TYPE Review

PUBLISHED 14 April 2023

DOI 10.3389/fgene.2023.1080176

Taxonomic revision of Physalis

OPEN ACCESS in Mexico
EDITED BY
Rocio Deanna,
University of Colorado Boulder, Mahinda Martínez 1,2, Ofelia Vargas-Ponce 3,4* and
United States Pilar Zamora-Tavares 3,4
REVIEWED BY 1
Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Querétaro, Mexico, 2Laboratorio
Gloria E. Barboza,
Nacional de Identificación y Caracterización Vegetal, Querétaro, Mexico, 3Instituto de Botánica,
CCT CONICET Córdoba, Argentina
departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias,
Milo Pyne,
Universidad de Guadalajara, Guadalajara, Jalisco, Mexico, 4Laboratorio Nacional de Identificación y
University of North Carolina at Chapel
Caracterización Vegetal, Guadalajara, Mexico
Hill, United States

*CORRESPONDENCE
Ofelia Vargas-Ponce,
ofelia.vargas@gmail.com
Physalis (Solanaceae, Solanoideae) is an American genus of ca. 90 species, with its
SPECIALTY SECTION
This article was submitted to Plant diversity centered on Mexico. We recognize 61 species within the country, for
Genomics, a section of the journal which we provide a generic morphological description, an artificial key to
Frontiers in Genetics
determine species, and brief descriptions. We include distributions, habitats,
RECEIVED 25 October 2022 diagnostic characters, phenology, and uses. Distribution maps and field
ACCEPTED 27 February 2023
PUBLISHED 14 April 2023
photographs are also provided. We include conservation status as evaluated by
the IUCN.
CITATION
Martínez M, Vargas-Ponce O and
Zamora-Tavares P (2023), Taxonomic KEYWORDS
revision of Physalis in Mexico.
Front. Genet. 14:1080176. revision, artificial keys, conservation, descriptions, distribution, uses
doi: 10.3389/fgene.2023.1080176

COPYRIGHT
© 2023 Martínez, Vargas-Ponce and
1 Introduction
Zamora-Tavares. This is an open-access
article distributed under the terms of the Physalis (Solanoideae, Solanaceae) has been the subject of a large amount of research to
Creative Commons Attribution License
determine its generic limits (Whitson and Manos, 2005; Zamora-Tavares et al., 2016), to
(CC BY). The use, distribution or
reproduction in other forums is address taxonomic problems (Pretz and Deanna, 2020), and describe cryptic species (Pyne
permitted, provided the original author(s) et al., 2019). The plants are of economic interest because of their fleshy edible fruits, some of
and the copyright owner(s) are credited
which are cultivated [e.g., P. philadelphica Lam., Zamora-Tavares et al. (2015); P. peruviana
and that the original publication in this
journal is cited, in accordance with L., Muniz et al. (2014); and P. angulata L., Vargas-Ponce et al. (2016)], and others which are
accepted academic practice. No use, gathered from their wild populations as food or medicine (Kindscher et al., 2012; Martínez
distribution or reproduction is permitted
et al., 2022). As currently circumscribed, Physalis is an American genus, with its diversity
which does not comply with these terms.
centered on the United States (28 species; Pretz and Deanna, 2020), Mexico (61 species; this
work), Central America [22 species, two of which are endemic, and absent from Mexico;
Knapp et al. (2005)], and the West Indies (current estimates are not available). Two species
(P. viscosa L. and P. subilsiana Toledo) are restricted to South America. However, assigning
names to herbarium specimens remains complicated because many of the taxonomically
important characters (e.g., calyx shape and anther color) are lost during herborization, and
good field notes are usually lacking. In addition, many Mexican species were described by
Waterfall (1967) based on single collections, making it difficult to assess their validity and
morphological variation. Some of the Mexican species ignored in floristic works [e.g., P.
purpurea Wiggins and P. vestita Waterf. by Martínez et al. (2017)] have turned out to be valid
after new evidence has been gathered. Our study aims to provide a key, short descriptions,
and conservation status for the species known to occur in Mexico. We include field
photographs, diagnostic characters, and distribution maps.

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Martínez et al. 10.3389/fgene.2023.1080176

FIGURE 1
Corolla diversity of Physalis spp. (A) P. acutifolia, (B) P. angulata, (C) P. angustiphysa, (D) P. campanula, (E) P. cinerascens, (F) P. chenopodiifolia, (G) P.
coztomatl, (H) P. cordata, (I) P. crassifolia var. infundibularis, (J) P. glutinosa, (K) P. greenmanii, (L) P. hederifolia, (M) P. latiphysa, (N) P. microcarpa, (O) P.
nicandroides, (P) P. patula, (Q) P. pruinosa, (R) P. purpurea, (S) P. solanacea, (T) P. sulphurea, (U) P. tamayoi, (V) P. waterfallii.

2 Methodology suitable specimens are probably deposited in European

herbaria to which we have no access.
We reviewed plant material deposited in Mexican (ANSM, To elaborate the distribution maps, we included the specimens
CIIDIR, ENCB, IBUG, IEB, MEXU, QMEX, XAL) and from the afore mentioned herbaria and specimens available as
United States (TEX-LL, MO, SD) herbaria (herbarium codes images, such as via SEINET and MEXU, only if the specimen
follow Thiers, 2016) and confirmed or corrected previous could be confidently determined and the geographical
identifications. Extensive collections have been made for most coordinates either were given in the label or could be
Mexican states, with emphasis on the north, west, and center. unambiguously assigned.
Specimens were georeferenced whenever possible and included in
a dataset. The conservation status of most Mexican species was
evaluated in 2017 by Vargas and Martínez using the AOO and EOO 3 Morphology
methods based on our distribution dataset, applying the IUCN
criteria (Goettsch et al., 2021). The status of each species is 3.1 Habit
available on the IUCN website (https://www.iucnredlist.org/
search?query=physalis&searchType=species; IUCN, 2022). Plants are frequently annuals that complete their life cycle
Distributions outside Mexico were also taken into consideration. within a year [e.g., P. solanacea (Schltdl.) Axelius or P. patula
Type material designated in the protolog was located Miller] and appear after the summer rains in June or July.
directly at the various herbaria. Small herbaria were searched Perennial plants are rhizomatous types that die back as the
with TORCH. We include either the catalog or the barcode unfavorable season begins (e.g., P. cinerascens (Dunal) Hitchc. or
number. Specimens not located are noted as such. Only P. P. vestita); however, in mesic species, the whole plant might be
campechiana and P. lagascae remain without a type, since present throughout the year (e.g., P. volubilis Waterf.). Finally, a few

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Martínez et al. 10.3389/fgene.2023.1080176

the plant. Geminate leaves usually have a large leaf accompanied by

a smaller one, which is frequently half the size but has the same
shape. The leaves can be sessile or petiolate, and the base is
frequently oblique.

3.3 Vestiture

Trichomes can be simple eglandular, glandular, or branched.

Glandular trichomes lend a fetid smell to the plant (e.g., P.
nicandroides Schltdl. and P. glutinosa Schltdl.) or gather debris
and insects, giving the plant a dirty appearance (P. sordida
Fernald). Many specimens are poorly preserved because pressing
such plants is difficult. Although most species have some form of
vestiture, a few are glabrous (P. glabra Benth.) or puberulent, and the
trichomes cannot be detected by the naked eye. Several species are
glabrescent, and trichomes are only present in the younger parts. A
detailed description of hair types is provided by Seithe and Sullivan
(1990).

3.4 Flowers
Corolla shape is rotate to campanulate, 5-angled, or 5-lobed
(Figures 1A–C, E–I, O–R, T–V). Tubular corollas occur only in P.
campanula Standl. & Steyerm. (Figures 1D, 4E) and P. glutinosa
Dunal (Figures 1J, 6C), and urceolate corolla occurs only in P.
solanacea (Figures 1S, 14A). The color varies from almost white
(Figures 1A, N, P, Q, T) to orange (Figures 1D, 4E). However,
most species have yellow corollas (Figures 1B, C, E–I, L, M, U,
V). Only P. solanacea (Figures 1S, 14A) and P. purpurea (Figures
1R, 12H) have purple corollas. Maculae are present in many
species, varying in shape and color; a few species have compound
macula [e.g., P. coztomatl Dunal (Figures 1G, 6A) or P.
greenmanii Waterf (Figure 1K)], and other species lack
maculations (Figures 1I, N). A dense mat of white trichomes
FIGURE 2
Diversity of fruits of Physalis spp. (A) P. aggregata, (B) P. cordata, is present at the corolla throat or at the filament insertion in most
(C) P. gracilis, (D) P. grisea, (E) P. microcarpa, (F) P. chenopodiifolia, (G) species. Martínez (1998) presents diagrams of corolla shapes and
P. nicandroides; (H) cross-section of P. nicandroides fruit showing the
maculations.
dry pericarp.

3.5 Inflorescence
species are either shrubs [e.g., P. campechiana L. and P. melanocystis
(B. L. Rob.) Bitter] or weak suffrutex that lean on other plants (e.g., Most of the species have a solitary pendulous axillary flower. The
P. coztomatl Dunal and P. lignescens Waterf.). The species are mostly exceptions are P. aggregata Waterf. (Figure 4C), P. angustior
erect and can grow up to 2–3 m high (e.g., P. latiphysa Waterf.), but a Waterf., P. campechiana, and P. melanocystis (Figure 9H), which
few (P. gracilis Miers, P. queretaroensis M. Martínez & L. Hernández, have fascicles of two or more flowers. Flower peduncles are usually
and P. volubilis) are decumbent and form long stolons that root at elongated in fruit.
the nodes. Stems can branch profusely, especially in perennial
plants, as in the case of P. crassifolia Beth., or remain a single
stem or one with a few branches. Stems are mostly terete, with the 3.6 Fruits
exception of P. angulata, which has angled stems.
The fruit of Physalis is always a two-carpellate berry that is
usually juicy. These fruits mature in different colors: they can be
3.2 Leaves green (Figures 2A, B), yellow (Figures 2C, D), orange, purple
(Figure 2E), or almost black (Figure 2F). A few species present a
The leaves are alternate, but in many cases the upper leaves are dry pericarp [e.g., P. nicandroides (Figures 2G, H), P. patula Miller,
geminate, and on some occasions, they are geminate throughout and P. latiphysa], and the fruits are brown when mature. Flavors are

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FIGURE 3
Fruiting calyces of Physalis spp. (A) P. acutifolia, (B) P. gracilis, (C) P. glutinosa, (D) P. lagascae, (E) P. hastatula, (F) P. waterfallii, (G) P. aggregata, (H) P.
angustiphysa, (I) P. grisea, (J) P. nicandroides, (K) P. pubescens, (L) P. porrecta.

acidic, bitter, or sweet, and in a few species, the fruit is strongly P. pubescens L., and P. solanacea]. Seven species are distributed in
glutinous with a fetid odor. Seeds are commonly abundant, only P. Mexico and the southern portion of the United States [P. acutifolia
glutinosa (Figure 6C) and P. solanacea (Figure 14A) always have a (Miers) Sandwith, P. caudella Standl., P. cinerascens, P. crassifolia, P.
small number (5–10). Fruits formed at the end of the growing season hederifolia A. Gray, P. latiphysa, and P. spathulifolia (Torr.) B. L.
may also contain fewer seeds. Turner]. Eight grow in Mexico and Mesoamerica (P. angustiphysa
Fruits are covered by an accrescent calyx, which takes different Waterf., P. campanula, P. campechiana, P. lassa Standl. & Steyerm., P.
shapes (see Martínez, 1998 for a diagram). Sometimes the calyx is leptophylla B.L. Rob. & Greenm., P. melanocystis, P. nicandroides, and P.
evidently 10-costate (Figures 3A–F), and in other cases it is strongly porrecta Waterf.). The remainder (36 species) are endemic to Mexico,
5-angled (Figures 3G–L). However, a few species have an 18 of which are micro-endemic to one state (e.g., P. greenmanii, P.
intermediate position in which some of the nerves are more glabra, P. mcvaughii Waterf., P. minimaculata Waterf., P. purpurea, and
prominent than others. The insertion of the peduncle can be P. vestita), to a few locations (P. hintonii Waterf. and P. pennellii
strongly invaginated, but invagination might be absent in other Waterf.), or to a mountain range (P. angustior Waterf., P. hunzikeriana
species. The fruiting calyx is green in immature fruits and becomes M. Martínez, P. lignescens Waterf., P. longiloba O. Vargas, M. Martínez
yellow (Figure 3A, B) or brown (Figure 3J) when the fruits mature. & Dávila, and P. longipedicellata Waterf.). See Table 1 for species
Only P. campechiana and P. melanocystis (Figure 9H) have purple or distribution and conservation status in Mexico.
almost black fruiting calyces. The tip of the fruiting calyx lobes is
constricted, but it remains open in P. purpurea (Figure 12H).
5 Economic importance, current or
potential
3.7 Seeds
In Mexico, sixteen wild species are locally used as food and for
Seeds of all the species of Physalis have a reniform shape and are medicinal purposes (Vargas-Ponce et al., 2016; Martínez et al.,
compressed. The color varies from yellow to dark brown, and the size 2022). The edible fruits of some Physalis species are nutritionally
varies from 1 to 3 mm in Mexican species. The testa is described as important (Puente et al., 2011; Zamora-Tavares et al., 2015;
foveolate. The etched seeds have undulated anticlinal walls and holes at Valdivia-Mares et al., 2016). Physalis acutifolia, P. angulata, P.
the bottom, and have been described in some detail by Axelius (1992). chenopodifolia Lam., P. cinerascens, P. cordata, P. coztomatl, P.
gracilis, P. philadelphica, and P. pubescens are harvested from their
natural populations for consumption (Santiaguillo Hernández
4 Distribution and Blas, 2009; Vargas-Ponce et al., 2011; Martínez et al.,
2022). The flavor of the fruits can be sweet or sour, and they
Of the 61 species that we recognize as part of the Mexican flora, are consumed fresh or in jams, sauces, and other dishes
ten have a wide distribution that includes the United States, Central (Santiaguillo Hernández and Blas, 2009; Vargas-Ponce et al.,
and South America, and the West Indies [P. angulata, P. cordata Miller, 2016). In particular, P. philadelphica is cultivated extensively in
P. grisea (Waterf.) M. Martínez, P. ignota Britton, P. lagascae Roem. Mexico (Peña-Lomelí et al., 2001; Zamora-Tavares et al., 2015)
& Schult., P. microcarpa Urb. & Ekman, P. philadelphica, P. pruinosa, and, to a lesser extent, in Guatemala and Nicaragua (López et al.,

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TABLE 1 Species distribution and conservation status of Physalis in Mexico.

Species Conservation status Distribution in Mexico

P. acutifolia LC B.C., B.C.S., Chih., Chis., Col., Camp., Jal., Mich., Nay., Sin., Son., Zac.

P. aggregata VU Jal., Oax.

P. ampla LC Col., Chih., Dgo., Gto., Jal., Mich., Nay., Oax., Qro., Sin., Son.

P. angulata LC Ags., B.C.S., Camp., Chis., Chih., Col., Dgo., Gto., Jal., Mich., Nay., N.L., Oax., Qro., Tab., Tamps., Sin., Son., Ver., Zac.

P. angustior DD Mor.

P. angustiphysa LC Chis., Gto., Jal., Mich., Nay., Son., Tamps.

P. campanula NT Chis., Hgo., Oax., Ver.

P. campechiana LC Camp., Chis., Oax., Q. Roo, Tab., Tam., Ver., Yuc.

P. caudella LC Chih., Coah., Dgo., Hgo., Gto., N.L., Qro., S.L.P., Son., Tamps., Zac.

P. chenopodiifolia LC Ags., Chih., Cd. Mex., Dgo., Edo. Mex., Gro., Gto., Hgo., Jal., Mich., N.L., Pue., Qro., S.L.P., Tlax., Zac.

P. cinerascens LC Ags., Camp., Chis., Coah., Edo. Mex., Gto., Hgo., Jal., Mich., N.L., Oax., Pue., Qro., S.L.P., Tamps., Ver., Yuc., Zac.

P. cordata LC Camp., Chis., Col., Gro., Jal., Mich., Nay., Oax., Sin., Tab., Ver., Yuc.

P. coztomatl LC Cd. Mex., Edo. Mex., Gro., Hgo., Jal., Mich., Mor., Nay., Oax., Pue., Qro., Tlax., Ver.

P. crassifolia LC B.C., B.C.S., Sin., Son.

P. glabra LC B.C.S.

P. glutinosa LC Ags., Chih., Cd. Mex., Dgo., Gto., Hgo., Jal., Zac.

P. gracilis LC Camp., Chis., Hgo., Jal., Mor., Nay., Oax., Pue., Qro., Q. Roo, S.L.P., Tab., Tamps., Ver.

P. greenmanii EN Ver.

P. grisea LC Jal., Mich.

P. hastatula EN Ags., Gto., Jal., Zac.

P. hederifolia LC Chih., Coah., Dgo., Gro., Jal., N.L., Oax., Qro., S.L.P., Son., Tamps., Zac.

P. hintonii LC Edo. Mex., Mich., Ver.

P. hunzikeriana DD N.L.

P. ignota LC Chis., Oax.

P. lagascae LC Ags., Camp., Chis., Chih., Col., Dgo., Edo. Mex., Gto., Gro., Jal., Mich., Mor., Nay., N.L., Oax., Pue., Qro., Q. Roo, Sin., Tab.,
Ver., Yuc., Zac.

P. lassa LC Chis., Col.

P. latecorollata DD Oax.

P. latiphysa LC Gto., Mich., Son.

P. leptophylla LC B.C.S, Chih., Col., Dgo, Edo. Mex., Gro., Jal., Mich., Nay., Oax., Sin., Son.

P. lignescens EN Jal.

P. longiloba LC Jal.

P. longipedicellata LC Jal.

P. mcvaughii NT Jal.

P. melanocystis LC Camp., Chis., Col., Gro., Hgo., Jal., Oax., Qro., S.L.P., Tab., Tamps., Ver.

P. microcarpa LC Chis., Edo. Mex., Jal., Son.

P. minimaculata VU Mich.

P. minuta LC Col., Chis., Gro., Jal., Nay., Oax.

(Continued on following page)

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TABLE 1 (Continued) Species distribution and conservation status of Physalis in Mexico.

Species Conservation status Distribution in Mexico

P. nicandroides LC Ags., B.C.S., Chis., Col., Dgo., Edo. Mex., Gto., Gro., Hgo., Jal., Mich., N.L., Mor., Nay., Oax., Pue., Qro., S.L.P., Sin., Son.,
Tamps., Ver., Yuc., Zac.

P. orizabae LC Ags., Cd. Mex., Chih., Dgo., Edo. Mex., Gto., Gro., Hgo., Jal., Mich., Mor., Nay., N.L., Oax., Pue., Qro., S.L.P., Ver., Zac.

P. parvianthera DD Mor.

P. patula LC Ags., Chih., Col., Cd. Mex., Dgo., Edo. Mex., Gto., Hgo., Jal., Mich., Oax., Pue., Qro., Sin., Tlax., Ver., Zac.

P. pennellii NT S.L.P.

P. philadelphica LC Present in all the Mexican states except Camp., Tlax., and Yuc.

P. porrecta DD Chis., Gro., Oax.

P. pringlei LC Cd. Mex., Dgo., Oax.

P. pruinosa LC Chis., Chih., Edo. Mex., Col., Gro., Jal., Mor., Nay., Oax., Qro., Sin., Son., Tamps., Ver., Yuc., Zac.

P. pubescens LC BCS., Camp., Chis., Chih., Coah., Col., Dgo., Gro., Hgo., Jal., Mich., Nay., NL., Oax., Pue., Qro., Q. Roo, Sin., Son., Tab.,
Tamps., Ver., Yuc.

P. purpurea DD Son.

P. queretaroensis LC Qro.

P. rydbergii DD Gto., Mich, Qro., Sin.

P. sancti-josephi DD Hgo., Jal, Nay., Qro., S.L.P.

P. solanacea LC Chih., Coah., Dgo., Gto., Hgo., Jal., Mich., N.L., Oax., Pue., Qro., S.L.P., Tamps., Ver., Zac.

P. sordida LC Coah., Dgo., Edo. Mex., Gto., Hgo., Jal., Mich., NL., Oax., Pue., Qro., S.L.P., Tam., Zac.

P. spathulifolia LC Tamps

P. subrepens LC Edo. Mex., Hgo., Jal., Ver.

P. sulphurea LC Cd. Mex., Dgo., Gto., Jal., Mich., Qro.

P. tamayoi NT Jal.

P. tehuacanensis CR Pue.

P. vestita Probably VU Sin.

P. volubilis LC Mich., Jal.

P. waterfallii LC Jal., Mich., Qro.

2009; Samuels, 2009). Mexico has a per capita consumption of P. 6 Taxonomic history
philadelphica of 5.3 kg; this is the fifth horticultural crop in
Mexico (SIAP 2019). It is also exported to the United States The taxonomic history of Physalis is complex due to controversies
of America, the United Arab Emirates, the United Kingdom, regarding the diagnostic characters traditionally used for its
France, Poland, Saudi Arabia, Canada, Switzerland, and classification. From Linnaeus (1753) to Rydberg (1896), the
Jamaica (SIAP, 2019). In recent years, P. angulata has gained characters commonly used to subdivide the genus included habit,
importance at the regional scale (Vargas-Ponce et al., 2016). flower arrangement, shape of the fruiting calyx, and pubescence
The cultural tradition of harvesting, the preference for small, (Martínez 1998). Waterfall (1967) monographed the vast majority of
acid-flavored fruits, and the high price and high yield make P. species. His study was followed by several nomenclature changes and
angulata a species with the potential to be cultivated extensively the discovery of new species. Martínez (1999) proposed an infrageneric
(Morales-Saavedra et al., 2019). classification based on habit, flower arrangement, flower and fruit
The roots, leaves, stems, fruits, and fruiting calyces of several morphology, trichome micromorphology, habitat, and geographical
species (P. philadelphica, P. angulata, P. chenopodifolia, P. distribution. According to Martínez (1999), Physalis has four subgenera:
cinerascens, P. pubescens, and P. coztomatl) are used by local 1) Physalis L., 2) Physalodendron (G. Don.) M. Martínez, 3) Quincula
populations as remedies. They are used as diuretics and against (Raf.) M. Martínez, and 4) Rydbergis Hendrych. The latter includes nine
headaches, stomach pains, diarrhea, skin rash, and diabetes sections: Angulatae (Rydb.) Menzel, Campanulae M. Martínez,
(Martínez et al., 2022). The various species of Physalis also have Carpenteri M. Martínez, Coztomatl M. Martínez, Epeteiorhiza G.
a reputation as fodder for cattle. Don, Lanceolata (Rydb.) Menzel, Rydbergae M. Martínez,

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Martínez et al. 10.3389/fgene.2023.1080176

Tehuacanae M. Martínez, and Viscose (Rydb.) Menzel. Regional floristic 3. Plants variously pubescent, but not minutely
studies have further advanced knowledge of Physalis. They have puberulent. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . . 4
included the southeastern United States (Sullivan 2004), 4. Plants with simple eglandular trichomes. . .. . .. . .. . .Key 4
Mesoamerica (Knapp et al., 2005), and Mexico (Martínez et al., 4. Plants with glandular trichomes, sometimes mixed with
2017; Martínez 2020). The molecular evidence does not support the simple eglandular trichomes . . .. . .. . .. . .. . .. . .. . .. . .Key 5
proposal of Martínez (1999), and this concept of the genus Physalis is
paraphyletic (Zamora-Tavares et al., 2016; Deanna et al., 2018). Key 1: Plants shrubby or suffrutescent
However, the Physalis subgenus Rydbergis is monophyletic (Zamora-
Tavares et al., 2016; Deanna et al., 2018) and includes 75 American 1. Corolla tubular or campanulate, flowers solitary. . .. . .. . .. . .. . .2
native species (Martínez 1999). The paraphyly of Physalis has been 2. Corolla tubular, limb not expanded, apex minutely 5-dentate,
partially resolved by segregating species as for Physalis alkekengi L., as 1.5–2.0 cm long and 1 cm wide, orange-yellowish, immaculate,
Alkekengi officinarum Moench (Whitson, 2011) and P. carpenteri fruiting calyx glabrous. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. campanula
Riddell as Calliphysalis carpenteri (Riddell) M. Whitson (Whitson, 2. Corolla campanulate, limb expanded, 2.5–3.5 cm long and
2012). This implied the proposal of Physalis pubescens L. as the 2.5–3–5 cm wide when fully expanded, yellow, with five large
new nomenclatural type for the genus (Whitson, 2011). Physalis compound maculations, fruiting calyx pubescent. . .. . . . . .. . .. . .
lobata Raf., which was recognized by Martínez (1999) as a . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. glutinosa
monotypic subgenus, was rehabilitated as the genus Quincula 1. Corolla rotate, 5-lobed or not, flowers solitary or in
Raf. by Barboza (2000). However, further taxonomic changes fascicles. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .3
are needed to make Physalis a monophyletic group. 3. Flowers in fascicles, with (1) two to seven flowers per node,
fruiting calyx purple to almost black at maturity. . .. . .. . .. . .. . .4
4. Plants with simple trichomes. . . . . . . . .. . .. . .P. melanocystis
7 Generic description and species key 4. Plants with dendritic trichomes. . .. . .. . . .P. campechiana
3. Flowers solitary or rarely with a shortened internode that
Physalis L., sp. Pl. 1:182, 1753. Conserved type: Physalis aggregates one to three flowers, fruiting calyx green at
pubescens L. (Whitson 2011). maturity. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .. . .. . .5
Shrubs, suffrutex, annual, or perennial herbs forming rhizomes; 5. Internode shortened and then one to three flowers
pubescence of simple eglandular, glandular, or branching trichomes, aggregated. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .P. angustior
frequently a mixture of two types; leaves petiolate, seldomly sessile, 5. Internode long, flowers solitary. . .. . .. . .. . .. . . . . .. . .. . .. . .6
alternate but frequently geminate, especially on the upper branches, 6. Fruiting calyx five-angled. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . . 7
blades oblong, elliptic, ovate, or lanceolate, less frequently rhombic, 7. Corolla pale yellow, anthers dark purple, plants
trulate, or linear, margin entire or dentate; flowers solitary or aggregated restricted to Veracruz. . . . . .. . .. . .. . .P. greenmanii
in few-flowered axillary fascicles, pedunculate; calyx campanulate to 7. Corolla yellow, anthers dull blue, plants from
tubular-campanulate, 5-lobed, lobes triangular to acuminate; flower Jalisco. . .. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .8
buds plicate, corolla actinomorphic, rotate, campanulate, tubular- 8. Flowers on peduncles up to 1.7 cm long, corolla
campanulate, or urceolate, mostly yellow but some species with 1.5–2.1 cm in diameter, reflexed flowering calyx
purple, orange, or white petals, yellow flowers commonly with five lobes. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .P. lignescens
simple or compound purple strongly contrasting maculations, or 8. Flowers on peduncles up to 4.5 cm long, corolla
maculations greenish or brown, not strongly contrasting, sometimes 2.2–2.4 cm in diameter, erect flowering calyx
maculations lacking, corolla throat usually pubescent at the insertion of lobes. . . . . .. . .. . .. . .. . .. . .. . .. . .P. longipedicellata
filaments; filaments filiform or flattened; stamens 5, anthers dehiscing 6. Fruiting calyx 10-costate . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .9
longitudinally, oblong, linear-oblong or sagittate, green, yellow, blue, 9. Pubescence abundant, simple glandular and
blue-tinged, or purple; style filiform, stigma capitate or clavate; fruit a eglandular trichomes mixed throughout the
two-carpellate spheric berry, green, yellow, orange, purple, brown, or plant. . . . . .. . . . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .10
almost black at maturity, that is loosely and completely covered by the 10. Corolla 2.8–4.0 cm in diameter, fruiting calyx
accrescent calyx; fruiting calyces 5-angled or 10-costate; seeds usually 4.0–4.5 cm long, with lanceolate-ovate lobes
numerous, yellow or brown at maturity, laterally compressed, testa 11–15 mm long. . . . . .. . .. . .. . .. . .. . .. . . .P. coztomatl
foveolate. 10. Corolla 1.2–2.0 cm in diameter, fruiting calyx
Artificial key to the species of Physalis in Mexico 2.5–3.0 cm long, with triangular lobes 4–5 mm
long. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .P. sancti-josephi
1. Plants shrubby or suffrutescent. . . .. . .. . .. . .. . .. . .. . .. . .. . . Key 1 9. Pubescence absent, or vestiture of simple eglandular
1. Plants annual or perennial, but not shrubby. . . . . . . . . . . .. . .. . .2 and glandular trichomes scarce. . .. . .. . .. . .. . .. . .11
2. Plants with branched trichomes. . .. . .. . .. . .. . .. . .. . . . . .Key 2 11. Flower on peduncles 2.5–3 cm long; calyx with deltoid lobes,
2. Plants with simple glandular or eglandular trichomes; or 3–5 mm long. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .P. mcvaughii
plants glabrous, but branched trichomes absent. . .. . . . . .. . .3 11. Flower on peduncles 0.5–1.0 cm long; calyx with acuminate lobes,
3. Plants glabrous or minutely puberulent. . .. . .. . .. . .Key 3 1.5–2.5 mm long. . .. . .. . . . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. tamayoi

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Key 2: Plants with branched trichomes 11 Corolla with reddish-brown evident maculations,
plants endemic to the Mexican Plateau, in desert
1. Trichomes branched only twice, long multicellular trichomes also scrubs. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .P. hastatula
present. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .P. vestita 10 Leaves ovate or lanceolate, never hastate. . .. . .P. caudella
1. Trichomes branched several times, long multicellular trichomes
mostly absent. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .2 Key 4: Plants with simple eglandular trichomes
2. Leaves spathulate, plants restricted to coastal sand dunes in
Tamaulipas. . .. . .. . .. . . . . .. . . . . . . . .. . .. . .. . .P. spathulifolia 1. Flowers aggregated, with short internodes. . .. . .. . .P. aggregata
2. Leaves never spathulate, plants of other habitats. . . . . .. . .. . .3 1. Flowers solitary. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .2
3. Anthers yellow, leaf margins entire, plants of arid 2. Corolla purple, with yellowish-brown maculations. . .. . .
zones. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . . . .. . .P. cinerascens . . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .. . .. . .. . . .P. purpurea
3. Anthers purple, blue, or blue-tinged but not yellow, leaf 2. Corolla yellow, with almost black, reddish-brown, or dark
margins dentate or undulate; plants of mesic habitats, purple maculations, but never yellowish maculations. . .. . .3
mostly forests. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .4 3. Plants annual or biannual . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .4
4. Fruiting peduncles 1.0–1.6 cm long, plants from 4. Fruiting calyx 5-angled. . .. . . . . .. . . . . .. . .. . . .P. grisea
Michoacán and Estado de México. . .. . .. . .P. hintonii 4. Fruiting calyx 10-costate. . . . . .. . .. . .. . . . . .. . .. . . . . .5
4. Fruiting peduncles 2–3 cm long, plants from Nuevo 5. Trichomes on the fruiting calyx with a widened
León. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .P. hunzikeriana base. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .P. ampla
5. Trichomes on the fruiting calyx without a widened
Key 3: Plants glabrous or minutely puberulent base. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . . .P. lagascae
3. Plants perennial
1. Plants annual. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .2 6. Plants decumbent, rooting at the nodes. . .. . . . . . . . .7
2. Corollas urceolate or tubular-campanulate. . .. . .. . .. . .. . .. . .3 7. Flowering peduncles 3.5–8.5 cm long, fruiting calyx 5-
3. Corollas tubular-campanulate, whitish; plants prostrate to angled. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .P. volubilis
erect up to 40 cm tall. . .. . .. . .. . .. . .. . .. . .. . .P. microcarpa 7. Flowering peduncles up to 2.5 cm long, fruiting calyx
3. Corollas urceolate, purple; plants erect up to 1 m 5-angled or 10-costate. . . . . . . . .. . .. . .. . .. . .. . .. . . .8
tall. . .. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .. . .P. solanacea 8. Fruiting calyx 5-angled, plants pubescent
2. Corollas rotate-campanulate. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .4 throughout, internode 6.0–10 cm long, leaf
4. Corollas whitish with slightly contrasting maculations, margins entire or dentate. . .. . . . .P. queretaroensis
plants of inundated habitats, stems hollow. . .. . . . . .. . .. . .. . . 8. Fruiting calyx 10-costate, pubescence restricted to
. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .. . . . . .. . .. . .. . . .P. sulphurea young parts, internodes up 6 cm long, leaf margins
4. Corollas pale yellow or whitish but evidently maculated, entire. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . .P. gracilis
plants in dry areas, stems solid. . .. . .. . .. . .. . .. . .. . .. . .. . .5 6. Plants erect, not rooting at the nodes. . .. . . . . .. . .. . .9
5. Fruiting calyx strongly 5-angled . . .. . .. . .. . .P. cordata 9. Stigma capitate, flowering buds apiculate. . .. . .
5. Fruiting calyx 10-costate. . . . . .. . .. . .. . .. . .. . .. . .. . .. . .6 . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. waterfallii
6. Flowering peduncles 1.8–6 cm long, corolla whitish 9. Stigma clavate, flowering buds rounded. . .. . .. . .10
or pale yellow. . .. . .. . .. . . . . .. . .. . .. . . . .P. acutifolia 10 Plants with long multicellular trichomes 2–3 mm long, leaves
6. Flowering peduncles 0.3–1.0 cm long, corolla with entire margin. . .. . .. . .. . .. . . . . .. . .. . .. . .. . . . . . . . .. . .. . .11
yellow. . .. . .. . .. . . . . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .7 11. Flowering peduncles 5–7 mm. . . . . . . . .. . .P. latecorollata
7. Stems angled, anthers straight after dehiscence 11. Flowering peduncles 8–23 mm. . .. . .. . .. . .. . .. . .P. subrepens
. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. angulata 10. Plants with trichomes shorter than 2 mm, leaves with dentate
7. Stems terete, anthers convolute after dehiscence margin. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . . . . . . .. . .. . .. . .. . .. . .12
. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .P. philadelphica 12. Fruiting calyx lobes 8–14 mm long, narrow, acuminate
1. Plants perennial. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .8 . . .. . .. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .. . .. . . .P. longiloba
8. Fruiting calyx 5-angled. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .9 12. Fruiting calyx lobes 3–7 mm long, ovate to triangular
9. Plants erect, pubescent only on one side of the stem, restricted . . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .13
to mesophytic forests of Chiapas and Oaxaca . . .. . .. . .. . .. . . 13. Plants gray due to abundant pubescence, leaf margins
. . .. . .. . .. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .. . .. . . .P. porrecta with 3–6 teeth per side. . .. . .. . .. . .. . .P. chenopodiifolia
9. Plants creeping, glabrous, growing in sand dunes . . .. . .. . . 13. Plants green, leaf margins entire or with 2 or 3 teeth per
.. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . . .P. minuta side. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. orizabae
8. Fruiting calyx 10-costate. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .10
10. Leaves hastate. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .11 Key 5: Plants with glandular trichomes, sometimes mixed
11. Corolla with blue slightly contrasting maculations, with simple eglandular trichomes
plants endemic to Baja California Sur, in sand dunes
and oasis. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. glabra 1. Plants annual. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .2

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2. Fruiting calyx 5-angled. . .. . .. . . . . .. . .. . . . . . . . .. . . . . .. . .. . .3

3. Corolla with purple maculations. . .. . .. . . . . .P. pubescens
3. Corolla with usually green, olivaceous, or reddish-brown
maculations. . .. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .4
4. Stems, peduncles, and fruiting calyx densely pubescent
with gray glutinous short trichomes. . .. . .. . . .P. ignota
4. Stems not densely pubescent, trichomes glandular with
orange tips. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .5
5. Fruiting peduncles thick (1.1–2.0 mm), corolla
creamy whitish. . . . . .. . .. . . . . .. . .. . .P. nicandroides
5. Fruiting peduncles thin (0.5 mm), corolla yellow. . .6
6. Plants small, up to 0.8 m high, fruiting calyx
1.5–3.0 cm wide. . .. . .. . .. . .. . .. . .. . .. . .P. patula
6. Plants large, higher than 1 m, fruiting calyx
2.5–4.0 cm wide. . .. . .. . .. . .. . . . . .. . .. . . . . .. . . .7
7. Fruiting peduncles 1.0–1.5 cm long, corolla
maculations reddish-brown, stems with
trichomes of the same size. . .. . . .P. latiphysa
7. Fruiting peduncles up to 5.5 cm long, corolla
maculations pinkish-brown, stems with a
mixture of long and short trichomes. . .. . .. . .
. . . . . . . .. . .. . .. . . . . .. . .. . .. . .. . .. . .P. pruinosa
2. Fruiting calyx 10-costate. . .. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . .8
8. Anthers 1.5–2.5 mm long, blue or blue-tinged; fruiting calyx
1.5–2.5 cm long. . .. . .. . . . . .. . .. . . . . .. . .. . .. . .P. leptophylla
8. Anthers 4 mm long, purple; fruiting calyx 0.9–1.9 cm
long. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .P. minimaculata
1. Plants perennial. . .. . .. . .. . . . . .. . .. . . . . .. . .. . . . . .. . .. . .. . .. . . .9
9. Leaf margin with large teeth, both surfaces densely covered with
large trichomes, plants endemic to Tehuacán in Puebla
. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . . . . .P. tehuacanensis
9. Leaf margin entire or toothed, but the teeth small;
trichomes do not cover the leaf surface. . .. . .. . .. . .. . .. . .10
10. Anthers short, 1.5–1.8 mm long, plants small, shorter
than 40 cm high. . .. . .. . .. . .. . . . . .. . .. . .P. parvianthera
10. Anthers larger than 2 mm, plants larger than 40 cm FIGURE 4
high. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . . .11 Details of flowers and fruits of Physalis spp. (A) P. acutifolia, (B) P.
11. Flowering peduncle <10 mm long. . .. . .. . .. . . . . . .12 angulata, (C) P. aggregata, (D) P. ampla, (E) P. campanula, (F) P.
angustiphysa, (G) P. chenopodiifolia, (H) P. cinerascens, (I) P. caudella,
12. Anthers smaller than 3 mm. . .. . .. . .. . .. . .. . . .13 (J) P. cordata.
13. Plants prostrate, fruiting calyx 5-angled. . .. . .
. . .. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . . .P. angustiphysa
13. Plants erect, fruiting calyx 10-costate . . .. . .. . .. . .14
14. Plants small, <20 cm tall, endemic to dry
areas in San Luis Potosí. . .. . .. . .. . .. . . . . . 17. Fruiting calyx 5-angled. . .. . .. . . . . .. . .. . .. . ..P. pringlei
. . .. . .. . .. . .. . . . . .. . .. . .. . . . . .P. pennellii 17. Fruiting calyx 10-costate. . .. . . . . .. . .. . .. . .P. crassifolia
14. Plants larger than 50 cm, distributed in mesic
habitats in western and southern Mexico
. . .. . .. . .. . .. . .. . .. . .. . . . . .. . .. . .. . . .P. lassa 8 Species descriptions
12. Anthers larger than 3 mm. . .. . .. . .. . .. . .. . .. . .15
15. Plants densely glandular, dirty, corollas 1. Physalis acutifolia (Miers) Sandwith, Kew Bull. 14 (2): 232 (1960)
1.5–2.5 cm in diameter. . .. . . . . .P. sordida
15. Plants glandular but not dirty, corolla 0.7–1.2 cm in Type: United States. California, T. Coulter 593 (holotype: K
diameter. . .. . .. . .. . . . . .. . .. . .. . .. . .. . . . . .. . .. . .. . .. . .. . .16 Barcode K000759429!). Saracha acutifolia Miers, Ann. Mag. Nat.
16. Leaf blades ovate to subcordate, corollas immaculate Hist. ser. 2,3 (18): 449. 1849. (Figures 1A, 4A).
. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .. . .P. hederifolia = Physalis wrightii Proc. Amer. Acad. Arts x. (1875) 63. Type:
16. Leaf blades lanceolate to trulate, corollas evidently USA, Texas, prairies of the San Pedro river, C. Wright 1602
maculated. . . . . .. . .. . . . . . . . .. . . . . .. . .. . .P. rydbergii (holotype: GH barcode 00003296!, isotypes: NY barcode
11. Flowering peduncle >11 mm long. . .. . . . . .. . .. . .. . .. . . .17 00138864!, US barcode 00027376!, microfilm MEXU!).

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Erect annual herb up to 1 m tall; stem highly ramified, glabrous; flowers solitary to almost aggregate with short
glabrous; leaves alternate throughout, petioles 1.1–6.6 cm internodes on peduncles 3.0–9.5 cm long; calyx with ovate lobes
long; blades lanceolate, 2.8–12.6 cm long, 1.1–6.7 cm wide, ca. 1.5 mm long, corolla rotate, yellow, 8–11 mm in diameter, with
apex acute to attenuate, base oblique to cuneate, decurrent, five dark brown maculations, corolla throat pubescent; anthers blue,
margin sinuate-dentate to serrate, glabrous; flowers solitary on 1.5–3.0 mm long; fruiting peduncle 1–16 mm long, fruiting calyx 10-
peduncles 1.8–6.0 cm long; calyx with triangular lobes 2–3 mm costate, inflated at the base and contracted at the upper third,
long, corolla rotate, pale yellow to whitish, 7–17 mm in 2.8–3.9 cm long, 1.8–2.2 cm wide, glabrous; mature berry green,
diameter, with five brownish-to-greenish slightly contrasting ca. 1 cm in diameter with numerous yellow seeds ca. 1.5 mm in
maculations, corolla throat pubescent; anthers purple, diameter.
1.5–3.0 mm long, sometimes convolute after anthesis; fruiting Distribution and habitat: Open sites in oak forest or ecotone
peduncle 2.5–6.0 cm long, fruiting calyx 10-costate, almost with cloud forest at 1,800–2,100 m of elevation. Jal, Oax.
round, 2–3.4 cm long, 1.6–2.4 cm wide, glabrous; mature Figure 5B.
berry green, ca. 1.5 cm in diameter with numerous yellow Diagnostic characters: Flowers aggregated together, or almost,
seeds ca. 1.5 mm in diameter. by short internodes, but the mature fruits are solitary.
Distribution and habitat: Common in the Pacific slopes on Common names and uses: Tomatillo.
volcanic rocks near sandy riverbanks or coastal vegetation, tropical Phenology: Flowering in July and August, and the fruits
deciduous forest, or mangroves up to 1,500 m elevation. B.C., B.C.S., collected in March.
Chih., Chis., Col., Camp., Jal., Mich., Nay., Sin., Son., Zac. Conservation status: VU.
(Figure 5A). Representative specimens examined. MEXICO. Jalisco:
Diagnostic characters: A whitish corolla, serrate leaves, and Jocotepec, camino de ascenso al cerro Viejo, por Las Trojes, A.
long flowering and fruiting peduncles. Rodríguez 993 (IBUG, IEB, MEXU). Oaxaca: Santiago Juxtlahuaca,
Common names and uses: Tomatillo. The fruits are eaten in 1–2 km del poblado El Manzanal, carretera a Infiernillo. Distrito
Baja California. Juxtlahuaca, J. Calzada 20,724 (MEXU).
Phenology: Flowering and fruiting from September to March.
Conservation status: LC. 3. Physalis ampla Waterf., Rhodora 69: 219 (1967)
Representative specimens examined. MEXICO. Baja
California: Rocky slope above Ejido San Matias, Type: Sinaloa, vicinity of Culiacán, Cofradía, T. S. Brandegee s.n.
31.32911N, −115.54025W, R.Thorne 57,561 (SD). Baja (holotype: UC barcode 104119!). (Figure 4D).
California Sur: Loreto, río Huatamote en el km 58 de la Erect annual herb 10–30 cm high, stem pubescent with
carretera Loreto-Ciudad Insurgentes, M. Martínez 9525 eglandular multicellular divergent trichomes; petioles 0.3–1.5 cm
(QMEX). Chihuahua: Janos, border of Chihuahua and Sonora, long; blades ovate to rhombic, 2–4 cm long, 1.0–1.5 cm wide, apex
S. White 2576 (MEXU). Chiapas: Tonalá, tidal marsh near Puerto acute, base attenuate, margin entire to dentate with three to four
Arista, D. E. Breedlove 25,546 (MEXU). Colima: Tecoman, upper teeth per side, pubescent with multicellular eglandular trichomes on
edge of the lagoon near kilometer post 55 on the highway between both surfaces; flowers solitary on peduncles ca. 2 mm long; calyx
Cuyutlán and Armeria, 9 km W of the Rio Armeria bridge, near with deltoid lobes ca. 1 mm long; corolla rotate, yellow, ca. 3 mm in
18° 55′ N, 104° 01′ W, Sanders 11,921 (MEXU, MICH). diameter with five slightly contrasting darker maculations; anthers
Campeche: Palizada, E. Matuda 3867 (MEXU). Jalisco: green or blue, 1–1.2 mm long; fruiting peduncles 3–4 mm long,
Tomatlán, near the new road ca. 25 km northwest of Río san fruiting calyx 10-costate 1.5–2.0 cm long, 1.5 cm wide, pubescent
Nicolás and 20 km southeast of Tomatlán, R. McVaugh 25,310 with multicellular trichomes with a widened base; mature berry
(MEXU). Michoacán: Tepalcatepec, ejido Tepalcatepec, carretera spherical ca. 1 cm in diameter with numerous dark brown seeds
Coalcomán, Agundis 1065 (MEXU, ENCB). Nayarit: Santiago 1–2 mm in diameter.
Ixcluitla a 10 km al W de Unión de Corrientes, camino a Distribution and habitat: A small plant of grasslands, tropical
Mezcaltitán, O. Téllez 12,112 (MEXU, MICH, ENCB). Sinaloa: deciduous forest, and farming areas; it grows from sea level to
1 km al W de El Quemado, saliendo de El Quelite, municipio de 2,000 m. Col., Chih., Dgo., Gto., Jal., Mich., Nay., Oax., Qro., Sin.,
Mazatlán, M. Martínez 9934 (QMEX). Sonora: Mesa del Seri, Son. (Figure 5C).
desviación hacia Sahuaripa, municipio de Hermosillo, M. Diagnostic characters: A widened persistent base of the
Martínez 9834 (QMEX). Zacatecas: Jalpa, Huanasco, R. multicellular trichomes in the flowering and fruiting calyx.
Hernández 9651 (MEXU). Common names and uses: Tomate verde. No uses known.
Phenology: Flowers and fruits from August to October.
2. Physalis aggregata Waterf., Rhodora 69: 108 (1967) Conservation status: LC.
Representative specimens examined. MEXICO. Colima:
Type: Mexico, Oaxaca, rancho de Calderon, San Juan, L. C. Pueblo Juárez, J. Mallet 12 (MEXU). Chihuahua: Temósachic,
Smith 771 (holotype: GH barcode 3284!). (Figures 2A, 4C). Nabogame J. E. LaFerrière 1638 (MEXU). Durango: Canantlán,
Erect annual herb up to 1 m tall; stem ramified, glabrous; leaves ca. 3 km del pueblo sobre la carretera a Santiago Papasquiaro, M.
alternate throughout, petioles 1.2–5.0 cm long; blades ovate or Martínez 9745 (QMEX). Guanajuato: Santiago Maravatío, cerro
suborbicular, 2.3–9.5 cm long, 2.2–7.0 cm wide, apex acute to Prieto, cerca de La Leona, Rzedowski 40591 (IEB, MEXU).
acuminate, base oblique, decurrent, margin sinuate to dentate, Jalisco: Jocotepec: cerro Viejo, camino que sale del poblado de

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FIGURE 5
Geographic distribution of Physalis spp. (A) P. acutifolia and P. coztomatl; (B) P. aggregata, P. latecorollata, and P. spathulifolia; (C) P. ampla; (D) P.
angulata; (E) P. campechiana and P. rydbergii; (F) P. campanula and P. volubilis.

Zapotitlán de Vadillo a El Aguacero, O. Vargas 833 (IBUG). camino a san Juan Peyotán, 22° 30′ N, 104° 30′ W, Flores 1213
Michoacán: alrededores de Sanguijuelas, municipio de (MEXU). Oaxaca: San Miguel Chimalapa, camino Benito Juárez-
Churintzio, J. Rzedowski 51986 (IEB). Nayarit: Nayar: 8 km aL La Ciénega, al E de la división, ca. 8 km al E de Benito Juárez, ca.
NW de san Juan Peyotán, en los límites de Nayarit y Durango, 37 km en línea recta al NNE de San Pedro Tapanatepec, S. Maya

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Martínez et al. 10.3389/fgene.2023.1080176

3849 (MEXU). Querétaro: Los Manantiales, 1 km aL SE de la Santiago Ixcuintla, O Tellez 12112 (MO). Nuevo León: San Francisco
presa El Batán, municipio de Villa Corregidora, L. Hernández de los Blancos, afuera de Galeana, M. Martínez 9636 (QMEX). Oaxaca:
3609 (QMEX). Sinaloa: vicinity of Culiacán, Cofradía, T. Sola de Vega, 25 km al S de Sola de Vega, J. Linares 4452 (MEXU).
Brandegee s.n. (UC). Sonora: Yécora, Puerto de la Cruz, north Querétaro: Alrededores de La Llave, Municipio de San Juan del Río,
base of Mesa del Campanero, T. Van Devender 96–477 (MEXU). J. Rzedowski 50,167 (IEB, QMEX). Tabasco: Jonuta, R. Fernández 1479
(MO). Tamaulipas: km 124 de la carretera Victoria-Miquihuana, ejido
4. Physalis angulata L., sp. Pl. 1: 183 (1753) José María Morelos, M. Martínez 9494 (QMEX). Sinaloa: Culiacán, a
20 km al N de Culiacán, carretera Culiacán a Guamúchil, R.Vega 6360
Lectotype (designated by D’Arcy in Ann. Missouri Bot. Gard. 60 (UAS). Sonora: Rancho Santa Bárbara, 42.3 km, ENE de Álamos, T.
(3):662. 1974): Anon. s.n. (LINN 247.9!). (Figures 1B, 4B). Van Devender 2006–1267 (ARIZ) Veracruz: Veracruz de Ignacio de la
Synonyms in Pretz and Deanna (2020). Llave, 1.6 km del poblado La Yerbabuena, sobre la carretera de La
Erect annual herb 30–40 cm high; stem angular, puberulent with Yerbabuena a El Ojital, M. Vázquez s. n (XAL). Zacatecas: 3 km al W
few eglandular trichomes to almost glabrous; basal leaves caducous, de Pueblo Viejo, Sierra de Morrones, cerro de Piñones, J. Balleza 8821
petioles 0.5–3.5 cm long; blades elliptic to elliptic-ovate, 2.5–5.0 cm (MEXU).
long, 1.5–3.0 cm wide, apex acute, base attenuate, margin dentate
with two to seven teeth per side, puberulent on both surfaces; flowers 5. Physalis angustior Waterf., Rhodora 69: 114 (1967)
solitary on peduncles 3–5 mm long; calyx with deltoid lobes ca.
2 mm long; corolla rotate, yellow, ca. 10 mm in diameter with five Type: Mexico, Morelos, Sierra de Tepoztlán, C. G. Pringle 7731
not strongly contrasting blue maculations, corolla throat almost (holotype: VT, isotype: US barcode: 00027313!)
glabrous; anthers yellow with blue lines ca. 2 mm long; fruiting Shrub or suffrutescent, 1–2 m high, densely covered with
peduncle ca. 1 cm long, fruiting calyx 10-costate, 2.5–3.0 cm long, articulated and partly glandular trichomes; petioles 3–5 cm
1.5–2.5 cm wide, puberulent; mature berry spherical 1.0–1.5 cm in long; blades ovate, 6–10 cm long, 4–8 cm wide, apex
diameter, green at maturity with numerous yellow seeds ca. 2 mm in acuminate, base attenuate, margin entire or with one or two
diameter. teeth per side; flowers solitary or 2–3-aggregated on peduncles
Distribution and habitat: A common weed associated with 12–20 mm long; calyx lobes lanceolate-tapered 5–9 mm long;
cornfields, rivers, grasslands, pine–oak forests, or tropical corolla yellow, 20–29 mm wide with five dark maculations,
deciduous forests; it grows from sea level up to 2,400 m. anthers violet, 3–4 mm long; fruiting peduncles 15–30 mm
Ags., B.C.S., Camp., Chih., Chis., Col., Dgo., Gto., Jal., Mich., long, fruiting calyx 25–30 mm long and 15–20 mm wide;
Nay., N.L., Oax., Qro., Tab., Tamps., Sin., Son., Ver., Zac berries 12–15 mm in diameter.
(Figure 5D). Distribution and habitat: Tropical deciduous forest; grows at
Diagnostic characters: Almost glabrous plant, with a small 2,200 m asl. Mor.
corolla and 10-costate fruiting calyx. Diagnostic characters: Suffrutescent with large acuminate
Common names and uses: Tomate, tomate de bote, tomate leaves.
ajuareño, tomate milpero, tomatillo, miltomate, tumat’cho, tomate Common names and uses: Unknown.
silvestre, tomate de tierra fría, tomate cuarentano, tuyuki (huichol). Phenology: Collected from November to May with flowers and
The species is self-pollinated, and spontaneous plants are tolerated fruit.
and harvested in Jalisco, Querétaro, Chihuahua, and Tamaulipas. Conservation status: Data deficient (DD), possibly extinct as no
The fruits are smaller than those of P. philadelphica but are recent collections are known of.
commonly sold in marketplaces. Representative specimens examined. MEXICO. Morelos:
Phenology: Flowering and fruiting from June to March. Sierra de Tepoztlán, C.G. Pringle 7731 (MEXU).
Conservation status: LC.
Representative specimens examined. MEXICO. Aguascalientes: 6. Physalis angustiphysa Waterf., Rhodora 69: 228 (1967)
Calvillo, área six Calvillo, Meza, Soto y León 505 (IBUG). Baja
California Sur: Comondú, Sierra La Giganta, Rancho Viejo El Type: Guatemala, Huehuetenango La Sierra, Steyermark 51977
Metate, 15 km al E. de Los Llanos de San Julio, M. Domínguez (holotype: US barcode 00027314!, isotype: F catalog 1148724!, US
3359 (HCIB). Campeche: Calakmul, a 17 km al S de la caseta de barcode 02824504!). (Figures 1C, 3H, 4F).
entrada a Calakmul, E. Martínez 28,930 (MEXU). Chiapas: Prostrate perennial herb 20–100 cm high; stems densely villous
Acapetahua, ejido Matamoros a 15 km aL sureste del poblado de with multicellular glandular trichomes; blades ovate to deltate,
Acapetagua, R. Trujillo 20 (MEXU). Chihuahua: arroyo Batopilas, 3–10 cm long, 3.5–7.0 cm wide, apex acute, base cordate, margin
a orillas del río, M. Martínez 9673 (QMEX). Colima: Colima, entire, sometimes dentate, villous on both surfaces; flowers solitary
Comaquitla, Sánchez-Ken 436 (MEXU). Durango: Puente on peduncles 4.5–9.0 mm long; calyx with triangular lobes
Morteros, al NE de Durango, González 2734 (CIIDIR-DGO). 2.5–3.5 mm long, corolla rotate, yellow, 1.0–1.5 cm in diameter
Guanajuato: Yuridia: 8 km al E de Yuridia, sobre la carretera a with five strongly contrasting purple maculations, corolla throat
Salvatierra, Rzedowski 40,314 (IEB). Jalisco: Jocotepec, camino de densely pubescent; anthers blue, 2.8 mm long; fruiting peduncle
ascenso al cerro Viejo, por Las Trojes, Rodríguez 994 (IBUG, IEB, 4.5–9.0 mm long, fruiting calyx strongly 5-angled, 2–3 cm long, ca.
MEXU, WIS). Michoacán: El Cerrito, 1 km al E de Jerécuaro, 1.5 cm wide, villous; mature berry spherical 1.5 cm in diameter with
municipio de Zinapécuaro, M.J. Jasso 1441 (IEB, QMEX). Nayarit: numerous yellow seeds ca. 2 mm in diameter.

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Distribution and habitat: Tropical deciduous forest; pine and Mexico, Campeche Houston s.n. (BM). Type not designated.
oak forest; grows from 800 to 2,100 m. Chis., Gto., Jal., Mich., Nay., Original material category “figure” according to Natural History
Son., Tamps. Museum (2022).
Diagnostic characters: A rare perennial, villous, and glandular = Physalis arborescens L., sp. Pl., ed. 2. 1: 261 (1762). Lectotype
plant with a narrow fruiting calyx. designated by Waterfall (1967). Physalis in Mexico, Central
Common names and uses: Chush-gutz, tomate, tomate zope, America, and the West Indies. Rhodora 69 (777): 95. The name
miltomate, chivol-antivo. No known uses. is illegitimate according to the Natural History Museum (2022).
Phenology: Flowering and fruiting from August to November. = Physalis mayana Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
Conservation status: LC. 8: 42 (1930). Type: Mexico: Yucatán: without definite locality, G.F.
Representative specimens examined. MEXICO. Chiapas: San Gaumer 24,381 (holotype: F barcode V0077207F!, isotype: US
Cristóbal de las Casas-Joigelito, Zinacatán, B. López 641 (MEXU). barcode 00027339!).
Guanajuato: Ocampo, mas ó menos 4 km al S de Santa Bárbara, E. Shrub with few ramifications 80–450 cm high; stems with
Carranza 4314 (IEB, MEXU). Jalisco: Autlán de Navarro, Las Galeras, abundant dendritic trichomes, glabrescent; blades elliptic to
Sierra de Manantlán, Vargas 861 (IBUG). Michoacán: Cerro la ovate, 5–14 cm long, 2–7 cm wide, apex acute, base decurrent,
Alberca, municipio de Villa Jiménez, E. Pérez y E. García 2590 margin entire, rarely crenate-dentate with six teeth per side, with
(IEB, QMEX). Nayarit: A 7 km al W del Venado, camino a San dendritic trichomes on both surfaces; flowers aggregated in two to
Pedro Ixcatán y El Zopilote. O. Téllez Valdés y Gabriel Flores F (TEX). seven flowered fascicles, peduncles ca. 1 cm long; calyx with
Sonora: El Divisadero, 1 km SE of El Llano on road to Bermúdez, T. R. triangular lobes, tomentose with short, branching trichomes;
Vandevender 97–662 (QMEX). Tamaulipas: Jaumave, Balcón del corolla 5-lobed, yellowish, 8–12 mm long, the lobes 3–5 mm
Chihue, M. Martínez 9475 (QMEX). long, with dark maculations; anthers yellow, ca. 4 mm long;
fruiting peduncles up to 1.2 cm long, fruiting calyx 10-costate,
7. Physalis campanula Standl. & Steyerm., Publ. Field Mus. Nat. purple ca. 3 cm long, ca. 2 cm wide; mature berry bright red
Hist., Bot. Ser. 23: 18 (1943) 7–11 mm in diameter with numerous yellow seeds 1.4–1.6 mm in
diameter.
Type: Guatemala: San Marcos: dry banks along upper part of Distribution and habitat: Tropical deciduous forest; it grows
Quebrada Canjula, between Sibinal and Canjula, Volcán Tacaná from sea level up to 400 m. Camp., Chis., Oax., Q. Roo, Tab., Tam.,
Steyermark 36067 (holotype: F catalog 151991!). (Figures Ver., Yuc. (Figure 5E).
1D, 4E). Diagnostic characters: Along with P. melanocystis, it presents a
= Physalis constricta Waterf., Rhodora 69: 99 (1967). Mexico: shrubby habit and fascicled flowers, but P. melanocystis has simple
Oaxaca, Ghiesbreght s.n. (holotype: P MNHN-P-P00387526) trichomes while P. campechiana has branched ones.
Shrub 30 cm high; stem with multicellular glandular trichomes; Common names and uses: Tomatillo, tamaliyo, tomate. No uses
blades ovate, 3–6 cm long, 2.5–4.0 cm wide, apex acute to acuminate, known.
base attenuate, margin entire to dentate with one to five teeth per Phenology: Flowering and fruiting throughout the year.
side, glandular villous on both surfaces; flowers solitary on Conservation status: LC.
peduncles 2.0–4.5 cm long; calyx with triangular lobes ca. 3 cm Representative specimens examined. MEXICO. Campeche:
long, ca. 3 mm wide; corolla tubular, orange or yellow, 1.5–2.0 cm Calakmul, a 37 km al S de la Caseta de entrada a Calakmul, E.
long, 1 cm wide, without contrasting maculations; anthers 4.5 mm Martínez 28,151 (MEXU). Chiapas: D. Breedlove 11800 (DS).
long; fruiting calyx 10-costate, 2–3 cm long, 1.5 cm wide, densely Oaxaca: Agua Tibia, 500 m al N de Nizanda, E. Pérez 1105
glandular-pubescent; mature berry yellowish, spherical ca. 1.5 cm in (MEXU). Quintana Roo: Tulum Sakbe’ 3 a San Pedro R. López
diameter. 276 (QMEX). Tabasco: Tenosique, a 2.6 km arriba del Cerro, en el
Distribution and habitat: Pine and oak forest, riparian campamento de la Escollera y 5 km de Tenosique hacia E. Zapata M.
vegetation; it grows from 2,200 to 3,000 m. Chis., Hgo., Oax., A. Magaña 843 (MEXU). Tamaulipas: Soto la Marina, 27.5 km del
Ver. (Figure 5F). camino de Soto la Marina a la Pesca, J. Jiménez 454 (MEXU).
Diagnostic characters: A rare shrub with orange, immaculate Veracruz: Tepetzintla, San José de Copaltitla, 7 km al NE de
tubular corollas with the limb obscurely dentate. Tepetzintla G. Castillo 2296 (QMEX). Yucatán: Río Lagartos
Common names and uses: None known. 2 km E del crucero Río Lagartos rumbo a Las Coloradas, P. Simá
Phenology: Flowers and fruits from February to April. 2332 (MEXU).
Conservation status: NT.
Representative specimens examined. MEXICO. Chiapas: 9. Physalis caudella Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
Motozintla, on the north and west slopes of the Cerro Mozotal 17: 273 (1937)
below the microwave tower along the road from Huixtla to El
Porvenir and Siltepec, D. Breedlove 25,859 (MEXU) Hidalgo: Type: Mexico, Chihuahua, Cajurichi, Rio Mayo, H.S. Gentry
Tenango de Doria, km 9 carr. 154, 8.8 km aL O de Cruz 2710 (holotype: F barcode V0072995F9!, isotypes: ARIZ catalog
Tenango P. Zamora Tavares et al. 260 (IBUG) Oaxaca: 274368!, GH catalog 77344!, MEXU-27636, MO 503577, S, UC
Ghiesbreght s.n. (P) Veracruz: Chiconquiaco, Arroyo Resbaloso, 582052, US barcode 27318!). (Figure 4I).
Ventura 4882 (ENCB, XAL). Synonyms in Pretz and Deanna (2020).
Erect perennial herb with long and short trichomes; petioles
8. Physalis campechiana L., Syst. Nat., ed. 10. 2: 933 (1759) 5–20 mm long, blades ovate-lanceolate to linear-lanceolate, 4–8 cm

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long, (6–) 15–40 mm wide, apex acute, base attenuate, decurrent, margins (holotype: US barcode 00027321!, isotypes: F barcode V0072998F!,
entire to irregularly undulate; flowers solitary on peduncles 5–10 mm GH 00077346!, NY barcode 138868!, UC 140093!).
long; calyx with acuminate lobes 5–8 mm long, corolla rotate, yellow, ca. = P. chenopodiifolia var. exigua Waterf,. Rhodora 69 (777):
2.2 cm wide with five reddish-blue or purplish maculations not strongly 110–111 (1967). Type: Mexico, San Luis Potosí, Charcas. C.
contrasting, or lacking; anthers bluish, ca. 3 mm long; fruiting peduncle Lundell 5398 (holotype: MICH 1109909!, isotype: US barcode
1.4–1.9 cm long, fruiting calyx 10-costate, 2.5–5.0 cm long and 00027320!).
2.5–3.0 cm wide, glabrous; mature berry yellow or orange, 1.0–1.3 cm = P. chenopodiifolia var. glandulosa Waterf., Rhodora 69 (777):
in diameter with numerous yellow seeds ca. 2 mm in diameter. 111 (1967). Type: Mexico, Querétaro, cliffs on steep mountainsides,
Distribution and habitat: Open sites in pine–oak forest and 22 miles NE of Zimapan, U.T. Waterfall 14170 (holotype: OKLA,
farming areas; it grows from 2,000 to 2,200 m asl. Chih., Coah., Dgo., isotypes: MICH, US. None of the cited specimens were located.
Hgo., Gto., N.L., Qro., S.L.P., Son., Tamps., Zac. (Figure 7A). Erect perennial herb, up to 70 cm tall, with dense, short
Diagnostic characters: A perennial with bluish anthers, pubescence throughout the plant, giving it a cinereous
purplish and not strongly contrasting maculations, and vestiture appearance; petioles 0.6–2.3 (−3.2) cm; blade ovate to lanceolate
of spreading, jointed trichomes, long and short intermixed. 1.7–6.7 cm long, 0.7–4.1 cm wide, apex acute, suddenly attenuated,
Common names and uses: Tomatillo, tomate borracho. No uses base oblique, decurrent, margin entire or with three to six teeth per
known. side; flowers solitary on peduncles 9–15 mm long; calyx with narrow
Phenology: Flowering and fruiting from July to October. triangular to ovate, attenuate or subulate lobes 1.5–4.2 mm long;
Conservation status: LC. corolla rotate campanulate, yellow, 1.6–3 cm in diameter, with five
Representative specimens examined. MEXICO. Chihuahua: strongly contrasting reddish-brown maculations, corolla throat
camino Ocampo-Moris, km 53, Municipio de Moris, M. Martínez densely pubescent; anthers blue, 2–3.5 mm long; fruiting
9666 (QMEX). Coahuila: Tanque Nuevo, M. Martínez 9605 peduncle 0.6–2.2 cm long, fruiting calyx 10-costate often with five
(QMEX). Durango: Cuencamé, ejido Las Mercedes, Campo more prominent angles, 2.2–3.4 cm long, 1.7–2.4 cm wide; mature
experimental de Zonas Áridas, aprox. 19 km al N de Cuencamé, berry green or purple-tinged, 1.3 cm in diameter with numerous
carr. 40 Durango-Torreón, 24° 52′ N y 103° 42′ W, A. Rodríguez yellow seeds ca. 2 mm in diameter.
1501 (IEB). Hidalgo: El Arenal, 1 km al E de el Arenal, M. López 103 Distribution and habitat: Open sites or pine–oak margins,
(MEXU). Guanajuato: Ocampo, 4 km aL SW de La Escondida, around crops or roads, and hillsides with woody vegetation or in
J. Rzedowski 52,206 (IEB, MEXU). Nuevo León: Doctor Arroyo, secondary grasslands. It develops between 2,000 and 3,100 m asl.
km 29 carretera a Doctor Arroyo, M. Martínez 9468 (QMEX). Ags., Chih., Cd. Mex., Dgo., Edo. Mex., Gro., Gto., Hgo., Jal., Mich.,
Querétaro: Municipio de Landa de Matamoros, misión de N. L., Pue., Qro., S.L.P., Tlax., Zac. (Figure 7B).
Tilaco, a 25 km de Landa de Matamoros, M. Martínez 2790 Diagnostic characters: A perennial plant with cinereous
(QMEX). San Luis Potosí: 2 km al W de Estación Catorce, hacia appearance and strongly contrasting corolla maculations. The
Santa María del Refugio, M. Martínez 9557 (QMEX). Sonora: plants grow in different conditions, and the species is highly
Puerto de Los Aserraderos, S. White 3207 (MEXU). Tamaulipas: variable. However, we do not consider the variations to merit
km 23 a Miquihuana, M. Martínez 9463 (QMEX). Zacatecas: San different names, as proposed by Waterfall (1967).
Tiburcio, ca. 3 km al E del entronque hacia Cedral, M. Martínez Common names and uses: Oxcones, tomate, tomatito,
9830 (QMEX). chapindikua, jaltomatillo. Roots, stems, and fruiting calyces are
used in traditional medicine (Santiaguillo Hernández and Blas,
10. Physalis chenopodiifolia Lam., Tabl. Encycl. 2 (3.1): 28 (1794) 2009; Valdivia-Mares et al., 2016). The fruit is eaten fresh in the
states of Mexico, Puebla, and Tlaxcala by the Mazahua population.
Type: Peru, Anon. s.n. (holotype: P-LAM) (Figures 1F, Phenology: Flowering and fruiting from June to October.
2F, 4G). Conservation status: LC.
= Physalis puberula Fernald, Proc. Amer. Acad. Arts 36: 502 Representative specimens examined. MEXICO.
(1901). Type: Mexico, State of Mexico, Sacro Monte, Amecameca, Aguascalientes: San José de Gracia, 1/2 km antes de llegar a La
C.G. Pringle 9147 (lectotype designated by Pretz and Deanna (2020) Congoja, O. Vargas 773 (IBUG). Chihuahua: Balleza, 46 km al E. de
MEXU barcode MEXU 00027679!, isolectotypes: CAS barcode Cabórachi, entre Cabórachi y Baquiriachi R. Hernández 8963
0004034, F barcode V0073008F, GH barcode 00077364, MICH (MEXU). Ciudad de México: Coyoacán, Reserva del Pedregal de
barcode 1109898, NMC 42525, RM barcode RM0004396, US San Ángel L. Mera 144 (MEXU. Durango: Nombre de Dios,
barcode 00027360, VT barcode UVMVT026427). Pretz and González-Elizondo 1042 (CIIDIR-DGO, MEXU). Estado de
Deanna interpreted the species as a synonym of P. hederifolia A. México: Huixquilucan Cerca de la presa “El Capulín,”
Gray. However, the leaves of P. puberula have entire margins and Fraccionamiento La Herradura J. Rzedowski 25,874 (MEXU).
angled fruiting calyces similar to P. chenopodiifolia. Furthermore, P. Guerrero: Ayutla de los Libres, Roca Colorada, A. Díaz 192
hederifolia grows mostly in the northern states of Mexico. (MEXU). Guanajuato: 4 km aL SW de La Escondida, municipio
= P. chenopodiifolia var. chenopodiifolia forma immaculata de Ocampo, J. Rzedowski 52,206 (IEB, QMEX). Hidalgo: Santiago de
Waterf., Rhodora 69 (777): 110 (1967). Type: Mexico, Tlaxcala, Anaya, A. Soriano 177 (MEXU). Jalisco: Acatic, 100 m adelante del
sunny banks between cultivation, Mt. Malinche, Balls 4861 entronque de la carretera Tepatitlán-Acatic, O. Vargas 900 (IBUG).
(holotype: US barcode 00027354!) Michoacán: cerca de la Cima, municipio de Epitacio Huerta,
= P. chenopodiifolia var. viridis Waterf., Rhodora 69 (777): J. Rzedowski 49,579 (IEB, QMEX). Nuevo León: 14 km al N de
110–111 (1967). Type: Mexico, Durango, Otinapa, E. Palmer 420 San Lázaro, 3 km de la desviación hacia La Joya, M. Martínez 9572

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Common names and uses: Agush, costomate, jaltomate, oshcon

de milpa, tabardillo blanco, tomatillo, xpajacan, pak-canul, xpakil,
tomate, tomate silvestre, tomate de monte, pichandra. The plant is
used against gastrointestinal ailments and skin infections and eaten
fresh or used for the preparation of sauces (Santiaguillo Hernández
and Blas, 2009; Martínez 2020).
Phenology: Flowering and fruiting from October to May.
Conservation status: LC.
Representative specimens examined. MEXICO.
Aguascalientes: Rincón de Romos: Pabellón, S. M. L. 728
(IBUG). Campeche: 3 km al S de Bolonchen de Rejón en los
alrededores de la gruta de Xtacum-bilxuan, E. Cabrera 8544
(MEXU). Chiapas: La Encañada, Ocozocuautla, F. Pimentel 77
(MEXU). Coahuila: Tanque Nuevo, M. Martínez 9607 (QMEX).
Estado de México: Carreta Coacalco-Tultitlán, 1 km de Tultitlan,
FIGURE 6
cabecera de municipio, F. Espinosa 535 (MEXU). Guanajuato:
Details of flowers and fruits of Physalis spp. (A) P. coztomatl, (B) P. Acámbaro: 5 km al W de Iramuco, sobre el camino a Santa Ana
crassifolia, (C) P. glutinosa, (D) P. greenmanii, (E) P. gracilis, (F) P. Maya, J. Rzedowski 44,860 (IEB, ENCB). Hidalgo: SW of Zimapan
hastatula, (G) P. hederifolia, (H) P. grisea.
ca. 1.5 km at crossing of Arroyo Santiago and the road to Estanzuela,
M. Mayfield 847 (MEXU). Jalisco: Lagos de Moreno, Est. Pedrito,
Lagos de Moreno, a la orilla de la vía del ferrocarril, Flores Nava 35
(QMEX). Puebla: supercarrtera a la altura de Estación, W. Boege 166 (IBUG). Michoacán: Morelia, Francisco Mujica, 9 km aL NE de
(MEXU). Querétaro: 7 km aL SE de Tancoyol, municipio de Jalpan, Morelia, carretera a Maravatío, J. Soto Núñez 6450 (ENCB). Nuevo
R. Fernández 3488 (ENCB, IEB). San Luis Potosí: 4 km hacia León: 9.5 km al N de Aramberri, M. Martínez 9624 (QMEX).
Alamillos, Sierra de Catorce, M. Martínez 9559 (QMEX). Oaxaca: Ciudad Ixtepec, al pie del cerro La Pedrera. 14 km al N
Tlaxcala: Ixtenco, 3 km al W de San Juan Ixtenco sobre el de la Ventosa, M. Martínez 1905 (MEXU). Puebla: Zinacatepec,
camino para la Malinche, D. Williams 112 (MEXU). Zacatecas: 3 km al E de Zinacatepec, P. Tenorio 17184 Querétaro: 3–4 km al N
San Pedro Piedra Gorda: cerro Prieto, Trujillo ZA-119 (ENCB). de las Flores, rumbo al río Santa María, municipio de Jalpan, E.
Carranza 3154 (IEB, QMEX). San Luis Potosí: 5 km al N de
11. Physalis cinerascens (Dunal) Hitchc., Key Spring Fl. Matehuala, hacia entronque San Roberto, M. Martínez 9833
Manhattan (1894) 32 (QMEX). Tamaulipas: Matamoros, parque industrial en avenida
División del Norte, A. Herrera 9705 (QMEX). Veracruz: 2 km al N
Type: Mexico, Matamoros, circa Matamoros urbem, J. L. de Paso del Toro o Concepción, 5 km aL NE de 6 de Enero, M.
Berlandier 2316 (lectotype designated by Waterfall Rhodora 60 Martínez 1906 (MEXU). Yucatán: Dzidzantún, paraje a orilla del
(713)136: GH barcode 00077295!, isolectotypes: F barcode poblado San Francisco Manzanilla, J. Santiaguillo 8 (QMEX).
V0073006F!, NY barcode 00138882!). Physalis pensylvanica var. Zacatecas: 15 km aL SE de Sain Alto, M. Martínez 9784 a (QMEX).
cinerascens Dunal, Prodr. 13:435–436. 1852. (Figures 1E, 4H).
Synonyms in Pretz and Deanna (2020). 12. Physalis cordata Mill., Gard. Dict., ed. 8. n. 14 (1768)
Prostrate to slightly erect perennial herb, branched from the base,
densely pubescent, with stellate trichomes throughout; leaves alternate Type: Mexico, Veracruz, Houston s.n. (holotype: BM = BH neg
or geminate, petioles 0.5–3.5 cm long; blades ovate, suborbicular, or 5099!). (Figures 1J, 2B, 4J).
triangular-ovate, 1.5–6.7 cm long, 1.3–5.4 cm wide, apex obtuse to Synonyms in Martínez (1998). Pretz and Deanna (2020)
acute, base cuneate to truncate, decurrent, margin entire; flowers included P. porrecta Waterf. as a synonym; however, the original
solitary on peduncles 0.6–2.2 cm long; calyx with deltoid lobes description states that the plant has articulated trichomes, whereas
1–2 mm long, corolla rotate, yellow, 8–13 mm in diameter, with five P. cordata is glabrous.
dark-brown-to-black maculations; anthers yellow, 3–4 mm long; Erect annual herb, branched, up to 90 cm tall, glabrous; leaves
fruiting peduncle 1.3–6 cm long, fruiting calyx 10-costate, nearly alternate, petiole 1.5–8.6 cm long; blades ovate 8.5 (−12) cm long,
cylindrical, 1.4–3 cm long, 1.1–2.1 cm wide; mature berry yellow, ca. 2–7.8 (−9) cm wide, apex acuminate, base oblique to cuneate,
1 cm in diameter with numerous yellow seeds ca. 2 mm in diameter. decurrent, margin serrate, glabrous; flowers solitary on peduncles
Distribution and habitat: The species thrives in dry places, on 0.6–1.7 cm long; calyx with narrow triangular lobes 3.0–5.2 mm
limestone slopes with grassland, scrub, or tropical deciduous forest, long, corolla campanulate rotate, yellow, 1.1–2.1 cm in diameter,
in disturbed areas; it grows at 1,700–2,100 m asl. Ags., Camp., Chis., with five purplish-brown maculations, corolla throat densely
Coah., Edo. Mex., Gto., Hgo., Jal., Mich., N.L., Oax., Pue., Qro., pubescent; anthers blue or purple, 2.2–4.0 mm long; fruiting
S.L.P., Tamps., Ver., Yuc., Zac. (Figure 7C). peduncle 2.2–3.2 cm long, fruiting calyx 5-angled, 2.2–4.1 cm
Diagnostic characters: Densely pubescent throughout with long, 1.5–3.0 cm wide, glabrous; mature berry spherical to ovoid,
stellate trichomes; dark maculations that contrast with the 1.2–1.3 cm in diameter, green or purple with yellow seeds 2–3 mm
yellowish corolla. in diameter.

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FIGURE 7
Geographic distribution of Physalis spp. (A) P. caudella, (B) P. chenopodiifolia, (C) P. cinerascens, (D) P. cordata, (E) P. crassifolia and P. glabra, (F) P.
glutinosa.

Distribution and habitat: A common plant of sandy Camp., Chis., Col., Gro., Jal., Nay., Mich., Oax., Sin., Tab.,
soils near humid places, such as riparian or secondary and Ver., Yuc. (Figure 7D).
ruderal vegetation, associated with tropical deciduous or Diagnostic characters: In herbarium specimens, Physalis
sub-deciduous forest; it develops from sea level to 1,300 m. cordata is recognized by the shiny, finely toothed leaves, the

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Martínez et al. 10.3389/fgene.2023.1080176

FIGURE 8
Geographic distribution of Physalis spp. (A) P. gracilis, (B) P. greenmanii, (C) P. hastatula and P. pennellii, (D) P. hintonii and P. latecorollata, (E) P.
hederifolia, (F) P. ignota and P. minuta.

glabrous 5-angled fruiting calyx, and peduncles up to Phenology: Flowering and fruiting from September to February.
3.2 cm long. Conservation status: LC.
Common names and use: Tomatillo miltomate; in Jalisco, the Representative specimens examined. MEXICO. Campeche:
fruits are called corazón de gato and corazón de pollo according to Tuxpeña, C. Lundell 936 (CAS; F). Chiapas: Esperanza, Escuintla, E.
their size. Matuda 16,484 (F). Colima: Tecomán, Cofradía de Morelos, Maillet 425

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Martínez et al. 10.3389/fgene.2023.1080176

(MEXU). Guerrero: 16 km of Acapulco on Rte 95, J. Miller 295 MEXU). 16 de la brecha El Fresnito-La Joya, Nevado de Colima,
Jalisco: Cihuatlán, coastal plain near the highway to Autlán, four miles Ramírez Delgadillo 1730 (IBUG, WIS). Michoacán: Sierra de
north of Bahía Navidad, R. McVaugh 20,843 (MEXU, MICH). San Joaquín, 11 km al S de Tlalpujahua, municipio de
Michoacán: 5 km camino Aquila-La Placita, B. Guerrero 166 Tlalpujahua, S. Moreno 199 (IEB). Morelos: Huitzilac al N
(MEXU). Nayarit: Bahía de Banderas, 5 km al N del río Ameca, del Volcán Quimitepec, Sierra de Chichinautzin, Y. González
Puga 9708 (IBUG). Oaxaca: vicinity of Cafetal Concordia, C. Morton 1985 (MEXU). Nayarit: Nayar, arroyo de los Negros, ejido
2556 (US). Sinaloa: Santa Fe, Ortega 4384 (US). Tabasco: Balancan Ojo Colorado de la Mora, a 500 m del poblado, 21 ° 47′ N, 104 °
de Agua, J. Santiaguillo 1 (MEXU). Veracruz: San Andrés Tuxtla, 38′ W, Álvarez 187 (MEXU, MICH). Oaxaca: km 27 on road
Estación de Biología Tropical Los Tuxtlas, G. Ibarra 1989 (MEXU). from Teotitlán to Huautla, at Puerto Soledad, D. Spooner 953a
Yucatán: Izamal, Gaumer s.n. (MEXU). (MEXU). Puebla: faldas del Ixtlacihuatl, F. Salazar 1914
(MEXU). Querétaro: 1 km aL NE de Pinal de Amoles,
13. Physalis coztomatl Mociño & Sessé ex Dunal, Prodr. [A. P. de municipio de Pinal de Amoles, A. Herrera 203 (ENCB, IEB,
Candolle] 13 (2): 450 (1849) MEXU). Tlaxcala: atrás de las cabañas del CREA en Ixtenco, V.
Sánchez 297 (MEXU). Veracruz: Calcahualco, 9 km W of
Type: Drawing 48 of Sessé and Mociño at the Hunt Institute Escola, Nee 23,189 (F, XAL).
(holotype Copy 916 of Icones florae mexicanae ineditae). (Figures
1G, 6A). 14. Physalis crassifolia Benth., Bot. Voy. Sulphur [Bentham] 40
= Physalis stapelioides (Regel) Bitter, Repert. Spec. Nov. Regni Veg. (1844)
18: 5 (1922). Saracha stapelioides Regel, Suppl. Hort. Bot. Petropol. 18.
1864. Type not located, could be at LE or ZT (Thiers 2016). Type: Mexico, Baja California Sur, Lower California, Bay of
= Physalis acuminata Greenm. Proc. Amer. Acad. Arts 35: 311 Magdalena R.B. Hinds s.n. (lectotype designated by Pretz and
(1900). Type: Mexico: State of México: collected on the Sierra de las Deanna 2020 K barcode K000042380!). (Figures 1I, 6B).
Cruces, C. G. Pringle 5315 (holotype: GH 00077342!, isotypes: F Synonyms in Pretz and Deanna (2020).
barcode V0072994F!, MEXU-28024). Perennial herb, sometimes woody-stemmed, 10–40 cm tall,
Suffrutescent perennial branched from the base, up to 2 m high, variously vestite with short and glandular trichomes; petioles 1–4
pubescent with eglandular and glandular trichomes; leaves alternate (−6) cm long; blades ovate (1−) 2–6 (−10) cm long, 1–5 (−7) cm
or geminate, petioles 1.4–6.5 cm long; blades ovate to suborbicular wide, apex acute to acuminate, base truncate to cordate, margins entire
ovate 5.7–19 cm long, 3.9–17 cm wide, apex acuminate, base cuneate, or toothed; flower solitary on peduncles 1–3.3 cm long; calyx with
truncate to subcordate, decurrent, margin entire or with one or a few triangular lobes 2–3 mm long, corolla campanulate, rotate to
teeth, sometimes serrated; flowers solitary on peduncles 14–28 mm funnelform, yellow, 8–30 mm long and 8–20 mm wide, immaculate,
long; calyx with lanceolate-ovate lobes 11–15 mm long, corolla rotate, or with slightly darkened maculations; anthers yellow, 2.5–3.5 mm long;
yellow, 2.8–4.0 cm in diameter with five reddish-brown-to-purple fruiting peduncles 2.2–7 cm long, sometimes coiling, fruiting calyx 10-
compound maculations, corolla throat densely pubescent; anthers costate, 1–6 cm long, 8–35 mm wide; mature berry green, spherical,
purple, 3–5 mm long; fruiting peduncle 1.8–4.0 cm long, fruiting calyx 1 cm in diameter, with numerous brown seeds ca. 2 mm in diameter.
10-costate, pubescent, somewhat coriaceous, 4.0–5.7 cm long, Distribution and habitat: It grows in coastal dunes and sandy
2.8–3.4 cm wide; mature berry green, ca. 2 cm in diameter with places, protected between rocks near cliffs and oases, from 0 to
numerous yellow seeds ca. 2 mm in diameter. 200 m asl. B.C., B.C.S., Sin., Son. (Figure 7E).
Distribution and habitat: In pine, oak, Abies, or cloud forest, Diagnostic characters: A perennial plant of desert areas,
from 2,300 to 3,100 m. Cd. Mex., Edo. Mex., Gro., Hgo., Jal., Mich., immaculate corollas that are sometimes funnelform, referred to
Mor., Nay., Oax., Pue., Qro., Tlax., Ver. (Figure 5A). as var. infundibularis I. M. Johnston.
Diagnostic characters: A large fruticose plant with dense Common names and uses: Tomatillo. No uses known.
pubescence, with large corollas and compound maculations; it Phenology: Flowering and fruiting throughout the year.
grows at high altitudes. Conservation status: LC.
Common names and uses: Coztomate; the plant is used as a Representative specimens examined. MEXICO. Baja
diuretic and against stomach inflammation, diarrhea, and flatulence. California: Cataviña, en el arroyo, M. Martínez 9545 (QMEX).
Applied directly to the chest area, it helps against asthma Baja California Sur: Comundu, poblado de Banderitas a 9.6 km al E
(Hernández 1943). de San Carlos, M. Martínez 9524 (QMEX). Sinaloa: en Bagresitos, a
Phenology: Flowering and fruiting throughout the year. 39 km al N de Culiacán, E. Martínez 4122 (MEXU). Sonora:
Conservation status: LC. Pinacate Region Hourglass Canyon, two miles NE of Tinaja
Representative specimens examined. MEXICO. Ciudad de Huarache (=Pyramid Tank) on the west side of Sierra Pinacate,
México: Parque Nacional Desierto de los Leones, E. Maldonado R. Felger 19,175 (MEXU).
7 (MEXU). Estado de México: Tlalmanalco, San Rafael E.
Matuda 27,578 (MEXU). Guerrero: Chilpancingo de los 15. Physalis glabra Benth., Bot. Voy. Sulphur [Bentham] 39
Bravo, 6 km aL NW de Omiltemi. Brecha Chilpancingo- (1844)
Oniltemi-Las Joyas, P. Tenorio 2588 (MEXU). Hidalgo:
Tenango de Doria, 10 km al W de Tenango de Doria, R. Type: Mexico, [Baja California Sur] Cape Lucas Hinds, s.n. 1841
Hernández 4303 (MEXU). Jalisco: Ciudad Guzmán, km (holotype: K specimen K000042387!)

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= P. hastata Rydb. Mem. Torrey Bot. Club 4: 363 (1896). Type: Representative specimens examined. MEXICO. Aguascalientes:
Mexico, Baja California Sur, Todos Santos T. S. Brandegee 422 (holotype: San José de Gracia, camino al cerro de La Ardilla, cuesta La Gloria, De
UC not located, isotypes: GH not located, US catalog 00027327!) la Cerda García 730 (CIIDIR-DGO). Chihuahua: Majalca, D. LeSueur
Perennial herb, 10–100 cm long, glabrous or minutely 894 (F). Ciudad de México: Eslava, E. Lyonnet 714 (MEXU).
puberulous; petioles 12–25 mm long, blades lanceolate 2–4 cm Durango: mountainside five miles southwest of Durango, U.
long, 0.7–3.0 cm wide, apex acute, base truncate to hastate, margin Waterfall 15,409 (MICH). Guanajuato: 4 km al S de Santa
undulate with five to seven teeth; flowers solitary on peduncles Bárbara, municipio de Ocampo, E. Carranza y Col. 4314 (IEB,
10–20 mm long; calyx with deltoid lobes ca. 2 mm long; corolla QMEX). Hidalgo: Metepec, C. Pringle 13,125 (MEXU). Jalisco:
rotate, yellow, 9–20 mm in diameter, with blue slightly contrasting Mezquitic, km 50 camino Bolaños-Tenzompa, al NE de Pinos
maculations; anthers yellow, 2.0–2.5 mm long; fruiting peduncles Altos, comunidad indígena de Santa Catarina, González
17–30 mm long; fruiting calyx 10-costate, 10–23 mm long, Villarreal 3176 (CIIDIR-DGO, IBUG, IEB). Zacatecas: 15
11–20 mm wide; mature berry spherical 10–12 mm in diameter, (air) miles NE of Estación Camacho on NW slopes near
with numerous yellow seeds ca. 2 mm in diameter. granitic summit of Pico de Teyra, J. Henrickson 13,456 (MEXU).
Distribution and habitat: Coastal dunes and oases, B.C.S.
(Figure 7E). 17. Physalis gracilis Miers, Ann. Mag. Nat. Hist. ser. 2, 4 (19): 37
Diagnostic characters: A glabrous woody perennial with small (1849)
hastate leaves.
Common names and uses: None known. Type: Mexico [Hidalgo] Real del Monte, T. Coulter 1222
Phenology: Flowering and fruiting from December to March. (holotype: K barcode K000042379!, isotype: GH 00077352!).
Conservation status: LC. (Figures 2D, 3B, 6E).
Representative specimens examined. MEXICO. Baja Perennial herb, slightly erect to prostrate, up to 90 cm, rooting at
California Sur: Los Cabos, A. Anthony 353 (MEXU). the lower nodes, with short, eglandular, septate trichomes; leaves
geminate throughout; petiole 8–15 mm long, blades ovate to sub-
16. Physalis glutinosa Schltdl., Linnaea 19 (3): 310 (1846) orbicular, 4.2–9.2 cm long, 2.1–5.2 cm wide, apex acute, base cuneate
or oblique, narrowly decurrent, margin entire; flowers solitary
Type: Mexico, Hidalgo, Cuesta Blanca, Mineral del Monte, C. on peduncles 1.3–2.5 cm long; calyx with deltoid to lanceolate
Ehrenberg 585 (there are two specimens deposited at HAL; we lobes 3–4 mm long; corolla rotate-campanulate, yellow,
designate specimen barcode 042,219! as the lectotype, as it has more 1.5–2.0 cm in diameter, with five reddish-brown maculations;
material and is better preserved). (Figures 1J, 3C, 6E). anthers purple-blue, 3.0–3.5 mm long; fruiting peduncle
= Physalis cinerea Waterf., Rhodora 69: 102 (1967). Type: Mexico, 1.3–2.6 cm long, fruiting calyx 10-costate, 2.6–3.4 cm long,
San Luis Potosí, in the region of San Luis Potosí, C. C. Parry & Ed. 1.1–2.1 cm wide; mature berry spherical, yellow, 8–10 mm in
Palmer 649 (holotype: NY barcode 138,869!, isotype: GH 00077347!). diameter with numerous yellow seeds ca. 1.5 mm in diameter.
= Physalis eximia Standl., Fieldiana, Bot. 27: 273 (1937). Type: Distribution and habitat: Open areas of pine–oak or cloud
Mexico, Chihuahua, Majalca, D. H. LeSueur 894 (holotype: F forest, or as a weed in crops associated with the forest. 500–1,700 m
barcode V0073000F!, isotypes: GH 00077351, UC not located). asl. Camp., Chis., Hgo., Jal., Mor., Oax., Pue., Qro., Q. R., S.L.P., Tab.,
= Physalis glutinosa var. eximia (Standl.) Waterf., Rhodora 69: Tamps., Ver. (Figure 8A).
101 (1967). Diagnostic characters: A perennial with a prostrated habit,
Erect perennial herb, sometimes spreading, up to 1.5 m tall, rooting at the basal nodes.
branched, densely pubescent, glutinous, fetid; leaves alternate or Common names and uses: Costomate, costomatillo, tomate,
geminate, petioles 1.2–4.7 cm long; blades ovate, subcordate or tomatillo, tomate borracho, tomatito amarillo, tomate de bolsa,
suborbicular, 3.3–8.2 cm long, 2.0–5.6 cm wide, apex acute to miltomate de monte, chihol ch’o, chucky chan, moen’cho,
acuminate, base oblique, margin dentate, sinuate, or entire; flowers chuchan. The sweet fruit is eaten fresh, and the plant is medicinal.
solitary on peduncles 1.5–2.0 cm long; calyx 1.1–1.7 cm long, 6–11 mm Phenology: Flowering and fruiting in July–September, probably
wide, corolla tubular campanulate, yellow or cream yellow, 22–35 mm throughout the year.
long, 25–35 mm in diameter, with diffuse purple maculations; anthers Conservation status: LC.
purple, 4–5 mm long; fruiting peduncle 1.6–2 cm long, fruiting calyx Representative specimens examined. MEXICO. Campeche:
10-costate, reticulate, 2.4–4.2 cm long, 1.6–4.3 cm wide; mature berry Calakmul, en Narciso Mendoza D. Álvarez 17 (MEXU). Chiapas:
spherical, 1.2–1.5 cm in diameter with very few (1–5) yellow seeds ca. Ocozocoautla de Espinosa, 1 km al N de Buenavista (El Suspiro)
2.7 mm in diameter. E.Martínez 31,288 (MEXU). Hidalgo: Molango de Escamilla,
Distribution and habitat: An infrequent large plant that grows Ismolintla, 5 km aL norte de Molango R. Hernández 7094
near xerophytic scrubs and dry oak forests at 2,000–2,500 m asl. (MEXU). Jalisco: Cuautitlán de García Barragán, Ayotitlán, cerro
Ags., Chih., Cd. Mex., Dgo., Gto., Hgo., Jal., Zac. (Figure 7F). El Pinacate, 1/2 km adelante de san Miguel, O. Vargas 805 (IBUG).
Diagnostic characters: A large plant, densely glutinous, Morelos: wet banks, barranca near Cuernavaca, C. Pringle 6319
pubescent, with large tubular-campanulate corollas. (MEXU). Oaxaca: San Lucas Ojitlán, del poblado El Zapotal a Mata
Common names and uses: Tomatillo. No uses known. Caña, J. Calzada 14,314 (MEXU). Puebla: Xicotepec, 3 km adelante
Phenology: Flowering and fruiting from July to April, probably de Xicotepec de Juárez rumbo a Poza Rica, M. Cházaro 562 (MEXU).
throughout the year. Querétaro: en el pueblo de Concá en los nacimientos, municipio de
Conservation status: LC. Arroyo Seco, M. Martínez 3456 (QMEX). Quintana Roo: Othón P.

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FIGURE 9
Details of flowers and fruits of Physalis spp. (A) P. hintonii, (B) P. lagascae, (C) P. latiphysa, (D) P. leptophylla, (E) P. lignescens, (F) P. longiloba, (G) P.
longipedicellata, (H) P. melanocystis, (I) P. microcarpa, (J) P. minuta, (K) P. nicandroides, (L) P. orizabae.

Blanco en Tomás Garrido, E. Cabrera 17,013 (MEXU). San Luis 00077353!, MEXU!, NY 138871!, UC 104116!, VT not located).
Potosí: Xilitla, a 3 km aL NW de Huicihuayán, P. Tenorio 63 (Figures 1K, 6D).
(MEXU). Tabasco: Paraíso, M. Magaña 20 (MEXU). Perennial herb, 0.8–2 m long, weak and reclining on other
Tamaulipas: Ejido Allende 30 km aL NW de Ocampo, L. vegetation; stem viscid-pilose with articulating trichomes;
Hernández 1450 (MEXU). Veracruz: Rancho Guadalupe, 3 km W petiole 1.5–5.5 cm long, blades ovate, 5–10 cm long,
of Jalapa, J. Calzada 1893 (MEXU, XAL). 3.0–6.5 cm wide, apex acuminate, base cordate, margin entire
or usually with one to two irregular, obtuse teeth per side;
18. Physalis greenmanii Waterf., Rhodora 69: 226 (1967) solitary flowers on peduncles 5–15 mm long; calyx 1.2–1.4 cm
long; corolla rotate, pale yellow with five clusters of purple
Type: Mexico, Veracruz, Jalapa, C. G. Pringle 8104 (holotype: maculations, 15–28 mm wide, velvety-pubescent on the throat;
US barcode 00,027,326!, isotypes: F barcode V0073002F!, GH anthers purple, 2.8–3.5 mm long; fruiting peduncles 7–10 mm

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FIGURE 10
Geographic distribution of Physalis spp. (A) P. lagascae; (B) P. lassa, P. parvianthera, and P. tehuacanensis; (C) P. latiphysa, P. porrecta, and P.
purpurea; (D) P. leptophylla; (E) P. longiloba, P. lignescens, and P. tamayoi; (F) P. mcvaughii, P. minimaculata, and P. pringlei.

long, fruiting calyx 5-angled; 3.0–4.5 cm long, 2.5–3 cm wide, Diagnostic characters: A perennial herb with cordate leaves,
mature berry spherical, ca. 1.5 cm in diameter, seeds ca. 1.8 mm spotted maculations, corollas, and long purple anthers.
in diameter. Common names and uses: None known.
Distribution and habitat: Cloud forest, 1,000–2,000 m asl., Phenology: Flowering and fruiting from June to April, probably
endemic to Ver. (Figure 8B). throughout the year.

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Conservation status: EN. Distribution and habitat: It develops in thorny forests with
Representative specimens examined. MEXICO. Veracruz: Opuntia, Acacia, and Mimosa, or grasslands on rhyolitic rocks or
Xico, Cascada de Texolo, P. Zamora-Tavares et al. 212 (IBUG). close to bodies of water in sandy places. It grows from 1,880 to
2,020 m asl. Ags., Gto., Jal., Zac. (Figure 8C).
19. Physalis grisea (Waterf.) M. Martínez, Taxon 42 (1): 104 Diagnostic characters: Hastate leaves, almost prostrated habit,
(1993): (1993) and cinereal appearance of the plant.
Common names and uses: None known.
Type: USA, Massachusetts, Middlesex Co., Cambridge, W. Phenology: Flowering and fruiting from August to September.
Deane s.n. (holotype: GH barcode 00003293!, isotype: NY!). Conservation status: EN.
Physalis pubescens var. grisea Waterf. Rhodora 60 (714): 167–168. Representative specimens examined. MEXICO.
1958. (Figures 3I, 6H). Aguascalientes: Rincón de Romos, shrub-covered, nearly treeless
Erect annual herb, sericeous with non-glandular multicellular mountainsides ca. 20 km east of Rincón de Romos, road to Asientos,
trichomes; petioles 5–7 cm long, blades ovate to lanceolate, between cerro Altamira and cerro San Juan, R. McVaugh 23,758
10–20 cm long, 5.0–5.4 cm wide, apex acute to acuminate, base (MICH). Guanajuato: cerca de la Quebrada, municipio de Ocampo,
oblique, margins coarsely dentate; flowers solitary on peduncles J. Rzedowski 52,320 (IEB, QMEX). Jalisco: km 36 de la carretera
4–6 mm long; calyx with triangular lobes 1.5–2.2 mm long, corolla Ojuelos-Lagos de Moreno, cercano al arroyo del poblado La Troje,
rotate, yellow, 0.7–1.0 mm in diameter with simple dark purple O. Vargas 771 (IBUG). Zacatecas: Pinos, Los Alpes, 200 m al E del
maculations; filaments purple, anthers yellow or with a faint blue poblado, O. Vargas 769 (IBUG).
tinge, 1.1–2.0 mm long; fruiting peduncles 6–10 mm long, fruiting
calyx strongly 5-angled, 2.5–3.0 cm long, 1.0–2.2 cm wide, sericeous 21. Physalis hederifolia A. Gray, Proc. Amer. Acad. Arts 10: 65
throughout; mature berry yellow, spherical, 1.0–1.5 cm in diameter, (1874)
with numerous brown seeds 1.5–2.0 mm in diameter.
Distribution and habitat: A spontaneous plant in disturbed Type: United States. New Mexico, Western Texas to El Paso C.
areas. The plant is common in the United States but has been Wright 528 (lectotype selected by Pretz and Deanna (2020) GH
collected only recently in Mexico at 1,500 m asl in oak forest and barcode 00077275!, isolectotypes: BM barcode BM 000995493, GH
crop fields. Jal., Mich. barcode 00077273, K barcode K759425, US barcode 01178071 right-
Diagnostic characters: Annual, the leaves are orange-tinged. hand specimen only). (Figures 1L, 6G). Synonyms in Pretz and
Common names and uses: Tomatillo. The fruits are edible and Deanna (2020), except for P. puberula Fernald; see comments under
mostly sweet. P. chenopodiifolia Lam.
Phenology: Flowering from May to October. Perennial herb 60–80 cm high, branched from a woody base,
Conservation status: Not evaluated, but the species is widely pubescent with eglandular and short, glandular trichomes; leaves
distributed. alternate, petioles 1.6–2.6 cm long; blades ovate to subcordate,
Representative specimens examined. MEXICO. Jalisco: 2.4–3.5 cm long, 2.1–3.0 cm wide, apex acute, base cuneate, oblique
Zapopan, cerro El Tepopote, frente al poblado de La Venta del to truncate, margin serrated; flowers solitary on peduncle 5–8 mm long;
Astillero, ladera de exposición este, A. Rodríguez 6578 (IBUG, UAQ, calyx with ovate-to-triangular lobes 2–4 mm long, corolla campanulate
MEXU). Michoacán: Sahuayo, áreas de cultivo de Sahuayo, rotate, yellow, 0.7–1.2 cm in diameter, with five pale maculations;
J. Sánchez 122 (IBUG, UAQ, MEXU). anthers yellow, 3.5–4.0 mm long; fruiting peduncle 9–13 cm long,
fruiting calyx 10-costate with five more prominent angles, 2.5–2.8 cm
20. Physalis hastatula Waterf., Rhodora 69: 111 (1967) long, 1.7–2.2 cm wide, mature berry green to orange, ca. 9 mm in
diameter, with numerous yellow seeds 1.6–1.8 mm in diameter.
Type: Mexico, Jalisco, Cerro La Campana, near km 36 SW of Distribution and habitat: Desert areas with Larrea and
Ojuelos on the road to Aguascalientes, R. McVaugh 16778 Flourensia, disturbed places with calcareous soils. It grows from
(holotype: MICH barcode 1000037!; isotypes: ENCB!, LL!, OKLA 1,400 to 1,500 m asl. Chih., Coah., Dgo., Gro., Jal., N.L., Oax., Qro.,
not located). (Figures 3E, 6F). S.L.P., Son., Tamps., Zac. (Figure 8E).
Prostrate perennial herb, branched from the base, glabrous or Diagnostic characters: Densely pubescent, with almost
almost puberulent, with eglandular, short trichomes; leaves subcordate leaves, short peduncles, and almost immaculate
solitary or geminate, petiole up to 11 mm long; blades corollas, yellow anthers.
lanceolate to linear-lanceolate, often hastate, sometimes Common names and uses: None known.
asymmetrical, 1–5 cm long, 0.7–2.4 cm wide, apex acute, base Phenology: Flowering from April to June and fruiting in August.
oblique, cuneate, decurrent, margin entire or with one to two Conservation status: LC.
teeth on both sides; flowers solitary on peduncles 6–10 mm long; Representative specimens examined. MEXICO. Chihuahua:
calyx with narrow triangular lobes 3–5 mm long, corolla pasando San Juditas hacia San Buenaventura, M. Martínez 9641
campanulate rotate, yellow, 0.8–2.1 cm in diameter, with simple (QMEX). Coahuila: Las Margaritas, km 190 carretera San Pedro-
dark brown to yellowish-brown maculations; anthers yellow, Cuatro Ciénegas, M. Martínez 9616 (QMEX). Durango: Tayoltita:
2.0–2.5 mm long; fruiting peduncle 8–9 mm long, fruiting calyx El Pino, 20 km del entronque a Sapioris con la brecha Coyotes-san
10-costate, with five more prominent angles, 1.4–2.5 cm long, Miguel de Cruces, 24° 31′ N, 105° 49′ W, P. Tenorio 6318 (MEXU).
1.0–1.7 cm wide; mature berry green ca. 10 mm diameter, with Guerrero: Zihuatanejo de Azueta en El Bálsamo, J. Soto 8660
yellow seeds ca. 1.8 mm in diameter. (MEXU). Jalisco: Ixtlahuacán del Río, puente Paso de Guadalupe,

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Martínez et al. 10.3389/fgene.2023.1080176

Carrillo Reyes 3590 et al. (IBUG). Nuevo León: 3 km al S de la pubescent; anthers blue or yellow with blue lines 2.5–4 mm long;
desviación a Cerralvo, camino a Los Herrera, M. Martínez 9819 fruiting peduncle 20–30 mm long, fruiting calyx 5-angled,
(QMEX). Oaxaca: en las ruinas de Monte Albán entre los edificios, 35–50 mm long, 25–30 mm wide; mature berry spherical, 20 mm
en lugares húmedos, M. Martínez 1903 (MEXU). Querétaro: km in diameter, with numerous yellow seeds ca. 2 mm in diameter.
203 carretera a México, E. Argüelles 781 (MEXU). San Luis Potosí: Distribution and habitat: Pine–oak forest from 1,600 to
Salinas, gravelly desert along mex. 49, 33.4 miles E of Salinas 2,270 m asl. N.L.
(ENCB). Sonora: Naco, desviación a Naco, de la carr, Cananea- Diagnostic characters: A perennial herb with a mixture of
Agua Prieta, P. Tenorio 13,622 (MEXU). Tamaulipas: km 5 de la eglandular, glandular, and dendritic trichomes throughout the plant.
carretera vía corta a Ciudad Victoria, 31 km aL NE de Jaumave, M. Common names and uses: None known.
Martínez 9890 (QMEX). Zacatecas: Melchor Ocampo, near and at Phenology: Flowering and fruiting in July; no other collection
Sierra del Yeso, almost due west of La Presa de los Angeles, 25° 04′ N, dates are known.
102° 08′ 30″ W, Johnston 11,531 (MEXU). Conservation status: DD; the plant is known only from one
mountain range.
22. Physalis hintonii Waterf., Rhodora 69: 226 (1967) Representative specimens examined. MEXICO. Nuevo León:
municipio de Zaragoza, cerro El Viejo, J. Hinton 22,255 (Herb.
Type: Mexico, Estado de Mexico, Tejupilco, Dist. of Hinton).
Temascaltepec, G. B. Hinton 8457 (holotype: NY barcode 477364
!, isotypes: F 1498378!, CAS 4031!, ENCB not located, GBH not 24. Physalis ignota Britton, Mem. Torrey Bot. Club 16: 100 (1920)
located, GH 00077354, OKLA not located, US barcode 00027332!).
(Figure 9A). Type: Cuba Santa Clara Rio Arimao, Britton and Wilson 5767
Erect perennial herb, 30–120 cm tall; stem with few branches (holotype: NY barcode 620785!).
with dense dendritic trichomes up to 1 mm long throughout; basal = Physalis pentagona S.F.Blake, Contr. U.S. Natl. Herb. 24: 20
leaves geminate, petioles 10–35 mm long, blades ovate, 4–9 cm long, (1922). Type: Guatemala, Izabal, Los Amates, S.F.Blake 7313
4–5 cm wide, apex acute, base truncate to cordate, margin undulate (holotype: US barcode 00027352!)
to dentate; solitary flowers on peduncles 2.5–5.0 cm long; calyx with Erect annual herb, 10–100 cm high, usually densely covered
deltoid lobes 5–10 mm long, corolla campanulate-rotate, 17–20 mm with short glutinous, multicellular trichomes throughout;
wide, with five dark brown maculations; anthers violet, 3–4 mm petioles 2–7 cm long, blades ovate to rhombic-ovate, 4–12 cm
long; fruiting peduncle 10–16 mm long, fruiting calyx 10-costate, long, 3–8 cm wide, apex acuminate, base oblique to cordate,
open at the apex, 25–35 mm long and 18–22 mm wide; mature berry margin entire to repand-dentate; flowers solitary on peduncles
ca. 1 cm in diameter with numerous yellow seeds ca. 1 mm in 3–7 mm long; calyx with triangular lobes; corolla campanulate,
diameter. yellowish, 6–10 mm in diameter, sparsely hairy in the throat,
Distribution and habitat: Pine–oak forest; it grows at immaculate; anthers yellow or bluish tinged, 2.0–2.5 mm long;
1,200–2,100 m asl. Edo. Mex., Mich., Ver. (Figure 8D). fruiting peduncle 7–15 mm long, fruiting calyx strongly 5-
Diagnostic characters: P. hintonii is easily recognized by its angled, 2.8–5 cm long, 2–4 cm wide, usually densely and
spreading dendritic (several-branched) trichomes and its large evenly covered with short, erect, jointed trichomes; mature
flowering calyx with broad sepals. berry spheric to ovoid, 9–15 mm in diameter with numerous
Common names and uses: None known. yellow seeds ca. 1.5 mm in diameter.
Phenology: Flowering and fruiting from March to July. Distribution and habitat: In wet or moist scrub, sometimes on
Conservation status: LC. stony or rocky slopes from sea level to 1,250 m. Chis., Oax.
Representative specimens examined. MEXICO. Estado de México: (Figure 8F).
Tenancingo, al O de Exhacienda San José Tenería, P. Zamora-Tavares Diagnostic characters: 5-angled fruiting calyces, the stem,
et al. 272 (IBUG). Michoacán: Presa la Yerbabuena, municipio de peduncles, and fruiting calyx densely pubescent with gray, short
Tlazazalca, E. Pérez 1600 (IEB, QMEX). Veracruz: Jilotepec, Piedra trichomes.
de Agua, F. Ventura 9820 (ENCB, F, XAL). Common names and uses: Vejiga de perro, tomatillo, soplón,
farolito. No uses known.
23. Physalis hunzikeriana M. Martínez, Kurtziana 27 (2): 383 Phenology: Flowering and fruiting throughout the year.
(−385, 1999) Conservation status: LC.
Representative specimens examined. MEXICO. Chiapas:
Type: Mexico, Nuevo León, Cerro el Viejo, J. Hinton et al., 22104 Amatenango de la Frontera, steep slopes and dry ravines along
(holotype: GBH!). Río Cuilco between Nuevo Amatenango and Frontera Comalapa D.
Erect perennial herb 30–100 cm high, stems with eglandular and Breedlove 41,575 (MEXU). Oaxaca: Santo Domingo Tehuantepec,
glandular trichomes mixed with dendritic; leaves alternate, petioles Llano de Lumbre, Santiago Lachiguirí, Jalapa del Marqués E. Cruz
1.5–5.0 cm long; blades ovate, 6–11 cm long, 4.5–5.5 cm wide, apex 97a (MEXU).
acute to acuminate, base rounded to attenuate, margin dentate with
three to six teeth per side, pubescent with glandular and dendritic 25. Physalis lagascae Roem. & Schult., Syst. Veg., ed. 15 bis
trichomes; flowers solitary on peduncles 5–8 mm long; calyx with [Roemer & Schultes] 4: 679 (1819)
lanceolate lobes 10 mm long, corolla rotate, yellow, 22–25 mm in
diameter with five purple maculations, corolla throat densely Type: based on P. parviflora Lagasca. (Figures 3D, 9B).

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Martínez et al. 10.3389/fgene.2023.1080176

= Physalis lagascae var. glabrescens O.E.Schulz, Symb. Antill. M. Magaña 1918 MEXU). Veracruz: Coatzintla, Palamar de Zapata,
(Urban). 6 (1): 147 (1909). Type: Cuba, provincia de la Habana Cortés 125 (MEXU). Yucatán: Hacienda Cancabchen A. Mizrahi s.
prope Calabazar, Wright 3636, (holotype: US barcode 00027335!) n. (UADY). Zacatecas: Jalpa, pastured hills five miles southwest of
= Physalis parviculea S.F.Blake, Contr. U.S. Natl. Herb. 24: 20 Jalpa, R. McVaugh 18,513 (MICH).
(1922). Type: Guatemala, Izabal, Los Amates, S.F.Blake
7318 holotype: US barcode 00027348!) 26 Physalis lassa Standl. & Steyerm., Publ. Field Mus. Nat. Hist.,
Erect annual herb 15–70 cm tall; stem branched, with adpressed Bot. Ser. 23: 19 (1943)
whitish trichomes; leaves geminate, petioles 1.3–4.6 (–8) cm long;
blades ovate to suborbicular ovate, 2.6–7.0 cm long, 1.3–4.2 cm Type: Guatemala, Dept. Jalapa, grassy thickets, between Jalapa
wide, apex acuminate, base oblique to subcordate, margin entire and Montaña Miramundo, J. Steyermark 32868 (holotype: F barcode
to sinuate lobed; flowers solitary on peduncle 3–8 mm long; calyx 0077205F!).
with ovate lobes 1.5–2.5 mm long, corolla campanulate, rotated to Erect to prostrate perennial herb, branched, up to 50 cm tall,
pentagonal, yellow, 6–8 mm in diameter, with five almost black densely pubescent with eglandular septate trichomes mixed with
maculations; corolla throat pubescent; anthers yellow or blue 2 mm some glandular, especially on the fruiting calyx; leaves alternate to
long; fruiting peduncle ca. 4 mm long, fruiting calyx 10-costate, geminate, petioles 1.0–3.8 cm long; blades ovate to suborbicular
1.2–2.0 cm long, 1.2–2.0 cm wide; mature berry ca. 10 mm diameter ovate, 3.5–7.5 cm long, 1.7–4.9 cm wide, apex acute to obtuse, base
with numerous yellow seeds ca. 2 mm in diameter. oblique to truncate, margin entire to sinuate or with few teeth;
Distribution and habitat: The species is generally found in flowers solitary on peduncles 5–7 mm long; calyx with triangular
secondary vegetation as a weed, in grasslands, or associated with lobes ca. 3 mm long, corolla rotate-campanulate, yellow, 10 mm in
tropical deciduous forest. It grows from 0 to 2,100 m asl. Ags., diameter, with five very dark purple maculations; anthers blue,
Camp., Chis., Chih., Col., Dgo., Edo. Mex., Gto., Gro., Jal., Mich., 2.5–3.0 mm long; fruiting peduncle 0.9–1.7 cm long, fruiting
Mor., Nay., N.L., Oax., Pue., Qro., Q. Roo, Sin., Tab., Ver., Yuc., Zac. calyx 10-costate, 2.6–3.5 cm long, 1.9–2.2 cm wide, pubescent;
(Figure 10A). mature berry 1.0–1.5 cm in diameter with numerous yellow seeds
Diagnostic characters: Densely pubescent at the stem base. The ca. 2 mm in diameter.
species is easily recognized by the presence of eglandular, white Distribution and habitat: It grows at the margins of the
conspicuous trichomes in flower buds and calyces, as well as small mountain–cloud forest at an altitude of 1,500 m. Chis., Col.
corollas with contrasting dark, triangular maculations. (Figure 10B).
Common names and uses: Tomatillo, vejiga de perro, Diagnostic characters: Corolla large, with contrasting
chimpululu. No uses known. maculations, large fruiting calyx. The long, glandular, dense
Phenology: Flowering and fruiting from June to December. pubescence gives P. lassa a wooly appearance with reddish-brown
Conservation status: LC. coloration.
Representative specimens examined. MEXICO. Common names and uses: Tomatillo; no uses known.
Aguascalientes: 6.5 km aL SW de Tanque de Los Jiménez, Phenology: Flowering and fruiting in November.
J. Sierra 881 (QMEX). Campeche: en las Palomas, al sur de Conservation status: LC.
Candelaria, sobre el Río Candelaria, E. Cabrera 4809 (MEXU). Representative specimens examined. MEXICO. Chiapas: La
Chiapas: D. Breedlove 26,384 (MO). Chihuahua: Temósachic, Grandeza, E. Matuda 1556 (MEXU). Colima: Comala, rancho El
Nabogame, J. LaFerrière 1551 (MEXU). Colima: Comala: Jabalí, 20 km (airline) N of Colima in the SW foothills of the Volcán
Noguera, más o menos 13 km aL norte de Comala, M. Arreguín de Colima, El Agostadero, ridgetop in hills above the shrine 4.9 km
754 (MEXU). Durango: 5 km aL NW de Yerbanis, km 128 carretera SE of Hac. San Antonio on the road to Comala, J. Sanders 11,807
Yerbanís-Cuencamé, M. Martínez 9782 (QMEX). Estado de (MEXU, MICH).
México: Temascaltepec, Pantoja, G. B. Hinton 8606 (MEXU).
Guanajuato: Abasolo: 35 km aL SW de Cuerámaro, sobre el 27. Physalis latecorollata Waterf., Rhodora 69: 117 (1967)
camino a la Barranca del Chilar, J. Rzedowski 45,015 (CIIDIR-
SGO, ENCB, MICH). Guerrero: Rincon de la Via, H. Kruse 541 Type: Mexico, Oaxaca, orilla de arroyo, campamento Rio Molino
(MEXU). Jalisco: Tala, márgenes del arroyo Caliente, 100 m antes de cerca de San Miguel Suchitepec, Garcia s.n. (holotype: ENCB!).
su nacimiento, Bosque Escuela, La Primavera, A. Rodríguez 1471 Perennial herb, 30 cm tall; stems villous, with articulated
(IBUG, IEB, MEXU). Michoacán: Apatzingán, 6 km aL noroeste de trichomes up to 2–3 mm long; petioles 15–25 mm long; blades
Apatzingán, O. Vargas 853 (IBUG). Morelos: Amacuzac, noroeste ovate, 5–6 cm long and 3.8–4.2 cm wide, apex acute, base
de Amacuzac, L. Estrada 1295 (MEXU). Nayarit: La Yesca, La cuneate, margin entire; flowers solitary on peduncle 5–7 mm
Manga, Bugarín s. n. (CHAPA, IBUG). Nuevo León: Mt. long; calyx 8–9 mm long and 7–8 mm wide; corolla 33 mm in
Obispado, J. Roybal 1071 (MEXU). Oaxaca: Yomillin Cañon, C. diameter with violaceous maculations; anthers purple, ca. 3 mm
Pringle 7004 (MEXU). Puebla: Matamoros, F. Miranda 2390 long; fruiting calyx, mature berry and seeds unknown.
(MEXU). Querétaro; San Juan del Río, km 175.5 carretera Distribution and habitat: Riparian vegetation, 2,500 m asl.
Querétaro-México en la desviación a San Clemente, M. Martínez Oax. (Figure 8D).
2990 (QMEX). Quintana Roo: a 18 km al W de Ucum, S de E de Diagnostic characters: The species is only known from the type
Nachi Cocom, O. Tellez 2412 (MEXU). Sinaloa: Culiacán, Tierras specimen, which lacks fruits. Its woody rhizome, pubescence, and
Blancas 2 km al N de El Rincón, brecha a Tamazula, P. Tenorio 8299 the shape of the leaves distinguish it from P. orizabae.
(MEXU). Tabasco: centro por el km 11 de Villahermosa a Cardenas, Common names and uses: None known.

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Martínez et al. 10.3389/fgene.2023.1080176

FIGURE 11
Geographic distribution of Physalis spp. (A) P. melanocystis, (B) P. nicandroides, (C) P. orizabae, (D) P. patula, (E) P. philadelphica, (F) P. pruinosa.

Phenology: Fflowering in September. 28. Physalis latiphysa Waterf., Rhodora 60: 169 (1958).
Conservation status: DD.
Representative specimens examined. MEXICO. Oaxaca: orilla Type: USA, Arizona. Pima County. Ronstadt Ranch, plain east of
de arroyo, campamento Rio Molino cerca de San Miguel Suchitepec, Baboquivari Mountains, T.H. Kearney & R.H. Peebles 14425 (holotype:
Garcia s.n. (ENCB). ARIZ catalog 90142!, isotype: OKLA 101862). (Figures 1M, 9C).

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Martínez et al. 10.3389/fgene.2023.1080176

Annual herb, 15–200 cm tall, branched, more-or-less villous or Ocampo-Moris, km 48.5, M. Martínez 9663 (QMEX). Colima:
glandular villous; petioles 1.5–7.0 cm long; blades ovate, 5–7 cm Manzanillo, steep hills with occasional rock outcrops, ca.
long, 3–5 cm wide, apex acuminate, base truncate, sometimes 14 miles west-north-west of Santiago, road to Cihuatlán, Jalisco,
oblique, margin entire to few-toothed; flowers solitary on R. McVaugh 20,782 (ENCB, MICH). Durango: 10 mi N of
peduncles 3–8 mm long; calyx with narrowly lanceolate to Tamazula, A. Gentry 5270 (CAS, NY). Estado de México:
acuminate lobes, corolla rotate, yellowish 3–4 mm in diameter Temascaltepec, at Bejucos, G. Hinton 5204 (US). Guerrero:
with five small, dark maculations; fruiting peduncles 1.0–1.5 cm Leonardo Bravo 14 km camino Chichihualco-Filo de Caballos,
long, fruiting calyx strongly 5-angled, sparsely appressed-hairy, J. Calónico 4395 (MEXU). Jalisco: Puerto Vallarta, along the
2.5–4 cm long, 3–4 cm wide; mature berry brown, 13–17 mm in highway in the gorge of the Rio Horcones, south of Puerto
diameter with numerous dark brown seeds ca. 2 mm diameter. Vallarta, about 11 km from Puerto Vallarta, Dieterle 4083
Distribution and habitat: It grows in tropical deciduous forest (ENCB). Michoacán: Lázaro Cárdenas: steep hills about
at an altitude of 1,500–2,000 m. Gto., Mich., Son. (Figure 10C). 45–48 km south of Arteaga, 12–15 km north of Playa Azul,
Diagnostic characters: It is distinguished from other species McVaugh 22,605 (ENCB, MICH). Nayarit: La Yesca, Paso de los
that have linear-subulate calyx lobes by its corollas with dark Bueyes, río Santiago, 12 km al E de Mojarras, brecha a Huajimic, 21°
maculations and fruiting peduncles longer than those of P. 29′ N, 104° 32′ W, P. Tenorio 6841 (MEXU, MICH). Oaxaca: Arroyo
nicandroides, but shorter than those of P. pruinosa. de Piedra ladera W, entrando por el Mármol, Cerro Guiengola.
Common names and uses: None known. Distr. Tehuantepec, L. Torres 233 (MEXU). Sinaloa: Concordia, El
Phenology: Flowering and fruiting from May to November and Cantil 32 km aL NE de Concordia, carretera Mazatlán-Durango, P.
probably throughout the year. Tenorio 2957 (MEXU). Sonora: Río Mayo Region arroyo Las
Conservation status: LC. Rastras, southwest edge of the Sierra de Alamos, T. Van
Representative specimens examined. MEXICO. Guanajuato: Devender 95-134A (ASU).
4 km al N de Uriangato, municipio de Uriangato, J. Rzedowski 51820
(IEB, QMEX). Michoacán: 7 km al E de Villa Jiménez, sobre el camino a 30. Physalis lignescens Waterf., Rhodora 69: 231 (1967)
Copándaro, J. Rzedowski 40745 (IEB, QMEX). Sonora: 19 km al E de
Yécora, orillas de la carretera, M. Martínez 9704 (QMEX). Type: Mexico: Jalisco, NE slopes of the Nevado de Colima,
below Canoa de Leoncito, R. McVaugh 13454 (holotype: OKLA
29. Physalis leptophylla B. L. Rob. & Greenm., Proc. Amer. Acad. 100185, isotypes: MICH barcode 1109905!, US barcode 00027336,
Arts 29: 389 (1894) MEXU barcode 114184!). (Figure 9E).
Erect perennial herb up to 50 cm and then becoming
Type: Mexico, Sinaloa, near Mazatlán, Wright 1252 (lectotype prostrate up to 1 m long, with few branches, often leans on
designated by D’Arcy, Ann. Missouri Bot. Gard. 60: 6641973 GH other plants; pubescent with eglandular articulated trichomes of
barcode 00003285!). (Figure 9D). different lengths; leaves alternate at the base, geminate toward
Annual herb, branched, spreading, up to 1 m tall, with eglandular, the apex, petiole 1–3.4 cm long; blades ovate to suborbicular
brown glandular trichomes; leaves alternate, petioles 0.9–3.8 (−7.2) cm ovate, 4.5–9.0 cm long, 2.4–6.1 cm wide, apex acute to short
long; blades ovate to suborbicular ovate, 2.7–8.0 cm long, 1.8–4.8 cm acuminate, base oblique, decurrent, or cuneate, margin entire,
wide, apex acuminate, base oblique, margin entire; flowers solitary on repand, or with several acute, irregular teeth; flowers solitary on
peduncles 4–5 mm long; calyx with ovate lobes 3–4 mm long, corolla peduncles 8–17 mm long; calyx 4–5 mm long, corolla
campanulate to rotate, whitish to pale yellow, 0.7–1.4 cm in diameter, campanulate rotate, yellow, 1.5–2.1 cm in diameter, with
with five simple, reddish-brown to dark purple-brown maculations; brown compound maculations; anthers purple 2.5–3.5 mm
anthers blue, 1.5–2.5 mm long; fruiting peduncle 6–8 mm long, fruiting long; fruiting peduncle 0.8–1.7 cm long, fruiting calyx 5-
calyx 10-costate, 1.5–2.5 cm long, 1.4–1.7 cm wide, green, translucent angled 2.0–4.1 cm long, 1.4–2.6 cm wide, reticulate, pubescent
when dry; mature berry spherical, 4–9 mm in diameter, with numerous only on the lobes; mature berry 6–12 mm in diameter, yellow
yellow seeds ca. 1 mm in diameter. seeds, 2–2.5 mm in diameter.
Distribution and habitat: Tropical deciduous and dry oak Distribution and habitat: Abundant along the paths and
forest along the Mexican Pacific coast from 50 to 800 m asl. margins of pine–oak and cloud forest, in clearings or ravines. It
B.C.S, Chih., Col., Dgo., Edo. Mex. Gro., Jal., Mich., Nay., Oax., grows between 2,000 and 2,300 m asl. Endemic to Parque Nacional
Sin., Son. (Figure 10D). Nevado de Colima, Jal. (Figure 10E).
Diagnostic characters: Entire leaves, acuminate and translucent Diagnostic characters: prostrate habit, lignified base, short
when dry, small dark maculations, contrasting with the corolla, and flower calyx with reflexed lobes, corolla with compound
calyx 10-costate in fruit. The pubescence is glandular. maculations and 5-angled fruiting calyx.
Common names and uses: Tomate, tomatillo, tomate del Common names and uses: None known.
monte, shiquipiltzi. The ripe fruits are cooked in Sonora. Phenology: It flowers and bears fruit abundantly from July to
Phenology: Flowering and fruiting from October to February. August. Later, it is found with flowers or scarce fruits.
Conservation status: LC. Conservation status: EN.
Representative specimens examined. MEXICO. Baja Representative specimens examined. MEXICO. Jalisco:
California Sur: 2 km aL NE de Misión de San Javier, km 32 de Venustiano Carranza: puerto El Floripondio, camino a la estación
la carretera, M. Martínez 9966 (QMEX). Chihuahua: camino de Microondas Las Víboras, A. Rodríguez 913 (ENCB, IBUG, MEXU).

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31. Physalis longiloba O. Vargas, M. Martínez & Dávila. Brittonia long, 2.0–2.5 cm wide, very large for the berry; mature berry
53 (4): 505–507 (2001) 1–1.2 mm in diameter.
Distribution and habitat: It develops in protected places, on
Type: Mexico. Jalisco: Autlán de Navarro, Sierra de Manantlán, slopes or margins of the cloud forest. The populations are small, with
passing Rincón de Manantlán along the creek, 19 35′55″ N, only one to three plants found at each location. It grows at an
104 12′35″ W, O. Vargas 873 (holotype: IBUG!, isotypes: altitude range of 2,100–2,200 m. Jalisco (endemic). Known only
MEXU!, NY barcode 468345!, ENCB!, IEB!, ZEA!). (Figure 9F). from the Sierra de Manantlán.
Prostrate perennial herb, 1 m long, branched, glabrescent or Diagnostic characters: Its prostrate habit and large plant size.
with eglandular, short, appressed, puberulent trichomes; leaves The leaves with long, glossy acumens and the long flowering and
alternate at the base but soon geminate, petiole 1.5–3.6 (−4.6) fruiting peduncles, as well as the wide corollas with compound
mm long; blades ovate to suborbicular ovate, 3.6–7.9 (–10.2) cm maculations and the 5-angled calyx in the fruit.
long, 2.1–6.1 cm wide, apex acuminate, base oblique, cuneate to Common names and uses: None known.
subcordate, margin entire, wavy, or toothed; flowers solitary on Phenology: It flowers and bears fruit from September to April.
peduncles 6–15 mm long; calyx lobes 5.5–9.0 mm long; corolla Conservation status: LC.
rotate campanulate, reflected, yellow, 1.5–2.2 cm in diameter, Representative specimens examined. MEXICO. Jalisco:
with compound maculations, brown, corolla throat pubescent; Cuautitlán de García Barragán, Sierra de Manantlán, 500 m antes
anthers purple, blue when dry, 3–4 mm long; fruiting peduncle de llegar a Tierras Blancas, por el camino a Llanos de San Miguel, O.
0.8–1.4 (−2.2) cm long, fruiting calyx 5-angled, 3.0–4.2 cm long, Vargas 868 (IBUG)
1.3–2.0 cm wide, puberulent; mature berry ca. 11 mm in diameter
with yellow seeds 1–1.5 mm in diameter. 33. Physalis mcvaughii Waterf., Rhodora 69: 104 (1967)
Distribution and habitat: It grows on slopes, open areas, or
paths in cloud forest, very humid pine–oak forest, or near bodies of Type: Mexico, Jalisco, two to three miles NW of San Miguel de la
water; abundant at 2,100 m. Jalisco (endemic). Known only from the Sierra, R. McVaugh 22,034 (holotype: MICH barcode 1109903!
Sierra de Manantlán, Jal. (Figure 10E). isotypes LL not located, NY barcode 138876!).
Diagnostic characters: The shape and length of the floriferous = Physalis jaliscensis Waterf., Rhodora 69: 231 (1967). Type:
calyx lobes and the 5-angled calyx in fruit, which reaches a large size, Mexico, Jalisco, steep mountainsides and barrancas one to two miles
in peduncles smaller than 2 cm. N of sawmill “La Cumbre” on the divide above the headwaters of Rio
Common names and uses: None known. Mascoto [Mascota], 25–30 km SE of Talpa de Allende, R. McVaugh
Phenology: It flowers and bears fruit from September to May. 21512 (holotype: MICH barcode 1109906!, isotypes: ENCB!, OKLA
Conservation status: LC. not located, US barcode 00027334!).
Representative specimens examined. MEXICO. Jalisco: Autlán Shrub up to 2 m tall, glabrous or glabrescent, eglandular
de Navarro, predio Las Joyas, cerca del letrero del Sendero ecológico, trichomes; leaves alternate or geminate, petiole 1.5–5.5 cm long;
O. Vargas 862 (IBUG, MEXU, ENCB). blades ovate to rhombic-ovate, 7–13 cm long, 4–9 cm wide, the apex
gradually attenuated, base oblique, decurrent, margin entire or with
32. Physalis longipedicellata Waterf., Rhodora 69: 230 (1967) some teeth; solitary flower on peduncle 2.5–3 cm long; calyx
8–15 mm long and 7–8 mm wide, deltoid lobes 3–5 mm long;
Type: Mexico: Jalisco, Cuautitlán de García Barragán, Sierra corolla campanulate, yellow, 1.1–1.4 cm long, 2.0–2.5 cm in
de Manantlán near Aserradero El Cuartón, 15–20 miles SE of diameter, compound maculations, brown; anthers violaceous,
Autlán, R. McVaugh 13828 (holotype: MICH barcode 1109904!, 2.7–3.5 mm long; fruiting peduncle 4.0–5.5 cm long, fruiting
isotypes: BRIT 23921!, G barcode G00343149!, K barcode calyx 10-costate, 2.5–4.5 cm long, 2.3–3.5 cm wide; mature berry
K000042281, OKLA not located, MEXU!, US barcode 1.5–2 cm in diameter.
00027337!). (Figure 9G). Distribution and habitat: It grows in cloud mountain forest at
Erect perennial herb, or with the main stem prostrate when it 1,700–2,000 m. Jalisco (endemic). It is known only from the Sierra
reaches dimensions of about 2 m long, pubescent with de Manantlán and Cacoma (Figure 10F).
eglandular, short, white trichomes; leaves geminate, petioles Diagnostic characters: Large glabrous to glabrescent plant, wide
3.6–6.7 (−9.2) cm long; blades sub-orbicular ovate, corollas with compound maculations. The length of the peduncles in
8.5–16.5 cm long, 6.6–11 cm wide, apex acute to acuminate, flower and fruit is a diagnostic character, as is the 10-costate fruiting
base oblique, cuneate to subcordate, decurrent, winged, with a calyx.
central groove where numerous trichomes accumulate, margin Common names and uses: None known.
entire, sinuate, or with some irregular, shallow, acute teeth; Phenology: Flowers and fruits from November to April.
flowers solitary on peduncles 1.7–4.5 cm long; calyx 8–11 mm Conservation status: NT.
long, 6–8 mm wide at the base of the triangular acuminate to Representative specimens examined. MEXICO. Jalisco: Ayutla,
ovate-deltoid lobes; corolla rotate campanulate, yellow, headwaters of Rio Mascota [20–25 km, airline, southeast of Talpa de
2.2–2.4 cm in diameter, with dark purple or brown compound Allende], narrow valley of steep mountain stream ascending to the west
maculations; anthers purplish-blue, 2.0–3.5 mm long; fruiting from a point 12–13 km above (south of) El Rincón, on the road to
peduncle (1.7) 4–6 cm long, fruiting calyx 5-angled, 3.3–4.0 cm Aserradero La Cumbre, R. McVaugh 23467 (ENCB, MICH).

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Martínez et al. 10.3389/fgene.2023.1080176

1.9–2.5 cm long, 1.5–2.0 cm wide, somewhat leathery, green or

purple; mature berry orange, dark green in herbarium specimens,
1.0 cm in diameter with numerous yellow seeds ca. 1.5 mm in
diameter.
Distribution and habitat: In tropical deciduous, oak, and
cloud forest. It is found at altitudes of 850–1,500 m. Camp.,
Chis., Col., Gro., Hgo., Jal., Oax., Qro., S.L.P., Tab., Tamps.,
Ver. (Figure 11A).
Diagnostic characters: A shrubby habit, narrow leaves, and
aggregated, greenish-yellow flowers. The berries are an intense
orange color, and the fruiting calyx is purple to almost black.
Common names and uses: None known.
Phenology: Flowering and fruiting through the year.
Conservation status: LC.
Representative specimens examined. MEXICO. Campeche:
Calakmul, núcleo arqueológico de Calakmul, C. Gutiérrez 5291
(MEXU). Chiapas: Tzimol, a 24 km aL este de Pujiltic, sobre la
carretera Venustiano Carranza-Tzimol, E. Cabrera 3018 (MEXU).
Colima:18 km aL SE de Colima, carretera Colima-Manzanillo,
Santana Michel 904 (IBUG). Guerrero: Zihuatanejo de Azueta
800 m aL SO del caserío La Vainilla, camino al mirador, C.
Gallardo 663 (MEXU) Hidalgo: Chapulhuacán, el Capulín 6 km,
del entronque de la brecha a Pisaflores con la carretera Jacala-
Tamazunchale, P. Tenorio 2372 (MEXU). Jalisco: Ahualulco de
Mercado, Piedras Bolas, Corona Oceguera s.n. (IBUG). Oaxaca:
Salina Cruz a Santa. Clara, en la subida de Cerro Marimba, al NO de
Salina Cruz, 22 km aL NO de Morro Mazatán, R. Torres 8933
FIGURE 12
Details of flowers and fruits of Physalis spp. (A) P. patula, (B) P.
(MEXU). Querétaro: km 6 de la brecha de Agua Zarca a
philadelphica, (C) P. porrecta, (D) P. pringlei, (E) P. pruinosa, (F) P. Neblinas, municipio de Landa de Matamoros, S. Zamudio 6744
pubescens, (G) P. queretaroensis, (H) P. purpurea. (IEB, QMEX). San Luis Potosí: Tamasopo, Puerto Verde, 64 km W
of Cd. Valles on Hwy. to Rioverde, P. Fryxell 3599 (MEXU).
Tabasco: en el campamento de la escollera y 5 km de Tenosique
hacia E. Zapata (ENCB). Tamaulipas: Gómez Farías, a 1 km de Alta
34. Physalis melanocystis (B. L. Rob.) Bitter, Repert. Spec. Nov. Cima en el camino a Rancho del Cielo, M. Martínez 1940 (MEXU).
Regni Veg. 20: 369 (1924) Veracruz: Dos Ríos, Rinconada, Ventura 3997 (ENCB, F).

Type: Mexico, San Luis Potosi, C. G. Pringle 3285 (holotype: GH 35. Physalis microcarpa Urb. & Ekman, Ark. Bot. 21A (5): 59
barcode 00003297!, isotypes: A 00003298!, AC not located, BR not (1927).
located, COLO not located, E not located, HBG not located, MO
barcode 022277, MA not located, MU not located, NDG not located, Type: Hispaniola, Santo Domingo in Llanura de Vega prope
NY 172306!, OKLA not located, PH PH00029454, RSA not located, Monte-Cristi, Ekman 5841 (lectotype designated here S 04-2808!).
US barcode 00027310!, barcode 00027311!). Withania melanocystis (Figures 1N, 2E, 9I).
B. L. Rob., Proc. Amer. Acad. Arts 26:171–172. 1891. (Figure 9H). Erect, branched annual herb 12–40 cm tall, glabrous or
= Physalis melanocystis var. melanocystis. pubescence with few eglandular trichomes; leaves alternate,
= Physalis melanocystis var. cernua (Donn. Sm.) Waterf. petioles 0.8–1.0 (−2.5) cm long; blades lanceolate to ovate,
Rhodora 69: 99 (1967). 1.4–9.0 cm long, 0.4–1.8 cm wide, apex acute to acuminate, base
= Physalis porphyrophysa Donn. Sm., Bot. Gaz. 61: 377 (1916). oblique, margin entire, pubescent; flowers solitary on peduncles
Type: Guatemala, Zacapa, H. Pittier 1754 (holotype: US barcode 2–4 mm long; calyx, lobes ca. 2 mm long, deltoid; corolla
00027356!; isotypes: F barcode V0077209F!, NY not located). campanulate to tubular, whitish, 1.2–2.0 mm in diameter,
Erect shrub up to 1.5 m tall, branched, glabrous in appearance, immaculate; anthers blue, 1.4–2.1 mm long; fruiting peduncle
with short, eglandular trichomes; leaves solitary, soon geminate, 4–5 mm long, fruiting calyx 10-costate, 1.1–1.5 cm long,
lanceolate to ovate-lanceolate, 4.0–7.0 cm long, 1.0–3.0 cm wide, 0.7–1.0 cm wide, glabrous; mature berry ca. 7 mm in diameter,
apex acute to acuminate, base cuneate, margin entire; flowers with yellowish seeds.
fascicled with one to three flowers, peduncle less than 1.0 cm Distribution and habitat: In mangroves, riparian, and pine–oak
long; calyx with deltoid lobes 2.5–3.0 mm long; corolla forests, in secondary vegetation, and near corn fields. It grows at an
campanulate stellate, greenish-yellow 0.8–1.1 cm in diameter, altitude range of 750–1,400 m. Chis., Edo. Mex., Jal., Son.
inconspicuous maculations; anthers yellow, 3 mm long; fruiting Diagnostic characters: The species has a fragile appearance and
peduncles 1.0–1.2 cm long, thickened, fruiting calyx 10-costate is small; the leaves are narrow, lanceolate. The fruiting calyx is

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Martínez et al. 10.3389/fgene.2023.1080176

almost transparent. Corollas are tiny, white, immaculate, almost campanulate, yellow, translucent, 4–9 mm diameter, immaculate
tubular, and not longer than 4 mm. or with five separate reddish-brown maculations; anthers yellow,
Common names and uses: Miltomate de culebra, tomatillo. No 1–2 mm long; fruiting peduncle 7–9 mm long, fruiting calyx 5-
uses known. angled with a pentagonal base, 1.1–2.2 cm long, 1.1–1.8 cm wide,
Phenology: Flowering and fruiting in August, probably until pubescent; mature berry ellipsoid ca. 8 mm diameter, yellow seeds
October. 1.5–1.8 mm.
Conservation status: LC. Distribution and habitat: It grows in coastal dunes and sandy
Representative specimens examined. MEXICO. Chiapas: places, protected between rocks near cliffs along the Pacific Ocean
Motozintla, outwash plain below Motozintla, D. Breedlove 40,607 coast. It is usually scarce, but it can cover spaces of up to 3 m. It
(MEXU, DS). Jalisco: Cuautitlán de García Barragán, Ayotitlán, develops at an altitude range of 0–200 m. Col., Chis., Gro., Jal., Nay.,
Piedra Gorda, 1.5 km por la vereda que va de Las Guayabillas a El Oax. (Figure 8F).
Maguey, O. Vargas 801 (IBUG). Sonora: Yécora, mesa del Diagnostic characters: A small light-yellow corolla and tiny
Campanero, T. Van Devender 96-387A (QMEX). maculations with reddish tones, 5-angled fruiting calyx, herbaceous,
with thin peduncles.
36. Physalis minimaculata Waterf., Rhodora 69: 219 (1967) Common names and uses: None known.
Phenology: Flowering and fruiting from June to April, so
Type: Mexico, Michoacán, old lava flows four miles NW of probably throughout the year; its presence increases after the
Apatzingán, R. McVaugh 17902 (holotype: MICH barcode rainy season.
1109901!, isotypes: ENCB!, US barcode 00027343!) Conservation status: LC.
Erect to prostrate annual herb, up to 60 cm long, branched, Representative specimens examined. MEXICO. Colima:
the entire plant pubescent, trichomes eglandular or glandular; Manzanillo, along the road between El Ciruelo and Cuyutlán,
leaves geminate, petiole 0.4–1.7 cm long; blades ovate, 1.4–2.6 cm Gilly 11 (MICH). Chiapas: Tuxtla Gutiérrez, D. Breedlove 19,991
long, 1.0–1.8 cm wide, apex acute, base oblique, margin sinuate- (MO). Guerrero: Agua de Obispo, Mochitlán, H. Kruse 813
lobate to toothed; flowers solitary on peduncle 5–6 mm long; (MEXU). Jalisco: Puerto Vallarta: a lo largo de la playa entre
calyx with narrow triangular lobes ca. 2 mm long; corolla Villa Varadero y la desembocadura del rio Ameca, Cházaro 6331
campanulate rotate, with, yellow, 0.7–1.0 cm in diameter, with (IEB, WIS). Nayarit: Bahía de Banderas, cerro de Punta Mita,
conspicuous simple dark purple-brown maculations; anthers Ramírez 4024 (IBUG). Oaxaca: Santo Domingo Tehuantepec,
deep purple, up to 4 mm long; fruiting peduncle 5–8 mm long, Distrito de Tehuantepec, Buenos Aires, 15 km al W de
fruiting calyx 10-costate 0.9–1.9 cm long, 0.5–1.5 cm wide; Tehuantepec, C. Martínez 671 (MEXU).
mature berry ca. 1.2 cm in diameter, with numerous brown
seeds ca. 1.5 mm in diameter. 38 Physalis nicandroides Schltdl., Linnaea 19: 311 (1846)
Distribution and habitat: It grows in secondary vegetation of
deciduous tropical scrub, or near rivers, on rocky soils. It develops at Type: Mexico, Hidalgo, Atotonilco el Grande, Ehrenberg 760
an altitude range of 200–350 m. Herbarium specimens of this species (lectotype designated by Martínez (1998): HAL 0042230!). (Figures
are scarce. Mich., endemic. (Figure 10F). 1O, 2G, 2H, 3J, 9K).
Diagnostic characters: A small plant with long anthers of intense Erect annual herb up to 1 m tall, branched, glutinous, fetid, with
purple color, small maculations, and dense glandular pubescence. eglandular trichomes, some glandular; leaves alternate, petioles
Common names and uses: None known. 1.0–9.0 cm long, blades ovate, broadly ovate to cordate,
Phenology: Flowering and fruiting from October to February. 3.5–18.4 cm long, 2.5–16.0 cm wide, apex acute to slightly
Conservation status: VU. acuminate, base oblique, truncate to subcordate, margin entire to
Representative specimens examined. MEXICO. Michoacán: serrated; flowers solitary on peduncle 6–8 mm long; calyx lobes
Mugica, km 161 carretera Nueva Italia-La Huacana en el puente La acuminate 1.5–4.5 mm long; corolla campanulate, creamy whitish,
Pastora, M. Martínez 9457 (QMEX). 4–6 mm in diameter, with five simple greenish maculations; anthers
blue or blue-green, 2–3 mm long; fruiting peduncle 1.2–1.4 cm long,
37. Physalis minuta Griggs, Torreya 3: 138 (1903) thickened, fruiting calyx 5-angled, 3.5–4.3 cm long, 2.5–2.6 cm wide,
strongly cordate, caudate lobes subulate, up to 1.5 cm long, golden
Type: Mexico: Guerrero, Acapulco, E. Palmer 304 (lectotype brown at maturity and coriaceous after drying, mature berry dark
designated here: MO barcode 1968849!, isolectotypes: US barcode brown, up to 1.5 cm in diameter, containing numerous dark brown
00027344!, GH 00077290!). (Figure 9J). seeds ca. 2.5 mm in diameter.
= Physalis mimulus Waterf., Rhodora 69:211 (1967). Type: Distribution and habitat: A common weed along roads or near
Mexico, Colima, Socorro Island, C. H. T. Townsend s.n. crop fields. It develops at an altitude range of 120–2,000 m. Ags.,
(holotype: US barcode 00027342!) B.C.S., Chis., Col., Dgo., Edo. Mex., Gto., Gro., Hgo., Jal., Mich., N.L.,
Perennial herb, slightly erect to prostrate, spreading, branched, Mor., Nay., Oax., Pue., Qro., S.L.P., Sin., Son., Tamps., Ver., Yuc.,
up to 60 cm long, pubescent with eglandular, short trichomes; leaves Zac. (Figure 11B).
alternate, petioles 0–9–2.6 cm long; blades ovate, slightly succulent, Diagnostic characters: The fruiting calyx is strongly 5-
1.8–4.0 cm long, 0.9–2.1 cm wide, apex obtuse to acute, base angled, cordate, and leathery; corollas are cream-white with
oblique, margin entire; flowers solitary on peduncles 4–5 mm pale maculations, short peduncles, and subulate caudate calyx
long; calyx with deltate lobes 1.0–1.5 mm long; corolla lobes.

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Common names and uses: Costomate, miltomate, tomatillo, V0073011F!, OKLA not located, UC 104036, US barcode 00027366!,
matapulgas, tomate zope, tomate de culebra, tomate de perro, barcode 01014183! VT not located).
tomate loco, tomatillo loco, tomatón, vejiga de perro, yucu-quise. Erect or spreading perennial herb, the rhizome about 30 cm
Reported as medicinal in Morelos; the species is left in the fields as an long, 2.0 cm thick, branching from the base, pubescence highly
insect trap in Central Mexico. The fruits are occasionally eaten in variable in length and density, the trichomes eglandular, septate,
Morelos. white; alternate or geminate leaves, petiole 0.8–4.0 cm long; blades
Phenology: It flowers and bears fruit from June to April. ovate to lanceolate 1.7–10.0 cm long, 1.2–5.6 cm wide, attenuated
Conservation status: LC. toward the acute or obtuse apex, base truncate, oblique, shortly
Representative specimens examined. MEXICO. decurrent, margin entire or with two to three teeth per side; flowers
Aguascalientes: San Francisco de los Romo, 1.8 km al ESE de la solitary on peduncles 0.8–2.0 cm long; calyx with ovate-to-attenuate
colonia Macario J. Gómez, 823b (MEXU). Baja California Sur: a lobes 4–7 mm long, corolla yellow, 1.5–2.6 cm in diameter, rotate
10 km de la desviación a San Antonio, cerca de Las Termópilas, M. campanulate, with simple-to-compound, reddish-brown to purple
Martinez 9519 (QMEX). Chiapas: municipio de Berriozábal, flats maculations; anthers 2–4 mm long, purple or blue; fruiting peduncle
near Berriozábal, D. Breedlove 52,384 (CAS, MEXU). Colima: 0.8–2.1 mm long, fruiting calyx 10-costate with five more
Comala, rancho El Jabalí, 20 km (airline), NNW of Colima in conspicuous angles or nearly round, 2.3–3.7 cm long, 1.5–3.0 cm
the SW foothills of the Volcan de Colima, ridgetop in hills near a wide; mature berry 1.0–1.2 cm in diameter with numerous yellow
shrine 4.9 km SE of Hacienda San Antonio on the road to Comala, seeds 2–2.5 mm in diameter.
near 19° 26.2′ N, 103° 41.8′ W, J. Sanders 10,277 (MEXU, MICH). Distribution and habitat: On slopes and margins of disturbed
Durango: Canelas, proximidades a Canelas, terrenos de la UAF oak forests and adjacent clearings of pine–oak forests, secondary
Topia, Montemojino 2370 (CIIDIR-DGO). Estado de México: grasslands, and subtropical scrub. From 1,850 to 2,700 m. Ags., Cd.
15 km aL SW de Tejupilco, E. Guizar 205 (MEXU). Mex., Chih., Dgo., Edo. Mex., Gto., Gro., Hgo., Jal., Mich., Mor.,
Guanajuato: Mesas del Tigre, municipio de Victoria, E. Ventura Nay., N.L., Oax., Pue., Qro., S.L.P., Ver., Zac. (Figure 11C).
8560 (IEB, QMEX). Guerrero: municipio de Tlapa, a 8 km al N de Diagnostic characters: The plant has entire leaves, large corollas
Tlapa, E. Martínez 2665 (MEXU, TEX). Hidalgo: Ajacuba, “La with strongly contrasting maculations and purple or blue anthers, a
mesa chata,” cerro al N W del poblado Santiago Tezontlale, sierra large fruiting calyx, and peduncles. The species can be confused with
del “Mexe,” ejido Santiago Tezontlale, I. Díaz 1206 (MEXU). P. chenopodiifolia; however, the latter is easily distinguished by its
Jalisco: Acatlán De Juárez, Cuesta de san Marcos, above and to cinereous appearance and different leaf form.
the east of Laguna de san Marcos, ca. 15 km. south-southeast of Common names and uses: Vonsummate, tomate cimarrón,
Acatlán de Juárez, R. McVaugh 342 (ENCB, MICH). Michoacán: tomate de mata, tomatillo, xapindikua. The plant is used
Ario de Rosales, en san José de Las Cañas, 20 km aL SW de Ario de medicinally, and the fruit is eaten.
Rosales, carretera a la Huacana, Soto Núñez 3464 (ENCB, MEXU). Phenology: It flowers and bears fruit from June to September.
Morelos: 3 km aL SE de La Joya, entre Cuernavaca y Yautepec, L. Conservation status: LC.
Hernández 2488 (TEX). Nayarit: Ahuacatlán, a 4 km al S de Representative specimens examined. MEXICO.
Ahuacatlán, por el camino a Amatlán de Cañas, 21° 01′ N, 104° Aguascalientes: Calvillo: Sierra del Laurel, near the
30′ W, O. Téllez 9947 (MEXU). Nuevo León: 3 km aL NE del Jalisco–Aguascalientes border, ca. 10 miles southeast of Calvillo
poblado El Potosí, hacia San José de la Joya, M. Martínez 9785 (3 h by horse from rancho de Los Adobes), McVaugh 18,439
(QMEX). Oaxaca: Valle del Etla, L. Smith 720 (MEXU). Puebla: (ENCB). Ciudad de México: Tlalpan, Área Natural Protegida:
municipio de Huehuetlán el Chico, ejido Tzicatlán, Arreola TZ 570 Volcán Pelado, J. Rivera 3783 (MEXU). Chihuahua: Guachochi,
(TEX). Querétaro: El Chacal, 1 km aL poniente de la Mesa de alrededores de Guachochi, R. Hernández 8625 (MEXU). Durango:
Fortín, municipio de Landa de Matamoros, H. Rubio 156 (IEB, Pueblo Nuevo, 35 km al W de El Salto, Hernández 7768 (MEXU).
QMEX). San Luis Potosí: 64 mi NE of San Luis Potosí, U. Waterfall Estado de México: National Park Lagunas de Zempoala, R.
15,712 (F, NY). Sinaloa: municipio de Culiacán, Santa María, McAdams 52 (MEXU). Guanajuato: más o menos 6 km aL SW
González 6694 (CAS, F, US). Sonora: Yécora, Rancho Los de San franco, municipio de San Diego de la Unión, E. Carranza
Jacales, ca. 6.2 km (by air) northwest of Santa Rosa, T. Van 5060 IEB, QMEX). Guerrero: 15 km W-SW de Chichihualco, M.
Devender 2008–545 (MEXU). Tamaulipas: Miquihuana, M. Mayfield 998 (MEXU). Hidalgo: roadside near Real de Monte above
Martínez 9462 (QMEX). Veracruz: municipio de Acultzingo, Pachuca, A. Sharp 4457 (MEXU). Jalisco: Acatic, 6 km al suroeste de
5 km ENE of center of Acultzingo, M. Nee 33,132 (NY). Acatic, Reynoso Dueñas 84 (IBUG). Michoacán: Cerro La Alberca,
Yucatán: Izamal, Gaumer 1446 (F, US). Zacatecas: cerro La cerca de Villa Jiménez, municipio de Villa Jiménez, J. Rzedowski
Cantarilla, 8 km al S de Moyohaua, E. Enriquez 1412 (MEXU). 40208, 40192 (IEB). Morelos: km 47 carretera de cuota México-
Cuernavaca, límites entre DF y Morelos, A. Rodríguez 2782
39. Physalis orizabae Dunal, Prodr. [A. P. de Candolle] 13 (1): 452 (MEXU). Nayarit: Acaponeta, La Ciénega on ridge
(1852) approximately 10 miles northwest of mesa del Nayar, Norris
14,575 (MICH). Nuevo León: General Zaragoza, Sierra El
Type: Mexico, Veracruz, Mount Orizaba, C. J. W. Schiede 607 Soldado, camino a Puerto Pinos J. Villarreal 4989 (MEXU).
(holotype: HAL 0042231!). (Figure 9L). Oaxaca: Santiago Textitlán, distrito: Sola de Vega, Paraje Tierra
= Physalis subintegra Fernald, Proc. Amer. Acad. Arts 35: 567 de Yunta, A. Sánchez 1692 (MEXU). Puebla: 5 km al E de
(1900). Type: Mexico, Edo. de Mexico, Sierra de Las Cruces, C. G. Xalitzintla, P. Tenorio 15,931 (MEXU). Querétaro: 4–5 km al S
Pringle 8225 (holotype: GH 00077373!, isotypes: BR not located, F de El Parador Santa Martha, municipio de Landa de Matamoros, E.

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Carranza 1799 (IEB). San Luis Potosí: 21 mi al E de San Luis Potosí, wide, 1.8–2.2 cm long; mature berry brown, 1.0–1.2 cm in
L. Dorr 1984 (MEXU). Veracruz: El Limón, Ramos 233 (MEXU). diameter with numerous dark brown seeds ca. 2 mm in diameter.
Zacatecas: Valparaiso, road to Huejuquilla el Alto, Jal., one mile Distribution and habitat: Dry or semi-arid areas, in thorny or
west of the road junction 18 miles south of Valparaiso on the road subtropical thickets, grasslands or hillsides near pine or oak forests,
to Mezquitic, Jal, 22° 38′ N, 103° 48′ W, R. McVaugh 17763 or as weeds in plots within the forest. It grows at an altitude range
(MICH). of 900–2,700 m. Ags., Chih., Col., Cd. Mex., Dgo., Edo. Mex., Gto.,
Hgo., Jal., Mich., Oax., Pue., Qro., Sin., Tlax., Ver., Zac.
40. Physalis parvianthera Waterf., Rhodora 69: 204 (1967) (Figure 11D).
Diagnostic characters: A fetid annual plant with dense and
Type: Morelos: Sierra de Chalchi, cerca de Tepoztlán, Paray glutinous pubescence, toothed leaves, 5-angled fruiting calyx, and
1614 (holotype: ENCB!). wide corollas with clear or diffuse maculations.
Erect herb, 25–40 cm tall; stems villous, with articulated more or Common names and uses: Tomate, tomate de burro, tomatillo,
less viscid trichomes; petioles 10–14 mm long; leaves deltoid-ovate, hierba del zopilote, jaltomate hediondo. No uses known.
ovate or lanceolate, more-or-less villous, 25–30 mm long, 20–30 mm Phenology: It flowers and bears fruit from July to November and
wide; apex obtuse to acute, base cuneate to truncate, decurrent, bears fruit until March.
margin entire; flowers solitary on peduncles 3–5 mm long; calyx Conservation status: LC.
lobes lanceolate 1–1.5 mm long; corolla campanulate, yellow, Representative specimens examined. MEXICO.
immaculate, 15–20 mm in diameter; anthers violet 1.5–1.8 mm Aguascalientes: Aguascalientes, highway to Ojuelos, Jal., nine
long; fruiting peduncles 6–8 mm long, fruiting calyx 10-costate, miles east of Aguascalientes, R. McVaugh 16,672 (MICH).
8–9 mm long, 5–7 mm wide. Chihuahua: en el camino a Cumbres de Majalca, municipio de
Distribution and habitat: Probably in pine–oak forest, such as Chihuahua, M. Martínez 9655 (QMEX). Colima: Comala, rancho El
the forest in Tepoztlán, at 1,700 m. Mor. (Figure 10B). Jabalí, 20 km (airline), N of Colima in the SW foothills of the Volcan
Diagnostic characters: The small, nearly 10-costate de Colima, El Agostadero, ridgetop in hills above the shrine 4.9 km
fruiting-calyx, immaculate yellow corollas, small violet SE of Hacienda San Antonio on the road to Comala, near 19° 26.2′ N,
anthers, and soft vestiture. The deltoid-ovate leaves are 103 \° 41.8′ W, J. Sanders 11,806 (MEXU, MICH). Ciudad de
uncommon in Physalis. México: Tláhuac, vertiente E del cerro de Santa Catarina,
Common names and uses: None known. J. Rzedowski 26,037 (CAS, MEXU, NY). Durango: Nombre de
Phenology: Flowering in August. Dios, al SW de La Parrilla, S. González 165 (CIIDIR-DGO).
Conservation status: DD. Estado de México: Huixquilucan, 2 km al W de Río Hondo,
Representative specimens examined. MEXICO. Morelos: J. Rzedowski 29,569 (MEXU). Guanajuato: 5 km al S de San
Tepoztlán, Sierra de Chalchi, cerca de Tepoztlán, Paray 1614 Felipe, sobre la carretera a Dolores Hidalgo, municipio de San
(ENCB). Felipe, J. Rzedowski 47,319 (IEB, QMEX). Hidalgo: Ajacuba
Jagüey “El Palo Seco,” planicie de Tulancalco, aprox. 4 km sobre
41. Physalis patula Mill., Gard. Dict., ed. 8. n. 12 (1768) el camino de terracería que se inicia al S de Emiliano Zapata, ejido
Tecomatlán, I. Díaz 1200 (MEXU). Jalisco: Ojuelos, potrero La
Type: Mexico, Veracruz, 1731, Houston s.n. (holotype: BM Colorada, 5 km al W de la carretera Ojuelos-Lagos de Moreno, M.
000775431!). (Figures 1P, 12A). Alcocer 62 (MEXU). Michoacán: Zamora, halfway between Jacona
= Physalis foetens Poir. Encycl. (J. Lamarck and al.) Suppl. 2. 348 and Zamora, along Mexico highway 15, 19° 55′ N, 102° 18′ W, Iltis
(1811). Type: Mexico, A. Bonpland s.n.; 1803–1804 but reportedly 648 (ENCB, WIS). Oaxaca: La Loma Pachona, 6 km al NE de Gpe.
from “Peru” without further locality (FI not located). Cuautepec o 1 km al E del entronque carr. Huajuapan de León-
= Physalis foetens var. neomexicana (Rydb.) Waterf., Rhodora 60 Tehuacán y la brecha de Cuautepec, A. Salinas 3680 (MEXU).
(714): 168 (1958). Type: United States, New Mexico, A. Fendler 678 Puebla: Xochitecatl 2300 cerca de Texmelucan, W. Boege 154
(lectotype, designated by Waterfall 1958 GH 00003288!). (MEXU). Querétaro: Santa Rosa, casa de Juan Antonio Isla,
= Physalis subulata Rydb., Bull. Torrey Bot. Club 22 (7): 306 municipio de Querétaro, E. Argüelles 1410 (ENCB, MEXU).
(1895). Type: Mexico, Chihuahua, waste grounds Guerrero, C. G. Sinaloa: Culiacán a 20 km al N de Culiacán, carretera Culiacán a
Pringle 1344 (holotype: GH 00003294!, isotypes: F barcode Guamúchil, ladera NE del cerro La Chiva, a 1 km aL O de Mirasoles,
V00073013F!, MEXU not located, NY barcode 138890!, OKL not R. Vega 6263 (UAS). Tlaxcala: Tzompantepec, San Juan
located). Quetzalcuapan, H. Vibrans 184 (MEXU). Veracruz: Puerto del
Erect annual herb up to 50 cm tall; stem branched, glutinous, Aire, carretera Orizaba-Tehuacán, Rosas 808 (MEXU). Zacatecas:
fetid, densely pubescent, with eglandular and glandular trichomes; Jalpa, pastured hills five miles southwest of Jalpa, R. McVaugh
leaves alternate, petioles 1.0–3.5 cm long; blades ovate, 3.5–7.0 cm 18508A (ENCB, MICH).
long, 2.0–4.6 cm wide, apex acute, base oblique to cuneate decurrent,
margin sinuate or serrated; flowers solitary on peduncles 3–7 mm 42. Physalis pennellii Waterf., Rhodora 69: 116 (1967)
long; calyx with subulate lobes 1.5–3.5 mm long, corolla
campanulate, pale yellow, 1.0–2.5 cm in diameter, with five Type: Mexico San Luis Potosí rocky limestone west of Santa
simple yellowish-brown to olive-green diffuse maculations; Ana, above Potrero, Sierra de Catorce F. W. Pennell 17523
anthers yellow or blue when dry, 1.5–4.0 mm long; fruiting (holotype: US barcode 00027351!, isotypes: NY barcode 138880!,
peduncle 0.4–1.2 cm long, fruiting calyx 5-angled, 1.5–3.0 cm PH not located).

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Perennial herb, 5–20 cm tall; articulated trichomes, long and Erect annual herb, 3–100 cm high, stem with adpressed trichomes
short to 1.5 mm long, partly glandular; petioles 5–11 mm long, 0.5 mm long; leaves alternate, petioles 1–2.5 cm long, ovate to lanceolate
blades ovate or broad-ovate 13–32 mm long, 12–25 mm wide, 2–9 cm long, 1–5 cm wide, apex acute, base attenuate, sometimes oblique
apex obtuse to acute, base cordate, margin undulate or entire; up to 1.5 cm, margin dentate with 8–10 teeth per side, and almost
solitary flowers on peduncles 3–6 mm long; calyx with deltoid or glabrous; flowers solitary on peduncles up to 5 mm long; calyx with
ovate-deltoid lobes densely pubescent; corolla campanulate, yellow, triangular lobes ca. 2 mm long, marginally pubescent; corolla rotate,
7–13 mm long, 10–14 mm wide, maculations dark brown; anthers yellow, 0.8–2.5 cm in diameter with five purple diffuse maculations,
violet, ca. 2.5 mm long; fruiting peduncles 9–15 mm long; fruiting corolla throat densely pubescent; anthers blue, 2–5 mm long, convolute
calyx 10-costate, 15–20 m long, 11–13 mm wide; mature berry after anthesis; fruiting peduncles 7–10 mm long, fruiting calyx 10-costate,
8–12 mm wide with numerous yellow seeds ca. 1.5 mm in diameter. 1–3 cm long, 1.8–2.0 cm wide; mature berry spherical, 1–5 cm in
Distribution and habitat: rocky limestones, S.L.P. (Figure 8C). diameter, green, yellow, or purple-tinged at maturity, with numerous
Diagnostic characters: Physalis pennellii is most similar to P. yellow seeds ca. 2 mm in diameter. In large-fruited specimens, the berry
sordida Fern. but is a smaller plant, with a mixture of long trichomes tears the fruiting calyx.
on the leaves and a more densely long-hairy flowering calyx. Distribution and habitat: A common weed of cities, roads, and
Common names and uses: None known. agricultural fields; it grows up to 2,500 m asl. Present in all the
Phenology: Flowering in July. Mexican states except Camp., Tlax., and Yuc. (Figure 11E).
Conservation status: NT. Diagnostic characters: Due to its weedy tendency and economic
Representative specimens examined. MEXICO. San Luis importance, Physalis philadelphica is the most widely distributed
Potosí: 3 km pasando Alamillos, al S de Real de Catorce, M. species in the country. It is also cultivated in some regions of Mexico
Martínez 9560 (QMEX). for its edible acidic fruit. The species is characterized by the large
globose calyx in the fruit, corollas with simple maculations,
43. Physalis philadelphica Lam., Encycl. Méth. Bot. 2: 101, 1786. convolute anthers, and the margin of the leaves with short teeth.
Lam. herb. P, IDC 6207. 471: I.5) Common names and uses: Tomatillo, tomate, tomate verde,
tomate de hoja, tomatillo de cáscara, tomate cuarentano, tomate de
Type: Cultivé en 1784 au Jardin du Roi; nous le croyons bolsa, yashal tumat, quelite, miltomate, tomate fresadilla, fresada.
originaire de l’Amérique septentrionale (lectotype designated by The fruit is commonly used in the preparation of sauces, either raw,
Waterfall Rhodora 69 (778): 213. 1967 P barcode P00357705!). fried, or boiled.
(Figure 12B). Phenology: Flowers and fruits mainly after the rainy season;
= Physalis chenopodifolia Willd., sp. Pl., ed. 4 [Willdenow] however, specimens with flowers were examined in March.
1https://www.ipni.org/n/77093275-12): 1023 (1798). Conservation status: LC.
= Physalis philadelphica var. immaculata Waterf., Rhodora 69: Representative specimens examined. MEXICO.
215 (1967). Type: Mexico, Chihuahua, silty flat in desert 14 miles NE Aguascalientes: San José de Gracia, 200 m adelante de El
of Parral, U.T. Waterfall 16,115 (holotype: OKLA not located, Milagro, en la desviación a Santiago, O. Vargas 779 (CUCBA,
isotypes: GH not located, United States not located). MEXU). Baja California: San Quintín, productora de plantas
= Physalis philadelphica subsp. ixocarpa (Brot. ex Hornem.) “Baja Plants,” M. Martínez s/n (QMEX). Baja California Sur:
Sobr.-Vesp. and Sanz-Elorza, Acta Bot. Malac. 32: 233 (2007). Todos Santos antes del pueblo cerca del río, M. Martínez 9523
= Physalis philadelphica var. minor Dunal, Prodr. [A. P. de (QMEX). Chiapas: arroyo de San Nicolás, está al Este de San
Candolle] 13 (1): 450 (1852). Type: Mexico, Tamaulipas, Tampico, Cristóbal de las Casas, A. Méndez 8473 (MEXU). Chihuahua:
J. L. Berlandier 20 (lectotype designated by Pretz and Deanna 2020 G Cuauhtemoc, Napavechia Valley, K. Adams 30 (ARIZ). Ciudad
barcode G00343158, isolectotype: BM barcode BM000775484). de México: Tláhuac, Mixquic, V. Popper 105 (MEXU). Coahuila:
= Physalis philadelphica var. parviflora Waterf., Rhodora 69: 215 pasando Ramos Arizpe, ca. 3 km al E hacia Monterrey, M. Martínez
(1967). Type: Mexico, Nayarit, Cerro Sanganguey 12 miles SE of 9793 (QMEX). Colima: Manzanillo, cerro de La Cruz, Campos 7
Tepic, Feddema 582 (holotype: MICH barcode 1109899!, isotype: (IBUG). Durango: Santiago Papasquiaro, Escuela Jose Ramon
OKLA not located). Valdez, R. Worthington 11,315 (UTEP). Estado de México:
= Physalis philadelphica f. pilosa Waterf., Rhodora 69 (no. 778): 214 Naucalpan de Juárez, Barranca cercana a El Huizachal,
(1967). Type: Guatemala, Finca La Alameda near Chimaltenango, P. C. J. Rzedowski 25109 (MEXU). Guanajuato: 2 km al S de San José
Stanley 79899 (holotype: F barcode V0077208F!). de Tránsito, municipio de Silao, cerca del aeropuerto, J. Rzedowski
= Physalis laevigata M. Martens and Galeotti in Bull. Acad. Brux. 49,832 (IEB). Guerrero: La Unión de Isidoro Montes de Oca,
xii. I. 131 (1845). Type: Mexico, presa de Morelia de Michoacán, H. Vallecitos, G. Hinton 11,449 (MEXU). Hidalgo: Mineral de El
G. Galeotti 1188 (lectotype designated by Pretz and Deanna 2020 BR Chico, calles aledañas al poblado, A. Rodríguez 2773 (CUCBA,
barcode 000005529483, isolectotypes: BR barcode 000005528332, P MEXU). Jalisco: Acatic, brecha crucero viejo de Acatic, Reynoso
barcode P00409133). 66 (IBUG). Michoacán: Crucero a Curimeo, carretera Zacapu-
= Physalis cavaleriei H.Lév., Repert. Spec. Nov. Regni Veg. 11: Purándiro, municipio Panindícuaro, E. Pérez 1839 (IEB).
295 (1912). Type: China, Guizhou, Kouy-Tehéou, Kouy-Yang sous Morelos: Curva La Pera, carretera de cuota México-Cuernavaca,
les rochers, J. Cavalerie 3800 (lectotype designated by Pretz and A. Rodríguez 2786 (CUCBA, MEXU). Nayarit: Santa Teresa, camino
Deanna (2020) E barcode E00284483, isolectotypes: E barcode a La Mesa del Nayar, a 1 km aL SW, 22° 30′ N, 104° 48′ W, Flores
00284484, G barcode G00343185, K barcode K000759437, P 1381 (MEXU). Nuevo León: Iturbide, Loma La Bandera, NW of
barcode P000061015). Ejido Santa Rosa, 4.1 mi S of Iturbide, J. Sullivan 2578 (TEX).

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Martínez et al. 10.3389/fgene.2023.1080176

FIGURE 13
Geographic distribution of Physalis spp. (A) P. pubescens, (B) P. solanacea, (C) P. sordida, (D) P. sulphurea, (E) P. queretaroensis and P. waterfallii.

Oaxaca: poblado de Santa María Ixcatlán (UTM 1974844, 14Q Davidse 20,028 (MEXU). San Luis Potosí: Rioverde, between Rio
691,794) E. Rivera 36 (MEXU). Puebla: San Andrés Cholula, a Verde and Plazuelo (km 89.9) MEX 69 km 93 Robert Bye 18944
1.2 km aL sur de San Antonio Cacalotepec, Y. Guarneros 348 (MEXU). Sinaloa: Concordia, Santuario El Palmito (La Chara
(MEXU). Querétaro: Municipio de Landa de Matamoros, Misión Pinta), La Liebre, J. Rubio 107 (CIIDIR). Sonora: Sirebampo
de Tilaco, a 2.5 km de Landa de Matamoros, M. Martínez 2788 vicinity. 9.5 km south on Mexico 15 from Las Bocas Road
(QMEX). Quintana Roo: 1 km NW of Puerto Morelos, Campo turnoff, 3.5 km west on Sirebampo Rd., 11.5 km south (by air)
Agropecuario del Centro de Investigaciones de Quintana Roo, G. San Jose de Masiaca, S. Friedman 094–95 (ASU). Tabasco:

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Martínez et al. 10.3389/fgene.2023.1080176

Huimanguillo, hacia la Laguna de los Limones y 5 km de la three irregular teeth or small lobes, or entire, sparsely appressed-
desviación a Fco. Rueda, M. Magaña 738 (MEXU). Tamaulipas: hairy, often with more abundant trichomes along the veins; flowers
Miquihuana, en el vado que atraviesa la carretera, M. Martínez solitary on peduncles 0.9–1.5 cm long; calyx lobes ovate to lanceolate
9469 (QMEX). Veracruz: Chacaltianguis, Benito Juárez, 7–10 mm long; corolla rotate campanulate, yellow, 18–22 mm wide
M. Nee 29,270 (XAL). Zacatecas: Jalpa camino a la with five dark brown maculations, corolla throat pubescent; anthers
colonia Morelos, 3 km a la salida a Apozol E. Enríquez 634 bluish, 2.2–3.0 mm long; fruiting peduncles 1.5–2.0 cm long, fruiting
(MEXU). calyx 5-angled 2.5–4.0 cm long, 1.0–1.5 cm wide; mature berry
1.0–1.4 cm in diameter, dark yellow seeds 1.8–2 mm.
44. Physalis porrecta Waterf., Rhodora 69: 237 (1967) Distribution and habitat: In moist Abies and pine–oak forests.
Alt. 2,750–3,100 m. Cd. Mex., Dgo., Oax. (Figure 10F).
Type: Costa Rica, Prov. San Jose, vicinity of El General, A. F. Diagnostic characters: A rotate-pentangular corolla with dark
Skutch 2931 (holotype: US barcode 00027357!, isotypes: GH barcode maculations, fruiting calyx 5-angulated, dense pubescence with
00077361!, NY barcode 138883!) (Figures 3L, 12C). some glandular trichomes.
Erect perennial herb, 0.4–1.3 m tall; stem branched with Common names and uses: None known.
puberulent trichomes; petioles (2–) 3–7 cm long; blades ovate, Phenology: Flowering and fruiting in September and October.
5–11 cm long, 4–7 cm wide, apex acute, base decurrent, margin Conservation status: LC.
unevenly dentate or entire; flowers solitary on peduncles 4–5 mm Representative specimens examined. MEXICO. Cd. Mex:
long; calyx with narrowly tapered lobes 1.5–4.0 mm long, corolla Sierra del Ajusco C. G. Pringle 6216 (MEXU). Durango: San
rotate campanulate, yellow, 12–15 mm in diameter, with diffuse not Ramón, E. Palmer 114 (IBUG). Oaxaca: Sierra de Clavellinas C.
strongly contrasting maculations; anthers yellow or violet, G. Pringle 6001 (IBUG).
1.8–2.3 mm long; fruiting peduncles 6–15 mm long, fruiting calyx
5-angled, 3–5 cm long, 2.2–3.0 cm wide, glabrous; mature berry 46. Physalis pruinosa L., sp. Pl. 1: 184 (1753)
12–15 mm in diameter with numerous yellow seeds ca. 1 mm in
diameter. Type: Habitat in America (holotype: LINN 247.13!). (Figures
Distribution and habitat: Mountain cloud forest at 1Q, 12E).
1,200–1,500 m asl. Chis., Gro., Oax. (Figure 10C). = P. maxima Mill., Gard. Dict., ed. 8. n. 15 (1768). Type: Mexico,
Diagnostic characters: According to Waterfall (1967), the Veracruz without further locality, Houston s.n. (holotype: BM
species is characterized by its usually abruptly beaked fruiting 000775441!, Sloane collection, photoholotype BH!, OKLA!).
calyx, yellowish corollas varying from immaculate to yellowish = Physalis cordifolia Dunal, Prodr. [A. P. de Candolle] 13 (1):
maculations, and diffuse margin. The stem, branches, and 441 (1852). Type: from a cultivated plant in the Lugduno-Baravo
petioles present a variable density of pubescence. Gardens, De la Roche s.n. (holotype: G-DC SIB 160733/1!)
Common names and uses: Moench’o, miltomate, tomatillo. No Erect annual herb 1.0–1.5 m tall; stem branched, glutinous, fetid,
uses known. pubescent with a mixture of eglandular short and long trichomes,
Phenology: Flowering and fruiting from July to February. some glandular; petiole 0.8–4.7 cm long; blades broadly ovate to
Conservation status: DD. cordate, 3.6–13.0 cm long, 2.1–6.0 cm wide, apex acuminate, base
Representative specimens examined. MEXICO. Chiapas: oblique, truncate to subcordate, margin usually serrate; solitary
Unión Juárez, En el camino Talquian-Cima del Volcan Tacana, flowers on peduncles 1.2–4.3 cm long; calyx with subulate caudate
mitad del camino hacia La línea, frontera con Guatemala Pilar lobes up to 8 mm long; corolla campanulate, spreading to reflected,
Zamora-Tavares et al. 273 (IBUG). Guerrero: Chilpancingo de los yellow, 0.8–1.3 cm in diameter, maculations diffuse, light pinkish-
Bravo, Omiltemi, La Frutilla, camino a 3 cruces A. Méndez 302 brown; anthers yellow, 2.5–3.0 mm long; fruiting peduncle 2.8–5.5 cm
(MEXU). Oaxaca: vicinity of Cafetal Concordia, 1 April 1933, long, fruiting calyx 5-angled, 2.3–6.0 cm long, 1.1–4.0 cm wide,
Morton and Makrinius 2556 (US). initially green, becoming golden brown; mature berry brown,
1.5 cm in diameter with numerous brown seeds ca. 2 mm in diameter.
45. Physalis pringlei Greenm., Proc. Amer. Acad. Arts 35: 311 Distribution and habitat: A ruderal weed or in ravines, margins
(1900) of oak or tropical deciduous forest or their ecotones at 120–1,850 m.
Chis., Chih., Edo. Mex., Col., Gro., Jal., Mor., Nay., Oax., Qro., Sin.,
Type: Mexico, Oaxaca, Sierra de Clavellinas, C.G. Pringle 6001 Son., Tamps., Ver., Yuc., Zac. (Figure 11F).
(lectotype selected by Waterfall (1967) GH 00077362!, isolectotypes: Diagnostic characters: A long flower peduncle, pinkish-
MEXU barcodes 27677!, 28217!, 27675!, BM not located, BR not brown maculations, yellow anthers. The species can be
located, NY 138884!, UC 104028, US barcode 00027358!, VT not confused with P. nicandroides because both are pubescent,
located). (Figure 12D). glutinous, and very similar in plant habit and appearance;
= Physalis pringlei var. curtiloba Waterf., Rhodora 69 (778): 228 both have a strongly 5-angled fruiting calyx. However, P.
(1967). Type: Mexico, Durango, San Ramón, E. Palmer 114 pruinosa has a mixture of short and long trichomes in the
(holotype: US barcode 00027359!). stem and a thin fruiting peduncle.
Erect perennial herb 1.0–1.7 m tall; stem densely covered with Common names and uses: Miltomate, vejiga de perro, farolito,
glutinous jointed spreading trichomes of varying lengths; leaves chipin sox, tomate de culebra, tomate de hoja, tomate de bolsa,
geminate throughout, petioles 1–3 cm long; blades ovate, 4–8 cm tomate, tomatillo, tomate grande, papay-zeal, tumat’cho, tumal,
long, apex acute to acuminate, base attenuate, margin with one to tumat. The fruit is eaten.

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Sahuaripa M. Martínez 9838 (QMEX). Tamaulipas: 19 km al E de la

carr Zaragoza-Gonzalez, arroyo de Minas hacia ejido El Cabrito, C.
González 133 (MEXU, UAT). Veracruz: Totutla, Encinal, F. Ventura
7046 (ENCB, MEXU) Yucatán: sin más localidad, G. Gaumer 24,310
(F). Zacatecas: Jalpa, pastured hills five miles southwest of Jalpa,
McVaugh 18508A (ENCB, MICH).

47. Physalis pubescens L., Sp. Pl. [Linnaeus] 1: 183 (1753)

Type: “In India utraque,” without further locality, date, or

collector (lectotype selected by Waterfall 1958: LINN 247.11!).
(Figures 3K, 12F).
Synonyms in Martínez (1998) and Pretz and Deanna (2020),
except for P. latiphysa Waterf., which we consider a distinct species.
= Physalis angustiloba Waterf., Rhodora 69: 320 (1967). Type:
Mexico, Jalisco [Talpa de. Allende], hills two miles north of La Cuesta,
road to Talpa de Allende, R. McVaugh 21141 (holotype: MICH
1109911!, isotypes: OKLA not located, LL!, US barcode 00027312!)
Erect annual to biennial herb 10–150 cm tall; pubescent, and
pilose to glandular-pilose with multicellular trichomes up to 3 mm
long; petioles 1.5–7.0 cm long, blades ovate to deltoid, 2–12 cm long,
1–7 cm wide, apex acute to acuminate, base obtuse to truncate,
sometimes oblique up to 3 mm, margin irregularly dentate to almost
entire, villous especially on the lower surface; flowers solitary on
peduncles 2–7 mm long; calyx villose with triangular lobes
1.5–3.0 mm long, corolla rotate, yellow, 0.9–2 cm in diameter,
with five strongly contrasting obtuse maculations, corolla throat
densely pubescent; anthers purple or blue, seldom yellow, ca. 2 mm
long; fruiting peduncle 5–10 mm long, fruiting calyx strongly 5-
angled, 1.2–3.0 cm long, 1.8–2.5 cm wide, always longer than wide,
and villous throughout with multicellular trichomes; mature berry
FIGURE 14 green tinged with purple or yellow, ca. 1 cm in diameter with many
Details of flowers and fruits of Physalis spp. (A) P. solanacea, (B) P. brown seeds ca. 1.5 mm in diameter.
spathulifolia, (C) P. sordida, (D) P. subrepens, (E) P. tamayoi, (F) P.
Distribution and habitat: The plant grows on disturbed
sulphurea, (G) P. volubilis, (H) P. vestita, (I) P. waterfallii.
areas of streams, grasslands, and pine–oak forest, also
associated with crop fields, from sea level up to 2,600 m asl.
B.C.S., Camp., Chis., Chih., Coah., Col., Dgo., Gro., Hgo., Jal.,
Phenology: Flowering and fruiting from June to November. Mich., Nay., NL., Oax., Pue., Qro., Q.R., Sin., Son., Tab.,
Conservation status: LC. Tamps., Ver., Yuc. (Figure 13A).
Representative specimens examined. MEXICO. Chiapas: Diagnostic characters: Physalis pubescens can be recognized by
Comitán de Domínguez, 1 km al SE del enronque Tzimol- the pubescent stem, blue anthers, simple corolla maculations, and
Uninajab, camino a Uninajab, A. Reyes 1098 (MEXU). Chihuahua: five-angled fruit calyx.
La Cienegita, Rio Mayo, H. Gentry 2632 (F, MEXU). Colima: Comala, Common names and uses: Cocostomat, tomatillo, miltomate,
cerro Caleras, 4 km antes de Tecolapa, autopista Colima-Guadalajara, tomate de culebra, chichol antivo, tumat’cho. The fruit of weedy
O. Vargas 814 (IBUG). Estado de México: Luvianos, Temascaltepec, plants is used in the preparation of sauces or eaten raw as fresh fruit
G. Hinton 4592 (MEXU). Guerrero: Coyuca de Benítez, Santa in several areas (Martínez, 1998).
Bárbara, G. Hinton 8503 (MEXU). Jalisco: La Huerta: arroyo Phenology: It flowers and bears fruit mainly from June to
Colorado, cerca de Los Pozos, E. Lott 528 (MEXU). Michoacán: October, but there are some specimens collected in February that
Apatzingán, barranca of rio Cancita at the bridge nine miles southeast bear fruit.
of Apatzingán, R. McVaugh 17,981 (ENCB, MICH). Morelos: 2.5 km Conservation status: LC.
al N y 4 km al W de Huautla, R. Ayala 198 (MEXU). Nayarit: Bahía de Representative specimens examined. MEXICO. Baja
Banderas, seaward-facing mountainsides 1–1.5 miles above La California Sur: Isla Cerralvo, en el arroyo Las Delicias, M.
Cucaracha, 12–13 miles south of Las Varas, R. McVaugh 19,226 Martínez 9500 (QMEX). Campeche: Carmen, 2 km antes de llegar
(ENCB, MICH). Oaxaca: 5 km al NW de Totolapan, M. Martínez al límite del estado de Campeche con Tabasco, C. Chan 6137 (MEXU).
1904 (MEXU). Querétaro: hacia Casa de Máquinas, Cadereyta, A. Chiapas: Angel Albino Corzo steep, heavily wooded slope near the
Herrera 200 (QMEX). Sinaloa: Escuinapa, 18 km al E de Escuinapa. Rancho Viejo of the Finca Prusia, A. Méndez 3607 (MEXU).
Brecha Escuinapa-Corral de Piedra, P. Tenorio 2913 (MEXU). Chihuahua: Guachochi, Cabórachi, 20 km al E. de Guachochi, R.
Sonora: Rancho Cuadritos, ca. 7 km al E de Mesa del Seri hacia Hernández 8562 (MEXU). Coahuila: Tanque Nuevo, M. Martínez

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Martínez et al. 10.3389/fgene.2023.1080176

9606 (QMEX). Colima: Comala, rancho el Jabalí, 22 km (airline), N of Common names and uses: None known.
Colima in the SW foothills of the Volcan de Colima, Colima/Jalisco Phenology: Flowering and fruiting from October to March.
line passes through the ranch, near 103° 1.8′ W, 19° 26′ N, El Conservation status: DD.
Agostadero, ca. 5 km S of San Antonio, J. Sanders 11,330 (MEXU, Representative specimens examined. MEXICO. Sonora: Cerro
MICH). Durango: Mezquital, 1 km aL NE de santa María de Ocotán, Algodones, 5 km al W de San Carlos, M. Martínez 9844 (QMEX).
S. González 1455 (CIIDIR-DGO). Guerrero: Temixco, second
barranca E of Temixco, Y Mexia 8760 (CAS, F, NY, US). Hidalgo: 49. Physalis queretaroensis M. Martínez and L. Hern., Acta Bot.
Eloxochitlán, 6.5 km al E, O. Alcántara 2172 (MEXU). Jalisco: Autlán Mex. 46: 73 (1999)
de Navarro, south- and west-facing slopes, precipitous rocky
mountain sides 11–12 miles southwest of Autlán (approximately Type: Mexico, Querétaro, 2–5 km al S del Parador Santa
2 miles below the pass), McVaugh 857 (ENCB). Michoacán: Martha, municipio de Landa de Matamoros, bosque de encino
Aguililla, an area locally referred to as Cerritos de Agua, ca. three en ladera de cerro, E. Carranza 2585 (holotype: IEB!).
miles below lumber camp at Dos Aguas, nearly west of Aguililla, Lat. (Figure 12G).
approx. 18° 45′ N, long. 102° 56′ W, McVaugh 17,865 (ENCB, MICH). Decumbent perennial herb 70 cm long, sparsely hairy, with a
Nayarit: Ixtlán, km 7–10 terracería a Cacalotán, que empieza a woody base; stems creeping, rooting at the nodes, internodes very
500 m al S del límite de los estados Nayarit-Jalisco, 21° 07′ 00″ N, long, 6.5–10 cm long; leaves geminate throughout, petioles 1.5–2.3 cm
104° 17′ 00″ W, Téllez 11,066 (MEXU). Nuevo León: Horsetail Falls, long; blades ovate to lanceolate, 3.0–5.5 cm long, 1.5–3.0 cm wide,
30 mi SE of Monterrey, Thompson 294 (TEX). Oaxaca: Santa María apex acuminate, base truncate to cordate, margin entire or with
Jacatepec en el poblado La Joya del ejido Corriente Ancha, J. Calzada 2–3 teeth per side, with a few eglandular trichomes on the upper
15,426 (MEXU). Puebla: Zapotitlán de Méndez, G. Villalobos 23 and lower sides; solitary flowers on peduncles 1.5–2.5 cm long; calyx
(MEXU). Quintana Roo: Felipe Carrillo Puerto, 100 m al E del with acuminate lobes 0.5 mm long; corolla rotate, yellow, 1–2 cm in
Ramonal, R. Durán 756 (MEXU). Querétaro: 1–2 km aL SW de diameter, with five purple maculations, slightly pubescent corolla
Barricales, municipio de Jalpan, E. Carranza 3335 (IEB, QMEX). tube; anthers blue or yellow with blue lines 3 mm long; fruiting
Michoacán: Cerro La Alberca, cerca de Villa Jiménez, municipio de peduncles 1.3–2.0 cm long, fruiting calyx 5-angled, 1.1–1.9 cm
Villa Jiménez, J. Rzedowski 40,204 (IEB). Sinaloa: Culiacán, poblado de long, 0.8–1.0 cm wide with a few trichomes, berry 0.5 cm in
Jesús María y cercanías de la presa Adolfo López Mateos (El Verjonal), diameter with numerous yellow seeds 1.5 mm in diameter.
R. Vega 3305 (QMEX). Sonora: La Huerta, approximately 2 km inland Distribution and habitat: Endemic to Qro.; it grows in pine,
from Ensenada La Manga, R. Felger 85-346B (MEXU). Tabasco: Paraíso oak, and cloud forests at 1,700–1,850 m asl. (Figure 13F).
dentro del rancho La Noria a 1 km de Paraíso hacia la Ría Moctezuma, Diagnostic characters: A perennial trailing herb that roots at
M. Magaña 1007 (MEXU). Tamaulipas: Rio Corona, 2 km al E de la the nodes, with few eglandular trichomes.
Hacienda Santa Engracia, M. Martínez 1945 (MEXU, UAT). Veracruz: Common names and uses: None known.
Hidalgotitlán, Rio Soloxuchil, campamento Hnos. Cedillo, Vázquez et al. Phenology: Flowering and fruiting from July to December.
145 (MEXU, XAL). Yucatán: Yaxcabá, Tixcacaltuyub, V. Rico 709 Conservation status: LC.
(MEXU). Representative specimens examined. MEXICO. Querétaro: El
Parador, 2 km al NW de El Madroño, Landa de Matamoros, E.
48. Physalis purpurea Wiggins Contr. Dudley Herb. 3: 74, tab. 19 González 969 (IEB).
(1940)
50 Physalis rydbergii Fernald, Proc. Amer. Acad. Arts 35: 569
Type: Mexico, Sonora low hills and flats near the tannery east of (1900)
Guaymas. Ira L. Wiggins 6352 (holotype: CAS 0004035!). (Figures
1R, 12H). Type: Mexico, Sinaloa, Imala, E. Palmer 1713 (holotype: GH
Erect perennial herb, branched, 25–75 cm tall, spreading- 00077369!, isotypes: GH 00077370!, NY barcode 138888!, US
puberulent, especially on the stems, petioles, and peduncles; barcode 00027363!).
petioles 2–5 cm long, blades ovate to ovate-deltoid, 2.5–5.0 cm Erect perennial herb, 20–60 cm tall; stem branched, glandular-
long, 2–5 cm wide, apex acuminate, base truncate to cordate, pubescent with trichomes 2–3 mm long; basal leaves deciduous, the
margins coarsely and irregularly dentate to sinuate dentate; upper ones geminate, petioles 1.0–3.5 cm long; blades ovate-
flowers solitary on peduncles 17–25 mm long; calyx with lanceolate to lanceolate or trulate, 2–7 cm long, 1.5–4.0 cm wide,
broadly lanceolate to deltoid lobes 1–2 mm long; corolla violet apex acute, base rounded to attenuate, margin toothed with
or purplish, 9–13 mm wide; anthers yellow, 1.5–3.0 mm long; 2–5 teeth per side, glandular-pubescent on both surfaces, but
fruiting peduncles 2–4 cm long, fruiting calyx 10-costate, especially on margin and veins; flowers solitary on peduncles
23–30 mm long, 17–24 mm wide; mature berry green or 2–3 mm long; calyx with deltoid lobes ca. 3 mm long, corolla
yellow, nearly spherical, 10–18 mm wide with numerous rotate, yellow, ca. 1 cm in diameter with five strongly contrasting
yellow seeds ca. 2 mm in diameter. purple maculations; anthers blue, ca. 3 mm long; fruiting calyx 10-
Distribution and habitat: Endemic to the humid canyons costate, slightly invaginated, 1.5–2.0 cm long, 1.0–1.5 cm wide,
around Guaymas, near creeks or waterfalls, from sea level to glandular-pubescent; berry ca. 1 cm in diameter with numerous
160 m elevation. Son. (Figure 10C). yellow seeds ca. 2 mm diameter.
Diagnostic characters: A rare perennial plant with purple Distribution and habitat: It grows in xerophytic scrub and low
flowers and dentate leaves; the fruiting calyx is open at the apex. deciduous forest, 80–2,100 m. Gto., Mich, Qro., Sin. (Figure 5E).

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Martínez et al. 10.3389/fgene.2023.1080176

Diagnostic characters: The species is distinguished by its 52. Physalis solanacea (Schltdl.) Axelius, Phytologia 79: 11 (1995)
glandular trichomes, perennial habit, and small flowers and
fruits. Our description differs from that of Waterfall (1967) in Type: Cult. in Horto Botánico Halensis 1838, “e seminis in
the color of the anthers, which he describes as yellow, and the Mexico locis calidioribus coll. C. Ehrenberg” (holotype: HAL
blade size, but he was only familiar with the material of the type. 0033693!). Margaranthus solanaceus Schltdl., Index Seminum
Common names and uses: None known. Hort. Hal. 1838: 8 (1838). (Figures 1S, 14A).
Phenology: Flowering and fruiting from October to November. Erect annual plant 6–100 cm tall; stem with few branches,
Conservation status: DD. glabrous at the base and with a few eglandular multicellular
Representative specimens examined. MEXICO. trichomes on the upper part; petioles 0.5–2.5 cm long, the upper
Guanajuato: 4 km al E de San Diego de la Unión, sobre el leaves almost sessile; blades ovate to lanceolate, 1.5–7.0 cm long,
camino a La Jaula, municipio de San Diego, J. Rzedowski 0.5–3.0 cm wide, acute apex, base acute, sometimes oblique up to
52,099 (IEB, QMEX). Michoacán: cerca de San José Itzícuaro, 5 mm, margin entire, sometimes slightly wavy or with three to four
municipio de Morelia, J. Rzedowski 45,382 (IEB, QMEX). teeth per side, glabrous or with a few eglandular trichomes on the
Querétaro: fondo del Sótano del Barro, Santa María de Cocos, margin and veins; flowers solitary, peduncles absent or up to 4 mm
municipio de arroyo Seco, G. Cifuentes 69 (QMEX). Sinaloa: long; calyx ca. 2 mm long with triangular, pubescent lobes, corolla
Imala, E. Palmer 1713 (GH). urceolate, purple to greenish-yellow, 2–4 mm long; anthers blue, ca.
1 mm long; fruiting peduncle ca. 4 mm long, fruiting calyx 10-
51. Physalis sancti-josephi Dunal Prodr. [A. P. de Candolle] 13 costate, 0.6–1.5 cm long, 0.6–1.1 cm wide, pubescent with
(1): 451 (1852) eglandular trichomes; mature berry purple, 4–6 mm in diameter
with few (4–10) yellow seeds ca. 2 mm in diameter.
Type: Mexico, Hidalgo, prope San José del Oro, reg. frig. C. J. W. Distribution and habitat: The species grows in desert scrubs,
Schiede (holotype: HAL 42233!). tropical deciduous forest, grasslands, roadsides, streams, and
Erect perennial herb 60–100 cm tall; stem with few branches, farmland. It blooms from August to December at 630–2,400 m.
pubescent with glandular multicellular trichomes, glabrescent; Chih., Coah., Dgo., Gto., Hgo., Jal., Mich., N.L., Oax., Pue., Qro.,
basal leaves alternate, upper leaves geminate, petioles 2.5–4 cm S.L.P., Tamps., Ver., Zac. (Figure 13B).
long, blades ovate, 4–8 cm long, 2.5–4.5 cm wide, apex acuminate, Diagnostic characters: A purple, urceolate corolla, small fruits,
base rounded, margin entire or with one to three teeth per side, 10-costate fruiting calyx. It is distinguished from P. lagascae and P.
pubescent with multicellular glandular trichomes on both surfaces; ampla, both of which bear small fruits, by its glabrous calyx.
flowers solitary on peduncles 1.0–1.4 cm long; calyx with Common names and uses: Tomatillo, mata pulgas. No uses
triangular lobes 4–5 mm long; corolla rotate, yellow, 1.2–2.0 cm known.
in diameter with strongly contrasting solid or compound blue Phenology: Flowering and fruiting from July to December.
maculations, corolla throat densely pubescent; anthers yellow or Conservation status: LC.
yellow with blue lines, ca. 3 mm long; fruiting peduncle 1.0–1.2 cm Representative specimens examined. MEXICO. Chihuahua:
long, fruiting calyx 10-costate, slightly invaginated, 2.5–3.0 cm km 98 de la carretera Camargo-Delicias, municipio de Saucillo,
long, 1.5–2.5 cm wide, glandular-pubescent; fruit 0.7–1.1 cm in M. Martínez 9680 (QMEX). Coahuila: 5 km antes de General
diameter with numerous yellow seeds ca. 1.5 mm in diameter. Cepeda, M. Martínez 9594 (QMEX). Durango: Mezquital, 3 km
Distribution and habitat: It develops in pine, oak, and cloud de Temohaya por el camino a Mezquital, González 1601
forests at 1,600–1,800 m. Hgo., Jal, Nay., Qro., S.L.P. (Figure 13E). (MEXU). Guanajuato: 15 km al W de Salvatierra sobre la
Diagnostic characters: The species is poorly characterized. It carretera a Yuriria, J. Rzedowski 38793 (IEB, MEXU).
resembles underdeveloped specimens of P. coztomatl, with which it Hidalgo: Cardonal, Tolantongo, J. Rzedowski 113347 (IEB).
shares habitat and distribution. They differ in that P. coztomatl has a Jalisco: Tala, a lo largo del arroyo Caliente y Los Letreros,
stem densely covered by trichomes up to 3 mm, whereas P. sancti- Bosque Escuela La Primavera, A. Rodríguez 1436 (IBUG).
josephi has much smaller trichomes and corolla, and the Michoacán: estación del tren, municipio de Charo,
maculations can be solid. J. Escobedo 2125 (IEB). Nuevo León: Montemorelos, en la
Common names and uses: None known. salida hacia General terán, ca. 3 km del centro del pueblo,
Phenology: It blooms all year round, but it has been collected M. Martínez 9808 (QMEX). Oaxaca: Heroica Ciudad de
with flowers and fruits from February to March and with fruit only Huajuapan de León, a 3 km del límite Puebla-Oaxaca, a un
in May. costado de la carretera 125 Tehuacán-Huajuapan de León.
Conservation status: DD. Distrito Huajuapan, L. Pérez 4 (MEXU). Puebla: Caltepec, La
Representative specimens examined. MEXICO. Hidalgo: Laguna, Cerro El Gavilán, E de Caltepec, P. Tenorio 21082
prope San José del Oro, reg. frig. C. J. W. Schiede (HAL). Jalisco: (MEXU). Querétaro: Misión de Tilaco, municipio de Landa,
Autlán de Navarro, arroyo de Las Juntas, 18–19 km aL SE de Autlán, M. Martínez 2789 (QMEX). San Luis Potosí: Guadalcázar, Los
3–4 km al N de El Zarzamoro, 19° 37′ 16″ N, 104° 16′ 36″ W, Aguajitos, 11 km aL NE de Guadalcázar, hacia Pozo de Acuña,
Guzmán 984 (ZEA). Nayarit: Tepic, 10 km al W de El Izote, brecha R. Torres 17,151 (MEXU). Tamaulipas: carretera Victoria-
al Cuarenteño, 21° 30′ N, 104° 58′ W, P. Tenorio 15,684 (ENCB, Llera, en el km 203 en la torre de telecomunicaciones,
MEXU, MICH). Querétaro: 11 km de el Cañón, camino a la Parada, municipio de Llera, M. Martínez 9894 (QMEX). Veracruz:
municipio de Jalpan, E. Carranza 1425 (IEB, QMEX). San Luis Pánuco, 4 km al S del límite del estado de Veracruz y
Potosi: Alvarez E. Palmer 220 (F, GH, MICH, NY, UC, US) Tamaulipas, 10 km aL SW de la carretera Tampico-Pánuco,

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Martínez et al. 10.3389/fgene.2023.1080176

L. Nevling 371 (MEXU). Zacatecas: 15 miles NE of Estación Type: United States, Texas, Rio Bravo, sea beach, A.C.V.
Camacho, in wash one mile NW of Pico de Teyra, in Arroyo de Schott 30. (holotype: NY 402103!, isotype: F barcode
Borrego, S of Cerro de Borrego J. Henrickson 13515 (TEX). V0072990F!). Physalis lanceolata var. spathulifolia Torr. in
Emory, Rep. U.S. Mex. Bound., Bot. [Emory] 153 (1858).
53. Physalis sordida Fernald, Proc. Amer. Acad. Arts 35: 568 (Figure 14B).
(1900) = Physalis cinerascens var. spathulifolia (Torr.) J.R. Sullivan, Syst.
Bot. 10 (4): 444 (1985).
Type: Mexico, Oaxaca, Boca de León, Telixtlahuaca, A. L. Smith = Physalis viscosa var. spathulifolia (Torr.) A. Gray Proc. Amer.
637 (holotype: GH 00077372!). (Figure 14C). Acad. Arts 10: 67 (1874).
Erect or prostrate perennial herb 20–60 cm tall; stem highly Decumbent perennial herb, 30–40 cm tall, with a long horizontal
branched, densely covered with glandular trichomes; basal leaves rhizome covered with branched trichomes throughout, mixed with
alternate, the upper ones geminate, petioles 0.7–2.0 cm long; blades glandular trichomes in young parts; leaves geminate, petioles
ovate to trulate, 1.5–4.5 cm long, 1–2.5 cm wide, apex acute, base 5–13 mm long or absent; blades spathulate, rarely ovate,
attenuate to rounded, margin toothed with four–six teeth per side, 5.5–8.0 cm long, 2.5–4.5 cm wide, apex obtuse, base attenuate,
sometimes almost entire, densely glandular-pubescent on both surfaces; margin entire; flowers solitary on peduncles 2.2–3.5 cm long;
flowers solitary on peduncles 5–7 mm long, glandular-pubescent; calyx calyx with triangular lobes ca. 2 mm long; corolla rotate-
with triangular lobes 3–4 mm long, glandular-pubescent; corolla rotate, campanulate, yellow, 1.3–1.5 cm in diameter with five solid
yellow, 1.5–2.0 (2.5) cm in diameter with strongly contrasting blue purple strongly contrasting maculations; anthers yellow, 3 mm
maculations, sometimes glandular-pubescent outside; anthers blue or long; fruiting peduncle 3.0–3.5 cm long, fruiting calyx 10-costate,
yellow with blue lines, ca. 3 mm long; fruiting peduncles 6–7 mm long, not invaginated at the base, 3.5–4.0 cm long, 2.5–3.0 cm wide,
fruiting calyx 10-costate, slightly invaginated, 1.5–2.5 cm long, 1–2 cm mature berry yellow, 2 cm in diameter, containing numerous
wide, glandular-pubescent; mature berry purple ca. 1.5 cm in diameter, yellow seeds ca. 2 mm in diameter.
glutinous with numerous yellow seeds ca. 2 mm diameter. Distribution and habitat: Restricted to the sandy shores of
Distribution and habitat: It grows on roadsides or secondary Tamaulipas, at sea level. Tamps. (Figure 5B).
vegetation of pine, oak, and xerophytic scrub, 1,800–2,000 m. Coah., Diagnostic characters: A perennial with branching trichomes
Dgo., Edo. Mex., Gto., Hgo., Jal., Mich., NL., Oax., Pue., Qro., S.L.P., restricted to sandy shores.
Tam., Zac. (Figure 13C). Common names and uses: None known.
Diagnostic characters: A dense glandular pubescent plant with Phenology: Flowering and fruiting from September to March.
blue anthers and a slightly leathery 10-costate fruiting calyx. The Conservation status: LC.
plant is commonly covered with dirt. Representative specimens examined. MEXICO. Tamaulipas:
Common names and uses: None known. Playa de La Pesca, municipio de Soto la Marina, M. Martínez 9482
Phenology: Flowering and fruiting from March to November. (QMEX).
Conservation status: LC.
Representative specimens examined. MEXICO. Coahuila: 55. Physalis subrepens Waterf., Rhodora 69: 229 (1967)
camino entre Sierra el Coahuilón y Sierra La Marta, M. Yañez
196 (MEXU). Durango: Along Hwy 40, 4 km NE of Yerbaniz, 21 km Type: Mexico, Hidalgo, Trinidad Iron Works, C. G. Pringle
SW of Cuencamé, G. Diggs (XAL). Estado de México: Texcoco, carr. 13,591 (holotype: VT not located; isotypes: F V0073012F!, GH
Mex.-D.F. km 25, L. Scheinvar 526 (MEXU). Guanajuato: San Luis 00077374!, MICH 1109896!, OKLA not located, TEX!, UC not
de la Paz R. Hernández 134a (MEXU). Hidalgo: Ajacuba La Mesa located, US barcode 00027367!). (Figure 14D).
Chata, cerro al NW del poblado Santiago Tezontlale, sierra del Mexe, Erect perennial herb up to 1.5 m tall, with a rhizome 20 cm long,
ejido Santiago Tezontlale I.Díaz 1221 (MEXU). Jalisco: Ojuelos, pubescent with eglandular trichomes; leaves alternate to geminate
Paso de La Troje, near km 36, southwest of Ojuelos on the road to almost from the base, petioles 1.3–2.5 cm long; blades ovate to
Aguascalientes, rocky slopes on and near Cerro La Campana, broadly ovate or suborbicular, 2.6–7.0 cm long, 2.2–4.3 cm wide,
McVaugh 16836 (MICH). Michoacan: A 13 km aL SE de Villa apex acute or slightly attenuate, base oblique, cuneate, somewhat
Madero, Carretera a Nocupetaro (IBUG). Nuevo León: road from decurrent to subcordate, margin entire or with few teeth, hairy with
San Rafael to 18 de Marzo and Galeana, ca. eight miles E of San scattered trichomes on veins and margins or glabrous; flowers
Rafael, 4.4 miles E of La Boca L. J. Dorr (TEX). Oaxaca: Teotitlán a solitary on peduncle 0.8–2.3 cm long; calyx with ovate to
6 km SE, on highway between Tehuacán and Telixtlahuaca, S. triangular lobes 3–4 mm long, corolla rotate campanulate, pale
Douglas (MEXU). Puebla: 3 km al N de Cuicatlán, O. Téllez yellow, 1.3–1.6 cm in diameter, with simple brownish
(QMEX). Querétaro: Cerro El Azteca (Los Cajones) L. maculations; anthers blue, 2–3 mm long; fruiting peduncle
Hernández 4855 (MEXU). San Luis Potosí: Chiefly in the region 1.3–2.5 cm long, fruiting calyx 5-angled, 1.6–3.3 cm long,
of San Luis Potosí, C. C. Parry (US). Tamaulipas: Sierra de 1.2–1.8 cm wide; mature berry 0.6–1.0 cm in diameter.
Guatemala. Barbara’s Patch. Halfway down Mine road toward Distribution and habitat: It develops in pine–oak or cloud
Mine lookout J. Sullivan s. n. (ENCB). Zacatecas: S slope of La forests, on slopes, or in open places. It grows at altitudes close to
Bufa R. Dressler (MO). 2,100 m. Edo. Mex., Hgo., Jal., Ver.
Diagnostic characters: The species is hairy, with roots at the
54. Physalis spathulifolia (Torr.) B. L. Turner, Phytologia 93 (2): basal nodes, geminate leaves, long peduncles in flower and fruit, and
264 (2011) a 5-angled fruiting calyx. Physalis subrepens can be confused with P.

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Martínez et al. 10.3389/fgene.2023.1080176

gracilis because of the geminate leaves, prostrate habit, and long Type: Mexico, Jalisco: Tapalpa, dirt road from Tapalpa to
flower and fruit peduncles, but P. gracilis has a 10-costate fruiting Chiquistlán, 2002′02´´N, 10,350′23´´W, O. Vargas 876 (holotype:
calyx. IBUG!; isotypes: MEXU!, NY!). (Figure 1U).
Common names and uses: Tomatillo; no uses known. Erect shrub up to 1.5 m tall, stems with eglandular trichomes
Phenology: It bears flowers and fruit from April to August, but mixed with glandular trichomes except in young parts; leaves
its phenology is probably longer. alternate at the base, soon geminate, petioles 1.0–3.8 cm long;
Conservation status: LC. blades ovate, 3.4–9.1 cm long, 2.3–6.7 cm wide, apex acuminate,
Representative specimens examined. MEXICO. Estado de base oblique to truncate, margin serrate; flowers solitary on
México: Contreras Galvan s.n. (MEXU). Hidalgo: Barranca near peduncles 0.5–1.0 cm long; calyx with acuminate lobes,
Honey Station, Pringle 13,440 (GH, MICH, US, VT). Jalisco: Autlán 1.5–2.5 mm long, corolla campanulate-rotate, yellow, 1.0–1.8 cm
de Navarro, ladera S en la falda del cuamil Zermeño, Las Joyas, in diameter, with dark brown simple-to-compound maculations;
Vázquez García 3266 (MEXU, ZEA). Veracruz: Huayacocotla, 1 km anthers purple or blue, 2.5–3.0 mm long; fruiting peduncle
SW of Ixtatetla, Nee and Taylor 26845 (XAL). 0.9–1.4 cm long, densely pubescent, fruiting calyx 10-costate,
2.4–3.5 cm long, 1.6–2.5 cm wide, velutinous, reticulate with
56. Physalis sulphurea (Fern.) Waterf., Rhodora 69: 224 (1967) purple veins; mature berry ca. 1.1 cm in diameter, yellow seeds,
1.5–1.8 mm in diameter.
Type: Mexico, Distrito Federal, the valley of Mexico, C. G. Distribution and habitat: Marginal vegetation of pine–oak
Pringle 8215 (lectotype selected by Waterfall (1967): GH 77375!) forest, under tejocote trees (Crataegus mexicana), or in ravines
Margaranthus sulphureus Fernald Proc. Amer. Acad. Arts 35: within the forest, at an altitude range of 1,910–2,150 m. Endemic,
566–567. 1900. (Figures 1T, 14F). Jal. (Figure 10E).
Erect or creeping annual herb 8–70 cm tall; stems with few Diagnostic characters: A bush or suffrutex with glandular
branches, angled, hollow, glabrous at the base, and puberulent with pubescence, small flowering calyx, and 10-ribbed fruiting calyx. It
eglandular trichomes on younger parts; basal leaves alternate, the shares with P. lignescens the small flowering calyxes and reflex lobes.
upper ones geminate, petioles 0.5–4.0 cm long; blades ovate to Common names and uses: None known.
lanceolate, 1.5–5.5 cm long, 0.8–3.0 cm wide, apex acute, base Phenology: Flowering and fruiting from July to November.
attenuate, decurrent, sometimes oblique up to 1 cm, margin Conservation status: NT.
entire to toothed, glabrous to sparsely puberulent with Representative specimens examined. MEXICO. Jalisco: Tapalpa,
multicellular trichomes on the margin and veins; flowers solitary dirt road from Tapalpa to Chiquistlán, P. Zamora 207 (IBUG).
on peduncles 0.8–2.0 cm long; calyx with triangular lobes ca. 1 mm
long; corolla rotate, whitish or pale yellow, 1.0–1.5 cm in diameter 58. Physalis tehuacanensis Waterf., Rhodora 69: 203 (1967)
with slightly contrasting brown maculations, corolla throat glabrous;
anthers blue, ca. 2 mm long; fruiting peduncles 0.8–1.5 cm long, Type: Mexico, Puebla Tehuacán city dump, C. E. Smith Jr,
fruiting calyx 10-costate, 0.8–1.4 cm long, 0.5–1.2 cm wide; mature Peterson and Tejeda 3992 (holotype: F barcode V0073015F!, isotype:
berry green ca. 0.5 cm in diameter with numerous yellow seeds ca. US barcode 00027369!)
3 mm in diameter. Perennial herb 22–27 cm tall with articulated capitate-
Distribution and habitat: In wet places around lakes, dams, and glandular trichomes up to 1.5–2.0 mm long; leaves geminate;
stream shores, at 1,700–2,000 m, Cd. Mex., Dgo., Gto., Jal., Mich., blades ovate or deltoid, 12–35 mm long, 12–35 mm wide, apex
Qro. (Figure 13D). acute, base truncate, margin unevenly coarsely dentate or sinuate-
Diagnostic characters: The whitish or pale-yellow corolla and dentate; flowers solitary on peduncles 4–6 mm long; calyx with
hollowed stem; it is the only aquatic species of Physalis. lanceolate or ovate-lanceolate lobes 2.0–2.5 mm long, corolla
Common names and uses: Jitomate, tomatillo. No uses known. rotate, yellow, immaculate, 10–11 mm wide; anthers yellow,
Phenology: Blooms from April to December. 2.7–3.5 mm long; fruiting peduncles 8–10 mm long, fruiting
Conservation status: LC. calyx 10-costate, 15–18 mm long, 14–16 mm wide, berry
Representative specimens examined. MEXICO. Ciudad de 10–11 mm in diameter.
Mexico: Xochimilco, Cienega Grande, E. Martínez 39,817 Distribution and habitat: Endemic to the Tehuacán area in the
(MEXU). Durango: Villa Unión, al E del 18 de Agosto, state of Puebla at 1800 m asl. Pue. (Figure 10B).
González 1234 (CHAPA, CIIDIR-DGO, ENCB). Guanajuato: Diagnostic characters: Long, coarse, multicellular trichomes
Salvatierra, 5 km aL NE de Salvatierra, sobre la carretera a intermixed with shorter ones, many of the trichomes tipped with
Celaya, Rzedowski 38572 (ENCB, MEXU). Jalisco: Chapala, reddish-brown glands, coarsely and irregularly dentate leaves, and
University of Guadalajara Country Club, Chapala, on Laguna yellow anthers.
de Chapala, 21 ° 17′ N, 103 ° 4′ W, Iltis 29,162 (IBUG, WIS). Common names and uses: None known.
Michoacán: alrededores de Copándaro, municipio de Villa Phenology: Collected with flowers in June.
Jiménez, J. Rzedowski 46491 (IEB). Querétaro: presa de Conservation status: CR.
Santa Catarina, A. Herrera 198 (QMEX). Representative specimens examined. MEXICO. Puebla:
Tehuacán city dump, C. E. Smith Jr, Peterson and Tejeda 3992 (F).
57. Physalis tamayoi O. Vargas, M. Martínez & Dávila, Brittonia
53 (4): 507–509 (2001) 59. Physalis vestita Waterf., Rhodora 69: 326 (1967)

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Martínez et al. 10.3389/fgene.2023.1080176

Type: Mexico, Sinaloa, vicinity of Mazatlán, J.N. Rose 13766 Phenology: Bears flowers and fruits from June to March.
(holotype: US barcode 00027372!). (Figure 14H). Conservation status: LC.
Erect perennial herb 15–65 cm tall from woody elongated Representative specimens examined. MEXICO. Jalisco:
roots; stems highly ramified, densely covered with branched Concepción de Buenos Aires, 5 km al S de Buenos Aires, brecha
trichomes throughout; leaves geminate, petioles 10–35 mm a Ciudad Guzmán, Villa 756 (IEB). Michoacán: Lado S de La Laguna
long; blades ovate, 25–40 mm long, 15–28 mm wide, apex Verde, zona geotérmica Los Azufres, municipio de Zinapécuaro,
acute, base truncate to oblique, margin entire or with two to M.J. Jasso 1094 (IEB).
three teeth per side, densely covered with gray branched
trichomes; flowers solitary on peduncles 4–6 mm long; calyx 61. Physalis waterfallii O.Vargas, M. Martínez & Dávila, Acta Bot.
with deltoid lobes 1 mm long, corolla rotate, yellow, Mex. 48: 22 (1999)
10–13 mm in diameter, with five purple maculations, corolla
throat densely pubescent; anthers violet, 3–4 mm long; fruiting Type: Mexico, Jalisco, municipio de Chiquilistlán, brecha
peduncle 7–13 mm long, fruiting calyx 5-angled, densely covered Tapalpa-Chiquilistlán, A. Rodríguez 986 (holotype: IBUG!;
with branched trichomes, 15–22 mm long, 13–18 mm wide; isotypes: ENCB!, MEXU 445306!). (Figure 14I).
mature berry green, 8–10 mm in diameter with numerous Perennial prostrate herb 60–100 cm long, lignified at the base,
yellow seeds ca. 1 mm in diameter. with eglandular trichomes ca. 3 mm long; leaves alternate at the base,
Distribution and habitat: Endemic to the rocky coast of soon geminate, petiole 1.3–4.0 cm long, blades ovate-lanceolate to
Sinaloa, known only from three localities at sea level. broadly ovate, 3.5–11.0 cm long, 2–7 cm wide, apex acute to caudate,
Diagnostic characters: A perennial plant densely covered base truncate, oblique, narrowly decurrent, margin entire, rarely
with branched trichomes. However, the trichomes are with one to three teeth, pubescence appressed; flowers solitary on
forked, whereas the trichomes of P. cinerascens have several peduncles 1.3–3.0 cm long; calyx with triangular lobes 0.6–1.0 cm
ramifications. long, corolla campanulate rotate, yellow, 1.4–2.0 cm long, 2–3 cm in
Common names and uses: None known. diameter, with simple to compound reddish-brown maculations;
Phenology: Bears flowers and fruits in June. anthers purple or blue, 3.0–3.5 mm long; fruiting peduncles
Conservation status: Not evaluated, but due to its restricted 1.5–2.8 cm long, fruiting calyx 5-angled, 2.2–3.8 cm long,
distribution, probably VU. 1.2–2.6 mm wide; mature berry green to yellow ca. 1 cm in
Representative specimens examined. MEXICO. Sinaloa: barra diameter with numerous yellow seeds ca. 2 mm in diameter.
de Piaxtla, en el faro, M. Martínez 9712 (QMEX). Distribution and habitat: In pine–oak and cloud forests, in
clearings and agricultural fields near the forest at 1,700–2,450 m. Jal.,
60. Physalis volubilis Waterf., Rhodora 69: 229 (1967) Mich., Qro. (Figure 13F).
Diagnostic characters: A long flower peduncle, the 5-angled
Type: Mexico, Michoacán, two miles above Tancitaro, Wm. C. fruiting calyx, and the long calyx lobes in flower and fruit up to
Leavenworth 519 (holotype: GH 00077383!; isotypes: F barcode 1.0 cm in length.
V0073021F!, MICH 138898!, NY 1109892!, MEXU!). (Figure 14G). Common names and uses: None known.
= Physalis viridoflava Waterf., Rhodora 69: 230 (1967). Type: Phenology: It flowers and bears fruit from July to November.
Mexico, Jalisco, on route 110 at km 59–60, 20 miles due WSW of Conservation status: LC.
Jiquilpan, and several miles S of Mazamitla, D. P. Gregory, G. Eiton 99 Representative specimens examined. MEXICO. Jalisco:
(holotype: MICH barcode 1109893!, isotype: NY barcode 138896!). Chiquilistlán, brecha Tapalpa-Chiqulistlán, Rodríguez 986 (ENCB,
Decumbent perennial herb up to 80 cm long, lignified at the base, IBUG, MEXU). Michoacán: municipio de Zinapécuaro, El Cerrito,
rooting at the basal nodes, glabrous or with few eglandular, septate, 1 km al E de Jeráhuaro, M. Jasso 199 (IEB, QMEX). Querétaro: entre
appressed trichomes; leaves alternate throughout, petioles 1.2–7.1 cm San Joaquín y Corral Blanco, municipio de San Joaquín, S. Zamudio
long; blades ovate to broadly ovate, 1.0–5.0 cm long, 1.0–4.5 cm wide, 7945 (IEB).
apex obtuse to acute, base truncate to cordate, margin entire; flowers
solitary on peduncle 3.5–8.5 cm long; calyx with narrow triangular
lobes 4–7 mm long, corolla campanulate, rotate-reflected, yellow, 9 Doubtful and excluded names
2.0–2.8 cm in diameter, with dark, purple-brown, compound
maculations, corolla throat pubescent; anthers yellow, blue-tinged, Physalis ixocarpa Brot. ex Hornem, Hort. Bot. Hafn. Suppl. 26
2.5–4.0 mm long; fruiting peduncles 1.7–6.0 cm long, fruiting calyx 5- (1819).
angled, 2.0–2.6 cm long, 1.1–1.4 cm wide, mature berry 0.8–1.2 cm in
diameter with yellow seeds ca. 1.5 mm in diameter. Type: Mexico (BR 0000013069179!).
Distribution and habitat: It develops mainly on slopes, Annual herb up to 130 cm high, stems angled, puberulent,
margins, or clearings of pine–oak and cloud forests. It grows at glabrescent; leaves alternate, petioles 1.8–4.1 cm long; blades
an altitude range of 1,390–2,700 m. Mich., Jal. (Figure 5F). linear to rhombic or ovate, 4.5–14 cm long, 2.3–5.1 cm wide,
Diagnostic characters: A prostrate habit, ovate leaves, a short apex acute, base attenuate, sometimes oblique, margin dentate;
rhizome, and long peduncles in flower and fruit. flowers solitary on peduncle ca. 5 mm long; calyx with triangular
Common names and uses: None known. lobes 1.7 mm long, corolla rotate, yellow, maculate, 0.7–1.2 cm in

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Martínez et al. 10.3389/fgene.2023.1080176

diameter; anthers green or blue-tinged, 1–2 mm long; fruiting because P. muelleri has a fruiting calyx almost filled by the
peduncle 1 cm long, fruiting calyx 10-costate, 1.6 cm long, 3.4 cm berry and opened apically. However, the characters may be
wide, berry 1–2 cm in diameter. pressing artifacts.
Waterfall (1967) did not include this species in his treatment.
Under our concept, the plant is similar to P. philadelphica, from which Physalis philippensis Fernald, Proc. Amer. Acad. Arts 35: 568
it differs in that the lectotype does not have convoluted anthers. (1900).
Additionally, a capitate stigma was referred to P. ixocarpa (Fernandes,
1974), whereas P. philadelphica has a clavate stigma. Plants from Type: Mexico: Oaxaca, Sierra de San Felipe C.G. Pringle 5621
Southern Mexico (Tabasco and Chiapas) can be referred to this taxon (holotype: GH 00003292!, isotype: VT not located).
and were treated as such for Flora Mesoamericana (unpublished). The Perennial herb, 15–20 cm long, several-branched; vestiture of
name P. ixocarpa is misapplied to cultivated husk tomato. According flattened, spreading, jointed trichomes of varying sizes; petioles
to molecular data, both species form a clade behaving as sisters 6–12 mm long, blades ovate to rhombic ovate to hastate-ovate,
(Deanna et al., 2018). We consider that P. ixocarpa could be 15–20 mm long, 15–20 mm wide, apex obtuse to acute, base
synonymous with P. philadelphica; however, more extensive work decurrent, margin usually with one to three irregularly shaped
is needed to address this issue. teeth, spreading-hairy to appressed-hairy on both sides; solitary
flowers on peduncles 5–8 mm long; calyx lobes ovate-lanceolate to
Physalis microphysa A. Gray, Proc. Amer. Acad. Arts 21: 402 deltoid; corolla slightly lobed, 20–25 mm wide, with violet or purplish
(1886). compound maculations, corolla throat tomentose; anthers bluish or
violet, oblong, 3–4 mm long; fruit and seeds unknown.
Type: Mexico, near the city of Chihuahua: Santa Eulalia The species was described over 120 years ago, but no recent
Mountains, limestone cliffs, C. G. Pringle 317 (holotype: GH 3286!, collections fit the description. The lack of fruit further complicates
isotypes: BR not located, F barcode V0073004F!, MEXU-2762!). the species concept.
= Physalis campanulata Brandegee, Univ. Calif. Publ. Bot. 4: 278
(1912). Type: Mexico, San Luis Potosí, C. A. Purpus 5313 (holotype:
UC not located, isotypes: F not located, GH 00077343!, MEXU- Author contributions
27635, US barcode 00027317!).
Perennial herb from a woody base up to 60 cm tall; trichomes a MM: concieved the paper, worked on the manuscript, located
mixture of glandular and eglandular trichomes; leaves alternate, type material, elaborated the key and the descriptions, reviewed the
petioles 5–12 mm long; blades reniform to ovate, 5–12 mm long, manuscript. OV: concieved the paper, worked on the manuscript,
10–23 mm wide, apex acute, base attenuate, margin dentate; flowers located type material, elaborated the key and the descriptions,
in pairs on peduncles 2 mm long; calyx with triangular lobes reviewed the manuscript. PZ: located type material, located
2–3 mm long, corolla rotate, yellow, immaculate, 8–10 mm in previously published material, elaborated the key and the
diameter; fruiting calyx 10-costate, open apically, 9–12 mm long; descriptions, concieved and structured the figures, reviewed the
fruit dry, dehiscent through irregular lines; seeds few, black. manuscript.
The species is to be removed from Physalis because molecular
evidence places it closer to Chamaesaracha and Quincula than to the
rest of Physalis (Zamora-Tavares et al., 2016). Morphologically, it Funding
differs from Physalis in its open fruiting calyx, dry dehiscent fruit,
and black seeds. CONABIO funded recent collections through the grant
SNIB-CONABIO, Proyecto no. RG002/Proyecto Agrobiodiversidad
Physalis muelleri Waterf., Rhodora 69: 115 (1967). Mexicana GEF 9380 awarded to the first author.

Type: Mexico, Nuevo León, Diente Canyon, mountains near

Monterrey, C. H. Mueller 129 (holotype: GH 77357!, isotype: F Acknowledgments
V0073005F!)
Perennial herb, 25–45 cm tall; trichomes short, up to 0.5 mm long, Marco Antonio Anguiano Constante tested our species key and
partly capitate-glandular; petioles 15–35 mm long, leaves ovate, 4–8 cm made suggestions to improve it. Yolanda Pantoja Hernández
long, 2.8–5.5 cm wide; flowers solitary on peduncles 5–6 mm long; elaborated the distribution maps. Luis Hernández-Sandoval
calyx 6–8 mm long and ca. 5 mm wide at the base of the ovate-deltoid reviewed earlier drafts of the paper. Alan Herrera detected
or lanceolate lobes; corolla campanulate, yellow, 10–12 mm long, several errors in the species distributions and improved the final
12–15 mm wide with dark maculations; anthers yellow, 3.5–4 mm version of the manuscript.
long; fruiting peduncles 10–15 mm long, fruiting calyx 10-costate,
with short trichomes, partly capitate-glandular, 17–22 mm long and
15–18 m wide; mature berry 10–14 mm wide, seeds unknown. Conflict of interest
The species is only known from the type specimen. The plant is
a perennial with coarsely dentate leaves and glandular trichomes, The authors declare that the research was conducted in the
but all those characters are also present in P. hederifolia. In his key, absence of any commercial or financial relationships that could be
Waterfall (1967) distinguished P. muelleri from P. hederifolia construed as a potential conflict of interest.

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Martínez et al. 10.3389/fgene.2023.1080176

Publisher’s note organizations or those of the publisher, the editors, and the
reviewers. Any product that may be evaluated in this article, or
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