers, K.L. 720, 771 (
Chan, C. 45, 100, 1827, 2689, 3166, 3983, 4835 (
Chapiro, G. 41 (
Charlton, D. 3259 (
Chaudhary, S.A. E455 (
Chavelas P., J. ES-3971 (
Chávez Montes 82 (
Chehaiber, T. 201 (
Chiapella, J. 1804, 2175, 2186 (
Chiarini, F. 40, 462 (
Christ, J. 991 (
Cirujano, S. R-10431 (
Claraz Schenkung, A. 81 (
Clark, W.H. 3289, 3304 (
Clausen, J. 595 (
Clausen, R.T. 4597 (
Clokey, I.W. 5937 (
Clover, E.U. 165 (
Cochrane, T.S. 8516 (
Cocucci, A.A. 220, 369, 439, 440, 977, 1011, 1014, 1364, 3848, 5014, 5041, 5760 (
Cocucci, A.E. 14 (
Colunga, P. 155 (
Compling, M. 9 (
Constance, L. 3116 (
Copp, J.F. 70-7 (
Cory, V.L. 52468 (
Cosa, M.T. 44, 108, 115, 124, 162, 182, 186 (
Coulter, T. 1246, 1247, 1248 (
Crampton, B. 7136 (
Croat, T.B. 39945 (
Cronemiller, F.P. 3061 (
Crosswhite, F.S. 720 (
Cruden, R.W. 1174 (
Cruz Durán, R. 4737, 5483 (
Cuesta, L.R. 57 (
Cuezzo, A.R. 872, 898, 1058, 1302, 2227, 2254 (
Cuming, H. 169, 1090 (
Curtiss, A.H. 5913 (
D’Arcy, W.G. 44A, 1974, 4861, 11689, 12027 (
D´Angelo, C. 285 (
Daniel, T.F. 6784, 6827 (
Daveau, J. 2260 (
Davidse, G. 9252 (
Davis, P.H. 66299 (
Davis, T. 677, 1000 (
Deaver, C.F. 6439 (
Del Vitto, L.A. 850, 3558 (
Dells, A. 3247 (
Demaree, D. 7674, 10857, 17916 (
Di Fulvio, T.E. 799, 823, 825, 831, 941, 942, 949, 1110, 1112, 1126 (
Díaz Luna, C.L. 4141 (
Díaz Vilchis, I. 124, 438 (
Dinelli, E. 654 (
Dixon, D. 3636 (
Doebley, J. 203 (
Domingo, M.V.Z. 524 (
Domínguez Cadena, R. 1237 (
Dorantes, A. 34 (
Dorantes, J. 251, 260, 827, 1033, 1127, 1224, 1333, 1416, 5209, 5260 (
Dottori, N.M. 138, 139, 145, 146, 148, 169, 179, 180, 186 (
Drouet, F. 3695 (
Drummond, J.R. 93, 266 (
Dullas, W. 135 (
Dunn, D. 23177 (
Dunnett, D. 13 (
Durán, R. 1581, 1626B, 3328 (
Dusén, P.K.H. 5217 (
Dwyer, J.D. 14151 (
Eastwood, A. 1040, 5958, 8550, 8627, 9301A, 11763, 13946, 15419, 15646, 17097, 18099 (
Eaton 57 (
Eggleston, W.W. 7424 (
Eggli, U. 1958 (
Ehlers, J.H. 6410 (
Ekman, E.L. 13942 (
El Ghani, M.A. 6908 (
Elizondo, J.L. 319 (
Elliot, B. 11929c (
Elliot, C. 62 (
Els, P. 3118 (
Encarnación, R. 32 (
Enríquez E., E.D. 114, 465 (
Enríquez, O.G. 98, 686 (
Equipo 5 (
Ertter, B. 2939 (
Escalante R., S. (
Escalante, S. 240, 549, 672, 761, 854 (
Escobedo, J.M. 928, 1594 (
Espejel, I. 99 (
Espejo, A. 1023 (
Ewan, J.A. 15, 680 (
Eyerdam, W.J. 23093, 23735 (
Fabris, H.A. 8357 (
Farnsworth, E.L. 310 (
Felger, R.S. 86-400, 85-795 (
Feliciano Kú y Yam, Q.B.A. 203, 254, 626, 751 (
Felippone, F. 5092, 6166 (
Fendler, A. 672 (
Fernández Casas, F.J. 11029, 11162 (
Fernandez, J.R. 156 (
Ferrari, R.V. 313 (
Ferris, R.S. 1153, 2302, 2720, 3133 (
Ferrucci, M.S. 2953, 2979 (
Fertig, W. 22568 (
Filippa, E.M. 79 (
Fischer, W. 238 (
Fisher, G.L. 5089 (
Flores F., G. 2122 (
Flores Martínez, A. 1328, 2037, 4751 (
Flores, J.S. 8431, 8819 (
Flores, R.M. 2 (
Flores-Franco, G. 400, 421, 512 (
Floyer [Mrs] 1 (
Fortuna, J. 22 (
Fortunato, R.H. 9916, 9942 (
Fosberg, F.R. S-3399 (
Foster, R. 3935 (
Frenkel, R.E. 841A (
Fritsch, P. 1325 (
Frye, T.C. 2383 (
Fryxell, P.A. 3706 (
Fuller, T.C. 8567, 9706, 9796, 11226, 19758 (
Gaitón Hinojosa, M.A. 2013-05 (
Galeotti, H.G. 1160, 1168 (
Galetto, L. 24, 214, 232, 237, 239, 260 (
Gallegos Harking, F. 81 (
Galván, M. 618 (
Galván, R. 2761, 3614 (
García Bielma, M.A. 536 (
García E., J.D. 17 (
García M., E. 322 (
García P., J. 670 (
García R., L.A. 1181 (
García Torres, F. 34, 78, 105, 148, 183, 315 (
García, A. 40 (
García, E. 85 (
Garcia, E.M. 413 (
García, R.J. 23 (
García-Mendoza, A. 9016 (
Gardner, M.F. 58 (
Gaumer, G.F. 135, 366, 1708, 23325, 23622, 24060 (
Gaviño, G. 13 (
Gay, C. 834 (
Genelle, P. 819 (
Gentry, A. 48912 (
Gentry, H.S. 357 (
Germán, M.T. 207 (
Ghiesbreght, A.B. 120 (
Giacomelli, A. 88900 (
Gillespie, J.W. 5222 (
Gillett, J.M. 17013 (
Gillies, J. ‘Solanum2’, 28, 669 (
Gilmartin, A.J. 1677 (
Giorgis, M. 693 (
Glass, V. 10674 (
Goddard, D.R. 836 (
Gold, D.B. 365 (
Gómez Lorence, F. 3 (
Gómez, A. 3, 13 (
Gómez-Pompa, A. 1857 (
Góngora, E. 59, 301, 694 (
González E., M. 13B.C. (
González G., J. 133 (
González H., J.E. 267 (
González Medrano, F. 8003, 8013 (
González Ortega, J. 56, 425, 716, 5609, 6578, 6743, 6805, 7249 (
González, M. 10 (
González-Medrano, F. 39, 115 (
Goodman, G.J. 1142 (
Graham, E.H. 3021, 3619 (
Grant, G.B. 5550 (
Griffiths, G. 216, 267 (
Gross, C.A. 82 (
Guaglianone, E.R. 1555, 1571 (
Guerra, O. 601 (
Guízar N., E. 4031 (
Gullion, G.W. 383 (
Gust, G. 2206 (
Gutiérrez B., C. 436, 723, 977 (
Guzmán C., U. 2121 (
Guzmán, C.A. 11, 44 (
Guzmán, R. 999 (
Haene, E. 61 (
Hahn, W. 1420 (
Hall, E. 498 (
Halse, R.R. 568 (
Hammerly, B.J. 75, 216 (
Hanekom, W.J. 554 (
Hannington, C. 23 (
Hansen, B. 1392 (
Hansen, C.J. 4421 (
Hansen, G. 1174 (
Hardham, C.B. 2628, 2804 (
Harrington, H. 7966 (
Hartman, C.V. 243 (
Hartman, R.L. 65234 (
Hartweg, K.T. 203 (
Harvey, D.R. 585 (
Hastings, J.R. 64-51 (
Hawkes, J.G. 1166, 1463 (
Helberg, J.J. 2528 (
Heller, A.A. 1511, 3733, 4185 (
Henderson, D.M. 6848 (
Henrickson, J. 5708, 6673a, 7444, 7917 (
Hernández A., C. 66 (
Hernández A., F. 567 (
Hernández Magaña, R. 207 (
Hernández X., E. 677 (
Hernández, A.C. 66 (
Hernández, L. 1168, 1174 (
Herter, W.G. 7290 (
Hespenheide, H.A. 8 (
Hick, P.M.L. 22, 30, 33 (
Hieronymus, G. 288 (
Higgins, L.C. 14434, 20971, 26756 (
Hill, M. 225 (
Hmama, H. 890 (
Hopkins, M. 897 (
Horr, W.H. 3521 (
Hosseus, C.C. 65, 210, 233, 500, 1259, 2007, 2050, 2097, 2638 (
Howell, J.T. 10610, 10611, 10643 (
Huckins, C.A. 6912-3002 (
Huidobro, A.M. 3514 (
Hunziker, A.T. 20, 1030, 3252, 15924, 16035, 17347 (
Hunziker, J.H. 66, 280, 347, 4097 (
Hutchison, P. 701 (
Ibarrola, T.S. 151, 2955 (
Igic, B. 07s20 (
Illanes, A. 104 (
Ingram, J. 650 (
Instituto de Investigaciónes Ecológicas Chiloé 502 (
Irvine, F.R. 276 (
Isern, J. 8096 (
Iwen, F.A. 355 (
Jafri, S.M.H. 3737 (
Jiménez, J. 426 (
Johansen, D.A. 551, 615 (
Johnson, D. 76, 130 (
Johnson, R.L. 12-029 (
Johnston, I.M. 2158 (
Johnston, M.C. 256, 10443 (
Jones, M.E. 4119 (
Jörgensen, P. 1070 (
Joyal, E. 1339, 1423, 1520 (
Juliani, H.R. 34, 35, 36, 37, 39 (
Jury, S.L. 8776, 20869 (
Jussel, M.S. 69 (
Kadhum, H. 37772 (
Karwinski, W.H. 583bis (
Keck, D.D. 2507 (
Kelley, H. 21 (
Kiesling, R. 1049, 3006, 4241, 4652, 4827 (
King, D.O. 148 (
King, R.M. 2669, 2701 (
Kipping, J. 308 (
Kishler, J. 145, 238, 579 (
Knapp, S. IM-10078, 10470 (
Knobloch, I.W. 70 (
Koch, S.D. 7493 (
Kolberg, H. 925 (
Krapovickas, A. 5950 (
Kriebel, R. 8208 (
Krishnappa, D.G. 162 (
Kugel, A.R. 2080 (
Kuhn, B. 2605 (
Kurtz, F. 209, 911, 2604, 4222, 4224, 4744, 5117, 5209, 7901, 7993, 8034, 8086, 8945, 9503, 13117, 14206, 14406, 15732, 16036, 16090, 16090a (
La Barrera, J.M. de 18, 20 (
La Seta, A.V. de 255 (
La Sota, A. de 132 (
La Torre, C. de 47 (
Lajtha, K. 123 (
Lamond, J.M. 1399 (
Landrum, L.R. 5409 (
Lanfranchi, A.E. 35, 1315 (
Langman, I.K. 2915 (
Laukkonen, P. 333 (
Leal, J. 29, 210 (
Legler, B. 10779 (
Legname, V. 221 (
León de la Luz, J.L. 10161, 10591 (
Leonard, S.W. 3394 (
Leuenberger, B. 2889 (
Leuenberger, B.E. 4459 (
Levin, G.A. 2159 (
Linden, J.J. 243 (
Lindheimer, F.J. fasc. I-135, III-135, 1041 (
Liogier, A.H. 9732 (
Lira Charco, E.M. 1379 (
Lizama M., J. 1479 (
Long, S. 1100 (
López, ? 713 (
López, L.E. 85 (
López Felix, ? 16 (
López-Forment, W. 292, 422 (
López Franco, R.M. 974, 1100 (
Lorentz, P.G. 9, 98, 99, 106, 320, 533, 905, 908, 1243 (
Lossen, W. 12 (
Lot, A. 454, 483, 604, 633, 646, 1877, 1922 (
Lott, E.J. 73 (
Lourteig, A. 1019 (
Luchetti, A.M. 4, 5, 6 (
Luckow, M. 2837 (
Luna M., V.E. 36, 149, 262 (
Luna, F.E. 1021 (
Lundell, C.L. 5145 (
Luz, J.L. de la 6121 (
MacDaniels, L.H. 443 (
MacDonald, J.R. 7551 (
MacKee, H.S. 10974 (
Maillet, J. 14 (
Makings, E. 1102 (
Mandon, E. 1010 (
Manning, W.E. 531244 (
Manríque, ? 1255 (
Marais, W. 28703 (
Marcus, E.J. 26245 (
Marín, R. 150 (
Marino, G. 201, 1914, 1942, 2047 (
Marquez, M. 1095 (
Marsh, E.G. 895 (
Martin, H.C. 272 (
Martínez Marín, J.L. 416, 464 (
Martínez O., E. 413 (
Martínez R., C. 104 (
Martínez S., E.M. 4083, 39209 (
Martínez, C. 1796 (
Martínez, G.J. 314, 486, 524, 1321 (
Martínez, J.L. 683 (
Martínez, M. 861 (
Mason, H.L. 1903 (
Matesevach, M. 3 (
Mathews, A. 238 (
Matuda, E. 18953, 19545, 28754 (
Mayfield, M.H. 301 (
McClatchie, A.J. 1282 (
McDaniel, T.F. 1856 (
McGregor, E.A. 857 (
McKee, ? 10974 (
Mearns, E.A. 122, 135, 619, 1698 (
Medina A, M.E. 24 (
Medina C., M. 2036 (
Medina, V. 241 (
Medina, W.A. 57 (
Meglioli, S. 33 (
Méndez, M. 261, 812 (
Mereles, F. 4010, 5109, 9095 (
Merello, M. 250 (
Mericle, L.W. 500 (
Mertz, S.L. 148, 205 (
Mertz, S.M. 15 (
Metz, M.C. 3164 (
Meyer, T. 565, 3506, 4046, 4067, 9863 (
MFB 53 (
Michener, D.C. 4253 (
Miehe, G. 277 (
Miers, J. 258 (
Miller, J.S. 7291 (
Miranda, F. 1418 (
Miranda, R. FESC-RE-38 (
Mooers, B.H.M. 854 (
Molinar, R. 47 (
Moncada, F. 4741 (
Monroy M., R. 4 (
Montenegro, M.A. 380 (
Montero Castro, J.C. 136 (
Moore, H.E. 1640 (
Mora-López, J.L. 189 (
Moran, R. 3518, 3603, 3740, 3779, 7218, 8030, 9266, 11411, 22371 (
Moreno, P. 150, 494, 794 (
Morong, T. 1190 (
Moscone, E.A. 34, 70, 71, 109, 111, 161, 180, 183, 191, 245 (
MRVG 503 (
Mulford, A.I. 684 (
Muller, F. 182 (
Nagahama, N. 129, 130, 131, 132 (
Naranjo, C. 937 (
Narváez C., J. 2 (
Narváez, M. 339, 590 (
Navarrete de la Paz, M. 33 (
Nee, M. 11787, 11815, 18972 (
Nelson, A. 1651, 11558 (
Nelson, E.W. 6429 (
Nevling, L. 544, 1043 (
Newman, M. 240 (
Nicora, E.G. 1646, 2101, 4162, 4189, 4282, 8114 (
Nixon, K.C. 838, 893 (
Norris, D.H. 20104 (
Novara, L.J. 8248, 8504 (
Novoa L., C.P. 54A (
O’Donell, C.A. 224, 1712, 1815, 1910 (
Ocampo, G. 874 (
Ocampo, R. 39 (
Ochoa Avalos, S. 188 (
Orbigny, A. d’ 223 (
Orcutt, C.R. 680, 1270 (
Orea L., L. 153, 206, 274, 407, 440 (
Orellana, R. 119BB (
Orozco H, J. 9957, 10757 (
Ortega O., R.V. O-720 (
Ortega, R. 427, 708 (
Osterhout, G.E. 2064 (
Oswald, P.H.O. 15 (
Palacios, ? 1695 (
Palmer, E. 25 (
Panti, M.A. 841 (
Parish, S.B. 8104 (
Parish, W.F. 172 (
Parker, H.M. 379 (
Parry, C.C. 536, 636 (
Paula-Souza, J. 7856, 8046 (
Pedersen, T.M. 5789, 5790, 8235 (
Peebles, R.H. 7410 (
Peña-Chocarro, M.C. 407, 1076 (
Pensiero, J.F. 2424, 6063, 7209, 7374 (
Pérez C., E. 3359 (
Pérez H., S. 86 (
Pérez Lara, M.A. 68, 94, 186, 612, 761, 860, 892 (
Pérez S., G. 318 (
Perkins, A.E. 3377 (
Perrill, B. 5513 (
Peterson, J. 075 (
Piccinini, ? 2046 (
Pineda C., J.L. 477 (
Plowman, T. 2711 (
Poeppig, E.F. 26, 140, Diar540 (
Popov, G.B. GP69266 (
Price, T. 338 (
Pringle, C.G. 1566 (
Proctor, G.R. 30960, 49664 (
Puch, A. 447, 825, 849, 1319 (
Pujalte, J.C. 210, 259 (
Purpus, C.A. 4548 (
Quick, C.R. 69-15 (
Ragonese, A.M. 6390 (
Ramírez, O.P. 33 (
Ramírez, R. 1050 (
Randolph, D. 173, 394 (
Rangel Ramírez, R. 130, 330 (
Rashid, A. 3692 (
Rasp, A.E. 68 (
Raven, P.H. 12484 (
Reales, A. 952, 968, 1615, 1678, 2015 (
Rechinger, K.H. 29735, 29752, 63645 (
Reddick, D. 157 (
Reed, F.M. 6190 (
Reeder, J.R. 6772 (
Reina G., A.L. 97-297, 2001-598, 98-2082, 98-2123 (
Reko, P.B. 4462 (
Reyes de los Santos, E. 147, 272, 367, 367, 477, 545, 662, 788, 871 (
Reynoso Dueñas, J.J. 634 (
Rico Arce, L. 378 (
Rico-Gray, V. 421 (
Riedel, J. 54 (
Ringuelet, E.J. 255 (
Rivera, R. 90 (
Robles H., L. 397, 398 (
Roderick, W. 3136 (
Rodríguez C., A. 2107, 5076 (
Rodríguez González, H. 143 (
Rodríguez, ? (V)869 (
Rodríguez, E. 42 (
Rodríguez, J.P. 43 (
Rojas, T. 1992, 13678 (
Rollins, R.C. 2749 (
Román Miranda, M.L. C-742, 1337 (
Romero, S. 1405, 1406 (
Rosas J., F. 1 (
Rosas R., M. 876 (
Rose, J.N. 2308, 10073 (
Rose, L.S. 65121 (
Ross, T.S. 6995, 7032 (
Roybal, J.J. 904 (
Rubtzoff, P. 1321 (
Ruiz Huidobro, A.M. 324 (
Ruiz, L. 53398 (
Runyon, R. 4194 (
Ryder, V.P. 248 (
Rzedowski, J. 16116 (
Sagalyn, J.L. 69 (
Salinas T., A. 7104 (
Sallee, E.H. ES-54 (
Sánchez Mejorada R, H. 80 (
Sánchez, E.V. 46 (
Sanders, A.C. 19612, 25026 (
Santana Michel, F.J. 546, 2955 (
Saravia Toledo, C. 2336a (
Sargent, F.H. 513 (
Saucedo, E. 338 (
Sauleda, R. 2649 (
Sayago, M. 570 (
Scaldaferro, M. 21, 25 (
Schaffner, J.G. 695 (
Schallert, P.O. 540 (
Schickendantz, F. 80, 82 (
Schinini, A. 37059 (
Schlottmann, A. 269 (
Schmitz, A. 166[a], 166[b] (
Scholl, M.R. 26 (
Schott, A.C.V. 310 (
Schreiter, R. 12408 (
Schulz, C.L. 1363, 5590 (
Schulz, E.D. 97 (
Schumann, W. 970 (
Schwarz, G.J. 9758 (
Scora, R.W. 2285 (
Scott-Elliot, G.F. 442 (
Selander, R.B. 21, 4-53, 6-53 (
Serrano, A. 128 (
Sessé, M. 1532 (
Sherwin, P. 226 (
Shevock, J. 502, 12209 (
Shreve, F. 6446 (
Silva R., L. 184, 187 (
Simá, P. 38 (
Singh, V. 80328 (
Siqueiros, M. 515 (
Six, I.L. 3409-84 (
Skehan, J. 16 (
Small, J.K. 4841 (
Smith, G.L. 1030 (
Sohmer, S.H. 9372 (
Solbrig, O.T. 140 (
Soriano M., A.M. 144 (
Soriano, A. 548, 1143 (
Soto N. [Soto Nuñez], J.C. 9508 (
Sousa Peña, M. 57, 79, 102, 133,146, 175 (
Sousa S., M. 4836 (
Souviron, M.J. 27 (
Sparrow, C.E.H. 46 (
Spegazzini, P.L. 28174, 33629 (
Spencer, M.F. 1417 (
Spetzman, L. 1366 (
Sprankle, J.A. 96 (
St Pierre, M. 2601 (
Stamatiadou, E. 13340, 15571 (
Stanford, L.R. 273, 586, 802 (
Starlinger, F. 22-93 (
Steere, W.C. 1065 (
Steibel, P. 4092 (
Stephens, S. 10953 (
Stevens, G.W. 802, 889 (
Steyermark, J.A. 83071 (
Stiefkens, L.B. 4 (
Stimson, W.R. 1403 (
Stofella, ? 328 (
Stroh, A.L. 69.63.02 (
Strother, J.L. 258 (
Stuckert, T.J.V. 1239, 1720, 2153, 2409, 2981, 3347, 3510, 4027, 4157, 4300, 4696, 6152, 6517, 7049, 8103, 8335, 8591, 8592, 8689, 9286, 9337, 10520, 11278, 11590, 12070, 15263, 15264, 15745, 16858, 16968, 19448, 19468, 19933, 23189, 23783 (
Suárez Gracida, C.G. 2006-25 (
Subils, R. 2909, 3589b, 3699, 3881, 4414, 4463 (
Sueltenfuse, C. 193 (
Symon, D.E. 14318, 15128 (
Takaki, F. 326 (
Tapia, J.L. 1533 (
Taylor, C.M. 1898 (
Tejero Diez, D. 6256, 6517 (
Téllez V., O. 1576, 3766, 11198 (
Temple, L.C. 5513, 12151 (
Templeton, B. 8644 (
Tenorio L., P. 1122 (
Terribile, M. 808 (
Thackery, F.A. 226 (
Tharp, B.C. 52-78, 51-261 (
Thomas L., E. 13 (
Thomas, J.H. 111 (
Thomas, R.D. 96112 (
Thornber, J.J. 482 (
Thorne, R.F. 58047 (
Tilton, D. 218, 229, 236, 237, 241, 266, 267, 321, 330, 337, 353, 369, 374, 399 (
Tirado, N. 2, 51 (
Tolstead, W.L. 5859 (
Torke, K.J. 381 (
Torrecillas, E. 208 (
Torres C., R. 3141, 3247, 3561, 3939 (
Torres, P. 21 (
Torriglia, M. 327 (
Toumey, J.W. 398 (
Tracy, S.M. 7576 (
Traverse, A. 802 (
Trejo, L. 13, 53 (
Troncoso, N.S. 6030 (
Tun, F. 418, 443 (
Turland, N.J. 671 (
Tweedie, J. 29, 70 (
Twisselmann, E.C. 1266, 2248, 4791, 4918, 9088, 10232, 14887, 15599 (
Ucán Ek, E. 277, 394, 1249, 1826, 2332, 3021, 3451, 4629, 4635 (
Uh Cauich, R. 4 (
Unsicker, J. 134 (
Urdampilleta, J.D. 583 (
Valdés, J. 55-2, 15-1978 (
Valiente Banuet, A. 666 (
Valov, D. 2004-218 (
Van Devender, T.R. 2005-10 (
Vanni, R.O. 2620 (
Vara, A. 157 (
Varela E., L. 96-184, 96-348, 97-01 (
Varela, F. 713 (
Varela, F.J. de 585 (
Vargas, C. 68 (
Vasey, G.R. 351 (
Vázquez Yanes, C. 653 (
Vázquez, J. 2164 (
Vega Aviña, R. 267, 1467, 1595, 9884 (
Ventura A., A. 1361, 1509, 1595 (
Ventura A., F. 3385, 10560, 11718 (
Ventura, E. 6112, 6812, 8022, 8329 (
Ventura, M.A. 127 (
Venturi, S. 63, 1091, 7132, 7678, 9882 (
Vilcapoma, G. 520, 540 (
Villafañe, A. 937, 1028, 1265 (
Villafañe, M. 36, 903, 993, 1234, 1271 (
Villalobos-Mejía, ? 254 (
Villaneuva G., R. 11 (
Villarreal de Puga, L.M. 11507, 16135 (
Villegas F., E. 219, 329 (
Vine, R.S. 81 (
Volponi, C.R. 240, 241 (
Wagner, R.J. 1559, 1646, 1880 (
Ward, D.B. 5742, 5771 (
Wawra, H. 673 (
Way, M.J. WSSB-3 (
Weaver Jr., R.E. 2044 (
Webster, G.L. 15557, 17551 (
Welsh, S.L. 25320 (
Werdermann, E. 142 (
Whalen, M.D. 13, 16, 182 (
Wheeler, H.-.M. 5859 (
White, D. 96 (
White, L.D. 6 (
White, S.D. 4526 (
White, S.S. 183, 1180, 1712, 2052, 2526, 2574 (
Whiting, A.F. 905 (
Whitson, M. 803, 817 (
Wickens, G.E. 828 (
Wiggins, I.L. 78 (
Wilbur, R.L. 2469 (
Wilczek, C. 239 (
Williams, M.J. 82-26-1, 79-176-1 (
Wisura, W. 4843 (
Wittman, R.C. 18438 (
Wolf, C.B. 2485, 2558, 3086 (
Wooton, E.O. 59, 3127 (
Worth, C.R. 8815 (
Wright, C. 1590 (
Wright, W.R. 1289 (
Wynd, F.L. 76, 780 (
Xantus de Vesey, J. 84 (
York, C.L. 52-300 (
York, H.H. 49 (
Zamora C., P. 143, 282, 541, 598, 634 (
Zamudio, S. 2097, 2212 (
Zanoni, T. 33168 (
Zardini, E.M. 36391 (
Zohner, O. 34 (
Zola B., M.G. 578, 1584, 2135, 2179 (
Zuloaga, F.O. 9373 (