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AND REVISION Nancy and Hensold^ Ana Maria THE REDEFINITION OF GiuliettP GENUS RONDONANTHUS HERZOG (ERIOCAULACEAE)^ Abstract The genus Rondonanthus Herzog now has until contained only the species R. roraimae and R. micropetalus, and has been defined by free petals in both sexes of flowers and the dioecious condition. here redefined according It is to a large suite of characters, especially the presence of long filamentous staminodes in the pistillate flowers, and by the adnation of the filament base to the corolla in the staminate flowers. Five additional species, including the monotypic genus Wurdackia, are transferred to Rondonanthus, and /f micropetalus removed to Paepalanthus. proposed is It is . Rondonanthus that is a highly primitive paepalanthoid genus, to be taken as a primitive outgroup of Syngonanlhus, The Eriocaulaceae are a pantropical monocot paepalanthoid genera. These are Rondonanthus family of about 13 genera and 1,200 Herzog Comanthera Lyman species (Krai, (1931), Smith B. Except 1989). in rare cases, the flowers are uni- (1937), Carptotepala Mold. and Wur- (1951), sexual and the plants monoecious, with the flowers dackia Mold. (1957). borne in scapose heads. Following the lead of Ruhland (1903), of all Ruhland (1903) divided the family into two sub- these genera were described with reference only to families, the Eriocauloideae, characterized by two a few floral characters. None contain more than whorls of stamens and a simple gynoecium, and two species, and none were discussed by their au- the Paepalanthoideae, with a single whorl of sta- thors as to their relationships with other genera or mens and "appendages" vascularized secretory in- implications for phylogeny. Recently, the avail- on serted the style alternate with the nonvascu- ability of abundant collections from the Venezuelan The larized functional style branches. Eriocauloideae tepuis and their study for Julian Steyermark's Flo- are truly pantropical and contain many aquatic a of the Venezuelan Guayana r {R^n^yoXA, in ^re^.) species, while the Paepalanthoideae are almost en- have acutely emphasized the need study and for New World some tirely limited to the (with African reevaluation of these endemic genera. & and more disjuncts), xerophytic. In another work Hensold, (Giulietti 1991), The we Paepalanthoideae, which contain 11 of the discuss the systematic position of Comanthera 13 genera, have two centers of diversity, one in and Carptotepala, concluding that they are syn- the Central Brazilian Plateau, especially the Cadeia onyms of Syngonanthus Ruhl. In work, we this do Espinha^o in Minas Gerais and Bahia, and one find the genus Rondonanthus be though to valid, Guayana in the Highland, especially Venezuela. in need of redefinition, and include within five it Botanical exploration of Guayana, in particular the additional endemic Guayanan species currently summits tepui of Venezuela, has only been under- placed in three other genera, including the mono- taken recently relative to that of the more acces- typic genus Wurdackia, mountains and Rondonanthus sible of central Brazil, has turned as here recognized a genus of is many up taxa of interest, including four endemic great interest from the standpoint of phylogeny in We VEN thank MO, NY, PORT, the curators of F, K, LL, and for the loan of specimens. The work was ' completed while the junior author was a visiting research scientist under the Missouri Botanical Garden Postgraduate Fellowship Program, supported by the Jessie Smith Noyes Foundation. Habit by Bruno Manara illustrations were provided by the Flora of Venezuelan Guayana Project. Department Museum 2 of Botany, Field of Natural History, Roosevelt Road at Lakeshore Drive, Chicago, Illinois 60605-2496, U.S.A. Departamento ^ de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, C.P. 1.461-05499 Sao Paulo 1 SP, Brazil. Ann, Missouri Bot. Card. 441-459. 78: 1991. 442 Annals of the Garden Missouri Botanical Eriocaulaceae. appears to combine characters rollas, with the corollas of the stamlnate flowers It of the Eriocauloideae with characters of two ''core" tubular and those of the pistillate flowers connate paepalanthoid genera, Paepalanthus and Syngo- at the middle and free at the base and the apex. mird nan thus. As (1985) concluded (limiting his Stiitzel known observations R. {Wurdackia) flabelllformis\ species of the family with bisexual flowers, to it Rondonan- appears represent a primitive outgroup to the All of these taxa are here treated in to large genus Syngonanthus. As a genus, also thus, though their diff'erences in corolla fusion it and caused shows unusual variability with respect to certain characteristics flower sexuality originally characters, such as fusion of perianth parts them to be assigned to four diff"erent genera, floral Rondo- now The and which have been only other species described in flower sexuality, until given a priori value in the definition of genera of nanthus since the type species, R. micropetalus Eriocaulaceae. Mold. (1951), reputedly with free petals in the synonymy removed staminate here flowers, to is Paepalanthus Excluded Ruhl. Species). (see in Taxonomic History The genus Rondonanthus was described by Generic Definition and Affinities Herzog (1931) on the basis of a single Luetzelburg collection of R. roraimne (Oliver) Herzog, based The single character that best distinguishes Ron- on Paepalanthus roraimae Oliver. donanthus from other genera of Eriocaulaceae all ma- Herzog found only staminate flowers in his the presence of long linear staminodes in the is known and defmed the genus by the free petals of flowers. Staminodes are in Pae- terial pistillate unknown the male corolla, which he believed to be palanthus, but they are small and scalelike, never Paepalanthus. Because he did not see the type linear. The only species of Rondonanthus in which in depended on material, he Oliver's illustrations for flabellifc description of the flowers, which he they are replaced by functional stamens which his pistillate described correctly as having the petals free and produce and release pollen in apparently normal Herzog incorrectly as having simple style branches. quantities. he was Rondonanthus by mixture also noted that the limited material studied further defined a is dioecious and suggested this as an additional char- of characters individually found in other genera, acter of the genus, also encountered rarely in the especially Syngonanthus and Paepalanthus, but family. not in combination with each other. Ruhland comprehensive treatment The Rondonanthus appear (1903), in his affinities of to lie of the family, had in fact erected Paepalanthus most closely with Syngonanthus, for a number of subg. Bostrychophyllum Ruhl. to accommodate reasons. In R. acopanensis, R. caulescens, and known R P. capillaceus Klotzsch ex Koern., the single Jlabelliformis, the petals of the pistillate flowers . species of Paepalanthus with free petals in the are fused in the middle but free at the base and staminate flowers. This species a vegetatively apex, which considered to be one of the defining is is specialized submerged aquatic also native to Ro- characters of .SjT/g^onan^/zu5. Elsewhere in the Er- raiina, and here treated as a species of Ron- iocaulaceae, this known only in the genus Phil- is is donanthus. (Ruhland had placed P. roraimae in odice C. Martius (2 spp.), which considered to is ''Species dubiae" since he had observed no material be derived from Syngonanthus, and in Mesan- of the species.) theniuni Koern., which with two whorls of stamens The same year that Herzog described Rondo- and an unmodified gynoecium quite distant from is nanthus, Gleason (1931) described Paepalanthus Rondonanthus. duidae from Cerro Duida in Arnazonas, noting the This character, however, not stable in the is close similarity to P. roraimae Oliver, and diff"er- genus. In R. roraimae and R. capillaceus, the entiating from that species only by minor char- petals of the pistillate flowers are always free, and it acters of gross morphology, and without mention in the variable species /?. tZui^ae, most populations of structure. have free petals, but a single population has been floral Rondonanthus were Later, additional species of found (at Aprada-tepui) with brief petal fusion. In described from the Chimanta Massif and placed in the genus Syngonanthus, the character also shows Syngonanthus acopanensis Mold., 1948, and a small degree of instabiHty. In the reduced, dim- {S, S. obtusifolius Mold., 1957), and in a new mono- erous species S. minutus (Mold.) Hensold (1991), typic genus W^urdackia (JF. jiabelliformis Mold., for example, the petals are free, and this situation 1957). All exhibit typical SyngonanthuS'\v\.e. co- is also occasionally found in sect. 77rjsa«oce/>Aa/t/5 Volume Number 78, 2 Hensold & 443 Giulietti 1991 Revision Rondonanthus of (Koern.) Ruhl., as In 5. ye/ima«ii(Gleason) Giulietti coloration of the involucral bracts, and frequently & Hensold (1991). Petal fusion probably repre- also the upper parts of the perianth; the of tufts sents a primitive state in Rondonanthus, as is also trichornes on the tips of the sepals and floral bracts; considered to be the case in Syngonanthiis, the clavate or subclavate shape of these perianth Fusion of the corollas of the staminate flowers trichornes, which sometimes {R. capillaceus) are is a fairly universal condition In the Paepalanthoi- densely ornamented on the internal wall; and, in deae, but character breaks down Ron- this also in three species, the ciliate margins of the staminate donanthus. Of the species with free petals in the flower corollas. pistillate flowers, R. ra/?t7/acea5 also has free petals All of these characteristics are common the in in the staminate flowers, R. roraimae has petals Eriocauloideae. The not known from any last is connate early in development and then separating, species of SyngonanthuSy but common some is in and R. duidae, except in very rare cases, has of the less specialized members of Pae/>a/a/z^/iu.s, The petals persistently connate. remaining three notably subg. Xeractis. Inner ornamentation wall which species, all exhibit fusion of corollas in the in floral trichomes has also not been described from have Syngonanthus pistillate flowers, similarly fused corollas of but is the rule in Paepalanthus. the staminate Bearded known flowers. sepal apices are Syngonanthus in Other characters which indicate an alliance with (especially Dimorphocaulon, hum- in sect. e.g., 5. Syngonanthus are the following: roots pale and boldtii (Kunih) Ruhl. diamantinensis S. Silveirsi), , aerenchymatous; hairs present on the floral axis at but not common, and involucral bract pigmenta- ovary base; base color of the perianth creamy tion, when occurs, usually more or less trans- it is white; pubescence of malpighian hairs on vegeta- lucent and reddish arenarius (Gardner) Ruhl., (5. tive parts. All of these characters except the last S. anomalus (Koern.) Ruhl.). An exception S. is may also be found in the genus Leiothrix Ruhl., niger Silveira, with long black linear-lanceolate but this genus diff'erentiated by specialized mor- bracts. is phology gynoecium, of the seed testa striations, Furthermore, in R. acopanensis, the outermost and basifixed anthers (Giulietti, 1984). bracts, as well as the inner bract apices, are ap- Stiitzel (1988) found similar root anatomical parently green and leaflike when young, which may structure In /?. roraimae^ Syngonanthus {Carp- be interpreted as a lack of specialization. This same Thy found some Paepa- situation is also in species of Mesanthemum. A diverj lanthus subg. Xeractis Koern. {P. digitiformis matous morphology was observed chrysan- Hensold, P. dianthoides Koern.). in S, Ruhh thus (Bong.) (sect. Dimorphocaulon Ruhl.), Thus, certain parallels can be drawn between S. caulescens (Poiret) Ruhl. (sect. Carphocepha- Rondonanthus and Paepalanthus suhg. Xeractis, and lus Ruhl.), Philodice. which are both interpreted as primitive within their Aerenchymatous unknown roots are in the genus respective lineages (Hensold, 1988). Both possess, Paepalanthus, however, and we have never also in their less specialized species, lanceolate, leaflike observed malpighian hairs in this group. The pres- bracts which surpass the head. In subg. Xeractis ence commonly of hairs at the base of the ovary a character these bracts are pubescent on the inner is that has not been systematically surveyed in the surface, and this is also found occasionally in Ron- family but which we have not encountered in Pae- donanthus, as also in Mesanthemum. Both also The palanthus. white base color of the perianth have typically pilose corollas of the staminate flow- and often also the involucral bracts a conspic- ers, though this also an extremely rare char- is is common uously character in Syngonanthus, but acteristic elsewhere in the Paepalanthoideae, known probably rare (if at all present) in Paepalanthus. only in a few species of Paepalanthus closely re- On number the other hand, a of other characters lated to subg. Xeractis. occur Rondonanthus which much more Another Rondonan- in are exceptional character of commonly observed in Paepalanthus. The most thus the fusion of the filament base with the is notable the deeply branches found This occurs is bifid style in corolla. in four of the five species of species of Rondonanthus and most Paepa- Rondonanthus. R. all in (In capillaceus, the only spe- lanthus, but never described from Syngonanthus cies in the genus with an ''androphore," or elon- or Leiothrix. gated above floral axis the sepals, the petals are Certain characteristics are shared by Rondo- inserted at the base of the androphore while the rta«/Au5 with Pae/?a/artf/iu5 and also the putatively stamens are inserted at apex, and therefore its more primitive Eriocauloideae {Eriocaulon L. and adnation of the filaments impossible.) Adnate is Mesanthemum). common These opaque are: the blackish filaments are in the Eriocauloideae (both 444 Annals of the Garden Missouri Botanical Eriocaulon and Mesanthemum) but not heretofore however, does not create problems for Stiitzel's (We we suppose documented Paepalanthoideae. have phylogeny, especially that free petals the if in them occur Syngonanthus are a derived character in the genus. In addition, also found to in sect. Rondonanthus shown be Cnrphoccphalus and amazonicus Mold.) the styles of are to s.l. in S, rather than simple, and the filaments adnate bifid, which characters are com- to the corolla, of all Phylogeny Implications for patible with Stiitzers system. Rondonanthus thus combines characteristics of This three-subfamily system problematic. is Paepalanthus and Syngonan- however, not phylogenetic interpretation of the Eriocauloideae, in its thus. apparently has strong affinities to Syn- "' [Furt^ac Arm,'* but in the cladistic emphasis places It it ^onn-^^/Aus but differs from that genus in characters on "syngonanthoid" fusion of the pistillate flower that seem more primitive in the subfamily or family corolla. Stiitzel cited scanning electron microscope (SEM) as a whole. Moreover, presents variation in flower developmental studies of flowers (Stiitzel, it sexuality, fusion of the petals, and presence of 1984) which have revealed the pistillate flower Syngonanthus trichome wall thickenings, characters which have corolla of to be congenitally fused. all been used to define genera. while that of Paepalanthus shown to be con- is He concluded that the flowers of genitally free. fl Paepalanthus cannot be derived phylogenetically IHfi between Syngonanthus and Mesanthemum. Mes- from those of Syngonanthus, and that the genus anthenium also shows Syngonanthus-\\ke fusion must therefore have a separate origin in the Er- of the pistillate flower corolla, but like Eriocaulon iocauloideae, perhaps in Eriocaulon, in which pet- possesses two whorls of stamens and an unmodified als of both sexes of flowers are free. gynoecium. As evidence, Stiitzel cited (a) the bifid This classification, however, assumes that evo- style branches of Wurdackia^ which he interpreted lution of the highly modified gynoecium with its as primitive relative to the simple style branches *'stylar appendages," not to mention the reduction of Syngonanthus; and (b) the adnation of the of stamens from two whorls to one, must occur fil- Wu The assume twice evolution. alternative to in is Mesanthemum, but believed by him to be lacking either that (a) the syngonanthoid corolla arose twice Paepalanthus was in Syngonanthus. in separate lines, or (b) origi- from syngonanthoid did not consider the bisexual flower of nally derived a species with a Stiitzel ^''Wurdackia'''' to represent a necessarily primitive corolla, in spite of the lack of ontogenetic evidence. The seem observing that eriocauls differentiation of alternative does not unlikely. state, in first floral sexuality occurs late in development (Stiitzel, especially in view of the finding that corolla fusion much 1985) and might be readily subject to evolutionary a character showing less stability, es- is inodification. Having found that bisexual flowers pecially in the primitive genus Rondonanthus, than (1) same The occur together with staminate flowers in the that of gynoecial modification. paepalanthoid head; ^''Wurdackia'''' has very close sympatric gynoecium furthermore must be thought of as in- (2) relatives with normal pistillate and staminate flow- volving more than one major evolutionary change, an and these related species possess long the modification of the ancestral style into ers; (3) fila- i.e., we mentlike staminodes in the pistillate flowers, ''appendage" (nectary) and the formation of non- agree that the bisexual flowers of ^'''W^urdackia^^ vascularized functional styles at a position rotated 60 are probably secondarily derived. degrees from the position of the original styles. In the same publication, Stiitzel (1985) proposed As far as we know there are no intermediate forms a new three-subfamily system of classification for in existence, and no cases in the Paepalanthoideae the Eriocaulaceae, in which the Syngonanthoideae where the gynoecium has reverted to the primitive {Mesanthemum, Wurdackia, Syngonanthus, and state. The second alternative also not hard to is Philodicc) and the Paepalanthoideae s. str. {Pae- visualize, if we do not assume that ontogeny fault- palanthus, Lachnocaulon Kunth, Tonina Aublet, recapitulates phylogeny. lessly We Rondonanthus) Syngonanthus and seem Rlastocaulon and are agree Ruhl., in- that its allies may dcpendently derived from the Eriocauloideae, con- to constitute a natural group which be usefully from Paepalanthus and taining only Eriocaulon. distinguished its allies. What Rondonanthus (represented only by R. rorai- most necessary present to survey the at is is mae) was evidently placed the Paepalanthoideae subgenera and sections of these two large, poly- in because of the free petals in the pistillate flowers. morphous, and probably unnatural genera in an The union of Wurdackia with Rondonanthus, effort to better characterize the taxa involved and Volume Number & 2 Hensold 445 78, Giulietti Rondonanthus 1991 Revision of to more fully identify and survey characters of abaxially at apex. Flowers 3-merous, either sta- evident taxonomic importance. minate and or staminate and bisexual, the pistillate Moreover, Rondonanthus taken as a prim- two types usually found in diflFerent zones in the if is outgroup Syngonanthus, may same capitulum roraimae) most itive of aid in the or rarely ca- (/?. it assessment of whether Syngonanthus mono- or pitula and plants unisexual. Staminateflowers. Ped- is mm, how polyphyletic, and the latter, to be prop- icels 0,3-^2.5 sometimes equaling or even if it is erly divided to reflect phylogeny. exceeding the flowers in length. Sepals free, elliptic We more feel practical and parsimonious to spatulate, pubescent abaxially at apex. Petals it is to retain for the present Ruhland's two-subfamily free or connate into a 3-lobed tube, and then some- much classification while acknowledging that more times separating after anthesis, glabrous or pilose information needed to build a strong phylogenetic at upper margin. Androphore usually lacking; when is understanding of Eriocaulaceae. present, the petals free and borne at base. Sta- its mens when adnate to corolla at base, except sep- Taxonomy arated from corolla by androphore; anthers dor- and sifixed, white or cream. Pistillate bisexual Rondonanthus Herzog, Feddes Repert. Spec. mm ^^^^^^ 2_2.0 p^j^^^lg q long. Sepals free, Nov. Regni Veg. 29: 210. 1931. type: Ron- ^^^^^ pigmented ^^ ^^^-^^-^ ^^ spatulate, as in the donanthus roraimae Herzog. (Oliver) staminate flowers or lighter, pubescent as in the Wurdackia Mem. New York staminate flowers, expanding and thickening with Mold., Bot. Card. 413. 9: 1957. TYPE: Wurdackia flabelliformis Mold. maturation of the fruit, especially at base. Petals Paepalanthus Bostrychophyllum subg. Ruhl., Pflanzenr. connate and free or distally free at the base, gla- IV. 30: 220. 1903. type: Paepalanthus capilla- brous or pilose at upper margin and abaxially. ceus Klotzsch ex Koern. more Linear staminodes, or rarely developed fully Plants rosulate or caulescent cespitose peren- stamens, present opposite petals, the staminodes nials branching at ground level. Roots creamy white usually about half to twice the height of the ovary to pale brown, porous (aerenchymatous), brittle, and bearing two tiny brownish lobes at the apex, not spongy, usually sparingly branched, the Ovary usually subtended by a ring of long hairs, much branches evidently smaller in diameter than the style much shorter than the style branches; the mm 2-10 main Stems roots. ca. diam., woolly with appendages infundibular, stalked, papillose, insert- fine cream to tan-colored filamentous hairs. Leaves ed on style at or commonly slightly below the level arranged spirally or distichous, filiform to linear or of divergence of the style branches; the style young ligulate, the leaves attaining full width long branches strongly bifid. Fruit a loculicidal capsule, mar- before full length; the bases also with parallel Seeds ellipsoidal, reticulate, the longitudinal stria- gins, but dilated with respect to the blades, at least tions of the testa raised and more conspicuous than somewhat and distally, scarious frequently woolly the transverse. Floral trichomes subacute to sub- with hairs like those of the stem. Pubescence of clavate, with or without granular thickenings of leaves and peduncles when present mostly mix- a the internal wall, ture of filamentous and malpighian hairs, the latter usually "retrorse" (i.e., strongly asymmetric with Key to the Species of Rondonamiius end the larger proximal); capitate hairs lacking. ^^ submerged p,^^^^ aquatics; leaves filiform-se. numerous Inflorescences single to per age class heads taceous, usually surpassing the peduncles; mm ± 3-5 and then usually synchronous. Peduncle sheaths diam.; floral bracts and sepals short- ± hairy capillaceus membranous /?. with respect to the leaves, often lb. Plants terrestrial; leaves linear to ligulate, al- pale and scarious at least toward the upper margin, mm 6-19 ways surpassed by peduncles; heads closed over the young inflorescences and splitting diam.; bracts and sepals densely bearded floral down one side from just below the apex, the opening with long hairs. Leaves oblique with the upper margins remaining minutely 2a. distichous. Stems 3a. leafy to base; flowers bisexual and involute often eventually lacerate. Peduncles and staminate R. flabelliformis mostly 3- or 6-costate, sometimes apparently more. Stems 3b. leafy only at apex; flowers pis- Capitula turbinate to hemispheric at maturity, rare- and staminate R. acopanensis tillate ly subglobose. Involucral bracts, or at least their 2b. Leaves spirally arranged. 4a. Peduncles 3-costate; involucral bracts visible apices, triangular to lanceolate, acute to of upper series acuminate, the acu- subacute, gold to nearly black, or variegated. Re- men R. coidescens colorless ceptacle pilose. Floral bracts present, usually about 4b. Peduncles 6-costate; involucral bracts equaling the flowers, linear to spatulate, pubescent of upper series subacute to acute, black 446 Annals the of Garden Missouri Botanical 4 ^ mm \ » I 50|jm 4 M : j b 4 i + ^ I. » I- -, I' ' r ' I 0.1mm fr L mm OS & Figure Rondonanthus acopanensis (Mold.) Hensold acopanensis {Steyermark 128872 1. Giulietti var. — — — MO). E-G. A. Habit, B. Inflorescence. C, D. Involucral bracts. Staminate flower. E. Flower with bract, past — — anthesis (corolla involute at upper margin). F. Sepal, adaxial view. G. CoroUa tube, opened to show stamens and — — pistillodes. H, I. Pistillate flower. H. Flower with bract, past anthesis. Petal, with hatched lines showing where I. — — ^K. fused with adjacent petal, and staminode adhering at base. J. Gynoecium (fruiting). Seed. L, M. Trichome of bract apex. floral 1 Volume Number & 78, 2 Hensold 447 Giulietti 1991 Rondonanthus Revision of at apex and margins, though some- and usually ciliate hirsute abaxially, the inner times the midvein pale. densely bearded at the apex, at least abaxially and Leaves 5a. sharp-cuspidate to aris- sometimes also adaxially, glabrate with age. Floral tate; petals of staminate flowers commonly bracts exceeding persistently connate and the flowers, linear to ciliate at upper margin R. duidae linear-oblanceolate, acuminate, sometimes with a 5b. Leaves obtuse subacute; to petals long ''whip tip," blackened distally, tufted with of staminate flowers connate ear- hairs abaxially at apex, Staminate flowers. Pedicels but separating by ly anthesis, gla- mm 0.7-1.3 long. Sepals obovate-elliptic to spat- brous R. roraimae ulate, attenuate-acute to long-acuminate, 2.3-3.5(- mm mm 0.6-0.9 4.0) long, wide, white at base, & Rondonanthus acopanensis (Mold.) Hensold blackened distally except for white midvein and comb. nov. Syngonanthus acopa- Giulietti, apex, the apex tufted abaxially with divergent tri- nensis Mold., Phytologia 41. 1948. TYPE: 3: chomes like a bottlebrush. Corolla connate into a Venezuela. Bolivar; Chimanta Massif, Cerro androphore 2.0- 3-lobed tube, the larking, the tube Acopan, ,900 m, Cardona 2280 (holotype, mm 1 3.5(-4.5) long, the lobes triangular-obtuse, US; isotype, NY). Figure 1. upper margin and the abaxial lobe apices ciliate- Plants short-stemmed cespitose perennials. Main pilose, involute and retracting the stamens after mm 0.5-1.2 roots ca. diam., sparingly branched, anthesis. Stamens with filaments adnate to corolla mm cream to pale orange-brown. Stems 1-4 cm long at base, ± included; anthers ca. 0.30.35 mm mm 2-5 at flowering, ca. diam., leafy only near long. Pi5ft7/afe/ou;er5. Pedicels 0.6-0.8 long, apex, but the lower portions covered with tightly Sepals to ovate-elliptic, concave, acuminate, elliptic mm mm persistent flabelliform leaf bases (these making the 3.5-4.0(-4.7) long, 1.0-1.5 wide, cream- mm 5-20 stem appear laterally compressed and colored or pigmented as in staminate flowers though wide), woolly with cream-colored Leaves more weakly apex hairs. dis- usually so, tufted at the as in mm 3-13 cm tichous, long, l-4(-6) wide, the base the staminate flowers, thickening at base at ma- and blade abruptly diff*erentiated, the base linear, turity. Petals connate into a tube except at base, ± dilated with respect to blade, scarious, densely en- the segments oblong, rounded apex, 2.8- at mm mm folded with long cream-colored woolly hairs (often 3.5(4.7) long, 0.6-0.9 wide, densely and black with dirt), persistent, a transverse ab- ciliate at upper margin and frequently also pilose scission layer apparently forming at the juncture abaxially at apex or in submarginal bands. Stam- mm with the blade; the blades linear, subacute to broad- inodes linear, 0.6- .3(2. 5) long. Ovary ringed mm 0- ly rounded or truncate depending on width, car- by hairs at base, 0.4-0.7 long, the style mm mm tilaginous and rigid, glabrous or pubescent with 0.3 long, the appendages ca. 0.4 long, malpighian when young retrorse hairs especially the stalk thick, the papillose portion orange-brown, adaxially, 5-30-veined, the veins of equal size. the style branches diverging slightly above the ap- mm Inflorescences single per age class, the old ones pendages, 1.5-2.3 long, conspicuously bifid, 211 often persistent on stem. Peduncle sheaths Floral trichomes subacute to rounded, lacking cm long, twisted, mostly striate-pubescent with re- granular thickenings of the internal wall. trorse malpighian hairs, the apex acute to rounded This species endemic to the Chimanta Massif is (corresponding in shape to leaf apex), green or in Bolivar, Venezuela, where represented by it is scarious, frequently lacerate with age. Peduncles two sympatric varieties, heretofore recognized as cm I2-45(-60) sometimes They commonly long, 6-costate, ap- species. are about equally col- parently more by division, pubescent with retrorse lected and are distinguishable only by their size malpighian hairs and long filamentous hairs fringing and robustness, as well as possible habitat diff"er- 7.5-12(15) the base of the capitulum. Capitula ences. This sort of variation crops up not uncom- mm diam., turbinate to hemispheric at maturity. monly in eriocauls (as in Pnepalanthus supcrbus 3-5 Involucral bracts in series, the outer narrowly Ruhl. and P. chlorocephalus Silveira in the Serra triangular to lance-linear, subacute to acute, erect do Cipo, Brazil; Hensold, 1988), and possibly is mm to slightly recurved-spreading, 2.5-5.5 long, associated with polyploidy or some other macro- mm 0,6-1,0 wide at base, the inner bracts oblan- mutation. This would imply that the larger form mm 4-7 may ceolate, broadly acuminate, erect, long, be '*polyphyletic," having been derived sev- mm 0.8-1.3 wide, mostly surpassing the head by eral times from the smaller form. (An alternative mm; ca. 0.5-1.5 the outer bracts pale brown to view would be that the smaller form may have been when castaneous, or greenish fresh, the inner black derived from the larger form.) However, because except for the pale tip and midvein; the outer bracts we lack cytological data, and since each type is 448 Annals of the Garden Missouri Botanical 12- about equally widespread and forming uniform pop- Stems, including attached leaf bases, ca. mm mm 11- ulations, we have treated them as varieties. 20 wide. Leaves (2-)2.8-6 wide, ca. mm mm I9(-30)-nerved, rounded to nearly truncate, very 1-2 7.5-10 Leaves wide; heads diam.; la. Huber chartaceous firm-coriaceous (only firmly in involucral bracts mostly ca. 3-seriate Rondonanthus acopanensis var. acopanensis 11522), the margins, especially near the apex, mm 10-14( Leaves (2-)2.8-6 wide; heads 15) lb. Heads 10-14(- often shiny and deep brown-tinged. 4-5- mtn diam.; involucral bracts mostly ca. mm (3-)4-5 15) diam. Involucral bracts in series, seriate mm 3.5-6 0.7- the lowest lance-Hnear, ca. long, Rondonanthus acopanensis var. obtusifolius mm nun 3.5-6.2 Male wide. Floral bracts long. .0 1 mm 0.6- Rondonanthus acopanensis acopanensis flowers with sepals 3.0-3. 5(-4.0) long, var. mm mm 0.9 wide, corolla 3.0-3.5(-4.5) long. 5-10 mm Stems, including attached leaf bases, ca. p^^^j^ fl^^^rs with sepals ca. 3.5(-4.7) long, mm mm Leaves 1-2 5-7(-9)-nerved, mm mm wide. wide, ^^ 1.3- 3.0-3. 5(-4.7) 1.5 wide, corolla firm, green, subacute, the margins not turning shiny |^^ mm brown. Heads 7.5- 10 diam. Involucral bracts mm Phenology. January 2.5-3.5 Collected in flower late 3 the lowest lanceolate, long. in series, mm mid-March. 0.6-0.9 3.5-4.7 nun to wide. Floral bracts long, mm 2.3-3.3 Staminate flowers with sepals long, Habitat and distribution. Locally abundant mm mm 0.6-0.8 and 2.0-2.7 wide, corollas long, among ^^^^ ^^^ savanna and sandstone out- j^ mm 3.5-4.0 flowers with sepals long, Pistillate crops, at 1,800-2,200 m. Venezuela. Bolivar: Chi- mm mm 1.0-1.4 and 2.8-3.2 wide, corollas long. ^^^^^^ ^^^^^^ (Abacapa-, AcopAn-, Apacara-, and Chimanta-tepuis). PIleno logy. Collected in flower mostly from January to March, but also occasionally in June Additional specimens examined. VENEZUELA, boli- & and August. Chimanta S Huber var: Massif, Apacara-tepui, sector, 128428 Steyermark 7061 (LL, NY), Steyermark (LL, Habitat and distribution. Locally abundant ^^^ Steyermark 128519 (LL, MO); Apacara- et at. on wet sandy depressions and banks on shallow SE 8854 Huber tepui, sector, lluher et at. (LL, NY), et open scrub 1,900-2,600 aL 8878 (LL, NY), Huber 11522 (MO), Huber 11524 soils, in the or in forest, NE & Huber Steyermark (NY); Chiinanta-tepui, sector, m. Venezuela. Bolivar: Chimanta Massif (Acopan-, ^, »# 666'^ (LL, NY); Chiiiianta-tepui, central-NE sector, S/(?r- A. pa^cara-, Lhn.nanta-' , (/TL--ruoda1 -lt\ epuis• )\. m- ' iVlurey- ,oo/.o/wmv ^ ' ' t crmark, , . 128090 (NY. M\;Oi)o\; Lhimanta-t*epui,• cen- -^ ^ -' / et at. //ui.rc/ aL (MQ); Abacapa-tepd sector Additional specimens examined. VENEZUELA. ^^^^^ W L^^'"^ * 8620 {NY); Acopan-tepu, N ^ ^^'^'^^ Huber ^^<^^^>^^ bolivar: Chimanta Massif, Apacara-tepui, section, !^'y/^ & & & (Anmri-tepm), Huber Steyermark 7082 (LL 8685 NY, VEN), Huber 8737 Colella (LL, Colella f^^^o'' ^^N), Steyermark 128453 (LL MO). Steyermark ct at. Steyermark (LL, NY); Apicara-tepui, E-central section, ^U NE Huber & Acopan-tepui, secXor e^ 75925 Apicara-tepui, S section, Huber Steyer- '\^[- (F); ;?^!^:t W65 SE SSE '0152 (LL NY), & (NY), sector //.6.r 6970 «^- mark 6964 MO), Huber Steyermark (LL, (LL); Steyermark al 129909 VEN). (LL, ^^^^^^^ et SE Steyermark 75850 Apacara-tepui, section, (F); Chi- manta Murey- Massif (Eru^^^^^^ Huber 11522 specimen included here ^.j^^ ^ is Steyermark 158l8'A (LL, VEN); Chimanta Massif, ^m Clii- 1 r c • some from i ^^th reservation. differs typical var. 128165 It Steyermark manta-tepui, central-NE sector, et al. (LL, MO, NY, VEN); Chimanta-tepui, E sector, vicinity obtusifolius not only in its overall larger dimen- & Cano Huber Steyermark 160 MO), of del Grillo, 7 {LL, sions, but in lacking stem elongation, a distichous & Steyermark Wurdack 804 NY), Steyermark (F, leaf arrangement, and coriaceous-thickened leaves. 128938 128872 MO), Steyermark Chimanta (LL, (LL); Normal {Huber 1524) been var. obtusifolius has 1 Wurdack 34213 NY, VEN), Massif, Churi-tepui, (F, ceii- from same Huber 8934 NY), Huber collected the locality. tral-SE section, <& Colella (LL, km 9275 (MO); 10 SE Chimanta Angasiiua-tepui, of Huber 1688 Massif, {MO). Rondonanthus 1 capillaceus (Klotzscb ex Koern.) & Hensold comb. Paepalan- nov. Ciulietti, Rondonanthus acopanensis var. obtusifolius thus capillaceus Klotzscb ex Koern. C. In & (Mold.) Hensold Giulietti, comb, et stat. nov. Martins, Fl. Bras. 3(1): 415, 53, fig. 11. t. Mem. New Syngonanthus Dupatya obtusifolius Mold., 1863. capillacca (Klotzscb ex York Bot. Card. 9: 410. 1957. TYPE: Vene- Koern.) Kuntze, Revis. Gen. PI. 2: 745. 1891. zuela. Bolivar: Chimanta Massif, central sec- TYPE: Guyana: savannas of Mount Roraima & 1222 1,940 m, 4 Feb. 1955, Steycnnark and Humiridia, April-May, Schotnburgk tion, Wurdack 406 NY; B (holotype, isotypes, F, GH). (holotype, not seen). Figure 2. Volume Number 2 Hensold & 449 78, Giulietti Rondonanthus 1991 Revision of — & Krai 72131 MO). Figure Rondonanthus capillaceus (Klotzsch ex Koern.) Hensold Giulietti A. 2. (/?. — — — Whole Habit. B. Inflorescence. C, D. Involucral bracts, lower and upper. E-I. Staininate flower. E. flower, with — — — J-M. bract.— G. Flower with perianth spread open. H. Sepal, abaxial view. Petal. floral F. Floral bract. I. — — — M. Petal.— flower.- Flower with bract. K. Flower with perianth spread open. L. Sepal, abaxial view. Pistillate J. N, 0. Trichomes of bract apex. floral 450 Annals of the Garden Missouri Botanical mm Paepalanthus 0.3-0.5 hippairichophyllus Herzog, Feddes Re- flowers. Pedicels ca. long. Sepals ob- 193L pert. Spec, Nov. Regni Veg. 29: 208. TYPE: ovate to spatulate, obtuse to rounded, concave, Brazil. Arnapa: Igarape Cre-cru of the Rio Oyapock. mm mm somewhat 1.1-1.4 0.45-0.6 fleshy, long, 21408 Laetzelhurg (holotype, M). wide, cream-colored to pale gray-brown, and ciliate *alanthus capillaceus var. proUferus Gleason, Bull. Torrey Bot. Club 58: 328. 1931. Paepalanthus short-hairy abaxially toward apex. Corolla of 3 capillaceus forma prolifvrus (Gleason) Mold., Phy- separate petals (these separate from earliest visible tologia 44: 384. 1979. type: Venezuela. Amazonas: stage of development), inserted at the base of the Cerro Duida, Central Camp, 4,800 Tate 552 ft., androphore (intercalary prolongation of the floral NY). (holotype, above Paepalanthus New ^^is the calyx), or at variable distances above capillaceus var. spiralis Mold., Mern. York Bot. Card, 9: 279. 1957. type: Guyana. Up- the base, but usually below the insertion of the per Mazaruni River, Karaurieng River, Maipuri Falls, filaments and androphore 0.1-0.5 pistillodes; the & 1,250 m, Maguire Fanshawe 32292 (holotype, mm mm 0.40.7 1.1-1.5 long, the petals long, NY; isotype, M). Note: This taxon, distinguished by mm wide, obtrullate obovate, rounded toward to known strongly spiral-twisted leaves, only from its is the type locality. Though may be valid as a variety, apex and abruptly apiculate, glabrous. Stamens it mm it is here regarded conservatively as a synonym. with filaments 0.7-1.2 long, usually inserted at apex of androphore, though occasionally one or Plants short-stemmed aquatic perennials, the more inserted at base, included, the anthers ca. mm vegetative portions submerged and fully infl 0.15-0.2 long. Pistillateflowers. Pedicels 0,2 mm cences mostly emergent and Main erect. roots 0.4 long. Sepals ovate' concave, to elliptic, mm abundant, 0.5-1.0 diam.,much.branched, mm mm ca. acute to obtuse, 1.2-1.5 long, 0.5-0.8 cm gray-brown, contorted. Stems up and to 7 long ^ide, colored as in staminate flowers, short-ciliate mm 15 diam., leafy only near the apex, but the from middle apex and to short-hairy abaxially in lower portions covered by and persistent roots leaf this region, coriaceous-thickening at maturity, es- bases, woolly with brownish Leaves hairs. spirally pecially at base and sometimes throughout. Petals arranged, green (2.5)7-30 cm pale to blackish, free, spatulate, usually acute or apiculate, 1.2-1.6 0.2-0.4 inm mm mm long, wide, the base slightly dilated, 0.40.6 long, wide, glabrous. Staminodes ephemerally woolly, coarsely filiform-setaceous, firm, linear to somewhat broad and nearly scalelike, sometimes mm corkscrew-twisted, the acute but tips 0.2-0.6(-l Ovary .4) long. with a few seat- usually broken glabrous (though often with sand mm off, 0.4-0.7 tered hairs at base, long, the style particles and algae adhering in dried material), 0.1-0.4 mm mm appendages 0.4-0.5 long, the traversed by a single central vein surrounded by long, the stalk thick, the papillose portion orange- mesophyll (Ruhland, 1903). Inflorescences nu- brown, darker than the the branches stalk, style merous, synchronous, commonly numbering 10 or diverging at the same level as the appendages, 0.7- more. Peduncle 3-5 cm mm sheaths long, strongly 0.9 long, bifid half to nearly their length, all twisted, glabrous, the apex acuminate, scarious, Floral trichomes clavate and thick-walled with dense cm often lacerate. Peduncles (5.5)9.0-20.0 long, granular thickening of the internal wall. 3-costate or apparently 6-costate by collapse of main 3- Phenology. the centers of the ribs, glabrous. Capitula Collected in flower year-round. mm 5 diam., depressed-turbinate depressed- to and Habitat Adhering distribution. rocks to hemispheric maturity, sometimes at vegetatively streambeds 100-1,600 of in rapidly flowing water, from proliferating the center. Involucral bracts in m. Venezuela. Amazonas: Cerros Duida and Mara- ca. 4 series, both the outer and inner narrowly h south Gran to Neblina; Bolivar: Sabana. commonly triangular, subacute to obtuse, erect, Guyana: Upper Mazaruni and River Kaieleur Pla- mm mm 1.1-1.4 0.50.7 the outer ca. long, wide, Amazonas: Amapa: teau. Brazil. Serra Araca; Rio mm mm the inner 1.9-2.1 0.50.7 long, wide, Oyapock (Oiapoque). about equaling the head; pale gold-brown to all castaneous or nearly black, fleshy or coriaceous- Additional specimens examined. VENEZUELA. amazonas: Cerro Duida, Farinas 420 (NY), Ma- thickened especially toward the base, the margins et al. ± .u.^a^;^. S^^^^' <^ ^^m^^S^^^u^^^^ec ^2:9^1i5^3^ ^(iM^iOvy,, iN^Yx);,, Mifaiguu^uiurte, et uail.. --^^---^^ i^i of at least the inner bracts --^ Pneeilliuucciida sshnrreeddddimngg ^g^jj 751-82 (NY); Cerro Mara- ^p^y); Tillett et al. . wi.t,h age; gl, a,brous or t,he mner bracts short-cdiate & huaca, Steyermark Hoist 130470 (MO); Cerro Ara- & & along distal margins. Floral bracts shorter than or camuni, Liesner Carnevali 22308 {}A0\ Wurdack Adderley 43602 about equaling the (NY); base of Cerro Neblina, Davidse flowers, spatulate, obtuse, strongly k,ee1led and somewhat f ^1^"^' cucullate apex, at t.^'TJ^?'^^ ^'''^^f.^?:!^.A'^?^^^"''^ Liesner 15737 (MO); Liesner 16653 (MO). BOLiVAR: , cream-colored to pale b, rown, t,he apex h, i. rsute Canmma, 7745 ESE 5rA;z^^ (MO); Rio Yune, of Churi- abaxially with short subclavate Staminate hairs. tepui, Ruber 9764 (MO); headwaters of Rio Venamo,

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Revision and Redefinition of the Genus Rondonanthus Herzog (Eriocaulaceae) PDF

19 Pages·1991·15.5 MB·English
by  Nancy Hensold
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Preview Revision and Redefinition of the Genus Rondonanthus Herzog (Eriocaulaceae)

AND REVISION Nancy and Hensold^ Ana Maria THE REDEFINITION OF GiuliettP GENUS RONDONANTHUS HERZOG (ERIOCAULACEAE)^ Abstract The genus Rondonanthus Herzog now has until contained only the species R. roraimae and R. micropetalus, and has been defined by free petals in both sexes of flowers and the dioecious condition. here redefined according It is to a large suite of characters, especially the presence of long filamentous staminodes in the pistillate flowers, and by the adnation of the filament base to the corolla in the staminate flowers. Five additional species, including the monotypic genus Wurdackia, are transferred to Rondonanthus, and /f micropetalus removed to Paepalanthus. proposed is It is . Rondonanthus that is a highly primitive paepalanthoid genus, to be taken as a primitive outgroup of Syngonanlhus, The Eriocaulaceae are a pantropical monocot paepalanthoid genera. These are Rondonanthus family of about 13 genera and 1,200 Herzog Comanthera Lyman species (Krai, (1931), Smith B. Except 1989). in rare cases, the flowers are uni- (1937), Carptotepala Mold. and Wur- (1951), sexual and the plants monoecious, with the flowers dackia Mold. (1957). borne in scapose heads. Following the lead of Ruhland (1903), of all Ruhland (1903) divided the family into two sub- these genera were described with reference only to families, the Eriocauloideae, characterized by two a few floral characters. None contain more than whorls of stamens and a simple gynoecium, and two species, and none were discussed by their au- the Paepalanthoideae, with a single whorl of sta- thors as to their relationships with other genera or mens and "appendages" vascularized secretory in- implications for phylogeny. Recently, the avail- on serted the style alternate with the nonvascu- ability of abundant collections from the Venezuelan The larized functional style branches. Eriocauloideae tepuis and their study for Julian Steyermark's Flo- are truly pantropical and contain many aquatic a of the Venezuelan Guayana r {R^n^yoXA, in ^re^.) species, while the Paepalanthoideae are almost en- have acutely emphasized the need study and for New World some tirely limited to the (with African reevaluation of these endemic genera. & and more disjuncts), xerophytic. In another work Hensold, (Giulietti 1991), The we Paepalanthoideae, which contain 11 of the discuss the systematic position of Comanthera 13 genera, have two centers of diversity, one in and Carptotepala, concluding that they are syn- the Central Brazilian Plateau, especially the Cadeia onyms of Syngonanthus Ruhl. In work, we this do Espinha^o in Minas Gerais and Bahia, and one find the genus Rondonanthus be though to valid, Guayana in the Highland, especially Venezuela. in need of redefinition, and include within five it Botanical exploration of Guayana, in particular the additional endemic Guayanan species currently summits tepui of Venezuela, has only been under- placed in three other genera, including the mono- taken recently relative to that of the more acces- typic genus Wurdackia, mountains and Rondonanthus sible of central Brazil, has turned as here recognized a genus of is many up taxa of interest, including four endemic great interest from the standpoint of phylogeny in We VEN thank MO, NY, PORT, the curators of F, K, LL, and for the loan of specimens. The work was ' completed while the junior author was a visiting research scientist under the Missouri Botanical Garden Postgraduate Fellowship Program, supported by the Jessie Smith Noyes Foundation. Habit by Bruno Manara illustrations were provided by the Flora of Venezuelan Guayana Project. Department Museum 2 of Botany, Field of Natural History, Roosevelt Road at Lakeshore Drive, Chicago, Illinois 60605-2496, U.S.A. Departamento ^ de Botanica, Instituto de Biociencias, Universidade de Sao Paulo, C.P. 1.461-05499 Sao Paulo 1 SP, Brazil. Ann, Missouri Bot. Card. 441-459. 78: 1991. 442 Annals of the Garden Missouri Botanical Eriocaulaceae. appears to combine characters rollas, with the corollas of the stamlnate flowers It of the Eriocauloideae with characters of two ''core" tubular and those of the pistillate flowers connate paepalanthoid genera, Paepalanthus and Syngo- at the middle and free at the base and the apex. mird nan thus. As (1985) concluded (limiting his Stiitzel known observations R. {Wurdackia) flabelllformis\ species of the family with bisexual flowers, to it Rondonan- appears represent a primitive outgroup to the All of these taxa are here treated in to large genus Syngonanthus. As a genus, also thus, though their diff'erences in corolla fusion it and caused shows unusual variability with respect to certain characteristics flower sexuality originally characters, such as fusion of perianth parts them to be assigned to four diff"erent genera, floral Rondo- now The and which have been only other species described in flower sexuality, until given a priori value in the definition of genera of nanthus since the type species, R. micropetalus Eriocaulaceae. Mold. (1951), reputedly with free petals in the synonymy removed staminate here flowers, to is Paepalanthus Excluded Ruhl. Species). (see in Taxonomic History The genus Rondonanthus was described by Generic Definition and Affinities Herzog (1931) on the basis of a single Luetzelburg collection of R. roraimne (Oliver) Herzog, based The single character that best distinguishes Ron- on Paepalanthus roraimae Oliver. donanthus from other genera of Eriocaulaceae all ma- Herzog found only staminate flowers in his the presence of long linear staminodes in the is known and defmed the genus by the free petals of flowers. Staminodes are in Pae- terial pistillate unknown the male corolla, which he believed to be palanthus, but they are small and scalelike, never Paepalanthus. Because he did not see the type linear. The only species of Rondonanthus in which in depended on material, he Oliver's illustrations for flabellifc description of the flowers, which he they are replaced by functional stamens which his pistillate described correctly as having the petals free and produce and release pollen in apparently normal Herzog incorrectly as having simple style branches. quantities. he was Rondonanthus by mixture also noted that the limited material studied further defined a is dioecious and suggested this as an additional char- of characters individually found in other genera, acter of the genus, also encountered rarely in the especially Syngonanthus and Paepalanthus, but family. not in combination with each other. Ruhland comprehensive treatment The Rondonanthus appear (1903), in his affinities of to lie of the family, had in fact erected Paepalanthus most closely with Syngonanthus, for a number of subg. Bostrychophyllum Ruhl. to accommodate reasons. In R. acopanensis, R. caulescens, and known R P. capillaceus Klotzsch ex Koern., the single Jlabelliformis, the petals of the pistillate flowers . species of Paepalanthus with free petals in the are fused in the middle but free at the base and staminate flowers. This species a vegetatively apex, which considered to be one of the defining is is specialized submerged aquatic also native to Ro- characters of .SjT/g^onan^/zu5. Elsewhere in the Er- raiina, and here treated as a species of Ron- iocaulaceae, this known only in the genus Phil- is is donanthus. (Ruhland had placed P. roraimae in odice C. Martius (2 spp.), which considered to is ''Species dubiae" since he had observed no material be derived from Syngonanthus, and in Mesan- of the species.) theniuni Koern., which with two whorls of stamens The same year that Herzog described Rondo- and an unmodified gynoecium quite distant from is nanthus, Gleason (1931) described Paepalanthus Rondonanthus. duidae from Cerro Duida in Arnazonas, noting the This character, however, not stable in the is close similarity to P. roraimae Oliver, and diff"er- genus. In R. roraimae and R. capillaceus, the entiating from that species only by minor char- petals of the pistillate flowers are always free, and it acters of gross morphology, and without mention in the variable species /?. tZui^ae, most populations of structure. have free petals, but a single population has been floral Rondonanthus were Later, additional species of found (at Aprada-tepui) with brief petal fusion. In described from the Chimanta Massif and placed in the genus Syngonanthus, the character also shows Syngonanthus acopanensis Mold., 1948, and a small degree of instabiHty. In the reduced, dim- {S, S. obtusifolius Mold., 1957), and in a new mono- erous species S. minutus (Mold.) Hensold (1991), typic genus W^urdackia (JF. jiabelliformis Mold., for example, the petals are free, and this situation 1957). All exhibit typical SyngonanthuS'\v\.e. co- is also occasionally found in sect. 77rjsa«oce/>Aa/t/5 Volume Number 78, 2 Hensold & 443 Giulietti 1991 Revision Rondonanthus of (Koern.) Ruhl., as In 5. ye/ima«ii(Gleason) Giulietti coloration of the involucral bracts, and frequently & Hensold (1991). Petal fusion probably repre- also the upper parts of the perianth; the of tufts sents a primitive state in Rondonanthus, as is also trichornes on the tips of the sepals and floral bracts; considered to be the case in Syngonanthiis, the clavate or subclavate shape of these perianth Fusion of the corollas of the staminate flowers trichornes, which sometimes {R. capillaceus) are is a fairly universal condition In the Paepalanthoi- densely ornamented on the internal wall; and, in deae, but character breaks down Ron- this also in three species, the ciliate margins of the staminate donanthus. Of the species with free petals in the flower corollas. pistillate flowers, R. ra/?t7/acea5 also has free petals All of these characteristics are common the in in the staminate flowers, R. roraimae has petals Eriocauloideae. The not known from any last is connate early in development and then separating, species of SyngonanthuSy but common some is in and R. duidae, except in very rare cases, has of the less specialized members of Pae/>a/a/z^/iu.s, The petals persistently connate. remaining three notably subg. Xeractis. Inner ornamentation wall which species, all exhibit fusion of corollas in the in floral trichomes has also not been described from have Syngonanthus pistillate flowers, similarly fused corollas of but is the rule in Paepalanthus. the staminate Bearded known flowers. sepal apices are Syngonanthus in Other characters which indicate an alliance with (especially Dimorphocaulon, hum- in sect. e.g., 5. Syngonanthus are the following: roots pale and boldtii (Kunih) Ruhl. diamantinensis S. Silveirsi), , aerenchymatous; hairs present on the floral axis at but not common, and involucral bract pigmenta- ovary base; base color of the perianth creamy tion, when occurs, usually more or less trans- it is white; pubescence of malpighian hairs on vegeta- lucent and reddish arenarius (Gardner) Ruhl., (5. tive parts. All of these characters except the last S. anomalus (Koern.) Ruhl.). An exception S. is may also be found in the genus Leiothrix Ruhl., niger Silveira, with long black linear-lanceolate but this genus diff'erentiated by specialized mor- bracts. is phology gynoecium, of the seed testa striations, Furthermore, in R. acopanensis, the outermost and basifixed anthers (Giulietti, 1984). bracts, as well as the inner bract apices, are ap- Stiitzel (1988) found similar root anatomical parently green and leaflike when young, which may structure In /?. roraimae^ Syngonanthus {Carp- be interpreted as a lack of specialization. This same Thy found some Paepa- situation is also in species of Mesanthemum. A diverj lanthus subg. Xeractis Koern. {P. digitiformis matous morphology was observed chrysan- Hensold, P. dianthoides Koern.). in S, Ruhh thus (Bong.) (sect. Dimorphocaulon Ruhl.), Thus, certain parallels can be drawn between S. caulescens (Poiret) Ruhl. (sect. Carphocepha- Rondonanthus and Paepalanthus suhg. Xeractis, and lus Ruhl.), Philodice. which are both interpreted as primitive within their Aerenchymatous unknown roots are in the genus respective lineages (Hensold, 1988). Both possess, Paepalanthus, however, and we have never also in their less specialized species, lanceolate, leaflike observed malpighian hairs in this group. The pres- bracts which surpass the head. In subg. Xeractis ence commonly of hairs at the base of the ovary a character these bracts are pubescent on the inner is that has not been systematically surveyed in the surface, and this is also found occasionally in Ron- family but which we have not encountered in Pae- donanthus, as also in Mesanthemum. Both also The palanthus. white base color of the perianth have typically pilose corollas of the staminate flow- and often also the involucral bracts a conspic- ers, though this also an extremely rare char- is is common uously character in Syngonanthus, but acteristic elsewhere in the Paepalanthoideae, known probably rare (if at all present) in Paepalanthus. only in a few species of Paepalanthus closely re- On number the other hand, a of other characters lated to subg. Xeractis. occur Rondonanthus which much more Another Rondonan- in are exceptional character of commonly observed in Paepalanthus. The most thus the fusion of the filament base with the is notable the deeply branches found This occurs is bifid style in corolla. in four of the five species of species of Rondonanthus and most Paepa- Rondonanthus. R. all in (In capillaceus, the only spe- lanthus, but never described from Syngonanthus cies in the genus with an ''androphore," or elon- or Leiothrix. gated above floral axis the sepals, the petals are Certain characteristics are shared by Rondo- inserted at the base of the androphore while the rta«/Au5 with Pae/?a/artf/iu5 and also the putatively stamens are inserted at apex, and therefore its more primitive Eriocauloideae {Eriocaulon L. and adnation of the filaments impossible.) Adnate is Mesanthemum). common These opaque are: the blackish filaments are in the Eriocauloideae (both 444 Annals of the Garden Missouri Botanical Eriocaulon and Mesanthemum) but not heretofore however, does not create problems for Stiitzel's (We we suppose documented Paepalanthoideae. have phylogeny, especially that free petals the if in them occur Syngonanthus are a derived character in the genus. In addition, also found to in sect. Rondonanthus shown be Cnrphoccphalus and amazonicus Mold.) the styles of are to s.l. in S, rather than simple, and the filaments adnate bifid, which characters are com- to the corolla, of all Phylogeny Implications for patible with Stiitzers system. Rondonanthus thus combines characteristics of This three-subfamily system problematic. is Paepalanthus and Syngonan- however, not phylogenetic interpretation of the Eriocauloideae, in its thus. apparently has strong affinities to Syn- "' [Furt^ac Arm,'* but in the cladistic emphasis places It it ^onn-^^/Aus but differs from that genus in characters on "syngonanthoid" fusion of the pistillate flower that seem more primitive in the subfamily or family corolla. Stiitzel cited scanning electron microscope (SEM) as a whole. Moreover, presents variation in flower developmental studies of flowers (Stiitzel, it sexuality, fusion of the petals, and presence of 1984) which have revealed the pistillate flower Syngonanthus trichome wall thickenings, characters which have corolla of to be congenitally fused. all been used to define genera. while that of Paepalanthus shown to be con- is He concluded that the flowers of genitally free. fl Paepalanthus cannot be derived phylogenetically IHfi between Syngonanthus and Mesanthemum. Mes- from those of Syngonanthus, and that the genus anthenium also shows Syngonanthus-\\ke fusion must therefore have a separate origin in the Er- of the pistillate flower corolla, but like Eriocaulon iocauloideae, perhaps in Eriocaulon, in which pet- possesses two whorls of stamens and an unmodified als of both sexes of flowers are free. gynoecium. As evidence, Stiitzel cited (a) the bifid This classification, however, assumes that evo- style branches of Wurdackia^ which he interpreted lution of the highly modified gynoecium with its as primitive relative to the simple style branches *'stylar appendages," not to mention the reduction of Syngonanthus; and (b) the adnation of the of stamens from two whorls to one, must occur fil- Wu The assume twice evolution. alternative to in is Mesanthemum, but believed by him to be lacking either that (a) the syngonanthoid corolla arose twice Paepalanthus was in Syngonanthus. in separate lines, or (b) origi- from syngonanthoid did not consider the bisexual flower of nally derived a species with a Stiitzel ^''Wurdackia'''' to represent a necessarily primitive corolla, in spite of the lack of ontogenetic evidence. The seem observing that eriocauls differentiation of alternative does not unlikely. state, in first floral sexuality occurs late in development (Stiitzel, especially in view of the finding that corolla fusion much 1985) and might be readily subject to evolutionary a character showing less stability, es- is inodification. Having found that bisexual flowers pecially in the primitive genus Rondonanthus, than (1) same The occur together with staminate flowers in the that of gynoecial modification. paepalanthoid head; ^''Wurdackia'''' has very close sympatric gynoecium furthermore must be thought of as in- (2) relatives with normal pistillate and staminate flow- volving more than one major evolutionary change, an and these related species possess long the modification of the ancestral style into ers; (3) fila- i.e., we mentlike staminodes in the pistillate flowers, ''appendage" (nectary) and the formation of non- agree that the bisexual flowers of ^'''W^urdackia^^ vascularized functional styles at a position rotated 60 are probably secondarily derived. degrees from the position of the original styles. In the same publication, Stiitzel (1985) proposed As far as we know there are no intermediate forms a new three-subfamily system of classification for in existence, and no cases in the Paepalanthoideae the Eriocaulaceae, in which the Syngonanthoideae where the gynoecium has reverted to the primitive {Mesanthemum, Wurdackia, Syngonanthus, and state. The second alternative also not hard to is Philodicc) and the Paepalanthoideae s. str. {Pae- visualize, if we do not assume that ontogeny fault- palanthus, Lachnocaulon Kunth, Tonina Aublet, recapitulates phylogeny. lessly We Rondonanthus) Syngonanthus and seem Rlastocaulon and are agree Ruhl., in- that its allies may dcpendently derived from the Eriocauloideae, con- to constitute a natural group which be usefully from Paepalanthus and taining only Eriocaulon. distinguished its allies. What Rondonanthus (represented only by R. rorai- most necessary present to survey the at is is mae) was evidently placed the Paepalanthoideae subgenera and sections of these two large, poly- in because of the free petals in the pistillate flowers. morphous, and probably unnatural genera in an The union of Wurdackia with Rondonanthus, effort to better characterize the taxa involved and Volume Number & 2 Hensold 445 78, Giulietti Rondonanthus 1991 Revision of to more fully identify and survey characters of abaxially at apex. Flowers 3-merous, either sta- evident taxonomic importance. minate and or staminate and bisexual, the pistillate Moreover, Rondonanthus taken as a prim- two types usually found in diflFerent zones in the if is outgroup Syngonanthus, may same capitulum roraimae) most itive of aid in the or rarely ca- (/?. it assessment of whether Syngonanthus mono- or pitula and plants unisexual. Staminateflowers. Ped- is mm, how polyphyletic, and the latter, to be prop- icels 0,3-^2.5 sometimes equaling or even if it is erly divided to reflect phylogeny. exceeding the flowers in length. Sepals free, elliptic We more feel practical and parsimonious to spatulate, pubescent abaxially at apex. Petals it is to retain for the present Ruhland's two-subfamily free or connate into a 3-lobed tube, and then some- much classification while acknowledging that more times separating after anthesis, glabrous or pilose information needed to build a strong phylogenetic at upper margin. Androphore usually lacking; when is understanding of Eriocaulaceae. present, the petals free and borne at base. Sta- its mens when adnate to corolla at base, except sep- Taxonomy arated from corolla by androphore; anthers dor- and sifixed, white or cream. Pistillate bisexual Rondonanthus Herzog, Feddes Repert. Spec. mm ^^^^^^ 2_2.0 p^j^^^lg q long. Sepals free, Nov. Regni Veg. 29: 210. 1931. type: Ron- ^^^^^ pigmented ^^ ^^^-^^-^ ^^ spatulate, as in the donanthus roraimae Herzog. (Oliver) staminate flowers or lighter, pubescent as in the Wurdackia Mem. New York staminate flowers, expanding and thickening with Mold., Bot. Card. 413. 9: 1957. TYPE: Wurdackia flabelliformis Mold. maturation of the fruit, especially at base. Petals Paepalanthus Bostrychophyllum subg. Ruhl., Pflanzenr. connate and free or distally free at the base, gla- IV. 30: 220. 1903. type: Paepalanthus capilla- brous or pilose at upper margin and abaxially. ceus Klotzsch ex Koern. more Linear staminodes, or rarely developed fully Plants rosulate or caulescent cespitose peren- stamens, present opposite petals, the staminodes nials branching at ground level. Roots creamy white usually about half to twice the height of the ovary to pale brown, porous (aerenchymatous), brittle, and bearing two tiny brownish lobes at the apex, not spongy, usually sparingly branched, the Ovary usually subtended by a ring of long hairs, much branches evidently smaller in diameter than the style much shorter than the style branches; the mm 2-10 main Stems roots. ca. diam., woolly with appendages infundibular, stalked, papillose, insert- fine cream to tan-colored filamentous hairs. Leaves ed on style at or commonly slightly below the level arranged spirally or distichous, filiform to linear or of divergence of the style branches; the style young ligulate, the leaves attaining full width long branches strongly bifid. Fruit a loculicidal capsule, mar- before full length; the bases also with parallel Seeds ellipsoidal, reticulate, the longitudinal stria- gins, but dilated with respect to the blades, at least tions of the testa raised and more conspicuous than somewhat and distally, scarious frequently woolly the transverse. Floral trichomes subacute to sub- with hairs like those of the stem. Pubescence of clavate, with or without granular thickenings of leaves and peduncles when present mostly mix- a the internal wall, ture of filamentous and malpighian hairs, the latter usually "retrorse" (i.e., strongly asymmetric with Key to the Species of Rondonamiius end the larger proximal); capitate hairs lacking. ^^ submerged p,^^^^ aquatics; leaves filiform-se. numerous Inflorescences single to per age class heads taceous, usually surpassing the peduncles; mm ± 3-5 and then usually synchronous. Peduncle sheaths diam.; floral bracts and sepals short- ± hairy capillaceus membranous /?. with respect to the leaves, often lb. Plants terrestrial; leaves linear to ligulate, al- pale and scarious at least toward the upper margin, mm 6-19 ways surpassed by peduncles; heads closed over the young inflorescences and splitting diam.; bracts and sepals densely bearded floral down one side from just below the apex, the opening with long hairs. Leaves oblique with the upper margins remaining minutely 2a. distichous. Stems 3a. leafy to base; flowers bisexual and involute often eventually lacerate. Peduncles and staminate R. flabelliformis mostly 3- or 6-costate, sometimes apparently more. Stems 3b. leafy only at apex; flowers pis- Capitula turbinate to hemispheric at maturity, rare- and staminate R. acopanensis tillate ly subglobose. Involucral bracts, or at least their 2b. Leaves spirally arranged. 4a. Peduncles 3-costate; involucral bracts visible apices, triangular to lanceolate, acute to of upper series acuminate, the acu- subacute, gold to nearly black, or variegated. Re- men R. coidescens colorless ceptacle pilose. Floral bracts present, usually about 4b. Peduncles 6-costate; involucral bracts equaling the flowers, linear to spatulate, pubescent of upper series subacute to acute, black 446 Annals the of Garden Missouri Botanical 4 ^ mm \ » I 50|jm 4 M : j b 4 i + ^ I. » I- -, I' ' r ' I 0.1mm fr L mm OS & Figure Rondonanthus acopanensis (Mold.) Hensold acopanensis {Steyermark 128872 1. Giulietti var. — — — MO). E-G. A. Habit, B. Inflorescence. C, D. Involucral bracts. Staminate flower. E. Flower with bract, past — — anthesis (corolla involute at upper margin). F. Sepal, adaxial view. G. CoroUa tube, opened to show stamens and — — pistillodes. H, I. Pistillate flower. H. Flower with bract, past anthesis. Petal, with hatched lines showing where I. — — ^K. fused with adjacent petal, and staminode adhering at base. J. Gynoecium (fruiting). Seed. L, M. Trichome of bract apex. floral 1 Volume Number & 78, 2 Hensold 447 Giulietti 1991 Rondonanthus Revision of at apex and margins, though some- and usually ciliate hirsute abaxially, the inner times the midvein pale. densely bearded at the apex, at least abaxially and Leaves 5a. sharp-cuspidate to aris- sometimes also adaxially, glabrate with age. Floral tate; petals of staminate flowers commonly bracts exceeding persistently connate and the flowers, linear to ciliate at upper margin R. duidae linear-oblanceolate, acuminate, sometimes with a 5b. Leaves obtuse subacute; to petals long ''whip tip," blackened distally, tufted with of staminate flowers connate ear- hairs abaxially at apex, Staminate flowers. Pedicels but separating by ly anthesis, gla- mm 0.7-1.3 long. Sepals obovate-elliptic to spat- brous R. roraimae ulate, attenuate-acute to long-acuminate, 2.3-3.5(- mm mm 0.6-0.9 4.0) long, wide, white at base, & Rondonanthus acopanensis (Mold.) Hensold blackened distally except for white midvein and comb. nov. Syngonanthus acopa- Giulietti, apex, the apex tufted abaxially with divergent tri- nensis Mold., Phytologia 41. 1948. TYPE: 3: chomes like a bottlebrush. Corolla connate into a Venezuela. Bolivar; Chimanta Massif, Cerro androphore 2.0- 3-lobed tube, the larking, the tube Acopan, ,900 m, Cardona 2280 (holotype, mm 1 3.5(-4.5) long, the lobes triangular-obtuse, US; isotype, NY). Figure 1. upper margin and the abaxial lobe apices ciliate- Plants short-stemmed cespitose perennials. Main pilose, involute and retracting the stamens after mm 0.5-1.2 roots ca. diam., sparingly branched, anthesis. Stamens with filaments adnate to corolla mm cream to pale orange-brown. Stems 1-4 cm long at base, ± included; anthers ca. 0.30.35 mm mm 2-5 at flowering, ca. diam., leafy only near long. Pi5ft7/afe/ou;er5. Pedicels 0.6-0.8 long, apex, but the lower portions covered with tightly Sepals to ovate-elliptic, concave, acuminate, elliptic mm mm persistent flabelliform leaf bases (these making the 3.5-4.0(-4.7) long, 1.0-1.5 wide, cream- mm 5-20 stem appear laterally compressed and colored or pigmented as in staminate flowers though wide), woolly with cream-colored Leaves more weakly apex hairs. dis- usually so, tufted at the as in mm 3-13 cm tichous, long, l-4(-6) wide, the base the staminate flowers, thickening at base at ma- and blade abruptly diff*erentiated, the base linear, turity. Petals connate into a tube except at base, ± dilated with respect to blade, scarious, densely en- the segments oblong, rounded apex, 2.8- at mm mm folded with long cream-colored woolly hairs (often 3.5(4.7) long, 0.6-0.9 wide, densely and black with dirt), persistent, a transverse ab- ciliate at upper margin and frequently also pilose scission layer apparently forming at the juncture abaxially at apex or in submarginal bands. Stam- mm with the blade; the blades linear, subacute to broad- inodes linear, 0.6- .3(2. 5) long. Ovary ringed mm 0- ly rounded or truncate depending on width, car- by hairs at base, 0.4-0.7 long, the style mm mm tilaginous and rigid, glabrous or pubescent with 0.3 long, the appendages ca. 0.4 long, malpighian when young retrorse hairs especially the stalk thick, the papillose portion orange-brown, adaxially, 5-30-veined, the veins of equal size. the style branches diverging slightly above the ap- mm Inflorescences single per age class, the old ones pendages, 1.5-2.3 long, conspicuously bifid, 211 often persistent on stem. Peduncle sheaths Floral trichomes subacute to rounded, lacking cm long, twisted, mostly striate-pubescent with re- granular thickenings of the internal wall. trorse malpighian hairs, the apex acute to rounded This species endemic to the Chimanta Massif is (corresponding in shape to leaf apex), green or in Bolivar, Venezuela, where represented by it is scarious, frequently lacerate with age. Peduncles two sympatric varieties, heretofore recognized as cm I2-45(-60) sometimes They commonly long, 6-costate, ap- species. are about equally col- parently more by division, pubescent with retrorse lected and are distinguishable only by their size malpighian hairs and long filamentous hairs fringing and robustness, as well as possible habitat diff"er- 7.5-12(15) the base of the capitulum. Capitula ences. This sort of variation crops up not uncom- mm diam., turbinate to hemispheric at maturity. monly in eriocauls (as in Pnepalanthus supcrbus 3-5 Involucral bracts in series, the outer narrowly Ruhl. and P. chlorocephalus Silveira in the Serra triangular to lance-linear, subacute to acute, erect do Cipo, Brazil; Hensold, 1988), and possibly is mm to slightly recurved-spreading, 2.5-5.5 long, associated with polyploidy or some other macro- mm 0,6-1,0 wide at base, the inner bracts oblan- mutation. This would imply that the larger form mm 4-7 may ceolate, broadly acuminate, erect, long, be '*polyphyletic," having been derived sev- mm 0.8-1.3 wide, mostly surpassing the head by eral times from the smaller form. (An alternative mm; ca. 0.5-1.5 the outer bracts pale brown to view would be that the smaller form may have been when castaneous, or greenish fresh, the inner black derived from the larger form.) However, because except for the pale tip and midvein; the outer bracts we lack cytological data, and since each type is 448 Annals of the Garden Missouri Botanical 12- about equally widespread and forming uniform pop- Stems, including attached leaf bases, ca. mm mm 11- ulations, we have treated them as varieties. 20 wide. Leaves (2-)2.8-6 wide, ca. mm mm I9(-30)-nerved, rounded to nearly truncate, very 1-2 7.5-10 Leaves wide; heads diam.; la. Huber chartaceous firm-coriaceous (only firmly in involucral bracts mostly ca. 3-seriate Rondonanthus acopanensis var. acopanensis 11522), the margins, especially near the apex, mm 10-14( Leaves (2-)2.8-6 wide; heads 15) lb. Heads 10-14(- often shiny and deep brown-tinged. 4-5- mtn diam.; involucral bracts mostly ca. mm (3-)4-5 15) diam. Involucral bracts in series, seriate mm 3.5-6 0.7- the lowest lance-Hnear, ca. long, Rondonanthus acopanensis var. obtusifolius mm nun 3.5-6.2 Male wide. Floral bracts long. .0 1 mm 0.6- Rondonanthus acopanensis acopanensis flowers with sepals 3.0-3. 5(-4.0) long, var. mm mm 0.9 wide, corolla 3.0-3.5(-4.5) long. 5-10 mm Stems, including attached leaf bases, ca. p^^^j^ fl^^^rs with sepals ca. 3.5(-4.7) long, mm mm Leaves 1-2 5-7(-9)-nerved, mm mm wide. wide, ^^ 1.3- 3.0-3. 5(-4.7) 1.5 wide, corolla firm, green, subacute, the margins not turning shiny |^^ mm brown. Heads 7.5- 10 diam. Involucral bracts mm Phenology. January 2.5-3.5 Collected in flower late 3 the lowest lanceolate, long. in series, mm mid-March. 0.6-0.9 3.5-4.7 nun to wide. Floral bracts long, mm 2.3-3.3 Staminate flowers with sepals long, Habitat and distribution. Locally abundant mm mm 0.6-0.8 and 2.0-2.7 wide, corollas long, among ^^^^ ^^^ savanna and sandstone out- j^ mm 3.5-4.0 flowers with sepals long, Pistillate crops, at 1,800-2,200 m. Venezuela. Bolivar: Chi- mm mm 1.0-1.4 and 2.8-3.2 wide, corollas long. ^^^^^^ ^^^^^^ (Abacapa-, AcopAn-, Apacara-, and Chimanta-tepuis). PIleno logy. Collected in flower mostly from January to March, but also occasionally in June Additional specimens examined. VENEZUELA, boli- & and August. Chimanta S Huber var: Massif, Apacara-tepui, sector, 128428 Steyermark 7061 (LL, NY), Steyermark (LL, Habitat and distribution. Locally abundant ^^^ Steyermark 128519 (LL, MO); Apacara- et at. on wet sandy depressions and banks on shallow SE 8854 Huber tepui, sector, lluher et at. (LL, NY), et open scrub 1,900-2,600 aL 8878 (LL, NY), Huber 11522 (MO), Huber 11524 soils, in the or in forest, NE & Huber Steyermark (NY); Chiinanta-tepui, sector, m. Venezuela. Bolivar: Chimanta Massif (Acopan-, ^, »# 666'^ (LL, NY); Chiiiianta-tepui, central-NE sector, S/(?r- A. pa^cara-, Lhn.nanta-' , (/TL--ruoda1 -lt\ epuis• )\. m- ' iVlurey- ,oo/.o/wmv ^ ' ' t crmark, , . 128090 (NY. M\;Oi)o\; Lhimanta-t*epui,• cen- -^ ^ -' / et at. //ui.rc/ aL (MQ); Abacapa-tepd sector Additional specimens examined. VENEZUELA. ^^^^^ W L^^'"^ * 8620 {NY); Acopan-tepu, N ^ ^^'^'^^ Huber ^^<^^^>^^ bolivar: Chimanta Massif, Apacara-tepui, section, !^'y/^ & & & (Anmri-tepm), Huber Steyermark 7082 (LL 8685 NY, VEN), Huber 8737 Colella (LL, Colella f^^^o'' ^^N), Steyermark 128453 (LL MO). Steyermark ct at. Steyermark (LL, NY); Apicara-tepui, E-central section, ^U NE Huber & Acopan-tepui, secXor e^ 75925 Apicara-tepui, S section, Huber Steyer- '\^[- (F); ;?^!^:t W65 SE SSE '0152 (LL NY), & (NY), sector //.6.r 6970 «^- mark 6964 MO), Huber Steyermark (LL, (LL); Steyermark al 129909 VEN). (LL, ^^^^^^^ et SE Steyermark 75850 Apacara-tepui, section, (F); Chi- manta Murey- Massif (Eru^^^^^^ Huber 11522 specimen included here ^.j^^ ^ is Steyermark 158l8'A (LL, VEN); Chimanta Massif, ^m Clii- 1 r c • some from i ^^th reservation. differs typical var. 128165 It Steyermark manta-tepui, central-NE sector, et al. (LL, MO, NY, VEN); Chimanta-tepui, E sector, vicinity obtusifolius not only in its overall larger dimen- & Cano Huber Steyermark 160 MO), of del Grillo, 7 {LL, sions, but in lacking stem elongation, a distichous & Steyermark Wurdack 804 NY), Steyermark (F, leaf arrangement, and coriaceous-thickened leaves. 128938 128872 MO), Steyermark Chimanta (LL, (LL); Normal {Huber 1524) been var. obtusifolius has 1 Wurdack 34213 NY, VEN), Massif, Churi-tepui, (F, ceii- from same Huber 8934 NY), Huber collected the locality. tral-SE section, <& Colella (LL, km 9275 (MO); 10 SE Chimanta Angasiiua-tepui, of Huber 1688 Massif, {MO). Rondonanthus 1 capillaceus (Klotzscb ex Koern.) & Hensold comb. Paepalan- nov. Ciulietti, Rondonanthus acopanensis var. obtusifolius thus capillaceus Klotzscb ex Koern. C. In & (Mold.) Hensold Giulietti, comb, et stat. nov. Martins, Fl. Bras. 3(1): 415, 53, fig. 11. t. Mem. New Syngonanthus Dupatya obtusifolius Mold., 1863. capillacca (Klotzscb ex York Bot. Card. 9: 410. 1957. TYPE: Vene- Koern.) Kuntze, Revis. Gen. PI. 2: 745. 1891. zuela. Bolivar: Chimanta Massif, central sec- TYPE: Guyana: savannas of Mount Roraima & 1222 1,940 m, 4 Feb. 1955, Steycnnark and Humiridia, April-May, Schotnburgk tion, Wurdack 406 NY; B (holotype, isotypes, F, GH). (holotype, not seen). Figure 2. Volume Number 2 Hensold & 449 78, Giulietti Rondonanthus 1991 Revision of — & Krai 72131 MO). Figure Rondonanthus capillaceus (Klotzsch ex Koern.) Hensold Giulietti A. 2. (/?. — — — Whole Habit. B. Inflorescence. C, D. Involucral bracts, lower and upper. E-I. Staininate flower. E. flower, with — — — J-M. bract.— G. Flower with perianth spread open. H. Sepal, abaxial view. Petal. floral F. Floral bract. I. — — — M. Petal.— flower.- Flower with bract. K. Flower with perianth spread open. L. Sepal, abaxial view. Pistillate J. N, 0. Trichomes of bract apex. floral 450 Annals of the Garden Missouri Botanical mm Paepalanthus 0.3-0.5 hippairichophyllus Herzog, Feddes Re- flowers. Pedicels ca. long. Sepals ob- 193L pert. Spec, Nov. Regni Veg. 29: 208. TYPE: ovate to spatulate, obtuse to rounded, concave, Brazil. Arnapa: Igarape Cre-cru of the Rio Oyapock. mm mm somewhat 1.1-1.4 0.45-0.6 fleshy, long, 21408 Laetzelhurg (holotype, M). wide, cream-colored to pale gray-brown, and ciliate *alanthus capillaceus var. proUferus Gleason, Bull. Torrey Bot. Club 58: 328. 1931. Paepalanthus short-hairy abaxially toward apex. Corolla of 3 capillaceus forma prolifvrus (Gleason) Mold., Phy- separate petals (these separate from earliest visible tologia 44: 384. 1979. type: Venezuela. Amazonas: stage of development), inserted at the base of the Cerro Duida, Central Camp, 4,800 Tate 552 ft., androphore (intercalary prolongation of the floral NY). (holotype, above Paepalanthus New ^^is the calyx), or at variable distances above capillaceus var. spiralis Mold., Mern. York Bot. Card, 9: 279. 1957. type: Guyana. Up- the base, but usually below the insertion of the per Mazaruni River, Karaurieng River, Maipuri Falls, filaments and androphore 0.1-0.5 pistillodes; the & 1,250 m, Maguire Fanshawe 32292 (holotype, mm mm 0.40.7 1.1-1.5 long, the petals long, NY; isotype, M). Note: This taxon, distinguished by mm wide, obtrullate obovate, rounded toward to known strongly spiral-twisted leaves, only from its is the type locality. Though may be valid as a variety, apex and abruptly apiculate, glabrous. Stamens it mm it is here regarded conservatively as a synonym. with filaments 0.7-1.2 long, usually inserted at apex of androphore, though occasionally one or Plants short-stemmed aquatic perennials, the more inserted at base, included, the anthers ca. mm vegetative portions submerged and fully infl 0.15-0.2 long. Pistillateflowers. Pedicels 0,2 mm cences mostly emergent and Main erect. roots 0.4 long. Sepals ovate' concave, to elliptic, mm abundant, 0.5-1.0 diam.,much.branched, mm mm ca. acute to obtuse, 1.2-1.5 long, 0.5-0.8 cm gray-brown, contorted. Stems up and to 7 long ^ide, colored as in staminate flowers, short-ciliate mm 15 diam., leafy only near the apex, but the from middle apex and to short-hairy abaxially in lower portions covered by and persistent roots leaf this region, coriaceous-thickening at maturity, es- bases, woolly with brownish Leaves hairs. spirally pecially at base and sometimes throughout. Petals arranged, green (2.5)7-30 cm pale to blackish, free, spatulate, usually acute or apiculate, 1.2-1.6 0.2-0.4 inm mm mm long, wide, the base slightly dilated, 0.40.6 long, wide, glabrous. Staminodes ephemerally woolly, coarsely filiform-setaceous, firm, linear to somewhat broad and nearly scalelike, sometimes mm corkscrew-twisted, the acute but tips 0.2-0.6(-l Ovary .4) long. with a few seat- usually broken glabrous (though often with sand mm off, 0.4-0.7 tered hairs at base, long, the style particles and algae adhering in dried material), 0.1-0.4 mm mm appendages 0.4-0.5 long, the traversed by a single central vein surrounded by long, the stalk thick, the papillose portion orange- mesophyll (Ruhland, 1903). Inflorescences nu- brown, darker than the the branches stalk, style merous, synchronous, commonly numbering 10 or diverging at the same level as the appendages, 0.7- more. Peduncle 3-5 cm mm sheaths long, strongly 0.9 long, bifid half to nearly their length, all twisted, glabrous, the apex acuminate, scarious, Floral trichomes clavate and thick-walled with dense cm often lacerate. Peduncles (5.5)9.0-20.0 long, granular thickening of the internal wall. 3-costate or apparently 6-costate by collapse of main 3- Phenology. the centers of the ribs, glabrous. Capitula Collected in flower year-round. mm 5 diam., depressed-turbinate depressed- to and Habitat Adhering distribution. rocks to hemispheric maturity, sometimes at vegetatively streambeds 100-1,600 of in rapidly flowing water, from proliferating the center. Involucral bracts in m. Venezuela. Amazonas: Cerros Duida and Mara- ca. 4 series, both the outer and inner narrowly h south Gran to Neblina; Bolivar: Sabana. commonly triangular, subacute to obtuse, erect, Guyana: Upper Mazaruni and River Kaieleur Pla- mm mm 1.1-1.4 0.50.7 the outer ca. long, wide, Amazonas: Amapa: teau. Brazil. Serra Araca; Rio mm mm the inner 1.9-2.1 0.50.7 long, wide, Oyapock (Oiapoque). about equaling the head; pale gold-brown to all castaneous or nearly black, fleshy or coriaceous- Additional specimens examined. VENEZUELA. amazonas: Cerro Duida, Farinas 420 (NY), Ma- thickened especially toward the base, the margins et al. ± .u.^a^;^. S^^^^' <^ ^^m^^S^^^u^^^^ec ^2:9^1i5^3^ ^(iM^iOvy,, iN^Yx);,, Mifaiguu^uiurte, et uail.. --^^---^^ i^i of at least the inner bracts --^ Pneeilliuucciida sshnrreeddddimngg ^g^jj 751-82 (NY); Cerro Mara- ^p^y); Tillett et al. . wi.t,h age; gl, a,brous or t,he mner bracts short-cdiate & huaca, Steyermark Hoist 130470 (MO); Cerro Ara- & & along distal margins. Floral bracts shorter than or camuni, Liesner Carnevali 22308 {}A0\ Wurdack Adderley 43602 about equaling the (NY); base of Cerro Neblina, Davidse flowers, spatulate, obtuse, strongly k,ee1led and somewhat f ^1^"^' cucullate apex, at t.^'TJ^?'^^ ^'''^^f.^?:!^.A'^?^^^"''^ Liesner 15737 (MO); Liesner 16653 (MO). BOLiVAR: , cream-colored to pale b, rown, t,he apex h, i. rsute Canmma, 7745 ESE 5rA;z^^ (MO); Rio Yune, of Churi- abaxially with short subclavate Staminate hairs. tepui, Ruber 9764 (MO); headwaters of Rio Venamo,

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