Table Of ContentCompositae of the Guayana Highland—XI. Tuberculocarpus
gen. nov. and Some Other Ecliptinae (Heliantheae)
John F. Pruski
United States National Herbarium, Department of Botany, MRC-166, Smithsonian Institution,
Washington, D.C. 20560, U.S.A.
ABSTRACT. A new genus from Venezuela, Tuber¬ The following novelties and adjustments involve
culocarpus, is described and illustrated, and the taxa primarily from the Guayana Highland of north¬
combination T. ruber is made. Thelechitonia and eastern South America. These novelties are proposed
Complaya are placed in synonymy of the formerly prior to the appearance of many of them in the Com¬
unispecific Sphagneticola, and a lectotype is chosen positae of the Guayana Highland, the Guide to the
for the generitype S. ulei and for Verbesina calen- Vascular Plants of Central French Guiana, the Flora
dulacea. Sphagneticola includes cultivated Wedelia of St. John, U.S.V.I., and the Flora of the Venezuelan
trilobata, and the combinations S. brachycarpa, S. Guayana. A map (Fig. 1) is given for the six Gua¬
calendulacea, S. gracilis, and S. trilobata are made. yana Highland endemics proposed or transferred
Wulffia is a taxonomic synonym of Tilesia, and com¬ here. The genera treated herein are restricted to the
binations in Tilesia are proposed for the three spe¬ New World, with the exception of Sphagneticola 0.
cies and one variety formerly placed in Wulffia: Ti¬ Hoffmann. Generic restructuring proposed here in¬
lesia baccata, T. macrocephala, T. rubens, and T. cludes reduction of Complaya Strother and Thele¬
baccata var. discoidea. The basionym of the most chitonia Cuatrecasas to synonymy of Sphagneticola,
widespread species of Tilesia (Coreopsis baccata) has which necessitates the transfer of the species of Wed¬
previously been attributed to Linnaeus filius, but was elia trilobata (L.) Hitchcock group to Sphagneticola:
published earlier by Linnaeus. Elaphandra moriana the reduction of Wulffia Necker ex Cassini to syn¬
from French Guiana is described, and the combi¬ onymy of Tilesia G. Meyer, including the transfer of
nation Elaphandra lehmannii is made. Oyedaea wur- Wulffia baccata (L.) Kuntze to Tilesia: and the de¬
dackii is described from Venezuela and is illustrated, scription of Tuberculocarpus Pruski, a generic seg¬
and the combination Oyedaea tepuiana is proposed. regate of the Aspilia Thouars-Wedelia Jacquin alli¬
The combination Verbesina ligulata is provided, and ance.
V. guianensis is lectotypified and is an earlier name
for V. schomburgkii. Elaphandra sucrensis is reduced Elaphandra
to synonymy of E. verbesinoides, E. lucidula is re¬
Strother (1991) recently proposed the segregate
duced to E. ulei, Oyedaea blakeana is reduced to 0.
Elaphandra from the Aspilia-Wedelia alliance. The
scaberrima, and Wulffia trujilloi is reduced to Tilesia
limits of formerly unispecific Elaphandra were ex¬
macrocephala.
panded by Robinson (1992; 1994) to include 11
additional species (only 10 of which are recognized
The Ecliptinae Lessing (including Verbesininae here), many of which were suggested by Strother
Bentham) of the Heliantheae (Compositae) are dif¬ (1991; in sched.) as belonging to Elaphandra. The
ficult taxonomically, both at the generic and specific combination proposed here and the description of
levels. The traditional generic and subtribal classi¬ Elaphandra moriana Pruski raises to 13 the num¬
fications of this group by Bentham (1873) and Hoff¬ ber of species in the genus.
mann (1890—1894) have been much revised in the The species of Elaphandra have sterile ray flo¬
recent past. The classification used here is that of rets (when present) and would previously have fit
Robinson (1981), but recent modifications by Rob¬ into the concept of Aspilia, now a synonym of Wed¬
inson (1984a, 1992, and literature cited therein) and elia. Elaphandra was not reported (nor was Aspilia)
Strother (1991, and literature cited therein) are ac¬ in the Guianas by Funk (1991), and E. moriana is
cepted. Features most commonly employed in clas¬ the first species of the genus known in the Guyana
sifications include anther, style, cypsela, and pappus Highland. The type of E. moriana was listed by
characters, and fertility of the ray florets. Karis and Cremers and Hoff (1995) as W. fruticosa Jacquin,
Ryding (1994) used the name Verbesininae for much which is unknown in French Guiana. Elaphandra
of the Ecliptinae. is diagnosed by eglandular leaves, by rays sterile
Novon 6: 404-418. 1996.
Volume 6, Number 4 Pruski 405
1996 Guayana Highland Compositae
when present (one species is discoid), by ovate Scandent shrub to 2 m tall; stems weak, subterete,
eglandular anther appendages, these commonly strigose, the intemodes 2.5-7(-ll) cm long, usually
abaxially black though sometimes tan within or dis- much shorter than the subtending leaves. Leaves
tally, by erect or laxly recurved (not strongly coiled) simple, opposite, petiolate; petioles 0.5—1.5 cm long,
papillose style branches, by rostrate (necked) cyp- strigose-hispidulous, thin, not obviously canalicu¬
selas that have a reduced or aristate pappus, and late; blades elliptic-lanceolate to elliptic, 4—12 X
by lack of obvious carpopodia and elaiosomes. 0.8-5.2 cm, chartaceous, weakly triplinerved from
well above base, basally cuneate or sometimes at¬
Elaphandra lehmannii (Hieronymus) Pruski, tenuate, the margins serrulate, apically acute to acu¬
comb. nov. Basionym: Aspilia lehmannii Hi¬ minate, the surfaces eglandular, without black dots,
eronymus, Bot. Jahrb. Syst. 28: 605. 1901. the adaxial surfaces green, scabrous, the abaxial sur¬
Wedelia lehmannii (Hieronymus) B. L. Turner, faces pale green, hispidulous, especially so on larger
Phytologia 72: 393. 1992. SYNTYPES: Co¬ veins. Capitulescences cymose or sometimes mono-
lombia. Risaralda: in silvis prope Arrayanal ad cephalous, terminal or axillary from the most distal
fluvium Rfo Risaralda, 1400-1800 m, 24 Oct. node, of 1-3 long-pedunculate capitula held slightly
1883, Lehmann 3282 (B destroyed [photo¬ above the subtending leaves; peduncles erect, 2-5
graph, NY, US]); locality not indicated, Triana cm long, densely hispid-strigose. Capitula radiate,
1363 (not seen) and Triana 1365 (not seen). 30-34-flowered; involucres campanulate, 6-9 X 6-
Lectotypification is deferred, because not all 8 mm; phyllaries 10-13, in ca. 2 series, imbricate,
syntypes were studied. subequal, herbaceous or inner ones scarious at base,
oblong, 5-9 X 2—2.5 mm, strigose, not black-
Distribution. Andean Colombia and presum¬ streaked, the apices broadly acute to rounded; re¬
ably extending southward into Ecuador. ceptacles flat, 2.5-3 mm diam., paleate, the pales
This species is one of four placed in Elaphandra conduplicate, 5-6.5 X 1—1.4 mm, glabrous, strami¬
Group I of Robinson (1992), and possibly includes neous, the apices acuminate. Ray florets (immature)
the very similar Aspilia patentipilis S. F. Blake. 5 or 6, sterile, lacking styles; corollas yellow, 5.5—
Specimens in US determined by Blake as A. paten¬ 6.5 mm long, the tubes ca. 1 mm long, glabrous, the
tipilis tend to have hairier stems, fewer capitula, limbs 4.5-5.5 X 2.5-3 mm, eglandular, shortly bi-
these with very long outer phyllaries, and longer lobed at the apices, 6-8-veined with 2 veins larger
peduncles. If these distinctions are consistently than the others, these abaxially puberulent distally.
found in future collections, then a combination in Disk florets 24—29, bisexual; corollas actinomorphic,
Elaphandra may be needed for A. patentipilis. It yellow, not black-streaked, tubular-funnelform, ca.
seems best to view E. lehmannii in a broad sense, 4.2 mm long, the tubes ca. 1.1 mm long, glabrous,
presently including A. patentipilis. Furthermore, E. the throats ca. 2.4 mm long, glabrous, the lobes 0.7-
quinquinervis (S. F. Blake) H. Robinson, which has 0.9 mm long, long-triangular, weakly puberulent; an¬
5-veined leaves, is very closely related to E. leh¬ thers weakly exserted, ca. 2.3 mm long, the thecae
mannii and may not prove to be distinct. If the two black, the apical appendages ovate, black at base,
above species are placed in synonymy of E. leh¬ tan distally, eglandular, the filaments 1.2 mm long;
mannii the distribution of E. lehmannii would style branches ca. 1.5 mm long, papillose, slightly
range from the eastern to the western cordilleras of spreading, with paired stigmatic lines, the apices
Colombia and southward into Ecuador. narrowed. Ray ovaries sterile, 2.8—3 mm long, ob-
conic, 3-angled, ciliate on the ribs, 3-aristate, the
Elaphandra moriana Pruski, sp. nov. TYPE: aristae 1.8-2.4 mm long, subequal to unequal. Disk
French Guiana. Inselberg granitique h l’ex- cypselas 4-4.2 X 1.5-2 mm, obovate, slightly com¬
tremits nordouest des Monts de la Trinity, 4 pressed, black, weakly puberulent on necks and
Aug. 1981, J. de Granville 4741 (holotype, US; shoulders, lacking elaiosomes and carpopodia; pap¬
isotype, US). Figure 1. pus 2^1-aristate, the aristae subequal to unequal,
1.5-3.2 mm long, fragile and easily knocked off,
Suffmtex scandens; caules strigosi; folia simplicia opposita
petiolata, lamina 4—12 X 0.8-5.2 cm chartacea eglandulosa; arising from a low fimbriate crown ca. 0.6 mm tall.
capitulescentia cymosa vel monocephala; capitula radiata pa-
leacea, flosculis 30-34; phyllaria strigosa herbacea basi sca- Distribution and ecology. This species is known
riosa; flosculi radiati 5 vel 6 steriles, corolla luteola 5.5-6.5 only from the type collection made on a granitic
mm longa; flosculi disci 24-29 hermaphroditi, corolla luteola inselberg (laja) in the northwestern sector of Monts
ca. 4.2 mm longa: aruherae nigrae, appendicibus apicalibus
de la Trinity in French Guiana. It was collected in
stramineis eglandulosis; achaenia rostrata nigra, aristis 2—4
fragilibus caducis 1.5-3.2 mm longis. flower in August.
406 Novon
Elaphandra moriana
Oyedaea tepuiana
Oyedaea wurdackii
Tilesia rubens
Tuberculocarpus ruber
Verbesina ligulata
Figure 1. Distributions of Elaphandra moriana Pruski, Oyedaea tepuiana (V. Badillo) Pruski, Oyedaea wurdackii
Pruski, Tilesia rubens (Alexander) Pruski, Tuberculocarpus ruber (Aristeguieta) Pruski, and Verbesina ligulata (Maguire
& Wurdack) Pruski.
This is the first Elaphandra known in the Gua- Elaphandra verbesinoides (DC.) H. Robinson, Phy-
yana Highland. Elaphandra verbesinoides and E. ulei tologia 72: 148. 1992. Gymnopsis verbesinoides
are the nearest geographical relatives. They differ DC., Prodr. 5: 561. 1836. Aspilia verbesinoides
from E. moriana by membranous leaves that are (DC.) S. F. Blake, Proc. Biol. Soc. Wash. 34:120.
black-dotted, and by black-streaked phyllaries or co¬ 1921. TYPE: Trinidad. Without date, J. Lockart
rollas, which place them in Group II of Robinson s.n. (holotype, G-DC not seen [photograph, US;
(1992). Elaphandra moriana has chartaceous non¬ IDC microfiche 800. 941.111.6]; isotypes, GH not
black-dotted leaves, radiate capitula, non-hispid seen [photograph, US], K not seen).
pales that are apically acuminate, and non-black-
streaked phyllaries and corollas, which place it as Aspilia sucrensis Aristeguieta, Acta Bot. Venez. 1(2): 80.
the fourth species referable to Group I of Robinson 1966. Syn. nov. Elaphandra sucrensis (Aristeguieta)
V, Badillo, Revista Fac. Agron. (Maracay) 45: 95.
(1992). It differs from the three Andean species of 1994 [1995]. TYPE: Venezuela. Sucre: Peninsula de
this group by smaller capitula, a well-developed Paria, Cerro Patao, norte de Puerto de Hierro, nord-
aristate pappus, and distally tan anther appendages. este de Guiria, fila seca en la cumbre, 875 m, 25—
I am happy to name this new species for Scott 26 July 1962, Steyermark & Agostini 91337 (holo¬
type, VEN; isotypes, NY, US—2).
Mori, Director of the Institute of Systematic Botany
of The New York Botanical Garden. Scott pointed Distribution. Venezuela (Sucre) and Trinidad.
my eyes eastward in the Guayana Highland through Elaphandra sucrensis is added to the synonymy
his fine collections made for his forthcoming Guide of E. verbesinoides given by Robinson (1992). Ela¬
to the Vascular Plants of Central French Guiana. phandra verbesinoides is closely related to Amazo¬
Elaphandra moriana is named in conjunction with nian (Bolivian and Brazilian) E. ulei [E. lucidula
my treatment of the Compositae in this flora. (S. F. Blake) H. Robinson, syn. nov.], the latter dif-
▲
Volume 6, Number 4 Pruski 407
1996 Guayana Highland Compositae
fering by shorter petioles, often by more rounded 5a. Leaves tomentose abaxially; Mona-
leaf bases, and by fewer capitula, these frequently gas.O. verbesinoides var.
hypomalaca Steyermark
on longer peduncles. It is possible that E. verbesi- 5b. Leaves pubemlent abaxially.
noides or the related E. ulei will eventually be 6a. Ray limbs 10-13 X 2 1.5 mm;
found in the Venezuelan Guayana or the Guianas. Amazonas and Bolfvar (includes O.
Elaphandra verbesinoides and E. ulei belong to Ela- blakeana Steyermark, syn. nov.) . .
phandra Group II of Robinson (1992), which con¬ . . 0. scaberrima (Bentham) S. F. Blake
6b. Ray limbs 18—20 X 5—5.5 mm; An-
tains four species characterized by black-spotted zoategui and Sucre .
leaves and black-streaked phyllaries or corollas. .0. maculata S. F'. Blake
Oyedaea Oyedaea tepuiana (V. Badillo) Pruski, comb. nov.
Basionym: Zexmenia tepuiana V. Badillo, Emstia
Oyedaea DC. is largely Andean and contains 30: 1. 1985. TYPE: Venezuela. Bolivar: Dtto.
about 15 species, 6 in Venezuela, and 3 of these in Piar, Karun-tepui, 17 km E of Canaima, 6°14'N,
the Guayana Highland. The Brazilian species once 62°43'W, 1050 m, 31 Aug. 1983, Prance & Hu¬
referred to Oyedaea were transferred by Robinson ber 28382 (holotype, MY; isotypes, K, MO, MYF
(1984b) to Dimerostemma Cassini, which includes not seen, NY, VEN). Figure 1.
not only the 11 species treated by Robinson, but also
Distribution. This species is endemic to Au-
the invalid D. oblonga (Baker) Barroso, not keyed
by Robinson. Oyedaea and Dimerostemma have ster¬ ydn-tepui, Cerro Venado, and Kurun-tepui in Bo¬
lfvar, Venezuela, from 1050 to 1360 m elevation,
ile ray florets and winged disk cypselas, but Dime¬
but should be looked for in neighboring Brazil and
rostemma differs from Oyedaea by its distinctly foliar
outer series of phyllaries and coiled style branches. Guyana.
This species has sterile ray florets typical of Oye¬
Oyedaea is further distinguished by its disk cypselas
daea and is misplaced in the sister genus Zexmenia
with two slender awns and an apical neck.
La Llave, which has pistillate ray florets.
Oyedaea is expanded here to include Zexmenia
tepuiana V. Badillo and a newly described species,
Oyedaea wurdackii Pruski, sp. nov. TYPE: Ven¬
both endemic to the Venezuelan Guayana. Addi¬
ezuela. Amazonas: Dpto. Atures, carretera Pto.
tional collections are needed to accurately place 0.
Ayacucho hacia Samariapo, km 27, 6 km al
verbesinoides var. hypomalaca Steyermark. This tax¬
sur Tobogdn de la Selva, 5°24'N, 67°37’W, 28
on should not be recognized as a variety of O. ver¬
Sep. 1993, A. Groger 1125 (holotype, VEN;
besinoides, but rather included in the synonymy of
isotype, US). Figures 1, 2A-F, 3.
0. maculata S. F. Blake (also of northeastern Ven¬
ezuela), or, alternatively, elevated to the species Flosculis radiatis sterilibus necnon disci achaeniis
level. The key to the Venezuelan taxa of Oyedaea compressis alatis biaristatis cum genere Oyedaea perfecte
given by Aristeguieta (1964) is modified here to congrua, sed a congeneribus omnibus corollis (tarn disci
quam radiatis) albis (nec luteis) diversa.
include the new synonymy and two novelties.
Shrubs 0.5-3 m tall; stems subterete or subhexa-
Key to the Venezuelan Taxa of Oyedaea gonal, strigose, the intemodes 1-6 cm long, shorter to
very much shorter than the subtending leaves. Leaves
la. Leaves strongly triplinerved from above base.
simple, opposite, petiolate; petioles 5-10 mm long,
2a. Corollas white; Amazonas . . 0. wurdackii Pruski
2b. Corollas yellow. strigose, stout and much broadened at base, those of
3a. Involucres 10-14 mm tall; phyllaries stri- the same node often connected by a narrow rim;
gose, outer ones lanceolate; ray limbs 15- blades elliptic-lanceolate, 4.5-13 X 1.2-5.6 cm, char-
20 X 4—6 mm; widespread in the Carib¬ taceous, triplinerved from well above base, basally
bean coastal ranges, less frequent in the
Andes, also Trinidad, Colombia, Panama, cuneate to attenuate, the margins serrulate to serrate,
and Costa Rica (includes 0. verbesinoides apieally narrowly acute to attenuate, the surfaces
var. glabior Steyermark, syn. nov.) .... eglandular, the adaxial surfaces green, scabrous, the
... 0. verbesinoides DC. var. verbesinoides abaxial surfaces pale green, strigose, especially so on
3b. Involucres 5-6.5(—10) mm tall; phyllar¬ larger veins, also with scattered moniliform hairs.
ies puberulent, outer ones obpyriform;
ray limbs ca. 10 X 3 mm; Bolivar . . . Capitulescences cymose, open, terminal or axillary
. O. tepuiana (V. Badillo) Pruski from the distal nodes, of 9-15 capitula held above the
lb. Leaves ± pinnately veined. subtending leaves, at anthesis somewhat congested,
4a. Inner phyllaries rounded at the apices; Lara rounded, ca. 5—9 cm tall and broad, in fruit some¬
and Trujillo. O. obovata S. F. Blake
times more open; peduncles 0.5—3.5 cm long, hispid
4b. Inner phyllaries apieally acute.
408 No von
Figure 2. A—F. Oyedaea wurdackii Pruski. —A. Capitulum with ray corollas removed. —B. Phyllary. —C. Ray
floret. D. Pale. E. Disk floret. —F. Disk cypsela. G—L. Tuberculocarpus ruber (Aristeguieta) Pruski. —G. Capitulum
with front ray corollas removed. —H. Involucre. —I. Ray floret. —J. Pale. —K. Disk floret. —L. Disk cypsela. A-C
from Groger 625 (US); D-F from Maguire et al. 30583 (NY); G, 1-K from Gentry & Berry 14567 (NY, US); H from
Davidse & Huber 15381 (US); L from Huber 3787 (NY).
Volume 6, Number 4 Pruski 409
1996 Guayana Highland Compositae
Figure 3. Oyedaea wurdackii Pruski (type collection Groger 1125). Photo taken by Andreas Groger in Amazonas.
Venezuela.
to strigose. Capitula radiate, 35-53-flowered; involu¬ 10, sterile, lacking styles; corollas white, 10.5-12.3
cres narrowly campanulate, 7—10 X 5-10 mm; phyl- X 2-2.5 mm, the tubes 1.2-1.8 mm long, glabrous,
laries ca. 18, in ca. 3 series, imbricate, graduate, lan¬ the limbs 9.3-10.5 X 3-4 mm, shortly 2- or 3-lobed
ceolate, 6-10 X 1.5-3 mm, rigid, broad and scarious at apices, ca. 7-veined with 2 veins larger than the
at base, the apices herbaceous, strigose, narrowly at¬ others, abaxially puberulent. Disk florets 25-43, bi¬
tenuate to cuspidate, commonly reflexed; receptacles sexual; corollas actinomorphic, white, tubular-funnel-
flat, ca. 5 mm diam., paleate, the pales conduplicate, form, 7.5-8.3 mm long, the tubes 2-2.2 mm long,
often keeled near apices, to 10 X 1.4 mm, firm, scar¬ glabrous, the throats 5.1-5.3 mm long, glabrous, the
ious basally, the apices subherbaceous, sometimes lobes 0.7-0.8 mm long, triangular, puberulent-papil-
suffused with purple, stiffly attenuate. Ray florets ca. lose within, sometimes weakly puberulent on outer
410 Novon
surfaces; anthers slightly to much-exserted, ca. 3 mm Sphagneticola
long, the thecae black, the apical appendages elon¬
gate, black, eglandular, the connectives black when In the course of routine work for neotropical flo-
ristic treatments I saw a photograph of a plant
young to cream-colored, the filaments 2.S-3.5 mm
seemingly the same species as the widely known
long; style branches ca. 1 mm long, papillose, laxly
Wedelia trilobata (L.) Hitchcock. This photograph,
recurved, with paired stigmatic lines, the apices nar¬
however, was labeled as a phototype of Sphagneti¬
rowed. Ray ovaries sterile, 2.5-3.5 mm long, obconic,
cola ulei 0. Hoffmann, a name with which I was
3-awned, the awns 1.5-3(-4.5) mm long, subequal to
unfamiliar. Robinson’s (1981) observation that the
unequal. Disk cypselas compressed, winged, 5-6 X
black anthers and the cypselas with a fimbriate co-
1.5-2.5 mm, the body black, weakly puberulent, the
roniform pappus found in unispecific Sphagneticola
wings ciliate, cream-colored, ca. 0.5 mm broad at ma¬
O. Hoffmann “indicate a probable place in the
turity; pappus stoutly 2-awned, the awns scabridulous, Ecliptinae” and the placing of W. trilobata in a new
subequal to unequal, 2-3.3 mm long, these exserted genus (Complaya) by Strother (1991) furthered my
from the involucre and often reaching to the top of interest. I have now seen type material of S. ulei,
the pales, the squamellae between the awns ca. 5, to which indeed proves to be conspecific with the
I mm long. widespread plant known in cultivation in the trop¬
ics and subtropics of both hemispheres as W. tri¬
Distribution and ecology. This species is known lobata (n.v. Wedelia, Creeping ox-eye. Wild mari¬
only from Amazonas, Venezuela, at elevations from gold, etc.). Sphagneticola is not only squarely
100 m to under 1000 m (J. Wurdack, pers. comm.), positioned within Ecliptinae sensu Robinson
at the escarpment base and slope of the northwest¬ (1981), but it is a generic name (earlier than Com¬
ern part of Cerro Yapacana, in secondary growth on playa and Thelechitonia) for W. trilobata', thus the
recently burned granitic lajas of Cerro Caldero and combination S. trilobata (L.) Pruski is proposed.
Cerro Danto, and in secondary areas at Tobog&n de Sphagneticola trilobata differs from most wede-
la Selva near Coromoto. It is known to flower in lias by being a stoloniferous herb rooting at the
January, September, and December. The species is nodes with stems elongating sympodially, thereby
to be expected from adjacent regions of Bolivar, laterally displacing the terminal capitula; by abax-
Venezuela, and Vichada, Colombia. ially glandular ray limbs; by black ovate anther ap¬
This new species matches yellow-flowered Oye- pendages; and by tuberculate cypselas without
daea as circumscribed by Blake (1921) except in well-developed carpopodia and without elaiosomes
(white) corolla color. All other oyedaeas have yellow or pappus awns. It and two other species were re¬
corollas. The triplinerved leaves and open capitu- cently treated at the generic rank (as Complaya) by
lescence of O. wurdackii most closely resemble Strother (1991). A fourth species of this group was
those of 0. tepuiana. treated by Cuatrecasas (1954) as Thelechitonia
I am pleased to name this Oyedaea after one of its muricata Cuatrecasas. Angel Cabrera noted that
discoverers, my friend and colleague John Julius Wur¬ Thelechitonia muricata is conspecific with W. bra-
dack. John is a co-discoverer of Cerro de la Neblina chycarpa Baker (J. Cuatrecasas, pers. comm.), and
and one of the foremost authorities on the plants of the name Thelechitonia was treated as a synonym
Guayana. In the 1950s as a staff member of The New of Wedelia by Robinson (1981). Robinson and Cua¬
York Botanical Garden he published or co-published trecasas (1992), prompted by Strother’s publication
6 new genera and 36 new species of Compositae from of Complaya, resurrected Thelechitonia and treated
Complaya as a generic synonym of Thelechitonia.
Guayana, thus the dedication of this new composite
They expanded Thelechitonia to include four spe¬
to him is appropriate and well-deserved.
cies.
The types of Sphagneticola ulei and Wedelia tri¬
Paratypes. VENEZUELA. Amazonas: Dpto. Atures,
lobata are conspecific, and the earlier generic name
Rfo Sipapo, 5 km de su desembocadura al Rfo Orinoco,
Cerro Caldero, 5°01'N, 67°46'W. 17 May 1992 (late fruit), must be used for this group of four species. I Pr0'
A. Groger & J. Barcroft 413 (US, VEN); Dpto. Atures, vide an updated generic synonymy and four new
carretera Pto. Ayacucho hacia Samariapo, km 19, Cerro combinations in Sphagneticola. The generic and
Danto al lado oriental de la carretera, 5°30'N, 67°3.V\Y. specific circumscriptions of Strother (1991) and
II Dec. 1992, A. Groger 625 (US, VEN); NW slopes of Robinson and Cuatrecasas (1992) are followed. An
Cerro Yapacana, 1 Jan. 1951, B. Maguire, R. Cowan & J.
Wurdack 30583 (NY); Cerro Yapacana, base of NW es¬ earlier epithet (blocked in Wedelia, but available
carpment, 2 Jan. 1951, B. Maguire, R. Cowan & J. Wur¬ here) is used for the paleotropical plant most widely
dack 30638 (NY). known as W. chinensis (Osbeck) Merrill.
Volume 6, Number 4 Pruski 411
1996 Guayana Highland Compositae
Sphagneticola 0. Hoffmann, Notizbl. Kdnigl. Bot. The leaves of this species are unlobed, and the
Gart. Berlin 3: 36. 1900. TYPE: Sphagneticola plants are more erect than the two other New World
ulei 0. Hoffmann [= Sphagneticola trilobata species. Many specimens of this species have been
(L.) Pruski]. determined previously as W. brasiliensis (Sprengel)
S. F. Blake (a synonym of S. trilobata). Strother
Wedelia sect. Stemmodon Grisebach, FI. Brit. W. I. 371.
1861. TYPE: not designated, but including only (1991) noted that such specimens “may represent
Wedelia carnosa Richard, nom. illeg. [= Sphagneti¬ a.. .species of Complaya.”
cola trilobata (L.) Pruski] and Wedelia gracilis Rich¬
ard [= Sphagneticola gracilis (Richard) Pruski], Sphagneticola calendulacea (L.) Pruski, comb. nov.
Thelechitonia Cuatrecasas, Bull. Soc. Bot. France 101:
242. 1954. Syn. nov. TYPE: Thelechitonia muricata Basionym: Verbesina calendulacea L., Sp. PI.
Cuatrecasas [= Sphagneticola brachycarpa (Baker) 902. 1753. Jaegeria calendulacea (L.) Sprengel,
Pruski], Syst. Veg. 3: 590. 1826. Wedelia calendulacea
Complaya Strother, Syst. Bot. Monogr. 33: 10. 1991. (L.) Lessing, Syn. Compos. 222. 1832, non Per-
TYPE: Silphium trilobatum L. [= Sphagneticola tri¬ soon 1807. Seruneum calendulaceum (L.) Kun¬
lobata (L.) Pruski],
tze, Revis. Gen. PL 1: 365. 1891, nom. illeg.
Sphagneticola contains four species that are (Seruneum Rumphius ex Kuntze, 1891, includes
common at lower elevations throughout much of the the type of Wedelia Jacquin, 1760.) TYPE: “Zey-
tropics and subtropics (though not known to occur lona:” s.d., Hermann [lectotype, chosen here.
in Africa), often near waterways or along coasts: S. Herb. Hermann, no. 311, vol. l,fol. 73 (BM not
trilobata, S. brachycarpa (Baker) Pruski, S. calen- seen [photograph, US]); possible isolectotypes.
dulacea (L.) Pruski, and S. gracilis (Richard) Pru¬ Herb. Hermann, no. 311, vol. 2,fol. 16 (BM not
ski. Two species, .S', trilobata and S. brachycarpa, seen [photograph, US]), Herb. Hermann, no. 311,
are known in the Guayana Highland of northeastern vol. 3, fol. 21 (BM not seen [photograph, US]),
South America. Two species, S. calendulacea and Herb. Hermann, no. 311, vol. 4, fol. 43 (BM not
S. trilobata, are used medicinally, and .S', trilobata seen [photograph, US]), Collectio plantarum Zey-
is also known to cause contact dermatitis (Lovell, lanicum.. .p. 65 (bottom center) (Bibliotheque de
1993; pers. obs.). l’lnstitut de France [photograph, US])]. Grierson
(1980) listed ‘TYPE: Herb. Herman [sic] (BM)”;
Sphagneticola brachycarpa (Baker) Pruski, however, 1 have seen photographs of four Her¬
comb. nov. Basionym: Wedelia brachycarpa mann specimens of this species scattered be¬
Baker, in C. Martius, FI. Bras. 6(3): 181. 1884. tween four of the five volumes of the Hermann
Seruneum brachycarpum (Baker) Kuntze, Re- collection. Thus, the statement by Grierson is not
vis. Gen. PI. 1: 365. 1891, nom. illeg. (Seru¬ specific enough for formal lectotypification (C.
neum Rumphius ex Kuntze, 1891, includes the Jarvis, pers. comm.). The Hermann specimen on
type of Wedelia Jacquin, 1760.) Stemmodontia p. 65 of the Burman herbarium folio in the Bib-
brachycarpa (Baker) Morong, Ann. New York liotheque de lTnstitut de France was presumably
Acad. Sci. 7: 147. 1893. Thelechitonia bra¬ the basis for tab. 22, fig. 1, in Burman, Thes.
chycarpa (Baker) H. Robinson & Cuatrecasas, Zeylan. 52. 1737 [1736], cited in the protologue
Phytologia 72: 142. 1992. TYPE: Paraguay. by Linnaeus (see Lourteig, 1966: 28), but be¬
Along the margins of Rfo Paraguay near As¬ cause it is unlikely that Linnaeus saw this sheet
sumption, 28 Apr. 1874, Balansa 855 (holo- it is not chosen as the lectotype. LINN 1021.6
type, K not seen [photograph and fragment, not seen (IDC microfiche 177. 610.III.6) is
US]; isotype, G not seen [photograph, US]). marked in Linnaeus’s hand “calendulacea 9”
(Savage, 1945), referring this sheet to the ninth
?Wedelia paludosa DC. var. villosa Baker, in C. Martius,
FI. Bras. 6(3): 181. 1884. Wedelia brasiliensis species of Verbesina in Species Plantarum, 1753.
(Sprengel) S. F. Blake var. villosa (Baker) S. F. Blake, However, LINN 1021.6 is also labeled “chin[a]”
J. Wash. Acad. Sci. 21: 332. 1954. TYPE: Guyana. (Savage, 1945), which does not agree with the
Ad fluv. Essequibo et Rupununi, s.d., Appun 2508 data given by Linnaeus in the protologue. Thus,
(holotype, K not seen nor found).
Thelechitonia muricata Cuatrecasas, Bull. Soc. Bot. France LINN 1021.6 was not chosen as the lectotype,
101: 242. 1954. TYPE: Colombia. Meta: Rfo Casanare, nor is it listed here as a possible isolectotype.
Esmeralda, 130 m, 19 Oct. 1938, Cuatrecasas 3902
(holotype, F not seen [photograph, US]; isotype, US). Solidago chinense Osbeck, Dagb. Ostind. Resa 241. 1757.
Syn. nov. Wedelia chinensis (Osbeck) Merrill, Philipp.
Distribution. Venezuela (Guayana and in the J. Sci. 12: 111. 1917. Complaya chinensis (Osbeck)
Llanos), Colombia, Guyana, Peru, Brazil, Bolivia, Strother, Syst. Bot. Monogr. 33: 14. 1991. Thelechitonia
Paraguay, and Argentina. chinensis (Osbeck) H. Robinson & Cuatrecasas, Phy-
412 Novon
tologia 72: 142. 1992. TYPE: China. Kwangtung: Da¬ made. Specimens from both localities cited in
ne’s Island, near Whampoa, 20 Oct. 1751, Osbeck s.n. the protologue match this species as circum¬
(holotype, LINN? not seen or S? not seen).
scribed here, and lectotype designation is de¬
Distribution. China including Taiwan, India, ferred until material in P is seen.
Indo-China, Japan, Malaysia, the Philippines, and
Distribution. Greater Antilles, Antigua, Barbu¬
Sri Lanka (Li, 1978).
da, Guadeloupe, Martinique, St. Barthelemy, and
I examined the photographs of the four Hermann
St. Martin (Howard, 1989).
specimens of Verbesina calendulacea L. in BM (one
This West Indian endemic is closely related to
of which is here designated as the lectotype), the
S. trilobata but differs by smaller leaves, capitula,
photograph of the Hermann specimen in the Bib-
and cypselas.
liotheque de l’lnstitut de France, and the proto-
logue, and agree with Merrills (1917) observation
that Solidago chinensis Osbeck and V. calendulacea Sphagneticola trilobata (L.) Pruski, comb. nov.
are conspecific. The Linnaean epithet was blocked Basionym: Silphium trilobatum L., Syst. Nat.,
in Wedelia by W. calendulacea Persoon, thus Merrill ed. 10: 1233. 1759. Wedelia carnosa Richard,
proposed the combination W. chinensis, which for in Persoon, Syn. Pl. 2: 490. 1807, nom. illeg.
much of this century has been the name used for Seruneum trilobatum (L.) Kuntze, Revis. Gen.
this plant. The epithet “calendulacea” was avail¬ PI. 1: 365. 1891, nom. illeg. (Seruneum Rum¬
able for this species in both Complaya and Thele- phius ex Kuntze, 1891, includes the type of
chitonia, but was not used. This species was sug¬ Wedelia Jacquin, 1760.) Wedelia trilobata (L.)
gested by Fosberg (1993) as perhaps being better A. Hitchcock, Annual Rep. Missouri Bot.
placed in Wollastonia DC. than in Wedelia, but is Gard. 4: 99. 1893. Stemmodontia trilobata (L.)
not properly placed in either genus. Small, FI. S.E. U.S. 1262, 1340. 1903. Com¬
Nicolson et al. (1988) listed opposite-leaved Ver¬ playa trilobata (L.) Strother, Syst. Bot. Monogr.
besina calendulacea L. and Solidago chinensis Os¬ 33: 14. 1991. Thelechitonia trilobata (L.) H.
beck as synonyms of “Wedelia chinensis (L.) Merrill,” Robinson & Cuatrecasas, Phytologia 72: 142.
a combination not made by Merrill. The “Wedelia 1992. TYPE: Tab. 107, fig. 2, in Plumier, PI.
chinensis (L.) Merrill,” in the sense of Nicolson et Amer. 1757 (lectotype, designated by Howard,
al. (1988) is based on Verbesina chinensis L. [= An- 1989).
isopappus chinensis (L.) Hooker f. & Amott], a spe¬
Buphthalmum strigosum Sprengel, Neue Entd. 2: 140.
cies with alternate leaves of tribe Inuleae. Nicolson 1821. Acmella brasiliensis Sprengel, Syst. Veg. 3:
et al. (1988) interpreted opposite-leaved Solidago 592. 1826, nom. superfl. Wedelia brasiliensis (Spren¬
chinensis Osbeck as either a “nom. illeg. or comb, gel) S. F. Blake, Contr. U.S. Natl. Herb. 26: 250.
nov.?” based on alternate-leaved Verbesina chinensis 1930, comb, illeg. TYPE: Brazil, s.d., Sello s.n. (ho¬
L., whereas I follow Merrill (1917) and treat Soli¬ lotype, P not seen)
Wedelia paludosa DC., Prodr. 5: 538. 1836. TYPE: Brazil,
dago chinensis as a different taxon based on a dif¬ s.d., Lund 611 (holotype, G-DC not seen [photo¬
ferent type described by Osbeck. graph, US; IDC microfiche 800. 935.1.3]).
Sphagneticola ulei O. Hoffmann, Notizbl. Kiinigl. Bot.
Sphagneticola gracilis (Richard) Pruski, comb, Gart. Berlin 3: 36. 1900. Syn. nov. TYPE: Brazil.
Rio de Janeiro: in Stimpfen der Restinga der Gavea,
nov. Basionym: Wedelia gracilis Richard, in Sep. 1899, Ule 3914pp (holotype, B destroyed, pho¬
Persoon, Syn. PI. 2: 490. 1807. Seruneum grac- tographs in NY, US; lectotype, designated here from
ile (Richard) Kuntze, Revis. Gen. PI. 1: 365. isotype, HBG). [Another sheet is also labeled Ule
1891, nom. illeg. (Seruneum Rumphius ex Kun¬ 3914 (HBG), but was collected in Apr. 1895 at Ja-
tze, 1891, includes the type of Wedelia Jacquin, car^pagud and is not type material.]
1760.) Complaya gracilis (Richard) Strother, Distribution. Wedelia is widespread in Guaya-
Syst. Bot. Monogr. 33: 14. 1991. Thelechitonia na (where it occurs from 119 to 900 m elevation)
gracilis (Richard) H. Robinson & Cuatrecasas, and in the rest of the Neotropics from Mexico and
Phytologia 72: 142. 1992. SYNTYPES: Antigua the West Indies south to Argentina; widely culti¬
and Santo Domingo, s.d., presumably Richard vated out-of-doors as an ornamental and natural¬
collections in P (not seen). The original Richard ized in eastern Australia, Malaysia, Pacific islands,
collection from Antigua was doubtfully referred and warmer regions of the New World. It is grown
to the species by Schulz (1911: 99-100), thus in at least four U.S. states: California (in Berkeley
defining “the type locality as Santo Domingo” fide J. Strother, pers. comm.), Florida (well known
(Howard, 1989). I am unaware, however, wheth¬ in south Florida; recently reported in Tallahassee
er a lectotype (sheet) designation has been (Anderson, 1995), near its northernmost limit), Ha
Volume 6, Number 4 Pruski 413
1996 Guayana Highland Compositae
waii, and Louisiana. The species has been noted in Elementa Botanica (1790) are “not to be treated”
Louisiana by Odenwald and Turner (1978), but was as genera (Art. 20.4 (b) Ex. 11, Greuter et al.,
not vouchered by them. It was not mentioned as 1988), and this work is listed in Appendix V (Ope¬
cultivated in New Orleans by Seidenberg (1993), ra Utique Oppressa) in Greuter et al. (1994).
although it is used as an ornamental there (Melinda Tilesia (1818) and Wulffia Necker ex Cassini
Taylor, pers. comm.; pers. obs.). Sphagneticola tri- (1823) were equated by Schultz-Bipontinus (1848)
lobata is here newly documented for Louisiana by and Bentham (1873: 367) under Wulffia, the later
the following out-of-doors collections from Baton genus. I do not wish to continue this misusage.
Rouge, Lafayette, and New Orleans: Harris 774 Thus, I herein propose transfers of three species
(LSU), Pruski et al. 3711 (NO, NY), Pruski 3724 and one variety from Wulffia to the earlier Tilesia.
(US), Pruski 3742 (K, LSU, MO, NO, US), Thomas Lastly, literature search reveals a misattribution
22247 (NLU), Thomas & Allen 74948 (NLU), Ur- of the authorship of Coreopsis baccata by Jackson
batsch 5155 (LSU). (1893—1895) in Index Kewensis to Linnaeus filius,
This species has variable leaf morphology. where it appears in Supplementum Plantarum (Lin¬
Thick-leaved forms, often cultivated, were previ¬ naeus filius, 1782). Additionally, Jackson noted
ously referred to W. carnosa, whereas longer, more that Coreopsis baccata also appears in Linnaeus’s
narrow-leaved forms were determined as either W. Amoen. Acad, as species no. Ill (Dalberg collec¬
brasiliensis or W. paludosa. A complete list of syn¬ tion no. 15) on p. 262, 1785. However, it is noted
onyms is given in Schulz (1911). here that Coreopsis baccata was earlier published
by Linnaeus (though not indexed as such by Jack-
son, 1893-1895) in Plantae Surinamenses (Linnae¬
TlLESIA
us 1775: 14) as an unnumbered taxon (Dalberg col¬
During preparation of fioristic treatments of neo¬ lection no. 15) and this name is cited below in the
tropical Compositae, Wulffia Necker ex Cassini (not transfer of that taxon to Tilesia.
Wolffia Horkel ex Schleiden, 1844, Lemnaceae)
was noted as a taxonomic synonym of Tilesia. Ac¬ Tilesia G. Meyer, Prim. FI. Esseq. 251. 1818.
cordingly, Wulffia is reduced to the synonymy of TYPE: Tilesia capitata G. Meyer [= Tilesia
Tilesia, and the three species and one variety of baccata (L.) Pruski var. baccata].
Wulffia are transferred to Tilesia.
Wulffia Necker, Elem. Bot. 1: 35. 1790, nom. illeg.; ex
Tilesia capitata G. Meyer was described as a new Cassini, in F. Cuvier, Diet. Sci. Nat. ed. 2, 29: 491.
genus and species in Primitiae Florae Essequeboen- 1823. Syn. nov. TYPE: Coreopsis baccata L. [= Ti¬
sis (Meyer, 1818). Tilesia capitata is the same as lesia baccata (L.) Pruski var. baccata].
the species subsequently called either Wulffia bac- Chatiakella Cassini, in F. Cuvier, Diet. Sci. Nat. ed. 2,
29: 491. 1823. TYPE: Chylodia sarmentosa Richard
cata (L.) Kuntze or W. stenoglossa (Cassini) DC. Al¬ ex Cassini [= Tilesia baccata (L.) Pruski var. bac¬
though the epithet “baccata” has priority, the name cata].
most widely used in Brazil and Venezuela is W. Chylodia Richard ex Cassini, in F. Cuvier, Diet. Sci. Nat.
stenoglossa (Cassini) DC. due largely to use by Ba¬ ed. 2, 29: 491. 1823, non Chilodia R. Brown, 1810.
TYPE: Chylodia sarmentosa Richard ex Cassini [=
ker (1884) and Aristeguieta (1964). Tilesia baccata (L.) Pruski var. baccata].
Five years after Meyer’s flora, Cassini (1823)
treated Coreopsis baccata (the basionym of Wulffia Tilesia is restricted to the Neotropics and con¬
baccata) as equal to the mononomial Wulffia of tains a somewhat wide-ranging species with two va¬
Necker (Elem. Bot. 1: 35. 1790). Also, Cassini not¬ rieties and two uncommon species. Two species (T.
ed that the unpublished genus and species Chylo- baccata var. baccata and T. rubens) occur in Gua¬
dia sarmentosa Richard was possibly near or the yana; one of them (T. rubens) is a Guayana High¬
same as Wulffia, and to avoid confusion with Chil- land endemic. Full synonymies are given for three
odia R. Brown (1810), he suggested Chatiakella as of the four taxa, but for Tilesia baccata var. baccata
a replacement generic name for Richard’s Chylo- only pertinent and regional synonymy is given. Ti¬
dia. Cassini was uncertain about the ultimate dis¬ lesia is partly characterized by eglandular leaves,
position of these names and treated them as poten¬ rays sterile (when present), strongly striate pales,
tial synonyms. He said if these three genera were and baccate fruits lacking pappus. Wedelia ambi-
indeed synonymous they should be united under gens S. F. Blake resembles Tilesia by strongly stri¬
the oldest available name, which he took to be ate pales and epappose cypselas. Wedelia ambi-
Wulffia Necker. Indeed, in Cassini’s time names of gens, however, has leaves glandular abaxially and
Necker were accepted. Unfortunately, Necker’s fertile ray florets, and therefore is retained in Wed¬
names are mononomials and now new names in his elia.
