A new combination in Elaphandra and a new variety of Eleutheranthera ruderalis (Compositae: Heliantheae: Ecliptinae) from Andean South America.
Se presenta la nueva combinacion Elaphandra patentipilis para una especie endemica de Colombia, anteriormente ubicada dentro de Aspilia. Se propone una clave para las especies del grupo de E. lehmannii, y E. quinquenervis se trata como una especie endemica de Colombia. Aspilia wedelioides se supone como un sinonimo de E. verbesinoides. Elaphandra se trata con 14 especies. Se describe Eleutheranthera ruderalis var. radiata de Venezuela y se discuten sus relaciones. Aspilia triplinervia se presume un sinonimo de Eleutheranthera tenella.
Palabras claves: Elaphandra, Eleutheranthera, Compositae, Asteraceae, Heliantheae, Ecliptinae, Colombia, Venezuela, Taxonomia.
The new combination Elaphandra patentipilis is made for an endemic Colombian species formerly placed in Aspilia. A key to the species in the Elaphandra lehmannii group is given, and E. quinquenervis is taken to be endemic to Colombia. Aspilia wedelioides is presumed to be a synonym of Elaphandra verbesinoides. Elaphandra is taken as currently containing 14 species. Eleutheranthera ruderalis var. radiata from Venezuela is described and its relationships are discussed. Aspilia triplinervia is presumed to be a synonym of Eleutheranthera tenella.
Key words: Elaphandra, Eleutheranthera, Compositae, Asteraceae, Heliantheae, Ecliptinae, Colombia, Venezuela, Taxonomy.
Ecliptinae Lessing, a subtribe of Compositae: Heliantheae has been delimited by Robinson (1981), although most of its genera were more recently treated under the name Verbesininae by Karis & Ryding (1994), who did not subtribally place Eclipta L. Generic additions and adjustments of Robinson (1984a, 1984b, 1992, and literature cited therein), Strother (1991 and literature cited therein), Pruski (1996, 1999, and literature cited therein), and Panero et al. (1999) are accepted as emendations to the group as defined by Robinson (1981). The subtribe Verbesininae, treated as a synonym of the Ecliptinae by Robinson, Strother, and Pruski, was recently resurrected by Panero et al. (1999) and includes only Verbesina L. and three small Mexican and Central American genera.
Here, a new combination is made in Elaphandra Strother for a sterile-rayed species formerly placed in Aspilia Thouars, and a key to species of the group centering about E. lehmannii (Hieron.) Pruski is provided. Also, a new variety with radiate capitula of pantropical Eleutheranthera ruderalis (Sw.) Sch. Bip. is described from Venezuela.
The following innovations in Elaphandra and Eleutheranthera Poit. ex Bosc are the result of both floristic work (e.g. Pruski, 1997) and routine identifications for species from South America.
Elaphandra Strother, Syst. Bot. Monogr. 33: 17. 1991.
Elaphandra was described by Strother (1991) as unispecific, and is typified by E. bicornis Strother, a discoid species from Panama. Strother (in sched.) annotated several northern Andean species as congeneric with E. bicornis, but he did not formally transfer these species to Elaphandra. More recently, several South American species have been transferred to (as earlier suggested by Strother) or newly described in Elaphandra (Robinson, 1992, 1994; Badillo, 1994; Pruski, 1996). Pruski (1996) reduced E. lucidula (S.F. Blake) H. Rob. to the synonymy of E. ulei (Hieron.) H. Rob. and also Pruski reduced E. sucrensis (Aristeg.) V.M. Badillo to the synonymy of E. verbesinoides (DC.) H. Rob. Elaphandra verbesinoides also presumably includes as synonymous the once-collected Aspilia wedelioides (DC.) S. F. Blake from Tobago. Most of the newer additions to Elaphandra have radiate capitula and were formerly placed in Aspilia.
Elaphandra is characterized by the technical characters of opposite, eglandular leaves; by sterile ray florets (when present); by ovate, eglandular anther appendages, these commonly black though sometimes tan within or distally; by erect or laxly recurved (not strongly coiled), papillose style branches; and by weakly pappose, rostrate (necked) cypselae, these without elaiosomes and without obvious carpopodia. Pruski (1996) cited Elaphandra as containing 13 species. The transfer of Aspilia patentipilis S.F. Blake to Elaphandra proposed here increases to 14 the number of species I recognize in Elaphandra.
Pruski (1996) cited both Aspilia patentipilis and Elaphandra quinquenervis (S.F. Blake) H. Rob. as closely related to E. lehmannii and perhaps not distinct from it. However, further study of the types, type photographs, and other specimens available to me of the species of the E. lehmannii group (in Group I of Robinson, 1992) show that the two former species, although clearly closely related to E. lehmannii, are distinct from it. In floral features all specimens of this species group are similar, and the species can be distinguished most readily by differences in indumentum of vegetative structures.
Lectotypification of A. lehmannii Hieron. was deferred by Pruski (1996) and similarly is delayed here until type material is found and studied. Unfortunately, syntype material of A. lehmannii (Lehmann 3282) was not found at BM (fide R, Vickery), nor at K (fide D. J. Nicholas Hind and S. J. Owens), and to date I have been able to examine only a photograph (NY!, US!) of the destroyed Berlin syntype of Lehmann 3282. Elaphandra lehmannii occurs in Andean Colombia and Ecuador, and is the most widespread of the three species of this group. It was described by Hieronymus as having stems "hirto-pilosis," thus differing from the species in the group with either strigose stems or long-patent, pilose stems.
I have been fortunate to have recently examined the holotype of Aspilia quinquenervis (K!), which has short-strigose hairs on the stems, leaf blades, and phyllaries; petioles and peduncles strigose to hirsute; and elliptic-lanceolate outer phyllaries. By strigose stems, phyllaries, and leaf blades this species (E. quinquenervis), a Colombian endemic (known from Antioquia, Caldas, Cauca, Tolima, Valle, and from along the Choco-Valle border), proves to be distinct from the two other species of the E. lehmannii group. The third species in the E. lehmannii group (Aspilia patentipilis) is characterized by its vegetative structures with long-patent, pilose hairs, and has yet to be transferred to Elaphandra. The needed combination for this species in Elaphandra is thus provided below.
Elaphandra patentipilis (S.F. Blake) Pruski & G. Mendez, comb. nov. Basionym; Aspilia patentipilis S.F. Blake, Contr. U.S. Natl. Herb. 22: 617. 1924. Wedelia patentipilis (S.F. Blake) B.L. Turner, Phytologia 72: 393. 1992. Fig. 1
Type. Colombia, TOLIMA: La Trinidad, Libano, 1100-1400 m, 21-25 Dec 1917, Pennell 3229 (holotype: NY!; isotype: US!).
Comments: The E. lehmannii group as circumscribed by Pruski (1996) includes two elements (A. patentipilis and E. quinquenervis), which are here removed from the provisional synonymy of E. lehmannii. This results in a better circumscription E. lehmannii, which is nevertheless still broadly defined and variable. One named element that I feel is best removed from E. lehmannii is the element including the type of A. patentipilis, namely the entity with long-patent, pilose hairs and long, elliptic-lanceolate outer phyllaries. This species is here formally transferred to Elaphandra. The type of A. patentipilis is the most extreme example of this form in that its hairs and outer phyllaries are the longest in the E. lehmannii group.
Monographic studies are needed to precisely define the taxonomic limits of E. patentipilis, which is an herb or vine and is a moderately common Colombian endemic. Plants provisionally referred here to E. patentipilis occur on all three Andean Cordilleras of Colombia in Antioquia, Boyaca, Cauca, Narino, Norte de Santander, Tolima, and Valle. Several specimens seem intermediate between E. lehmannii and E. patentipilis, but I nevertheless believe it is useful to recognize the two as distinct species. Elaphandra patentipilis is characterized by having long-patent, pilose stems, leaves, and phyllaries; and longer, elliptic-lanceolate outer phyllaries. It is thus distinguished from E. lehmannii, which commonly has substrigose, hispid, or sericeous leaves and phyllaries; stems shortly pilose or hispid; and shorter, elliptic-ovate outer phyllaries.
I am happy to validate this transfer with Gina Paola Mendez R. (of the COL herbarium), who is working on Colombian members of the taxonomically difficult subtribe Ecliptinae.
Key to the species centering about Elaphandra lehmannii 1. Stems with short, strigose hairs; Colombia E. quinquenervis (S.F. Blake) H. Rob. 1. Stem hairs not short-strigose. 2. Stems, leaves and phyllaries without long-patent, pilose hairs, indumentum type various, occasionally shortly pilose or sericeous; outer phyllaries often elliptic-ovate and about as longer as or shorter than the inner phyllaries; Colombia and Ecuador E. lehmannii (Hieron.) Pruski 2. Stems, leaves, and phyllaries commonly, with long-patent, pilose hairs, phyllaries less commonly sericeous; outer phyllaries often elliptic-lanceolate and longer than the inner phyllaries; Colombia E. patentipilis (S.F. Blake) Pruski & G. Mendez
Pantropical Eleutheranthera ruderalis is uniformly discoid throughout most of its range. However, in the Venezuelan coastal ranges plants with radiate capitula otherwise matching the typical form with discoid capitula of this species are occasionally found. The collections with radiate capitula are described here as a new variety of E. ruderalis as follows.
Eleutheranthera ruderalis (Sw.) Sch.-Bip. var. radiata Pruski, var. nov.
Type. Venezuela, DISTRITO FEDERAL: Hacienda Chichiriviche, ca. 300 m, Jul 1958, L. Aristeguieta 3233 (holotype: NY!: isotype: VEN n.v.).
A var. typica affinis, sed capitulis radiatis diversa.
Herbae annuae vel subperennes; folia opposita petiolata; lamina elliptica vel ovata 2.7-8.4 X 0.9-4.2 cm chartacea strigosa et glandulosa; capitulescentia paucicapitulata; capitula radiata, flosculus 9-12; involucrum campanulatum; receptaculum paleaceum; flosculi radiati steriles ca. 4, limbo ca. 5 X 3 mm glandulifero; flosculi disci hermaphroditi 5-8, corolla ca. 2.8 mm longa, lobis ca. 0.3-0.4 mm longis; antherae nigrae; stylus breviexsertus, ramis lineari-lanceolatis, ca. 0.5 mm longis; achaenia nigra, obovoidea, 3-3.5 mm longa, tuberculata, epapposa.
Erect herbaceous annuals to short-lived perennials, more than 30 cm tall. Stems moderately branched, striate, pubescent. Leaves simple, opposite, petiolate; petiole (0.3) 0.7-1.8 cm long; blade elliptic to ovate, 2.7-8.4 X 0.9-4.2 cm, chartaceous, weakly serrulate, 3-veined from near base, acute at apex, decurrent at base, strigose to weakly so, the lower surface also glandular, sometimes obscurely so. Capitulescence axillary, in few-headed cymes in the uppermost nodes or capitula solitary; peduncles short, commonly 2-2.5 cm long. Capitula radiate, 9-12-flowered; involucre campanulate; phyllaries weakly imbricate, 2-3-seriate, subequal or outer series commonly longer, the outer series foliar, elliptic to pyriform, 6.5-8 X 2.6-4 mm, thinly chartaceous, hirsute, the inner ones foliar, elliptic-lanceolate, hirsutulous, usually slightly smaller than the outer series; receptacle minute, paleate, the pales lanceolate, conduplicate, ca. 5.5 mm long, apically attenuate to apiculate, pilose to glabrate. Ray florets sterile, ca. 4; corolla yellowish, the tube ca. 0.7 mm long, glabrous, the limb ca. 5 X 3 mm, 5-nerved, abaxially glandular. Disk florets bisexual, 5-8; corolla tubular-funnelform, ca. 2.8 mm long, pale yellow, shortly 5-lobed, glabrous except for lobes hispid within, the tube ca. 0.4 mm long, the throat slightly broader, ca. 2 mm long, the lobes deltoid, 0.3-0.4 mm long, strongly hispid within and marginally; anthers very weakly apically exserted, black, 1.2-1.5 mm long, the appendages broad, often weakly glandular; style weakly exserted, the style branches erect or nearly so, linear-lanceolate, ca. 0.5 mm long, attenuate at apex, strongly pilose, with weakly paired marginal stigmatic lines. Ray ovaries sterile, narrowly cylindrical, apically setose. Disk cypselae black, obovoid, often angled, 3-3.5 mm long, tuberculate, puberulent at apex, otherwise glabrous, carpopodia indistinct, elaiosomes absent; pappus absent or less commonly a ciliate ring on top of the short neck of the cypsela.
Paratype: VENEZUELA, YARACUY: Distr. San Felipe: Municipio Veroes: Fila montanosa 5 kms S de Bella Vista, 10[degrees]23'N, 68[degrees]24'W, 200-450 m, 11 Jul 1973, G. Agostini, G. Morillo & B. Morillo 1737 (US!, VEN n.v.).
Comments: Eleutheranthera had been treated traditionally as a unispecific discoid genus until Robinson (1992) transferred to it a second species with tuberculate cypselae, the radiate-headed Gymnolomia tenella H.B.K. With the present description of E. ruderalis var. radiata, the genus now contains three taxa, two of these with radiate capitula. The discoid and widespread pantropical E. ruderalis var. ruderalis remains the most common taxon of the genus, however. The sterile-rayed entities are each much less widespread, one is endemic to Colombia, and the other is endemic to Venezuela. Eleutheranthera tenella (H.B.K.) H. Rob. is regionally common in the Colombian Andes, especially near Cali, at elevations of about 950-1900 m, but occasionally this species may be found at sea level on the Pacific coast. Eleutheranthera tenella presumably includes Aspilia triplinervia (H.B.K.) S.F. Blake as synonymous, although not cited as such by Robinson (1992). The second variety of E. ruderalis, described here, is endemic to the northeastern-most Andes, the Venezuela coastal range near Caracas, where it flowers in July and has been collected from 200-450 m elevation.
The new variety is treated as an variety of the otherwise similar E. ruderalis, from which it differs most noticeably by radiate (vs. discoid) capitula. By radiate capitula, the new variety, however, resembles E. tenella. However, E. tenella is a reduced herb with small leaves, thus differing from E. ruderalis var. radiata. Moreover, E. tenella differs from E. ruderalis var. radiata by terminal (vs. axillary) capitula that are about 35- (vs. 9-12-) flowered, these long(vs. short-) pedunculate and with outer phyllaries lanceolate (vs. elliptic to pyriform).
I thank Santiago Diaz, Dan Nicolson, Gina Paola Mendez R., Harold Robinson, and Gisela Sancho for reviewing an earlier draft of this paper; R. Vickery (BM), D. J. Nicholas Hind (K), and S. J. Owens (K) for searching for the type of Aspilia lehmannii in their respective herbaria; and Susan K. Hunter, Smithsonian Institution Botany Department Staff Photographer, for her photograph of the isotype of Aspilia patentipilis.
Badillo, V. M. 1994 , Enumeracion de las Compuestas (Asteraceae) de Venezuela. Revista Fac. Agron. (Maracay) 45: 1-191.
Karis, P. O. & O. Ryding. 1994. Chapter 22, Tribe Heliantheae. Pp. 559-624. In: Bremer, K. Asteraceae: Ciadistics & Classification. Timber Press, Portland, Oregon.
Panero, J. L., R. K. Jansen & J. A. Clevinger. 1999. Phylogenetic relationships of subtribe Ecliptinae (Asteraceae: Heliantheae) based on chloroplast DNA restriction site data. Amer. J. Bot. 86: 413-427.
Pruski, J. F. 1996. Compositae of the Guayana Highland-XI. Tuberculocarpus gen. nov. and some other Ecliptinae (Heliantheae). Novon 6: 404-418.
--. 1997. Asteraceae. Pages 177-393. In: J. A. Steyermark et al. (eds.), Flora of the Venezuelan Guayana. Vol. 3. Araliaceae-Cactaceae. Missouri Botanical Garden, St. Louis.
--. 1999. Novelties in Angelphytum and Oyedaea (Compositae: Heliantheae: Ecliptinae) from South America. Comp. Newsl. 34: 1-8.
Robinson, H. 1981. A revision of the tribal and subtribal limits of the Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51: i-iv, 1-102.
--. 1984a. Studies in the Heliantheae (Asteraceae). XXXI. Additions to the genus Dimemstemma. Proc. Biol. Soc. Wash. 97: 618-626.
--. 1984b. Studies in the Heliantheae (Asteraceae). XXXIV. Redelimitation of the genus Angelphytum. Proc. Biol. Soc. Wash. 97: 961-969.
--. 1992. New combinations in Elaphandra Strother (Ecliptinae-Heliantheae-Asteraceae). Phytologia 72: 144-151.
--. 1994. A new species of Oblivia and a new combination in Elaphandra from Ecuador (Ecliptinae-Heliantheae-Asteraceae). Phytologia 76: 24-26.
Strother, J. L. 1991. Taxonomy of Complaya, Elaphandra, Iogeton, Jefea, Wamalchitamia, Wedelia, Zexmenia, and Zyzyxia (Compositae-Heliantheae-Ecliptinae). Syst. Bot. Monogr. 33: 1-111.
John F. Pruski (1)
(1) United States National Herbarium Department of Botany, MRC-166 Smithsonian Institution Washington, DC 0560-0166, U.S.A.
Caption: Figure 1. Photograph of isotype of Aspilia patentipilis S.F. Blake (Pennell 3229, US).
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|Author:||Pruski, John F.|
|Publication:||Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales|
|Date:||Sep 1, 2001|
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