Malvaceae of Mexico

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LopiMiA MALACOPHYLLA (Link & Otto) Martius

(Koch et al. 79511).

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MALVACEAE OF MEXICO Paul A. Fryxell of Agriculture Department in cooperation with Texas A&M University College Station, Texas 77843

U.S.

Abstract. occurs

The one

inMexico, of Mexico.

"Malvaceae of

its centers

of Mexico" of floristic

treatment of the family as it floristic is a revisionary-level to an eventual it is intended as a contribution diversity;

ten infraspecific is genera taxa) are treated. Discussion (372 species including Fifty-five to endemism and diversity in the family and to taxonomic subdivision of the family, wherein as occurring one of which four tribes are recognized inMexico, is further subdivided into (the Malveae) of 16 generic alliances. the is with attention reviewed, family special Morphology given to specialized

Flora

devoted

and for each genus the following relevant to the family. A key to the genera is presented, a state is given: and derivation, synonymy, generic description, etymological typification ment on the size and distribution of the genus, citation of important references the genus, concerning and a key to the Mexican information is provided: species of the genus. For each species the following and synonymy names and uses with full bibliographical vernacular data, species description, typification terminology information

of the plant if any, distribution and elevational) and flowering citations period, specimen (geographical are included for most of the genera. Illustrations Scientific names and map. by states, and a distribution are fully indexed. Two new sections sect. Mexabutilon citations of Abutil?n and specimen (Abutil?n Abutil?n Blanchard cies,

are published, as are three new sections of Hibiscus described Pluriovulata) by O. sect. Muenchhusia, sect. Striati). Four new spe sect. Clypeati, Hibiscus Hibiscus are de Pavonia and Pavonia tarasoides, alia, Pavonia baumliana, durangensis and a new combination, Melochia is proposed. the new genus Also, (Sterculiaceae), spicata sect.

(Hibiscus Bastardiastrum

scribed, Allosidastrum Allosidastrum

and

the new

pyramidatum

combinations

Allosidastrum

hilarianum,

Allosidastrum

interruptum,

and

are published.

INTRODUCTION are incomplete, or treatments of the Malvaceae of Mexico outdated, Existing otherwise unsatisfactory for understanding the taxonomic diversity of this family in Of the more Mexico. is incomplete significant sources, Urbina's (1897) checklist and provides neither keys nor descriptions, are cited; Stand although specimens treatment deals with the ley's (1923) only woody species; Wiggins (1964) treats only more the species of the Sonoran Desert the detailed region; keys of Kearney concern the Other references, (1954a, 1954b, 1955a, 1955b) only larger genera. such as Ferris (1927), Martinez (1955), Cowan (1983), Sousa and Cabrera (1983), & Rzedowski Fryxell (in Rzedowski 1985), Lott (1985), and Breedlove (1986), are local in their coverage. None of these, including the generic treatments of Kearney recent advances in realigning (1951a) and Hutchinson (1967), has incorporated new and boundaries within the it is genera generic establishing family. Therefore, a as a as occurs to one treatment of the undertake whole it in Mexico, timely family of the regions of its richest development. are quite diversified with 382 taxa (372 species The Malvaceae of Mexico 21 and distributed varieties among 55 genera; in addition, a including subspecies) of the introduced Lagunaria single specimen (L. patersonii G. Don; Bright 5316) 1

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2

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

in Tijuana, Baja California. The family is principally has been seen from cultivation in origin, although it iswell represented in the American South American, probably inAustralia the Old World as well, with diversification Plagianthus (e.g., complex), Africa East the Mediterranean and Middle (Hibiscus, (e.g., Lavatera, Alcea), Anisodontea K. Presl), and Madagascar Mexico, however, ap (tribe Hibisceae). of the family separate from its South pears to represent a region of diversification American center; its mallow flora is richer than that of adjacent areas of Central and it contains a large number of endemic America, species and about a dozen so. genera that are endemic or essentially re workers have significantly studies of the family by contemporary Recent new genera, and otherwise established sharpened our generic boundaries, as treatments such of Available the Standley understanding (1923) and family. now a signifi are out of date. somewhat Moreover, Kearney (1951a) consequently cant number of new species and genera of Mexican Malvaceae have been described into the systematics of the family. in recent years that need to be incorporated in several respects. At one level, it has The family Malvaceae has significance and history of mankind had enormous through the genus impact on the economy cottons. This story is a long and the cultivated from which derive Gossypium vised

Silow & Stephens involved one, better told elsewhere 1947; Fryxell (Hutchinson, 1979b). The family has also had impact on human aesthetics by way of highly prized in such genera as Hibiscus, Malvaviscus, ornamentals Alcea, Phymosia, Lavatera, of tropical regions and others. Indeed, one of the most widely known ornamentals the world is undoubtedly rivaled Hibiscus rosa-sinensis, only by throughout and by another mallow, Bougainvillea spectabilis Willdenow (Nyctaginaceae) in temperate Malvaviscus regions Alcea rosea perhaps takes second penduliflorus; and Alcea are highly diverse in horticultural place only to Rosa itself. Both Hibiscus and variants. A few species such as Hibiscus lavaterioides, Lavatera assurgentiflora, for ornament. into occasional cultivation Pavonia candida have been introduced the Mexican representatives, Many others stand available for similar use. Among one might mention Phymosia spp. (flowering spp. (flowering shrubs), Robinsonella Pavonia and perhaps Pavonia macdougallii, trees), Anotea gentryi, flavida, Bakeridesia spp., and Abutil?n grandidentatum. of the Malvaceae include: a vegetable Other uses for members {Abelmoschus a a condiment and as forage (Malva parviflora), esculentus), (Hibiscus sabdariffa), a source of bast fibers for making rough cordage. At least ten Mexican species are two of known to be used for this last purpose, including three species o? Abutil?n, The and Hibiscus pernambucensis. Robinsonella mirandae, Sida, three of Hampea, also Hampea last two species (and sometimes spp.) are of minor importance as a source of wood for construction and other uses. the most notable in The Malvaceae also provide a significant number of weeds, Mexico cristata, Herissantia crispa, Urocarpidium being Anoda (and elsewhere) l?mense, Kearnemalvastrum lacteum, and various species of the genera Sida, Malva, and Malvastrum. Malachra, found inMexico The largest genera of Malvaceae (and their respective number and Sida Pavonia of species) are Abutil?n Hibiscus (35), (29). These four (45), (37), on a are of the worldwide basis. Intermediate also the genera family largest genera Anoda sized genera in Mexico include Sphaeralcea (20), (23), Gossypium (14), the exception of (14). With (11), Bakeridesia (10), and Robinsonella Hampea

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3

MALVACEAE OF MEXICO

1988

in Mexico these genera are principally developed Gossypium, centers of has additional areas; diversity nearby Gossypium as well as amajor center inMexico. Africa, and Australia,

and one or more in South America,

The next smaller genera include Allowissadula (9), Kosteletzkya (8), Phymosia center of has another and Malvaviscas Kosteletzkya (6). (7), Bastardiastrum (8), are to of rest the this either of endemic in but the Africa, genera group development or are principally developed there. The remaining twenty-nine genera have Mexico from one to four species each. Of the fourteen genera with only a single species in Mexico, Modiola,

only

five

are monotypic so caution must

Fryxellia), of genus ing frequency distribution having only two species inMexico, are genera of only two species.

Billieturnera, genera (Anotea, Neobrittonia, concern in drawing conclusions be excercised size from these data. Similarly, of the ten genera and Meximalva) only two (Kearnemalvastrum

ENDEMISM One hundred and eighty-four (almost half of the total) are species of Malvaceae as endemic Of the fifty-five genera, three may be characterized endemic toMexico. more as en to Mexico and and nine "almost (Anotea, Bastardiastrum, Per?ptera) distribution and variability within the country demic" or with their principal Fryxellia, Hors (Malvaviscus, Robinsonella, Phymosia, Billieturnera, Allowissadula, Of these twelve Mexican and genera, fordia, Anoda, characteristically Meximalva). seven are moderate-sized genera of 5-25 species each, showing significant diversifica tion inMexico.

DIVERSITY as delineated and described by Using the phytogeographic provinces of Mexico, and the distribution of that diver Rzedowski (1978), the diversity of the Malvaceae sity have been estimated on the basis of the number of genera and number of species present in each province (table 1). As Rzedowski points out, certain of the bound aries of these provinces are somewhat arbitrary. Moreover, the occurrence of a given a species in given province is not always known with certainty, because of the vagaries of distribution and of collecting, especially for the less common species. Therefore, in tabulating the figures in table 1, some relatively arbitrary decisions had to be made. In spite of these qualifications, I believe that the overall picture of diversifi however, cation presented in table 1 is reasonably sound. Probably the greatest problem is for Province 14 (Depresi?n del Rio Balsas), where the figures given for encountered number of genera and species may be significantly underestimated. This possible underestimation results from the relative deficiency of collections from this province, as a result of the precipitous terrain and inaccessibility that characterize much of it, Hinton and Rzedowski (1972,1975). graphically described by are most richly developed 1 indicates that the Malvaceae Table in the lower elevations along both coasts, in the Costa Pac?fica and the Costa del Golfo prov inces. The diversity found in these provinces is markedly reduced in the adjacent of of and Peninsula de California Yucat?n, elevation, provinces Baja comparable are operative. Moreover, which indicates that factors other than elevation the

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4

SYSTEMATIC BOTANY MONOGRAPHS

1. Distribution

Table

of

the Malvaceae

(cf. Rzedowski

province

phytogeographic

in Mexico

by

1978).

Genera 1. California

VOLUME 25

Species

3 6

2.

Isla Guadalupe 3 5 3. Sierra Madre Occidental

4. Sierra Madre

11 10

Oriental

21

13

5. Serran?as Meridionales

61 23

6. Serran?as

30

Trans?stmicas

7. Baja California 8. Planicie Costera

9. Altiplanicie 10. Planicie

del NW

Costera de Tehuac?n

12. Costa

Pac?fica

del NE

48 20

9 9

13.

Islas Revillagigedos 14. Depresi?n del Balsas

33 118 8 12 4 5

4 6

15. Soconuso

del Golfo

17. Peninsula Total

58 21

18 54

11. Valle

16. Costa

12

14 41

de Yucat?n

for Mexico

26 88 36

18

372

55

in spite of being at markedly province Serran?as Meridionales, higher elevation, to that of the Costa del Golfo. shows diversity comparable The various provinces into which Mexico is divided are quite dissimilar in area. It is therefore probably unfair to compare the diversity of provinces as dissimilar in or the Serran?as size as, for example, and the Altiplanicie, the Valle de Tehuac?n Trans?stmicas and the Peninsula de Yucat?n. This problem may be circumvented by so that species diversity estimates of the areas of these provinces, using reasonable Such estimates are difficult to make and compared. per unit area can be computed in defining the boundaries of certain of the provinces. in view of the difficulties

SUBDIVISION OF THE FAMILY divides the family into five tribes. Four of these Contemporary understanding five tribes occur inMexico; the fifth tribe (the Decaschistieae Fryxell) occurs in the from India and Indochina to northern Australia. The distinctions among Old World, the tribes are indicated in table 2. The four tribes that occur inMexico are stated in detail below, following the family description. nomenclatural and bibliographical The key to genera attempts to be a synoptic key rather than an artificial key, such that major taxa (in this case tribes) are keyed out as units and so that juxtaposi tion of genera in the key is an indication of alliance. This objective has, of course, been only imperfectly achieved. The same objective has in most cases guided the are to construction of the keys found under the individual genera. species that In Mexico, and Gossypieae each have few genera? the tribes Hibisceae in the Hibisceae, and Abelmoschus and Gossypium, Ham Hibiscus, Kosteletzkya, in the Gossypieae. and Thespesia The tribe Malvavisceae in pea, Cienfuegosia, Mexico includes seven genera (Malvaviscus, Anotea, Pavonia, Peltaea, Lopimia, Malachra,

and Urena),

but the greatest

diversity

is in the tribe Malveae

with about

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5

MALVACEAE OF MEXICO

1988

Table

2. Distinctions

Malveae Fruit

schizocarp

Mericarps

per fruit

3 to many free,

Styles

3 to many

among

the tribes of the Malvaceae.

Malvavisceae

Hibisceae

Gossypieae

schizocarp

capsule

capsule

5 3 to 5

5

free,

Decaschistieae

10

free, 5

connate,

capsule 10

free,

10

stigmatic lobes 3-5 Ratio:

1

stigmas/

1

mericarps Apex of staminal column

Gossypol

glands

antheriferous

absent

5-toothed

absent

5-toothed

absent

5-toothed

5-toothed

present

absent

are so great in this group that it is useful forty genera. The taxonomic complexities to divide the tribe into generic alliances, (1970), by following Bates and Blanchard also using data on palynology but with modifications data slight using cytological and comparative morphology (table 3). to aperture number of the pollen grains (e.g., If we give special attention Hashmi 1986a, 1986b), we find that many of the 1970; S?nchez 1982; Christensen alliances low have aperture numbers (3-6), a few have intermedi generic relatively ate numbers two It seems and have very (8-35), high aperture numbers (50-200). to suggest that low aperture number is primitive and high aperture reasonable some anomalous number specialized There Christensen are, however, (cf. 1986b). groups that need special attention. Most striking is the case of the Sida alliance in values have been reported for different genera which both low and intermediate and in which the genus Sida itself has had aperture numbers reported from 3 to 35 be rewarding and (-90). A more detailed study of this alliance would doubtless could well lead to a better understanding of these taxa. Other anomalous groups are the Malvastrum alliance, with intermediate aperture numbers, except for the South which has 4-5 apertures, and the American M. spiciflorum (Hassler) Krapovickas, Anoda also with intermediate alliance, aperture numbers except for A. pentas chista, which has only 3 apertures. In view of our still incomplete and developing of these plants, understanding are presented as an aid in understanding these generic alliances only tentatively and in grouping the genera, but not as a formal taxonomic interpreta relationships tion.

SPECIALIZED CHARACTERS AND TERMINOLOGY Pubescence types. Pubescence types are important in characterizing members is not always precise or the Malvaceae. Pubescence however, terminology, on usage of to comment and it is therefore necessary generally agreed upon, on I trichome in the work. have relied Payne (1978) and present terminology

of

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SYSTEMATIC BOTANY MONOGRAPHS

6

Table

3. Generic

alliances

as represented

in the tribe Malveae chromosome

Base

VOLUME 25

by Mexican

number

Pollen

genera.

aperture

alliance:

Sphaeralcea

5 3-5

Sphaeralcea Tarasa

5

Abutil?n

4-5

10 8-9

Urocarpidium 10 3 Eremalche alliance:

Abutil?n Allowissadula

15 3

Bastardiastrum

1 3-6

Bastard?a

1 3-5

Hochreutinera Billieturnera

8 3 alliance: 7 3-4

Herissantia M odi?la

3-4

7 3-5

Wissadula

Herissantia

*

7,8

8 3-4

alliance:

Modiola

Fryxellia Fryxellia

9

4-6

7

5-10

alliance: 3

Batesimalva

alliance: 16

Batesimalva

3-4

15 3

Horsfordia Briquetia Dirhamphis Anoda

alliance:

15 Anoda

Bakeridesia

30-70

(3-)

13 18-35

Per?ptera alliance:

Bakeridesia

15 3

Sida alliance: Sida Meximalva

7,8

7 or

Rhynchosida Sidastrum Malvella Dendrosida

Gaya

(-90)

8 4-6

Krapovickasia

Allosidastrum

3-35

8 8-12

8

?

12-16

16 15-20 16 (probably)

3-4

7 3 (-4) 21

alliance: Gaya

Robinsonella Robinsonella

6 10-25

alliance: 16 3-5

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number

7

MALVACEAE OF MEXICO

1988

Table Kearnemalvastrum

alliance: 3-4 16

Kearnemalvastrum Malvastrum

3 continued

alliance: 6 8-30

Malvastrum

[except M.

in ]

alliance:

Phymosia

17 3-7

Phymosia Malacothamnus

17 4 (3-)

Neobrittonia Malva

4-5

spiciflorum

16

alliance: Malva

21

60-180

Alcea

21 -

100 or more

Lavatera Sidalcea

50-60

or

more

alliance: Sidalcea Callirho?

10 150-200 14,15

100-200

in choos (1986), and various botanical glossaries (1971), Wurdack I have also consulted that discuss trichomes terms; papers ing appropriate specifi 1960; Ramayya & Raja Shanmukha Rao cally in the Malvaceae (e.g., Ragonese Rao & Ramayya 1977; Hill 1982; Inamdar et al. 1983; 1976; Raja Shanmukha Lander 1984). of the Malvaceae Stellate hairs are characteristic (and families such as Euphor consulted

Roe

Solanaceae, biaceae, Cruciferae, Melastomataceae, among others). Hill (1982, p. 41, fig. 5) illustrates the types of stellate hairs to be found in the genus Malvastrum a six-rayed stellate and suggests phyletic pathways of specialization. He proposes amultipli hair as "unspecialized" and from this derives three lines of specialization: cation of ray number, a coalescence of rays, and a reduction of ray number. The distinctive "H-shaped hairs" (Hill, loc. cit., fig. 5e) are found in certain unrelated and coulteri, Malvastrum species (e.g., Sida ciliaris, Hibiscus coromandelianum) are characteristically and preferentially oriented symmetrical, 4-rayed, bilaterally (generally parallel with the stem axis). An additional is that borne type of stellate hair (not found in Malvastrum) above the epidermis on a stalk or stipe (e.g., Fig. 115). Such hairs are called stalked or stipitate hairs (sometimes called dendritic hairs, but perhaps incorrectly so); they a generally have a relatively large number of rays and thus may be considered of the line of ray proliferation. specialization Stipitate hairs are found in such genera as Bakeridesia, Malacothamnus, Tarasa, and Phymosia, Sphaeralcea, Wissadula. Coalescence of the rays of stellate hairs is carried to the extreme form of peltate scales in a few Mexican such as Malvella Sphaeralcea sagittifolia, lepto species, a and in certain other and (on minute samaricarpa, phylla, scale) Robinsonella species in other parts of the world, often in arid or saline habitats (cf. Lander 1984). Such hairs are variously termed peltate, lepidote, or squamose (Fig. 74). further

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SYSTEMATIC BOTANY MONOGRAPHS

8

VOLUME 25

or A number of species of Malvaceae in various genera have viscid pubescence the various types of glandular hairs. No attempt will be made here to distinguish glandular hairs (Inamdar et al. 1983) but only their presence or absence. Glandular hairs are unknown in many genera of Malvaceae but are variously present among the species in other genera. They may be generally distributed throughout the plant or restricted to certain organs. In extremely viscid species (e.g., Bastard?a viscosa, the plants are malodorous, Bastardiastrum Abutil?n mucronatum) wissaduloides, are sticky to handle (when fresh), and often discolor the papers in which they are pressed.

in the Malvaceae, often in combination with Simple hairs also are common stellate hairs or glandular hairs (or sometimes with both). Pubescence of simple hairs (like stellate hairs) may be soft, bristly, scabrid, or pungent (urticating). in some species of Hibiscus Pungent hairs grade into prickles or aculeae, especially sect. Furcaria, such as H. uncinellus (Fig. 55). a In number of genera of Malvaceae (as well as in other families) specialized are in narrow lines decurrent from the stipules hairs found on the stems distributed and also on the adaxial sides of the petioles and pedicels. These hairs are generally on simple, short, and recurved. They are more or less well developed, depending the species, hair types.

and their presence

seems

to be

independent

of the presence

of other

The term pedicel refers to the stalk that bears an individual Pedicel/peduncle. ismore generalized and less precise; it refers to the stalk flower. The term peduncle that bears a group of flowers, but may refer to one that bears a single flower. In some Malvaceae the distinction between peduncle and pedicel is reasonably clear in Phymosia used. For example, cut, and the terms are appropriately umbellata, the axillary peduncle divides distally where flowers are borne in axillary umbels, into three or four pedicels, each bearing a flower. Many Malvaceae, however, have the flowers borne singly in the leaf axils, in what may be interpreted as a uniflorate on an often articulated stalk. To call this structure a peduncle below inflorescence, a the articulation is considered and and pedicel above the articulation pedantic, such usage is not followed here. Rather, for plants with solitary flowers, the two terms may be used interchangeably, although pedicel is perhaps to be preferred as less subject to misinterpretation. Involucel. The terms involucel and involucre are sometimes confused, but gen that subtend either an individual flower or erally refer to bracts or similar elements an inflorescence, In the present work the term involucel is used for the respectively. that immediately subtend the calyx, although in the literature specialized elements em of the Malvaceae, the terms involucel, involucre, or epicalyx are variously or a structure. For bracts that subtend group of ployed for this specialized elements flowers (or inflorescence), for which the term involucre might arguably be used, I Such floral bracts in some species prefer to me floral bract(s) to avoid confusion. a occur case at subtend the base of the pedicel, not at its single flower, in which they apex.

or absence of an involucel generally presence among the distinguishes but two in of these (Malvella and Callirhoe), the presence or absence on the genera, involucel distinguishes among the species within the genus. The number of elements in the involucel is of taxonomic significance. In many the number of is two elements three in reduction genera (or rarely uniformly by The

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1988

9

MALVACEAE OF MEXICO

are more numerous in other genera, often 8-10, the elements whereas Malva), as as is important in characterizing elements 20. The of number many occasionally the species in such genera, as is the degree to which they may be connate or distinct. the is extreme Where the degree of connation pernambucensis), (e.g., Hibiscus involucel takes the form of a cup with a certain number of lobes or teeth. In some species the involucel more or less conceals the calyx, e.g., in Lopimia malacophylla. bracts is highly variable. The bracts may be The form of the involucellar the entire enclosing and concealing broadly foliar (as in Gossypium gossypioides), the bracts may be inconspicuous, bud prior to anthesis. At the opposite extreme, a aridum, Hibiscus denudatus), (e.g., Gossypium triangular or subulate elements are or subulate lanceolate fraction of the length of the calyx. Commonly they or spreading. structures to the calyx, appressed In a few cases the subequal or peltate Hibiscus Peltaea involucellar bracts are stipitate uncinellus, (e.g., trinervis,

Phymosia

umbellata);

rarely,

they are deciduous

(Hampea

tomentosa,

Phymosia rosea). In most Malvaceae is whorled the involucel, when present, (e.g., Fig. 50); a in Malvella few rovirosae, however, species (e.g., Hampea leprosa) the bracts of the involucel are irregularly inserted (Fig. 74) in such a way that the whorled seems to have been derived from a spiral disposition. The more or less arrangement relictual, occurs only rarely in the family. spiral arrangement, presumably Nectaries. Many angiosperms have nectaries within the flower, generally at the base of the calyx internally, which provide a food reward for flower visitors that serve the plant as pollinators. of the Malvaceae Certain members (and of a few other families) also have nectaries located outside of the flower, and the distribu tion and structure of these extra-floral nectaries have taxonomic value. The ecologi cal role of such nectaries is less clear. These extra-floral nectaries are of three types: and foliar, involucellar, calyx nectaries. All exude a sugary nectar. Foliar nectaries occur on the abaxial (lower) surface of the leaves, singly on the midrib or several on the principal nerves. In some cases (e.g., Thespesiapopulnea), the foliar nectary is simply an aggregation of secretory cells that are exposed along the surface of the vein. More commonly, the secretory tissue ismore or less sunken and is surrounded by raised protective the tissue, the whole structure constituting are in nectaries found of the tribe Foliar many genera nectary (Figs. 47, 55). in some genera of the tribe Hibisceae, and in one genus (Urena) of the Gossypieae, tribe Malvavisceae. Foliar nectaries are absent in the large tribe Malveae. Even in are that of foliar nectaries characterized the genera presence by (e.g., Gossypium), they may be absent in particular species (e.g., G. gossypioides). Involucellar nectaries are of more restricted occurrence. As the name implies, at the distal end of associated with the involucel, positioned they are immediately the pedicel below of the the involucellar insertion bracts. The immediately involucellar nectaries are always trimerous and are differentiated into a more or less raised margin and a sunken secretory zone. Involucellar nectaries are found only in the tribe Gossypieae, which inMexico includes the genera Gossypium, Cienfuego and these nectaries may be suppressed in particular sia, Hampea, Thespesia. Again, species within these genera. Calyx nectaries are found on the midribs of the calyx lobes and are thus always is con pentamerous. They are usually bordered and prominent. Their occurrence fined to species of Hibiscus sect. Furcaria and Hibiscus sect. Azanzae; in the former

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SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

in certain species (e.g., H. costatus). InH. calyphyllus, section they are suppressed are located at the tips of the calyx lobes and may be homologous with nectaries those found on the midribs of the calyx lobes. of gossypol glands is confined to and charac Gossypol glands. The occurrence structures are lysigenous of the tribe Gossypieae. These teristic of the members are most of the which scattered parts of the plant. above-ground throughout glands, are suberized, and contain a mixture of pig They are approximately spherical, is the characteristic of which gossypol ments, (a sesquiterpene-like compound) constituent. Typically these glands occur beneath the epidermis and appear as black In leaf tissue they are best seen by transmitted (or sometimes translucent) punctae. is helpful. In the leaf blade the gossypol glands light, and low power magnification are usually found one per areola among the smallest veins. In other plant parts, the calyx and the carpel wall (e.g., Fig. 41), these glands are often larger especially and more prominent and easily observed. is particularly fruit morphology Fruit morphology. In the Malvaceae, important a certain amount of special in distinguishing among the genera and consequently Fruits may be either capsules (tribes Hibisceae and ized terminology has developed. on or schizocarps and Malvavisceae), (tribes Malveae depending Gossypieae) is principally loculicidal or septicidal. The morphology of whether the dehiscence so the following the capsules is relatively commentary pertains straightforward, primarily to the schizocarps. of as few as three to as fruits of Malvaceae may be composed Schizocarpic many as forty carpels. In many cases these matured separate, so carpels ultimately is the dispersal unit or disseminule; in that the carpel wall and contained seed(s) In any event it is convenient is only partially accomplished. others, this separation to use the term mericarp for this segment of the fruit; it is considered more precise in a than the term carpel, which is roughly the same as pistil (or its equivalent a are not generally fruit) and thus includes the style and stigma, which compound part of the mature fruit. in terms of dorsal and ventral aspects. The ventral The mericarp is described is (or was) attached to the side is the adaxial side, the region where the mericarp central column. The dorsal or abaxial side refers to the "outside" of the fruit. In to refer to the lateral walls of the mericarp, in contrast certain cases it is appropriate are those walls that were to the dorsal wall. The lateral walls of the mericarp in in the developing fruit. immediate contact with those of adjacent mericarps the mericarp In a number of genera of the tribe Malveae, is divided into an a a lower cell in upper and variety of ways (e.g., Figs. 37, 58). The lower cell, and its lateral walls are often reticulately is one-seeded and ind?hiscent, generally, or less dehiscent, more and its lateral walls ornamented. The upper cell is generally are generally smooth. The upper cell often contains one or more seeds (commonly two), or the ovules of the upper cell may be abortive. are separated by a slight constric In these genera the two cells of the mericarp tion of the lateral walls, or they are separated by a partial or complete partition The term endoglossum between them called an endoglossum. applies to a variety of structures in different genera that are probably not homologous. When Fries (1908) used its presence or absence to define genera, its diversity and distribution were not the term was later coined by Hochreutiner yet well known. When (1920a), the structure was still not well understood. is a protrusion into the The endoglossum

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1988

MALVACEAE OF MEXICO

11

interior of the mericarp that (in different genera or species) may originate from the it dorsal or the ventral wall; itmay be a complete partition or a vestigial outgrowth; may be a single or a double structure; in the genus Gaya it is not a partition but a more complex structure that encircles the solitary seed (Fig. 40); finally, in some so that the one-seeded mericarps are in fact one genera the upper cell is suppressed that the uni-ovulate celled, but a vestigial endoglossum remains, suggesting carpel was derived was present ancestors in which an endoglossum from pluri-ovulate (e.g., Allosidastrum, glossum morphology

A comparative Malvastrum). would probably yield valuable

developmental insights.

study of endo

VERNACULAR NAMES names are given wherever Vernacular available. These have been taken princi and from labels, primarily from MEXU, pally from two sources: from herbarium the recent compilation of Martinez from these two (1979). The names extracted sources overlap in large measure, but inmany cases names from one source are not found in the other, so that combining both sources provides a more comprehensive list. a vernacular name with a particular species and its correct botanical Equating a difficult name is sometimes task. A name taken from a herbarium label, by one can circumvent is a name tied to a voucher specimen. Therefore, definition, of the plant or subsequent nomenclatural problems (such as misidentification of the specimen changes necessitated by improved taxonomy) by direct consultation itself. Those names that I have taken directly from herbarium labels are tied di as interpreted treatment. The majority in the present of rectly to the species, Martinez's from herbarium labels. Unfortu names, of course, are taken ultimately so nately he does not cite the source of individual names in an unequivocal manner, names. In many that most of the names that he presents are basically unvouchered because the species in question cases, there is little or no problem of identification, is clearly defined and not to sinensis), and nomenclatural of identity others, questions Martinez generally provides), a be considered confidence

rosa be confused with anything else (e.g., Hibiscus like doubtful synonyms are not at issue. In questions can be resolved by geographical information (which so that "Hibiscus can with tubiflorus (Yucat?n)" to Hibiscus Still others can be reference poeppigii. resolved by consideration of the brief plant description that Martinez provides. There are cases, however, that are almost unresolvable. For example, Martinez names to Malvastrum includes vernacular A. referring scoparium (L'H?ritier) a name that applies to a South American is Gray, plant (Hill 1982), but which to Malva in reference the basis for the name presumably scoparia Jacquin, ex Small. How Malvastrum in Lamarck) Britton corchorifolium (Desrousseaux and M. ameri ever, in the past other species of Malvastrum (M. coromandelianum and specimens have canum) have been frequently confused with M. corchorifolium, been misidentified because of this confusion. Hence, it is difficult to be certain asMalvastrum the species referred to by Martinez concerning scoparium. Wherever to determine I have used plausible the species to which Marti possible reasoning names refer; where doubts remain concerning nez's vernacular species identity, I have omitted the names.

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SYSTEMATIC BOTANY MONOGRAPHS

12

VOLUME 25

SPECIMEN CITATIONS are cited by collector and collector's number. They are grouped by Specimens are not more precisely but localities indicated nor are the herbaria of deposit states, listed to conserve of space. By way exception, type specimens are cited fully. The is indicated by the standard acronyms given herbarium of deposit of type specimens et al. (1981), with the additional inHolmgren indication "pf" being used to refer to the author's herbarium, in Station, Texas. Generally College kept speaking, speci mens lacking collection numbers (s.n.) are omitted, unless they are types or unless rare species or geographically collections. they represent noteworthy Specimens collected by three or more collectors are generally cited either by the first collector " only or with an "et al. appended (i.e., either "Ortega 2T or "Ortega et al. 2T). No to has been made different collectors with the same surname. attempt distinguish names as such and each represent more than a single collec Smith Gonz?lez Thus, tor among collections cited here. For Hispanic surnames, a citation problem exists, because a given individual's name may be cited on a herbarium label in various Gonz?lez O., Gonz?lez Ortega, Gonz?lez-Ortega, ways, for example as: Gonz?lez, or even (incorrectly) as Ortega. Certain collectors are not always consistent in how an own names. not Where I do know individual and there cite their collector is they correct such inconsistency, I am left with some uncertainty the citation concerning of the person's name. It has not been possible to be consistent in this matter. I have on I find and cited what herbarium when there the have been label, generally several variants, have tried to use the fuller citation (in the preceding example: to those individuals whose names I have I can only apologize "Gonz?lez-Ortega"). cited incorrectly. For certain of the more active or historically more of important collectors an are resources are to Mexican that there aid about plants, bibliographical learning their itineraries and collecting localities and often biographical details about their careers. Some of the principal are cited in such references lives and botanical 1. Appendix Specimens (especially important herbaria, types) cited from certain historically G-DC B-W P-HBK LINN, (de Candolle), (Humboldt & principally (Willdenow), P-LA and S have P-JU been con (Lamarck), (Linnaeus), (Jussieu), Bonpland), and widely distributed photographs sulted in microfiche. The well-known of types from European herbaria of the Field Museum, & Plowman (Grim? Chicago 1986), are cited by negative number as: F-23734, for example, the type of Abelmoschus achanioides Turczaninow.

TAXONOMY Malvaceae

Jussieu, Gen.

pl. 271.

1789.?Type:

Malva

Linnaeus.

some roots fibrous or woody, shrubs, or trees, often stellate-pubescent; Herbs, or stems erect times fleshy in perennial sometimes herbs; repent. procumbent, Leaves alternate, stipulate (the stipules rarely suppressed), ovate, lanceolate, some with hairs that are stellate, simple, sometimes times lobed or dissected, prickly, sometimes glandular, or rarely lepidote. Flowers solitary or fasciculate in the axils

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1988

13

MALVACEAE OF MEXICO

sometimes in inflorescences of the leaves or grouped (usually racemes or panicles, or or involucel or umbels absent; present heads); spikes scorpioid cymes, rarely or truncate, 5-lobed, 5, distinct, 5-toothed, petals 5-parted; calyx gamosepalous, usually clawed, adnate to the staminal column at base; androecium monadelphous; to 5); pollen spheroidal, numerous anthers reniform, echinate; (rarely reduced or truncate, 1-40; stigmas styles capitate, 3-40-carpelled; superior, gynoecium a berry; seeds reniform or or capsular, sometimes Fruit schizocarpic decurrent. or glabrous, arillate. Base chromosome rarely (in Hampea) 8,9,10,11,13,14,16,17,18,19,20,21,25,28. The family includes more than 100 genera and perhaps 2000 species, principally from tropical and subtropical regions but with a few genera from temperate regions.

turbinate, pubescent numbers: x = 5,6,7,

References: Medikus (1787), de Candolle (1824),Garcke (1849,1896), Alefeld (1862), Bentham (1862), Gray (1887), Baker (1890-1894), Kuntze (1891), Rolfs (1892), Urbina (1897), Hochreutiner (1920a), Standley (1923), Webber (1934), Edlin

and Hess

(1935), Record

and Peebles

(1937), Kearney

(1942),

and

Standley

Steyermark (1949), Kearney (1951a), Martinez (1955), Wiggins (1964, 1980), Robyns (1966), Hutchinson (1967), Bates (1968a), Fryxell (1968a), Bates and Blanchard

of Tribal

Synopsis

Bot.

Alefeld, naeus.

Gossypieae

and Johnston

(1970), Correll

(1970). of the Malvaceae

Classification 19: 301.

(Berlin)

Zeitung

1861

inMexico

(as "Gossypiidae").?Type:

Lin

Gossypium

References: Alefeld (1861), Ulbrich (1914),Hutchinson (1947), Fryxell (1968a, 1979b). Included Hibisceae

Mexican

genera:

Reichenbach,

Gossypium,

Cienfuegosia,

reg. Consp. 1862.

veg.

202.

Hampea,

1828.?Type:

Thespesia.

Hibiscus

as sub

Linnaeus.?Recognized

family by Alefeld,

?sterr. Alefeld, K. Presl.

Kosteletzkyeae Kosteletzkya References: Included Malvavisceae

Zeitschr.

(1951a), Hutchinson Kearney Mexican genera: Abelmoschus,

K. Presl, Bentham

Reliq. Haenk. & Hooker,

(1967), Hibiscus,

12:

147.

1862

(as

(1968a,

Fryxell

"Kosteletzkydae").?Type:

1975a).

Kosteletzkya.

2:135.1835

Malvaviscus Fabricius. (as "Malvaviscaceae").?Type: 1: 205. 1862; Bentham, Proc. Linn. Soc. London 6: 99. as subfamily Urena Linnaeus.?Recognized Bot. 1862.?Type: ?sterr. by Gray ex Alefeld, Zeitschr. 12: 147. 1862.

Ureneae

Pavonieae References: Included Malveae

Bot.

Alefeld,

?sterr.

Bot.

Gen.

pi.

Zeitschr.

12: 147.

1862.?Type:

Pavonia

(1978). Kearney (1967), van Heel (1951a), Hutchinson Mexican Malvaviscus, Anotea, Malachra, genera: Lopimia,

St.-Hilaire,

FI. Bras.

mer.

1: 173. 1827.?Type:

Malva

Cavanilles.

Pavonia,

Peltaea,

Linnaeus.?Recognized

Urena.

as subfamily

Alefeld (1862) and as subtribe by Kearney (1951a). Linnaeus. Reichenbach, reg. veg. 201. 1828.?Type: Malopeae Consp. Malope Sideae K. Presl, Reliq. Haenk. 2: 104. 1835.?Type: Sida Linnaeus. Abutilieae Bot. Zeitschr. 12: 147. 1862 (as "Abutilidae").?Type: Miller. Abutil?n ?sterr. Alefeld, Alceae Alefeld, 12: 146. 1862 (as "Alceidae").?Type: Bot. Zeitschr. Alcea Linnaeus. ?sterr. Malvastreae

Bot. Zeitschr. ?sterr. Alefeld, A. Gray. Bot. Zeitschr. ?sterr. Alefeld, Plagiantheae thus J. Forster & G. Forster.

12: 147. 1862 (as "Malvastridae").?Type: 12: 146.

1862

(as "Plagianthidae").?Type:

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Malvastrum

Plagian

by

SYSTEMATIC BOTANY MONOGRAPHS

14

References:

Baker

(1951a),

Kearney

(1890-1894),

Hutchinson

VOLUME 25

Bates

(1967),

(1968a),

Bates

and

Blanchard (1970). Included Billieturnera,

Mexican

Abutil?n

genera:

Hochreutinera,

alliance

Wissadula).?Anoda alliance (Batesimalva,

(Bakeridesia).?Batesimalva alliance alliance (Fryxellia).?Gaya trum alliance (Kearnemalvastrum).?Malva

(Abutil?n, alliance

Allowissadula, (Anoda,

Bastard?a,

Bastardiastrum, alliance

Per?ptera).?Bakeridesia

Briquetia,

Dirhamphis, Horsfordia).?Fryxellia alliance (Herissantia).?Kearnemalvas alliance Lavatera, Malva).?Malvastrum alliance Neobrittonia, (Malacothamnus, Phy alliance (Allosidastrum, Dendrosida, Krapovick

(Gaya).?Herissantia alliance (Alcea,

alliance (Malvastrum).?Modiola (Modiola).?Phymosia alliance (Robinsonella).?Sida mosia).?Robinsonella asia, Malvella, Sida, Sidastrum).?Sidalcea Meximalva, Rhynchosida, Sphaeralcea alliance Tarasa, Urocarpidium). (Eremalche, Sphaeralcea,

alliance

(Callirho?,

Sidalcea).?

are presented The relationships in among the genera as currently understood capsule form in the preceding synopsis and in table 3. In the following treatment, the genera are treated in alphabetical however, sequence for ease of use. note: Achille Richard published a number of names for Cuban Bibliographical on of Ram?n de la Sagra in a work that was published collections in plants based a a both French and Spanish edition (cf. Stafleu & Cowan 1983, p. 763, no. 9150). differs in the two editions, and since "it is unclear which Since the pagination loc. cit.), both references are cited for the edition has priority" (Stafleu & Cowan, names concerned. In the present work, such citations are found for the names (or synonyms)

of species

no's.

Key 1. Fruits

staminal

capsular;

and 313.

128,163,180,189,238,

to the Genera column

inMexico

of Malvaceae

surmounted

by 5 apical

teeth;

carpels

involucel

3-5;

present

(rarely suppressed). 2. Stigmas usually petals, calyx, 3. Perennial 3.

in decurrent (or if capitate, carpels 3); carpels 3-5; gossypol glands present or herbage; bractlets of involucel 3-9; tribe Gossypieae. ca. 9 herbs or subshrubs less than 1 m tall; foliar nectaries involucel absent;

corolla yellow; carpels (in ours) 3; seeds pubescent. or trees more than 1m tall ;abaxial foliar nectaries usually

parted; Shrubs

Cienfuegosia. ; involucel usually present or glabrous. ;corolla white or cream, yellow, or rose ;carpels 3-5 ;seeds pubescent in often dioecious (or large shrubs), Hampea rovirosae); (perfect-flowered

3-parted 4. Trees flowers

5 cm or less in diameter; or

4. Trees

Gossypium subglabrous 5. Capsule 2.

shrubs,

petals

perfect-flowered;

pallid, flowers

seeds glabrous, arillate. reflexed; Hampea. 5 cm or more in diameter in (except or

to bright thurberi); yellow petals pale to pubescent, exarillate. ind?hiscent. (in ours) oblate, Thespesia. to ovoid, dehiscent. orbicular Gossypium.

5. Capsule Stigmas carpels 5; gossypol glands absent; bractlets capitate; tribe Hibisceae. more; 6. Capsule seeds solitary; plants often depressed, 5-winged; often

fruits),

in wet

ovoid or elongate types of vestiture,

6. Capsule various 7. Calyx fruits herbs

asymmetrical,

absent.

not winged; (fusiform), from wet or dry habitats; spathaceous,

splitting

usually elongate (fusiform); or subshrubs ; foliar nectaries

7. Calyx symmetrical, nous or cultivated, 1. Fruits

foliar nectaries

habitats;

pentamerous, mostly

plants absent.

rose,

of

not

involucel

scabrid

seeds

reflexed;

4 to usually

or hispid

8 or

(especially

Kosteletzkya. seeds several foliar

laterally

per locule; plants with nectaries sometimes present.

at anthesis,

sometimes introduced, Abelmoschus.

deciduous naturalized,

in fruit; robust

fruits more or less ovoid; plants persistent; indige or absent. foliar nectaries obscure, prominent, Hibiscus.

shrubby;

but (rarely pseudocapsular, schizocarpic at apex; carpels apical teeth or filamentiferous absent.

then 3-40,

involucel

staminal column with 5 absent); often more than 5; involucel present or

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1988

15

MALVACEAE OF MEXICO

column surmounted 5, dry, viscid, or fleshy, 1-seeded; by 5 apical teeth; mericarps or as mericarps); foliar nectaries 10 (twice as many present (in Urena) styles and stigmas tribe Malvavisceae. involucel usually present, absent; 5-20-parted; 9. Leaves with prominent abaxial nectaries; bractlets of involucel 5, alternate with calyx lavender. Urena. lobes; mericarps glochidiate spines; corolla (in ours) with numerous Staminal

9. Leaves

bractlets

nectaries;

lacking

absent; Malachra) various colors.

mericarps

with

of

than usually more barbed spines or without

involucel

3 retrorsely

5,

sometimes

spines;

(in of

corolla

10. Flowers

inflorescences into capituliform and subtended by specialized, aggregated or less ovate bracts; mericarps smooth, dry, spineless. or setose; 11. Specialized floral bracts often basally dissected involucel absent or (in

more

or lavender, incon bracts; corolla white, radiata) of subulate yellow, or showy. Malachra. 11. Specialized floral bracts bracts of serrate, cordate-ovate, stellate-pubescent; corolla involucel Peltaea. (in ours) yellow and showy. (in ours) bifurcate; or sometimes or in umbellate, 10. Flowers headlike racemose, solitary paniculate, Malachra

spicuous

involucellate but not subtended inflorescences, bracts; mericarps by specialized fleshy or dry, 3-spined or smooth. or 3-spined; or 12. Mericarps reticulate, dry and smooth, winged, plants herbaceous staminal column usually included (sometimes Pavonia. shrubby; exserted). 12. Mericarps viscid or fleshy, umn usually exserted.

unornamented;

plants

usually

shrubby;

staminal

col

fruits blackish, viscid. (in ours) 15 or more; Lopimia. fewer than 15; fruits fleshy. 14. Petals with basal auricle, usually bright red (sometimes staminal white); column 1-2 times the length of the corolla; mature fruits (at anthesis) Malvaviscus. usually bright red. 14. Petals without staminal column 3-4 auricle, (at anthesis) cream-yellow;

13. Bractlets

of involucel

13. Bractlets

of involucel

the length of the corolla; mature fruits blue-black. Anotea. at summit; mericarps 3-40, dry, 1-5 (-7)-seeded; styles or absent, as mericarps; foliar nectaries involucel if absent; present

times Staminal and

column

stigmas

as many

present usually 15. Mericarps circle

filamentiferous

(2-)

in Alcea); tribe Malveae. (except 6-7-parted or reniform in a indurate, 1-seeded, (with ventral notch), fruit; stigmas capitate column, forming a depressed (oblate) or less palmately ovate or rotund and more lobed or dissected;

3-parted

horseshoe-shaped a central surrounding

or decurrent; leaves involucel usually present. 16. Staminal

column

involucel

absent; column

16. Staminal

divided into an inner and outer apically raceme. inflorescence usually an elongated as above), the simple (not differentiated

series

of

filaments;

Sidalcea.

all arising filaments from near the apex of the column; involucel present; inflorescence various. equally 17. Bractlets of involucel flowers connate; (in ours) 6-7, partially (some showy in spiciform times double) introduced inflorescence; (in ours) 20-40; carpels Alcea. garden cultigens. 17. Bractlets of involucel 3; flowers showy or inconspicuous, disposed; variously carpels 6-35. 18. Bractlets

of involucel connate; partially large shrubs with showy axillary 6-10. Lavatera. flowers; mericarps to filiform, 18. Bractlets of involucel lanceolate distinct; herbs or small shrubs; flowers various; mericarps 7-35. or yellow; erect herbs or small shrubs; mericarps 8-13. mature mericarps climax leaves 3-7-lobed; black 1500-3100 m. Kearnemalvastrum. smooth; ish, glabrous, 20. Corolla climax leaves ovate or lanceolate, meri unlobed; yellow; the walls often ridged or carps reddish brown, more or less setose, Malvastrum. armed; sea level to 2600 m.

19. Corolla 20.

white

Corolla

19. Corolla cending,

white;

rose, purplish, burgundy, or erect herbs; mericarps

or pale

lavender;

procumbent,

7-35.

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as

16

SYSTEMATIC BOTANY MONOGRAPHS

21.

VOLUME 25

Procumbent

herbs (in ours) with large taproots; corolla perennial red or burgundy, 3-5 cm in diameter; often exceed pedicels 14-22; beaked; Coahuila, ing the leaves; mericarps obtusely Nuevo Le?n. Callirho?. deep

or erect annual or perennial 21. Ascending herbs; corolla pale laven der or purplish, less than 1.5 cm in diameter usually larger (except in Eremalche and Malva shorter rotundifolia sylvestris); pedicels than the leaves (except in Eremalche rotundifolia); mericarps 35, not beaked. 22. Flowers in scorpioid and fruits subsessile cymes; mericarps 14; usually above 2200 m. 23. Mericarps

7 7

with irregular transverse ind?hiscent, glabrous, rounded calyx hairs stellate (not apically;

dorsally,

ridges

stipitate). 23. Mericarps dehiscent

Urocarpidium. hairs, (and sometimes stipitate) smooth or reticu ventrally (and later dorsally), late laterally, more or less acute apically; calyx often with Tarasa. hairs (the stipe dark-pigmented), stipitate-stellate

22.

Flowers

24.

and not

mose, m.

with

fruits scorpioid;

Stigmas fornia,

manifestly

8-35;

usually

on style; mericarps to erect, introduced,

decurrent unilaterally or ascending decumbent Malva.

capitate; Sonora.

mericarps Eremalche.

or race axillary 2000 below

pedicellate,

mericarps

zone

Stigmatic

8-15; plants often weedy. 24.

stellate

12-35;

erect; Baja Cali

plants

not forming a depressed 15. Mericarps fruit made up of horseshoe-shaped otherwise, seg or dilated not decurrent; ments, apically, l-5(-7)-seeded; usually capitate stigmas leaves various; involucel present or absent. rounded and more or less inflated. thin-walled, or large shrubs; mericarps sometimes around the 1-seeded, collapsed an involucel, to form a samara; flowers seed at maturity small, without aggre Robinsonella. axillary fascicles or panicles. gated into showy multiflorate to large shrubs; mericarps 26. Low herbs inflated 1-4-seeded, always (though

25. Mericarps 26. Trees

if numerous); flowers large or small, with in few-flowered inflorescences, tary, axillary racemes. thin

or without or

an

soli involucel, or in panicles

sometimes

the seed encircled 1-seeded, by a pectinate endoglossum; absent; herbs or low subshrubs with solitary flowers. Gaya. one- to several-seeded, 27. Mericarps without involucel present endoglossum; or absent; perennial herbs to large shrubs, the flowers solitary or in various types of inflorescences.

27. Mericarps involucel

28.

in panicles, Involucel of 3 bractlets; flowers 7-40; small to bels, rarely solitary; mericarps or 29. Bracts of involucel (linear,) spatulate, flowers manifestly phyllous; pedicellate,

or axillary racemes, shrubs. large

um

sometimes ovate, gamo large and in few-flowered or small and in panicles; mericarps

axillary umbels (rarely solitary) 10-40 ; seeds 2-3 per carpel. Phymosia. 29. Bracts of involucel linear, distinct; flowers aggregated densely cothamnus foliosus); 28.

nia, Involucel herbs

absent; or shrubs.

flowers

into racemiform

usually inflorescences

mericarps Sonora.

7-14;

usually

solitary;

seeds

(except

solitary;

mericarps

often

subsessile,

inMala

Baja Califor Malacothamnus.

8-14;

perennial

Shrubs 2-3 m tall; mericarps 8-12, with 2 long (6-14 mm) spines at base of each; corolla purplish, 3-4 cm in diameter. Neobrittonia. or shrubs less than 1.5 m tall; mericarps 30. Herbs 10-14, spineless; corolla white, 1-1.5 cm in diameter. Herissantia. 30.

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1988

MALVACEAE OF MEXICO

17

or smooth-walled, if thin-walled 25. Mericarps and indurate, often reticulate-walled not inflated, often apically acute or otherwise differentiated, 31. Mericarps twice as long as the column, rounded but usually acute or sometimes as a result the spine single, becoming double only at maturity apically, apiculate more or less elliptic of dehiscence, in cross section and often dorsally keeled in Bakeridesia); involucel absent. (winged 32. Mericarps 5-8,1-seeded. more 33. Leaves than 2 cm long; cordate-ovate, 5-8. erect, 0.5-1.5 (-3) m tall; mericarps

subulate; stipules Bastard?a.

plants

cm long; stipules broadly oblanceolate; 0.5-1.5 cun?ate, plants low and branching, less than 0.1 m tall; mericarps 5; Nuevo generally Billieturnera. Le?n, Tamaulipas.

33. Leaves

32. Mericarps 3-30,2-7-seeded. not differentiated 34. Mericarps

into upper and lower cells. 35. Uppermost leaves below the inflorescence) sessile (immediately and amplexicaul; 5-seeded with small endoglossum be mericarps Hochreutinera. seed, blackish at maturity. an endoglos without 2-7-seeded, petiolate; mericarps at maturity. sum, usually tan or brown (sometimes blackish) or 36. Leaves entire with toward so, nearly tendency es venation; pubescence camptodrome usually ferrugineous, low lowermost

35. All

leaves

in meristems, the stellate hairs often pecially stipitate; often blackish, often with erose wings on rounded, mericarps dorsal sometimes often with margin (wings suppressed), rather than dorsal dehiscence. Bakeridesia. ventro-apical 36. Leaves

to manifestly serrate, at the margins; pubescence acute to acuminate mericarps

subentire

terminating

the principal veins fer seldom various, or spinescent (some with dorsal usually

rugineous; times rounded),

without wings, Abutil?n. differentiated into a 1-seeded by a constriction

dehiscence. 34. Mericarps and a 2-seeded

upper

cell

(the seeds

tive). 37. Mericarps 5-11; constriction of mericarps sometimes

of the upper

of mericarps separated

lower

cell sometimes

cell abor

cells obscure; a double by Abutil?n sect. Anasida.

externally internally

endoglossum. 37. Mericarps 3-6;

constriction of mericarps manifested; externally present endoglossum (and simple) or absent. 38. Calyx less than half the length of the mature fruit, not promi

to glabrate; mericarps glandular-puberulent seeds (especially the lowermost) often Wissadula. pubescent. 38. Calyx the mature fruit, more or less approximately equaling nently

nerved;

endoglossum

absent;

nerved; mericarps prominently pubescent; ent or absent; seeds glabrous or pubescent. 39. Mericarps 5, apically rounded, acute, glossum

present

orange

corolla

pale

pres

or apiculate; endo corolla yellow sub manifest,

(sometimes absent); staminal column white);

(rarely to filaments; 39. Mericarps 3 (-4), equal

endoglossum

seeds

glabrous.

Allowissadula.

rounded; absent; apically endoglossum to almost white; lavender staminal column much shorter than the filaments; the lower

rudimentary, most seed often

on the hilum. Bastardiastrum. pubescent 31. Mericarps to twice as long as the column, approximately rounded, equaling in cross section with differentiated dorsal acute, or 2-spined trigonal apically, and lateral walls; involucel present or absent. 40. Involucel present absent inMalvella). (sometimes or procumbent 41. Prostrate herbs.

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SYSTEMATIC BOTANY MONOGRAPHS

42. Leaves dissected; divided

or less palmately more orbicular, symmetrical, to brick-red; mericarps corolla orange setose, and 1 seed into upper lower cells (with

Modiola. endoglossum. Leaves flabellate asymmetrical,

42.

smooth-walled, Malvella.

ish; mericarps 41. Erect

VOLUME 25

to triangular; unornamented,

corolla

or

parted apiculate, each)

pale

1-seeded,

by

yellow without

endoglossum. herbs or subshrubs.

into reticulate 43. Mericarps lower portion and usually differentiated at least apically smooth 1-3-seeded, dehiscent; upper portion, not scorpioid; inflorescences usually below 2000 (-2500) m. Sphaeralcea. or as above, not differentiated 43. Mericarps ind?hiscent 1-seeded, in scorpioid flowers cymes; ventrally; tardily dehiscent usually 2200 m.

above 40.

Involucel

asa.

Tar

absent.

44. Carpels in Batesimalva 3-ovuled (but 1-seeded by abortion of 2 ovules and sometimes inHorsfordia). the inflorescence and am 45. Leaves below sessile immediately

a leafless

inflorescence 46. Upper

45.

but

spike or panicle.

not

cell

upper

present;

yellow; plexicaul; petals endoglossum 1-2-seeded. mericarp 46. Upper cell of mericarp dehiscent

flaring

of

widely;

Briquetia. at maturity; flowers

cell of mericarp soli flaring widely Colima. tary in the leaf axils; Guerrero, Dirhamphis. or yellow-orange; all petiolate; lavender Leaves bluish petals 0-2-seeded. present or absent; upper cell of mericarp endoglossum with two abor upper cell of mericarp present; not developing to absent; seeds; stipules vestigial not developing Nuevo Coahuila, apical wings; mericarps Batesimalva. Le?n, Tamaulipas.

47. Endoglossum tive ovules,

47. Endoglossum ing 1-2 seeds ally (except nia, Sonora.

absent;

upper

cell of mericarps

(rarely none); stipules inH. exalata) developing

usually

develop

usu mericarps apical wing; Baja Califor

manifest;

Horsfordia. 1-seeded. uniovulate; mericarps encircled by elaborate pectinate endoglossum in ventral wall of mericarp; fruit somewhat inflated.

44. Carpels uniformly 48. Seeds closely nating

48. Endoglossum ous structure);

usually absent (if present, fruit not inflated.

then a minute,

origi Gaya.

inconspicu

more or less disintegrat walls of mericarps evanescent, the dorsal wall usually with an umbo or spine ing at maturity, at the dorsal angle (rarely absent); leaves sometimes with an irregular purple blotch along the midvein.

49. Lateral

50.

Petals mas

50.

49.

erect, red; androecium cl?vate or knobshaped.

Petals

yellow, spreading, androecium shorter than

and

stigmas

exserted;

stig

Per?ptera. or purplish; white, lavender, to equaling the corolla; stigmas Anoda.

flattened. usually capitate, or reticulate; of mericarps Lateral walls smooth persistent, or at umbos absent dorsal in present (except spines angle but apical spines sometimes leaves usually present; Fryxellia), without

purple

51. Low

blotch

caespitose

along midvein (except in Sida anodifolia). reticulate herbs; mericarps laterally, dor with a small endoglossum; Coahuila. Fryxellia.

sally spinescent, 51. Erect shrubs or subshrubs herbs

but

(rarely with sal spine.

not

caespitose);

an endoglossum)

(sometimes

prostrate perennial smooth laterally mericarps or reticulate, lacking a dor

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MALVACEAE OF MEXICO

with

52. Mericarps walls.

smooth

19

(or weakly

lateral

reticulate)

53. Plants

herbs; corolla pale yel prostrate perennial low with a rose flush. [Note: Sida jussieana may but incorrectly key out here.] fortuitiously 54. Leaves involucel sometimes asymmetrical; or less to more stellate pubescence nor inflated; calyx neither accresent lepidote; in saline habitats. northern Mexico Malvella. present;

54. Leaves

involucel absent; pubes symmetrical; accrescent be stellate; calyx notably and reticulate-membranous inflated coming at maturity; Nuevo San Luis Potos?, Le?n, cence

in oak forest. Tamaulipas, Krapovickasia. corolla usually trees, shrubs, or subshrubs; rarely purple. yellow, [Note: a few species o? Sida whose mericarps have smooth lateral walls (e.g.,

53. Erect

S.

may

rzedowskii)

out

key

here

sought in couplet 59.] 55. Trees or large shrubs 3-10 low, 1.5-4 cm long; calyces fruits

indurate,

cence Oaxaca, 55. Shrubs

with

m

tall; petals

Chiapas, Dendrosida.

(-5) m in Sidastrum

subshrubs

yel

10-ribbed; septicidal dehis

capsular); 1-3

yellowish (or purple tum), less than 1 cm

be

should

basally

imperfect

(functionally Guerrero. or

but

tall; petals panicula

long; calyces ecostulate, fruits fragile, schizocarpic,

rounded; basally the mericarps ind?hiscent. essentially 56. Mericarps with smooth lateral small broadly 56. Mericarps

present;

endoglossum ovate.

walls; leaves

Allosidastrum.

sometimes

weakly

reticulate leaves

absent; endoglossum rowly elliptic to ovate. with reticulate lateral walls. erally;

lat nar Sidastrum.

52. Mericarps 57. Mericarps

ind?hiscent, fragile-walled, essentially reticulate or smooth Sidastrum. laterally. re 57. Mericarps the lateral walls indurate, strongly ticulate (except in Sida rzedowskii et aff.). 58. Calyx accrescent; notably plants decumbent weakly

to ascending herbs with blackish roots; mericarps

with

ally reticulate-fenestrate, zontal rostrum. 58. Calyx

not

accrescent erect

plants

prostrate

herbs; mericarps

rostrum). or 59. Calyx 10-ribbed the 5 commissural

posed; rose or purplish); spined or muticous.

hori

or

subshrubs

lat

usually some dehiscent, horizontal (lacking

-angled nerves

the midribs;

the foliage corolla white

tap later

Rhynchosida. in Sida jus

various,

reticulate, erally apically times with 2 apical spines

than

prolonged

(except herbs or

sieana);

prominent or herbs,

large perennial at maturity,

at

the base, more

often

subshrubs

usually spirally dis or yellow (rarely

mericarps Sida.

apically

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2

20

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

erect ecostate, rounded; basally the foliage and branching pattern or purple; corolla lavender distichous; with double-walled mericarps apical " "horns. Meximalva.

59. Calyx

shrubs,

Abelmoschus Medikus, & Millspaugh,

45. 1787.?Lectotype, Malvenfam. designated 1920: Abelmoschus moschatus Medikus. Bamia R. Brown ex Sims, Bot. Mag. 41: t.1702. 1815.?Lectotype, nated: Hibiscus manihot Linnaeus.

by Britton here desig

or hispid. herbs or subshrubs, often tomentose annual or perennial Robust, Leaves long-petiolate, the blades palmately lobed, crenate or dentate, lacking foliar nectaries. Flowers solitary in the leaf axils; involucel of 4-16 bractlets, usually at sometimes caducous, calyx spathaceous, persistent; splitting asymmetrically anthesis and falling with the corolla; corolla large, yellow with a dark red center; androecium included, apically 5-toothed; style single with 5 sessile capitate stigmas. or hispid, 5-carpelled, with numerous dehiscent, Capsules elongate, pubescent or squamous. Base chromosome num seeds; seeds reniform, glabrous, pubescent, ber not known but large; reports have been made of 2n = 56, 58, 68, 72, 130, and other numbers. to Don (1831), Abelmoschus Etymology. According bic name kabb-el-misk, which signifies musk-seed. The schus exhale a scent like musk." Jaeger (1950) derives

is "latinized from the Ara seeds of Hibiscus abelmo the name from the Arabic

father of musk. Genaust abu-al-misk, (1976) agrees with Don rather than Jaeger. is an Asiatic Abelmoschus genus of six or seven species, some of which have been in cultivation, introduced elsewhere sometimes naturalized. becoming is widely cultivated Abelmoschus for the immature fruits, eaten as a esculentus vegetable ("okra" or "chimbomb?"). References:

Hochreutiner

Borssum Waalkes

(1951a),

(1924), Kearney

(1966),

Bates (1968b). Key 1. Pedicels 1. Pedicels

0.5-2 1.5-9

cm

to the

long; capsules

cm long; capsule

Species with hairs

of Abelmoschus

hairs 0.5 mm

stiff,

1-3 mm

inMexico

long; bractlets long; bractlets

less than 1mm

of involucel of involucel

wide.

1. A.

more

than 1mm

esculentus.

wide. 2. Hairs

of capsule

spreading;

hairs

of pedicel

(and

stem)

antrorse;

bractlets

of

involucel

mm wide. 2. Hairs

of capsule

appressed;

hispid

hairs of pedicel

(and stem) retrorse; bractlets mm wide. 1.5-2.5

5-8

2. A. manihot. of involucel 3. A. moschatus.

1. Abelmoschus esculentus Methodus 617. 1794. Hibiscus (Linnaeus) Moench, esculentus Linnaeus, "esculentus 15" (holotype: Sp. pl. 696. 1753.?Type: A of the is Molinari LINN-875.31!). type photo (1965, pl. reproduced by 15). Erect herbs 1-2 m tall, the stems sparsely 10-15 cm long, often cordate, palmately mostly

Leaf blades hispid to glabrescent. lobed or parted, crenate, sparsely

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1988

MALVACEAE OF MEXICO

21

the sometimes hispid above and beneath especially on nerves; petioles exceeding cm long, sparsely hispid, becoming stout in fruit (5 mm in blades. Pedicels 0.5-2 at base, to 10 mm in diameter diameter incrassate involucel of ca. 9 distally); mm after anthesis; calyx 15 bractlets 10-15 long, 0.5 mm wide, hispid, deciduous cm long; anthers yellow; stigmas purple. Cap 17 mm long, hispid; corolla 3.5-4.5 sules up to 13 cm long, 3-4 cm in diameter, seeds 4 5-angular, minutely pubescent; mm long, glabrous. Chromosome numbers: 2n = 72,108,120,122,130,132. names: Vernacular cocoa, chimbomb?, okra, algalia, ang?, chimbinvoy, quimbomb?. This species

the immature fruits being eaten in grown as a vegetable, and Martin (1982) give further informa various ways. Martinez (1959, pp. 226-228) tion on this useful species. Abelmoschus esculentus occurs throughout the world (in warmer the It presumably in southern Asia but is parts) in cultivation. originated not certainly known in the wild. It is cultivated inMexico in home gardens and as an agricultural crop (Fig. 1). Hinton (in herb. no. 4727) notes that the seeds are used as a coffee substitute. Johnston (1949, p. 195) comments on its ability to naturalize on sites of former cultivation.

Specimens

Examined.

4727.?Guerrero: Men?ndez

is widely

Palmer

404; Ortega

Baja 282.?

California: Oaxaca:

D?az Rzedowski

s.n.? Michoac?n: 31327;

Zizumbo

1718. ?M?xico: Hinton Fryxell & Colunga 482, 513.?Tabasco:

908.

2. Abelmoschus

manihot Malvenfam. 46. 1787. Hibiscus (Linnaeus) Medikus, manihot Linnaeus, 696.1753. Sp. pi. Hibiscuspapyriferus Salisbury, Prodr. nom. 384. Borssum Waalkes 1796, 1966, p. 97): stirp. (cf. superfl.?Type in herb. Clifford specimen (lectotype: BM); second specimen in herb. Clif ford (isotype: BM); "13 manihot" (isotypes: LINN-875.30! S!).?Other are Borssum Waalkes synonyms given by (1966).

Subshrubs to 2.5 m tall, the stems sparsely antrorsely pubescent with short rigid hairs. Leaf blades mostly 8-12 cm long, slightly wider than long, usually 5-lobed, coarsely dentate, glabrate above and beneath; petioles usually shorter than the blades. Pedicels 1.5-7 cm long, antrorsely hispid; involucel of 4-6 lanceolate to ovate bracts, each 5-8 mm wide, more or less persistent; cm 2-2.5 soft calyx long, corolla 6 cm long; anthers pallid; stigmas purple. Capsules 5-6 cm long, pubescent; 2.5 cm in diameter, fusiform, apically acuminate, hispid with spreading simple hairs 3 mm long; seeds 3.5 mm long, minutely num pubescent, striped. Chromosome bers: 2n = 68,130,194. 2. Fig. names: flor de algod?n, hierba de la culebra, pazija, viuda. Vernacular The masticated seeds, applied to the site of the bite, are said to be effective snakes. Abelmoschus manihot occurs from southern Asia to New against poisonous Guinea and Australia; it is introduced in cultivation and sometimes naturalized in near human in humid lowlands Mexico, habitation, hot, generally usually (Fig. 1). It flowers throughout the year. Specimens Veracruz:

Examined.

Inzunza

Nayarit:

69; Ortega

204,

Rose

1911.?

342; Zol?

Puebla:

without

collector,

date,

or number

42, 567.

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(US).?

22

systematic

botany

monographs

VOLUME 25

\

i \

? Abelmoschus

Abelmoschus esculentus

manihot

Abelmoschus

Abutil?n abutiloides

moschatus

Abutil?n

Abutil?n

andrieuxii

barrancae

}4

Abutil?n bastardioides FIG.

Abutil?n berlandieri 1. Distribution

of species

of Abelmoschus

^

and Abutil?n.

3. Abelmeschus

moschatus Malvenfam. 46. 1787. Hibiscus abelmoschus Medikus, moschatus Linnaeus, Sp. pl. 696. 1753. Hibiscus (Medikus) Salisbury, Prodr. Cat. horti Endlicher, stirp. 387. 1796. Abelmoschus of?cinalis Vindob. 2: 253. 1842, nom. superfl. Abelmoschus abelmoschus (Linnaeus) Fl. 157. Deutschi. ed. ii. 2: 1894.?Type: in herb. Clif Karsten, specimen ford (holotype: BM).

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MALVACEAE OF MEXICO

1988

FIG.

2. Abelmoschus

manihot.

Fruit

(left),

portion

of

23

flowering

branch

(center),

mature

leaf

(right). (Ortiz342.) Hibiscus

collinsianus

Nuttall

"collinsiana").?Type: designated: PH!).

ex Torrey & Gray, FI. N. Amer. West U.S.A. Florida, Ware s.n.

1: 237.

1838 (as here

(lectotype,

1-3 m tall, the stems retrorsely hispid. Leaf blades mostly 8-12 cm to palmately the lobes often narrowly lobed or parted, linear, long, hastately crenate-serrate, hispid above and beneath; petioles subequal to the blades. Pedicels 5-9 cm long, retrorsely hispid, incrassate distally; involucel of ca. 9 bractlets 8-20 mm long, 1.5-2.5 mm wide, in fruit, hispid, usually decidu sometimes persisting cm mm corolla 7-8 30-35 ous; calyx long; anthers yellow; stigmas long, hispid; cm cm in 3-4 6-7 fusiform, diameter, antrorsely hispid, long, purple. Capsules number: seeds 4 mm long, glabrous. Chromosome hairs 1-2 mm long, appressed; 2n = 72. Erect

herbs

names: algalia, caf? extranjero, Vernacular chimbomb?, la culebra, lirio. for its showy flowers and as a This species is cultivated for snake bite and as The seeds are used as a remedy occurs in southeast Asia, Malesia, moschatus Abelmoschus in the tropics. InMexico it is cultivated introduced elsewhere in hot, humid localities (Fig. 1). ralized, generally Specimens Hern?ndez

&

Examined. Cedillo

1255;

San

Luis

Morales

Potos?: 10;

Soto

Owens

s.n.?Veracruz:

& Horvitz

46.?Oaxaca:

Do?a

Elvira,

flor de

source of bast fibers. a coffee substitute. It is and Australia. and sometimes natu

467; Dorantes Mart?nez-Calder?n

Ch?zaro

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3593; 197.?

SYSTEMATIC BOTANY MONOGRAPHS

24

Tabasco:

Fryxell

730; Gonz?lez

100; Gonz?lez

& P?rez

4184.?Chiapas:

VOLUME 25

Breedlove

7711,12880,

28873;

Calzada 2751; Fryxell 3217; Gilly & Hern?ndez 432;Mart?nez 7998;Montes de Oca 337; T?llez 879; Ton 1587; Valdivia

2420.

Miller, Gard. diet, abridg. ed. iv. 23. 1754.?Lectotype, designated by ex A. Gray, 1852: Abutil?n See discus Shuttleworth theophrasti Medikus. sion following the generic description. ex A. Gray, Smithsonian Contr. Knowl. 3 (art. 5, PI. Belo?re Shuttleworth 1909: Belo?re pt. 1): 21.1852.?Lectotype, Wright, designated by Hassler, ex A. Shuttleworth Gray. cistiflora R. E. Fries, Kongl. Svenska Vetenskapsakad. Handl. 43(4): 96. Pseudabutilon 1908.?Lectotype: Pseudabutilon scabrum (K. Presl) R. E. Fries. (See discussion of this choice of lectotype under Allowissadula).

Abutil?n

or pubescent, sometimes shrubs to small trees, glabrescent or the blades Leaves ovate, cordiform, glandular-pubescent. petiolate, elliptic, sometimes lobed or parted, usually crenate or serrate (rarely subentire), lacking into racemes or foliar nectaries. Flowers solitary in the leaf axils or aggregated Subshrubs

or

involucel absent; calyx lobes lanceolate, into umbels; less often white, often yellow or orange, lavender, rose, included the column fila (rarely exserted), usually at apex; styles 5 to many; mentiferous stigmas capitate. Fruits schizocarpic (but 5 to many, twice the length of the column, sometimes pseudocapsular); mericarps usually apically acute or usually 1-celled (rarely partitioned by an endoglossum), less commonly panicles, ovate, or cordate; corolla or purplish; androecium

to spinescent acuminate (rarely rounded), usually with 3-6 seeds each, not inflated, more or less elliptic in cross section, dorsally dehiscent; seeds glabrous or slightly = number: x 7, 8. pubescent. Base chromosome cites Britton and Millspaugh Index Nominum Genericorum (1920) as choosing as lectotype, although Britton and Brown had earlier done Sida abutil?n Linnaeus so (cf. Jones 1960). However, Shuttleworth (ex A. Gray, 1852, p. 21) even earlier as had chosen Abutil?n avicennae J. Gaertner (based on Sida abutil?n Linnaeus) It lectotype. (Both of these names are referable to Abutil?n theophrasti Medikus.) to establish, is necessary that one of these names can be explicitly con however, of one of the fifteen species included by Miller nected with the description (without Both A. avicennae and A. theophrasti in the original generic description. on Sida abutil?n. None of Miller's species can be explicitly related to 5. abutil?n, but several (no's. 1, 2, 3) probably represent this species, and no. 1 clearly does, described by Miller as "very common in Virginia, and most of the other Parts of America; where it is called by some of the Inhabitants Marsh-Mallow, because are in this Sort." See Spencer soft and woolly. There is no great Beauty the Leaves the history of the early introduction of this species to the United (1984) concerning binomials) are based

States.

It has been variously interpreted. The origin of the name Abutil?n Etymology. or Arabic in Don has been said to be Greek, (1831) origin. "probably Oriental," and Sweet (1839) said that Abutil?n "is the Greek name for the mulberry tree; in the shape of the leaves." Conzatti resemblance (1903) says it is an "ancient but gives no derivation. Gray (1897) attrib name" meaning "without appendage," an origin described in greater detail by Iljin (in uted the name to Avicenna,

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1988

25

MALVACEAE OF MEXICO

and physician Avi Shishkin, philosopher 1974): "Name given by the Bukharian . . .The name Abutil?n cenna or Ibn-sina (980-1037) is to [Abutil?n theophrasti]. the derived from the Greek words a not, bous bull, and tilos watery excrement, in cattle." Genaust (1976) denies the plant being used as a remedy for diarrhea and asserts the name to be Arabic. The name could conceivably Greek derivation be derived from the Arabic abu, father of, and the Persian tula or tulha, Malva or mallow.

includes about Abutil?n 45 occur inMexico.

200

from

species

the Americas,

Africa,

and

Asia,

Australia;

References: Baker (1893), Garcke (1893), Standley (1923), Kearney (1951a, 1955a),Wiggins (1964,1980), P. A. Fryxell (1976b), J. E. Fryxell (1983). of Sectional

Synopsis

of Abutil?n

Classification

sect. Abutil?n.

Abutil?n

Included

Mexican

sect.

Abutil?n

A.

species:

Anasida

hirtum. Fl.

Grisebach,

Brit. W.I.

78.

Abutil?n

1859.?Type:

umbellatum

(Linnaeus)

Sweet. ser. Umbellata

Abutil?n

in Martius,

Schumann

Fl. Bras.

366.

12(3):

1891.?Type:

Abutil?n

um

Sweet. (Linnaeus) sect. Abutilastrum J. Bot. 31: 71. 1893. Pseudabutilon E. G. Baker, Wissadula subg. Abutilastrum Handl. Svenska Vetenskapsakad. (E. G. Baker) R. E. Fries, Kongl. 43(4): 102.1908.?Type: Wissadula scabra K. Presl. bellatum

Included

Mexican

sect.

Abutil?n

Armata

Presl Included

K.

Presl,

A.

divaricatum

mollicomum,

A.

reventum,

A.

sect. Mexabutilon

Suffrutices

floribus

var.

A.

barrancae,

Reliq.

(cf. Fryxell 1976b). Mexican species: A.

divaricatum,

Abutil?n

A.

species:

Fryxell,

A.

andrieuxii, A.

hintonii,

1835.?Lectotype: A.

bastardioides,

axillaribus;

Abutil?n

A.

A. berlandieri, A. californicum, A. species: A. abutiloides, A. permolle, A. palmeri, A. pinkavae, A. procerum, hypoleucum, A. wrightii. sphaerostaminum, A.

sect. Oligocarpa Prodr. Candolle,

Abutil?n

J. Fryxell, (de Candolle) 1: 467. 1824.?Lectotype, Urban.

tum ( Jacquin) Mexican species: A. coahuilae, A. parvulum, A. percaudatum, A. trisulcatum. Included

Abutil?n

K.

divaricatum

var.

macvaughii,

A.

Abutil?n calyx

A. Gray. hypoleucum in alabastro quinquangularis

umbellatum.

haenkeanum

A.

bracteosum,

haenkeanum, A. xanti.

atque plicatis ad basem. Included Mexican

florum,

A.

thurberi,

A.

tehuantepecense,

sect. nov.?Type:

A.

orientale,

grandidentatum, A.

straminicarpum,

plerumque

2: 114.

Haenk.

A.

ellipticum,

30: 84.

Madro?o designated

A. fruticosum,

A.

1983.

Sida

by J. Fryxell, incanum,

ob

lobis

cordatis

dugesii, A. glabri A. A. simulans,

sect. Oligocarpae de 1983: Abutil?n trisulca

A. malacum,

A. mucronatum,

sect. Pluriovulata

Fr?tices unguiculatis; = 8. Included

sect. nov.?Type: Abutil?n Fryxell, purpusii Standley. vel grandes, interdum arbores parvae; flores comp?rate grandes, petalis fructibus plerumque chromosomatum rotundatis; carpidia 4 vel pluribus ovulis, parvi

Mexican

species:

A.

hulseanum,

A.

jaliscanum,

A. purpusii,

Species

of Abutil?n

A.

striatum,

A.

saepe longi x numerus tridens, A.

vexillarium.

Key 1. Mericarps with numerous than 2.5 cm long (flowers

to the Mexican

seeds; calyx often 2 cm long or longer; (4 or more) sect. Pluriovulata. smaller in A. hulseanum);

petals

often more

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SYSTEMATIC BOTANY MONOGRAPHS

26

VOLUME 25

2. Calyx 1.2-1.5 cm long; petals ca. 2 cm long, pale yellow drying pale rose or lavender; A. 21. hulseanum. with simple hairs 2-4 mm long. cm long, yellow or orange, cm long; petals 2.5-4.5 often long-clawed; 2. Calyx 1.5-3.5 hairs less than 2 mm long or absent. 3. Androecium

to corolla;

subequal

petals

yellow;

mericarps

apically

stems stem leaves

rounded;

unlobed. 4. Petals

3.5-4.5

cm long; mericarps

cm long; central

2-2.5

and southern

Mexico.

cm long; mericarps 2.0 cm long; Jalisco. 24. or yellow, to greatly 3. Androecium sometimes exserted; slightly petals orange rounded or acute; leaves lobed or unlobed. veins; mericarps 4.

5.

Petals

2.5-3.0

ovate-sessile, often

Stipules tube,

inflated,

manifestly

purplish, 5. Stipules

with

red

the lobes much shorter than the prominent; calyx tubular, red or purplish, androecium corolla; nearly yellow equaling in cultivation. leaves unlobed; exserted; plants subglabrous; 47. A. vexillarium.

to elliptic; neither inflated nor half-divided, calyx campanulate, androecium half the length of the corolla; pallid, variously

subulate

ca.

purplish,

usually

exserted; 6. Leaves

pubescence

leaves

various;

unlobed,

cium

36. A. purpusii. A. jaliscanum.

markedly about twice

exserted,

long; Chiapas. 6. Leaves deeply

lobed or unlobed.

androe discolorous; petals orange; plants pubescent; cm the length of the petals; mericarps 2.5-3.5 spinose, 44. A. tridens.

concolorous; plant glabrate; rounded, slightly exserted; mericarps 41. A. striatum. sometimes naturalized. lobed,

androecium

petals yellow with red veins; less than 2 cm long; in cultiva

tion, calyx less than 2 cm long; petals 2.5 cm long or less. Mericarps (2-) 3-seeded; 7. Mericarps to acute; corolla orange-yellow with dark center; obtuse 20-25, 20. A. hirtum. with long simple (2-5 mm) hairs; sect. Abutil?n.

stems

viscid

and

or spinose to acuminate corolla usually yellow (sometimes obtuse); of stem various. dark center ;pubescence orange, purplish), usually without or herbs of vigorous sometimes 2 m or arborescent, commonly growth, more an ample often terminal leafless panicle tall; inflorescence (or raceme); petals more or less reflexed; sect. Armata. sometimes

7. Mericarps

9.

acute

5-15,

white, shrubs

(sometimes 8. Robust

and mericarps

Styles

or purplish; violet, arborescent.

5-8; calyx 10-15 mm long; stems often viscid; petals yellow, staminal column (at least basally); pubescent plants commonly

stems minutely not viscid; and mericarps 5; petals yellow; puberulent, I.A. bastardioides. 5-ribbed; Colima. calyx prominently stems viscid; calyx obscurely 10. Styles and mericarps 6-8; petals violet or purplish; ribbed.

10. Styles

3-4 mm calyx 12-15 mm long; filaments purplish, externally glabrous; stems with often weakly dentate, 3-lobed; long simple long; leaves coarsely to glandular hairs; Oaxaca. hairs (1-2 mm long) in addition 18. A. grandidentatum.

11. Petals

violet, stellate-pubescent abaxially on proximal 2-2.5 mm long; filaments long; leaves denticulate stems glandular-puberulent, lacking long non-glandular

11. Petals

9.

Styles

and mericarps

obscurely reflexed

8-12;

third; calyx 10-12 mm to subentire, unlobed; hairs; Oaxaca.

42. A. tehuantepecense. (-13) mm long; stems usually not viscid (or or orange, sometimes with dark basal spot, buff, yellow, or pubescent; robust herbs or column glabrous plants calyx

3-11

so); petals white, or not; staminal only rarely arborescent.

shrubs, 12. Calyx

divided

in fruit; petals long, strongly 13. Stems and somewhat

almost

to the base,

or buff, yellowish keeled dorsally. petioles retrorse;

with petal

the lobes narrowly lanceolate and fully reflexed with basal spot; mericarps 12-20 mm

sometimes

dense,

simple hairs, Veracruz

spot present;

or long, spreading and Chiapas. var. divaricatum. 12. A. divaricatum 1-2 mm

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1988

27

MALVACEAE OF MEXICO

13. Stems

and petioles hairs,

glandular 12. Calyx

minutely

lacking the

^2-%-divided,

hairs;

lobes

spreading without basal

or orange, yellowish, keeled. strongly 14. Stipules and subulate

white,

sometimes

stellate-pubescent,

long simple

also with minute

spot absent; M?xico. 13. A. divaricatum

petal

or somewhat

reflexed 7-12

spot; mericarps

var. hintonii.

in fruit; petals mm long, not

or white, yellow petals abaxially staminal column (unknown straminicarpum); glabrous glabrous. and minutely often 15. Stems viscid-puberulent dark-pigmented, sparsely inconspicuous;

inA.

to glabrate; mericarps and stellate-pubescent to Oaxaca. long; Sonora and Chihuahua

obtuse;

mm

15. Stems

usually

petals 37.

A.

or puberulent usually densely pubescent acute or acuminate-apiculate; usually petals

green,

viscid; mericarps 5-9 mm long (unknown inA. straminicarpum). white, 16. Stems and petioles with long simple hairs 2-3 sometimes calyx lacking in the inflorescence); 27. 5-8 mm long, yellowish.

9-15

yellow,

(-4) mm 4-6 mm

reventum.

but never to

yellow

(these

long

long; petals A. mollicomum.

or -tomentose, and petioles lack densely stellate-puberulent ing long simple hairs; calyx 4-8 mm long; petals 7-9 mm long, yellow ish to greenish white inA. straminicarpum). (unknown 17. Pubescence of stem 0.5 mm or longer; mericarps dark brown or cm at maturity; blackish leaves often often 8-15 3-lobulate, A. 49. xanti. long; Baja California.

16. Stems

17. Pubescence maturity; 14. Stipules

broad

of leaves

stem

0.1 mm

unlobed,

at straw-colored long; mericarps less than 5 cm long; Puebla.

usually

40. A. straminicarpum. and prominent, falcate to auriculate-clasping (but caducous); or orange, often with minute abaxial staminal pubescence;

petals yellow at least at base. column stellate-pubescent 9-20 mm 18. Calyx 8-13 mm long; petals yellow, 2-4 mm long. reflexed; filaments

long,

slightly

or not at all

mm mm 8-10 inflorescence 18-20 long; calyx long; of stem stellate, minute; branched; paniculate, pubescence stipules, and calyx eciliate; leaf surface subglabrous floral bracts, upper (but with a few minute 25. A. macvaughii. stellate hairs); Jalisco.

19. Petals

mm mm 9-15 9-13 inflorescence long; calyx long; stem hairs simple, 1-2 mm long; stipules, racemiform, unbranched; floral bracts, and calyx ciliate; leaf surface with appressed upper to Chiapas. 9. A. bracteosum. simple hairs; Colima

19. Petals

4-10 mm 18. Calyx long; petals 1-1.5 mm long. filaments 20. Mericarps or acute puberulence 20. Mericarps

orange,

6-14

mm

long,

reflexed;

clearly

(hairs 0.2-0.5 mm long), relatively blunt short-puberulent stems leaves and with stellate apically; (beneath) mm long; Oaxaca 0.2-0.5 and Chiapas. 5. A. andrieuxii.

1-1.5 mm long-hirsute (hairs long), and stems whitish with puberulence (beneath) to Chiapas. Sinaloa 19. than 2 m tall (except A. procerum and sometimes

Plants usually less and A. trisulcatum); inflorescence glabriflorum, and A. trisulcatum). incanum, A. mucronatum, 21. Calyx lobes usually cordate and overlapping

various;

petals

not

acuminate; ca. 0.1 mm A.

leaves long; haenkeanum.

A. barrancae, reflexed (except

A. A.

the bud thus plicate-pentangular; basally, 6-15; sect. Mexabutilon. mericarps 22. Stems markedly viscid; seeds glabrous. 23. Calyx lobes 5-9 mm wide, with glandular hairs intermixed with stellate leaf surface usually with both stellate and glandular upper pubescence; central Mexico and Baja California. hairs; leaves somewhat discolorous; 14. A.

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dugesii.

SYSTEMATIC BOTANY MONOGRAPHS

23. Calyx lobes 9-15 mm wide, with minute stellate lar hairs, often flushed with purplish pigment

22.

VOLUME 25

pubescence, lacking glandu in part; upper leaf surface

leaves markedly discolorous, hairs; lacking glandular stellate-pubescent, whitish central Mexico. 38. A. simulans. beneath; in A. berlandieri, viscid (except glandular hairs present Stems not or obscurely inA. pinkavae); seeds minutely pubescent. sparsely present up to 4 m

Plants

24.

Plants

seldom

blades

less than 12 cm long. soft to the touch,

than 2 m

more

tall,

discolorous

palmeri). 26. Petals

26.

35. A. procerum. leaf

sometimes

procumbent-ascending;

the leaves often often whitish, in A. pinkavae concolorous and A.

the hairs

25. Herbage

markedly

than 12 cm long; Tamaulipas.

tall; leaf blades more

24.

(except

nearly

tomentose,

17-22 mm white-stellate white, long; calyx densely also with the hairs less than 0.5 mm long [sometimes

long (2 mm)

simple

pure

hairs];

anthers

ca. 100; eastern Mexico. 17. A.

glabriflorum. or orange (sometimes rose), 6-25 mm long; calyx not or if so then petals anthers fewer than white-tomentose, yellow; 100. Petals

yellow

2-4 cm long; styles and mericarps 6-9; plants procum or erect, less than 0.5 m tall, stems slender. bent, ascending, or ascending; leaves markedly discolor 28. Plants procumbent as long; corolla about as wide ous, prominently dentate,

27. Leaves

28.

Mexico. 48. A. wrightii. pale yellow; northeastern Plants erect; leaves nearly concolorous, crenate, obscurely Nuevo corolla orange; Coahuila, Le?n, longer than wide; 34. A. pinkavae. Chihuahua.

than 4 cm long; styles and mericarps 9-15; usually more plants erect, 0.5-2 m tall, stems stout. 29. Leaves often as wide as long; stems with simple hairs 1-2 mm long; northern Mexico. 30. A. palmeri.

27. Leaves

29.

Leaves

than wide;

longer

stems usually

lacking

long simple

hairs. 30.

cor 13-15; calyx lobes notably Styles and mericarps than 8 mm wide; date and often accrescent, often more to Puebla northeastern Mexico, extending principally 22. A. hypoleucum. and lobes 9-12; calyx only slightly Styles mericarps cordate and accrescent, less than 8 mm wide. generally and styles 31. Pedicels less than 3 cm long; mericarps and Oaxaca.

30.

39. A. sphaerostaminum. 9-11; Veracruz. 31. Pedicels more than 3 cm long; mericarps and styles 33. A. permolle. and Yucatan 10-12; Chiapas peninsula. to the touch, the hairs often yellowish, the leaves not or

25. Herbage

rough only slightly discolorous. hairs intermixed 32. Glandular northeastern 32. Glandular

with

stellate 8.

Mexico.

hairs

on most

plant

A

parts; berlandieri.

hairs absent.

seed pubescence of carpel wall uniform; recurved, 4. A. abutiloides. in reticulate bands; northern Mexico. arranged seed pubescence 33. Pubescence of carpel wall of two size classes; to appressed, erect distributed; Baja California, uniformly 10. A Sonora. californicum. 33.

21. Calyx 5-10.

lobes

Pubescence

lance-ovate

or

triangular,

not

overlapping;

mericarps

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MALVACEAE OF MEXICO

29

or less cylindrical more or caskshaped, sometimes than wide; longer or acute less than 1 mm (beaks finely puberulent mericarps long); herbage sect. Oligocarpa. seeds short-pubesent; tomentulose; stems sometimes stems prominently 35. Young in viscid, especially trisulcate; fruits columnar, medially inflorescence; constricted; widespread.

34. Fruits

45. A. stems terete, 35. Young constricted inmiddle.

stems

fruit 9-12 mm without

staminal

not

pet

column);

at base; long, yellowish 32. A. percaudatum. than 7 mm long; herbage

long; central dark center

or petals less not viscid (except A. mucronatum);

or puberulent, pubescent mm long; fruit 6-10 mm

trisulcatum.

or caskshaped,

fruits columnar

center (and purplish viscid; calyx 6-8 mm and southern Mexico.

yellow with purple als 8-15 mm long; herbage

36. Corolla

36. Corolla

or not;

viscid

calyx

3-8

long.

but leaf surface visible (at least on mature stellate-pubescent often decumbent; leaf margin the hairs; plants through basal spot; dentate; coarsely petals 4-7 mm long, pinkish without flowers and fruits solitary in the axils, few and scattered; northwest

37. Leaves

leaves)

ern

37. Leaves leaf

31. A. parvulum. and beneath, the the stellate hairs; plants petals 3-15 mm long, yellow

or tomentulose softly puberulent of surface obscured by the density

Mexico.

above

serrate to crenate; erect; leaf margin or pink; flowers and fruits often aggregated (at least in well-grown plants). and styles 6-9; northeastern 38. Mericarps and styles uniformly 5. 38. Mericarps

into terminal Mexico.

39. Calyx 6-8 mm long, erect in fruit; terminal panicle; northern Mexico. 39. Calyx 3-6 mm fruits scattered

panicles

16. A. fruticosum. a compact 26. A. malacum.

inflorescence

in fruit; flowers long, erect or reflexed or in open terminal panicles.

and

10-15 mm basal rotate, long, yellow without and fruits in open in fruit; flowers calyx reflexed south-central Coahuila. H.A. coahuilae. panicles;

40.

Petals spot;

40.

Petals

3-6

reflexed,

spot; calyx usually tered or paniculate. 41. Petals obtuse,

column

squamose, pillate, 42. Mericarps apiculate

nearly glabrous; leaf axils; Sonora. flowers

herbage

at apex, the beaks usually more than 2mm long, not constricted absent, the mericarps basally. with long, simple hairs (2-4 mm long) or sometimes leaves cordate-ovate; flowers in the solitary or paired 43.

leaves truncate stellate-pubescent; into inflorescences. aggregated

44. Leaf

Mexico. viscid; west-central 28. A. mucronatum.

or spinose

endoglossum and petioles

Stems

pallid;

about

subglobose, rounded; herbage

43.

staminal

as wide as long; mericarps or acute to often apiculate or long-hirsute; seeds verruculate, pa stellate-pubescent or glabrous; sect. Anasida.

34. Fruits

divergent; 43. Stems

basal long with or without in fruit; flowers and fruits scat

basal yellow or pink with prominent tomentu column purplish; herbage Mexico. 23. A. incanum. lose; northwestern Petals mucronate, basal spot ; staminal pink without spot;

41.

mm

erect

mm

blades basally cordate, in axillary long; flowers

Leaf

blades

sometimes umbels;

A. thurberi. or cordate,

weakly widespread

ovate

to elliptic;

fruits 5-9 tabulate; at low elevation. 46. A.

44.

13-14 spread

mm

basally

truncate, unlobed, flowers in terminal

long; at high elevation.

more racemes 15.

or

less elliptic; or panicles; A.

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umbellatum. fruits wide ellipticum.

30

SYSTEMATIC BOTANY MONOGRAPHS

to acute at apex, 42. Mericarps if beaked rounded the mericarps double present, endoglossum long; basally. 45. Mericarps

8-10;

plants

erect;

stems

lacking

VOLUME 25

the beaks more

long simple

less constricted

hairs;

Chiapas. 45. Mericarps (4-) 5 (-6); plants with simple hairs (1-2 mm to Sinaloa Oaxaca.

1 mm

less than

or

Sonora 6. A.

to barrancae.

the branch tips drooping; stems scandent, to stellate pubescence; in addition

long)

29.

A.

orientale.

4. Abutil?n abutiloides Bot. Porto Rico 5: ( Jacquin) Garcke ex Britton & Wilson, 547. 1923. Sida abutiloides Jacquin, Obs. 1:17. t.7. 1764. Abutil?n jacquinii 1: 503. 1831, nom. hist. G. Don, Gen. superfl. Abutil?n decarpum nom. 1: 40. fl. Amer. in cult, from New 1836, superfl.?Type: Rafinesque, Jamaican seeds Lavatera americana

(BM). Linnaeus,

Syst. nat. ed. x. 1148. 1759. Sida americana americanum (Lin (Linnaeus) Linnaeus, Sp. pi. 963. ed. ii. 1763. Abutil?n non 53. Hort. Abutil?n brit. americanum 1826, Sweet, Panzer, naeus) 1797.?Type: "americana" (holotype: LINN-871.7!). 1: 28. t.6.f.2. 1785. Abutil?n Sida lignosa Cavanilles, Diss. lignosum (Cava 1: 501. 1831.?Type: G. hist. Gen. Santo Do Don, nilles) [Hispaniola] no. as MA P-JU 12312!; photo F-29783!). mingo (holotype: isotype: ex R. Hort. Paris Sida tricuspidata Cavanilles, Diss. 1: 30. t.6.f.5. 1785.?Type: as MA (P?, photo F-29792!). Sida crassifolia L'H?ritier, Stirp. nov. 2: 125. t.60. 1789. Abutil?n crassifolium

hist. 1: 504. 1831.?Syntypes: G. Don, Gen. Hispaniola, (L'H?ritier) Thierry, Dubour(G-DC\). Sida tomentosa Cavanilles, Descr. in Hort. Madrid pi. 164. 1801.?Type: as MA photo F-29791!). (holotype: Bull. Soc. Nat. 31: 205. Abutil?n Turczaninow, domingense Imp. Moscou s.n. Santo 1858.?Type Jaeger Domingo, [Hispaniola] (holotype: KW!; isotype: OXF!). Proc. Amer. Acad. Arts 24: 41. 1889.?Type: scabrwn S. Watson, Abutil?n 4-30 Jul 1887, Palmer 97 (lectotype, Mexico. Sonora: Guaymas, here

designated GH!; isolectotypes: BM! K! NY! US!). Abutil?n dentatum Rose, ico. Chihuahua, Oct

Contr.

U.S.

1885, Pringle

Nati. Herb. 8: 318. 306 (holotype: US!;

1905.?Type: Mex BM! BR! isotypes:

F! GH! K! MASS! MICH! NA! NY! PENN! PH! VT! pf!). to 1.5 m tall, moderately Subshrubs branched, roughly stellate-pubescent (not both the often and stipitate. Leaf blades 2-10 cm hairs sessile viscid), yellowish, cm wide, cordate to ovate, basally truncate to cordate, obscurely to long, 1.5-5.5 sur the lower crenate-serrate, acuminate, discolorous, usually slightly prominently than the upper; petioles 1-4 (-6) cm long, usually face more densely pubescent shorter than the blades; stipules 6-8 mm long, subulate. Flowers solitary in the leaf 1.5-3 (-5.5) cm long, articu axils or aggregated into racemes or panicles; pedicels lated 3-7 mm below the flowers, calyx 9-12 mm long, subequal to the petioles; the lobes basally cordate and overlapping densely stellate-pubescent, (plicate in corolla rotate, without a dark center, the petals 10-12 mm long, bud), acuminate; on margin of claw, otherwise glabrous; staminal stellate-pubescent orange-yellow, 1.5-2.5 mm long, the column 3-4 mm the filaments long, stellate-pubescent,

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1988

31

MALVACEAE OF MEXICO

Fruits 9-14 mm long, yellow; styles 8-10, glabrous, exceeding androecium. or the 10-16 mm in diameter, 8-10, calyx; mericarps slightly exceeding equaling with minute stellate hairs acuminate, 3-seeded, uniformly nearly always apically and with longer (0.5-1 mm) simple hairs on dorsal margin; seeds 3 mm long, with minute recurved scabridulous hairs arranged in a reticulate pattern. Chromosome number: 2n = 28. anthers

names: malva Vernacular colotahue, rasposa, misbil, colotag?e, pelotazo branco, sak-xiu (maya), yax-holch? (maya). occurs from Oaxaca to Sonora (and adjacent Arizona), Abutil?n abutiloides across northern Mexico and and in the West Indies. It adjacent Texas, (Fig. 1) occurs generally 1000 m in open, arid habitats and flowers more or less below throughout the year. The name for this species is often cited as A. abutiloides (Jacquin) Garcke; Garcke however, Garcke used the binomial only as a nomen provisorium. simply cited Schumann the name provisional, and stated that he and, like him, considered use name to A. for the this The earliest preferred crassifolium species. acceptable and Wilson, is that by Britton cited above. The no usage of A. abutiloides confusion has been discussed by Strong (1977), as has the confusion of menclatural A. abutiloides and A. californicum. Specimens Hartman

239,

111-76,

114-76,

Examined.

Sonora:

935; Hastings 125-76,

Breedlove

& Turner

137-76,

149-76;

15952;

65-82, Lehto

65-131, 18602,

538, 3057, 3081; Gentry 1038, 19284; 69-101, 69-115, 72-32; Johnson 147-75, 19489; Palmer 97, 662; Rose 12659, 12763;

Fryxell 65-20U,

19313,

Sanders et al. 2476, 2531, 2615, 2681; Shreve 7526;White 3277, 4494;Wiggins 6394, 7243, 7526, 8244; & Rollins 345. ?Chihuahua: 2884; Palmer 172; Pringle Wiggins Bye 2853, 5671, 7084, 9240; Gentry 306.?Coahuila: Breedlove 7186.?Sinaloa: 1589, 35571; Fryxell 136, 78017; Fryxell & Bates Barkley

2109; Gonz?lez-Ortega 5479, 5908, 6084; Jones 22393, 22855, 22857; Lehto 19512;Mahler & Thieret 5988; Moran

7590; Narv?ez-Montes

13387,

13389, 13489, 13643, & Fern?ndez 18148; Gonz?lez

& Salazar

14548; Sanders 2178.?Zacatecas:

5; Palmer et al. 4268; Rose

10; Rose 13363, 1254, 1687; P?rez 10; Robles 40.?Durango: Shapiro Fryxell 3030; Gentry 2425.?Nuevo Le?n: Dorr 2617. ?Tamaulipas:

Elliot 11; Fisher 3394; Fryxell 710; Fryxell & Bates 817, 821; Johnston 4303B, 4908; Nee 22237; Palmer 410, 574; Runyon 794; Ferris 5583 (Tres Mar?as

Strong

156,

157.?

San

Luis

Islands).?Aguascalientes:

Potos?: Rzedowski

Feddema 3792.?Nayarit: 1274; Fryxell & McVaugh 1254.?Jalisco: Cervantes

& V?zquez 150;Dieterle 3074; Fryxell 1034,1037,1601; McVaugh 22142, 25935; Rose 2886; V?lareal de & Wilbur Puga 6, 1212, 2607; Wilbur Conzatti & Gonz?lez 1085; Ghiesbreght

2387.?Colima:

Ferris

6031.?

Puebla:

Miranda

2991.?Oaxaca:

348.

5. Abutil?n andrieuxii Hemsley, Mex Diagn. pi. nov. mexic. 2: 24. 1879.?Type: ico. Oaxaca: Tlacolula, 522 (holotype: K!; isotypes: G as photo Andrieux F-23764! K! OXF!). The type locality is at 16?57'N, 96?29'W, at 1600 m,

fide Sousa (1979). Abutil?n reticulatum Rose, Contr. U.S. Nati. Herb. 5:171.1899.?Type: Mex ico. Oaxaca: Monte Alb?n, 23 Nov 1894, Pringle 6062 (lectotype, here as BR! F! F-56132! GOET! K! US!; designated: isolectotypes: photo

MASS! MEXU! MICH! MIN, MO! MSC! PH! UC! VT! pf!). Abutil?n calderonii J. Washington Acad. Sei. 14: 99. 1924.?Type: Standley, El Salvador. San Salvador, 1923, Calder?n 1639 (holotype: US!). Robust puberulent

shrubs (hairs

2-5 m stellate,

tall, often with several trunks, the herbage minutely mm 0.2-0.5 long), with glandular hairs only in the

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32

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

cm long, orbicular to ovate, 10-15 mostly deeply or somewhat the blades cordate, obscurely acuminate; petioles equaling Inflo caducous. shorter; stipules broadly foliar, auriculate-clasping, asymmetrical, rescence an ample terminal panicle; pedicels 6-20 mm long; flowers subtended by floral bracts of stipular origin, these prominent but caducous; calyx 4-8 mm long, ca. half-divided, a dark center, the petals 8-14 without in corolla fruit; spreading mm long, orange, reflexed; staminal column 5-6 mm long, densely pubescent, the inflorescence.

blades

Leaf

serrate,

1-1.5 mm long, crowded at the apex, the anthers yellow; styles ca. 10, slender, reddish. Fruits 8-9 mm long, 8-11 mm in diameter, puberulent; mericarps ca. 10, 8-10 mm to acute apically; seeds 2 mm long, rounded long, 3-seeded, Chromosome number: 2n = 14. blackish, sparsely pubescent. Abutil?n andrieuxii occurs in southern Mexico (Fig. 1) and adjacent parts of El in deciduous Central America Salvador, forest, often Honduras) (Guatemala, on steep slopes and in ravines at 600-1600 m. It flowers principally from November toMarch. filaments

Specimens Pringle

6062;

examined. Smith

320,

Oaxaca: 322.?

Andrieux

Chiapas:

522; Conzatti 1639; Fryxell Breedlove 23727, 30590; Fryxell

et al. 78432; 901; Seler & Seler

& Bates

904; Koch

& Bates

894,

1911; Ton 3772, 3909. 6. Abutil?n barrancae M. E. Jalisco: Guadalajara,

Jones, Contr. W. Bot. La Barranca, 19 Nov

18: 57. 1933.?Type: 1930, Jones 27162

Mexico. (lectotype,

POM-192430!; isolectotypes: BM-2! NY! POM-193309! UC!). The type locality is discussed by McVaugh (1972, p. 253). 2: 117. t.69. 1835. Pseudabutilon Wissadula scabra K. Presl, Reliq. Haenk. scabrum (K. Presl) R. E. Fries, Kongl. Svenska Vetenskapsakad. Handl. scabrum S. Watson, 1889.?Type: Mexico. 43(4): 103. 1908, non Abutil?n Haenkes.n. (holotype: PR). Wissadula paniculata Rose, Contr. U.S. Nati. Herb. 5: 178. 1899. Pseudabuti lon paniculatum Svenska Vetenskapsakad. (Rose) R. E. Fries, Kongl. Handl. 104. 1908, non Abutil?n Handel-Mazzetti, paniculatum 43(4): 1933.?Type:

Mexico.

Sinaloa:

Ymal?,

25 Sep-3

Oct

1891, Palmer

1743

(holotype: US!; isotypes:BM! NY!). inornatum Standley Pseudabutilon Bot. Ser. 23: 15. 1943.?Type:

& Steyermark, Guatemala.

Publ, Field Mus. Nat. Hist., near Amatitl?n, Guatemala:

29 Dec 1938, Standley 61425 (holotype: F! as photo F-56242! F-56243!; isotype: US!). 1-2 (-3) m tall, minutely the older branches becom stellate-pubescent, to with reddish brown bark. Leaf blades 8-12 cm long, gray ing glabrate mostly truncate to somewhat ovate, serrate, acute to acuminate, mi cordate, obscurely nutely stellate-pubescent; petioles V2-V3 the length of the blades; stipules ca. 3 mm or paniculate, Inflorescence racemiform often long, subulate. simple or more or subsessile; flowers branched; calyx 3-6 mm long, ca. half short-pedicellate Shrubs

the base of the fruit, with triangular lobes; divided, basally rounded and enclosing a dark center, the petals 5-10 mm long, yellow; staminal corolla rotate, without to glabrous, column 3-5 mm long, sparsely pubescent the filaments almost as long

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1988

33

MALVACEAE OF MEXICO

as the column, the anthers yellow; styles 8-10, slender, pallid. Fruits 4-6 mm long, about as wide, 8-10, 3-seeded, mericarps usually apically stellate-pubescent; with double internal endoglossum; seeds 2 rounded to acute, basally constricted, mm long, reddish brown, papillate. Chromosome number unknown. Vernacular

Abutil?n ous

forest, Guatemala

name:

malva.

inMexico barrancae occurs widely (Fig. 1), in seasonally dry decidu in more open habitats, sometimes below 1200 m, and in generally to December. It flowers principally from October and Honduras.

Specimens 1317.? Sinaloa: Examined. Sonora: Breedlove 35584; Fryxell & Bates 2117; Gentry & Lyons 15628.? 6607; Kimnach 700; Palmer 1743; Webster & Breckon Gentry 4966; Gonz?lez-Ortega Durango: Jones 27162; hangman Breedlove 43921.?Jalisco: 25042, 3159; McVaugh 25841; Mexia Martinez 5479.?Veracruz: 1239, 1293; Pringle 1721; Rose 2928.? Guerrero: 2239; Ventura Purpus & Anderson 256? Oaxaca: Anderson 2655, 2903, 10554, 12032; Villanueva 5346; Fryxell & Bates 903; et al 78349; MacDougall & Lott 3397; Koch 4158. ?Chiapas: 5; Rzedowski 3707; Torres Fryxell et al 30056; Fryxell & Bates 895; Fryxell & Lott Breedlove 20401, 24637, 30570, 41774, 56295; Davidse 3236, 3373.

7. Abutil?n E. G. Baker bastardioides 1895.?Type: Mexico. Colima, US!; isotype: NY!).

ex Rose, Contr. U.S. Nati. Herb. 1: 306. 27-28 Feb 1891, Palmer 1314 (holotype:

Shrubs 2-3 m tall (rarely small trees to 6 m tall, the trunk 7-8 cm in diameter), with minute stellate puberulence. Leaf blades mostly 6-10 cm long, cordate to as as serrate about wide broadly ovate, (sometimes long, obscurely so), short as as Vt-\ times the acuminate, blades; stipules 6 slightly discolorous; petioles long 8 mm long, linear, deciduous. Flowers solitary or paired in the leaf axils, or in short cm long, slender, articulated near the middle, 3.5-4.5 axillary racemes; pedicels and with long (1-1.5 mm) simple hairs intermingled, the buds puberulent mm 12-15 at the 5-ribbed nodding; calyx long, glandular-puberulent, prominently ca. a the lobes corolla without dark the half-divided, base, center, acuminate; petals 1.5-2 cm long, yellow, reflexed; staminal column 8-10 mm long, pallid, stellate

minutely

pubescent basally, glabrous distally; the filaments pallid, 4-6 mm long, the anthers yellow; styles 5, reddish, with dark red stigmas. Fruits ca. 6 mm long, minutely 5, usually 2-seeded, muticous, puberulent; mericarps (Mature fruits and seeds un = 28. 2n Chromosome number: known.) Abutil?n is endemic to Colima (Fig. 1), where it occurs in decidu bastardioides ous woodland at 300-500 m and flowers inOctober. Specimens Cervantes

Examined.

Colima:

Fryxell

1612; McVaugh

& Koelz

1042;

Palmer

1314;

Santana

&

850.

8. Abutil?n berlandieri A. Gray ex S. Watson, Proc. Amer. Acad. Arts 20: 358. 1885.?Type: Mexico. San Oct 1835, Berlandier Fernando, Tamaulipas:

3050 (holotype: GH!; isotypes:MO! NY! PH! US!).

Shrubs hairs often

0.5-1.5

m

yellowish.

the tall, with both glandular and rough stellate pubescence, cm long, 2-8 cm wide, ovate, basally Leaf blades 2.5-10

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VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

34

to cordate, serrate to obscurely crenate, acute or sometimes acuminate, cm shorter than the 1-7 discolorous; blades; long, stipules 3-10 slightly petioles mm long, subulate. Flowers solitary in the leaf axils or forming racemes or panicles; 3-10 mm below the flowers, shorter than to 1.5-3 cm long, articulated pedicels mm 8-13 with the long, densely stellate-pubescent slightly exceeding petioles; calyx cordate at the the and lobes hairs base, acuminate; basally overlapping, glandular a dark center, the petals ca. 1 cm long, orange-yellow, corolla rotate, without on margin of claw; staminal column 2-3 mm long, stellate stellate-pubescent mm 2-3 the filaments long, the anthers yellow; styles 8-10, glabrous, pubescent, mm long, 10-13 mm in diameter, 10-13 androecium. Fruits exceeding subequal to truncate

8-10, 3-seeded, calyx; mericarps apically acuminate, with both stellate and glandu and evenly pubescent. number: lar hairs; seeds 3 mm long, minutely Chromosome 2n = 42. Abutil?n berlandieri occurs in northeastern Mexico (Fig. 1) and southern Texas at relatively low elevations in relatively open, arid habitats. The species is sympatric but it is clearly distinguished with and commonly from confused with A. abutiloides, as shown by Strong (1977). it both morphologically and cytologically, Specimens Coahuila: Examined. 1486, 2668.?Nuevo Fryxell 1941 [some sheets mixed with Allowissadula 2464; Pringle patrick Berlandier 3050, 3108; Fryxell 3731; Gonz?lez-Medrano 91; Runyon 172.

Le?n:

Fryxell

holosericea], 416; Strong

1214; Fryxell & Kirk 10262.?Tamaulipas: 150,152,154,164,171,

9. Abutil?n

32: 262. 1980.?Type: bracteosum Mexico. Oaxaca: Fryxell, Brittonia 6 km al N de Puerto Escondido San Pedro Mixtepec, sobre la ter Mpio. 140 m, 22 Nov 1978, Koch & Fryxell 78408 racer?a a San Gabriel Mixtepec,

(holotype: CHAPA!; isotypes:CTES! ENCB! MICH! NY! pf!). Shrubs to 2 m tall, the stems sparsely hirsute (hairs simple, 1-2 mm long), more Leaf blades mostly 8-14 cm long, cordate, ovate, densely so at base of petioles. as as almost and shallowly with wide denticulate, acuminate, long, remotely beneath; petioles Yi-Y* the length appressed simple hairs above, stellate-pubescent of the blades on midstem shorter upwards; stipules 8-20 mm leaves, progressively foliaceous, long, 5-10 mm wide, sessile, ciliate, falcate to auriculate-clasping, api the buds in the inflo cally laciniate, becoming paired floral bracts, enclosing an interrupted raceme (sometimes caducous. Inflorescence rescence, branched) surpassing the leaves; pedicels 0.3-2 cm long; calyx 9-13 mm long, V^-^-divided, the lobes erect or spreading, minutely and prominently ciliate; stellate-puberulent a dark center, corolla without the petals 9-15 mm long, orange-yellow, slightly staminal column 5-7 mm long, pallid, densely the reflexed; stellate-pubescent, ca. 50; styles 10-11, slender, pallid, filaments 2-3 mm long, the anthers yellowish, with sometimes purplish stigmas. Fruits 7-9 mm long, 8-10 mm in diameter, mi 10-11, 8-9 mm long, nutely and evenly stellate-pubescent; mericarps seeds 3 mm long, minutely scabridulous. Chromosome rounded-acute;

3-seeded, number

unknown.

occurs along the Pacific coast of Mexico Abutil?n bracteosum from Chiapas to Colima (Fig. 3), generally below 500 m in seasonally dry deciduous forest, sometimes to December. in evergreen forest or in disturbed sites. It flowers from October

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MALVACEAE OF MEXICO

1988

35

Vx.

XV

Abutil?n

Abutil?n

californicum

bracteosum

? X \ x'? -^

Abutil?n coahuilae

-?

+-^

Abutilon V_

divaricatum^^ VAr divaricatum

x

^x^*

x>;\ K., O

Abutil?n V_ divaricatum vARhintonii

^-~

5'

Abutil?n ellipticum

Abutil?n dugesii

Abutil?n fruticosum FIG.

3. Distribution

of species

o? Abutil?n.

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36

SYSTEMATIC BOTANY MONOGRAPHS

Specimens Fryxell

Colima:

Ferris

6241

.?Guerrero:

& Fryxell

78408,

79467,

79533.?

Examined.

& Lott3398;

Koch

Hinton Chiapas:

VOLUME 25

11010; Breedlove

742.?Oaxaca: Langlass? 44442; Croat 46277.

Bot. voy. Sulphur 8. 1844.?Type: californicum Mexico. Bentham, s.n. (holotype: K!). See 1841, Hinds Bay, Baja California: Magdalena Raven (1964). Proc. Amer. Acad. Arts 20: 357. 1885.?Type: Abutil?n lemmonii S. Watson, "Tia Mexico. California: here desig Juana," Orcutt 676 (lectotype, Baja nated: GH!; see Strong, 1977).

10. Abutil?n

Abutil?n nia:

fragile Cape

Zoe

Brandegee,

Sierra

Region,

5: 156. 1903.?Type: Nov de la Trinidad,

Mexico. 1902,

Baja Califor s.n. Brandegee

(holotype: UC!). to 1.5 (-2) m tall, roughly pubescent, the stellate hairs mostly sessile, cm Leaf blades 2-8 than often ovate, wide, yellowish. long, longer rarely stipitate, to or sometimes cordate truncate, crenate-serrate, obscurely prominently basally acute or acuminate, slightly discolorous; petioles usually shorter than (or subequal to) the blades; stipules 4-7 mm long, subulate. Flowers solitary in the leaf axils or 1-4 cm long, articulated 3-5 mm into racemes or panicles; pedicels aggregated below the flowers, subequal to the petioles or longer; calyx 6-11 mm long, densely Shrubs

in bud), the lobes basally cordate and overlapping (plicate rotate, without a dark center, the petals 12-15 mm long, orange on margin of claw, otherwise glabrous; staminal column yellow, stellate-pubescent 3-4 mm long, sparsely pubescent apically, the filaments ca. 2 mm long, the anthers Fruits 12-15 mm long, 10-13 androecium. 7-10, yellow; styles glabrous, exceeding mm in diameter, the 7-10, nearly always 3-seeded, calyx; mericarps exceeding stellate-pubescent, acuminate; corolla

the hairs dimorphic apically acuminate, stellate-pubescent, (smaller hairs 0.1-0.2 mm long, larger hairs 0.4-0.6 mm long); seeds 3 mm long, sparsely and evenly number unknown. pubescent. Chromosome on occurs more or less throughout Baja California, Abutil?n californicum western coast of and the Socorro Island of the Revillagigedo Archipelago, along at low in arid habitats Isla Tibur?n) Sonora (including open, (Fig. 3), generally on available the year depending it flowers more or less throughout elevations; confused A. californicum with A. abuti moisture. The literature has commonly loides, but Strong (1977) showed them to be clearly distinct both morphologically from Socorro Island have a different aspect and geographically. The specimens to and from California those Sonora, including denser pubescence Baja compared and more prominent leaf serration. More detailed study may indicate that these at infraspecific rank. plants should be recognized 381; Cruz-Cisneros Carlquist (Revillagigedo Archipelago): 184; Berry 43; Carter 1928, 2284, 3616, 332; Barkelew Anthony 3172; Eastwood 12420; Ferris 8566; Fryxell 1981, 821; Collins 69; Constance 4808, 4920, 5928; Chambers & Turner, 63-267, 67-70, 67-110, 500; Hastings 1992a, 1996, 2000; Gentry 7562, 7721; Guzm?n-Huerta 1056; Mason 1911; Moran 5727, 109-75; Jones 24212; L?pez-Forment 31065; Johnson 71-109; Howell Specimens

1555; Miranda

7075, 21970,

Examined.

8698.?Baja

25226;

Nelson

Isla

Socorro

California:

& Goldman

7201; Orcutt

373,

676; Palmer

840; Porter

299, 322,

345; Purpus

54, 293, 505; Rose 16451,16476,16563,16734,16892; Rzedowski 26556; Sanders 1197,1265, 3318, 5559; Solis 5; Spjut & Edson 5338; Thomas 8306; Whitehead 922; Wiggins 3731, 4453, 5442, 11341, 14662, 15215,

15396,

15586,

17711,

17943,

18119;

Wiggins

& Ernst

524; Wiggins

& Thomas

281.?

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Sonora:

1988

19316;

Feiger

37

MALVACEAE OF MEXICO

Fryxell

Hansen

3073;

19316;

Moran

8799;

Shreve

6544;

Turner

79-62.-Isla

Tibur?n

(Sonora):Alcocer & Sosa 4;D?az 2286; Gold380. 11. Abutil?n coahuilae Kearney, huila: Cerro de Santiago

12: 115. 1953.?Type: Madro?o near Bol?var, Apr 1910, Purpus

Mexico. Coa 4617 (holotype

UC!; isotypes: F! US!). Shrubs to 1 m tall, the stems densely and minutely the hairs tomentulose, stellate, yellowish. Leaf blades mostly 4-6 cm long, ovate, truncate to cordate (the sinus open), tomentulose, nearly concolorous; sharply serrulate, acute, minutely than half the length of the blades; stipules 3-5 mm long, subulate. petioles more Flowers generally borne in an open terminal panicle; pedicels 10-15 (-20) mm mm mm 2-3 below articulated 5-6 the the flowers; long, calyx long, puberulent, a lobes lanceolate, in corolla without dark reflexed the rotate, center, fruit; fully on margins of claw, otherwise glabrous; petals 10-15 mm long, yellow, pubescent staminal column ca. 3 mm long, pallid, with a few stellate hairs apically, the fila ments 1-2 mm long, the anthers yellow; styles 5, glabrous. Fruits 8-10 mm long, 6 8 mm in diameter, more or less cylindrical, green maturing stellate straw-colored, mm 2 seeds 3-seeded, 5, pubescent; mericarps apically long-acuminate; long, mi number unknown. Chromosome nutely pubescent. occurs Abutil?n coahuilae in open, arid habitats in central and southern ca. m. at Coahuila most It 800-1400 of the year. flowers (Fig. 3) apparently Specimens Purpus

Examined.

4617; Rodriguez

Coahuila:

Fryxell

78077;

Fryxell

et al

1499,

1507, Nee

& Diggs

25346;

1026.

12. Abutil?n divaricatum Turczaninow, Bull. Soc. Imp. Nat. Moscou 31: 204. 1858, var. divaricatum.?Type: Veracruz: Mirador, Mexico. 1840, Galeotti 4071 1976b: KW!; BR! G as by Fryxell, designated (lectotype, isolectotypes: photo F-23765?K!). Abutil?n chiriquinum A. Robyns, Ann. Missouri Bot. Gard. Type: Panama. Chiriqui: vicinity of El Boquete, 3 Feb-15 Maurice 705 (holotype: US!; isotype: MO!).

52: 574. 1966.? Mar 1938, Bro.

2-3 m tall, usually pilose with simple hairs 1-2 mm long, the stem hairs slightly retrorse, and with smaller stellate hairs. Leaf blades mostly 8-12 cm long, above ovate, deeply cordate, serrate, acuminate, slightly discolorous, pubescent so beneath; and more densely more than the of half the blades petioles length (rarely subequal); stipules 15 mm long, 10 mm wide (smaller in inflorescence), a terminal panicle; pedicels caducous. Inflorescence auriculate, asymmetrically cm 0.5-2 mm to the base, the lobes 4-8 divided almost (-3) long; calyx long, mm reflexed in 7-9 fruit; petals narrowly lanceolate, fully long, pale yellowish or a with at dark diffuse the sometimes buff, base, reflexed; staminal column 4-5 spot mm long, densely pubescent mm the filaments 1 basally, long, the anthers yellow; mm Fruits 12-20 slender, glabrous. styles 8-12, long, 12-15 mm in diameter, Shrubs

keeled dorsally, basally rounded, hirsute; mericarps 8-12, 3-seeded, prominently mm 2 seeds num apically acute; long, blackish, essentially glabrous. Chromosome ber

unknown.

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SYSTEMATIC BOTANY MONOGRAPHS

38

VOLUME 25

var. divaricatum occurs in Veracruz and Chiapas (Fig. 3), Abutil?n divaricatum as far as Panama, in pine-oak forest and in secondary and through Central America m. to It flowers 2000 up throughout the year. In the past this vegetation apparently with A. been has confused giganteum species ( Jacquin) Sweet, a species that does not occur inMexico, but the matter was clarified by Fryxell (1976b). Examined.

Specimens 1468; Ventura

Veracruz:

9535.?Chiapas:

Galeotti

4071; 8984; Matuda

Breedlove

var.

13. Abutil?n divaricatum Mexico. M?xico:

Distr.

hintonii

Linden

1378; Purpus

8313,

16702; Rosas

16649,

1919.

Fryxell,

Brittonia Pantoja,

Temascaltepec,

32: 265. in barranca,

1980.?Type: 26 Feb 1935,

Hinton 7431 (holotype: K!; isotypes:K! MICH!). var. hintonii differs from var. divaricatum in Abutil?n divaricatum principally on are retrorse hairs that the and the somewhat found stems, petioles, lacking long, It lacks the basal petal spot and also differs in pedicels of the nominate variety. distribution geographical (Fig. 3). Specimens

Examined.

M?xico:

Hinton

3341,

7431.

Proc. Amer. Acad. Arts 21: 447. 1886.?Type: dugesii S. Watson, Mexico. 1883, Dug?ss.n. Guanajuato, (holotype: GH!). Annuaire Jard. Bot. Gen?ve Conserv. Abutil?n tultitlanapense Hochreutiner, Puebla: San Luis Tultitlanapa, 21: 443. 1920.?Type: Mexico. Jul 1908,

14. Abutil?n

Purpus3254 (holotype: NY!; isotypes:BM! F!MO! UC! US!). more or less throughout. Leaf Viscid shrubs to 1.5 m tall, glandular-pubescent to crenate, acute blades mostly 6-10 cm long, ovate, cordate, obscurely denticulate or acuminate, the upper surface with at least some glandular somewhat discolorous, Vi-V? the length of the blades; stipules up to 12 mm long, subulate. hairs; petioles Flowers solitary in the leaf axils or aggregated into terminal panicles; pedicels 1-5 cm long, articulated 2-4 mm below the flowers; calyx 9-16 mm long, glandular the lobes 5-9 mm wide, cordate and overlapping, acuminate; ^-divided, pubescent, corolla rotate, without a dark center, the petals 10-12 mm long, yellow, pubescent on margins of claw, otherwise glabrous; staminal column 2-3 mm long, glabrous, mm 2 the filaments pallid, long, the anthers yellow; styles 8-10, glabrous. Fruits ca. 12 mm long, 11-15 mm in diameter, stellate- and glandular-pubescent; mericarps 8-10,3-seeded, some

apically

number

acuminate;

seeds 2.5 mm

long, blackish,

Chromo

glabrous.

unknown.

and desert zones in central Mexico, dugesii occurs in arid matorral in and and and in the Islas Tres between 1000 2000 m, generally Baja California to Novem Mar?as at lower elevations It flowers from principally August (Fig. 3). ber. The specimens lack glandular hairs on the upper leaf from Baja California surface and have broader leaves but otherwise appear to represent this species. Abutil?n

Specimens Carter

Examined.

& Moran

(Nayarit):

5659;

Mason

Baja

Fryxell 1771; Solis

California:

Carter

1992; Hammerly 74.?Guanajuato:

3113,

3380,

5134; Carter

& Ferris

3797,

199.-Islas 3569; Palmer 304; Moran Calzada s.n.; McVaugh 9422; Dug?s

3890, 4091; Tres Mar?as

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24273;

Rose

1988

MALVACEAE OF MEXICO

11519.?San

Luis

11237. ?Quer?taro: Altamirano Rzedowski 1687; Ars?ne 10303; Fryxell3788; Rose 9803, 11156. ?Michoac?n: Germ?n 1182.? Hidalgo: 695; Fryxell 5593.?Puebla: Conzatti 3254, 4011.?Oaxaca: 2364; Hern?ndez-Maga?a Purpus

Potos?:

1062; Matada

Fryxell

39

Gonz?lez-Quintero

18564;

1977;Delgadillo 132;Delgado 721; Ernst 2206; Smith 636. 15. Abutil?n

Linnaea 11: 368. 1837. Sida elliptica Schlechtendal, bot. ed. ii. 577. 1840.?Type: Mexico. Steudel, Nomencl. s.n. (holotype: 1830, Keerl HAL; Tlalpujahua, isotype:

ellipticum

(Schlechtendal) [Michoac?n:]

BR!). sidoides Hemsley, Diagn. pi. nov. mex. 2: 24. 1879, non Abutil?n sidoides Dalzell & Gibson, 1861. Abutil?n hemsleyanum Rose, Contr. U.S. Nati. Herb. 10: 123. 1906, nom. nov.?Type: San Luis Potos?, Mexico.

Abutil?n

1878,Parry & Palmer89 (holotype: K!; isotypes:BM! NY! US!). Abutil?n

attenuatum

Robinson & Seaton, Proc. Amer. Mexico. Jalisco: near Lake Ch?pala,

1893.?Type:

Acad. Arts 28: 104. 21 Nov 1892, Pringle

4354 (holotype: GH?; isotypes: BM! BR! F! as photo F-56142! GOET! K! MASS! MEXU! MIN, MO! MSC! NY! PH! UC! US! VT! pf!). E. G. Baker ex Rose, Contr. U.S. membranaceum 5 Jan-6 Feb Mexico. 1895.?Type: Nayarit: Tepic,

Abutil?n

Nati. Herb.

3: 312. 1959

1892, Palmer

(holotype: BM!; isotypes: F! GH! NY! US!). Shrubs 1-2 (-3) m tall, coarsely stellate-pubescent. Leaf blades mostly 5-11 cm as as to ovate about twice truncate, serrate, acute or wide, long, long elliptic, more or Va-Vi the less acuminate, discolorous; petioles length of the blades; stipules 5-9 mm long, subulate. Flowers in terminal racemes or panicles; calyx 6-8 mm more than the lobes ovate half-divided, long, stellate-pubescent, basally rounded, a mm corolla without the 8-15 dark rotate, center, acuminate; petals long, yellow or yellow-orange, staminal column 4-6 mm long, pallid, stellate broadly obovate; at base, the filaments ca. 2 mm long, the anthers yellow; styles 5-7, pubescent mm 13-14 Fruits stellate slender, pallid. long, 8-10 mm in diameter, coarsely and invested 5-7, 3-seeded, pubescent, by calyx; mericarps exceeding closely api the spines divergent, 1-4 mm long; seeds ca. 2 mm long, blackish, cally spinose, number: 2n = 16. squamose. Chromosome minutely name: colotag?e. Vernacular

Abutil?n ellipticum iswidely distributed inMexico at 1200-2600 m (Fig. 3) and also occurs in Guatemala. It grows in deciduous forest, pim-oak-Liquidambar and disturbed sometimes forest, sites, reaching cloud forest, and flowers more or less throughout the year. Kearney (1955a) retained A. ellipticum and A. hemsleya num as distinct species, but he distinguished them only inmericarp number (five in the former, or in any other six to seven in the latter) and not in distribution a character. The type of A. membranaceum to be appears juvenile, shade-grown the upper leaves are elliptic although the lower are specimen of A. ellipticum; broader.

Specimens

Examined.

Sinaloa:

Breedlove

1733, 18926; McVaugh 58264. ?Tamaulipas: Duke

Tryon 7783.?Jalisco: Iltis et al

Anderson

1328; Jones

& Anderson

27156;

et al 4372.?Durango: 1235, 5112; Sanders Bustos s.n.; Rollins & 637; Patoni 289.? Zacatecas: 1959. ?Aguascalientes: Palmer Rose & Painter

Gonz?lez-Ortega

23595; Palmer M3579.?Nayarit:

5957; Calzada & Nieves Lott & Magallanes 946; McVaugh

9468; Cervantes 13946,

23392,

80; Fryxell

& Bates

23470; McVaugh

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2153; & Koelz

VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

40

904;Nieves 260; Nieves & Trujillo 114; Pringle 4354, 4583; Rose & Painter 7672; V?zquez 994; V?zquez & Cervantes

148.?Sau 1498,

McVaugh

22722;

Pringle

80, 89; Rzedowski 4386, 5713. ?Quer?taro: Parry & Palmer Feddema s.n.; Koch 12913, 15722; Keerl 77397; 213; Hinton & McVaugh 555.?M?xico: Rzedowski Fryxell & Bates 2173; Garc?a

Potos?:

Luis

2007.?Michoac?n:

Arguelles

13920;

1483;Gold 284; Hinton 1943, 2916, 7028;Matuda 27434, 29520;Medell?n 127; Paray 240, 748; Pringle 11933; Rzedowski 23296, 27642, 29531, 29542, 29562, 32140, 35413, 35246, 37919; Ventura 182.? Distrito 3786.?

Federal:

Balls

Hidalgo:

Crist?bal

Flores

7283; Rzedowski 702, 925; Rose & Painter Bourgeau & Bates 3011, 2174; Gonz?lez-Quintero 3194; 7325, 7347, 9689, 13508; Ventura 3678.?Morelos: Pringle

5607;

97; Fryxell

23475; Moore 5458; Hinton 3586.? Guerrero:

V?zquez Nelson 1523; Rzedowski21754.?Chiapas:

11692;

Schwabe Breedlove

s.n.?Puebla: 49810; Garc?a

26052;

Ventura &

Krapovickas

Lyonnet2831; Hill 1852; Purpus 2604.?Oaxaca: Ton 409. 594; Matuda 87,1905;

Perrottet & Richard, in Guillemin, & Perrottet fruticosum Guillemin 16. Abutil?n D. Dietrich, Fl. Seneg. tent. 1: 70. 1832. Sidaperrottetiana Syn. pl. 4: 855. Senegal. Suffal, Leprieurs.n. 1847, nom. nov.?Type: (holotype: P). non Sida gracilis Richard, Sida gracilis R. Brown in Salt, Abyss. App. 65.1814, Ethiopia. 1822.?Type: 1805-1807, 1796, nee Elliott, 1792, nee Salisbury, Salt s.n. (holotype: BM?; isotype: MO!). 1: 231. 1838.?Type: U.S.A. nuttallii Torrey & Gray, Fl. N. Amer. Abutil?n Texas: on the Red River, Nuttall s.n. (lectotype, by J. Fryxell, designated

1983:NY!; isolectotypes: BM! GH! PH!). 1: 231. 1838.?Type: texense Torrey & Gray, Fl. N. Amer. U.S.A. Abutil?n 43 (holotype: Texas: between Brazoria and San Felipe, 1835, Drummond is not located; isotypes: BM! K-3! NY! OXF!). Note: The BM duplicate a death. Drummond's after dated 1836, year incorrectly fl. abyss. 1: 70. 1847. Abutil?n A. Richard, Tent. microphyllum var. Fl. W. Pakistan 130: Abedin, Richard) (A. microphyllum fruticosum Ethiopia. 58.1979.?Type: Chocho, Petit s.n. (holotype: P).

Abutil?n

Leaf blades 2 1-1.5 m tall, the stems terete, stellate-tomentulose. Subshrubs 10 cm long (often smaller), ovate, cordate, irregularly serrate, acute, densely Vi-iA the length of the blades; tomentulose; stipules ca. 2 mm long, petioles into terminal subulate. Flowers solitary in the leaf axils or more or less aggregated mm cm 2-5 below the articulated 0.5-2 flowers; calyx 3-5 long, panicles; pedicels erect in flower to reflexed in fruit; mm long, ca. half-divided, the lobes lance-ovate, corolla rotate, without a dark center, the petals 5-10 mm long, yellow, the margins of the claw glabrous; staminal column 2-3 mm tall, glabrous, pallid, the filaments ca. 1mm long, the anthers yellow; styles 6-9. Fruits 8-9 mm long, about as wide, 6-9, 3-seeded, tomentulose; apically acute or apiculate; mericarps caskshaped, seeds 2 mm long, blackish, number: 2n = 14. name:

Vernacular

minutely

pubescent

(appearing

Chromosome

glabrous).

pelotazo.

occurs in northeastern in relatively open, arid habitats fruticosum and Mexico (Fig. 3) and in adjacent parts of the United States in Texas, Oklahoma, the year. It also Arkansas (cf. J. Fryxell 1983). It flowers more or less throughout occurs disjunctly in northern Africa and the Levant countries. in the Old World, Abutil?n

Specimens

Examined.

Chihuahua:

Palmer

242.?

Coahuila:

Aguirre

& Reko

60; Dorr

2236;

Fryxell 1485; Gould 10670; Iltis & Lasseigne 20; Johnston & Muller 960;Marsh 389, 442, 1103, 1227, 1665, 2030;

Powell

1414; Waterfall

16665.?Zacatecas:

Rose

& Hay

5265.?Nuevo

Le?n:

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Ars?ne

6243;

1988

41

MALVACEAE OF MEXICO

Bennett 621; Dodge 130;Dorr 2330; Flook & Spears 83; Frye & Frye 2367, 2368; Fryxell 78072;Fryxell & & Mueller 69; Smith M234; 2466; Mueller Fryxell 3729; Fryxell & Bates 814; Gonz?lez-Medrano San Luis Potos?: Lemus 84; Purpus 5386.

Kirkpatrick

52269.?Tamaulipas: Steyermark 92, 8525; Kral 24754; Palmer

Bartlett 572; Runyon

10662; 415.?

17. Abutil?n Jard. Bot. Gen?ve Annuaire Conserv. Hochreutiner, glabriflorum of 21: 439. 1920.?Type: Mexico. Victoria, May-Jun Tamaulipas: vicinity 1907, Palmer 373 (holotype: NY!; isotypes: CM! F! as photo F-56137! and

F-56138! GH! K! MO! UC! US!). Note: The holotype is labelled "glabrifolium." Leaf blades mostly 9-12 cm long, 1.5-3 m tall, minutely soft-pubescent. to denticulate 7-9 ovate, deeply cordate, finely subentire, acuminate, palmately above and Vi-l nerved, beneath, discolorous; softly stellate-pubescent petioles mm or times the length of the blades; 6-8 lanceolate falcate, stipules long, caducous. Flowers solitary or paired in the leaf axils; pedicels 2-6 cm long; calyx 10-15 mm long, softly whitish tomentose less than 0.5 mm long, (hairs stellate, also with simple hairs 2 mm long), ca. half-divided, sometimes the lobes cordate a and overlapping without corolla dark the rotate, center, basally; petals 17-22 mm ca. 4 mm tall, on otherwise staminal column ciliate claw, long, white, glabrous; mm ca. 2-4 the filaments anthers the 100, pallid, glabrous, yellow; styles ca. long, Shrubs

the androecium the stigmas 13, slightly exceeding (half the length of the petals), mm ca. mm Fruits 10 in 14-18 diameter, purple. long, stellate-pubescent; mericarps ca. 13, 3-seeded, or short-spinescent; seeds ca. 3 mm apically acuminate long, number: 2n = 14. irregularly scabridulous. Chromosome occurs from Coahuila to Oaxaca Abutil?n and Tamaulipas glabriflorum (Fig. in deciduous forest and below 300 m; it open matorral, 4), principally generally flowers more or less throughout the year. This species is unique among the Mexican o? in Abutil?n that was curiously species having pure white corollas, a characteristic omitted

from the original

Specimens

Examined.

3652; Fryxell & Bates Veracruz: Dorantes Oaxaca:

description.

Kochetal

Coahuila: Palmer 110.?Tamaulipas: 3043, 3063; Fryxell 1223, Chapman 110A.? 2192; Hern?ndez 1122; Mart?nez 483; Palmer 230, 373.? Puebla: Maslin & Bates 15198.? 507; Fryxell 943; Purpus 5714, 11089; Ventura 10518, 8371, 78284, 78347.

18. Abutil?n 23: 324. 1976.?Type: grandidentatum Fryxell, Madro?o Oaxaca: 11 Dec Cerro Guiengola, H143 1971, MacDougall NY!; isotypes: F! as photo F-56136! NY! pf!).

Mexico. (holotype:

Large shrubs or small trees to 4 m tall, the stems densely and minutely glandular pubescent and with long simple hairs (1-2 mm) interspersed. Leaf blades mostly 10 15 cm long, almost as wide, ovate, often weakly 3-lobed, deeply cordate, coarsely or slightly discolorous, concolorous dentate, acuminate, evenly stellate-pubescent; petioles ca. ?$ as long as the blades; stipules 7-15 mm long, 2-6 mm wide, lanceolate to ovate, in the inflorescence often connate in pairs to form floral bracts, deciduous. Flowers generally in terminal panicles; pedicels 2-10 mm long, viscid; calyx 12-15 mm long, densely hirsute, obscurely ribbed, ca. half-divided; corolla without a dark on center, the petals 10-15 mm long, purplish, often reflexed, densely pubescent

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42

SYSTEMATIC BOTANY MONOGRAPHS

FIG.

4. Distribution

of species

VOLUME 25

o? Abutil?n.

of claw, otherwise staminal column 5-7 mm margin long, stellate glabrous; pubescent, pallid, the filaments 3-4 mm long, the anthers yellow; styles ca. 8, exceed ing the androecium, glabrous, purplish, the stigmas purple. Fruits 8-10 mm long, 10 ca. 8, 3-seeded, 12 mm in diameter, mericarps apically acute or glandular-hirsute; apiculate;

mature

seeds

unknown.

Chromosome

number

unknown.

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1988

MALVACEAE OF MEXICO

43

is endemic to the Isthmus of Tehuantepec Abutil?n grandidentatum is the allied A. tehuantepecense, and grows in dry habitats on hillsides. toMay. from November Specimens H143,

H270,

Examined.

Oaxaca:

Dorado

1626; Fryxell

& Lott3382;

Koch

(Fig. 4), as It flowers

et al. 78324; MacDougall

H442.

19. Abutil?n

2: 115. 1835. Sida presliana D. K. Presl, Reliq. Haenk. nom. Mexico, "in terris mexi 4: nov.?Type: Dietrich, 856,1847, Syn. pi. s.n. (holotype: PR!; isotypes: BM! MICH! canus occidentalibus," Haenke haenkeanum

MO !PR!W as photo F-32631 !). or rarely small trees 2-3 (-6) m tall, the stems softly and minutely 10-15 cm long, broadly ovate, Leaf blades mostly stellate-puberulent. deeply serrate, acuminate, cordate, obscurely discolorous, softly puberulent (hairs ca. 0.1 mm long); petioles equaling the blades or sometimes longer; stipules 1.5-2 cm long, but early deciduous, and foliaceous sessile, asymmetrically auriculate, prominent an ample terminal panicle; pedicels acuminate. 1-1.5 cm long; calyx Inflorescence 6-10 mm long, ca. half the length of the fruit (smaller in specimens from the Islas Tres Mar?as), the lobes hirsute, the hairs 0.5-1 mm long, more than half-divided, Shrubs

a dark center, the petals 6-9 to spreading; corolla without more or somewhat fleshy, less reflexed; staminal column 3-4 mm at at least 1-1.5 mm long, the base, the filaments long, pallid, stellate-pubescent anthers usually purplish; styles 8-10, slightly exceeding androecium. Fruits ca. 1 cm ca. and 8-10, wide, blackish; mericarps 3-seeded, long apically acuminate, promi the hairs 1-1.5 mm long; seeds 2 mm long, minutely verruculate. nently hirsute,

lanceolate-acute, mm long, orange,

erect

number unknown. name: hoja de plata. Vernacular Abutil?n is found inwestern Mexico from Sinaloa to Chiapas (Fig. haenkeanum It forest in and below 500 m and disturbed grows along 4). margins vegetation to June. It is closely allied to A. andrieuxii, which has a flowers from December more southerly distribution and occurs at higher elevations. (Oaxaca to Honduras) The specimens from the Islas Tres Mar?as tend to have smaller calyces than those from the mainland conform to the species. Rose (1899b) incorrectly but otherwise a of this species inA. reventum. collection placed (Nelson 4203) Chromosome

Specimens Examined. Sinaloa: Tres Mar?as 5644.?Islas Gentry (Nayarit): 4203; Rovirosas.n?Nayarit: 1049,1054.?Michoac?n: Fryxell4942.?Colima: Fryxell Koch 22604. ?Morelos: 77487; McVaugh 1109, 1366, 3024, 3030, 3038.? V?zquez et al. 405.?Chiapas: Matuda 16340. 2296; Paray 1829; Tenorio 11783; Nelson

98; Nelson Maltby Hinton 16214; Hinton Guerrero:

20. Abutil?n

hirtum (Lamarck) Sweet, Hort. brit. 53. 1826. Sida hirta Lamarck, 1: 7. 1783. Abutil?n indicum var. hirtum (Lamarck) Grisebach, Fl. Encycl. Brit. W.I. 78. 1859. Abutil?n in graveolens var. hirtum (Lamarck) Masters Hooker s.n. India 1: 327. 1874.?Type: f., Fl. Brit. India, Sonnerat

(holotype: P-LA!). Sida pilosa L'H?ritier,

Stirp.

nov.

130. 1789, non Sida pilosa Miller,

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1768, nee

44

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

1825.?Type: 1785, nee Vellozo, 1781, nee Cavanilles, India, s.n. (holotype: P). ex Hornemann, Hort. Hafn. Sida graveolens Roxburgh suppl. 77. 1819. ex & Catal. pi. Abutil?n graveolens Arnott, Hornemann) (Roxburgh Wight s.n. (holotype: C; isotypes: BR! K!). 13.1833.?Type: India, Roxburgh ex A. Gray, Smithsonian Contr. Knowl. 3 (art. Belo?re cistiflora Shuttleworth Florida: Feb 21. 1852.?Type: U.S.A. West, 5, PL Wright, Key pt. 1): Retzius, Sonnerat

1846, Rugel94

(holotype:

GH!;

isotypes: K-2!).

Robust herbs ca. 1m tall, the stems viscid and with long simple hairs 2-5 mm long. Leaf blades mostly 5-7 cm long, subrotund, cordate, finely serrate, acuminate, to above; petioles beneath, more sparsely pubescent equaling softly tomentose mm recurved. 7-9 Flowers the blades; stipules long, lanceolate, greatly exceeding becoming a terminal panicle; pedicels solitary in the leaf axils but the inflorescence corolla 2-3.5 cm long; calyx 12-27 mm long, ca. half-divided, stellate-tomentulose; mm a the 18-20 staminal with red dark center, rotate, long, orange-yellow; petals the filaments 4 mm long, the anthers yellow; styles column 7 mm long, pubescent, the calyx; 20-25. Fruits 12-14 mm long, ca. 2 cm in diameter, oblate, exceeding ca. 1 to the hairs obtuse acute, stellate-hirsute, 20-25, 3-seeded, mericarps apically mm =

long; seeds 2.4-2.8

mm

long, minutely

scabridulous.

Chromosome

number:

2n

42.

names: bot?n de oro, malva lisa. Vernacular Abutil?n hirtum is an Old World species that occurs in tropical parts of Africa, in the West and It is adventive Australia. Indies, southern Florida, Mexico, Asia, Its occur and Central America, and parts of South America (Peru, Venezuela). rence inMexico is sporadic in the eastern lowlands (Fig. 4). Specimens Schott

Examined.

Veracruz:

Palmer

71.?Tabasco:

Matuda

3280; Rovirosa

359.?

Yucat?n

121.

21. Abutil?n hulseanum (Torrey & Gray) Torrey ex A. Gray, Mem. Amer. Acad. 23. 1849. Sida hulseana Torrey & Gray, Fl. N. Arts n.s. 4 (PI. Fendl.): s.n. Florida: Tampa Bay, H?lse U.S.A. Amer. 1: 233. 1838.?Type:

(holotype: NY!). Abutil?n leucophaeum Hochreutiner, Puerto Rico. 15. 1902.?Type: tensis3738 (holotype: G?).

Annuaire Gu?nica:

Jard. Bot. Gen?ve 6: Conserv. et Salinas, Sin inter Montalba

1-2 m tall, the stems and petioles minutely Perennial herbs or subshrubs and with long simple hairs 2-4 mm long. Leaf blades mostly 6 stellate-tomentulose above and 10 cm long, ovate, cordate, crenate, rounded-acute, softly tomentulose or to blades somewhat shorter; beneath, more or less discolorous, subequal petioles stipules ca. 8 mm long, filiform. Flowers solitary in the leaf axils; pedicels up to 12 cm long, articulated 6-24 mm below the flowers; calyx 12-15 mm long in flower, more than half-divided, accrescent to 15-20 mm long in fruit, stellate-tomentulose, corolla rotate, without a dark center, the petals the lobes cordate and overlapping; ca. 2 cm long, yellowish staminal column 4-5 mm long, the fading rose-lavender; filaments 2-3 mm long, the anthers yellow; styles ca. 12. Fruits 12-15 mm long, 2 ca. 12, 1.5 cm long, 4-6-seeded, 2.5 cm in diameter; mericarps apically apiculate,

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:

45

MALVACEAE OF MEXICO

1988

hirsute, the hairs 1-2 mm long; seeds ca. 2 mm prominently number: 2n = 14. Chromosome long, minutely pubescent. Texas to Tabasco (Fig. 4) and in Abutil?n hulseanum occurs from southernmost and the West Indies (Puerto Rico, Jamaica, Cuba, and the Florida, Honduras, commuta auctt. non H.B.K. and Abutil?n Lesser Antilles). Abutil?n pedunculare to this species. This species apparently are referrable turn auctt. non Schumann blackish

at maturity,

flowers principally Specimens Guti?rrez

from February Puebla:

Examined.

260, 302.-Tabasco:

toMay.

Harriman

Cowan

2138,

15614.?Veracruz: 2814,

Burke

1046; Castillo

etal.

1619,1635;

2951.

hypoleucum A. Gray, Smithsonian Contr. Knowl. 3 (art. 5, PI. Wright. s.n. Nuevo Le?n: Monterrey, Berlandier Mexico. pt. 1): 20. 1852.?Type: 752: cf. 1923, p. K!). Standley, (lectotype, Abutil?n selerianum Ulbrich, Repert. Spec. Nov. Regni Veg. 12: 227. 1913.? 12 Oct 1895, Seier & Seler 1096 Type: Mexico. Nuevo Le?n: Monterrey,

22. Abutil?n

isotypes: GH! US!). (holotype: B, destroyed; Annuaire Jard. Bot. Gen?ve Conserv. Abutil?n subsagittatum Hochreutiner, 1May-13 Mexico. of 21: 444. 1920.?Type: Victoria, vicinity Tamaulipas: K! Jun 1907, Palmer371 US!). (holotype: NY!; isotypes: with Shrubs 0.5-1.5 m tall, the stems sparsely to densely stellate-pubescent, or without 5-11 cm long, narrowly ovate longer simple hairs. Leaf blades mostly (up to twice as long as wide), deeply cordate, crenate, acute or slightly acuminate, discolorous above, white 7-9-nerved, beneath), (green palmately markedly V2-I times soft-tomentose beneath; petioles above, densely sparsely scabridulous the length of the blades; stipules 2-4 mm long, subulate. Flowers solitary in the the flowers, with 4-8 mm below leaf axils; pedicels 2-8 cm long, articulated similar to that of stem; calyx 1-1.5 cm long in flower, softly white pubescence the the lobes broadly cordate tomentose, basally, (overlapping deeply 5-lobed, in fruit (up to 2.5 cm long, the lobes to 2 cm wide); accrescent buds 5-angled), cm long, yellow or a dark center, the petals 1.5-2.5 corolla rotate, without staminal column 5 mm tall, ciliate on claw, otherwise glabrous; yellow-orange, the filaments 3-4 mm long, ca. 10-nerved, pallid, glabrous, apically antheriferous, the androecium. Fruits ca. 50, the anthers yellow; styles 13-15, pallid, exceeding 1.5 cm long and wide and exceeded by calyx), stellate-pubescent; (enclosed seeds 2.5 mm acuminate; 3-seeded, 13-15, long, minutely apically mericarps number: 2n = 14. scabridulous. Chromosome name: tzacotxojol The use of this species for me Vernacular (or tzacotzojol). no. dicinal purposes is cited by Alcorn 2582). (in herb. and Texas south to Oaxaca is found from Coahuila Abutil?n hypoleucum (Fig. sea and disturbed level to 1300 m. It apparently sites from 4), in dry matorral flowers

throughout

the year.

Coahuila: Henrickson 10297; Marsh 93, 420, 562a, 597a.? 13100; Johnston 7168; Carlson 2701, 2718; Crockett 8061; Dorr 1981,1998, 2012, 2020, 157; Barkley 2328, 2482, 2621; Edwards 1208, 1696, 2439, 2465; Johnston 294; 305A; Fryxell 4625; Lacas 62; LeSueur Mueller & Mueller 664; Seler & Seler 1096; 84, 1128, 1201; Nelson 6111; Rollins & Tryon 5887; Roybal Specimens

Nuevo

Le?n:

Examined.

Alanis

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SYSTEMATIC BOTANY MONOGRAPHS

46

Smith M522;

Stevens

2246;

Thompson

267; Weaver

569. ?Tamaulipas:

VOLUME 25

Bartlett

10155;

Berlandier

3107;

Fryxell 640, 3674; Fryxell & Bates 815, 820; Fryxell &Magill 2258, 2354; Gaumer & Clark 19;Hansen & Nee 7331;Higgins 2614; Johnston 4909, 4997D; King 45m; LeSueur 278;Nee et al. 19444;Nelson 6661; Palmer 371; Puig 3694; von Rozynski 31; Runyon 988; Stanford et al 2330; Viereck 634; Whiffen & Rodr?guez 201.?San Luis Potos?: Chapman 3091; Fryxell 654, 788, 3795; Lundell 5522; Pringle 7858; Urbinas.n.? & Hough 4879; Rzedowski 6453, 7017, 24606.? Hidalgo: Calzada 161; Fryxell 756; Ervendberg 10032,11227.?Veracruz: 4528; Dorantes Hill 1774. 551; Worth 87.? Oaxaca: 294; Nee & Diggs24636; S?nchez-Mejorada

Purpus & Rose

4900; Rose

Puebla:

Rose

1679; LeSueur

23. Abutil?n incanum (Link) Sweet, Hort. brit. 53. 1826. Sida incana Link, Enum. Hawaii, "in insulis Sandwich," alt. 2: 204. 1822.?Type: h?rt, berol. Link's J. 1983: description). Fryxell, by designated (lectotype, Jard. Bot. Gen?ve 6: 14. Conserv. Annuaire Abutil?n pringlei Hochreutiner, & J. 1902. Abutil?n incanum subsp. pringlei Lowe, (Hochreutiner) Felger Sierra Tucson, 21 U.S.A. Arizona: Arizona Acad. Sei. 6: 83. 1970.?Type: Apr 1884, Pringle s.n. (holotype: NY!; isotypes: K! US!). Annuaire Conserv. Jard. Bot. Abutil?n pringlei var. sinaloensis Hochreutiner, in vicinity of Topolo Mexico. Sinaloa: 21: 437. 1920.?Type: Rose 13272 Mar 23 1910, (holotype: NY!). bampo, 21: Annuaire Conserv. Jard. Bot. Gen?ve Abutil?n mochisense Hochreutiner, 23 Mar 1910, Rose Sinaloa: near Los Mochis, 447. 1920.?Type: Mexico. Gen?ve

13329 (holotype: NY!; isotype:US!). to 2 m tall (usually smaller), the stems minutely Subshrubs tomentulose, green or sometimes purplish. Leaf blades up to 6 cm long (often smaller), ovate, cordate, densely tomentulose; petioles irregularly serrulate or crenate, acute to acuminate, half the length of the blades or sometimes longer; stipules 3 mm long, subulate. Flowers solitary in the leaf axils or in open leafy panicles; pedicels 0.5-2 cm long, the 2-5 mm below the flowers; calyx 3-5 mm long, ca. half-divided, articulated in fruit; corolla with a dark lobes lance-ovate, reflexed) (or somewhat spreading center, the petals 4-6 mm long, yellow or pink, reflexed, the margins of the claws staminal column 2-3 mm tall, purplish, mi the apex obtuse; densely pubescent, or bifurcate hairs, the filaments 1-2 mm long, purple, nutely pubescent with stellate the anthers yellow; styles 5. Fruits 7-9 mm long, ca. 6 mm in diameter, more or less columnar or caskshaped, tomentulose; 5, 3-seeded, apically acute or mericarps mm pubescent (appearing glabrous). long, blackish, minutely apiculate; seeds ca. 2 number: 2n = 14. Chromosome names: pelotazo, Vernacular chico, tronadora. pelotazo incanum occurs in open, arid habitats (up to 1000 m) in Baja Califor Abutil?n in Hawaii. This and disjunctly Sonora nia, Sinaloa, (Fig. 4), adjacent Arizona, have been more and A. mucronatum, species and its close allies, A. fruticosum incanum appar and characterized by J. Fryxell (1983). Abutil?n clearly delimited in the year, although predominantly ently may flower more or less throughout stem fibers. It is sometimes used for its October and November.

Specimens

Examined.

Baja

California:

Carter

3213,

4295,

4481,

4809,

5051; Carter & Ferris

3802,

3889; Carter & Moron 5378; Carter & Sharsmith 4192; Collins 64; Fryxell 1980, 2011; Hammerly 116; Henrickson

2214;

Johnson

105-75;

Jones

24103,

27103,

27165; Montufar

29; Moran

7066,

7077,

12550,

12585, 18184, 18626, 18881, 20626; Nelson & Goldman 7250; Palmer 120; Sanders 3289; Thomas 8442;

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1988

47

MALVACEAE OF MEXICO

Whitehead 718;Wiggins 7945, 11449, 14500, 15536, 17359.? Isla Coronados (Baja California): Moran 9105.?

Sonora:

Ban

& Brewer

63-649;

Breedlove

18636; Daniel

938; Dawson

1066; Drouet

& Richards

3955; Fryxell 539, 78100, 78103; Gentry 233, 2983, 3543, 4700; Goldman 390;Hartman 201;Hastings & Turner

69-106,

72-35;

Johnson

156-75,

115-76,

121-76,

126-76,

128-76,

132-76,

147-76,

209-76,

219-76;

Jones 22398; Lehto 19364;Mahler & Thieret 6042; Maltby 213, 216;Moran 8779; Palmer 6, 104, 288, 650, 651, 768, 810; Rose 1254,12355, 12901; Sanders 1001, 4622; Shreve 6096, 6780; Smith 4718;White 4495;Wiggins 6067, 6328, 7195, 7310,8246; Wiggins & Rollins 343, 476.?Sinaloa: Fryxell 140,142, 541; Gibson & Gibson 2106; Lehto 19513,19593,19635; Palmer 220; Rose 13272,13329,13504. 24. Abutil?n jaliscanum Publ. Field Columbian Mus., Bot. Ser. 4: 229. Standley, 1929.?Type: Mexico. Jalisco: Hacienda de Ototal, Arroyo de los Palos 1842a (holotype: F! as 8 Mar 1927, Mexia Blancos, W of San Sebasti?n,

photo F-45085!; isotypes:BM! CAS! DS! GH! MICH! NY! US!). Trees 4-5 m tall, the branches Leaf stellate-tomentulose. slender, greenish, blades to ca. 25 cm long, broadly ovate, deeply cordate, minutely serrulate to stellate above, paler and densely subentire, acuminate, green and subglabrous as as on the almost lower blades shorter beneath; leaves, puberulent long petioles cm long, articulated near distally. Flowers solitary in the leaf axils; pedicels 4.5-6.5 base of the calyces; calyx 1.5-2 cm long, densely stellate-pubescent, deeply divided, the lobes acuminate, 1-nerved; corolla more or less tubular, without a dark center, cm long, pale yellow, the petals 2.5-3 rounded distally, broadly long-clawed; to the androecium column the filaments 4-5 mm long, subequal petals, glabrous, the anthers yellow; styles ca. 12, slender, exceeding the androecium. Fruits ca. 2 cm ca. 12, ca. 6 long and wide, densely stellate-pubescent, subglobose; mericarps number unknown. seeded, apically rounded; mature seeds unknown. Chromosome Abutil?n is known only from the type collection from dense wood jaliscanum land in Jalisco (Fig. 4). It is closely allied to (and sometimes in) A. submerged but is it from the in distinct latter flower and fruit charac purpusii, morphologically ters and occurs outside the range of A. purpusii (cf. Figs. 4 and 7). 25. Abutil?n

macvaughii

Fryxell, Madro?o

23: 332.1976.?Type:

Mexico.

Jalisco:

8

km E of Chamela, 8-10 Dec 1970,McVaugh 25118 (holotype: MICH!; isotypes:

pf! plus 7 duplicates

to be distributed).

Shrubs 1-3 m tall, the stems sparsely and minutely Leaf stellate-pubescent. blades mostly some 8-12 cm long, ovate, deeply cordate, subentire, acuminate, what discolorous, the upper surface sparsely and minutely pubescent (subglabrous); leaves, shorter distally; stipules petioles ca. Yi the length of the blades on midstem and foliaceous but early deciduous, sessile, asymmetrically prominent auriculate, in the inflorescence laciniate floral bracts. Inflo acuminate, forming racemose or paniculate, mostly above the leaves; pedicels 2-15 mm long; ca. half-divided, somewhat reflexed in calyx 8-10 mm long, minutely puberulent, a corolla without dark the petals 18-20 mm long, yellow; fruit; rotate, center, staminal column 4-5 mm long, pallid, stellate-pubescent at least at base, the fila ments 3-4 mm long, the anthers pallid; styles ca. 10, exceeding the androecium, stigmas purplish. Fruits 6-8 mm long, 8-9 mm in diameter, densely and minutely ca. 10, 3-seeded, stellate-puberulent; mericarps apically acute to obtuse; seeds 2 2.5 mm long, minutely scabridulous. Chromosome number unknown. Fig. 5. eciliate, rescence

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SYSTEMATIC BOTANY MONOGRAPHS

48

FIG. androecium,

5. Abutil?n gynoecium,

and mature branch macvaughii. Flowering mature fruit, seed. (McVaugh 25118.)

leaf;

VOLUME 25

at

right,

above

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to below:

1988

FIG.

MALVACEAE OF MEXICO

6. Distribution

of species

49

of Abutil?n.

Abutil?n macvaughii is confined to the coastal plain of Jalisco (Fig. 6) in decidu ous woodland to Febru and disturbed sites below 150 m; it flowers from November more occurs most to which is the A. It allied bracteosum, ary. southerly nearly to the coastal from Colima Oaxaca. along plain

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50

SYSTEMATIC BOTANY MONOGRAPHS

Specimens

Examined.

Jalisco:

Iltis & Nee

1592; Magallanes

VOLUME 25

313; McVaugh

20875,

25118,

25321,

26255;McVaugh & Koelz 1682; P?rez 627,1101; Sousa3889, 3913; Villareal 5921. 26. Abutil?n malacum U.S.A. Texas:

S. Watson, Proc. Amer. Acad. Arts 21: 446. 1886.?Type: El Paso, 10 Sep 1883, Jones 4193 (lectotype, by designated

J. Fryxell, 1983: POM-84966!; isolectotypes: BM! BR! F! NY! PH! POM the type locality, Jones states "My botanizing 84969!). Concerning on foot from El Paso to the north. Paso was mostly in the mountains went west a few miles" (Lenz 1986, p. 325).

at El I also

Herbs or subshrubs up to 1m tall, the stems minutely the stellate-tomentulose, hairs yellowish. Leaf blades mostly 3-7 cm long, often as wide as long, suborbicular to ovate, cordate, sharply serrate, acute, densely tomentulose; Vi-1 times petioles as long as the blades; mm a 5-9 subulate. Inflorescence long, stipules compact terminal panicle; 3-12 pedicels (-20) mm long; calyx 6-8 mm long, ca. half a dark the lobes lance-ovate and erect in fruit; corolla rotate, without divided, center, the petals 9-15 mm long, yellow, pubescent on margins of claw but other wise glabrous; staminal column 3-4 mm long, pallid, stellate-pubescent, the fila ments ca. 2 mm long, the anthers yellow; styles 5. Fruits 6-7 mm long, about as the hairs 0.5 mm long; wide, more or less cylindrical, coarsely stellate-pubescent, mm 2 seeds acute; 5, blackish, 3-seeded, mericarps long, minutely usually pubes cent (appearing glabrous). Chromosome number: 2n = 14. occurs in relatively arid habitats in northern Mexico Abutil?n malacum (Fig. 6) and also inArizona, New Mexico, and Texas, usually at elevations up to 1500 m. It the year. apparently flowers throughout Specimens

Examined.

Sonora:

Spooner

& Schilling

2427; White

4562.?

Chihuahua:

Dwyer

14221;

Knobloch 349; Lehto 21571, 22841, 22861; Nelson 6421; Pringle 164, 363; Randolph 207; Stewart 2605; Waterfall

6355.?Coahuila:

Chiang

8333A,

9529;

Fryxell

1487,

1489,

78076; Henrickson

5929,

6875,

7695,12082,12194,12291; Johnston &M?ller 842;Marsh 97; Palmer 112;Passini & Robert 5677; Pearce 2151;

Pinkava

5905;

Stewart

268,

345,

557,

616,

1309,

1837; Waterfall

16641; Wynd

746.?Durango:

Correll & Johnston 19979, 21433; Daniel 465; Fryxell 78078; Fryxell & Bates 2065; Iltis & Lasseigne 95; Koch 78139; McGill 8027; Pittier 488; Kirkwood So.?San Luis Potos?: Fryxell

13343, Schery 45; Waterfall 10811. 779; Rzedowski

15365.?

Zacatecas:

Banda

s.n.;

27. Abutil?n

mollicomum Sweet, Hort. brit. 54. 1826. Sida mollicoma (Willdenow) ex caldarii, Willde Enum. pi. hort. berol. 725. 1809.?Type: Willdenow, now s.n. (holotype: B-Wno. 12718!). Sida sericea Cavanilles, Descr. 1768.? pi. 165. 1801, non Sida sericea Miller, Type: described from a plant grown in the Madrid Botanic Garden, proba collected by N?e (holotype: MA as bly raised from seeds from Acapulco

photo F-29788!). sonorae A. Gray, Smithsonian Contr. Knowl. 5 (art. 6, PL Wright, pt. Abutil?n Sonora: Sonoita River, Mexico. 23.1853.?Type: 2): [16Sep] 1851, Wright

538 [899] (holotype: GH!; isotypes:K-2! NY! PH!). Shrubs 1-2 m tall, the stems erect, usually green, prominently hirsute, the hairs mm cm 2-4 Leaf 10-20 blades mostly simple, long, spreading. long, broadly ovate,

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1988

51

MALVACEAE OF MEXICO

or -lobulate, deeply cordate, acuminate, irregularly crenate-serrate, 3-5-angulate and above, discolorous, softly felted minutely evenly stellate-pubescent markedly mm to 5-9 the tomentose blades; beneath; stipules long, petioles subequal axils in the but in leaf subulate. Flowers sometimes generally solitary ample termi 4-8 mm 1-4 cm long, articulated nal panicles rising above the leaves; pedicels corolla below the flowers; calyx 4-6 mm long, 1/2-^3-divided, stellate-pubescent; a dark center, the petals 5-8 mm long, yellowish; staminal column rotate, without meri glabrous; styles ca. 10. Fruits 8-10 mm long and wide, stellate-pubescent; mm ca. ca. 2 seeds dark brown, 10, 3-seeded, carps minutely long, short-apiculate; verruculate. Chromosome number: 2n = 14. name: pintap?n cimarr?n. occurs principally and in adja in Sonora and Chihuahua mollicomum in Guerrero, and It also occurs disjunctly and New Mexico. Oaxaca, m at to 1500 arid habitats elevations It in Veracruz is found up relatively (Fig. 6). to December. and flowers from September name with a Sweet cited Cavanilles's In making in Abutil?n, the combination to de Candolle. The de Candolle "?" and attributed Sida mollicoma reference, so an Willdenow's citation of Sweet's combina is indirect name, however, simply Vernacular

Abutil?n cent Arizona

tion is correctly cited above. in the Madrid garden is a plant cultivated The type of Sida sericea Cavanilles that the type of Sida is from seeds collected by N?e in Acapulco. (It quite possible name is a sterile derived from the same source.) The type of Willdenow's mollicoma seen me the in Because lack microfiche specimens plant by only reproduction. of structures is and because the is it difficult microfiche poor quality, reproductive to be certain of the identity of the plant. However, the leaf shape conforms closely to that of the type of Sida sericea and to the better-known plants from Sonora a as sonorae. at F A. N?e collection from known Moreover, generally Acapulco to a Palmer collection conforms (no. 45) from the same locality. The pubescence in the available that is characteristic of this species cannot be clearly observed . . mollissime . tomen but Willdenow the (foliis photographs, descriptions given by . . . como for and Cavanilles los Sida mollicoma ramos, ("tallo peloso tosis") are for Sida sericea that described the indicate pec?olos, y ped?nculos") species they the one under discussion. A direct examination of the Palmer and N?e collections a collection from Acapulco this view. In addition, from confirms by Haenke was name the for the which basis Abutil?n s.n., (Haenke MO!), Acapulco giganteum sensu Presl 1835 [non ( Jacquin) Sweet, 1826], also supports this interpre tation. It was Presl's misapplication of the name that gave rise to statements inmore recent literature that A. giganteum occurs inMexico, which it does not (cf. Fryxell

1976b). Willdenow the carpel number of his plant as 9; Cavanilles describes says 6, but an examination of the type suggests 8 as a more accurate count. of the photograph These figures are in conformity with the interpretation presented here.

Specimens

Examined.

Sonora:

Drouet

& Richards

3507,

3749;

Fryxell

3086; M?ller

3615;

Sanders

1903, 1940; Shreve 6643, 6759; White 4681; Wiggins 7017, 7478;Wiggins & Rollins 373; Wright 899.? Chihuahua: 45.?Veracruz:

Gentry

2891;

Higgins

Palmer Haenke 71,171; Pringle 329, 944. ?Guerrero: et al. F-1487. 2573.?Oaxaca: Gonz?lez-Medrano

s.n.; N?e

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s.n.; Palmer

SYSTEMATIC BOTANY MONOGRAPHS

52

VOLUME 25

30: 89. 1983.?Type: mucronatum J. Fryxell, Madro?o Mexico. ca. 15 km E of Hwy 15 on the road from Navojoa to Alamos,

28. Abutil?n

Sonora:

(27?02'N, 109?30'W), 200m, 22Oct 1978,Fryxell &Whitacre 128 (holotype: TEX!; isotypes:ARIZ! CHAPA! ENCB! F! GH! MEXU! MO! NY! US! pf!). 1m tall, well branched, the stems minutely the fresh viscid-puberulent, Leaf blades up to 6 cm long (usually smaller), ovate, cordate, herbage malodorous. and minutely stellate-tomentulose serrate, acute to usually acuminate, densely above and beneath; petioles V2-I times the length of the blades; stipules 2-3 mm long, subulate. Flowers solitary in the leaf axils or sometimes paniculately disposed; 1-2 cm long, slender, viscid; calyx 3-5 mm long, glandular-pubescent, pedicels more than half-divided, the lobes triangular-acuminate, 1-nerved, erect in fruit; a dark center, petals 3-5 mm long, glabrous, corolla without strongly reflexed, Shrubs

or pink almost throughout, distally mucronate; staminal column 2-3 pink-margined mm long, pallid, with a few stellate scabrid hairs near the base, the filaments 1-2 mm long, the anthers yellow; styles and stigmas 5, with slightly darker pigmentation than androecium. Fruits 7-8 mm long, ca. 6 mm in diameter, cylindrical, minutely acute or stellateand glandular-pubescent; 5, 2-3-seeded, mericarps apically with minute whitish pubescence. seeds 2 mm long, dark-pigmented, apiculate; Chromosome number: 2n = 14. occurs in open, arid habitats from central Sonora to mucronatum Abutil?n Puebla (Fig. 6) below 1500 m. It flowers more or less throughout the year. The and the closely allied A. distinctions incanum and A. this species between are discussed by J. Fryxell (1983). fruticosum Examined. Sonora: 223; Fryxell & Whitacre 128; Gentry 19269; Palmer 650, Arguelles 456.? Chihuahua: & Rollins Breedlove 1536; 1281; Sanders 2682; Wiggins Bye 10051. ?Sinaloa: 141, 145, 162; Gentry 223; Gonz?lez-Ortega 5943; Rose Fryxell & Lane 7, 8, 10, 18; Fryxell & Whitacre & Cervantes 14447. ?Zacatecas: Rose 147.?Colima: Mexia 2449.?Jalisco: 489; 13193, V?zquez Palmer 1281.?Guanajuato: 2272.? Puebla: 1133; Fryxell & Lane 192, 194; Miranda Kenoyer Fryxell Specimens

653,

2990.

29. Abutil?n orientale Standley & Steyermark, Guatemala. Ser. 23: 173. 1944.?Type: and Cumbre Blanca de Chiquimula,

Publ. Field Mus. Nat. Hist., Bot. Zacapa: along road between Agua 350-500 m, 15 Oct 1940, Standley

74420 (holotype: F!). Abutil?n

demissum

Mpio.

Pochutla,

32: 263. 1980.?Type: Mexico. Oaxaca: Fryxell, Brittonia 1-2 km al oeste de Puerto Angel, 50 m, 22 Nov 1978,

Koch & Fryxell 78402 (holotype: CHAPA!; isotypes: BH! BM! CAS! CTES! ENCB! F! K! MEXU! MO! NY! TEX! WIS! pf!). subshrubs to 1m tall, the branches often scandent on other Widely branching the stems green, usually with both stellate and simple and drooping, vegetation hairs (1-2 mm long) interspersed. Leaf blades mostly 6-8 (-13) cm long, progres sively smaller toward branch tip, ovate, cordate, crenate, acuminate, slightly dis ca. half the of the blades on colorous, minutely petioles length stellate-pubescent; mm shorter midstem 4-8 leaves, progressively stipules long, subulate. upwards;

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1988

53

MALVACEAE OF MEXICO

1-2 mm Flowers usually solitary in the axils; pedicels 4-15 mm long, articulated below and glandular the flowers, with both stellate hairs; calyx 4 mm long, shallowly 5-lobed, the lobes triangu glandular- and stellate-pubescent, cupuliform, the sinuses broadly rounded; corolla rotate, without a dark center, lar-acuminate, the petals 5-7 mm long, yellow; staminal column 2-3 mm long, pallid, stellate 1.5 mm the filaments long, the anthers yellow; styles 5-6, slender, pubescent, the Fruits 5-6 mm long, about as wide, closely androecium. glabrous, exceeding stellate- and glandular-pubescent; minutely calyx, pseudocapsular, acute 5 and basally, mericarps (-6), apically forming spines, constricted (4-) a the lower cell double divided broad, internally by endoglossum, completely the upper cell loculicidally seeds ca. dehiscent, 2-seeded; ind?hiscent, 1-seeded, 1.6 mm long, glabrous. Chromosome number unknown. of seasonally dry deciduous forest Abutil?n orientale occurs in the understory below 200 m along the Pacific coast from Sinaloa to Oaxaca and probably also in in Guatemala. It flowers in October and Chiapas (Fig. 6); the type was collected invested

by

November.

Specimens Santana 79460,

Examined.

& Cervantes

Sinaloa:

1168.?Oaxaca:

Gonz?lez-Ortega & Lott

Fryxell

6785.?Jalisco: 3376;

Koch

3917.?Colima: Magallanes et al. 79454, & Fryxell 78402; Koch

79538.

30. Abutil?n palmen A. Gray, Proc. Amer. Acad. Arts 8: 289. 1873.?Type: Mex ico. Sonora: Yaqui River, 1869, Palmer s.n. (holotype: GH!; isotypes: NY!

US!). Proc. Amer. Acad. Arts 20: 357. 1885, non aurantiacum S. Watson, aurantiacum Linden ex Turczaninow, Mexico. 1858.?Syntypes: Todos Santos Bay, Jan 1883, Parry s.n.; Tijuana, May Baja California: 1883, Orcutt s.n. (K! MO! PH?). Rose & Standley, Contr. U.S. Nati. Herb. Abutil?n macdougalii 16: 13. pi. 4. Mexico. 1912.?Type: Sonora: Pinacate Mtn, 22 Nov 1907, MacDougal47 Abutil?n

Abutil?n

(holotype: US!). more or less to 1.5 (-2) m tall, the stems sometimes dark-pigmented, mm and sometimes also with hairs 1-2 stellate-pubescent simple long. Leaf blades cm or as as 4-8 often wide sometimes so, ovate, mostly long, long nearly broadly Shrubs

to finely dentate, cordate, 3-lobed, acuminate, weakly coarsely deeply softly tomentose above and beneath, nearly concolorous; petioles half the length of the blades to exceeding the blades, commonly subequal; stipules 6-9 mm long, filiform. Flowers solitary in the leaf axils or forming terminal more or less leafless panicles; pedicels 2-3.5 (-6) cm long; calyx 9-15 mm long in flower to 20 mm long in fruit, the lobes cordate and overlapping; corolla rotate, hirsute, ca. half-divided, a dark center, cm long, yellow-orange, on the petals 2-2.5 pubescent mm of 4 otherwise staminal column claw, margins glabrous; long, glabrous, pallid, ca. 50; styles ca. 10, slender, the filaments 2-3 mm long, the anthers yellow, the stigmas maroon. Fruits 10-12 mm long, 15-16 mm in diameter, glabrous, ca. hirsute; 10, 3-seeded, densely mericarps apically acute or apiculate; seeds ca. 3 mm long, brownish, minutely number unknown. Chromosome pubescent. densely without

Abutil?n

palmen

occurs

in open,

arid habitats

in Baja

California,

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Sonora,

VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

54

Sinaloa,

in coastal to April.

and also disjunctly and California, parts of Arizona m. sea to It 1000 flowers from October from level (Fig. 6)

adjacent

Tamaulipas Specimens

Examined.

California:

Baja

55; Carter

Brandegee

4901; Carter & Ferris

4066; Carter &

Heckard 5747; Collins 184; Fryxell 1984,1986; Gonz?lez 12; Jones 24041;Miller 36;Montufar 34;Moran 11463, 16908, Thomas 8457;

20479, Thome

20617,

(Baja California): 401.? Sonora: Palmer Wiggins

8310.?

31. Abutil?n pt.

Nelson

25123;

57766; Wiggins Moran 7199.?Isla Feiger

Sinaloa:

19728; 5435;

Gentry

& Goldman

7314; Orcutt

256, 9976, 14453, San Pedro M?rtir Johnson Jones

parvulum A. Gray, 1): 21. 1852.?Type:

16012,18112.-Isla

s.n.,

835; Palmer San Angel

90; Parry s.n.; de la Guarda

Johnston 3158; Moran 8806; (Baja California): s.n., 239; Webster 19791; 47; Palmer Fryxell 1228, 3657; LeSueur 286.

208-76; MacDougal 22860.?Tamaulipas:

3 (art. 5, PI. Wright. Smithsonian Contr. Knowl. U.S.A. Texas: calcareous hills of the San Felipe

and San Pedro Rivers, 7 July [1849],Wright 625 [56] (holotype: GH!; isotypes: BM! K! NY! OXF! US!). Note: all six cited specimens bear the number 56, which was a "distribution number" assigned by Gray; Wright's field number for this collection was 625 according toWright's field lists in

Shaw (1987). Perennial

herbs or subshrubs with

the stems sometimes dark trailing branches, and sometimes also with simple hairs 1mm

pigmented, minutely stellate-pubescent long. Leaf blades up to 5 cm long (usually smaller), broadly ovate, cordate, coarsely above and beneath acute, stellate-pubescent dentate, (the surface visible through the length of the blades; ?/2-l times the hairs), essentially concolorous; petioles mm in the 2-4 Flowers leaf subulate. axils; pedicels 1-2.5 cm solitary stipules long, mm ca. half-divided, 3-5 the long, minutely stellate-pubescent, long, slender; calyx lobes triangular, reflexed in fruit; corolla rotate, without a dark center, the petals 4-7 mm

of throughout glabrous (including margins long, more or less pinkish, mm mm 1 2 column the filaments the staminal tall, pallid, glabrous, claws); long, Fruits 8-9 mm anthers few (ca. 15), yellowish; styles 5, subequal to androecium. 5 (rarely 6), 3 long, 7-9 mm in diameter, more or less caskshaped; mericarps or acute 2 mm long, seeds seeded, minutely stellate-pubescent, apically apiculate; = 14. 2n number: Chromosome blackish, minutely pubescent. occurs in northern Mexico to Tamau from Baja California Abutil?n parvulum In the it in in habitats. States arid is found United California, lipas (Fig. 6) relatively and Texas. It apparently flowers more or less New Mexico, Colorado, Arizona, It is an inconspicu throughout the year, probably depending on available moisture. ous plant and thus not often Specimens Chihuahua: Tamaulipas:

Examined.

Baja

8695F; Chiang Garc?a & Garcia

32. Abutil?n percaudatum 438. 1920.?Type:

collected.

California:

Everitt

&

Moran

9839,13641,

VanDevender

s.n.;

24736,24955.?Sonora: Stearns s.n.?Coahuila:

Hartman Fryxell

6

?

3022.?

32.

Annuaire Conserv. Jard. Bot. Gen?ve 21: Hochreutiner, Mexico. San Luis Potos?: Rioverde, 2-8 Jun 1904,

Palmer 19 (holotype: NY!; isotypes:CM! F! GH! K! MO! US!). shrubs branched Widely 6-10 cm long, ovate, mostly

to 2 m

tall with

deeply

cordate,

glandular crenate,

pubescence. acuminate,

Leaf very

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blades sparsely

1988

MALVACEAE OF MEXICO

55

above and beneath; petioles subequal to the blades or somewhat shorter, pubescent Flowers viscid; stipules 3-5 mm long, subulate and inconspicuous. solitary in the leaf axils or in lax panicles, pedicels 1-4 cm long; calyx 6-8 mm long, half-divided, somewhat spread yellowish at base, enclosing base of fruit, the lobes lance-ovate, mm a maroon the with corolla 8-15 rotate, ing; spot at the petals long, yellowish the filaments 2-2.5 mm long, the base; staminal column 3-4 mm long, purplish, anthers orange; styles 5, pallid, slender, the stigmas purplish. Fruits 9-12 mm long, ca. 10 mm in diameter, more or less cylindrical, minutely puberulent (viscid?), 5, 3-seeded, loculicidally dehiscent suppressed); mericarps (septicidal dehiscence scabridulous. Chromo apically acute; seeds 2.8 mm long, dark brown, minutely some number: 2n = 28. Abutil?n percaudatum forest and matorral, occurs at widely grows in deciduous scattered in central and southern Mexico localities (Fig. 6), and has also been collected to January and is the only in Costa Rica. It flowers from October tetraploid species in sect. Oligocarpa ( J. Fryxell 1983). Specimens Veracruz: Chiapas:

Examined.

Ventura

San Luis

10762,

12020,

Potos?: 14653,

Palmer

19.?Hidalgo:

Krapovickas Breedlove 35909;

19192.-Oaxaca:

& Crist?bal

23479.?

Cruz-Cisneros

2535.?

Ton 3144.

33. Abutil?n

brit. 53. 1826. Sida permollis Sweet, Hort. permolle (Willdenow) Enum. Hort. Bot. Berol. Willdenow, pi. hort. berol. 728. 1809.?Type: (holotype: B-W no. 12689! as photo F-9794!). ex A. Gray, Smithsonian Contr. Knowl. 3 (art. Abutil?n peraffine Shuttleworth U.S.A. Florida: Key West, Feb 5, PI. Wright, pt. 1): 20. 1852.?Type:

1846,Rugel95b (holotype: GH?; isotypes:BM, pf!). Shrubs 1-2 m tall, softly stellate-pubescent. Leaf blades mostly 7-12 cm long, ca. half as wide, more or less dis ovate, serrate, cordate, acuminate, deeply whitish colorous, above, densely beneath; petioles sparsely pubescent pubescent Vi-1 times the length of the blades; Flowers stipules 5-7 mm long, lanceolate. or in more leaf in cm the axils than 3 solitary apical panicles; pedicels long; calyx 1 1.5 cm long, soft-puberulent, ca. %-divided, the lobes lance-ovate, the bases over lapping and plicate in bud; corolla rotate, without a dark center, the petals 10-18 mm long, yellow or yellow-orange; staminal column 3-4 mm long, pallid, glabrous, ca. 2 mm long, ca. 50, the anthers yellow; the filaments slender, styles 10-12, pallid. Fruits 8-10 mm long and wide, hirsute, about equaling the calyx, loculici 10-12, 3-seeded, dally dehiscent; mericarps apically acute; seeds 2 mm long, black number unknown. ish, papillate. Chromosome names: sakmizbil (maya), sakxiu or zakxiu (maya). Vernacular Abutil?n is found in the Florida Keys, the West Indies (Bahamas, permolle Cuba, Puerto Rico, etc.), the Yucatan peninsula, Chiapas, and Guatemala (Fig. 7), in thorn forest, evergreen forest, and disturbed vegetation, generally on limestone soils below 1200 m. It flowers principally from December to June, perhaps also at other times. Specimens Yucat?n:

Examined.

Bradburn

Chiapas:

& Dafwin

Breedlove

1281; Gaumer

24178, 357,

30528.?Campeche: 1499,

1506,

1691, 2222,

Goldman 2335,

? 949 472; Lundell 23481; Goldman 561;

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56

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

r

r _\

"

#.*S

i

Abutil?n

Abutil?n,

permolle

pinkavae

r \

Abutil?n

?

?

procerum

Abutil?n purpusii

^o

i^

*

Abutil?n

^^

Abutil?n

reventum

simulans

^*

r \

^ Abutil?n sphaerostaminum

Abutil?n straminicarpum FIG.

Greenman

398,

7. Distribution

of species

Roo:

of Abutil?n.

1067; Yam

pinkavae

37: 285. 1977.?Type: Mexico. Coahuila: Fryxell, Phytologia route 30 at N end of Sierra San Marcos, 14 Aug Basin, & Reeves 13044 (holotype: ASU!; isotype: pf!).

17.? Quintana

Cabrera

& Cabrera

409; Steere

211; Sousa

10901;

T?llez

1443.

34. Abutil?n

Cuatrocienegas 1975, Pinkava

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MALVACEAE OF MEXICO

1988

57

or spreading subshrubs 0.5 m tall, the stems slender, sparsely pubescent also with simple hairs 1-2 mm long, glabrate in with glandular hairs, occasionally cm cordate, indistinctly crenulate age. Leaf blades usually 2 (-4) long, oblong-ovate, or serrulate, acute or obtuse, stellate-pubescent or above and beneath, concolorous mm as as 4-6 the blades; stipules almost long nearly so; petioles long, subulate. Flowers solitary in the leaf axils; pedicels 1-4 cm long, usually exceeding the subtend articulated 3-5 mm below the flowers; calyx 8-17 mm long, with both ing petioles, Erect

the lobes cordate and somewhat plicate at the stellate and simple hairs, ca. ^-divided, a dark center, the petals 14-18 mm long, orange, base; corolla rotate, without on claw, otherwise staminal column 3 mm glabrous; long, stellate-pubescent mm long, orange, the anthers yellow; styles 6 glabrous, orange, the filaments 1-1.5 stigmas. Fruits 10-11 mm long and wide, equaling yellow, with maroon with both stellate and simple hairs 1-2 mm long; mericarps 6-7, 3 seeded, dorsally costate, apically spinescent (the spine 1-4 mm long); seeds ca. 2 mm number unknown. Chromosome long, blackish, muriculate.

7, glabrous, the calyces,

Abutil?n pinkavae Coahuila and adjacent from the Sierra Ju?rez Rinconada Canyon near Specimens 885; Pinkava

Examined. 5193,

5481,

is found in the deserts of the Cuatrocienegas Basin of areas (Fig. 7). The Worthington collection cited below is near Ciudad is from Ju?rez, and the Pringle collection Monterrey.

Chihuahua: 5745,13044.?Nuevo

Worthington Le?n:

35. Abutil?n

procerum Fryxell, Syst. Bot. Farias, 24 Nov pas: near G?mez

14534.?Coahuila:

Gregg

495; Johnston

& M?ller

s.n.

Pringle

10: 269.

1985.?Type: Mexico. Tamauli 939 Richardson 1968, (holotype: TEX!;

isotype: pf!). to 4 m tall, the stems densely and softly tomentose, the hairs up to 0.5 cm 16-20 Leaf blades long. mostly long, broadly ovate, deeply cordate, crenate to subentire, discolorous, acuminate, softly tomentose; petioles almost as long as the blades; stipules ca. 7 mm long, subulate. Flowers solitary in the leaf axils or in small axillary racemes; calyx 1 cm long, softly tomentose and with a few simple hairs (1-2 mm long) at base, more than half-divided, the lobes cordate-ovate, Shrubs

mm

the bases overlapping acuminate, slightly to make the bud 5-angled; corolla rotate, without a dark center, the petals ca. 12 mm long, yellow, copiously hirsute on claw; staminal column ca. 3 mm long, essentially the filaments ca. 2 mm long, glabrous, the anthers yellow, 50 or more. Fruits ca. 1.5 cm long and wide, subglobose becom ca. 12, 3-seeded, the ing oblate, hirsute, blackish at maturity; mericarps oblong, mm 3 and seeds Chromosome acute; apex expanded long, minutely pubescent. number

unknown.

Abutil?n procerum is known only from the type locality (Fig. 7) at 600-800 m in the lower reaches of rain forest vegetation. It appears to be allied with A. permolle, with which it shares a preference for limestone soils. Specimens

Examined.

Tamaulipas:

Diggs

& Nee

2427;

Johnston

36. Abutil?n Contr. U.S. Nati. purpusii Standley, Mexico. Veracruz: Barranca de Tenampa,

12794; Richardson

939.

Herb.

23: 750. 1923.?Type: Jan 1910, Purpus Zacuap?n,

4332 (holotype: US!; isotypes:BM! F! GH!). This content downloaded from 132.211.1.50 on Sun, 06 Sep 2015 20:17:37 UTC All use subject to JSTOR Terms and Conditions

58

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

Shrubs or jsmall trees 2-6 m tall, the stems densely stellate-pubescent (hairs to cm mm 0.5 12-18 (-25) long, ovate, cordate with an long). Leaf blades mostly acuminate, minutely beneath, very open sinus, serrate-crenate, stellate-pubescent sparsely so above; petioles mostly V4-V2 the length of the blades; stipules 5-7 mm solitary in the leaf axils; pedicels at least twice the length long, lanceolate. Flowers cm long, minutely of the subtending petioles; calyx 1.5-2.5 stellate-pubescent, ca. a dark corolla more or less tubular, without Vi-^?-divided; basally brownish, cm center, the petals 3.5-4.5 long, clear yellow, with long glabrous claws; stam inal column 2.5 cm long, glabrous, pallid, the filaments apical, 5-7 mm long, the anthers yellow; styles ca. 10. Fruits 2-2.5 cm long and wide, subglobose, minutely ca. 10, ca. 6-seeded, at maturity; mericarps blackish stellate-pubescent, apically mm number: 2n = 16. Chromosome blunt; seeds 3 long, short-pubescent. Abutil?n purpusii occurs in wet forests of central and southern Mexico (Fig. 7) at 750-1450 m. It flowers principally and in Central America to from November April. Specimens

Jones 67; Richardson Luis 1121, 1403.?San M3592; 124.?Puebla: Guti?rrez 923; Tenorio & Hern?ndez 152; Koch & Fryxell Beaman 690.? Veracruz: 7715; M?rquez 5616; Linden 842; Purpus 4332; Romero 63; Ventura 532, 872, 15052, 16937.? Oaxaca: Matuda 700; Koch & Fryxell 78401 .?Chiapas: 85; Delgado Purpus 6919; Ton 3612. Potos?:

Examined.

Rzedowski

Tamaulipas:

10494.?Hidalgo:

Duke

G?mate

reventum S. Watson, 37. Abutil?n Proc. Amer. Acad. Arts 21: 418. 1886.?Type: Mexico. Hacienda San Jos?, 25 mi S of Batopilas, Aug 1885, Chihuahua:

Palmer56 (holotype: GH!; isotype:US!). to 2 m tall, usually branching only in the Robust erect herbs or subshrubs stems usually reddish or purplish pigmented, inflorescence; minutely glandular to glabrate. Leaf blades mostly 8-10 (-20) cm long, broadly rotund pubescent above ovate, deeply cordate, acuminate, serrulate, obscurely velvety-pubescent and beneath, to the (whitish beneath); strongly discolorous petioles subequal an ample Inflorescence blades; stipules subulate (lost early and seldom observed). cm long, articulated 7-10 terminal panicle rising above the leaves; pedicels 0.5-3.5 mm below the flowers; Vi-Vi stellate-tomentose, calyx 3-6 mm long, minutely corolla rotate, without a dark center, the petals 9-15 divided, the lobes lance-ovate; mm long, pale yellow; staminal column 3 mm long, glabrous, pallid, the filaments 2 mm long, the anthers yellow; styles ca. 10, slender, the stigmas purplish. Fruits 7 10 mm long and wide, globose, minutely blackish at maturity; stellate-tomentose, ca. mm seeds 2.5-3 obtuse; 10, mericarps long, sparsely appressed apically the hairs arranged in reticulate lines. Chromosome number unknown. scabridulous, reventum occurs in open, arid habitats inwestern Mexico Abutil?n from Chihua at to hua and Sonora to Oaxaca it 1000 1500 also is in southern found m; (Fig. 7) to November. Arizona. It flowers principally from August Specimens Chihuahua: Henrickson

Examined.

56.?

1233; Lott & Rose 11157.?Guerrero: 2191.?

Sonora:

3076; Phillips Fryxell s.n.? Durango: Brandegee Estrada 22A; Fryxell B6348; Rose 2463.?Jalisco: et al. 2048; Puga 9327; Rose 2905; Wilbur & Wilbur Palmer

Oaxaca:

Fryxell

Sinaloa:

Fryxell

4163.? 467; Rose 12766; White 2770, Palmer 216.? Zacatecas: 499; Patoni Guzman & Puga 1035,1591, 1599,1600;

2315.?Quer?taro: 349; Rose Arguelles 13695. ?Puebla: 13694 Miranda 2171; Pringle [mixed], 112. 5746; Rose & Hough 4566; T?llez & Magallanes

& Bates

1150, 2569; Pringle

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59

MALVACEAE OF MEXICO

1988

Mexico. simulaos Rose, Contr. U.S. Nati. Herb. 8: 318.1905.?Type: near Cuernavaca, 21 Sep 1900, Pringle 8427 (holotype: US!; Morelos:

38. Abutil?n

isotypes:BM! CAS! F! as photo F-56131! GH! GOET! K! MASS! MEXU! MICH! MIN, MO! MSC! NY! PH! POM! RM! UC! VT! pf !).According to Davis tions case

(1936, p. 187) the type locality is "along the railroad in both direc and about a mile distant in each from the station [in Cuernavaca] . . . ."

Mexico. Abutil?n holwayi Rose, Contr. U.S. Nati. Herb. 8: 318. 1905.?Type: Oaxaca: near city, 10 Nov 1903, Holway 5374 (holotype: US!). 10: 123. Abutil?n durangense Rose & York ex Rose, Contr. U.S. Nati. Herb. Mexico. 1896, Palmer 587 (holotype: US!; pi. 40. 1906.?Type: Durango,

isotypes:BM! F! NY! UC!). some Shrubs 1-2 m tall, the stems often purplish pigmented, viscid-pubescent, times with long simple hairs. Leaf blades mostly 6-10 (-15) cm long, ovate, promi discolorous crenate-serrate, acuminate, nently cordate, markedly (whitish be above and beneath; petioles Vi-3/4 the length of the soft-tomentose neath), minutely blades; stipules ca. 2 mm long, subulate. Flowers solitary in the leaf axils or the more or 1-4 cm long, articulated 3-6 mm less paniculate; inflorescence pedicels cm more below the flowers; calyx 1-1.5 than half-divided, the lobes 9-15 mm long, accrescent in fruit, often flushed rose or wide, broadly cordate and overlapping, a dark center, the corolla rotate, without purplish, minutely stellate-pubescent; staminal column 2-3 mm the petals ca. 1.5 cm long (?), yellow; long, pallid, mm filaments 2 long, the anthers yellow; styles ca. 10. Fruits 1-1.5 cm long and enclosed by and only slightly exceeding the calyx, blackish at wide, stellate-hirsute, ca. 10, 3-seeded, the spine 2-3 mm long; maturity; mericarps apically apiculate, seeds 2.5 mm long, dark-pigmented, number: 2n ==28. glabrous. Chromosome to Abutil?n inwestern Mexico from Durango simulans is found in dry matorral Oaxaca to December (Fig. 7) at elevations up to 1500 m. It flowers from September or

later.

Specimens Rose

&

Painter

Examined. 7510.?

Pringle

8427.?Oaxaca:

Pringle

5780.

Durango: Michoac?n: Conzatti

Palmer Ars?ne 1526; Holway

587.?Jalisco: 1909, 5374;

39. Abutil?n Hochreutiner, sphaerostaminum ?ve 21: 441. 1920.?Type: Mexico. near Zacuap?n, Dec 1906, Purpus

3238,

4582; Fryxell & Bates 2151, 2152; Pringle 8697.?M?xico: 3586.?Morelos: Hinton

Krapovickas

& Crist?bal

23591;

Nelson

1261;

Conserv. Annuaire Jard. Bot. Gen Barranca de Santa Maria, Veracruz: 2234 (holotype: NY!; isotypes: GH!

UC!US!). m tall, the stems sparsely stellate-pubescent Shrubs 0.5-1.5 becoming gla brescent. Leaf blades 5-10 cm long, ca. half as wide, ovate, deeply cordate, or obscurely denticulate crenate to subentire, acuminate, palmately 9-nerved, dis so tomentose above and colorous, beneath; petioles Yi-l times the softly densely mm of 7-8 subulate. Flowers blades; length stipules long, solitary in the leaf axils or more or in cm long, less crowded 0.5-3 racemes; aggregated apically pedicels more articulated 2-5 mm below the flowers, densely so above stellate-pubescent, mm the articulation; 9-12 and calyx long, softly densely pubescent, more than half

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SYSTEMATIC BOTANY MONOGRAPHS

60

VOLUME 25

and overlapping, less than 8 mm wide, more or less the lobes cordate-ovate corolla rotate, without a dark center, the petals 1-1.5 cm long, yellow; acuminate; androecium included, pallid, the staminal column 3 mm long, the filaments 1.5 mm the anthers yellow; styles 9-11. Fruits 10 mm long, 10-12 mm in diameter, long, acute or short-apiculate, seeds 2 mm long, dark 9-11, 3-seeded, hirsute; mericarps divided,

verruculate. Chromosome number unknown. brown, minutely to Veracruz is it is Abutil?n endemic nearly sphaerostaminum (Fig. 7), where found at 600 to 1400 m, flowering May to November. It superficially resembles A. glabriflorum but differs in size and color of the corolla and in the number of carpels in the fruit. Specimens 1403, 9037,

Examined.

9381,12010;

40. Abutil?n

Tamaulipas: Zol?

straminicarpum

Puebla:

[Mpio.

Fryxell

785.?Puebla:

4956.?Veracruz:

Purpus

Purpus

2234,

2235,

8279;

Ventura

1261.

32: 264. 1980.?Type: Mexico. Fryxell, Brittonia San Sim?n] Tlacuilotepec, 6000-7000 ft, Jul

Caltepec,

1909, Purpus 4010 (holotype: F!; isotypes:BM! MO! NY! US!). the stems green, minutely Erect herbs or subshrubs, apparently unbranched, and densely stellate-puberulent, the hairs 0.1 mm long. Leaf blades up to 5 cm long, about as wide, dis acute, densely puberulent, cordate, broadly ovate, dentate, mm or to 1-4 the subulate few blades; colorous; petioles stipules long, subequal a terminal leafless panicle; pedicels 4-10 mm Inflorescence laciniate, caducous. half the length of the calyx 4-7 mm long, ca. half-divided, long; flowers unknown; more or mature less the lower part of fruit or less, minutely pubescent, enclosing the fruit. Fruits 7-9 mm long, somewhat wider; mericarps stellate 9-10, 3-seeded, ca. mm to at acute 0.2 obtuse (with apiculum pubescent, apically maturity long), num straw-colored, loculicidally dehiscent; mature seeds unknown. Chromosome ber

unknown.

is known only from the type collection from the Abutil?n straminicarpum Tehuacan Valley of Puebla (Fig. 7). (The type locality is at 97?30'W, 18?4'N.) Its nearest ally appears to be A. xanti. ex Lindley, Edward's Bot. Reg. App. 39. 1839. Sida striatum Dickson ex Lindley) D. Dietrich, striata (Dickson Syn. pi. 4: 852. 1847.?Type: Dickson inMaund, The Botanist 3: t.144. 1839. venosum Lemaire, FI. Serres 2(3): t.5. 1846.?Type: Abutil?n plate 5, Fl. 1846. Serres, Abutil?n 11: 7. 1885.?Type: thompsonii Andr?, Rev. Hortic. Belge Etrang?re the plate.

41. Abutil?n

Shrubs 2-4 m tall, largely glabrate. Leaf blades mostly 8-15 cm long, truncate to cordate, moderately to deeply 5-parted, 3-lobed the lobes lanceolate, serrate, V2-I times the length of the blades; concolorous; acuminate, essentially petioles Flowers solitary in the leaf axils; pedicels 6-15 stipules 5-9 mm long, lance-ovate. cm long, slender, articulated 2-9 mm below the flowers; calyx 1.5-3 cm long, ca. corolla more or less tubular, half-divided, sparsely and minutely stellate-pubescent; cm long, yellow with dark red claw and conspicuous the petals 3.5-4.5 dark-red

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1988

61

MALVACEAE OF MEXICO

androecium corolla, yellow, the filaments equaling or slightly exceeding ca. 1.5 cm long, ca. 2 cm in Fruits the anthers 8-10. yellow; styles long, less than 2 cm long, proba 8-10, diameter, minutely mericarps stellate-pubescent; number: 2n = 16. Chromosome rounded. 4-6-seeded, bly apically name: monacillo amarillo. Vernacular venation; 3-4 mm

in cultivation and is occasionally Abutil?n striatum occurs in Mexico natural in disturbed sites (Fig. 8). It is native to Brazil and perhaps other ized, generally and is cultivated or naturalized in various parts of Latin parts of South America and elsewhere cf. Mitchell America Australia, 1982). It is sometimes offered (e.g., as "flowering maple." in temperate zone horticulture Specimens Ascensio

Puebla:

Ars?ne

& Kennedy Reisfield

Jalisco:

Examined.

M?xico:

55; Hinton 375,

2331,

1705; Nevling 1336. ?Chiapas:

Ars?ne 798.? s.n.?Quer?taro: 10523; Basile Velasco-Reyes Federal: 29039.?Distrito 4878; Koch 7785; Rzedowski Paray 233, 239.? 14.?Veracruz: & Morales Ballesteros 171; Cant? 68; Foster 5298; Goldman & G?mez-Pompa

Breedlove24421;

285; Vel?zquez 167; Ventura Calzada 3695; Fryxell3204.

9728.?Oaxaca:

42. Abutil?n 23: 326. 1976.?Type: tehuantepecense Fryxell, Madro?o Oaxaca: Cerro de las Animas, 27 Dec 1971, MacDougall H166 NY!; isotypes: F! as photo F-56130! pf!).

Solheim

&

Mexico. (holotype:

Tree to 5 m tall, the twigs (of flowering shoots) densely glandular-puberulent, soon glabrate and striate. Leaf blades mostly 10-12 cm long, 7-9 cm wide, ovate, more or less cordate, weakly denticulate to subentire, acute or acuminate, strongly whitish Va-V2 the length of the discolorous, beneath; petioles felted-pubescent or Flowers borne on more or less leafless paniculate blades; stipules rudimentary. racemiform branches; pedicels 5-12 mm long, densely viscid; calyx 10-12 mm long, corolla viscid, deeply divided, ribbed, the lobes 1-nerved, lanceolate; obscurely without a dark center, the petals 10-11 mm long, violet, reflexed, densely pubes cent on margins of claw and abaxially; staminal column ca. 7 mm long, densely mm the filaments 2-2.5 the anthers yellow; stellate-pubescent, long, purplish, ca. or more the mm less Fruits 10 6, pallid, styles stigmas purplish. long, 10-11 mm in diameter,

ca. 6, 3-seeded, subequal to calyx, stellate-pubescent; mericarps api mature seeds Chromosome number unknown. unknown. cally acuminate; Abutil?n is known only from dry hills in the vicinity of tehuantepecense and La Oaxaca Ventosa, Tehuantepec (Fig. 8), at 600 to 800 m. It flowers from to February, November trees have defoliated with the advancing after the evidently dry

season.

Specimens

Examined.

Oaxaca:

Cedillo-Trigos

707; Koch

& Fryxell

78334;

MacDougall

H166,

H229.

43. Abutil?n thurberi A. Gray, Mem. Amer. Acad. 1854.?Type: Mexico. Sonora: Magdalena,

type:GH!; isotypes:K! MO!).

Arts n.s. 5 (PI. Thurb.): 307. Oct 1851, Thurber 911 (holo

sonorae Wiggins, Pseudabutilon Contr. Dudley Herb. 3: 70.1940.?Type: Mex ico. Sonora: along R?o de Los Alisos, 31 mi S of Nogales, 8 Sep 1934,

Wiggins 7034 (holotype: DS!).

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62

SYSTEMATIC BOTANY MONOGRAPHS

FIG.

8. Distribution

of species

VOLUME 25

of Abutil?n.

Erect or decumbent subshrubs less than 1m tall, the stems terete, green, promi mm long (or sometimes nearly glabrous). Leaf blades up 2-4 the hairs nently pilose, to 6 cm long, almost as wide, ovate, cordate, prominently and coarsely crenate with appressed above beneath and serrate, acute or acuminate, sparsely pubescent the length ca. mm Vi-\ stellate times 2 and occasional hairs hairs; petioles simple long

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1988

MALVACEAE OF MEXICO

63

of the blades; stipules 4-9 mm long, subulate to spatulate, ciliate, persistent. Flowers cm long, slender, pilose, articu solitary or paired in the leaf axils; pedicels 0.5-2.5 lated ca. 2 mm below the flowers; calyx 4-5 mm long in flower, to 7 mm long in fruit, basally rounded, with minute glandular and a few stellate hairs and prominently mm setose the lobes ecostate, hairs 1-3 ovate (the long), ca. half-divided, acuminate; corolla rotate, without a dark center, the petals 4-6 mm long, yellowish; androecium unknown; styles 5. Fruits subequal to calyx, 5-6 mm long and wide, at maturity; mericarps straw-colored 5, usually 3-seeded, with scattered hispidulous, stellate pubescence, the apically spinescent, spine 2-3 mm long; seeds 2-3 mm long, number: 2n = 16. Chromosome sparsely stellate-puberulent. Abutil?n thurberi occurs in dry, open habitats in northern and central Sonora (Fig. 8) and southern Arizona. Specimens Rollins

Examined.

Sonora:

Hastings

& Turner

72-73;

Thurber

911; Wiggins

7034; Wiggins

&

369.

44. Abutil?n 23:

tridens Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 173. 1944.?Type: El Progreso: El Progreso, Guatemala. between Finca Piamonte and the slopes southeast along a small creek, Steyermark 43439 (lectotype, here designated: F-1132511!; F-1132507!). isolectotype:

Shrubs 2-4 m tall, more or less densely pubescent with minute stellate hairs and simple hairs 1-2 mm long. Leaf blades up to 19 cm long, broadly ovate, deeply cordate, acuminate, discolorous, subentire, above, markedly sparsely pubescent densely canescent beneath; petioles usually ca. % the length of the blades; stipules linear or subulate. 10-11 mm Flowers often in 2-4-flowered umbels, long, 10-15 mm peduncles solitary, 11-18 cm long; pedicels 2.5-6 cm long, articulated below the flowers; calyx 1.5-3.5 cm long, densely brown-pubescent, campanulate, cm long, the petals 3-3.5 corolla tubular, half-divided, nerved; prominently the claw); androecium about exserted, orange, glabrous throughout (including cm long, glabrous, pallid, the twice the length of the corolla, the column 4.5-5.5 filaments the anthers yellow; 3-8 mm long, slender, styles 10-13, exceeding cm long, 3-4 cm in diameter, the stigmas purplish. Fruits 2.5-3.5 androecium, truncate with crown of divergent 10-13, 4-6 spines 2-4 mm long; mericarps mm number unknown. Chromosome seeded; seeds 4 long, minutely pubescent. Fig. 9. Abutil?n tridens occurs in Chiapas and Guatemala forests (Fig. 8) in pine-oak between to 1300-2100 m, often reaching cloud forest, and flowers from November March.

Specimens

Examined.

Chiapas:

Breedlove

9384; Laughlin

426; Matuda

89,

1592; Ton

4155.

45. Abutil?n trisulcatum Spec. Nov. Regni Veg. 16: 32. (Jacquin) Urban, Repert. 1919. Sida trisulcata Jacquin, Enum. syst. pi. 26. 1760. Sida triquetra Lin naeus, Sp. pi. ed. ii. 962. 1763, nom. superfl. Abutil?n triquetrum (Lin naeus) Sweet, Hort. brit. 53. 1826. Bastard?a triquetra (Linnaeus) Morales in M. G?mez, Anales Soc. Esp. Hist. Nat. 19: 218. 1890.?Type: [His ?nsula Domingo, Jacquin s.n. (specimen unknown). paniola]

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VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

64

FIG.

9. Abutil?n

tridens.

Flowering

branch

and mature

leaf (Ton 4155)',

flower

(center)

(right) (Breedlove 9384).

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and fruit

1988

MALVACEAE OF MEXICO

65

2:116. 1835. Sida ramosissima ramosissimum K. Presl, Reliq. Haenk. 4: Mexico. D. 1847.?Type: Guererro: 853. Dietrich, (K. Presl) Syn. pi. s.n. as Haenke BM! MO! W PR?; isotypes: photo F (holotype: Acapulco,

Abutil?n

32635!). Abutil?n nealleyi Coulter, Contr. U.S. Nati. Herb. 1: 32. 1890.?Type: Texas: Hidalgo Co., near Hidalgo, Nealley 50 (holotype: US!).

U.S.A.

herbs or shrubs 1-2.5 m tall, the stems prominently trisulcate when Robust in inflorescence. Leaf blades young, minutely usually viscid especially puberulent, 6-11 cm long, ovate, cordate, crenulate-serrulate, acuminate, mostly minutely velutinous above and beneath, petioles Va-Va times as long as slightly discolorous; the blade (sometimes longer on midstem leaves); stipules 2-4 mm long, subulate. Flowers usually in lax terminal panicles; pedicels 1-3 cm long; calyx 3-4 mm long, corolla often with a dark center, the lobes lance-ovate, half-divided, puberulent; the petals 4-6 mm long, yellow, often reflexed; androecium pallid, about equaling petals, the staminal column 2-2.5 mm long, the filaments 0.5-1.5 mm long, the anthers yellow; styles 5. Fruits 6-8 mm long, 5-8 mm in diameter, cylindrical but near center, minutely usually slightly constricted loculicidally stellate-puberulent, dehiscent dehiscence 5, ca. 3-seeded, (septicidal mericarps suppressed); apically seeds 2 mm long, minutely number: 2n = 14. Chromosome apiculate; pubescent. names: amantillo, malva cueste, pelotazo, Vernacular sak pelotazo manso, misbil, sak-xiu (maya), tronadora. trisulcatum occurs from Texas to Nicaragua Abutil?n and in the West Indies. It or is found widely in Mexico in dry matorral (Fig. 8), generally at low elevation to March. and flowers from September disturbed vegetation, It is a common road side weed of robust growth and is easily recognized by its prominently triquetrous stems.

Specimens Examined. 254.? Chihuahua: Sonora: 144, Pennington Bye 7080.?Sinaloa: Fryxell & Salazar 4; Palmer 161, 544; Gentry 5855, 6441; Jones 22397; Narv?ez-Montes 7119; Gonz?lez-Ortega Le?n: 25j?52.-Nuevo Dorr 2611.? 13649, 5023; Wiggins 1700; Rose 13875, 14546, 14733; White Tamaulipas: 17M106; Dunn & Harmon 17631; Fryxell 512, 514, 3701; Hern?ndez 799; Johnston Barkley Ferris 5562; Gentry 19646.?Jalisco: 5746; Martinez 622; Nee 22191; Palmer 160a; Puig 3347.? Nayarit: Cobi?n 238; Lott et al. 1567; Pringle 4591; V?zquez Emrick 777; Wilbur & Wilbur 2216, 2386.?Colima: 6086.?San Potos?: Palmer Luis 7472.?Michoac?n: Dieterle 20; Ferris 1037; Rzedowski 3148; & Spetzman s.n.; Koch et 210; Haenke Dorantes 662; Chiang 190; Cox MC-558; 1013, et al. 92; Higgins 3707, 5052; Fryxell 524, 713; G?mez-Pompa 4729; Hern?ndez 2571; Lot 482; M?rquez 1093; Nee et al. 22976; Ross 14-58; V?zquez-Yanes 651; Ventura 2658, 2703; Worth 114; Yong 44.? Oaxaca: Goldman 480. ?Yucat?n: 2091, 23554. Gaumer954,1573, Vargas 4L? Campeche:

McVaugh al. 79263;

17995.?Morelos: 50.?

Palmer

46. Abutil?n

9126.? Guerrero: Pringle & Dorantes Veracruz: Acosta

umbellatum

Linnaeus,

bellata"

Syst.

(Linnaeus) nat.

(holotype:

ed.

x.

Sweet, 1145.

Freeland

Hort.

1759.?Type:

brit.

53.

Jamaica,

1826. Sida Browne

umbellata s.n.,

"um

LINN-866.16!).

Shrubs 1-2 m tall, the stems densely stellate-pubescent. Leaf blades mostly 6 or weakly 12 cm long, broadly ovate acute or 3-lobulate, cordate, serrate, more so beneath than somewhat acuminate, minutely above, pubescent, densely discolorous; petioles Vi-l times the length of the blades; stipules 8-10 mm long.

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66

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

into a terminal these often aggregated umbels, cm 1-5 inflorescence, long, pedicels usually 1-2 cm long; compound peduncles mm the lobes triangular, prominently 6-8 half-divided, rounded, calyx long, basally a dark center, the petals 6-8 mm 3-4 without the hairs corolla rotate, hirsute; long; mm long, yellowish; staminal column 3-4 mm long, the filaments 1-2 mm long, the anthers yellow; styles 6-8. Fruits 5-9 mm long and wide, hirsute, with divergent Flowers

in few-flowered

axillary

seeds 2 6-8, 3-seeded, lacking endoglossum, apical spines 2-4 mm long; mericarps mm long, papillate. Chromosome number: 2n = 16. names: pelotazo peludo, sak-xiu. Vernacular occurs in dry forested areas and in disturbed sites in Abutil?n umbellatum and also in the West and Indies in and Central various parts of Mexico (Fig. 8), it flowers more or less throughout the year. South America; Specimens

Examined.

Sinaloa:

983.?Tamaulipas:

Gonz?lez-Ortega

Fryxell

510,

652,

1085,

3664,

3666, 3686, 3695; Fryxell & Bates 813, 819; Fryxell & Magill 2250, 2349; Johnston 5482; LeSueur 280; Nee

Palmer

22232;

Chiang

269; Koch

1148; Yucat?n:

Fryxell

78253, Darwinetal.

47. Abutil?n

Luis

411?Sax

vexillarium

Morren's

Potos?:

Nee

22289.?Jalisco:

Rose

14296; Ventura 7103, 8273, Purpus 4673. ?Campeche: 78289, 78342; Rose & Hough 2176b; Gaunter 1179; Valdez30.

& Bates

858;

E. Morren,

Belgique

& Painter

14: 289. f.16.

Hort.

7355.?Veracruz:

9425, 14585. ?Oaxaca: Stewart & Percival

Fryxell 84-18B.?

1864.?Type:

plate.

Erect shrubs 1m tall, the stems very sparsely and minutely stellate-pubescent to glabrate, the young stems often purplish. Leaf blades 3-6 cm long, 2.5 (-3) cm acute wide, triangularly and narrowly ovate, more or less cordate, crenate-serrate, or acuminate, ca. Va the length of the concolorous; petioles glabrous, essentially blades; stipules 6-9 mm long, 5 mm wide, sessile, broadly ovate and amplexicaul, caducous. Flowers often purplish, solitary in the leaf axils; subequal to petioles, cm 1.5-2 the slender, petiole, pedicels long, exceeding purplish; calyx 1.5-2.5 cm the lobes 2-4 mm and tubular inflated, 5-lobed, apically long; long, purplish, corolla tubular, the petals 2.5-3 cm long, 8-12 mm wide, yellowish; androecium the filaments 4-5 mm long, more or less erect, the exserted, manifestly purplish, included in androecium. Fruits unknown. anthers purple; styles ca. 8 (?), purple, Chromosome number: 2n = 16. is a South American in the Abutil?n vexillarium species that iswidely cultivated a as as a or is It shrub sometimes sometimes encountered tropics pot-plant. garden inMexican gardens though seldom in herbaria. Specimens Ortega

Examined.

1476; Ventura

Hidalgo:

9396.?Without

Hern?ndez Locality:

et al. 6913.? Berlin

Veracruz:

57("planta

& G?mez-Pompa Nevling exotica cultivada").

281 ;

48. Abutil?n wrightii A. Gray, Boston J. Nat. Hist. 6: 162. 1850.?Type: U.S.A. N center is 3-4 mi of Texas: Maverick Rio the of Creek Seco Co., [Seco

Eagle Pass], 4 Jul 1849,Wright 594 [54] (holotype: GH!; isotypes:BM! K! MO! NY! OXF! PH! UC! US!). or ascending perennial herbs or subshrubs less than 0.5 m tall, the Procumbent stems slender, sometimes purplish, with minute stellate pubescence and usually also

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1988

MALVACEAE OF MEXICO

long simple hairs ca. 1 mm long. acute deeply cordate, dentate, or whitish above, sparsely pubescent cence beneath; often petioles longer filiform. Flowers in the leaf solitary articulated 2-6 mm below the flowers;

with

ovate,

67

Leaf blades 1.5-4 cm long, about as wide, to obtuse, markedly discolorous, green and with soft matted gray-green densely pubes than the blades; stipules 3-4 mm long, 1-3 (-4) cm long, slender, axils; pedicels calyx 10-15 mm long in flower to 20 mm in

fruit, densely stellate-pubescent, deeply divided, the lobes cordate and overlapping at base, acuminate; corolla rotate, without a dark center, the petals 14-18 mm long, on margin of claw, otherwise glabrous; staminal column ca. pale yellow, pubescent 3 mm long, glabrous, pallid, the filaments ca. 2 mm long, the anthers yellow; styles the androecium, 6-9, pallid, glabrous, exceeding stigmas maroon or pallid. Fruits 1 cm long (shorter than calyx), slightly wider, densely hirsute with both stellate and seeds 2.6 mm long, 6-9, 3-seeded, simple hairs; mericarps apically acuminate; muriculate. Chromosome number: 2n = 14. Abutil?n wrightii is found in dry, open habitats in northern and eastern Mexico It (Fig. 8) and in southern and western Texas and adjacent parts of New Mexico. flowers

the year.

throughout

Specimens Henrickson

Examined.

Chihuahua:

14897; Marsh

525;

Purpus

Pringle

35,

574.?Coahuila:

4901; Waterfall

15809; White

Dorr 2227; 1389.?Nuevo

Fryxell Le?n:

1474, Dorr

3021; 2481;

Flook & Spears 59; Frye & Frye 2370; Fryxell 1325,1330; Fryxell & Kirkpatrick 2434, 2463;May 8910.? Tamaulipas: 4384G,

Berlandier Luis

4893B.?San

3046; Fryxell & Bates 3031; Chapman Potos?: Rzedowski 24633.? Veracruz:

947; Fryxell P?niche 33.

& Magill

2257;

Johnston

49. Abutil?n xanti A. Gray, Proc. Amer. Acad. Arts 22: 301. 1887.?Type: Mex ico. Baja California: 1859-Jan 9 1860, Xantus Cape San Lucas, Aug

(holotype: GH!; isotypes:NY! US!). Abutil?n

carterae Kearney, Madro?o 13:114.1953.?Type: de Tabor, Sierra de la Giganta, W

fornia: Arroyo

Mexico. Baja Cali of Puerto Escondido, 3

Dec 1950, Carter & Kellogg 2871 (holotype: UC!; isotypes: CAS! DS! MEXU! "isotype"

US!). Note: by Kearney

The CAS,

DS,

and MEXU

specimens

are labelled

(THK).

Erect shrubs 1.5-3 m tall, the stems green, densely and softly tomentose (hairs 0.5 mm long or longer). Leaf blades mostly 8-15 cm long, somewhat narrower to somewhat wider than long, broadly ovate or shallowly 3-lobed, deeply cordate, serrate to crenate, acuminate, discolorous, sharply indistinctly softly tomentose above and beneath; often the blades; stipules 5-10 mm long, petioles equaling to filiform. an ample subulate Inflorescence terminal panicle overtopping the cm or mm 1 6-8 leaves; pedicels less, pubescent; (-3) long calyx long, stellate the lobes ovate; corolla rotate, without a dark center, the Vi-ys-divided, pubescent, or greenish white, pubescent on margins of claw, petals 7-9 mm long, yellowish mm otherwise staminal column 3 glabrous; long, glabrous, pallid, the filaments 2 mm the ca. anthers 8, slender, pallid, notably long, styles yellow; exceeding androecium. Fruits 8-11 mm long, 9-13 mm in diameter, the calyx, exceeding on and suture with hairs ca. 8, 3 of stellate-pubescent dehiscence; mericarps simple or acute or at dark brown blackish seeds 3 mm seeded, apically apiculate, maturity; = 14. dark brown. number: 2n Chromosome long, banded-muriculate,

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SYSTEMATIC BOTANY MONOGRAPHS

68

VOLUME 25

xanti occurs in the deserts of the southern part of the Baja California Abutil?n and also in coastal Sonora and Sinaloa (Fig. 8). peninsula Specimens 1990; Gentry 3054.

Examined. 4111; Moran

Baja

California:

7331;

Thomas

Carter 2055, 2871, 4077, 4087, 9.?Sonora: 7780; Xantus Wiggins

4258, 5177; Fryxell 1982, 6051.?Sinaloa: Fryxell

Alcea Linnaeus, Sp. pi. 687. 1753; Gen. pi. ed. v. 307. 1754.?Lectotype, 1929: Alcea rosea Linnaeus. nated by Hitchcock and Green,

desig

or perennial erect, annual, biennial, herbs, with stellate pubescence Mostly sometimes mixed with long simple hairs, sometimes glabrate. Leaves long-petio ovate to lobed or deeply palmately the blades late, suborbicular, angled, weakly acute to obtuse, Inflorescence lacking foliar nectaries. parted, crenate or dentate, or spiciform, usually leafless; involucel shorter than to equaling calyx, connate below; calyx 5-lobed, more or less pubescent; petals usually 5-11-parted, large (more than 3 cm) and showy, apically notched, pink, white, purple, or yellow, at included, glabrous, antheriferous spreading to form a rotate corolla; androecium more zone than 15, the stigmatic apex, usually pallid and compact; styles introrsely decurrent. than 15 mericarps; Fruit a diskshaped schizocarp of more mericarps and circular with a prominent ventral notch, ind?hiscent, laterally compressed racemiform

smooth, wrinkled, winged, furrowed, etc.; seeds glabrous or pubescent, variously number: x = 21. solitary, glabrous or pustulate. Base chromosome Don (1831) states "from alke, remedy; the Alcea of the ancients Etymology. . . ."On the other hand, was a kind of mallow. Iljin (in Shishkin, 1974) says "from Greek alee, strength, probably referring to the vigorous growth of certain species, particularly A. rosea L." a genus of the Middle East and southwestern Asia, contains about 60 Alcea, species. One

of these

(Alcea

rosea) has become

almost

cosmopolitan

in cultivation.

References: Baker (1890), Kearney (1951a), Zohary (1963a, 1963b), Riedl (1976). rosea (Linnaeus) Cavanilles, 50. Alcea rosea Linnaeus, Sp. pi. 687. 1753. Althaea "rosea 1" ex Hort. Upsal. Diss. 2: 91. 1786.?Type: (holotype: LINN 869.1 !).A photo of the type is reproduced by Molinari (1965, plate 7). Althaea sinensis Cavanilles, Diss. 2: 92. t.29.f.3. 1786.?Type: cult. Roy. Bot. Gard. Paris (P?). Althaea mexicana Kunze, Linnaea canis" (specimen unknown).

19: 404.

1845.?Type:

"e seminibus

mexi

Leaf blades 7-15 cm long, 1-2.5 m tall, roughly pubescent. Strict perennial orbicular to 5-7-angled, cordate, crenate, roughly pubescent; petioles equaling or into the blades. Flowers terminal, leafless, exceeding showy, subsessile, aggregated involucel of 6-7 broadly triangular bracts, these connate spiciform inflorescences; the involucel; corolla 8-10 cm in diameter, exceeding basally; calyx tomentose, sometimes double, in a wide variety of colors; styles 20-40. Fruits disciform, pubes cent, enclosed by calyx; mericarps 20-40, 6 mm long, horseshoe-shaped, dorsally seeds solitary, minutely channelled and winged, pubescent; mosome number: 2n = 42. Fig. 10.

pubescent

in part. Chro

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69

MALVACEAE OF MEXICO

1988

FIG.

10. Alcea

rosea.

Flowering

branch

and mature

leaf.

(Hill

10371.)

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70

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

names: altea, amapola grande, malva rosa, vara de San Jos?. Vernacular as an ornamental. in cultivation Alcea rosea is almost cosmopolitan It is also as a source of mucilage for medicinal use, and as a grown as a source of dyestuffs, source of oil from the seeds. This species originated on the Aegean Islands and the to China), attributed sometimes erroneously adjacent Balkan Peninsula (although the world as an ornamental, in the tropics at but is now widely dispersed throughout and in the temperate and even boreal zones. In cultivation it is higher elevations It may be seen in gardens in with a complex ancestry. undoubtedly polymorphic, most parts of Mexico but is evidently seldom collected for the herbarium.

Specimens

Examined.

10532.?Morelos:

Sonora:

Torres

220;

Pennington

V?zquez

965,

98.?Chihuahua: 1014.?

Guerrero:

29.? Quer?taro: Ars?ne Pennington Hinton 6127.?Yucat?n: Guti?rrez

Rivasll.

gen. et stat. nov. Fryxell & Bates, (Hochreutiner) Krapovickas, Pseudabutilon Jard. Allosidastrum Annuaire Conserv. Hochreutiner, subg. Bot. Gen?ve 20: 118. 1917.?Type: Allosidastrum interruptum (Balbis ex de Candolle) Krapovickas, & Bates. Fryxell

Allosidastrum

Shrubs 1-5 m tall with pubescence of both stellate and simple hairs, sometimes or ovate, the blades viscid. Leaves lanceolate crenate, cordate, long-petiolate, or less leafless Flowers in terminal, more acuminate, lacking foliar nectaries. involucel absent; calyx basally rounded, twisted in bud, the panicles or racemes; lobes divergent; petals white to yellow, with or without a dark tips of the acuminate at apex; styles 5-9, the basal spot; androecium included, the column antheriferous of 5-9 oblate, stigmas capitate. Fruits schizocarpic, mericarps; mericarps essentially ind?hiscent, more or less rounded dorsally, with relatively lacking fragile walls an apical beak, with a small endoglossum; with or without seeds ornamentation, number: x = 7. solitary, glabrous. Base chromosome From the Greek allos, other, plus the generic name Sidastrum. Etymology. Allosidastrum includes four Neotropical species, three of which occur inMex ico (Fig. 12). References: Hochreutiner (1917), Krapovickas, Fryxell & Bates (1988). Key in an interrupted

1. Flowers floral 1. Flowers mm wide; 2. Petals long, 2 mm

raceme

more frequently a central depression.

bracts

around

to the Mexican

in a terminal

Species

of Allosidastrum

of axillary glomerules, the inflorescence sometimes 1 mm wide; with fruits of (7-) 8-9 mericarps

than 52.

much-branched

A. interruptum. not glomerulate; panicle, with apical beak or beakless.

fruits of 5-7 mericarps, 5-8 mm long, yellow or yellow-orange, and usually also with stellate-pubescent

with

floral

a conspicuous

bracts basal

linear,

branched; short spurs less than 1

spot; calyx 6-8 mm staminal column

hairs;

long (1-2 mm) simple a the filaments often purplish; scabrid fruits without long, densely (hairs 0.5 mm), 51. A. the dorsal wall flattened. hilarianum. beak, of 6-7 mericarps, or pale yellow 2. Petals 4-5 mm throughout (or rarely with an indistinct basal long, whitish mm and 3-5 sometimes with glandular hairs; staminal calyx long, spot); stellate-pubescent column 1.5 mm long, sparsely the filaments stellate-pubescent (hairs 0.1-0.2 mm), pallid; fruits beaked,

of 5-6

(-7) mericarps,

the dorsal wall

rounded

(the fruits thus lobulate). 53. A. pyramidatum.

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71

MALVACEAE OF MEXICO

1988

hilarianum 51. Allosidastrum Fryxell & Bates, comb. nov. (K. Presl) Krapovickas, 2: 107. 1835, non Sida hilariana Sida hilariana K. Presl, Reliq. Haenk. 1842.?Type:

Walpers,

(lectotype,

"in

Mexico,

designated

s.n.

Haenke

occidentali,"

parte

et al., 1988: PR!;

by Krapovickas

BM!

isolectotypes:

LE! MO! PR!). Mexico. Guer 2: 106. 1835.?Type: Sida kunthiana K. Presl, Reliq. Haenk. s.n. (holotype: rero: Acapulco, Haenke PR!; isotypes: MO! PR! W as photo F-32640!). var. Bot. voy. Sulphur 69. 1844. Sida dumosa Sida glanduligera Bentham, 30: 296. Nicara Bot. 1892.?Type: E. G. J. Baker, (Bentham) glanduligera gua. El Realejo, Sinclair s.n. (holotype: K!). cin?rea

Sida

E.

G.

ex

Baker

Contr.

Rose,

U.S.

Nati.

3:

Herb.

311.

1895.?

Nayarit: Tepic, 5 Jan-6 Feb 1892, Palmer 1835 (holotype: BM! F! GH! K! NY! US!). BM!; isotypes: Annuaire Conserv. Jard. Bot. Gen?ve Pseudabutilon langlassei Hochreutiner, Guerrero: rives du Rio Tecpan, 28 Jan 20: 120. 1917.?Type: Mexico. as K! P, G 817 G, F-23753!; 1899, Langlass? isotypes: photo (holotype: Type: Mexico.

US!). Shrubs 2-5 m tall, the stems minutely rescence with long simple hairs (0.5-1.5 mm 8-14

mostly

cm

long,

ovate,

truncate,

stellate-puberulent, long) and glandular

serrate,

acuminate,

glabrate

also in the inflo hairs. Leaf blades above,

minutely

concolorous; beneath, petioles Vi-\ times the length essentially stellate-puberulent linear stipuliform floral in terminal panicles with numerous of the blades. Flowers 1-5 (-12) mm long; calyx 6-8 mm long, with bracts less than 1mm wide; pedicels stellate hairs and usually also with simple hairs 1-2 mm long; petals 5-8 mm long, with a dark red spot at base; staminal column 2 mm long, yellow or yellow-orange mm scabrid (hairs 0.5 long), the filaments often purplish; styles 6-7. Fruits 5 mm in in the of 6-7 mericarps, diameter, beakless, mericarps apically stellate-pubescent; form of a semicircular wedge with relatively flat dorsal walls; seeds 1.5 mm long. number: 2n = 14. Fig. 11. Chromosome Allosidastrum hilarianum forest, oak forest, and evergreen grows in deciduous m to in and southern Mexico forest at elevations 1300 central up (Fig. 12) and in to March. as far south as Costa Rica. It flowers from October Central America Examined.

Specimens 24494.?Nayarit.

1513; Miller 12848.?M?xico: Hinton

& Koelz Mexia

8871;

Sonora:

495.?

Palmer

2240, 8672, Purpus Breedlove & Almeda

52. Allosidastrum Bates,

14200, 56893;

2647.?Guerrero:

Sinaloa: Breedlove 5653.? Durango: Gentry 1835; Rose 3131, 14179, 14438.?Jalisco: McVaugh Hinton 1249.?Michoac?n: 12846, 3174; Palmer hangman Croat s.n.; Hinton 817; 5684; Langlass? 45759; Haenke

Sanders

563; Lehto 24234; et al. 257.?Colima:

Lamb

1028.?

Palmer

s.n.? Veracruz: Guti?rrez Salazar 1142; Nee 16861.?Chiapas: 12191, 15005, 15724, 14247; Ventura 5959. Croat 46282; Matuda 17356; Miranda Puebla:

& Taylor Breedlove

29594; 24494;

interruptum Fryxell & (Balbis ex de Candolle) Krapovickas, Prodr. 1: 464. comb. nov. Sida interrupta Balbis ex de Candolle, G-DC

as

photo F-7997!; isotypes: F! US!). Annuaire Jard. Bot. Gen?ve Conserv. smithii Hochreutiner, Pseudabutilon Santa Marta, 119. 1917.?Type: Smith 489 (holotype: Colombia.

G;

1824.?Type:

Colombia.

Sanctae

Martae,

Bertero

s.n.

(holotype:

isotypes:A! BM! BR! F! GH! K! MICH! PH! US!). This content downloaded from 132.211.1.50 on Sun, 06 Sep 2015 20:17:37 UTC All use subject to JSTOR Terms and Conditions

20:

72

SYSTEMATIC BOTANY MONOGRAPHS

FIG. above),

11. Allosidastrum

mericarp

(right,

below).

hilarianum. (Breedlove

Flowering

and

fruiting

branch,

VOLUME 25

flower

(left),

24494.)

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fruit

(right,

73

MALVACEAE OF MEXICO

1988

y

.^

-...

Allosi((astrum hilarianum

Allosidastrum interruptum

?

l\ .s

Allosidastrum^?

/

.

?

pyramidatum

Allowissadula

chiang i i \ rv~?

^

Allowissadula floribunda

Allowissadula glandulosa

Allowissadula holosericea

Allowissadula lozanii

FIG.

12. Distribution

of species

of Allosidastrum

and Allowissadula.

the stems minutely 1-1.5 m tall, little branched, Subshrubs scurfy-puberulent to glabrate. Leaf blades mostly 6-11 cm long, broadly ovate (narrower upward), truncate

or

slightly

pubescent; petioles raceme occasionally

cordate,

crenate-dentate,

acuminate,

minutely

and

obscurely

an interrupted Inflorescence length of the blades. 1-4 in flowers branched; axillary glomerules; paniculately

1/2-2/3 the

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74

VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

1-5 mm long, with floral bracts lance-linear, often more than 1mm wide; pedicels mm 4-6 densely calyx petals 4-5 mm long, yellow, occa long, stellate-puberulent; the sionally with a basal spot; staminal column 1.5 mm long, sparsely pubescent, mm long; styles (7-) 8-9. Fruits 4.5-5 mm in diameter, minutely hairs 0.1-0.2 of (7-) 8-9 mericarps, with a ring of short divergent spurs stellate-pubescent, around a central depression; number unknown. seeds 1.8 mm long. Chromosome Allosidastrum interruptum is known from western Mexico (Fig. 12) in season forest below 200 m, often in shade, and also from Nicaragua and ally dry deciduous toMarch. It flowers from October Colombia. Specimens Michoac?n:

Examined.

Jalisco:

3179.?Oaxaca:

Dieterle

Iltis & Fryxell

? 25061 29144; Magallanes 3926; McVaugh V?zquez et al 78296; Koch & Fryxell & Lott 3377; Koch 78344,

78406.

53. Allosidastrum pyramidatum (Cavanilles) Krapovickas, Fryxell & Bates, comb, nov. Sida pyramidata Diss. 1: 11. t.l.f.10. 1786.?Type: Cavanilles, s.n. (holotype: P-JU no. 12272!). Santo Domingo, Desportes [Hispaniola] Sida dumosa Swartz, Prodr. 101.1788.?Type: Jamaica, Swartz s.n. (holotype: S!; isotypes: BM! B-W no. 12713!). Sida leiophylla s.n. (B?). Sida dumosa ela,

Sprengel,

Syst. veg. 3: 121. 1826.?Type:

Puerto

Rico,

Bertero

var. fendleri E. G. Baker, J. Bot. 30: 296. 1892.?Type: Venezu coloniam Tovar, 1854-1855, Pendler 70 (holotype: K!; isotype:

prope

GH!). Shrubs up to 2 m tall, the stems minutely Leaf blades stellate-puberulent. mostly 6-11 cm long, broadly ovate, truncate or slightly cordate, serrate or crenate, above and beneath or glabrate above, es acuminate, minutely stellate-puberulent Vz-Va the length of the blades. Flowers in much sentially concolorous; petioles branched terminal panicles with reduced leaves and linear stipuliform bracts less than 1 mm wide; pedicels 1-4 mm long; calyx 3-5 mm long, with stellate and or pale yellow sometimes short glandular hairs; petals 4-5 mm long, whitish throughout (or rarely with an indistinct reddish spot at base); staminal column 1.5 mm long, sparsely stellate-pubescent (hairs 0.1-0.2 mm), pallid throughout; styles 5-6 (-7). Fruits 3.5-4 mm in diameter, beaked, lobulate, stellate-pubescent; 5-6 (-7), with rounded dorsal walls; seeds 1.2 mm long. Chromosome mericarps number: 2n = 14. name: thipom (huastecan). Vernacular occurs in various parts of Mexico Allosidastrum pyramidatum (Fig. 12), espe in seasonally forest and in disturbed sites, cially eastern Mexico, dry deciduous usually below 500 m but occasionally higher. It also occurs more or less throughout to Colombia, the West Indies and through Central America Venezuela, Surinam, to April. It flowers from November Brazil, and Bolivia. Specimens

Examined.

Tamaulipas:

Duke

3630;

Jones

71.?Jalisco:

Iltis & Nee

1645?San

Luis

Potos?: Fryxell 1021; Fryxell & Anderson 3458; Hern?ndez-Maga?a ? Tenorio 6992; King 4381; Shiller 3068.?Guerrero: Jezepczuk

Paray

1361; Purpus

1795?Puebla: 15040;

Smith

Goldman 1028; V?zquez

44.?Veracruz: 444; Ventura

Chiang349; 4556; Worth

Fryxell & Bates 941; 36.? Oaxaca: Koch &

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1988

75

MALVACEAE OF MEXICO

78426; Fryxell 54540; Matada

859; Reko 3688.?Chiapas: Seler & Seler 1924; Ton 3189.?

Makrinius 2697;

Allowissadula

Bates, Gentes (Rose) Bates.

Herb.

Breedlove Quintana

11: 337.

Roo:

32573, Bye

1978.?Type:

33984; Breedlove et al. 11981.

Allowissadula

& Davidse

lozanii

often with or spreading subshrubs with soft or rough stellate pubescence, more or less 3 or ovate blades the viscid. Leaves sometimes petiolate, simple hairs, or foliar acute nectaries; acuminate, dentate, lobed, cordate, stipules lacking involucel ab filiform or subulate, caducous. Flowers usually in terminal panicles; or white (fading the lobes lanceolate; petals yellow-orange sent; calyx 10-nerved, column staminal otherwise the claws included, glabrous, ciliate, glabrous; rose), Erect

at apex,

frequently orange; styles 5, pallid, the anthers numerous, slender, the stigmas capitate. Fruits schizocarpic capsular), (but often functionally the lobes of the calyx; mericarps the lobes opposite 5, rounded 5-lobed, pubescent, or apiculate or less constricted also with an below (sometimes apically, more seeds the lower two the 1-seeded, cells, upper 2-seeded; making endoglossum) = 8. x number: Base chromosome reniform, glabrous. derives from allos, other or different, The name Allowissadula Etymology. name Wissadula. with the generic compounded and the taxo the name Pseudabutilon Nomenclatural surrounding problems to it necessary of this genus made nomic problems related to Fries's conception to reduce and of Wissadula the Fries's Pseudabutilon, genera conception fragment to synonymy and to propose a the name Pseudabutilon (see following paragraph), antheriferous

new name

for the group treated here (Bates 1978a). Hutchinson spicatum (H. B. K.) R. E. Fries as (1967) designated Pseudabutilon R. E. Fries. I believe this choice is based on a the lectotype species of Pseudabutilon on the basis of the genus Pseudabutilon Fries (1908) established misinterpretation. sect. Wissadulastrum two previously sections of the genus Wissadula: described E. G. Baker (type: W. scabrd). Schumann (type: W. spicata) and sect. Abutilastrum Fries did not elevate either section to generic rank but coined the new name, either sec for the combined taxon; nor did he explicitly designate Pseudabutilon, he did segregate is open. However, tion as "typical." Therefore, the choice as is indi to Abutil?n, because of its resemblance from Wissadula Pseudabutilon and in his statement cated in his formation of the name Pseudabutilon (op. cit. p. . . . einen neuen new und wollen wir "muss that the genus 15) [Namen] erhalten, R. E. Fr. vorschlagen, damit ihren Platz hier als solchen den Namen Pseudabutilon . . . ." It therefore seems best in auch in der N?he der Gattung Abutil?n andeuten, as the nominate accord with Fries's intent to consider sect. Abutilastrum section, of the sectional name and because of the morphology both because of the meaning is here of the plants included in the section. The lectotype therefore, species, to be Pseudabutilon R. E. So Fries. scabrum considered (K. Presl) typified, in the present treatment. is a synonym of Abutil?n Pseudabutilon two is a principally Mexican Allowissadula genus of nine species (Figs. 12,13), of which also occur in Texas. Reference:

Bates

(1978a).

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76

SYSTEMATIC BOTANY MONOGRAPHS

to the

Key

Species

VOLUME 25

of Allowissadula

to stellate with and glandular simple hairs 1-1.5 mm long in addition pubescence; stipules ca. 8 mm long. 2. Leaves up to 7 cm long; calyx 10-12 mm simple, long, the lobes acuminate-apiculate; lower seed cavity. 59. A. endoglossum completely covering pringlei. 2. Leaves more or less 3-lobulate, often more than 10 cm long; calyx 7-10 mm long, the lobes or absent. acuminate-caudate A. sessei. 62. ;endoglossum rudimentary

1. Calyx

1. Calyx

viscid

stellate-pubescent,

long. 3. Calyx

6-8 mm

4. Petals

14-18

or not,

long simple

hairs

absent;

stipules

not more

long; fruits 5-8 mm mm long (more

long, each mericarp dorsally rounded or apiculate. than twice the length of the calyx); calyx viscid; endo 55. A. floribunda. absent. glossum than twice the length of the calyx); calyx stellate-pubescent,

8-10 mm long (less hairs ;endoglossum 61. A. rosei. lacking glandular present. 3. Calyx 7-14 mm long; fruits 6-10 mm long, each mericarp with dorso-apical or or acute rounded sometimes long dorsally. merely 4. Petals

5. Calyx and pedicels stellate-pubescent, lacking glandular hairs. mm truncate 6. Petals 8-12 long, basally yellow-orange; calyx of plant coarse, yellowish; mericarps pubescence apically acute. or pale yellow; mm 6. Petals whitish calyx 7.5-9.5 long, basally veins;

than 6 mm

pubescence

apiculate. 7. Mericarps 1. Mericarps

of plant

soft, whitish;

apical spine 1.5 mm long. rounded or short-apiculate. 5. Calyx and pedicels viscid. or pale 8. Endoglossum present; petals white spine 1-1.5 mm long. 8. Endoglossum

mericarps

apically

with

absent;

petals

orange-yellow,

54.

12-20 56.

10-25

mm

strongly

1-1.5 mm

ribbed; 58. A. lozanii.

rounded, with raised or rounded rounded A.

60.

yellow,

and

spine

chiangii. racemosa.

A. mm

long; mericarps A.

long; mericarps apiculate.

with

glandulosa. rounded

57. A.

holosericea.

54. Allowissadula in Bates, Gentes Herb. 11: 352. 1978.? chiangii M. Johnston Type: Mexico. San Luis Potos?: 12 km N of El Cubo, 43 km N of Charcas 1700 m, 3 Jul 1972, Chiang, Wendt & Johnston 8232 (23?26'N, 101?0'W),

(holotype: LL!; isotype:MEXU!). Subshrubs up to 1 m tall, densely and softly pubescent with both stellate and some glandular hairs, the hairs whitish. Leaf blades mostly 4-8 cm long, broadly ovate (sometimes acute, softly pubes shallowly 3-lobed), cordate, crenate-dentate, the blades below (shorter up cent, discolorous; petioles equaling or exceeding ward); stipules 4-5 mm long, filiform. Flowers solitary in the leaf axils or aggre cm long, slender; calyx 8-10 mm gated into narrow, leafy panicles; pedicels 0.5-2.5 ca. half long, densely stellate-pubescent (glandular hairs sometimes present), the lobes apiculate, with a prominent midvein; divided, petals 12-14 mm long, white to yellowish; staminal column 6 mm long, the filaments 2-4 mm long, styles the androecium. Fruits 7.5 mm long, white-stellate-pubescent; exceeding mericarps with apical spine 1.5 mm long, with broad endoglossum completely dividing into two cells; seeds ca. 2.5 mm long, blackish. Chromosome number mericarp unknown.

to north-central Mexico Allowissadula chiangii is endemic (Fig. 12), in rela tively dry habitats at ca. 1700 m, but is as yet poorly known. This species and A. racemosa are very similar, and careful study of a series of specimens to is needed determine their relationship.

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77

MALVACEAE OF MEXICO

1988

Specimens Potos?:

Examined.

Chihuahua:

LeSueur

1413.?

Durango:

Tenorio

& Torres

4263.?San

Luis

et al. 8232.

Chiang

55. Allowissadula Brittonia 32: 265. 1980. floribunda Fryxell, (Schlechtendal) Linnaea 11: 366.1837. Abutil?n floribundum Sida keerleana Schlechtendal, nom. nov. Sida bot. ed. ii. 2: 578.1841 Steudel, Nomencl. (as "keerlena"), D. Dietrich, floribunda Syn. pi. 4: 853. 1847, non Sida (Schlechtendal) & Kunth, 1822.?Type: Mexico. Humboldt, floribunda Bonpland Michoac?n: 1830, Keerls.n. Tlalpujahua, (holotype: HAL; isotype: BR!). Rose ex R. E. Fries, Kongl. Svenska Vetenskapsakad. Wissadula microcalyx Handl. 43(4): 87. 1908. Allowissadula (Rose ex R. E. Fries) microcalyx 11: 345. 1978.?Type: Mexico. Puebla: near Tehua Bates, Gentes Herb.

c?n, 12 Sep 1906, Rose & Rose 11418 (holotype: US!; isotypes: BM! NY! US!). or shrubs to 2.5 m tall, stems softly pubescent, Subshrubs hairs yellowish, stellate and glandular. Leaf blades mostly 10-20 cm long (but much reduced in the inflorescence), broadly ovate, cordate (often deeply so), crenate-dentate, usually acute, sometimes acuminate, discolorous; petioles V2-I times as softly pubescent, a terminal panicle; long as the blades; stipules 5-6 mm long, filiform. Inflorescence cm to mm 3 6-8 viscid; up pedicels long (mostly shorter), calyx long, stellate or ca. and the lobes half-divided with viscid, less, pubescent apiculate, prominent staminal column 4 mm long, the midribs; petals 14-18 mm long, orange-yellow; filaments 3-4 mm long; styles slightly exceeding the androecium. Fruits 6-8 mm long, stellate- and glandular-pubescent; mericarps apically subacute to rounded, ca. mm seeds 2.3 black. Chromosome number: 2n = 16. absent; endoglossum long, occurs in open matorral Allowissadula in northeastern and eastern floribunda from Tamaulipas Mexico and Nuevo Le?n to Puebla (Fig. 12) at 850-1950 m and flowers from July to October. It is closely allied to A. holosericea. Specimens

Examined.

2910.?Tamaulipas: 5135.? Rose

Michoac?n:

Nuevo Le?n: Dorr 2027, 2610; Fryxell 1211, 1703; Gregg 168; hangman von Rozynski et al. 911; Viereck 890.? San Luis Potos?: Rzedowski 148; Stanford s.n.? Hidalgo: Keerl & Zavala Rose & 725.?Puebla: 48; Villase?or Montufar

11418.

56. Allowissadula

11: 352. 1978. Wissadula glandulosa (Rose) Bates, Gentes Herb. Contr. Nati. 123. 1906.?Type: U.S. Herb. 10: Mexico. glandulosa Rose, and San Pablo, 24 Aug between Higuerillas Quer?taro: 1905, Rose &

Painter 9809 (holotype:US!; isotypes:GH! MEXU! NY-2!). Subshrubs ca. 0.5 m tall (or more?), the stems densely and softly pubescent, the hairs both stellate and glandular. Leaf blades mostly 4-7 cm long, 3-6 cm wide, ovate, deeply cordate, acute, essentially crenate-dentate, concolorous, coarsely above and beneath, the hairs stellate and sometimes softly pubescent glandular; the blades below (shorter upwards). Inflorescences petioles equaling or exceeding narrow, terminal racemes or racemiform panicles; pedicels up to 3 cm long, slen der, viscid; calyx 10-14 mm long, densely pubescent with both stellate and glandu lar hairs, ca. half-divided, the lobes acuminate to caudate, with prominent midrib;

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78

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

and styles unknown. Fruits ca. 8 petals ca. 20 mm long, yellowish (?); androecium mm long, densely pubescent with both stellate and glandular hairs; mericarps with apical spine 1-1.5 mm long, with broad endoglossum completely dividing mericarp into 2 cells; seeds ca. 2.8 mm long, brownish. Chromosome number unknown. known from the collection is Allowissadula type glandulosa only (Fig. 12). 57. Allowissadula 11: 340. 1978. Abuti holosericea (Scheele) Bates, Gentes Herb. 471. 21: l?n holosericeum L?inaea 1848. Wissadula holosericea Scheele, 63: 124. Naturwiss. Zeitschr. 1890.?Type: U.S.A. (Scheele) Garcke, 354 Lindheimer ASU! BM! Texas, unknown; 1846, (holotype: isotypes:

GOET! K-4! NY! OXF! PH-2! US!). velutinum A. Gray, Gen. Amer. bor. 2: 67.1.125. 1849, non Abutil?n U.S.A. W. Texas, 4M velutinum G. Don, 1831.?Type: 1849, Wright 595

Abutil?n

[53] (holotype:GH!). Wissadula Handl. 43(4): insignis R. E. Fries, Kongl. Svenska Vetenskapsakad. Presidio 87. 1908.?Type: Texas: U.S.A. Co., Vieja Mtns, Oct 1883, Havards.n. (holotype: US!). 18: 683. 1937.?Type: Abutil?n marshii U.S.A. Standley, Amer. Midi. Nat. 172 Texas: Chisos Mtns, Aug 1935, Marsh (holotype: F!, as photo F-45084! andF-56774!). Subshrubs or shrubs 1-2 m tall, the stems yellowish pubescent, the hairs both stellate and glandular. Leaf blades mostly 7-15 cm long (reduced upward), broadly ovate or shallowly 3-lobed, deeply cordate, acute or acuminate, crenate-dentate, or more and and above less beneath, discolorous; densely softly pubescent petioles times the length of the blades; stipules 5-6 mm long, filiform. Inflorescence 1-2 (-3) usually an open terminal panicle, more or less above the leaves; pedicels cm long, viscid; calyx 8-12 mm long, densely stellate- and glandular-pubescent, ca. the lobes apiculate, with a prominent midrib; petals 10-25 mm long, half-divided, staminal column ca. 4 mm long, the filaments 4-6 mm long; styles orange-yellow; androecium. Fruits 7-9 mm long, stellate- and slender, glabrous, slightly exceeding rounded with a short (1 mm) apiculus, apically glandular-pubescent; mericarps mm blackish. seeds 2.5-3 number: 2n = Chromosome absent; endoglossum long, Vi-1

16.

occurs in open, dry habitats at elevations holosericea Allowissadula m in northeastern Mexico (Fig. 12) and in southern and western Texas, from June to November. It is closely allied to A. floribunda. Specimens & Stanford

Examined.

6836;

Kenoyer

643, 659; Fryxell Fryxell 4728. Rzedowski

Coahuila:

Fryxell 348; Leavenworth & Bates 824; Grant

2678; Palmer 777; Pringle 511; Johnston

111.?Nuevo

Le?n:

Fryxell

1945, 2675.?Tamaulipas: 4143B.?$a*? Luis Potos?:

up to 1500 and flowers

1322; Hitchcock Dressier 2295; Lukefahr

58. Allowissadula

s.n.;

lozanii (Rose) Bates, Gentes Herb. 11: 347. 1978. Wissadula 10: 124. t.41. 19%. Pseudabutilon lozanii Rose, Contr. U.S. Nati. Herb. lozanii (Rose) R. E. Fries, Kongl. Svenska Vetenskapsakad. Hand!. 43(4): near Monterrey, 100. 1908.?Type: Mexico. Nuevo Le?n: El Carrizo, 4

Sep 1904,Pringle 13443 (holotype:US!; isotypes:MEXU! VT! pf!).

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79

MALVACEAE OF MEXICO

1988

the hairs stellate, somewhat Shrubs 1-1.5 m tall, densely yellowish pubescent, cm to blades 6-11 the touch. Leaf mostly long (reduced upward), ovate to rough to dentate denticulate, cordate, usually acute, roughly irregularly triangular-ovate, above and beneath, discolorous; petioles stellate-pubescent subequal to the blades below but shorter upwards; stipules 4-5 mm long, subulate. Flowers solitary in the in terminal racemes or panicles with re leaf axils or more commonly aggregated duced leaves; pedicels 0.5-2 cm long, densely yellowish stellate-pubescent; calyx 8 truncate and strongly 10-ribbed at base, 12 mm long, densely yellowish pubescent, ca. half-divided, 13-20 mm the lobes carinate, apiculate; petals long, yellow mm mm the filaments 2-5 staminal 3-4 column orange; long, long; styles subequal to androecium, hir Fruits 8-10 mm long, densely yellowish the stigmas maroon. narrow mm and seeds 3 the acute, stout; sute; mericarps endoglossum long, apically dark brown. Chromosome number: 2n = 16. below 400 m in in open, dry habitats generally Allowissadula lozanii occurs It flowers from and southern Texas. June to Novem northeastern Mexico 12) (Fig. ber.

Specimens 13935.?

Examined.

Tamaulipas:

Nuevo

Fryxell3730,

2471; Johnston Fryxell & Kirkpatrick & Bates 2195; LeSueur293.

Le?n: 3732;

4651;

Pringle

13443,

Fryxell

11: 351. 1978. Wissadula pringlei (Rose) Bates, Gentes Herb. 3: Pseudabutilon Nati. Herb. 312. Contr. U.S. 1895. Rose, pringlei pringlei R. E. Svenska Handl. Fries, 43(4): 99. Kongl. Vetenskapsakad. (Rose) Mexico. 5 Sep 1894, Pringle 4872 Oaxaca: Monte Alb?n, 1908.?Type: as BR! F! BM! CM! US!; photo F-56183! GOET! K! isotypes: (holotype:

59. Allowissadula

MASS! MEXU! MIN, MO! NY! PH! UC! VT! pf!).

shrubs 0.5 (-1.5) m tall, the stems with dense glandular pubescence Spreading and also simple hairs 1-3 mm long. Leaf blades mostly 4-6 cm long, broadly ovate, acuminate, with stellate and simple deeply cordate (sinus almost closed), dentate, hairs above, dense stellate hairs beneath, nearly concolorous; petioles subequal to in the leaf axils or the blades; stipules 8-10 mm long, filiform. Flowers solitary cm to 4 racemiform into terminal inflorescences; up pedicels long aggregated (shorter upwards and often less than 1 cm long), viscid; calyx 10-12 mm long, viscid with glandular hairs and also with stellate hairs and long simple hairs 1-1.5 mm with a prominent the lobes acuminate-apiculate, midrib; long, ca. half-divided, mm or ca. cm 1 staminal column 2.5 white long, the petals long, pale yellow; filaments 1.5 mm long; styles exceeding Fruits 8-10 mm long, the androecium. the spine 2.5-3.5 mm long, with stellate-pubescent; mericarps apically spinescent, two into cells; seeds 2.5 mm long, dark mericarp endoglossum dividing completely brown.

Chromosome

number:

2n

=

16.

occurs in open, dry habitats at 1300 Allowissadula pringlei to Oaxaca Quer?taro (Fig. 13). It flowers from June to October.

to 2000 m

from

1185.?Hidalgo: 8196.?Puebla: Specimens Examined. Quer?taro: DArcy Arguelles Krapovickas 11964, Fryxell 183; Fryxell & Bates 924; Patoni 1127; Pringle 9520; Purpus 2603, 2603a; Rose 10131,11448; Weber 333.?Oaxaca: Bates & Vivaldi 4872; Rose 11297; Smith & Kitchen 3452; Fryxell 2570; Pringle 4770; Torke 407.

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80

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

r (

v

\

*.,>

\

Allowissadula

Alloyvis.sqdula pringlei

racemosa

r

1 # Allowissadula rosei

Allowissadula sessei

>

"^T

Anoda

Anoda

abutiloides

acerifolia

ax ' !>

Anoda

Anoda crenatiflora

albiflora FIG.

13. Distribution

of species

of Allowissadula

and Anoda.

racemosa (Schlechtendal) 60. Allowissadula Fryxell, Brittonia 32: 266. 1980. Abuti l?n racemosum 11: 367. 1837. Sida racemiflora Linnaea Schlechtendal, Steudel,

Michoac?n:

Nomencl.

bot.

Tlalpujahua,

ed.

ii.

2:

578.

1830, Keerls.n.

1841,

orth.

(holotype:

mut.?Type:

HAL;

Mexico.

isotype: BR!).

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1988

MALVACEAE OF MEXICO

81

11: 349. 1978.?Type: Herb. moorei Bates, Gentes 6.4 mi above Los Venados, Barranca de Metztitl?n, Hidalgo: Nov 1973, Bates & Vivaldi 3468 (holotype: BH).

Mexico.

Allowissadula

1850 m,

6

shrubs to 2 m tall, the stems (and much of the herbage) softly whitish the hairs stellate and somewhat arachnoid. Leaf blades mostly 4-6 cm or obscurely crenate to subentire, acute, strongly long, ovate, cordate, denticulate ca. half the above and beneath; discolorous, petioles softly stellate-pubescent length of the blades (?); stipules 3-5 mm long, subulate. Flowers solitary in the leaf axils or aggregated apically in compact racemes or panicles; pedicels up to 1.5 cm whitish tomentose; calyx 7.5-9.5 mm long, long (flowers sometimes subsessile), the lobes triangular, whitish stellate-tomentose, basally rounded, ca. half-divided, Erect

tomentose,

1-3-nerved; (or pale yellow); staminal column 3-4 petals 12-18 mm long; whitish mm long, the filaments 3-4 mm long; styles slender, slightly exceeding androecium, the stigmas maroon. Fruits 6-8 mm long (shorter than the calyx), densely stellate with rounded dorsum, the hairs 0.5-1 mm long; mericarps pubescent, scarcely into 2 cells; seeds the endoglossum umbonate, dividing the mericarp completely 2.5-3 mm long, blackish. Chromosome number: 2n = 16. racemosa is known from scattered localities in central Mexico Allowissadula and Novem (Fig. 13) in open, dry habitats at 1000 to 2500 m. It flowers in October

ber (?). Specimens & Vivaldi

Examined.

San Luis

3468; Hern?ndez-Maga?a

Potos?:

Rzedowski

3609; Moore

1865,

5714.?

Michoac?n:

s.n.? Hidalgo:

Keen

Bates

4197.

61. Allowissadula rosei (R. E. Fries) Bates, Gentes Herb. 11: 346.1978. Pseudabuti lon rosei R. E. Fries, Kongl. Svenska Vetenskapsakad. Handl. 43(4): 102. 1908.?Type: Mexico. Puebla: near Tehuac?n, 12 Sep 1906, Rose & Rose

11417(holotype: US!; isotype:NY!). Wissadula

Zoe 5: 250.1908.?Type: confer?a Brandegee, of Luis Cerro de la Yerba, San ity Tultitlanapa,

Mexico. Aug

Puebla:

vicin 1908, Purpus 2602

(holotype:UC!; isotypes:BM! DS! F!MO! NY! US!). or floccose, Shrubs 1-2 m tall, the stems densely and minutely the pubescent hairs soft, stellate, whitish. Leaf blades mostly 4-11 cm long, ovate, cordate, irregu soft discolorous, larly serrulate or crenate, acute or slightly acuminate, markedly as above and whitish Y2-I times beneath, beneath; wooly pubescent petioles long as mm a the blades; 5-6 filiform. Inflorescence terminal stipules long, compact raceme, or many of these forming an open panicle, more or less above the leaves; less than 1 cm long, the flowers sometimes subsessile; pedicels usually calyx 6-8 mm long, ca. half-divided, the lobes car?nate, apiculate, densely stellate-pubescent, the hairs sometimes stipitate; petals 8-10 mm long, yellow; staminal column 2.5-3 mm long, the filaments ca. 3 mm long; styles slightly exceeding the androecium. Fruits 5-7 mm long, stellate-pubescent; the mericarps apically rounded-apieulate, or more mm less the 2 into 2.3 seeds cells; endoglossum dividing mericarp long, dark brown. Chromosome number unknown. rosei occurs in east-central Mexico Allowissadula at 1000-2000 m. It flowers inOctober and November.

(Fig. 13) in open,

dry habitats

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VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

82

Specimens 6163.?

Ars?ne

Examined. Puebla:

2445a.?

Le?n:

Nuevo

2602,

Purpus

Fryxell & Kirkpatrick 5697; Rose & Rose 11417.

Quer?taro:

850, 851;

Arguelles

11: 343. 1978. Sida sessei sessei (Lagasca) Bates, Gentes Herb. sessei (Lagasca) sp. pl. 21. 1816, (as usesei"). Wissadula Lagasca, Gen. in cultiv. R. Hort. Madrid, 3: 22. 1969.?Type: Bonplandia Krapovickas, the See Krapovickas 1815 (MA). Note: (1969a, pp. 22-23) concerning type. Prodr. 1: 465. 1824, non Sida oxyphylla Wallich, Sida oxyphylla de Candolle, Pl. nov. Hisp. 110. 1889, nom. triloba Sess? & Moci?o, 1829. Sida acc. no. 412! Icones Florae Mexicanae (Torner Collection superfl.?Type:

62. Allowissadula

no.

6331.995,

Hunt

Institute).

F-30491

[Cf. photo

of G

copy

(not

the

type).] 2: 24. 1879. Wissadula Abutil?n trilobatum Hemsley, Diagn. pl. nov. mex. 5: 178. 1899.?Type: Herb. U.S. Nati. Contr. trilobata (Hemsley) Rose, Mexico. San Luis Potos?, 1878, Parry & Palmer 81 (holotype: K!; isotypes:

PH! US!).

BM!GH!MO!P, Wissadula

acuminata

ico. Hidalgo: isotypes: BM!

Rose,

near BR!

Contr.

U.S.

Nati.

Herb.

5:

144.

Mex

1897.?Type:

6610 (holotype: 24 Oct US!; 1896, Pringle Tula, K! F! as photo F-56184! GH! GOET! CM! ENCB!

MASS! MEXU! MIN, MO! NY! PH! UC! VT! pf!). Leaf blades mostly 8 Erect subshrubs ca. 1m tall with soft stellate pubescence. 13 cm long, more or less 3-lobed (or simply ovate distally), deeply cordate, dentate, V3-I times the length of the blades; somewhat acuminate, discolorous; petioles solitary in the leaf stipules 8-10 mm long, subulate or filiform. Flowers occasionally 1-4.5 cm long; calyx 7-10 mm long, axils, usually paniculately pedicels disposed; with long simple (1-1.5 mm) and short stellate and glandular densely pubescent to caudate; petals 1-2 cm long, white (fading rose) or the lobes acuminate hairs, column 3 mm long, the filaments 3 mm long; styles staminal sometimes yellowish; the androecium, pallid, the stigmas reddish. Fruits 1 cm in diame slightly exceeding ter, oblate; mericarps apically rounded, with a small apical umbo, indistinctly con or sometimes with a rudimentary an endoglossum stricted below, without one; number unknown. Fig. 14. seeds ca. 2.5 mm long, blackish. Chromosome sessei occurs in central Mexico Allowissadula (Fig. 13) in open matorral and dry to November. deciduous forest at 1800-2400 m and flowers from August Specimens Jalisco: 13850;

Examined.

531, 1673; Arguelles 27094, 29011, 29051; 7962.

Anoda

Durango:

17792.?Colima: McVaugh Sess? & Moci?o 3471.?San Pringle Paray

Palmer

10052; Reddick Rzedowski

241;

902; Patoni'276.? 34.?Guanajuato:

Aguascalientes: 287; Dug?s 81; Rzedowski

Rzedowski Kishler

25050.?

401;

Pringle 24604.? Ouer?taro: Parry & Palmer Garc?a 9605, 9656.?M?xico: 1484; Matuda 439; Rose 15874.?Hidalgo: Johnston 6610, 7269, 4756A; Pringle

Delgadillo Luis Potos?:

hastata Cavanilles 1: 38. 1785.?Lectotype: Anoda Diss. cristata Gray (1887) assigned all three (Linnaeus) Schlechtendal]. [=Anoda names to a single species, for which he chose the name A. of Cavanilles's hastata, and listed the others as synonyms. This choice constitutes lectotypifi

Cavanilles,

cation of the genus.

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MALVACEAE OF MEXICO

1988

FIG.

14. Allowissadula

leaf (left, below) rudimentary

(Rzedowski

endoglossum

sessei. 15874); (right,

Flowering mature

below)

and fruiting branch (McVaugh fruit (left, above) and mericarp

(Rzedowski

83

17792); opened

portion of mature to show position of

25050).

Cavanillea

Malvenfam. 19. 1787.?Type: Cavanillea hastata (Cava Medikus, Medikus. nilles) Sidanoda & Standley, Contr. U.S. Nati. Herb. 19: 427. (A. Gray) Wooton sect. Sidanoda A. Gray, Proc. Amer. Acad. Arts 22: 299. 1915. Anoda Sidanoda pentaschista & Standley. 1887.?Lectotype: (A. Gray) Wooton statement sect. Note: that Hochreutiner's is Sidanoda (1916, p. 39) on not A. correct. and based abutiloides is When Wooton & monotypic sect. Sidanoda to generic rank, they narrowed Gray's Standley elevated an action that to taxon the the of concept single species Anoda pentaschista, serves as the choice of a lectotype.

Annual or perennial herbs or subshrubs, erect or decumbent, hispid or stellate or -puberulent to glabrate. Leaves petiolate, the blades linear, lanceo pubescent to crenate or dentate lobed or palmately late, ovate, cordate, hastately divided, subentire, usually green, sometimes with an irregular purplish blotch along mid canescent in vein, sometimes beneath, lacking foliar nectaries; stipules generally and deciduous. Flowers solitary in the leaf axils or aggregated in open conspicuous accrescent involucel absent; calyx sometimes in fruit, apical racemes or panicles; lavender, basally rounded, usually 5- or 10-nerved; petals spreading, yellow, white, or purplish; androecium shorter than to equaling the petals, usually yellowish, at antheriferous the androecium apex; styles 5-20, slender, exceeding (barely so in A. thurberi), the stigmas usually abruptly capitate, glabrous. Fruits (Fig. 15) oblate

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84

SYSTEMATIC BOTANY MONOGRAPHS

A. A. polygyna. B. A. leonensis. FIG. 15. Fruits of Anoda. A. pedunculosa. F. A. reticulata. G. A. cris tata. H.A. acerifolia. A. pubescens. L. A. thurberi. M. A. hintoniorum. N. A. pr?stina.

VOLUME 25

D. A. pentaschista. C. A. paniculata. I.A. lanceo lata. J. A. guatemalensis.

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E. K.

1988

MALVACEAE OF MEXICO

85

or disklike, schizocarpic, to hispid; mericarps 5-20, with or without spur puberulent at maturity; or spine at dorsal angle, the lateral walls usually disintegrating seeds or in endo reticulate enclosed sometimes persistent pubescent, solitary, glabrous number: x = 15. carp. Base chromosome and nodus, knot; given to this Don (1831) states "from a, without, Etymology. " are which is remarked in Sida, the without articulation because the genus pedicels a derivation Schlechtendal also given by Sweet (1839). However, (1837) notes the more vernacular the from derivation name, anoda, given to a Ceylonese likely and taken Burman noted of Abutil?n, up by by (Thesaurus Zeylanicus), species Jos? Cavanilles Cavanilles. Cavanillea was named to honor Antonio (1745-1804), classis disserta?ones decent and other Spanish botanist and author of Monadelphiae of the generic names Sida and Anoda. is a combination works. Sidanoda in Mexico has its principal center and maximum The genus Anoda diversity occur in the A few southwestern United and States, 20). species (Figs. 13, 16, 18, is widespread and relatively and one species (A. cristata), with weedy tendencies, to States through Central America from the southern United common, occurring as a occurs waif else of cristata America. Anoda South many parts occasionally where, presumably by transport with agricultural planting seed. References:

Schlechtendal

(1837), Gray

(1887), Garcke

(1896), Hochreutiner

(1916),Wiggins (1964), Fryxell (1987), Bates (1987). to the Species

Key

yellow or pale yellow. and styles 5-8 ;endocarp 3-5 mm long; mericarps

1. Petals

bright 2. Mericarps 3. Calyx

absent

of Anoda

;corolla

yellow but sometimes fruits 4-5 mm styles 5-8; linear. in shape, often narrowly

fading pale reddish. in diameter; leaves 78. A. pentoschista.

and

variable discolorous, coriaceous, in diameter; leaves membra 3. Calyx and styles 5; fruits 6 mm 5-7 mm long; mericarps acuminate. 63. A. abutiloides. nous, concolorous, ovate, broadly corolla not changing color. 2. Mericarps and styles 8-15; endocarp usually present; or absent; stems and petioles 4. Endocarp 71. A. hirta. notably hirsute. developed weakly 4. Endocarp present lent to subglabrous. 5. Stems 5.

Stems

and petioles and petioles

6. Midstem

leaves

6. Midstem

leaves

7. Upper

in A.

(unknown nearly

succulenta);

stems

and petioles

pubescent

or puberu

stems succulent. 84. glabrous; or puberulent, stems not succulent.

A.

succulenta.

pubescent

deeply palmately ovate to hastate;

and styles lobed; mericarps and styles 10-16.

75. A. palmata.

8-13.

mericarps

with simple appressed with hairs; endocarp well developed 10-12 with radiating spines. 72. A. lanceolata. excrescences; mericarps to glabrate; leaf surface sparsely reticulate stellate-pubescent endocarp excrescences; 10-16, crested or spurred. mericarps

leaf surface

prominent 7. Upper lacking 8. Petals

8-12 (-20) mm long, bright yellow; mericarps 73. spurs virtually absent; plants 1-2 m tall. 8. Petals 6-8 mm long, pale yellow; mericarps 10-13, with than 1m tall. plants seldom more 9. Corolla

with dark center, the androecium base of calyx to 1mm long. dark center, the androecium 9. Corolla without mm 0.1-0.3 long. to white. rose, pink, or pale lavender

1. Petals

reddish

12-16, A. spurs

crested

but leonensis.

1-2 mm

or purplish;

long;

hairs

at

74.

A. maculata.

pallid; 66.

hairs at base of calyx A. crenatiftora.

purplish, 10. Petals 4-7 mm

calyx), blue-purple. long (barely exceeding and mericarps 6-8, the latter with spurs to 1mm absent. 85. A. thurberi. triangular; endocarp

11. Styles

long;

leaves

ovate

to hastate

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to

VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

86

the latter with rounded dorsum 10-11, (spurs completely absent); 3 narrow, linear lobes ;endocarp 82. A. reticulata. present. to white. to pale lavender long, purplish to petals; endocarp absent. subequal

11. Styles and mericarps midstem leaves with 10. Petals

6-28 mm

12. Androecium 13. Flowering

12-15

calyx

mm

22-28

long; petals

mm

staminal

long;

13. Flowering

6-12

calyx

mm

7-17

long; petals

mm

long;

staminal

ca. 20

column

mm long.

83. A.

column

7-15

speciosa. mm

long. 14. Petals

7-14 mm long, uniformly lavender (except whitish on claw); dorsal spur ca. 0.5 mm long. of mericarp A. 81. pubescens. 14. Petals 11-17 mm often with microscopic streaks dis long, lavender, purple ca. 1.5 mm long. 69. A. henricksonii. tally; dorsal spur of mericarp shorter than the petals; endocarp present or absent. 15. Upper leaf surface stellate-pubescent. 16. Mericarps and styles ca. 10; endocarp absent; calyx 7 mm to 10 mm accrescent 68. fruit. in

12. Androecium

16. Mericarps and styles not accrescent.

12-20;

reticulate

long in flower, A. guatemalensis. calyx 4-11 mm long,

present;

endocarp

17. Mericarps and styles 12-13; long; petals calyx 4-6 mm to lavender leaves white; pale sparsely pubescent. 17. Mericarps and styles 17-20; calyx 9-11 mm long; petals or leaves densely pubescent. pink pale lavender; 15. Upper leaf surface with appressed simple hairs or glabrate. 18. Mericarps with persisting lateral 12-13, apically dehiscent,

mm long, 77. A. pedunculosa. 15-23 mm long, 6-15

79. A. polygyna.

reticulations. 80. A. pr?stina.

18. Mericarps 19. Plants

7-19,

ind?hiscent

but

the

at maturity. disintegrating pink, or lavender.

lateral wails

erect, sometimes shrubby; corolla white, 20. Upper leaf surface glabrate; present. endocarp leaf surface with appressed 20. Upper simple hairs; 21.

Petals white,

21.

Petals

pink

20-25

mm

or lavender,

long; mericarps 14-18 mm

76. A. paniculata. endocarp

and styles

long; mericarps 10.

absent. ca. 15. 65. A. albiflora. and styles 8 70. A. hintoniorum.

19. Plants

corolla usually purplish. herbaceous; decumbent, frequently 22. Mericarps the dorsal spur absent or vestigial 7-11, (rarely to 1.5 mm not depressed, the mericarps the suture between the fruits long), therefore mately

seemingly

inflated;

lower

leaves

frequently

lobed.

ovate

to pal

64. A.

acerifolia. 22. Mericarps dorsal spine 1.5-4 mm 10-19, with a horizontal long, the suture between the mericarps lower leaves frequently trian depressed; 67. A. cristata. gular or hastate.

63. Anoda

abutiloides A. Gray, Proc. Amer. Acad. Arts 22: 300. 1887.?Type: Arizona: Santa Catalina Mtns, 7 Jun 1882, Pringle s.n. (holotype:

U.S.A.

GH!; isotypes: CM! F! MO! NY! PENN! US! VT! WIS!). Note: This is incorrectly attributed to S. Watson in Index Kewensis. & Proc. Amer. Acad. Arts 29: 382. Robinson Greenman, caudatifolia & & Green Robinson 1894. Anoda caudatifolia (Robinson Greenman) Nati. Herb. 5: 172. 1899.?Type: Mexico. Jalisco: man, Contr. U.S. 13 Oct 1893, Pringle 5445 (holotype: GH!; isotypes: Barranca of Tequila, binomial

Sida

ARIZ! CAS! F! as photo F-56124! K! MEXU! MICH! MO! NY! RM! SMU! UC! VT!). Anoda 110. 1923.?Type: urophylla Riley, Kew Bull. Rinc?n de la Casafera, May 1921, Gonz?lez-Ortega

Mexico. Sinaloa: El 888 (holotype: K!;

isotype: MEXU!).

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MALVACEAE OF MEXICO

1988

87

Erect perennial subshrubs ca. 1m tall, branching principally in the inflorescence, of shorter the stems with patent simple hairs 0.5-1 mm long and with understory more or less glandular hairs. Leaf blades up to 12 cm long, gradually reduced and mi much narrower upward, cordate, dentate, caudate-acuminate, rotund-ovate, above and beneath, concolorous; essentially nutely and softly tomentose petioles solitary in subequal to the blades or somewhat shorter upwards. Flowers occasionally 1-5 cm long, with the leaf axils but usually in an open terminal panicle; pedicels like stem, lacking articulation; calyx 5-7 mm long, less than half-divided, pubescence the lobes broadly triangular, more or less acuminate, with darkly pigmented midrib; a dark center, the petals ca. 1 cm long, pale yellow (drying rose), corolla without bearded on claw; staminal column ca. 6 mm long, with recurved hairs, prominently the filaments 3-4 mm long; styles 5, slightly exceeding androecium, subequal to reddish. Fruits ca. 6 mm in diameter, oblate, petals, pallid, the stigmas sometimes seeds 3 mm long, minutely 5, dorsally rounded; mericarps number: 2n = 30. without enclosing endocarp. Chromosome

minutely

pubescent; appressed-pubescent, Fig. 89. Anoda abutiloides in open, dry matorral examined.

Specimens Chihuahua:

Bye

Gonz?lez-Ortega

6939,

occurs

to Jalisco along the Pacific coast (Fig. 13) from Arizona to December. forest. It flowers from October

and deciduous

et al. 2690; White 485, 821, 1083; Sanders 3518, 3748.? Gentry & Torres 4452? Sinaloa: 10473; Gentry 660; Tenorio Bye & Linares 7594.?Jalisco: 6082; Moran Fryxell & Bates 2136; Pringle 5445. Sonora:

10054;

706, 888,

Anales Ci. Nat. 6: 336. 1803, non Anoda acerifolia Cavanilles, ex hort. Madrid, de 1803 [grown from 1824.?Type: Candolle, (Zuccagni) at Florence] from the Botanical Garden seeds obtained (holotype: MA as

64. Anoda

acerifolia

photo F-29756!). in Roemer, 148. 1809, non Sida acerifolia Sida acerifolia Zuccagni Collectanea nee Anoda 1816. 1787, acerifolia (Zuc (Cavanilles) Medikus, Lagasca, non 1: Prodr. Candolle de 459. Anoda 1824, acerifolia Cavanilles, cagni) 1803. Sida zuccagnii Sprengel, Syst. veg. 3: 121. 1826, nom. nov. Anoda ex hort. Florent. Aliso 11: 519. 1987.?Type: zuccagnii (Sprengel) Fryxell, s.n. not Zuccagni located). (holotype: FI, to prostrate, Herb or subshrub, annual or perennial, suberect or decumbent the or obscurely hispid or almost glabrous. Leaf blades highly vari stems prominently able in size and form, palmately lobed below to narrowly hastate above, subentire, with appressed acute, sparsely pubescent simple hairs to glabrate above and be neath, sometimes with an irregular purple blotch along the midrib; petioles usually less than half the length of the blades. Flowers in the leaf axils; long-pedicellate mm mm to 6-11 accrescent in 10-15 in often hispid; flower, fruit, calyx long long a dark center, corolla without the petals 12-18 mm long, purple or lavender; staminal column shorter than the petals, basally glabrous, apically hispid, the fila ments 2-3 mm long; styles 7-11. Fruits (Fig. 15H) 8-11 mm in diameter, oblate, with short spurs less than 1.5 mm (or these absent), densely hispid; mericarps 7-11, mm the walls lateral seeds 2.5 evanescent; ind?hiscent, gibbous, long, glabrous, number: 2n = 30. endocarp absent. Chromosome Anoda acerifolia iswidely distributed inMexico (Fig. 13) and occurs principally in disturbed habitats. It flowers more or less throughout the year.

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88

SYSTEMATIC BOTANY MONOGRAPHS

Specimens 5493?Sonora:

Baja

Examined.

5774; Gonz?lez-Ortega 5993; 576, 4326. ?Tamaulipas: Ortega Mexia

507.?Jalisco:

California:

& Rollins

Wiggins

Carter

195?Chihuahua:

2111, 5094; Gentry 4240; Moran; alo a: Fryxelll37, Palmer 234.?Sin

6991; Wiggins

3042; Gentry 1205. ?Durango: Gonz?lez 24168; Wright Croat 45391, 45395; Ferris 5770; Lehto 24246; & Puga 617; Lott & Magallanes 764; Gonz?lez 327; Oliva 46; Rinehart et al. 1614; Langman Wilbur & Wilbur 2221.-Colima: 3170; Fryxell

Lamb

Carvajal

VOLUME 25

458; Lane 2305; Romo 3.?Nayarit:

6417, 6431; 7060; Spellenberg & Fields 326.? San Luis Palmer 909; Thompson terle 4264; Fryxell 1728; Hinton 12523,12847,13663;

Lehto

Potos?:

Cottam

Die 10535; King 4246.?Michoac?n: 17915.-Morelos: Duncan 179; McVaugh et al. 234; Hinton et al. Koch 69; Vaquez 285,1062?Guerrero: 14175; Germ?n 10885,11632; Barkley et al. 78209; Nelson & Armbruster 79126.?Veracruz: Balls 4301; Holstein 20382; Koch 21, 23, 395; Rosas Mart?nez-Calder?n 442.?Chiapas: Breedlove 774; Sousa 1336.? Oaxaca: 33488; Carbon 2380; Thome

& Lathrop

65. Anoda

41044.?

Yucat?n:

Bradburn

Mart?nez

& Darwin

11: 493. Fryxell, Aliso Jan 1978, Lapes.n. pec, cultivated, albiflora

1229; Gaumer

1987.?Type:

23549.

Mexico.

(holotype: MICH!;

Jalisco:

Jocote

isotype: pf!).

or subshrub to 2 m tall, sparsely hirsute to glabrate. Leaf blades 3-7 cm narrower upward, sometimes weakly hastately ovate, long, becoming progressively the hairs mostly acute, sparsely hirsute, lobed, crenate-serrate, simple, on the or surface half the of the blades less. Flowers upper length petioles appressed; or or more in the axils less 1.5-6.5 cm leaf solitary aggregated apically; pedicels a mm corolla without dark 8-10 center, the hirsute; long; calyx long, prominently cm on 2-2.5 of hirsute claw, otherwise petals long, white, prominently margins mm 4 column hirsute toward staminal apex, the filaments 2 glabrous; long, pallid, Herb

3mm

ca. long; styles ca. 15, pallid. Fruits disciform with radiating spines; mericarps mm the 1.5 walls dorsal the lateral seeds 15, sparsely hispid, evanescent; long, spine in endocarp, minutely verruculate. 2.6-2.8 mm long, not enclosed Chromosome number

unknown.

Anoda Specimens

66. Anoda

albiflora Examined

is a garden (both

plant not known

from garden

cultivation).

from the wild

Chihuahua:

Vaid?s

(Fig. 13). 838.?Jalisco:

Lape

s.n.

Ortega, Nov. pi. descr. dec. 8: 96. 1798. Sida crenatiflora Persoon, Syn. pi. 2: 247. 1807. Anoda ortegae Sprengel, Nachtr. (Ortega) I. Bot. Gart. Halle 11. 1801, nom. superfl. Sida ortegae Steudel, Nomencl. inMadrid Botanical Gar bot. ed. ii. 2: 578. 1841, nomen superfl.?Type: crenatiflora

: den (holotype MA). Icon. 5: 19. t.431. 1799. Sida parviflora Cavanilles, (Cava parviflora Enum. pi. hort. berol. 726. 1809, non Sida parviflora nilles) Willdenow, ex herb. Pav?n s.n. (G as photo F Sess? & Moci?o, 1894.?Type: Mexico,

Anoda

23684! P). crenatiflora var. glabrata Rose, Contr. U.S. Nati. Herb. 5: 172. 1899.? near San Juan Capistrano, 20 Aug 1897, Rose Zacatecas: Type: Mexico. 2444 (lectotype, 1987: BM! US!; designated by Fryxell, isolectotypes:

Anoda

GH!). annual herbs to 1m tall, minutely the hairs 0.1-0.3 stellate-puberulent, so cm ovate to trun Leaf 3-9 blades hastate, long, long. narrowly upwards, to subentire, acute, minutely and obscurely pubes cate, coarsely crenate-dentate or stellate hairs above, stellate-pubescent cent, with simple, bifurcate, beneath; Erect

mm

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MALVACEAE OF MEXICO

1988

89

in the blades on lower leaves, shorter upward. Flowers nearly equaling petioles terminal racemes or panicles; pedicels 2-7 cm long; calyx 3-7 mm long in flower, a dark to 6-8 mm in fruit, densely accrescent corolla without short-tomentose; center, the petals 6-8 mm long, pale yellow, ciliate on claw; staminal column ca. 2 mm long, pallid, glabrous or with a few hairs apically, the anthers few (up to 10), the androecium. Fruits 7-9 mm in diame subsessile; styles 10-13, slightly exceeding ter, densely 10-13, dorsally spurred (the spur 1-2 mericarps stellate-pubescent; mm long), the lateral walls evanescent; seeds ca. 2.5 mm long, completely enclosed number: 2n = 60. by persistent reticulate endocarp. Chromosome name: pintapan del monte. Vernacular inMexico is widely distributed forest Anoda crenatiflora (Fig. 13) in deciduous at 800-2300 m. It flowers principally from August to November. Sonora: Drouet & Richards California: Gould 12163a.? 3946; Gentry & Rollins 421.? Sinaloa: 4398.? s.n.; Howe 7057; Wiggins Brandegee Henrickson & M?ller 562, 1073.? Coahuila: 975; 128; Pringle 15052; Johnston et al 1701, 1707.?Tamaulipas: Stewart 1135.?Zacatecas: Rose 2444.?Nuevo Le?n: Fryxell Fryxell s.n.? 15704. ?Guanajuato: 1115; Fryxell etal. 1693.? San Luis Potos?: Schaffner 158; Waterfall Dug?s Specimens

1091, 2429; Chihuahua:

Examined.

Wiggins Palmer

Baja

7046,

Jalisco:

2963; Rose 2916.?Hidalgo: 17347; Pringle McVaugh Fryxell & Bates 2177, 2180; Rose & et al. 79119.?Puebla: 7075. ?Morelos: 880.? Guerrero: Hinton 6923; Koch V?zquez Purpus 3256.?Oaxaca: & Gonz?lez Conzatti 1932; Conzatti 28, 1212; Fryxell 1143, 2571; Rose & Rose 11325; Rzedowski & Strother 46137; Fryxell & Lott Breedlove 37079; Smith 945.? Chiapas: 52347; Breedlove Painter

3323.

67. Anoda

cristata (Linnaeus) Linnaea 11: 210. 1837. Sida cristata Schlechtendal, lavateroides Medikus, Malvenfam. 19. Linnaeus, Sp. pi. 685. 1753. Anoda herb. Linnaeus 1787, nom. superfl.?Type: (holotype: LINN-866.31!). 1: 38. t.ll.f.2. Anoda hastata Cavanilles, Diss. 1785. Cavanillea hastata (Cava 19. 1787. Sida hastata (Cavanilles) Willde Malvenfam. nilles) Medikus, now, Sp. pi. 764. 1801, non Sida hastata St.-Hilaire, 1827, nee Sess? & s.n. (lectotype, designated Peru. Lima, Dombey 1889.?Type: Moci?o, by

1837: P-JUno. Schlechtendal, 12295-A!). Anoda Diss. 1: 39. t.lO.f.3. 1785. Sida quinqueangulata triloba Cavanilles, D. "cult, in h. Yssy 1785" Dietrich, Syn. pi. 4: 857. 1847, nom. nov.?Type: (holotype: P-JU no. 12296!; isotype?: MA as photo F-29759!). Anoda dilleniana Cavanilles, 1: 40. t.ll.f.l. Diss. 1785. Sida dilleniana (Cava "abutil?n lavaterae flore . . ." nilles) Willdenow, Sp. pi. 763. 1801.?Type: (holotype: P-JU no. 12298!). Sida mexicana Scopoli, Delic. fl. faun, insubr. 1: 22. t.9. 1786.?Type: plate 9. Sida deltoidea Hornemann, Enum. pi. hort. hafn. 36. 1807.?Type: unknown. Sida triangularis Willdenow, Enum. pi. hort. berol. 726. 1809. Anoda triangu laris (Willdenow) de Candolle, Prodr. 1: 459. 1824.?Type: & Humboldt s.n. (holotype: B-W no. 12722!; isotype: G as photo F-7993!). Bonpland Anoda Reichenbach, brachyantha Iconogr. bot. pi. crit. 1: 24. t.34. 1824. Anoda cristata var. brachyantha Annuaire Hochreutiner, (Reichenbach) Conserv. 20: 47. 1916.?Type: Jard. Bot. Gen?ve Reichenbach, op. cit., plate 34. Sida centrota Sprengel, Hort. Hal., 1826 Syst. veg. 4(2): 259. 1827.?Type:

(HAL?).

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SYSTEMATIC BOTANY MONOGRAPHS

90

VOLUME 25

Anoda populifolia Philippi, Linnaea 28: 313. 1856.?Type: cerca de Quillota, 41048 (holotype: Germaine SGO;

Chile.

Valparaiso: isotype: W as photo

F-32643!). Anoda arizonica

A. Gray, Proc. Amer. Acad. Arts 22: 298. 1887.?Type: Lemmon 599 (holotype: GH!). U.S.A. Arizona, 1881, Sep Anoda arizonica var. digitata A. Gray, Proc. Amer. Acad. Arts 22: 298. 1887. cristata var. digitata (A. Gray) Hochreutiner, Conserv. Anoda Annuaire 20: 47.

Jard. Bot. Gen?ve

1916.?Type:

U.S.A.

Arizona,

1881, Lemmon

517 (holotype: GH?). Anoda

var. albiflora Hochreutiner, Annuaire Conserv. Jard. Bot. Mexico. Oaxaca: Cordillera, 20: 48. 1916.?Type: 2300 m, 1840, 4108 (lectotype, here designated: G as photo F-23685!).

cristata

Gen?ve Galeotti

Suberect to sprawling perennial herbs, the stems usually hispid, the hairs patent or retrorse. Leaf blades mostly 3-9 cm long, variable (ovate, triangular, hastate, or above and rarely palmately lobed), crenate to subentire, acute, sparsely pubescent a the hairs mostly often with and blotch beneath, simple appressed, purple along also on margin; Vi-1 times as long as the blades, midvein, occasionally petioles hispid. Flowers solitary in the leaf axils, long-pedunculate; calyx 5-10 mm long in to 12-20 mm long in fruit, hispid; corolla without a dark center, flower, accrescent the petals 8-26 mm long, purplish or violet (rarely white); androecium shorter than the petals, pubescent; styles 10-19. Fruits (Fig. 15G) 8-11 mm in diameter (exclud a flattened 10-19, disk, densely hispid; mericarps ind?hiscent, with ing spines), mm on 1.5-4 dorsal the lateral walls seeds 3 mm long, evanescent; spine long angle, an enclosing endocarp, with or without 2n = 30, numbers: Chromosome glabrous. 60,90. Vernacular

names: alache, altea, alushi, amapola del campo, amapolita morada, itsukua tsipata (tarascan), malva de castilla, malvavisco, bimalva, pie del gallo, violeta, violeta del campo, violeta silvestre, yiwa t?io (mixtee). It is noted on several herbarium labels (e.g., Boege 254, 585; Messer 8, 197) that this species is used as a green vegetable. and widespread Anoda cristata is a common species, occurring throughout Mexico and the year. It is highly variable (Fig. 16) flowering essentially throughout and often occurs as a weed in disturbed areas. It is usually lavender-flowered but white-flowered sometimes occur. Its distribu variants (var. albiflora Hochreutiner) tion extends northward into the southern parts of the United States, and southward to much of South America. through Central America Specimens

Examined.

Baja

California:

Brandegee

s.n.; Carter & Chisaki

399.?

3529; Hammerly

Sonora: Gentry 428; Hartman 886; Shreve 6604; White 2627, 2892; Whitehead M70; Wiggins 6189; Wiggins

Sl Rollins

339.?Chihuahua:

Bye

4863;

Correll

& Gentry

22841;

Gentry

1829; Hartman

559;

Pennington 73; Powell & Edmondson 997; Pringle 1041; Rzedowski 32347; Vald?s 735; Weber & Charette

Latorre

11634.?Coahuila:

74; Marsh

1039;

Stewart

1175. ?Sinaloa:

Breedlove

& Thome

259, 5712; Mexia 182, 301; Palmer 1596; Sanders 176.? Zacatecas: Henrickson 581; Patoni 781; Longman 2957; Palmer Le?n: Dorr 2576; Fryxell et al. 1708.?Tamaulipas: Daniel B6328; Rose 3632.?Nuevo 320; Stanford 212.?Nayarit: Croat 45152; Jones 22866; Palmer 2033.?Jalisco: Barnes & Land 804; von Rozynski 158,159; 18172; Fryxell et al. 4568.?Durango:

Gentry Garc?a

6282; Gonz?lez-Ortega

206; Daniel 1104; Dieterle 3425; Germ?n 186;Harker & Mellowes 102; Jones 27143; Langman 3137; McVaugh Dug?s

287

10545; Rose 2523.?Colima: 13131, 16304; de Puga & Carvajal bis; Dunn 103, 111, 400, 471, 20533; Hern?ndez 2366; Kishler

Burch 472,

5136.?Guanajuato: 479.?San

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Luis

Potos?:

1988

FIG.

MALVACEAE OF MEXICO

16. Distribution

of species

91

of Anoda.

& Reko 332; Duke 3659; Parry & Palmer 77; Rzedowski 5220, 5371, 6199, 6843, 9913, 24411; 159; Waterfall 145, 269, 540, 555, 1496.? Michoac?n: 14276, 15703.? Quer?taro: Schaffner Arguelles Ars?ne 2844; Calzada 8210; Cutler 4067; Dieterle 4348; Gonz?lez-Quintero 1731; Hinton 13205; King & Soderstrom 743.? Mexico: 4629, 4706; Kishler 171, 526; McGregor 16570; Patrick 7290; Barkley 560; Carrera 11; Hern?ndez 256; D?az 2123; Hint?n 1154, 2564, 6796, 6987; Koch 76221; Bourgeau Aguirre

Matuda

19526,

21710,

21897,

21913;

Moore

1144;

Pineda

912;

Roe

1625;

Rose

&

Painter

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6612;

SYSTEMATIC BOTANY MONOGRAPHS

92

VOLUME 25

Rzedowski 34310; Trejo 91; Ventura 2250, 3357, 3398, 3667, 3797; Villegas 87, 206; Waterfall 16485. DistritoFederal: 235; Pascoe 75.?Hidalgo: Wood

Berlandier671

;Bourgeau292, 471, 559, 560; Garcia22, 450; LyonnetBO, 161; Paray Rzedowski V?rela 19; Russell & Souviron 7316; Rovirosa 44,158; 1852,16286; & 622, 777; Gonz?lez-Quintero 1615; Moore 36; Moore 1589; Karwinski Gimate-Leyva

200; Pringle Trevi?o

4301;

256; Gimate-Leyva

41; Ventura 217, 501.?Morelos: 81; Harris 33; Jackson 25; G?mez

Diaz 176; ?ores-Crespo 193; Estrada 112, 227, ? 441, 494, 2323, 3060 53; Patrick 368; V?zquez

Guerrero: Bell & Duke 17006; Cowan 4985; Croat 45527; Fryxell 609, 628; Hinton 6877, 6889, 9524, 9612,

11165,

11359,

8922. ?Tlaxcala:

14824; Mexia

14597,

Weber

693.?

Puebla:

Ars?ne

155,

1284,

1658, 2035; Basurto & Patron 239; Boege 254, 585; Chute 166; D'Arcy 11942; Dunn & Dunn 18761; 873,

Botteri 755, 2215, 2335; Purpus 852; Miranda 4391; Weber 24, 222, 348.?Veracruz: & Cochrane Cochrane 16; Mart?nez-Calder?n 8606; Gonz?lez-Quintero 2214; Hern?ndez et al. 99; Rosas 100, 278, 734, 938, 1452; Ventura 4626, 4781, 7114, 1801; Matada 18300; Ortega Croat 46312; 9186, 9463, 10047, 12077, 12401, 12403; Villavicencio 2; Zol? 162, 777.-Oaxaca: & Leake

Gallian

1135;

1033, 8892,

Garc?a 458, 728;Hill 1687;King 780, 1036;Koch 73161; Lorence 3697;Marcks & Marcks 1051;Messer 4778.?Tabasco: Schuhes ? Reko 103; Smith & Kitchen 8, 197; Nelson 1718; Rose M532; & Maga?a 1069. ?Chiapas: Breedlove 7542, 9453, 11899, 12853, 14467, 1958; Fern?ndez

Cowan 14613,

1920, 27806,

52378, 52647, 53062; Breedlove & Raven 13060; Calzada 3375; Croat 47697, 47829; Cruden 1198;Fryxell 1176, 2587; Laughlin 110, 735, 2113, 2410; Matuda 80, 5817; Roe 1141; Saunders 21; Ton 851, 1237, 1931, 4582; Webster

12880;

Zuill

254. ?Campeche:

Stewart

& Per civ al 84-85.?

Yucat?n:

Enr?quez

249.

11: 498. fig. 3. 1987.?Type: Guatemala. 68. Anoda guatemalensis Fryxell, Aliso of Sol?la and 21 Sep Los Robles border Depts, bridge, Chimaltenango

1971, Molina & Molina 26698 (holotype: ENCB!; isotypes: EAP, F! MICH!). and peduncles minutely scabridulous. Erect herbs 1m tall, the stems, petioles, smaller upward), blades up to 10 cm long (progressively basally cordate, to serrate the central lobe lance-ovate 3-lobed, linear-lanceolate, usually palmately to obscurely and crenate, acute, concolorous, evenly minutely stellate-pubescent; Y2-I times as long as the blades. Flowers solitary in the leaf axils or petioles the subtending somewhat aggregated apically; pedicels often equaling or exceeding to 10 mm long in fruit, evenly stellate leaves; calyx 7 mm long in flower, accrescent a dark center, the petals 11-14 mm long, lavender; corolla without pubescent; 1.5 mm long, the staminal column 7 mm long, pallid, densely hispid, the filaments

Leaf

styles ca. 10. Fruits (Fig. 15J) ca. 1 cm in diameter, ca. 10, dorsally spurred, the spur ca. 0.5 mm oblate, stellate-pubescent; mericarps the walls 3 mm long, endocarp absent. Chromosome lateral seeds evanescent; long, anthers

sometimes

number

unknown.

Anoda

lavender;

guatemalensis

occurs

in oak-pine

forest

inOaxaca,

Chiapas,

and Guate

mala (Fig. 16) at 1000 to 1800m. Specimens

examined.

Oaxaca:

Pringle

5646.?Chiapas:

Breedlove

41257; Nelson

1466,

3169.

53: 451. 1983.?Type: 69. Anoda henricksonii M. Johnston, Mexico. Phytologia Henrickson 13305 Zacatecas: Sierra del Astillero (24?37'N, 101?08'W), (holotype: LL!). herbs ca. 0.5 m tall, the stems densely pubescent becoming stellate. Leaf blades mostly 3-6 cm long, the hairs mostly sparsely pubescent, below to hastate-lanceolate hastate-deltoid above, deeply cordate, serrate, acute, 3-4 cm long, with pubescence somewhat discolorous, petioles mostly pubescent; Erect

branched

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MALVACEAE OF MEXICO

1988

93

like stem. Flowers solitary in the leaf axils or forming a terminal raceme; pedicels 2-4 cm long in flower, up to 6 cm long in fruit, slender; calyx 9-12 mm long, the lobes lance-oblong, stellate-tomentose, densely and minutely deeply 5-lobed, a dark center, the petals 11-17 mm long, more or less apiculate; corolla without with darker microscopic streaks distally); lavender (sometimes sub androecium mm to 10-15 the column unknown. Fruits oblate, long; styles equal petals, densely and minutely 10, with a dorsal spur ca. 1.5 mm long; mericarps stellate-pubescent; mature seeds unknown, number unknown. endocarp absent. Chromosome Anoda henricksonii is known only from Zacatecas in desert habitats (Fig. 16). It is very similar to A. pubescens from Hidalgo and M?xico, but differs principally in the characters given in the key. Specimens

Examined.

Zacatecas:

70. Anoda

Henrickson

hintoniorum Fryxell, S. Torricillas, Michoac?n:

Aliso Distr.

13305,

13405b;

Palmer

377.

11: 500. fig.4. 1987.?Type: Mexico. et al. 17 Dec 1938, Hinton

Coalcom?n,

12778 (holotype: US!; isotypes: BM! BR! DS! GH! K! MICH! NY! RSA! UC! pf!). to glabrate, often shrubs to 2.5 (-4) m tall, the stems sparsely puberulent cm Leaf blades 4-8 truncate, crenate mostly long, ovate-lanceolate, purplish. hairs minute stellate hairs serrate, narrowly acute, with appressed above, simple than less half the of the Flow blades. beneath, nearly concolorous; petioles length ers solitary a in the leaf axils but generally in aggregated apically congested inflorescence; pedicels mostly 2-3.5 cm long, slender, minutely stellate-pubescent; calyx 7-11 mm long, evenly pubescent with stellate and glandular hairs intermixed, ca. half-divided, the lobes long-acuminate, sometimes apically purplish; corolla without a dark center, the petals 14-18 mm long, pink or lavender; staminal column ca. 4 mm long, the filaments 3 mm long; styles 8-10, pallid, apically pubescent, Fruits in diameter, stellate- and oblate, minutely pinkish. (Fig. 15M) 8-9 mm Erect

the 8-10, with a small (0.5 mm) dorsal umbo, glandular-puberulent; mericarps lateral walls evanescent; seeds 2.5 mm long, minutely ab scabridulous, endocarp sent. Chromosome number unknown. occurs principally Anoda in oak-pine hintoniorum forest at 2500 to 2800 m in central Mexico to It November flowers from February. (Fig. 16). Specimens etal.

3036,

Examined.

7229.?

Morelos:

Michoac?n: V?zquez

Frye & Frye3098; 1843.

Hinton

etal.

12778,13554.?M?xico:

Hinton

71. Anoda hirta Fryxell, Aliso 11: 502. 1987.?Type: Mexico. 27 km Tamaulipas: SE of Miquihuana, road to Palmillas, 13 Aug 1941, Stanford, Retherford &

Northcraft878 (holotype:MO!; isotypes:ARIZ! DS! NY! OS! UC!). Erect herbs or subshrubs, the stems, petioles, hirsute when young, with slender simple hairs 2-4 mm long, and an understory stellate and of minute cm long (progressively smaller upward), triangular crenate,

acute, with

appressed

simple

and pedicels green and densely mm long, stout scabrid hairs 1 glandular hairs. Leaf blades to 8 or hastate, truncate, obscurely hairs above, stellate hairs beneath; petioles

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VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

94

subequal to the blades; stipules 5-6 mm long, filiform. Flowers solitary in the leaf axils, more or less aggregated apically; pedicels 2-3 cm long in flower to 8 cm long in fruit, 10-ribbed, in fruit; calyx 5-6 mm (to 7 mm) long, slightly accrescent mm long, yellow, a dark the 8-9 without corolla center, petals densely pubescent; on otherwise column ca. of staminal hirsute claw, glabrous; prominently margins 1-1.5 mm the anthers ca. 20, the filaments 4.5 mm long, pallid, densely hirsute, mm ca. ca. in 9 13. Fruits diameter, oblate, densely pubescent; long; styles ca. 13, with a dorsal spine ca. 1mm long, the lateral walls evanescent, the mericarps reticulate endocarp weakly developed and not enclosing the seed; seeds ca. 2.3 mm number unknown. long, glabrous. Chromosome from Baja California Anoda hirta is known only from the type and a collection in habitats. both arid evidently gathered (Brandegees.n.) (Fig. 16), & Arnott, Bot. Beechey lanceolata Hooker voy. 411. 1838. Sida nom. 4: D. Dietrich, nov.?Type: Mexico. 858. 1847, Syn. pi. s.n. to San Bias, 1828, Lay Nayarit: Tepic (holotype: K!). Anoda wrightii A. Gray, Smithonian Contr. Knowl. 5 (art. 6, PI. Wright, pt. 2): on the summit of mountains near 22. 1853.?Type: New Mexico: U.S.A.

72. Anoda

unidentata

the copper mines, 1851,Wright [894] (holotype: GH!; isotypes: BM! K! MO! NY! PH! US!). Leaf blades mostly and roughly pubescent. herbs or subshrubs, minutely cm to to truncate cun?ate, lanceolate, remotely (-12) long, ovate-triangular and obscurely dentate or crenate to subentire, acute, with simple appressed hairs Erect

3-7

somewhat discolorous; stellate hairs beneath, above, minute petioles usually less than 1 cm long but much longer on lower leaves. Flowers solitary in the leaf axils or stellate up to 4 (-6) cm long, minutely forming a terminal panicle; pedicels ca. mm or 5 below and articulated the with without scabrid hairs, flower; pubescent to 9 mm long in fruit, minutely calyx 6 mm long, accrescent stellate-pubescent; a dark center, the petals 9-16 mm long, yellow; staminal column corolla without stellate-pubescent; styles 10-12. Fruits (Fig. 151) ca. 9 mm in diameter, densely 10-12, dorsally spined (the stellate-pubescent (the hairs 0.5-1 mm long); mericarps seeds 3 mm the lateral walls evanescent; spine 1-1.5 mm), long, minutely on mar excrescences in reticulate with enclosed verruculate, prominent endocarp number unknown. gin. Chromosome in western Mexico Anoda lanceolata occurs principally (Fig. 16), extending to Arizona, northward New Mexico, and trans-Pecos Texas, often in dry habitats and disturbed It apparently the year. The type flowers throughout vegetation. stems in of A. A. and pedicels of differ that the lanceolata and specimens wrightii of the latter and are the former lack the stout scabrid hairs that are characteristic intermingled with the smaller stellate hairs found in both. There appear to be no other significant differences. Specimens 912.?Coahuila:

Examined. Purpus

6678; Jones 22854; Lay et al. 15328; McVaugh Fryxell 83148.

Baja

California:

5014?Sinaloa: s.n.; Palmer 17899.?San

Brandegee & Breedlove

2021.?Jalisco: Luis

Potos?:

Lottl425; Rzedowski

48. ?Chihuahua:

6859; White Spellenberg 17903. ?Nayarit: Gonz?lez-Ortega Hinton Pringle 4352, 5454.?Michoac?n:

Thome

4387.?Guerrero:

Hinton

7113; Koch

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&

1988

MALVACEAE OF MEXICO

73. Anoda leonensis Fryxell, Aliso Le?n: between Linares &

95

11: 504. fig.5. Mexico. 1987.?Type: Nuevo 15 Oct 1969, Fryxell 1207 (holotype:

Iturbide,

MEXU!; isotypes: BH! CTES! MICH! US! pf!). the stems minutely stellate herbs 1-2 m tall with spreading branches, cm to 4-9 Leaf ovate-cordate below blades pubescent. mostly long, hastately 3 truncate to cordate, coarsely crenate, acute, sparsely lobed and narrower above, to glabrate above and beneath; petioles V?-1 times as long as the stellate-pubescent Erect

an open terminal raceme or panicle; pedicels 2-6.5 cm long, Inflorescence corolla without a 5-6 slender; calyx yellowish pubescent; (-8) mm long, minutely dark center, the petals 8-12 (-20) mm long, yellow to yellow-orange, prominently ciliate on claw, otherwise glabrous; staminal column 2-3 (-5) mm long, pallid, with 5 vertical rows of setae, the filaments 1-2 (-4).mm long; styles 12-16. Fruits (Fig. mm 8-9 in diameter, oblate, hirsute; mericarps 12-16, dorsally 15B) densely

blades.

in rounded (crested but scarcely spurred), the midvein ("crest") darkly pigmented in a immature fruit, the lateral walls evanescent; seeds 2.5 mm long, enclosed number: 2n = 30. persistent reticulate endocarp. Chromosome occurs in and in deciduous forest in Nuevo Le?n Anoda matorral leonensis dry and Tamaulipas to the south in the northern extremity of the Sierra Madre Oriental m. at to of Monterrey 600 It flowers in 1200 October and November. (Fig. 16) Specimens 1697; Lacas

Examined.

520; Pringle

Nuevo 3016;

Le?n:

Dorr

et al. 1999,

Smith M580.?Tamaulipas:

2035, 2606; Fryxell Bartlett 10168,10378.

1207,

1212; Fryxell

et al

74. Anoda mac?lala 11: 506. 1987.?Type: Mexico. Guerrero: Mpio. Fryxell, Aliso carr. a Cd. Altamirano, 40 km NE of Vallecitos, de La Uni?n, 25 Oct 1982,

Koch & Fryxell 82191 (holotype: CHAPA!; isotypes: ENCB! F! MEXU! MICH! MO! US! pf!). herbs ca. 1 m tall, puberulent. Leaf blades mostly sively narrower and hastately late or remotely serrate, acute, as long as the blades. Flowers Erect

the stems and petioles minutely and evenly stellate 3-7 cm long, broadly ovate below, becoming progres 3-lobed above, truncate to cordate, ciliate and undu glabrate above, nearly so beneath; petioles V4?1 times

solitary in the leaf axils or forming a terminal leafless cm 2-8 hirsute (hairs at base raceme; pedicels long; calyx 6-8 mm long, prominently to 1mm long), the lobes with a prominent midrib; corolla with a dark red center, the petals 6-8 mm long, pale yellow or cream; staminal column ca. 2 mm long, reddish or ribbed, glabrous (or with a few apical hairs), the filaments less purplish, prominently than 1mm long, purple, the anthers reddish; styles ca. 10, pallid. Fruits ca. 7 mm in ca. 10, the diameter, oblate, hirsute, with radiating spurs ca. 1mm long; mericarps lateral walls evanescent, seeds ca. 3 mm long, completely in a persistent enclosed reticulate endocarp. Chromosome number unknown. Anoda maculata is known from central Mexico (Fig. 1300 to 2000 m. It flowers principally from July to October. Specimens 17333; Pringle

Examined. 4575.?

Durango:

Puebla:

Fryxell

Shreve

9164.?Quer?taro:

189,1137.?

Guerrero:

Koch

16) in deciduous

Arguelles & Fryxell

1434.?Jalisco: 82161,

82191.

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forest at

McVaugh

96

SYSTEMATIC BOTANY MONOGRAPHS

FIG. 79278);

17. Anoda

separate

larger

stem Habit and enlarged palmata. leaf (Koch & Fryxell 83144); separate

segment smaller

VOLUME 25

(Koch showing pubescence leaf (Fryxell et al. 1631).

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et al.

1988

97

MALVACEAE OF MEXICO

11: 507. fig.6. 1987.?Type: Mexico. Michoac?n: palmata Fryxell, Aliso ter km W 44-45 de La Uni?n, de al de Guerrero, Coahuayutla Mpio. 14 Nov 570 m, raceria a Coahuayutla, 1983, Koch & Fryxell 83144

75. Anoda

(holotype: CHAPA!; MO!pf!).

isotypes: BM! CTES! ENCB! F! MEXU! MICH!

and pedicels widely branching herbs 1-1.5 m tall, the stems, petioles, a sometimes and with also with few stellate-pubescent glandular hairs, long simple hairs. Leaf blades mostly 3-8 (-10) cm long, variable, the juvenile ones ovate or more deeply palmately 5-lobed (ultimately shallowly lobed, becoming progressively the lobes linear-lanceolate, subentire, with narrowly 3-lobed) upward, becoming more or less appressed hairs above, evenly stellate-pubescent beneath; peti simple oles Vi-1 times the length of the blades. Flowers solitary in the leaf axils, forming an open terminal raceme more or less above the leaves; pedicels 2-5 cm long, slender; densely covered with stellate and glandular calyx 5-8 mm long (not accrescent), a dark the petals 6-9 mm long, yellow; staminal center, hairs; corolla without Erect

2-3 mm long, pallid, prominently hirsute apically, the filaments 1-2 mm mm in 8-13. Fruits 8-9 diameter, oblate, densely stellate-pubescent; long; styles 8-13, dorsally umbonate mericarps (the spur 1mm long), the lateral walls evanes reticulate endocarp 2.5-3 mm long and completely cent, the persistent enclosing the seed. Chromosome number: 2n = 30. Fig. 17. and in southern Baja California Anoda palmata occurs in west-central Mexico m in 600 below forest and open, arid habitats. deciduous It (Fig. 18), generally to December. flowers from October column

Examined.

Specimens

Baja

California:

Brandegee

49; Carter

2590;

Carter & Kellogg

3160;

Carter

& Moran 5389;Moran 6892, 6924, 7064; Purpus 338; Sanders et al. 3437;Wiggins 15571,15658; Wiggins etal.

315, 502.? &. Fryxell

Koch

Nayarit: 77520,

Feddema 8375,

83144,

1228.?Jalisco: 83215;

Soto-N??ez

Fryxell et al. 1598.? Michoac?n: Fryxell et al. 1631; etal. 11256.? Guerrero: Koch & Fryxell82151,

83144.

76. Anoda

paniculata 1916.?Type:

Annuaire Conserv. Jard. Bot. Gen?ve 20: 53. Hochreutiner, Mexico. 13 Oct 1906, Pringle Guerrero: Iguala Canyon,

10323 (holotype: G; isotypes:BM! CM! DS-fragment! F! as photo F-56123! GH! K! MEXU! MIN, MO! NY! PH! UC! US! VT!). robust herbs to 2 m tall, the stems, petioles, and pedicels little-branched, and tomentose. Leaf blades mostly 6-8 cm long, broadly ovate minutely evenly or lobed below, progressively more deeply lobed to hastately pentangular shallowly Erect,

the margins and main veins beneath above, remotely serrate to subentire, otherwise sometimes with purplish blotch along scabridulous, minutely glabrate, mid vein; petioles V3-I times as long as the blades. Inflorescence an open terminal 1-3.5 (-6.5) cm long; calyx 6-9 mm long in flower, accrescent to raceme; pedicels 12 mm long in fruit, 10-ribbed, densely pubescent; a dark center, corolla without trilobed

long, white or rose drying to pink or lavender (the claw staminal column 3-5 mm hirsute remaining white); long, whitish, prominently mm the filaments 3-4 11-15. mm Fruits 8-9 in apically, long; styles (Fig. 15C) the petals

10-19

mm

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98

VOLUME 25

SYSTEMATIC BOTANY MONOGRAPHS

4a.

V\

-A

'A

^

?\

?i

Anoda paniculata

Anoda palma ta

Vfr.

icv >m

^

Anoda

Anoda pedunculosa

^

pentaschista

li.

Anoda

\

Anoda

polygyna

pr?stina

A

Anoda pubescens

Anoda reticulata

FIG.

diameter, eral walls 2.5-3

mm

18. Distribution

of species

oblate, stellate-tomentose; mericarps enclosed seeds completely evanescent; long.

Chromosome

number

of Anoda.

the lat 11-15, dorsally umbonate, in a persistent reticulate endocarp

unknown.

occurs in the Balsas Anoda paniculata open matorral at 300 to 1500 m and flowers

Basin (Fig. 18) in deciduous inOctober and November.

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forest and

1988

99

MALVACEAE OF MEXICO

Specimens

Examined.

Michoac?n:

Koch

& Fryxell8357.?M?xico:

Guizar

249; Hinton

?tal.

1926,

5025; Koch & Fryxell 82247.? Guerrero: Koch & Fryxell 82238; Mexia 8726; Pringle 10323, 13691, 13694.

Conserv. Annuaire Jard. Bot. Gen?ve 20: 11. Anoda pedunculosa Hochreutiner, near 5 Oct 6541 Mexico. 60. 1916.?Type: Tula, 1896, Pringle Hidalgo:

(holotype: G?; isotypes: CAS! CM! F! as photo F-56122! GH! K! MASS! MEXU! MICH! MIN, MO! NY! PH! UC! VT! WIS!).

and evenly stellate herbs or subshrubs 0.5-1.5 m tall, the stems minutely cm ovate 3-7 Leaf blades (juvenile leaves) to long (often smaller), puberulent. narrower crenate, 3-lobed, hastately upward, obscurely becoming progressively a with and sometimes acute, minutely purple blotch sparsely stellate-pubescent, as as the V4-1 times Flowers blades. solitary in the leaf long along midvein; petioles a or or raceme more in less above the leaves; often axils, panicle aggregated apically mm cm 1-4 4-6 pedicels long; calyx long (not accrescent), minutely puberulent; a dark center, the petals 6-15 mm long, whitish to lavender; stam corolla without inal column 3-4 mm long, pallid, with long transparent hairs distally, the filaments 2 mm long, the anthers and pollen yellow; styles 12-13, sometimes reddish. Fruits mm whitish 8-9 in diameter, oblate, 12-13, with a pubescent; mericarps (Fig. 15E) short umbo on dorsal wall, the lateral walls evanescent; seeds 3 mm long, com in a persistent reticulate endocarp. Chromosome number: 2n = 30. pletely enclosed Erect

occurs in eastern Mexico Anoda pedunculosa from Tamaulipas in matorral deciduous forest and open 18) generally below 900 m. or less throughout the year.

to Oaxaca

(Fig. It flowers more

et al. 734; Johnston examined. Tamaulipas: & Fryxell & Bates 946, 948; Hern?ndez Luis Potos?: Fryxell & Anderson 4557; Nee 22248.?San 3539; McGregor 16328; Rzedowski et al. 1709, 7020.? Hidalgo: & Justo 1436; Dorantes 6541?Veracruz: Calzada 6391; Ch?zaro Pringle et al. 1989; Fryxell 2551; Johnston & 1731, 1806; Dorr 53-687; Judziewicz $187; Linden 832; Nevling Velez 20; Ventura 2665,11765,14385.? 52; Ortega 437; Paray 2883; Purpus 1948,16311; G?mez-Pompa Specimens

Graham

Oaxaca:

Fryxell

1152; Koch

& Fryxell

78326.

78. Anoda

Contr. Knowl. 5 (art. 6, PI. Wright, pentaschista A. Gray, Smithsonian & Standley, pt. 2): 22. 1853. Sidanoda pentaschista (A. Gray) Wooton Contr. U.S. Nati. Herb. 19: 427.1915.?Type: U.S.A. New Mexico: valley

between Ojo de Gavil?n andCond?'s Camp, 30Aug 1851,Wright 385 [893] (holotype: GH!; isotypes:BM! GH! K! MO! NY! UC! US!). FI. Mex. 171. ed. ii. 1894, non Sida Sess? & Moci?o, integrifolia Mexico. 1860.?Type: Sess? et al. Guanabacoa, Montrouzier, integrifolia location "Sida unknown; integerrima" (holotype: [3498] isotype?: F!). Sida palmeri J. G. Smith, Rept. Missouri Bot. Gard. 6: 113. t.48. 1895, non var. obtusior Robin Sida palmeri E. G. Baker, 1892. Anoda pentaschista son in A. Gray, Syn. fl. N. Amer. U.S.A. Texas: 1(2): 320. 1897.?Type: 1894, Nealley s.n. (F! GH! US!). Corpus Christi, extrema Hochreutiner, Anoda Conserv. Jard. Bot. Gen?ve Annuaire 20: 64. 1916.?Type: "Nova Hispania 1827," ex herb. Pav?n s.n. sub. nom. Sidae heterophyllae (holotype: G as photo F-23687!). Sida

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SYSTEMATIC BOTANY MONOGRAPHS

100

1-2 m

VOLUME 25

but widely branching, the stems Leaf often becoming glabrescent, purplish pigmented. stellate-pubescent minutely or or more ovate and in form the lower leaves variable blades coriaceous, size, highly less 3-lobed, remotely and irregularly dentate, the upper leaves narrowly oblong or toothed at base, otherwise linear, sometimes virtually entire, markedly hastately Erect

subshrubs

tall, single-stemmed

and gray-green discolorous stellate-tomentose stellate above, densely (sparsely tomentose and whitish beneath); petioles commonly V4the length of the blades (but a lax panicle; pedicels 1.5-3.5 cm up to ?3 as long in juvenile leaves). Inflorescence long, slender, articulated near middle; calyx 3-5 mm long, minutely pubescent with corolla without a dark center, the petals ca. 1 cm long, darkly pigmented midvein; and sparsely scabrid toward yellow; staminal column ca. 3 mm long, pallid, minutely apex, otherwise glabrous, the filaments 1.5 mm long, the anthers yellowish; styles 5 Fruits (Fig. 15D) 4-5 mm in diameter, androecium. 8, pallid, slightly exceeding oblate, minutely walls evanescent; 30.

5-8, ind?hiscent, dorsally spurred, the lateral pubescent; mericarps seeds ca. 2 mm long, endocarp absent. Chromosome number: 2n =

occurs widely inMexico at low elevation pentaschista (Fig. 18), usually as high as 1500 m. It is commonly found in disturbed 600 m but occasionally situations, such as roadsides and agricultural fields, and flowers throughout the year. Anoda

below

Baja California: Carter 5067; Carter & Moran Specimens Examined. 5290, 5352, 5532; Moran & Thome Sonora: Breedlove 18625; 18632; Gentry 304, 11375, 11442, 15182, 15518.? 18864; Wiggins et al. 20071; Palmer 192.? & Rollins 306, 758, 19250; Norris 265, 661, 780; Rodr?guez 1618; Wiggins Sinaloa: 5928; Palmer 139, 540; Fryxell & Bates 2100, 2115; Gentry 7120; Gonz?lez-Ortega 198; Fryxell et al. 1179.?Zacatecas: 1314L? Durango: Rose 3762.?Tamauli 1594; Wiggins Rodriguez Rodriguez pas: Johnston

et al. 1616; Martinez 7632?-Colima: Rowell 4391, 4935B.?Jalisco: Fryxell [Barkley] Rose & Painter 6603; Michoac?n: 1727; Fryxell & Bates 2155; Pringle 8505.? Morelos: Fryxell et al. 5857, 6697.?Veracruz: 716.? Oaxaca: Hinton 3300.? Guerrero: 752, Fryxell Fryxell V?zquez 4432.?

1158;Fryxell & Bates 907; King 860. polygyna Fryxell, Aliso de Sinaloa y Vela, Mpio.

79. Anoda

11: 512. fig.7. Mexico. Sinaloa: 1987.?Type: at El Alamo, 3 Mar Sierra Surutato 1971,

Breedlove 19033 (holotype: CAS!; isotypesENCB! F! pf!). the stems, petioles, herbs or subshrubs, and pedicels little branched tomentose Leaf blades 3-6 densely yellowish (with glandular hairs interspersed). cm long, angularly ovate to hastately 3-lobed, entire, densely stellate- pubescent above and beneath, sometimes red-margined; petioles subequal to the blades. Inflo rescence a terminal more or less leafless raceme; pedicels 2-5.5 cm long; calyx 9-11 Erect

mm

tomentose with glandular hairs inter long (not accrescent), densely yellowish a dark center, the petals 15-23 mm spersed, the lobes 1-veined; corolla without or pale lavender; staminal column ca. 5 mm long, pallid, prominently long, pink hirsute, the filaments 3-4 mm long; styles 17-20, pallid, the stigmas reddish. Fruits oblate, densely hirsute; mericarps 17-20, dorsally (Fig. 15A) 8-9 mm in diameter, seeds 3.5 mm long, spurred (the spurs 0.5-1 mm long), the lateral walls evanescent; num in persistent reticulate endocarp. Chromosome enclosed glabrous, completely

ber unknown. Fig. 19. Anoda polygyna occurs in Sonora, ate elevations, in deciduous apparently

Sinaloa, forest.

and Durango

(Fig. 18) at intermedi

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1988

19. Anoda

FIG. 19033.)

See Fig.

Specimens

polygyna. 15A for fruit.

Examined.

MALVACEAE OF MEXICO

Flowering

Sonora:

Gentry

and fruiting

1234.?Sinaloa:

branches,

101

and separate

Breedlove

lobed

leaves.

19033.?Durango:

(Breedlove

Hurd

56.

Mexico. pr?stina Fryxell, Syst. Bot. 4: 253. 1979.?Type: Chiapas: Mpio. on Pinus of with and Quercus in the Kulak'tik, slope Tenejapa, paraje barrio of Chana', 7546 (holotype: DS!; 4800 ft, 24 Nov 1964, Breedlove isotypes: F! MICH!).

80. Anoda

Perennial herb or subshrub, the stems reddish, with short pungent some glandular hairs, becoming glabrate. Leaf blades to 7 cm long, wider

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hairs and than long

SYSTEMATIC BOTANY MONOGRAPHS

102

VOLUME 25

on and palmately 3 (-5)-lobed, subentire, acute, glabrate except appressed-ciliate antrorse than with shorter the blades, hairs; discolorous; margins, petioles stipules 2-3 mm long. Flowers solitary in the leaf axils; pedicels 1-9 cm long, evenly hispid, to 12-14 the hairs more or less antrorse; calyx 5-6 mm long in flower, accrescent mm in fruit, nearly glabrous, with 10 relatively prominent nerves; corolla without a ca. 4 mm dark center, the petals 7-9 mm androecium long, long, purplish; or more ca. mm 1 less the filaments long, glabrous, pallid, grouped into 5 fascicles; Fruits in diameter, oblate, styles 12-13. (Fig. 15N) 10-13 mm subglabrous; the lateral walls evanescent but with a persis 12-13, apically dehiscent, mericarps tent coarse seeds 3 mm but with short reticulum; long, seemingly glabrous number unknown. absent. Chromosome appressed pubescence, endocarp Anoda pr?stina is known only from the type (Fig. 18). 81. Anoda

pubescens

Schlechtendal,

Linnaea

11: 218.

1837.?Type:

Mexico.

Hidalgo: Mineral del Monte, Ehrenbergs.n. (holotype: HAL?). 33: 42.1974.?Type: Mex Per?ptera grandiflora Fryxell, Bol. Soc. Bot. Mexico ico. M?xico: Mpio. de Texcoco, 11 Nov 1951, Gold Cerro de Purificaci?n, s.n. (holotype: MEXU!). m tall, the stems, petioles, and pedicels 0.5-1.5 sparsely Leaf blades 3-8 cm long, broadly ovate below to more nar 3-lobed serrate, acute or above, deeply cordate, rowly triangular or hastately and above and acuminate, minutely beneath; petioles sparingly stellate-pubescent subequal to the blades below, shorter upward. Flowers solitary in the leaf axils or 1-5 cm long in flower (slightly longer in fruit); forming a terminal raceme; pedicels mm 6-12 the lobes 1 stellate-tomentose, calyx long, densely deeply 5-lobed, a more or dark less corolla without the center, nerved, apiculate; petals 7-14 mm mm on or staminal 7 column more, long, lavender (white claw); long subequal to Perennial

herbs

stellate-scabridulous.

the petals, whitish, apically lavender; styles 8-10. Fruits minutely stellate-pubescent; the lateral walls evanescent; some number: 2n = 30. occurs Anoda pubescens cent states

at 2100 to 2700 m and in adja in Hidalgo principally and flowers from July to November. (Fig. 18), primarily in dry matorral,

Specimens berg s.n.;

Examined.

Fryxell

23321, 23907, Gold s.n.

the filaments ca. 2 mm long, the anthers pubescent, mm in diameter, oblate, densely and 7-8 (Fig. 15K) 8-10, with a dorsal spur ca. 0.5 mm long, mericarps mm 3 seeds long, glabrous, endocarp absent. Chromo

San Luis

et al. 1675; Medina

24278,

26848,

29166,

Potos?: 1808, 29472;

Parry

& Palmer

2238; Pringle Ventura 266,

6969; 360,

78; Rzedowski

3991. ?Hidalgo:

Ehren

1685; Rzedowski 20621, 22952, Purpus 3671.? Puebla: 2610.-M?xico: Purpus

82. Anoda

Proc. Amer. Acad. Arts 17: 368. 1882.?Type: reticulata S. Watson, Arizona: Santa Catalina Mountains, U.S.A. May 1881, Lemmon & Pringle s.n. (holotype: GH! as photo CAS!).

Erect herbs ca. 1m tall, the stems and pedicels minutely scabridulous and with some glandular hairs. Leaf blades mostly 3-6 cm long, typically 3-lobed (rarely reduced the lobes narrowly linear (1-5 mm wide), progressively obscurely 5-lobed), in the bracts to to and leaves filiform upwards ultimately stipuliform simple

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1988

103

MALVACEAE OF MEXICO

to subentire, with purplish blotch and beneath; petioles often equal stellate-pubescent along midrib, minutely Flowers or shorter the blades, solitary in the leaf axils or upwards. exceeding ing cm to a 8.5 terminal slender; raceme; calyx ca. 5mm long long, pedicels up forming a corolla without with lobes the midrib; strong densely pubescent, (not accrescent), a dark center, the petals 5-6 mm long, bluish purple; staminal column ca. 2 mm long, glabrous or with a few hairs, pallid, the filaments purplish, the anthers yellow ish; styles 10-11. Fruits (Fig. 15F) 6-7 mm in diameter, oblate, stellate-pubescent; 10-11, the dorsum rounded (completely lacking spur or spine), the lat mericarps in a persistent enclosed the seed completely eral walls evanescent, coarsely reticu number unknown. late endocarp 2.5 mm long. Chromosome in Sonora is known from Sinaloa, Anoda reticulata (Fig. 18), and Arizona midstem

inflorescence;

leaves

remotely

serrate

above

desert habitats. Examined.

Specimens 7373A.?

Sinaloa:

83. Anoda

Sonora:

Gentry

818,

1143,

1355,

3043a,

19296;

White

3618;

Wiggins

Brandegees.n.

speciosa Fryxell, Aliso

11: 516. fig.8.

1987.?Type:

Mexico.

M?xico:

hills

N of Toluca, 23 Sep 1900, Pringle 9249 (holotype: VT!; isotypes: GH! US!). and pedicels evenly and minutely the stems, petioles, Erect herbs or subshrubs, on one side. Leaf blades up to sometimes purplish pigmented stellate-scabridulous, 10 cm long (smaller and narrower 3-lobed, hastately deeply cordate, upward), acute or acuminate, to subentire, crenate-dentate minutely stellate-pubescent above and beneath; petioles Va-Va as long as the blades. Flowers solitary in the leaf inflorescence; axils, more or less aggregated pedicels up to apically in a racemiform stellate pubescent, with 5.5 cm long; calyx (in flower) 12-15 mm long, minutely mm a the 22-28 dark without corolla lanceolate-acuminate center, lobes; petals mm to ca. androecium the the 20 column staminal subequal long, long, lavender; a the filaments 3-5 hairs few minute but with essentially glabrous, pallid, petals, mm Fruits lavender styles unknown. long, the anthers (?), the pollen orange; number unknown. Chromosome is Anoda known speciosa only from the type collection ca. at in oak-pine forest. 2600 m, 20) presumably

unknown.

84. Anoda

succulenta

Cieneguita,

11: 518. 1987.?Type: Fryxell, Aliso R?o Mayo, canyon, Upper Sonoran,

from near Toluca

(Fig.

Mexico. Chihuahua: La 10 Sep 1936, Gentry 2638

(holotype: ARIZ!). Erect succulent herbs 1.5 m tall, the stems hollow, essentially glabrous. Leaf blades up to 11.5 cm long (smaller upward), palmately 3-5-lobed, basally cordate lobe the sinuses central (the broadly ovate, basally narrowed (the narrowly acute), sinuses rounded), subequal to the blades; stipules 5-7 mm long, glabrous; petioles ciliate. Flowers solitary in the leaf axils; pedicels up to 13 cm long (subequal to the at anthesis); leaves calyx 10-12 mm long, externally glabrous or with a subtending few pungent hairs, internally woolly; corolla without on margins mm bearded densely long, yellowish,

a dark center, the petals 15-16 of claw, otherwise glabrous;

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SYSTEMATIC BOTANY MONOGRAPHS

104

FIG.

20. Distribution

of species

of Anoda,

Anotea

flavida,

and species

VOLUME 25

of Bakeridesia.

staminal column ca. 3 mm long, pallid, with a few hairs apically, filaments 2-3 mm styles pallid, of uncertain number. Fruits unknown. long, the anthers yellowish; Chromosome

Anoda

number

succulenta

unknown.

is known

only

from the type and Gentry

6518

from Sinaloa

(Fig. 20).

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1988

MALVACEAE OF MEXICO

105

85. Anoda

thurberi A. Gray, Proc. Amer. Acad. Arts 22: 299. 1887.?Type: Mex ico. Chihuahua: 3 Oct 1885, Pringle 283 (lecto Santa Eulalia Mountains, 1916: GH!; BM! BR! type, designated by Hochreutiner, isolectotypes:

COLO! DS-2! F! K! MASS! NY-2! PENN! PH! RSA! UC! US! VT! WIS! pf!). Erect herbs ca. 1m tall, the stems, petioles, and pedicels minutely stellate- and Leaf blades 5-8 cm long (often smaller), ovate-cordate to glandular-pubescent. to narrowly hastately 3-lobed triangular, sometimes with a purplish blotch along and a purplish margin, minutely midvein above and beneath, the hairs pubescent usually stellate; petioles Vi-i times as long as the blades. Inflorescence commonly an open terminal raceme or panicle borne more or less above the leaves; pedicels 1 3 cm long, slender; calyx 3.5-6 mm long in flower, accrescent to 6-8 mm long in the lobes somewhat inrolled in fruit; corolla with a pur fruit, stellate-pubescent, plish center, the petals 4-7 mm long, bluish lavender; staminal column 1.5-2 (-4) mm long, stellate-pubescent, less than 1mm long, pallid or lavender, the filaments the anthers few, lavender; styles 6-8, the stigmas barely exceeding the androecium, cl?vate. Fruits (Fig. 15L) 6-8 mm in diameter, oblate, minutely stellate minutely and glandular-pubescent; or up obscure 6-8, spur mericarps dorsally spurred (the to 1 mm seeds 2-2.5 mm long), the lateral walls evanescent; long, glabrous, numbers: 2n = 26, endocarp absent or fragile and incomplete at best. Chromosome 28. Anoda thurberi grows mostly in deciduous forest and dry matorral from Chihua hua to Tamaulipas south to Guerrero and Oaxaca (Fig. 20). It also occurs in southern Arizona and New Mexico. It flowers from August to November. Anoda thurberi may be a bridging species between the genera Anoda and Per?ptera (Bates

1987). Examined.

Specimens Shreve

8869; 1745.?Durango:

7744.?Jalisco:

Stewart

2296.?

Sonora: Coahuila:

372.?Chihuahua: Lloyd Henrickson 13245;

Fryxell & Bates 2066; Dieterle 3076. ?Guanajuato:

& Tryon 1997.? Michoac?n: Arguelles Potos?: Rzedowski 6768, 9442.?

Rollins

Fryxell

832?Hidalgo:

6536.?Morelos: 1270, 2067. ?Guerrero: Pringle V?zquez Puebla: Conzatti 1138; Purpus 4099, 4187.? Oaxaca: Fryxell

Anotea

Luis

13025;

283, 2349; Pringle et al. 1261; Wendt Rose & Painter

2558; Stewart Aguascalientes:

Tamaulipas: et al. 7983.?

et al. 1694.?San

Henrickson Paray 58286.?

Koch

& Fryxell

8356.?

Quer?taro:

548, Arguelles & Bates 2164; Koch

Fryxell & Gonz?lez

29; Smith

641.

Ind. sem. h?rt. bot. berol. 1846. 13. 1846. Malvavis Kunth, (de Candolle) cus sect. Anotea de Candolle, Prodr. 1: 445. 1824.?Lectotype: Anotea De Ulbrich. Candolle included four species in sect. flavida (de Candolle) Anotea. Of these, only A. flavida was retained by Ulbrich when he treated the taxon at generic rank. This choice constitutes lectotypification.

to Don (1831), Anotea derives from the Greek a, with Etymology. According "not sufficiently known." out, and notus, known, meaning Anotea is a monotypic Mexican genus. Kunth (1847) and Ulbrich (1915) in cluded a second Mexican A. was but it trans chlorantha, species, subsequently ferred to Pavonia.

References: Kunth (1847), Ulbrich (1915), Kearney (1951a), Fryxell (1968b).

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106

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

86. Anotea

flavida (de Candolle) Ulbrich, Repert. Spec. Nov. Regni Veg. 14: 109. 1915. Malvaviscusflavidusde "Hibis Candolle, Prodr. 1:446.1824.?Type: no. 376! (Torner Collection cus flavidus" Icones Florae Mexicanae ace. no. 6331.389, Hunt Institute). [Cf. photo F-30525 of G copy (not the type).] K. 2: 135. 1835.?Type: Malvaviscus Presl, Mexico, acerifolius Reliq. Haenk. as W BM! MO! Haenkes.n. PR; isotypes: photo F-32652!). (holotype:

2-3 m tall, with stellate puberulence. Leaf blades mostly 10-17 cm as as to wide about cordate, 5-lobed, shallowly long, pentagonal long, coarsely without foliar nectaries; petioles almost as long as blades below to crenate-dentate, race much shorter upward; deciduous. Inflorescence linear-lanceolate, stipules mose (or sometimes paniculate); involucel pedicels axillary; basally gamophyllous, the bractlets about equaling the calyx in fruit; calyx 12 10-12, linear-acuminate, mm long, half-divided; petals 25 mm long, cream-yellow with darker veins, without Shrubs

basal auricles, erect, forming a tubular corolla; androecium greatly exserted (twice the length of the corolla in bud to 4 times its length at anthesis), pallid, sparsely stamens 20-40, yellow, in upper half of column, the pubescent, apically 5-dentate; filaments 2-3 mm long; styles 10, diverging for 6-7 mm above androecium; stigmas capitate. Fruits schizocarpic (a fleshy berry), 1 cm in diameter, oblate, blue-black at number: 2n = 56. Fig. 21. the 5 carpels each 1-seeded. Chromosome maturity, is known only from central Guerrero Anotea flavida (Fig. 20) at an elevation of ca. 900 m (Fryxell & Koch areas in It in oak-pine forest and grows open 1987). to January. flowers from October Specimens

Examined.

Guerrero:

Croat

45715;

Fryxell

623; Fryxell

& Bates

2168; Germ?n

& Funk

622;Koch & Fryxell 83254; Koch et al. 79125;Kruse 138. Jard. Bot. Gen?ve Annuaire Conserv. Hochreutiner, Bakeridesia 1913.?Type: galeottii (E. G. Baker) Hochreutiner.

Bakeridesia

15/16:

297.

often ferrugineous. Shrubs or small trees 1-8 m tall, densely stellate-pubescent, Leaves petiolate, the blades broadly ovate, cordate (or narrower and truncate in acute or acumi the inflorescence), entire (or sometimes with vestigial dentation), more so beneath, above and nate, minutely beneath, densely stellate-pubescent or foliar nectaries. Flowers in the sometimes discolorous, solitary paired lacking leaf axils or aggregated into axillary or terminal inflorescences; involucel absent; sometimes twisted in bud; petals usually ribbed, calyx lobes usually prominently often with a dark large (1.5-6 cm long) and showy, white, yellow, or yellow-orange, at reddish spot at the base; staminal column included or exserted, antheriferous apex, glabrous or pubescent; styles 7-27, the stigmas capitate. Fruits schizocarpic, more or less blackish, 7-27, rounded, often with a stellate-pubescent; mericarps lacerate or erose wing on the dorsal margin, 2-7-seeded; seeds reniform, pubescent or glabrate. Base chromosome number: x = 15. was named to honor Edmund Gilbert Baker (1864 Bakeridesia Etymology. 1949), British botanist and author of "Synopsis of Malveae." and Central American Bakeridesia is a genus of thirteen Mexican species (subg. to Venezuela, and seven or more addi with one species extending Bakeridesia)

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FIG.

107

MALVACEAE OF MEXICO

1988

21. Anotea

flavida.

Flowering

branch,

mature

leaf, and mature

fruit.

(Koch & Fryxell

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83261.)

SYSTEMATIC BOTANY MONOGRAPHS

108

VOLUME 25

tional South American Ten species that may or may not be congeneric. occur inMexico (Figs. 20,23). References: Hochreutiner (1920b), Kearney (1951a), Bates (1973). to the Mexican

Key

Species

species

of Bakeridesia

less than 4 cm long; calyx 6-18 mm long. usually or absent. 2. Wing on dorsal margin of mericarp rudimentary 3. Petals 1.5-2 cm long, yellow staminal column throughout; 9 mm long; Yucat?n, Roo. B. 89. Quintana

1. Pedicels

3. Petals

cm

2.5-3.5

long,

yellow

with

9-11 mm

red spot at base; 94.

long; Chiapas. mericarps on dorsal margin 2. Wing of mericarp manifest, 4. Pedicels less than 1 cm long. usually 4. Pedicels

usually

6. Calyx

(10-)

ca.

10 mm

long;

pittieri.

lacerate. 95.

B.

mostly

pedicels

more 15-20

1-4 cm long, erect; Tamaulipas,

mostly

pedicels

5. Flowering 1. Pedicels

B.

6.5

long; mericarps

gaumeri. staminal column

subcordata.

cm long.

0.5-5

5. Flowering

4-6 mm

than 4 cm long; calyx mm

long,

cm

2.5-4.5

15-35 mm

(10-)

thin-textured;

flexuous;

long,

petals

25-40

San Luis Potos?, Veracruz. 91. B. integerrima. Jalisco,

Colima, Oaxaca.

long. mm

long; Veracruz lowland Oaxaca.

Guerrero, 87. B. bakeriana. and adjacent 88. B. ferruginea.

6. Calyx 20-35 mm long, thick-textured; petals 30-60 mm long. 7. Petals white (4 cm long, 1.5 cm wide), more or less (or pale yellow), narrowly spatulate Roo. B. yucatana. 96. Quintana erect; young growth orange-pubescent; 7. Petals rotate or reflexed; obovate (more than 2 cm wide), broadly yellow or orange, young growth more or less ferrugineous. 8. Petals

reflexed, pubescent; toward the base;

pubescent

pedicels

cm

9-15

calyx densely

wooly

long; staminal with complex

column

ca. 3 cm

stipitate

hairs;

long, southern

92. B. nelsonii. Chiapas. or glabrous; 3-12 cm long; staminal (not reflexed), pubescent pedicels cm long, glabrous or pubescent; column 1.1-3.5 calyx finely and densely pubescent. 9. Staminal column 11-20 mm long; margins of claw pubescent; basally pubescent,

8. Petals

rotate

southern

Veracruz, Oaxaca, Chiapas. column glabrous, 25-35 mm Luis Potos? and northern Veracruz.

9. Staminal

87. Bakeridesia bakeriana bakerianum Rose, Mexico.

Oaxaca:

90. long; margins 93.

B. of claw glabrous; B.

gloriosa. eastern

San

notolophium.

10: 473. 1973. Abutil?n (Rose) Bates, Gentes Herb. 5: 133. pi. 11. 1897.?Type: Contr. U.S. Nati. Herb. 1 Dec 1895, Pringle 6278 (lectotype, Tomellin Canyon,

designated by Bates, 1973:US!; isolectotypes: A! BM! BR! CAS! CM! ENCB! F! as photo F-56141, GH! GOET! K! MASS! MEXU! MIN, MSC! MO! NY! PH! UC! US! VT! pf!). The type locality is atAlmoloyas, "56mi from Oaxaca

as the train runs" (Davis

1936, p. 145).

Small trees to ca. 6 m tall, the young growth densely stellate-pubescent. Leaf blades mostly 6-11 cm long, ovate (broadly so and cordate below to narrower and or acute, slightly discolorous, truncate above), acuminate be stellate-pubescent ca. half the length of the blades neath, sparsely so to glabrate above; petioles in clusters on the ends of short lateral branches or (sometimes longer). Flowers cm long, flexuous, 2.5-4.5 stellate grouped apically; pedicels mostly mm ca. 7-12 half-divided; pubescent; calyx petals 2-3 cm long, stellate-pubescent, mm at sometimes reddish the staminal column 7-8 base; long, yellow, long, pallid, these

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MALVACEAE OF MEXICO

1988

109

I

2 cm

u FIG. mericarp

22. Bakeridesia (above)

(McVaugh

bakeriana.

Flowering

branch

(McVaugh

15724);

mature

fruit

(below)

and

15767).

the filaments 4-6 mm long; styles 11-15, glabrous. Fruits 16-17 stellate-pubescent, mm in diameter, 11-15, dorsally with narrow (1-2 mm) subglobose; mericarps lacerate wing bearing seeds 3.5 mm long, glabrate. stipitate hairs, 2-3-seeded; Chromosome number unknown. Fig. 22. Bakeridesia bakeriana in western Mexico grows in dry, deciduous woodland from Jalisco to Oaxaca (Fig. 20) at elevations up to 1400 m. The species is described of populations from three widely disjunct regions: by Bates (1973) as "composed It flowers from January to July. and Colima-Jalisco." Oaxaca, Guerrero, Specimens Magallanes Langlass?

Examined.

Jalisco:

3019, 3609; McVaugh Conzatti 249.?Oaxaca:

& Lott39; Bates 3149; Bullock & de Puga 909; 1551; Guzman 1718.?Colima: Guerrero: 15724, 15767, 23013.? McVaugh 4030; Pringle 6278; Smith 547.

Ayala & Koelz

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110

SYSTEMATIC BOTANY MONOGRAPHS

VOLUME 25

in Miller) Krapovickas, 88. Bakeridesia Bol. Soc. Argent. ferruginea (T. Martyn Bot. 7: 37.1957. inMiller, Fig. pi. gard. diet. ed. Sida ferruginea T. Martyn iii. 1: 3. t.3.f.l. de Candolle, 1824.?Type: 1798, non Sida ferruginea Miller's t.3.f.l.y reproduced by Bates (1973, fig. 2). See also McClintock

and Fryxell (1979). Abutil?n t.3.

planiflorum 1857.?Type:

K. Koch & Bouch?, Koch & Bouch?'s

Berliner plate

25: 98. Allg. Gartenzeitung 3 (lectotype, and designated

1973, fig. 1). reproduced by Bates, Abutil?n macranthum Peyritsch, Linnaea 30: 59. 1859, non Abutil?n macran thum St.-Hilaire, 1827. Abutil?n Standley, Contr. U.S. Nati. peyritschii Mexico. Veracruz: Zacuap?n, Hel Herb. 23: 751. 1923, nom. nov.?Type:

ler46 (holotype:W). G. Baker, J. Bot. 31: 73. 1893. Bakeridesia galeottii (E. G. galeottiiE. Jard. Bot. Annuaire Conserv. 15/16: 298. Gen?ve Hochreutiner, Baker) Mexico. 1913.?Type: Galeotti 4103 (lectotype, Veracruz, designated by BR! G). Bates, 1973: K!; isolectotypes:

Abutil?n

the hairs more or less 2-5 m tall, the young growth stellate-pubescent, cm 9-15 often Leaf blades mostly long, ovate (broadly so ferrugineous, stipitate. to narrower somewhat dis and cordate below and truncate above), acuminate, and above sometimes colorous, beneath, minutely glabrate stellate-pubescent ca. half the length of the blades or sometimes above; petioles longer. Flowers solitary or paired in the leaf axils; pedicels 4-9 cm long, slender, the buds nodding; Shrubs

the densely pubescent, deeply divided, calyx (10-) 15-20 mm long, thin-textured, a cm with 2.5-4 lobes narrowly prominent midrib; petals triangular-acuminate, on a claw with and reddish pubescent (adaxially base), long, yellow (rarely staminal column 10-14 mm long, densely pubes otherwise glabrous; abaxially), cent, tapering distally, the filaments 5-6 mm long; styles 13-15, slender, sometimes Fruits subglobose; mericarps 13-15, dorsally with a broad lacerate wing pubescent. seeds 3.2 mm long, pubescent. Chromosome bearing stipitate hairs, 3-5-seeded; number: 2n = 30. occurs in central Veracruz and eastern Oaxaca Bakeridesia ferruginea (Fig. 20) inmatorral and secon below 1000 m, often in stream beds and canyons, commonly dary vegetation, Specimens

occasionally

Examined.

Veracruz:

in oak woodland.

Bourgeau

It flowers

2120; Dorantes

& Ch?zaro

from January 1909; Fryxell

to April. & Bates

937;

Galeotti 4103;Heller 46;Matuda S-13, 1415; Pedraza 259; Purpus 2233, 4316, 7541; Smith 6209; Sousa 4563; Trejo26;

Ventura

15020,15197,15720,15735,16729.-Oaxaca:

Sousa

1286.

10: 480. 1973. Abutil?n (Standley) Bates, Gentes Herb. Bot. Publ. Ser. 8: 24. 1930 ? Columbian Field Mus., gaumeri Standley, Type: Mexico. Gaumer 24072 (holotype: F! as photo F-56140!; Yucat?n,

89. Bakeridesia

gaumeri

isotypes:A! BM! F! GH! MO! NY! US!). the hairs Shrubs up to 6 m tall, the young growth densely stellate-pubescent, or less ferrugineous. Leaf blades mostly 7-11 cm long, ovate (progressively mi smaller and narrower upward), somewhat discolorous, cordate, acuminate, half of the and above the beneath; length petioles usually nutely stellate-pubescent

more

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1988

MALVACEAE OF MEXICO

111

(-3) cm long, solitary in the leaf axils; pedicels 0.5-1.5 ca. mm the lobes half-divided, calyx pubescent; long, stellate-pubescent, cm on a 1.5-2 with midrib; long, yellow, pubescent petals prominent apiculate, mm 4-6 column the 2 of otherwise staminal filaments claw, long, margin glabrous; 4 mm long; styles 8-10, slender, glabrous. Fruits 10-12 mm in diameter, urceolate 8-10, 6.5-9 mm long, dorsally with a wing that is (fide Bates, 1973); mericarps or a to Chromosome reduced rudimentary ridge; seeds ca. 3 mm long, pubescent. blades

or less. Flowers 7-10

unknown.

number

names: yaxholch? (maya), sachol, yaaxmisib. occurs in the Yucatan Bakeridesia gaumeri peninsula (Fig. 20) but is also from a single station (Cort?s: 1 km N of Villa Nueva, known disjunctly in Honduras at low elevations and apparently flowers Molina 6792). It is found in dry matorral more or less throughout the year. Vernacular

Specimens 1812,

23480,

Examined. 24046,

Arias

Yucat?n:

24072;

Lundell

7407,

& Vara

42, 56; Darwin 189; Bequaert 1229; Gaumer 497, Roo: Cabrera 35.?Quintana 194; Moreno

7500; Whigham

249; T?llez 1311. 10: 452. 1973.?Type: Mexico. Herb. gloriosa Bates, Gentes on to road 20 N of Tuxtla Guti?rrez El km 16 Feb Sumidero, Chiapas: 9044 (holotype: F! as photo F-56215!; F! as BH, 1965, Breedlove isotypes:

90. Bakeridesia

photo F-56216!MICH!). Shrubs

or small

stellate-pubescent. cordate, acuminate,

trees up to 6 m tall, the young growth densely ferrugineous 7-14 cm long, broadly to narrowly ovate, blades mostly

Leaf

above and minutely slightly discolorous, stellate-pubescent ca. of half the blades. Flowers the beneath; petioles usually length solitary in the cm leaf axils; pedicels up to 9 cm long, stellate-pubescent; 2-3 calyx long, thick or more a keel-like with half the lobes textured, ferrugineous-pubescent, divided, cm some corolla the 3.5-6 rotate, midrib; petals long, broadly obovate, yellow, on margins of claw; staminal times with an orange or reddish base, pubescent column 11-20 mm long, basally pubescent; Fruits 16-28 styles 16-18, pubescent. mm in diameter, oblate; mericarps 16-18, dorsally with a broad (3-4 mm) lacerate wing bearing stipitate hairs, ca. 6-seeded (?); seeds 3.5 mm long, pubescent. Chro mosome

number

unknown.

Bakeridesia gloriosa and Veracruz Oaxaca, to April. December Specimens MacDougall

Examined. 598.S?Chiapas:

occurs (Fig.

in deciduous and evergreen forests in Chiapas, m at to elevations 1400 and flowers from up 20)

Veracruz: Breedlove

Nevling 9044;