Revision of the Genus Anoda (Malvaceae)

A revision of the genus Anoda is presented that recognizes 23 species, of which ten are described as new. The genus includes annual herbs, subshrubs, and occasionally shrubs and occurs from the southern United States to Bolivia, Argentina, and Chile. Most of this distributional range, however, is accounted for by a single species, A. cristata, which also occurs as an adventive in a few other parts of the world. The genus is principally Mexican, all of the species occurring within that country. Anoda is closely allied to the genus Periptera, also a Mexican genus, and the two genera share features of fruit structure. The two genera together seem to occupy a relatively isolated position within the family. Anoda is subdivided into six sections, and keys to the sections and to the species are presented.


INTRODUCTION
The genus Anoda is principally Mexican in its distribution and its diversity, but it may be found from the southern United States to Bolivia, Argentina, and Chile.All of the species occur in Mexico, and only six of the 23 species occur outside of Mexico.Three species are common weeds, and one of these, A. cristata (L.) Schlechtendal, accounts for the entire breadth of distribution of the genus from the United States to Argentina and sometimes to other parts of the world (e.g., Mitchell 1982).
The genus has been studied taxonomically by Schlechtendal (1837), Gray (1887), Baker (1892), Garcke (1896), andHochreutiner (1916), and the genus has been treated in various regional floras such as Kearney and Peebles (1942), Shreve and Wiggins (1964), Correll and Johnston (1970), and Wiggins (1980).The most recent comprehensive study is that of Hochreutiner (1916), whose study, however, was based on only a limited number of specimens (42 cited specimens, distributed among 14 species and 5 varieties).Numerous subsequent collections have revealed additional species or have clarified our understanding of species already known.Therefore, it is considered appropriate to present a revision of Anoda at this time.Ten species are described as new in the present work, and 23 species are recognized.A cytological study by Bates (1987) complements the present study.TAXONOMY ANODA Cavanilles, Diss.1:38. 1785.
Lectotype: Anoda hastata Cavanilles.Gray (1887) reduced the three species of Cavanilles to a single species, for which he chose to use the name A. hastata, giving the other names as synonyms.This choice constitutes a lectotypification of the genus.
Gray.-Note:The assertion by Hochreutiner (1916, p. 39) that sect.Sidanoda is monotypic and based on Anoda abuti/oides A. Gray is incorrect.When Wooton and Standley elevated sect.Sidanoda to generic rank, they narrowed Gray's concept of the taxon to the single species, Anoda pentaschista, an action that serves as the choice of a lectotype.
One of the characteristic features of Anoda is that leaf form is highly variable within individual plants (e.g., Fig. 6, 7).Basal leaves, mid-stem leaves, and leaves immediately below and in the inflorescence may be very different from one another in size and form on an individual plant.It is therefore appropriate to speak of a "leaf spectrum" in relation to these plants and to keep this fact in mind when interpreting the descriptions or identifying an unknown plant.Generally, the leaves become smaller and narrower upward.This phenomenon, of course, occurs in many other genera, but its expression is relatively marked in Anoda.
Another characteristic feature of A noda concerns its distribution.Certain species are found in the field in relatively large populations (e.g. , A. palmata Fryx., or the weedy species such as A. cristata and A. pentaschista).Other species, on the other hand, occur as widely scattered individuals, and collectors may see only one or two individuals at a given site.The collector is thus faced with the choice of making a unicate collection or no collection at all and often opts for the latter, unless he or she has a particular interest in the genus.Thus, some of the species of Anoda are probably under-represented in herbaria as a result of this characteristic of scattered distribution.
The name Anoda is derived from the vernacular Ceylonese name "anoda" given to a species of Abutilon, a name that was noted by Burman (Thesaurus Zeylanica) and adopted by Cavanilles.Schlechtendal (1837) divided the species of Anoda into two sections, but he did not give the sections names.Various subsequent authors (Gray 1887;Baker 1892;Hochreutiner 1916) have also divided Anoda into sections.More recently, Kearney (1951) noted the practical difficulties of applying the distinctions among these  Anoda. -A. A. polygyna (Breedlove 19033).-B. A. leonensis (Fryxell et al. 1697).-C. A. paniculata (Koch and Fryxell82 247).-D. A. pentaschista (Waterfall 6210).-E. A. pedunculosa (Hernandez et al. 734).-F. A. reticulata (White 3618).-G. A. cristata (Gentry 1829).-H. A. zuccagnii (Hinton 11632).-1. A. lanceolata (Pringle 5454).-J. A. guatemalensis (Nelson 3169).-K. A. pubescens (Pringle 6969).-L. A. thurberi (Pringle 6536).-M. A. hintoniorum (Hinton 7229).-N .A. pristina (Breedlove 7546).sections, at least using the characters given as diagnostic by these authors.As is often true, the matter of such taxonomic subdivision is better approached by grouping species with more or less clear overall affinities and subsequently seeking characters that support the grouping.This approach results in a less artificial imposition of characters on taxa and more natural sections, in spite of the essentially subjective nature of the process.Such an approach leads me to the following subdivision of the genus into six sections.
Distinctive characters: fruits in the form of disks with radiating spines; hispid fruits; endocarp sometimes poorly developed or absent; plants often annual (?).
The species that constitute the sections Pseudanoda and Sidanoda are sufficiently distinctive from the rest of the genus that they are clearly set apart and have no close allies.The inclusion of A. succulenta in section Pseudanoda is tentative, pending the acquisition of fruiting specimens.
Section Liberanoda has two clear subgroups: the one including A. pubescens, A. henricksonii, and A. speciosa; the other A. hintoniorum and A. guatemalensis.The affiliations of the other two species, A. abutiloides and A. thurberi, are less clear.On the basis of the cytological evidence (Bates 1987), another disposition for A. thurberi may be necessary.
Sect.Cleistanoda, as here interpreted, seems to be a very natural group.On the other hand, sect.Clausanoda seems less coherent and may be a heterogeneous group.In any case, all of the sections here proposed are presented in the belief that they are a reasonable interpretation of the genus, with the hope that this interpretation will provide a frame of reference (and a set of hypotheses concerning relationships) for studies in other areas, such as cytology and crossing behavior.
The genus most closely affiliated with Anoda is the genus Periptera (Fryxell 1974;Bates 1987).The two genera form an isolated group within the family (Bates and Blanchard 1970), apparently without close allies.They share various similarities, especially concerning pollen characters (Sanchez 1982), fruit structure, and plant habit.In both cases the fruits are oblate disciform schizocarps, each mericarp often with a spur (or spine) on the dorsal angle and with the lateral walls evanescent at maturity.The seeds are solitary, and in some species (at least of Anoda) the seeds are completely enclosed in a persisting reticulate endocarp, a feature not known elsewhere in the Malvaceae.In those species with a persisting endocarp, the dispersal unit is the seed and enclosing endocarp; in those species lacking the endocarp, the seed alone is the dispersal unit.
Anoda and Periptera are distinguished principally on chromosome number (Bates 1987) and floral characters.In Anoda the corollas are purplish to pale lavender, white, or yellow, and the disposition of the petals is campanulate to rotate.In Periptera, on the other hand, the petals are usually some shade of red (or orange-red), and the petals are erect and often narrowly spatulate, forming a tubular corolla similar to that of Malvaviscus, though on a smaller scale.In addition, the genitalia are included within the corolla in Anoda, but are manifestly exserted in Periptera.The two genera apparently also differ in the form of the stigmas, which are abruptly capitate in Anoda (with the possible exception of A. thurberi) and more or less clavate in Periptera.Cytologically, Anoda is characterized by a base number ofx = 15 (with the exception of A. thurberi) and Periptera byx= 13.
Future studies may indicate the desirability of combining these two genera, but for the present they are maintained as distinct because of their different chromosome number, stigma structure, and floral aspect, the latter presumably related to pollinator adaptation.Even if Anoda and Periptera were to be combined, it would still be appropriate to distinguish them in subgeneric or sectional rank, so that the question becomes one of the level at which to distinguish them.There does not appear to be any advantage to recognizing the distinction between these two groups in infra-generic rank rather than generic rank, and so no such change is proposed here.
The following key to the species of Anoda is an artificial key that does not follow the sectional breakdown previously discussed but is presented to facilitate the identification of specimens.In the subsequent treatment of individual species, specimens are cited according to the conventional herbarium acronyms with the exception of the lower-case "pf," which refers to the author's herbarium.Erect subshrubs ca. 1 m tall, branching principally in the inflorescence, the stems with simple spreading hairs 0.5-1 mm long and with shorter more or less glandular hairs.Leaves long-petiolate, orbicular-ovate, cordate, dentate, caudateacuminate, to 12 cm long, gradually reduced and much narrower upward, minutely and softly tomentose, essentially concolorous.Flowers sometimes solitary in the leafaxils but usually in open terminal panicles; pedicels 1-5 cm long, without articulation, with pubescence like that of the stem; calyces 5-7 mm long, the lobes broadly triangular, more or less acuminate, shorter than the tube, with the midrib dark-pigmented; petals pale yellow (drying rose), ca. 1 cm long, prominently bearded on the claw; staminal columns ca.6 mm long, with recurved hairs, the filaments 3-4 mm long; styles 5, slender, slightly exceeding the androecium, subequal to the petals, pallid, the stigmas abruptly capitate, sometimes reddish.Fruits oblate, ca.6 mm in diam, minutely pubescent; mericarps 5, dorsally rounded; seeds solitary, 3 mm long, with minute appressed hairs, lacking an endocarp surrounding the seed.Chromosome number: 2n = 30 (Bates 1987).Distinctive characters of Anoda abutiloides include the uniformly 5-carpelled fruits and the leaves with caudate apices.The pollen grains of this species have been described by Sanchez (1982, as A. urophylla).Fryxell, sp. nov.Type: MEXICO.JALISCO: Jocotepec, cultivated; half-shrubby 2 m high; flowers white; said to be native, Jan 1978, Lape s. n. (holotype: MICH; isotype; pf).
Herbs or sub shrubs up to 2 m tall, the stems sparsely hirsute (the hairs simple, spreading, 1-2 mm long) to glabrate.Leaf blades broadly ovate below (3-7 cm long), smaller and narrower upward, sometimes hastately lobed, basally truncate or cuneate, crenate-dentate, acute, sparsely hirsute on the upper surface (the hairs principally simple, appressed, ca. 1 mm long), the pubescence on the lower surface similar but denser, especially on the nerves; petioles half as long as the blades or less, with pubescence like that of the stems; stipules subulate, 3-5 mm long, hirsute, persistent.Pedicels solitary in the leafaxils with pubescence like that of the stems, slender, 1.5-6.5 cm long (equaling or exceeding the subtending leaves), more or less aggregated apically, where the subtending leaves are greatly reduced; calyces 8-10 mm long, prominently hirsute especially on the nerves (the hairs 1-2 mm long), more than half-divided, the lobes ovate to narrowly lanceolate, acute or acuminate; petals 2-2.5 cm long, 1-1.5(-2) cm wide, white, prominently hirsute on the margins of the claw, otherwise glabrous; staminal columns ca. 4 mm long, pallid, glabrous basally, hirsute distally, with filaments arising apically; filaments 2-3 mm long, the anthers and pollen pallid; styles ca.15, slender, pallid, glabrous, the stigmas abruptly capitate, ultimately exceeding the androecium.Fruits oblate, disciform with radiating spines; mericarps ca.15, sparsely hispid, with a spine on the dorsal angle 1.5 mm long, the lateral walls evanescent, the seed not enclosed in a reticulate endocarp; seeds solitary, 2.6-2.8mm long, minutely warty, otherwise glabrous.
Erect herbs or subshrubs of unknown height, the stems green, the young stems densely yellowish hirsute becoming sparsely hirsute late, the hairs polymorphic, including simple slender hairs 2-4 mm long, rigid scabrid hairs ca.I mm long (on younger stems), with understory pubescence of short stellate hairs and a few glandular hairs.Leaf blades triangular or hastate, to 8 cm long, basally truncate, obscurely crenate, acute, slightly discolorous, with appressed simple hairs 0.5-1 mm long above, uniformly and minutely stellate-pubescent beneath, the pubescence denser and yellowish on the nerves; petioles almost as long as the blades, with pubescence like that of the stems; stipules filiform, 5-6 mm long, hirsute.Pedicels 2-3 cm long in flower, accrescent to 6-8 cm long in fruit, with pubescence like that of the stem, solitary in the leafaxils, more or less aggregated apically; calyces 5-7 mm long in flower, accrescent to 7-9 mm long in fruit, lO-ribbed, densely yellowish pubescent, approximately half-divided; petals 8-9 mm long, pale yellow when dry (orange according to collectors' notes), prominently hirsute on margins of claw, otherwise glabrous; staminal columns ca.4.5 mm long, pallid, densely hirsute, the anthers ca.20, the filaments 1-1.5 mm long; styles glabrous, the stigmas capitate.Fruits oblate, 9-11 mm in diam, densely pubescent; mericarps 13-16, with a dorsal spur ca. 1 mm long, the lateral walls evanescent, the reticulate endocarp poorly developed and not enclosing the seed; seeds glabrous, ca.2.3 mm long.The disjunction between the two Mexican specimens cited is notable and remains to be explained, but the two collections appear to represent the same species and are so dealt with here.Future collections may reveal a continuous distribution across northern Mexico.The provenience of the cultivated specimens of M .S. Bebb (1833Bebb ( -1895) )  Erect herbs, minutely and roughly pubescent.Leaf blades mostly 3-7(-12) cm long, variable in form, the lower leaves triangular, smaller and narrower upward, lanceolate, truncate, or cuneate basally, obscurely dentate or subentire, more or less discolorous, with simple appressed hairs (0.3-1 mm long) above, with short stellate hairs beneath; petioles with pubescence like that of stem, to 10.5 cm long on lower leaves but usually less than 1 cm long; stipules inconspicuous.Pedicels solitary in the leafaxils (up to 6.5 cm long, subequal to the subtending leaves) or more commonly aggregated apically into leafless racemes, articulated 5 mm below the flowers; calyces 6 mm long in flower, accrescent to 8 mm in fruit, minutely stellate-pubescent, approximately half-divided, the lobes with prominent midribs, triangular in flower, the margins somewhat recurved in fruit; corollas yellow (sometimes reddish at base), 8-16 mm long, the petals densely ciliate on the margins of the claws; staminal columns densely pubescent, the hairs stellate, more or less retrorse.Fruits (Fig. 11) oblate, 9-11 mm in diam, densley stellate-pubescent, the hairs 0.5-1 mm long; mericarps 10-12, each with a dorsal spur ca. 1 mm long, the lateral walls evanescent; seeds solitary, minutely verruculate, 3 mm long, the endocarp usually well developed with excrescences on the margin (Fig. 2).Anoda lanceolata has a superficial resemblance to A. cristata, but differs most notably in its yellow corolla and in the elaborately developed endocarp surrounding the seeds.The pollen grains of this species have been described by Hashmi (1970).10.Anoda leonensis Fryxell, sp.nov.Fig. 1B, 5 Type: MEXICO.Nuevo Leon: between Linares and Iturbide, near K36, by large bas-relief on cliffside, alt.4000 ft, 15 Oct. 1969, Fryxell1207 (holotype: MEXU; isotypes: BH, CTES, MICH, US, pt).
androecium, the stigmas abruptly capitate, glabrous, pallid or reddish (?).Fruits (Fig. IB) oblate, 8-9 mm in diam, densely hirsute (the hairs 0.3-0.7 mm long, the longest hairs at the apex); mericarps 12-16, dorsally rounded (crested but lacking spurs), the central rib or crest with dark pigmentation in the immature fruit, the lateral walls evanescent; seeds solitary, each with a persistent reticulate endocarp surrounding the seed 2.5 mm long.Chromosome number: 2n = 30 (Bates 1987).Anoda leonensis is named for its principal occurrence in the state of Nuevo Leon, where it occurs at elevations of 500-1500 m.Many of the above specimens were initially determined as A. paniculata or A. wrightii, but these species are amply distinct.
Paralypes examined.-MEXICO.GUERRERO: Mpio.de La Union, carretera Zihuatanejo-Cd.Altamirano, 84 km al N del entronque con carr.Zihuatanejo-Utzaro-Cardenas.Bosque sabanoide de encino con elementos tropicales; suelo franco.Alt. 1550m, 23 Oct. 1982, Koch and Fryxell 82161  (CHAPA, ENCB, MEXU, US, pf) Anoda maculata, as the specific epithet indicates, is distinctive for the dark center of the flower.It is similar to A. crenatiflora in many particulars, but the two species may be distinguished by pubescence (especially that of the calyces and pedicels), perhaps by leaf shape and anther number, and by other characters.
Erect annual herbs 1-1.5 m tall, freely branched, the stems with stellate hairs, • 0.2-0.5 mm long and a few very short glandular hairs and sometimes also with long simple hairs 2-3 mm long.Leaf blades mostly 3-8(-10) cm long, the young leaves ovate or slightly lobed, truncate or cordate, coarsely dentate, upwards deeply palmately 5-10bed and ultimately 3-10bed, the lobes becoming narrowly linear-Ian ceo late, obscurely dentate to entire, acute, essentially concolorous, with stellate hairs 0.5-2 mm long and sometimes also appressed simple hairs above, with 6-8-rayed stellate hairs ca.0.5 mm in diam uniformly distributed beneath, rarely with irregular purplish blotch along midvein; petioles to 8 cm long (half the length of the blade to equaling the blade), with pubescence like that of the stems; stipules filiform, 3-5 mm long, caducous.Flowers solitary in the leafaxils, forming open terminal racemes more or less exceeding the leaves; pedicels 2-5 cm long, slender, minutely stellate-pubescent, articulated 5-8 mm below the flowers; calyces 5-8 mm long (not accrescent), basally rounded and yellowish or brownish, densely covered with stellate and glandular hairs, half or more divided, the lobes ovate-apiculate, I-ribbed, with lanate margins; petals yellow, 6-12 mm long, ciliate on the margins of the claws, otherwise glabrous, staminal columns 2-3 mm long, pallid, basally glabrous, prominently hirsute apically (the hairs 0.5 mm long), the filaments 1-2 mm long, the anthers and pollen yellow; styles slender, pallid, glabrous, slightly exceeding the androecium, the stigmas abruptly capitate, 0.3-0.5 mm in diam (at least 3 times the diameter of the style).Fruits oblate, 8-9 mm in diam, densely stellate-pubescent (the hairs 0.5-1 mm long); mericarps 8-13, each with a dorsal spur 0.5-1 mm long, the lateral walls disintegrating at Anoda palmata is characterized by notably lobed leaves, as the specific epithet indicates, and bright yellow flowers.It is a plant relatively common in Guerrero, Michoadm, lalisco, Nayarit, and Baja California, at times growing in large populations.It occurs from sea level to 1500 m elevation.
Robust erect herbs to 2 m tall, the stems little branched, minutely and uniformly tomentose.Leaf blades variable, the lower leaves more or less simple, angled or slightly lobed, mostly 6-8 cm long, as wide as long, basally cordate or truncate, coarsely serrate or undulate to entire, acute, scabridulous on the margins and the veins beneath, otherwise glabrous, essentially concolorous, sometimes with an irregular purplish blotch along the midrib, the upper leaves becoming more deeply palmately lobed (the sinuses rounded) and finally narrowly hastately lobed; petioles minutely pubescent like the stem (the hairs 0.1-0.2mm long), one-third as long as blade to equaling the blade.Inflorescences of open terminal racemes; pedicels 1-3.5(-6.5)cm long, with pubescence like that of stem, articulated 3-10 mm below the flower; calyces 6-9 mm long in flower, accrescent to 12 mm long in fruit, 10-ribbed, minutely and densely yellowish stellate-pubescent, ca.halfdivided; petals 10-19 mm long, prominently ciliate on the claws with hairs ca. 1 mm long, otherwise glabrous, white or rose drying to pale rose or purple with the claws remaining white; staminal columns whitish, 3-5 mm long, glabrous basally, prominently hirsute apically with hairs ca. 1 mm long, the filaments 3-4 mm long, the anthers and pollen yellowish; styles slender, pallid glabrous, the stigmas abruptly capitate.Fruits (Fig. 1 C) oblate, 8-9 mm in diam, stellate-tomentose; mericarps 11-15, with dorsal spurs 0.5-1 mm long, the lateral walls disintegrating; seeds solitary, each totally enclosed in a persistent black reticulate endocarp 2.5-3 mm long.Anodapaniculata is found in the Rio Balsas Depression from northern Guerrero to the southern part of the state of Mexico, at elevations of 300-1300 m.It is a robust plant with glabrous foliage and a variable corolla color-the petals may be white, lavender, or rose, generally drying to lavender or purple, with the claws remaining white.Erect annual herbs or subshrubs 0.5-1.5 m tall, the stems minutely and uniformly stellate-pubescent.Leafblades 3-7 cm long (often smaller), ovate (young leaves) to hastately 3-lobed, becoming narrower upwards, obscurely crenate, acute, minutely and sparsely stellate-pubescent, sometimes with an irregular purple blotch along the midrib.Flowers solitary in the leafaxils, long-pedunculate, often grouped apically in terminal racemes or panicles more or less exceeding the leaves; calyces 4-6 mm long, not accrescent, puberulent, half-divided; petals whitish or lavender, 6-15 mm long, prominently ciliate on the margins of the claws; androecium included, the columns pallid, with long transparent hairs apically; anthers and pollen yellow; styles glabrous, 12-13, sometimes reddish, the stigmas abruptly capitate.Fruits (Fig. 1 E) oblate, 8-9 mm in diameter, whitish-pubescent, the 12-13 mericarps each with a short dorsal spur, the lateral walls disintegrating at maturity; seeds solitary, each completely enclosed in a blackish reticulate endocarp (Fig. 2).Chromosome number: 2n = 30 (Bates and Blanchard 1970, as A. reticulata; Bates 1987).Anoda pentaschista occurs widely through many parts of Mexico and extends into the southern parts of Texas, New Mexico, and Arizona.It sometimes becomes weedy in agricultural fields, roadsides, and disturbed ground.The pollen grains of this species have been described by Hashmi (1970).
The distinctive characters of mericarp structure set A. pristina apart from all other members of the genus: mericarps that are apically dehiscent and that have a persisting reticulum in the lateral walls.In overall habit, the plant shows some resemblance to the common A. cristata but may prove to be allied to A. succulenta when both are better known.At present A. prist ina is known only from the type collection.
Perennial herbs 0.5-1 .5 m tall, the stems sparsely scabridulous, the hairs stellate, 0.2-0.4mm long.Leaf blades broadly ovate below to narrowly triangular or hastately 3-10bed above, deeply cordate, 3-8 cm long, serrate, acute or acuminate, minutely and sparsely stellate-pubescent above and beneath, essentially concolorous; petioles with pubescence like that of the stems, to 6 cm long but shorter upwards.Pedicels 1-5 cm long in flower, somewhat longer in fruit, articulated 2-4 mm below the flowers ; calyces 6-12 mm long, densely stellate-tomentose, deeply 5-10bed, the lobes lanceolate-oblong, I-nerved, more or less apiculate; petals 7-14 mm long, lavender (whitish on the claw), ciliate on the margins of the claws, otherwise glabrous; staminal columns whitish, ca.7 mm long, stellate-pubescent apically, the filaments ca. 2 mm long, the anthers lavender, subequal to the petals, the pollen yellow; styles slender, the stigmas capitate.Fruits (Fig. lK) oblate, 7-8 mm in diam, densely and minutely stellate-pubescent; mericarps 8-10, each with a dorsal spur up to 0.5 mm long, the lateral walls evanescent; seeds solitary, glabrous, 3 mm long, not enclosed in an endocarp.Chromosome number: 2n = 30 (Bates 1987).Anoda pubescens is clearly allied to A. henricksonii, from which it differs morphologically (see key) and geographically.Fresh flowers show a slightly geniculate staminal column, but this feature is poorly if at all preserved in a dried specimen, so it does not enter into the preceding description.The pollen grains of this species have been described by Sanchez (1982).Erect herbs ca. 1 m tall, the stems scabridulous (the hairs < 0.5 mm long, rigid, simple, bifurcate, or stellate) and with some glandular hairs.Leaves petiolate, ovate when young but later leaves deeply 3-10bed (rarely 5-10bed), the lobes narrowly linear, 1-5 mm wide, the central lobe mostly 3-6 cm long, the lateral lobes 2-4 cm long, reduced upwards to filiform leaves and ultimately in the inflorescences to stipuliform bracts; the larger leaves serrate to subentire, with an irregular purple blotch along the midrib, minutely stellate-pubescent above and beneath, more densely so beneath.Pedicels solitary in the leafaxils or forming terminal (essentially leafless) racemes, slender, up to 8.5 cm long, articulated 4-8 mm below the flowers, with pubescence like that of the stems, the hairs scabridulous and often somewhat antrorse; calyces ca. 5 mm long (not accrescent), densely stellate-pubescent, half-divided, the lobes with prominent midribs, apiculate; petals 5-8 mm long (slightly longer than the calyces), lavender or purple, ciliate on the claws; staminal columns ca. 2 mm long, pallid, glabrous or with a few hairs, the filaments purplish, the anthers yellowish; styles and stigmas clavate, yellowish, recurved.Fruits (Fig. 1 F) oblate, 6-7 mm in diam, 3-4 mm tall, stellatepubescent (the hairs ca.0.5 mm long); mericarps 10-11, the dorsum of each rounded (the spur totally absent), the lateral walls evanescent, the solitary seeds completely enclosed in a black reticulate endocarp 2.5 mm long.The distinctive features of A. reticulata are the total absence of spurs on the fruits and the distinctive trilobed leaves with linear lobes, at least on well-grown mid-stem leaves.It is sometimes confused with A. thurberi because ofthe similar small, bluish flowers, but these two species differ markedly in fruit characters (cf.Figs.IF and lL) and in leaf shape.The pollen grains of this species have been described by Sanchez (1982, as  Herbae vel suffrutices erecti, caulibus uniformiter stellato-scabridiusculis; foliis hastato-trilobis, profunde cordatis ad basem, minute stellato-pubescentibus; calycibus 12-15 mm longis; petalis lavandulis, 22-28 mm longis; columna staminali 20 mm longi, fere glabri; fructibus ignotis. Herbs or subshrubs of unknown height, the stems uniformly scabridulous (the hairs stellate), sometimes purplish on one side.Leaf blades hastately 3-lobed, deeply cordate at the base, crenate-dentate or subentire, to 10 cm long (smaller and narrower upwards), acute or acuminate, somewhat discolorous, minutely stellate-pubescent throughout; petioles to 6.5 cm long, 1f4-% as long as the blade, with pubescence like that of the stems; stipules 6-9 mm long, filiform.Pedicels solitary in the leafaxils, usually exceeding the subtending petioles, up to 5.5 cm long, more or less aggregated apically into racemiform inflorescences, with pubescence like that of the stems; calyces in flower 12-15 mm long, minutely stellatepubescent, 2h-divided, the lobes lanceolate-acuminate, I-nerved; petals 22-28 mm long, 15-18 mm wide apically, tapered to a narrow claw, ciliate on the margins of the claws; staminal columns ca.20 mm long, essentially glabrous but with a few minute hairs, pallid, bearing filaments apically; filaments 3-5 mm long, the anthers ca.40, lavender (?), the pollen yellow-orange; styles and stigmas not seen.Fruits unknown.
Erect succulent herbs, l.5 m tall, with thick hollow stems that are essentially glabrous.Leaf blades palmately 3-5-lobed, palmately 7-nerved, up to 1l.5 cm long, about as wide, basally cordate-hastate (the sinus narrowly acute), entire, acute, the central lobe broadly ovate, narrowed at the base making the sinuses characteristically rounded, glabrous above and beneath and on the margins; petioles subequal to the blades, with a narrow line of weak hairs on the adaxial side and a few spinescent hairs 0.5-1 mm long, otherwise glabrous, sometimes purplish on the adaxial side; stipules 5-7 mm long, ciliate, caducous.Pedice1s in the leaf axils, to 13 cm long (subequal to the subtending leaves at anthesis), with a narrow line of hairs and sometimes purplish on the adaxial side, otherwise glabrous; calyces 10-12 mm long, externally glabrous or with a few pungent hairs on the margins, internally wooly, half-divided or more, the lobes lanceo1ate, acute, I-nerved; petals yellowish, 15-16 mm long, densely bearded on the margins of the claws (hairs 1 mm long), otherwise glabrous; staminal columns pallid, ca. 3 mm long, with a few hairs apically; filaments 2-3 mm long, the anthers yellowish, numerous; styles slender, pallid, the stigmas capitate (of uncertain number).Fruits unknown.
As the specific epithet indicates, Anoda succulenta is distinctive (indeed apparently unique in the genus) for its hollow, succulent stems.It also has a leaf form (palmately lobed, with broad rounded sinuses) that is distinct within the genus.It may be allied to A. pristina.
Erect herbs ca. 1 m tall, the stems minutely pubescent with stellate and glandular hairs 0.2-0.4mm long.Leaf blades 5-8 cm long, heteromorphic (the lowermost leaves ovate-cordate, mid-stem leaves hastately 3-lobed, the uppermost narrowly hastate or triangular), sometimes with an irregular purple blotch along the midrib and sometimes also with purple margins, obscurely or manifestly serrate, moderately or sparsely pubescent above and beneath, the hairs minute and stellate or rarely simple on upper surface; petioles half length of blade to subequal to the blade.Rowers solitary in the leafaxils or more commonly forming terminal racemes or open panicles more or less exceeding the leaves, the pedicels slender, 1-3 cm long, articulated 1-4 mm below the flower, with pubescence like that of the stems; calyces 3.5-6 mm long in flower, accrescent to 6-8 mm long in fruit, stellate-pubescent, half-divided or more, the lobes with a pronounced midrib, acute or apiculate, somewhat revolute in fruit; corollas 4-7 mm long, bluish lavender, often with dark centers, ciliate on the claws, otherwise glabrous; staminal columns 1.5-2(-4) mm long, stellate-pubescent, pallid or lavender, the filaments < I mm long, the anthers few (l0-20), lavender, the pollen orangish; styles and stigmas barely exceeding the androecium, minutely clavate, yellowish.Fruits (Fig. 1 L) oblate, 6-8 mm in diam, minutely pubescent (the stellate and glandular hairs to 0.5 mm long); mericarps 6-8, with dorsal spurs very short (or up to 1 mm long), the lateral walls evanescent; seeds solitary, 2-2.5 mm long, glabrous, the endocarp incomplete and fragile or absent.Chromosome number: 2n = 26, 28 (Bates 1987).Anoda thurberi is a widespread species that is distinct for its small, blue-purple flowers, a characteristic it shares with A. reticulata, with which it is sometimes confused.These two species are clearly distinguished by leaf shape and fruit characters, however, and need not be confused.Bates (1987) has shown cytogenetic affinity of A. thurberi with Periptera punicea.Further study is needed to clarify the generic relations of Anoda and Periptera and the placement of A. thurberi.Annual or perennial herbs or subshrubs, erect or decumbent to prostrate, the stems prominently or obscurely hispid with the hairs spreading and retrorse, or almost glabrous.Leaves petiolate, palmately lobed below to narrowly hastate above, subentire, acute, sparsely pubescent to glabrate, sometimes with an irregular purple blotch along the midrib and sometimes also purple-margined.Flowers solitary in the leafaxils, with long peduncles; calcyes 6-11 mm long in flower, accrescent to 10-15 mm long in fruit, often more or less hispid; petals purple or lavender, 12-18 mm long; androecia included, the staminal columns apically hispid, basally glabrous.Fruits (Fig. I H) oblate, 8-11 mm in diam, without spines or with short spurs, densely hispid; mericarps 7-11, gibbous, indehiscent, the lateral walls evanescent; seeds solitary, glabrous, 2.5 mm long, lacking an endocarp surrounding each seed.Chromosome number: 2n = 30 (Bates 1987).two species are recognized as distinct in this study because there appear to be two distinct morphological modes (cf.distinctions given in key and Fig. IG and Fig. 1 H), although some overlap of certain character expressions exists, and a few individuals show intermediate expressions.For example, the number of carpels in A. zuccagnii varies between 7 and 11 with a modal value of 9; for A. cristata the range is 10 to 18 with a modal value of 13.Nevertheless, I recognize that genetic, cytological, and other studies may find that the recognition of two species cannot be supported.For the present, however, they appear distinct.
Distinctive characters: mericarps with apical dehiscence; lateral walls with persistent reticulum; endocarp not enclosing the seed; plants more or less glabrous.