The genus Fleischmannia in Argentina, Bolivia, Brazil and Paraguay (Eupatorieae, Asteraceae)

Abstract Species of the genus Fleischmannia from Argentina, Bolivia, Brazil and Paraguay are reviewed, and keys are provided that cover the species in each country. New taxa described are Fleischmannia hatschbachii, Fleischmannia matogrosensis, Fleischmannia microstemon var. paniculata from Brazil, Fleischmannia hassleri from Paraguay and Fleischmannia neei and Fleischmannia steinbachii from Bolivia, and one new combination for a Fleischmannia prasiifolia variety is provided. The additions bring the total known species of the genus to 102.


Introduction
Fleischmannia is a genus of annual to perennial, often scrambling, herbs in the tribe Eupatorieae concentrated in the Central America and the northern and central Andes. One species occurs in eastern North America and a few occur in eastern Brazil. Prior to the present study the genus contained 97 known species. Recent eff orts to clarify the species of have concentrated on Mesoamerica (Robinson 2015), and northern and western South America (Robinson 2001(Robinson , 2008. In these areas, the delimitation of species has proven more complicated than initially expected. Meanwhile, the study of the genus from farther south in South America has been subjected to the lowest form of taxonomy, a process that I refer to as identifi cation work. Some individual specimens have been put aside over the years and are now subjected to new studies in more detail. Th e species have again proven to be more complicated than expected. Even countries with few species such as Brazil and Paraguay have presented some diffi culties, and Bolivia has many names in use without clear distinctions and many more recent collections that do not match any of the described species.

Materials and methods
Specimens used in the study were those accumulated over the last 150 years in the U.S. National Herbarium. Studies were restricted to use of the dissecting and compound microscopes.

Results
Th e species of Fleischmannia in southern and eastern South America are reviewed here by country starting with Argentina, Paraguay and then Brazil, based on material accumulated over the years at the U.S. National Herbarium (US). Treated last is Bolivia with its 12 species. In the process, new species are described and keys are provided.

Argentina
Only three species are credited to the country, none of them endemic. Th e three can be distinguished by the following key:

Paraguay
Two species have been collected in Paraguay. One previously only reported from Argentina and the other undescribed. Th e specimens of both species have previously been mostly mistakenly identifi ed as Fleischmannia prasiifolia (Griseb.) R.M. King & H. Rob., a species now known not to be from Paraguay.
Th e species has previously been reported only from Entre Rios in Argentina. Th e Paraguay localities are farther up river.
Specimens have usually been determined in the past as Fleischmannia prasiifolia with which it shares many characteristics such as ascending branches, short peduncles in the infl orescence, purple color of the corollas and achenes with somewhat paler ribs. Th e new species diff ers by the smaller more densely pubescent leaves and smaller clusters of heads, and it occurs far to the east of the distribution of F. prasiifolia and at generally lower elevations. Th e presence of minute glandular dots on the abaxial leaf surfaces would also seem to be a distinction, but minute stipitate glands occur in Fleischmannia prasiifolia var. glandulifera as noted in the Bolivian treatment.  Description. Erect herb 0.6-0.7 m tall, stems brownish, terete, striated when dry, hispid with small hairs. Leaves opposite, petioles 0.5-1.0 cm long, puberulous; blades ovate, 2.5-5.0 cm long, 1.0-1.8 cm wide, widest at basal 1/4or 1/3, base obtusely to acutely cuneate, margins bluntly subserrate with 4-6 teeth on each side, apex acute, or slightly blunt at tip, adaxial surface sparsely pilosulous; abaxial surface with prominent pale primary and secondary scabrid veins, areolae glabrous, without evident glandular dots, triplinervate from base with strongly ascending secondary veins. Infl orescence a lax panicle with mostly opposite branches spreading at mostly ca. 45° angles; mostly glabrous; main axis with foliiform bracts 0.6-2.0 cm long and 0.3-1.1 cm wide; branches fi liform, peduncles 6-15 mm long, glabrous. Heads ca. 5 mm high, 3 mm wide; Involucral bracts ca. 25 in ca. 4 series, narrowly ovate to narrowly oblong, 1.0-3.5 mm long, 0.5-0.8 mm wide, with broadly scarious obtuse tips; fl orets ca. 17 in a head; corollas white, ca. 2.3 mm long, basal tube ca. 0.5 mm long, throat ca. 1.5 mm long, lobes ca. 0.5 mm long, with short monoseriate hairs on outer surface; anther thecae ca. 0.8 mm long; apical appendage ca. 0.3 mm long; style branches not broader at tips. Achenes submature, ca. 1.7 mm long. brownish, ribs slightly paler, setuliferous on ribs; pappus whitish, ca. 2 mm long, of ca. 25 slender scabridulous non-contiguous bristles.
Only the type collection is known. Fleischmannia hatschbachii is one of two new species from Mato Grosso described here with slender glabrous or nearly glabrous branches of the infl orescence. Th is species and F. matogrossensis, nevertheless, diff er too much in their leaves to be the same species. A third species with glabrous branches of the infl orescence is found farther west in Bolivia, F. neei. Th at diff ers by its less slender infl orescence branches, lavender corollas and its more glabrous leaf surfaces.  Figure 3).
With the habit of the typical variety, but diff ering by the rather abrupt short acuminatie tips on leaf blades, stipitate glands on the peduncles and branches of the infl orescence, and by the not or scarcely paler ribs on the submature achenes.
Paratypes Th e variety is evidently restricted to areas near the coast of northestern Brazil. Th e locality in Bahia and the shape of the leaf blades as seen in the photograph of the G-DC specimen make it almost certain the Schrader nom. illeg. represents this same variety. While there is no nomenclatural requirement for perpetuating the Schrader name it has been decided to do so. Specimens are usually described as vines. Stems have branches usually spreading at ca. 90° angles. Th e leaves have mostly 9 to 14 blunt or sharp teeth on each margin, and vary from ovate with acute apices to narrowly ovate or lanceolate with narrowly acuminate apices. Infl orescence with elongate panicle with widely spreading branches, heads closely clustered at ends of branches; peduncles 2-6 mm long, puberulous. Heads ca. 7 mm high, 4-5 mm wide; involucral bracts ca. 22 in ca. 4 series; all bracts with acute to acuminate tips, 2-6 mm long, outer bracts narrowly ovate, inner bracts narrowly oblong. Corollas are rather consistently described as white, but sometimes lavender, ca. 3 mm long, basal tube ca. 0.5 mm long, throat ca. 2 mm long, lobes ca. 0.4 mm long, glabrous outside; anther thecae ca. 0.9 mm long, apical appendage ca. 0.25 mm long; style branches broadened distally. Achenes ca. 2 mm long, not paler on ribs, weakly setuliferous on ribs; pappus whitish, of ca. 35 bristles ca. 3 mm long, slightly broadened and contiguous at base, often detaching in groups.
Specimens seen.  Th is is the most common species of Fleischmannia in eastern Brazil. It is distinct from other Brazilian species by the more densely corymbiform infl orescence and broader style branches.

Bolivia
Th e understanding of Fleischmannia in Bolivia has been particularly bad, with most of the specimens misidentifi ed. Th e key to the Bolivian species of Eupatorium by B.L. Robinson (1920), treated the species intermixed with other species that were included in Eupatorium at the time. It relied on characters such as number of fl owers in the heads which are useful in other elements then known in Eupatorium but usually not in Fleischmannia. It was also based on extremely inadequate collecting, especially in southern Bolivia. Th e Robinson (1920) treatment did not include the Argentinian species, Eupatorium prasiifolium Griseb. and E. schickendantzii Hieron., which are now known to occur in southernmost Bolivia. Among material previously misidentifi ed by the present author are specimens here described as two new species, and some other individual collections have been seen that suggest still other undescribed species of the genus occur in Bolivia. Certainly, more collections of members of the genus in Bolivia is necessary.
Characteristics that seem more reliable at the species level in Fleischmannia are the habit of the plants, the angle of branching, the pubescence, density of the infl orescence, width of the style branch tips, the basic color of the corollas and the width of the bases of the pappus bristles.
Twelve species are known from the country including two that also occur in Argentina and the widely distributed F. microstemon. One species that has been credited to Bolivia, F. marginata (Poepp.) R.M. King & H. Rob. is actually found only the Dept. Junin in central Peru. Th e species cited in most previous studies such as Robinson (1920)  Specimens seen that bear the name, have thin pale-green leaves with slightly paler abaxial surfaces and white main veins. Th e petioles in all four specimens are comparatively short, 1/5 the length of the blade or less. Th ey are unbranched with fl at-topped corymbiform infl orescences with strongly ascending infl orescence branches, and have at least the outer involucral bracts lanceolate. Th ese specimens do not show the squarrose-spreading tips of the outer involucral bracts cited by Robinson (1920), but this characteristic is probably only a feature of an individual specimen. Such recurved tips have been seen in occasional specimens of other species as a result of something done during preparation.
Four diff erent specimens seem to share these general characteristics. Th e one that seems to fi t the concept best is from Bolivia: Cochabamba, 2-3 ft. Cardenas 494 (US ex hb. Gray as Eupatorium prasiifolium. Th e latter three specimens have most involucral bracts with more obtuse and more extensive reddish puberulence on stems, branches of the infl orescence and involucral bracts. All four specimens seem to be snatchings from tops of plants, and the total habit and any vegetative branching is unknown. Th e ascending branches, viscid glandular pubescence and lavender colored corollas should distinguish the species. No specimens have been seen matching this description. Th e strongly ascending branches indicate relationship to F. prasiifolia, and some specimens of F. prasiifolia placed in this study in the var. glandulifera also have glands. However, the specimens determined below as F. prasiifolia var. glandulifera have very minute glands, much denser clusters of heads in the infl orescence, have petioles of the leaves much shorter and tips of the leaves narrowly acute. Further collecting in southern Bolivia should discover new material of this species, and a neotype can be established.

Note. See description in Brazilian treatment.
Specimens with typical obtuse to short-acute leaf shapes have been seen from higher elevations in La Paz. Specimens with more acuminate leaf tips has been seen from elevations of 500 m or less in Cochabamba and Santa Cruz.
Description. Erect to scrambling perennial herbs to 1.2 m tall, stems terete, scarcely striate when dry, weakly puberulous to glabrate or glabrous; branches spreading at ca. 45 to nearly 90° angles; leaves opposite; petioles slender, 0.3-1.7 cm long, fi nely puberulous; leaf blades ovate, 2.3-6.5 cm long, 1-3 cm wide, base broadly rounded, scarcely acuminate at petiole, margins of lower leaves with teeth often sharply acute, apex short to narrowly acute, adaxial surface mostly glabrous, slightly shiny, abaxial surface slightly paler, puberulous on larger veins, mostly glabrous between; triplinervate from base of blade, larger veins whitish. Infl orescence a lax pyramidal panicle with mostly elongate, opposite, spreading branches, bearing small clusters of heads at tips; below and in infl orescence with small foliiform bracts on main axis; axis and branches of infl orescence glabrous or nearly so; peduncles 3-7 mm long, glabrous. Heads narrowly to broadly campanulate, 6 mm high, ca. 4 mm wide; involucral bracts ca. 25 in ca. 5 series, gradate,1-4 mm long, 0.3-1.0 mm wide, with narrowly scarious margins and apex, most basal bracts narrowly ovate, inner bracts oblong with obtuse apices, outer surfaces glabrous; fl orets ca. 17 in a head, corollas pale violet or pink, 2.0-2.3 mm long, basal tube ca. 0.4 mm long, throat 1.5-1.7 mm long, lobes ca. 0.3 mm long with few uniseriate hairs outside; anther thecae ca. 0.5 mm long, apical appendage ca. 0.15 mm long; style branches not broader distally. Achenes ca. 1.8 mm long, ribs sparsely setiferous above scabrid below, not or scarcely paler; pappus white, bristles ca., Fleischmannia neei is most distinct among Bolivian members of the genus by the glabrous branches of the infl orescence. Th e Buchtien collection was annotated by Rusby as Eupatorium polopolense B.L. Robinson, probably because of the locality. Many of the other specimens were subsequently also given that name. Typical Fleischmannia polopolensis diff ers obviously in its large denser corymbiform infl orescence and its densely puberulous peduncles. Relationship of the new species might seem much closer to the newly described F. steinbachii, with which it cooccurs in Santa Cruz, but the latter again has densely puberulous peduncles, more pubescent leaf surfaces, and lacks larger acute teeth on the margins of the lower leaves. An additional distinction of the new species seems to be the shorter corollas and much shorter anther thecae.
Description. Reclining herbs to 1 m tall, stems green to pale brown, terete, striated when dry, hirsute or hirsutulous. Leaves opposite; petioles 0.7-1.5 cm long densely puberulous; blades ovate, 2-4 cm long, 1.5-2.0 cm wide, base obtuse, margins serrulate with 5-9 blunt teeth, apex narrowly acute, adaxial surface pilosulous, abaxial surface essentially concolorous, pilosulous on and between veins, triplinervate from base, with main veins prominent, usually whitish. Infl orescence, a lax panicle with many pairs of elongate mostly opposite widely divaricating branches spreading at 60-90° angles, distally with cymiform branching, each bearing 2-8 heads, main axis and main branches with foliiform bracts 1.0-2.5 cm long, 0.3-1.2 cm wide; branches and peduncles densely puberulous, peduncles 4-20 mm long. Heads ca. 6 mm high, 4 mm wide; involucre of ca. 25 strongly gradate bracts 1.0-4.5 mm long, mostly ca. 1 mm wide, few at base acute at tips, most bracts broadly rounded at tip, with narrowly scarcious margins, glabrous and with prominent pair of longitudinal veins outside. Florets ca. 17 in a head; corollas pale bluish to reddish tinged, ca. 3 mm long, basal tube ca. 0.5 mm long, throat ca. 2 mm long, lobes ca. 0.5 mm long, with few slender styliform cells on outer surface; anther thecae ca. 1 mm long; style branches not broader distally. Achenes ca. Th is new species would key in Robinson (1920) to what he called Eupatorium tamboense Hieron., and that is is where specimens may have been placed in the past. However, Fleischmannia tambonsis is described with reddish hairs and stipitate glands on the branches of the infl orescence. Some of the present paratypes have been identifi ed in the past as F. polopoloensis and F. schicktendanzii. Of these the former diff ers by the denser more corymbiform infl orescence with white corollas. Th e latter diff ers by the more alternate distal branching of the mature plants, the more slender peduncles, the fewer and non-contiguous bristles of the pappus, and the lack of hairs or styliform papillae on the outer surfaces of the corolla lobes.
Th e styliform papillae on the outer surfaces of the corolla lobes have been seen in all three specimens placed in the species. Th e position of these papillae is one often occupied in other species by a few broader multicellular monoseriate hairs. Such styliform papillae have not been noticed in any other species in the genus, and this may prove a valuable distinguishing characteristic. No specimens have been seen that match the description. Th e specimen that matches best is one cited by Robinson (1920) "Bolivia: La Paz, South Yungas, Sirupaya near Yanacachi, alt. 2100 m, Buchtien 191 (300) (NY, US)" distributed as Eupatorium stipuliferum. Th is specimen cited by Robinson has glands on the peduncles but no reddish hairs, and it is from La Paz, not Tarija. Th e other specimen cited by Robinson (1920) "Bolivia: Cochabamba, Río Juntas, alt. 900 m, April 1892, Kuntze s.n. (NY, US)." fi rst annotated as Eupatorium marginatum, has densely puberulous but eglanduliferous peduncles without reddish hairs. Th is latter specimen remains unplaced at this time.
Th e species described after the Robinson 1920 treatment, F. yungasensis (B.L. Rob.) R.M. King & H. Rob. seems closest to F. tamboensis with its reddish hairs and stipitate glands, but it diff ers by its lower stems and leaves also being densely covered with stipitate glands and by its white corollas. Description. Scandent or subscandent perennial herbs, with all but most distal branches spreading at 90° angles, nearly straight or curving toward apex of main stem; stems hirtellous with reddish hairs; internodes 10-15 cm long; internodes of branchlets 2-3 cm long. Leaves opposite, leaves of main branches with petioles 10-22 mm long, densely hirtellous; leaf blades deltoid, 3.5-4.5 cm long, 1.5-2.5 cm wide, widest in basal fi fth, truncate with only slight acumination at petiole; ca. 8 or 10 teeth on each margin, apex narrowly acuminate, adaxial surface hispidulous with hairs reddish near margin and pale farther from margins, abaxially spreading hirtellous on main veins, between veins sparsely pilosulous, triplinervate with strongly ascending lateral veins from basal acumination; branchlet leaves with petioles 3-5 mm long, blades 1.5-2.0 cm long, 1.0-1.7cm wide, 5-7 teeth on each margin, apex weakly acuminate to acute, pubescence as in leaves. Infl orescence with small clusters of heads terminal on elongate main stems and branches, branches of infl orescence and peduncles densely hirtellous with reddish hairs and intermixed stipitate glands; peduncles 3-7 mm long; heads ca. 6 mm high, 3-4 mm wide; involucral bracts ca. 15, in ca. 4 series, 1.5-5.0 mm long, 1.0-1.2 mm wide, basal bracts ovate and acute, more herbaceous and densely pubescent, middle and inner bracts oblong, greenish to stramineous, with mostly 2 strong longitudinal veins, margins and tips scarious, apices obtuse. Florets ca. 15-17 in a head; corollas white. ca. 3.5 mm long, basal tube ca. 0.5 mm long, throat ca. 2.5 mm long, lobes ca. 0.5 mm long, with few hairs outside, anther thecae ca. 0.8 mm long, apical appendages ca. 0.25 mm long; style branches broadened distally to 0.3 mm wide. Achenes ca. 1.7 mm long, ribs scarcely pale, sparsely setuliferous; pappus ca. 3 mm long, of ca. 35 slender bristles not broader at base, not or scarcely contiguous. Specimens seen. Bolivia: La Paz: Sud Yungas, Yanacachi, camino haxia la Chojlla a la derecho primer desvic, al borde del camino, hierba apoyandose -1.5 cm, fl ores blancas, fortelezas, irrigation canals and mysterious foundations cover this ridge which is 2 km NE of the mouth of the Río Aguilani at Lakachaka, and 2 km S of the junction of the Río Mikhailpurhua and Río Aduada. 11 km N of Choquetanga. Area of ancient ruins which has been overgrazed, burned and generally raped into semi-barren grassland with some small shrubs. Collected bordering woodlands and mattorales. 15°39'S 67°20'W, vine-like, with opposite leaves, infl orescence without rays, white, leaves rather dry and curled when collected. Shrubby woodland, 19 Nov.1991 in Peru. Fleischmannia tamboensis may be closely related, with its indument of red and glandular hairs, but that is from southernmost Bolivia in Tarija, and was described by Hieronymus (1897), one of its collectors, as suff ruticose to 4 m tall, and sparsely branched, with membranaceous leaves with upper surfaces "scabriusculis" and abaxial surfaces scabrid on the nerves, and with peduncles ca. 6 mm long. Th is latter habit agrees with the type photograph that has been seen.