Taxonomic Revision of Trichogonia (Eupatorieae, Asteraceae): A South American
Genus
Author(s) :Nádia Roque, Hortensia Pousada Bautista, and Aline Costa da Mota
Source: Systematic Botany, 37(2):525-553. 2012.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364412X635575
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and
environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published
by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of
BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries
or rights and permissions requests should be directed to the individual publisher as copyright holder.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research
libraries, and research funders in the common goal of maximizing access to critical research.
Systematic Botany (2012), 37(2): pp. 525–553
© Copyright 2012 by the American Society of Plant Taxonomists
DOI 10.1600/036364412X635575
Taxonomic Revision of Trichogonia (Eupatorieae, Asteraceae): a South American Genus
Nádia Roque,1,4 Hortensia Pousada Bautista,2 and Aline Costa da Mota3
1
Universidade Federal da Bahia, Instituto de Biologia, Depto. de Botânica, Rua Barão de Geremoabo s. n.,
campus Universitário de Ondina, 40.171-970, Salvador, Bahia, Brazil.
2
Universidade do Estado da Bahia, Depto. Ciências da Vida, Rua Silveira Martins, 2555, 41.150-000, Salvador, Bahia, Brazil.
3
Programa de Pós-Graduação em Botânica, Depto. de Ciências Biológicas, Universidade Estadual de Feira de Santana,
Km 03, BR 116, 44031-460, Feira de Santana, Bahia, Brazil.
4
Author for correspondence: ([email protected])
Communicating Editor: Mark P. Simmons
Abstract—Eupatorieae is one of 43 tribes circumscribed in Asteraceae and it is characterized mainly by opposite leaves, discoid heads,
tubular, white, pink, or lilac florets, style branches linear to clavate, and cypsela walls with black or carbonized resin-like deposits in mature
stages. The tribe is essentially Neotropical, found in Mexico, Central America, and South America, with 29 genera, of which Trichogonia is
one of them. Trichogonia is found only in South America and is characterized by stems with prominent longitudinal ridges and the presence
of dense pubescence on the upper throat and lobes of the corolla. The present work is a taxonomic revision of Trichogonia with illustrations
and geographic distribution maps for every species. Morphological data were obtained from the field and from specimens from 26 herbaria.
From a total of 20 recognized species, 17 occur in Brazil, one is endemic to Bolivia (T. capitata), and two are endemic to Paraguay (T. chodatii
and T. phlebodes). From the species currently known to occur in Brazil, 14 are endemic, while T. salviifolia Gardner and T. arguta have
widespread distributions. Narrow endemics are found in the states of Bahia (T. harleyi, T. spathulifolia, T. tombadorensis) and Minas Gerais
(T. hirtiflora). Other species share a distribution between Pernambuco and Bahia (T. heringeri), Tocantins, Piauı́ and Bahia (T. campestris),
Bahia and Goiás (T. cinerea and T. laxa), Bahia and Minas Gerais (T. villosa), Goiás and Minas Gerais (T. grazielae). Trichogonia occur mostly in
savannas cerrados, caatingas, campos gerais, and campos rupestres. Thirteen lectotypifications are designated and 14 new synonyms are
proposed in this paper.
Keywords—Compositae, morphology, nomenclature, taxonomy, Trichogonia.
site or alternate, spiral leaves, eximbricate or slightly subimbricate phyllaries, heads with 10 or more florets, and
pappus of many capillary or sometimes plumose bristles.
This subtribe is considered to be related to a paraphyletic
Eupatoriinae subtribe (Robinson et al. 2009).
The Gyptidinae are distributed in South America, with the
greatest concentration of genera in Brazilian savannas. Out of
a total of 29 genera (ca. 150 species), 25 can be found in Brazil,
20 of which are in northeastern Brazil, with eight genera
entirely restricted to the campos rupestres (fields with rocky
outcrops) of the state of Bahia (King and Robinson 1987;
Hind and Miranda 2008).
Trichogonia (DC.) Gardner belongs to subtribe Gyptidinae
and is characterized by stems with prominent, longitudinal
ridges, the presence of dense pubescence on the corolla
throat and lobes, cypselae with setose ribs, and by a plumose
pappus. Trichogonia was first mentioned by De Candolle
(1836) as a section of Kuhnia L., with four species with trichomes on the cypselae angles (ribs), in opposition to section
Leiogonia where trichomes on cypselae angles are absent.
Gardner (1846) transferred De Candolle’s section Trichogonia
into a new genus, and added three species from Brazil. Later,
Baker (1876) in his treatment for the Asteraceae in Martius’s
Florae Brasiliensis added three species for a total of ten. In
1951, Barroso recognized 18 species of Trichogonia for Brazil.
Studies of Eupatorieae started with B. L. Robinson (1913)
and continued by King and Robinson (1977, 1980), where
new arrangements were proposed within the tribe. In addition, generic concepts were redefined, or dismembered into
smaller genera. As a result, Trichogonia and related genera
were more tightly circumscribed, and the following combinations were made (King and Robinson, 1972, 1978, 1980):
T. chodatii (Hassler) R. M. King & H. Rob., T. phlebodes (B. L.
Rob.) R. M. King and H. Rob., Trichogonia cinerea (Gardner)
R. M. King & H. Rob., and T. dubia (B. L. Rob.) R. M. King &
H. Rob. In addition, a few species were transferred into
Asteraceae are the biggest family of angiosperms with ca.
24,000 species and 1,600–1,700 genera with a cosmopolitan
distribution (Funk et al. 2009b). Recent molecular studies of
Asteraceae recognize 12 subfamilies and 43 tribes (Panero and
Funk 2008; Funk et al. 2009a), a higher number of subfamilies
and tribes compared to traditional concepts (Bremer 1994).
Generally, the Eupatorieae is characterized by opposite
leaves, homogamous and discoid heads, tubular, white, pink,
or lilac florets, style branches lineate or clavate, and cypsela
walls with black or carbonized resin-like deposits in mature
stages (King and Robinson 1987). The tribe is essentially Neotropical, however, a few species are found in North America,
around the tropics or in the Old World (Bremer 1994).
The Eupatorieae was divided by King and Robinson (1987)
into 18 subtribes based on micromorphological, macromorphological, and anatomical features. In addition, Robinson
et al. (2009) proposed the new subtribe Piqueriinae based
on molecular studies, and published a phylogeny of the
Eupatorieae based on genera from 15 subtribes. Although
the phylogenetic positions of these subtribes are not well
resolved, the cladogram has the Mexican genera in basal
position, suggesting that the South American taxa are the
result of a recent radiation.
In a later study, Ferreira (2010), using ITS and trnL-F,
concluded that genera of the Gyptidinae sensu King and
Robinson (1987) do not constitute a monophyletic group.
Their analyses suggest that six other genera from other tribes
also belong there. In their studies, the basal genera (which
include Trichogonia, Campuloclinium, Barrosoa) are widely
distributed in South America; while smaller genera as
Agrianthus, Lasiolaena, Stylotrichium, have restricted distributions, suggesting recent origins due to adaptation to new
habitats such as the open fields of northeastern Brazil.
Subtribe Gyptidinae is one of the largest and most complex
subtribes of Eupatorieae (King and Robinson 1987; Hind
1999, 2000; Hind and Robinson 2007), characterized by oppo525
526
SYSTEMATIC BOTANY
Trichogoniopsis R. M. King & H. Rob., Platypodanthera R. M.
King & H. Rob., and Campuloclinium DC. In Trichogoniopsis,
the corolla throat and lobes have sessile glandular trichomes and the stem is nearly smooth; in Platypodanthera
the receptacle is convex, the stem is smooth and the corolla
is glabrous; and in Campuloclinium, the receptacle is convex
and the corolla has different pubescence when compared
with Trichogonia (King and Robinson 1980). In summary,
King and Robinson (1972, 1978, 1980, 1987) recognized 30 species in Trichogonia, 21 of which were considered endemic
to Brazil.
Ferreira (2010) recognized Trichogoniopsis as a sister group
of Platypodanthera and Trichogonia, a monophyletic clade
with high bootstrap support. The morphological characters
were mapped on the cladogram and it was found that
the plumose to sub-plumose pappus are synapomorphous,
a character state apparently unique to this clade in the
Subtribe Gyptidinae. In addition, Ferreira (2010) suggested
the stipitate cypsela as a synapomorphous character for
Platypodanthera and Trichogonia and the presence of stems
with prominent, longitudinal ridges and corolla throat and lobes
with dense pubescence as autopomorphous for Trichogonia.
The present taxonomic revision of Trichogonia recognizes
20 species and provides an identification key, descriptions,
illustrations, geographic distribution maps, and taxonomic
discussions for every species.
Materials and Methods
Morphological data were obtained from the analysis of material
collected in Brazil (T. attenuata, T. campestris, T. hirtiflora, T. salviifolia,
T. tombadorensis and T. villosa), and from specimens borrowed from
26 herbaria (Holmgren et al. 1990) as follows: ALCB, CEN, CEPEC, EAC,
ESA, F, G, HB, HEPH, HRB, HRCB, HUEFS, IBGE, INPA, IPA, K, MBM,
MBML, MG, R, RB, SP, SPF, UB, UEC, and US.
Species delimitations are based on the study of type material (ca.
170 specimen types, including digital images), original descriptions, available specimens (ca. 460), and main references in the genus (cited throughout the manuscript). Specimens were analyzed and illustrated by using
an Olympus SZH10 stereomicroscope with attached camera lucida.
The terms ‘regular’ versus ‘irregular’ (or defective) are used to describe
the pappus, the former refers to the presence of pappus in all florets,
while the latter to the absence of pappus (florets epappose) or if present,
only in the inner florets. The term “defective pappus” was proposed for
Trichogonia by King and Robinson (1980).
Species are treated in alphabetical order, their descriptions are accompanied by an illustration that highlight diagnostic characters. Morphological terminology follows Radford et al. (1974), Roque and Bautista
(2008), and Roque et al. (2009).
Taxonomic Treatment
Trichogonia (DC.) Gardner, London J. Bot. 5: 459. 1846.
Kuhnia L. sect. Trichogonia DC., Prodr. 5: 126. 1836.—
LECTOTYPE: Trichogonia arguta (Kunth) Benth. & Hook.
ex Klatt (designated by King and Robinson 1987).
Erect or rarely prostrate herbs, subshrubs, weakly
branched above base; stems terete, with prominent longitudinal ridges, sometimes striate, pubescent, often with glandular trichomes, viscous or not. Leaves alternate, spirally
arranged, or alternate, spirally arranged, usually lower
leaves opposite and similar to upper leaves, or larger than
upper leaves, or alternate, fasciculate or congested (cluster
[Volume 37
of leaves with a short internodes), commonly all with axillary and congested branches and leaves, coriaceous,
membranaceous or chartaceous, concolorous or discolorous,
petiolate, subsessile or sessile; laminae linear, elliptic, lanceolate, triangular, ovate, or obovate, acute, obtuse, or rounded
at apex, entire, crenate, or serrate on margins, attenuate,
truncate, or cordate at base; venation pinnate, 3–5-nervate or
inconspicuous. Capitulescences corymbose-paniculate, lax,
with homogamous and discoid, subsessile to pedicellate
heads; involucres campanulate; receptacles flat or convex,
glabrous; phyllaries 12–34, 2–3-seriate, eximbricate, subequal, abaxial surface with prominent veins, puberulous,
pubescent, tomentose, glandular trichomes sessile or
stipitate. Florets 18–100, corolla pink or lilac, becoming
whitish with age, narrowly funnelform, densely pubescent on
limb including the lobes, lobes triangular, conspicuous or
hidden by a dense pubescence, corolla tube glabrous or pubescent, glandular trichomes sessile and/or stipitate; anther connective appendages obtuse, retuse or acute, as long as wide
or longer, anther collars cylindrical, thickened; style bases
not enlarged, glabrous, style branches clavate, mamillose, lilac
with white apex. Cypselae prismatic, cylindrical, 5-ribbed,
long-, short- or not-stipitate, villous or setuliferous at least on
ribs or less often glabrous; carpopodium annular or the ribs
decurrent into it; pappus plumose to sub-plumose, persistent,
bristles 14–30, in one series, regular, defective or irregular
(absent at least on the outer florets) or absent. Chromosome
number x = 10 (Robinson et al. 1989).
Trichogonia is a South American genus of 20 species, 14
of which are endemic to Brazil, one to Bolivia (T. capitata
(Rusby) B. L. Rob.), and three to Paraguay (T. chodatii (Hassler)
R. M. King & H. Rob. and T. phlebodes (B. L. Rob.) R. M. King &
H. Rob.). The remaining three species are more widely distributed, with T. salviifolia Gardner found in Bolivia, Brazil,
Colombia, Paraguay, and Venezuela, T. arguta (Kunth) Benth. &
Hook. f. ex Klatt in Brazil, Colombia, and Venezuela, and
T. hassleri Mattf. known from Paraguay and Brazil.
The Brazilian endemics are distributed in the states of
Ceará, Brası́lia, Espı́rito Santo, Goiás, Mato Grosso, Mato
Grosso do Sul, Minas Gerais, Pernambuco, Piauı́, Rio de
Janeiro and São Paulo (Fig. 1). Of these, three, T. harleyi R. M.
King & H. Rob., T. spathulifolia Mattf. and T. tombadorensis
R. M. King & H. Rob., are found only in Bahia and one,
T. hirtiflora (DC.) Sch. Bip. ex Baker, in Minas Gerais. The
remaining species are distributed in two or more states. The
distribution of each species is shown in the maps in the individual treatment of each species. The species occur mostly in
open savanna-like vegetation such as cerrado, campos gerais,
campos rupestres (rocky fields), and caatinga (deciduous low
forest and thorny thicket vegetation) (Veloso et al. 1991).
The name Trichogonia refers to the presence of trichomes
along cypselae angles (Fig. 2D, G). The species of Trichogonia
can be classified into two groups, one whose heads present
convex receptacles and glandular stipitate trichomes on
the corolla tube (T. cinerea, T. eupatorioides, T. hassleri, and
T. prancei); a second group includes most species and is characterized by flat receptacles and glabrous corolla tube. Variations on leaf characters (phyllotaxy, texture, shape, size,
and indumentum), number of florets per head, cypselae
(base, indumentum, carpopodium) and pappus (regular
or absent in some or all cypsela) are diagnostic only at the
specific level.
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
527
Fig. 1. South American geographical distribution of Trichogonia (DC.) R. M. King & H. Rob.
Key to the Species of TRICHOGONIA
1.
Receptacles convex (Figs. 6D, 13E ); corolla tube with glandular stipitate trichomes (Figs. 6C, 9E–F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Pappus regular (present in all cypselae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Leaves ovate, margin dentate, cordate at base, adaxial surface puberulous, venation trinervate;
florets 25–30; cypselae cylindrical; pappus 3–4 mm long (Paraguay, Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. hassleri
3. Leaves elliptic, margin crenate, base cuneate, adaxial surface tomentose, venation pinnate;
florets 60–80; cypselae short-stipitate; pappus shorter than 0.6 mm long (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. prancei
2. Pappus absent or irregular (absent at least in the marginal florets) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Lamina oblong to elliptic, margin crenate; florets 60–100; pappus absent (florets epappose) (Fig. 6C) (Brazil) . . . . . . . . . . . . . . . . . . . T. cinerea
4. Lamina lanceolate, margin serrate; florets ca. 40; pappus irregular or defective
(absent in marginal florets and present in the inner) (Fig. 8C–D) (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. eupatorioides
Receptacles flat (Fig. 18D); corolla tube glabrous or sparsely covered with glandular sessile trichomes (Figs. 14C, 17C, 18C) . . . . . . . . . . . . . . . . . . . 5
5. Lower leaves opposite, decussate, larger than upper leaves; lamina 4–6 2–4 cm, ovate,
apex obtuse, margin broadly crenate, base attenuate, venation trinervate, glabrescent (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. rhodotricha
5. Lower leaves opposite, decussate, similar in size to upper leaves, or alternate and spiral or fasciculate;
lamina, venation and indumentum not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Leaves alternate, fasciculate and congested (Figs. 9A, 17F), laminae orbicular or obovate, 0.3–0.9 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Laminae orbicular, crenate on margins, rounded at base (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. harleyi
7. Laminae obovate, entire to serrulate on margins, attenuate at base (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tombadorensis
6. Lower leaves usually opposite and upper alternate, rarely fasciculate and congested (T. villosa),
laminae over 1.5 cm long (if smaller, laminae triangular or ovate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Herbs, laminae membranous, hyaline, margins irregularly deeply serrate (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. heringeri
8. Subshrubs, laminae chartaceous, subcoriaceous or coriaceous, margins entire,
serrate, crenate, dentate and not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Leaves sessile to subsessile (petiole up to 5 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Laminae narrow-elliptic, oblanceolate, linear or elliptic; margins entire, dentate or serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Laminae narrow-elliptic to oblanceolate; florets 20–25, 7–8 mm long, corolla tube glabrous;
pappus 2–3 mm long (Colombia, Venezuela, Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. arguta
11. Laminae linear to elliptic; florets 45–75, 10–12 mm long, corolla tube pubescent at the apex;
pappus 4–6 mm long (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. villosa
10. Laminae obovate, triangular to ovate; margins crenate to crenate-dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
+
1.
528
SYSTEMATIC BOTANY
+
Laminae 3.5–6(–7) 3–4.5 cm, obovate, adaxial surface non-bullate (veins not sunken);
cypselae not-stipitate (Paraguay) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. chodatii
12. Laminae 0.5–2.5 0.5–2 cm, ovate to triangular, adaxial surface bullate;
cypselae stipitate (Fig. 12D–F) (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. hirtiflora
9. Leaves petiolate (petiole > 6 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Branches griseo-puberulous; laminae subcoriaceous, oblanceolate, with
rounded apex and entire margins (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. spathulifolia
13. Branches puberulous, pubescent or tomentose; laminae chartaceous, shape ovate, lanceolate,
triangular or oblong, with obtuse to acute apex, and crenate, serrate,
dentate, or rarely entire margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Laminae ovate to lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Leaves sessile, laminae subcordate to rounded at base, 5-nervate; cypselae
not-stipitate (Paraguay) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. phlebodes
15. Leaves petiolate, laminae attenuate or truncate to subcordate at base, venation
pinnate or 3(4)-nervate; cypselae stipitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Laminae serrate on margins, attenuate at base; venation pinnate; petiole 1–2 cm long;
pappus 2–2.5 mm long (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. grazielae
16. Laminae dentate to crenate-serrulate on margins, truncate to subcordate at base; venation
3–4-nervate; petiole (2–)3–5(–7) cm long; pappus 3–4 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Laminae lanceolate to triangular, truncate and symmetric at base; florets 20–25;
anther connective appendages longer than wide (Venezuela, Colombia,
Paraguay, Bolivia, Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. salviifolia
17. Laminas ovate, truncate to subcordate and asymmetric at base; florets 60–100;
anther connective appendages as long as wide (Bolivia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. capitata
14. Laminas elliptic, narrow-elliptic or oblong (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Laminae oblong, slightly revolute at margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. laxa
18. Laminas elliptic or narrow-elliptic, planar at margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Subshrubs 0.8–1.5 m tall; laminae elliptic 4–8(–11.5) 1.2–2.5 cm, entire to
serrulate at margins; anther connective appendages longer
than wide (Fig. 2F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. attenuata
19. Subshrubs 0.3–0.6 m tall; laminae narrow-elliptic 1.5–2.5(–3) 0.2–0.5 cm,
crenate at margins; anther connective appendages wider
than long (Fig. 2C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. campestris
+
+
+
12.
[Volume 37
Aug 1949, Idrobo & Fernández 128 (US); Valle del Cauca, Cerro de las
Cruces, 21 May 1944, Killip & Cuatrecasas 38416 (US). Popayan: 16 Nov
1990, Ruiz & Rengifo 1088 (US). Santander del Norte: 28 Sep 1969,
Cuatrecasas & Rodrı́guez 27982 (US); 21 Jul 1974, Garcı́a-Barriga & Jaramillo
M. 20578 (US).
BRAZIL. Mato Grosso: Alto do Garça, 19 Dec 1992, Silva & Rodrigues
Jr. 366 (US); 18 Dec 1992, Silva & Rodrigues Jr. 612 (SPF, US); Chapada
dos Guimarães: Casa de Pedras, 2 Feb 1995, Dubs 1833 (ESA).
Erect subshrubs 0.2–1 m tall; branches scabrous,
puberulous, with glandular, sessile trichomes. Leaves alternate, spirally arranged, chartaceous, slightly discolorous,
subsessile, petioles 3–5 mm long; laminae 2–4
0.2–0.4 cm,
narrow-elliptic, acute at apex, serrate on margins, flat to revolute, cuneate at base, adaxial surface glabrescent, abaxial
surface scabrous, with glandular sessile trichomes, venation
ca. 0.8 cm;
inconspicuously pinnate. Involucres 0.5–0.8
receptacles flat; phyllaries ca. 18, 2-seriate, apex pinkish,
the outer 3–4
1 mm, oblanceolate, acute, the inner 5–6
0.5 mm, narrow-spatulate, obtuse, tomentose. Florets 20–25,
7–8 mm long, corolla 3–3.5 mm long, corolla lobes ca. 0.3 mm
long, conspicuous, corolla tube 2–2.5 mm long, glabrous;
anther connective appendages retuse, as long as wide;
styles ca. 6 mm long. Cypselae ca. 4 mm long, stipitate to
long-stipitate (1–2 mm long), setose, especially along ribs;
carpopodium annular; pappus 2–3 mm long, regular.
Figure 2A–D.
Distribution and Habitat—Trichogonia arguta is known
from Brazil (Mato Grosso), Colombia and Venezuela, in
savannas with rocky soils, between 1,000–2,500 m elevation.
The disjunction in the distribution of this species is probably
due to the lack of adequate collecting sample. Figure 3.
Notes—Trichogonia arguta is recognized by its subsessile
leaves, with narrow-elliptic lamina, serrate margins, and
stipitate to long-stipitate cypselae. Trichogonia arguta is morphologically similar to T. campestris, an endemic species from
northeastern Brazil (Tocantins, Piauı́ and Brazil), however
T. arguta can be distinguished from T. campestris by its short
petioles 0.3–0.5 mm long (vs. 1–1.5 cm long), heads with 20–
25 florets (vs. 30–40) and by the cypselae with stipe 1–2 mm
long (vs. 0.5–1 mm long).
+
+
Erect subshrubs 0.8–1.5 m tall; branches puberulous, glandular trichomes sessile and stipitate. Leaves alternate, spirally arranged, usually lower leaves opposite and similar to
upper leaves, chartaceous, discolorous, petiole 1–1.5 cm long;
laminae 4–8(–11.5) 1.5–2.5 cm, elliptic, acute at apex, entire
to serrulate on margins, attenuate to decurrent at base, adaxial surface glabrescent, abaxial surface pubescent, glandular
sessile trichomes, venation pinnate. Involucres 0.5–0.7 0.8–
1 cm; receptacles flat; phyllaries ca. 18, acute at apex, 2–
2 mm, oblong to elliptic, the inner
seriate, the outer 5–6
6–7 1 mm, spatulate, pubescent on upper third, puberulous
below, ciliate at margins, glandular trichomes sessile, sparse.
Florets 22–30, 0.9–1.2 cm long, corolla 3–3.5 mm long, corolla
lobes 0.2 mm long, inconspicuous, corolla tube 2–3 mm long,
+
Representative Specimens Examined—VENEZUELA. Bolivar: Piar,
17 km E of Canaima, 31 Aug 1983, Prance & Huber 28392 (US); Santa
Elena, 30 May 1946, Lasser 1961 (US).
COLOMBIA. Cali: 1 Dec 1947, L. A. Bermúdez & F. A. Barkley 17C841
(US); 25 Dec 1959, Cuatrecasas et al. 25715 (US). Cauca: El Ramal to Rı́o
Sucio, West Popayan, 3 Jul 1922, Pennel & Killip 8140 (US); El Tambo,
Trichogonia attenuata G. M. Barroso, Arch. Jard. Bot.
Rio de Janeiro 11: 14. 1951.—TYPE: BRAZIL. Minas
Gerais, Ituiutaba, 6 Feb 1949, A. Macedo 1598 (holotype:
RB!; isotypes: US!, NY!).
+
+
+
+
+
Trichogonia arguta (Kunth) Benth. & Hook. f. ex Klatt,
Bot. Jahrb. Syst. 8: 33. 1886. Kuhnia arguta Kunth in F. W.
H. A. von Humboldt, A. J. A. Bonpland & C. S. Kunth,
Nov. Gen. Sp. 4: (folio ed.) 82. 1818.—TYPE: COLOMBIA.
Popayan, Sep 1801, Humboldt and Bonpland 1913 (holotype: P; isotypes: B, F (digital JSTOR-image)!, Photo: US!).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
529
Fig. 2. Trichogonia arguta (Kunth) Benth. & Hook. f. ex Klatt. A. Floriferous branch. B. Floret. C. Stamen. D. Cypsela long stipitate and plumose
pappus. (Dubs 1833, ESA). Trichogonia attenuata G. M. Barroso. E. Floriferous branch. F. Stamens, retuse and rounded anther connective appendage.
G. Cypsela stipitate and pappus. (Santi and Santos F 15894, UEC).
Fig. 3.
SYSTEMATIC BOTANY
[Volume 37
Distribution of T. arguta, T. capitata and T. hassleri.
glabrous; styles ca. 5 mm long, anther connective appendages obtuse to retuse, longer than wide. Cypselae 3.5–5 mm
long, stipitate to long-stipitate (1–2 mm long), villose, glandular trichomes sessile; carpopodium annular, the ribs
decurrent into it; pappus 2–3 mm long, regular. Figure 2E–G.
Distribution and Habitat—This species is endemic to
Brazil, recorded from the states of Goiás, Minas Gerais, Mato
Grosso, and São Paulo. The species predominantly occurs in
cerrado and forest margins along the Atlantic coast of Brazil.
Figure 4.
Representative Specimens Examined—BRAZIL. Goiás: Catalão, 30 Jan
1990, Arbo et al. 3070 (HRCB). Mato Grosso: Barra do Garça, 13 Jan 1968,
Philcox & Bertoldo 3998 (UB). Minas Gerais: Belo Horizonte, Serra do
Curral, 2 Jan 1959, Magalhães s. n. (UB 20092); Buenópolis, 27 Jul 1976,
Davis et al. 2324 (UEC); Campina Verde, 6 Feb 1944, Macedo 255 (SPF);
Lavras, 2 Jan 1977, Gibbs 4109 (UEC); Paraopeba, Faz. Itapoã, 26 Jan 1999,
Silva s. n. (BHCB 45767); Passos, 2 Mar 1984, Santi & Santos Filho
15894 (UEC); Perdizes, Galheiros, 14 Feb 2003, Amorim et al. 470 (SPF);
São Sebastião do Paraı́so, Feb 1945, Vidal I-287 (R). São Paulo: Altinópolis,
Morro do Forno, 1 Mar 2004, Oliveira & Godoy 452 (SPF); Bauru, Nov
1941, Santos s. n. (R 36960); Botucatu, 21 Mar 1975, Gottsberger 6R-21375
(UB); Cajuru, 8 Feb 1990, Sciamarelli & Coffani-Nunes 490 (SPF, UEC);
Mogi-Guaçu, Fazenda Campininha, 2 Feb 1984, Noronha 310 (HRCB,
UEC); Pedregulho, Parque Estadual das Furnas do Bom Jesus, 28 Jan
1993, Macedo 75 (SPF); Piraçununga, 31 Mar 1995, Aragaki & Batalha
331 (SP); Porto Ferreira, 27 Jan 1981, Bertoni 18661 (CEPEC, INPA).
Notes—Trichogonia attenuata resembles T. grazielae, but can
be distinguished from it by the elliptic, entire to serrulate
leaves (vs. lanceolate, serrate) and by its shorter cypsela stipe
(0.2–0.5 mm long vs. 1–2 mm long).
Trichogonia campestris Gardner, London J. Bot. 5: 459.
1846.—TYPE: BRAZIL. Piauı́, near Oieiras, Mar 1839,
G. Gardner 2212 [lectotype: here designated: BM
(541026)!; isolectotypes: BM (541027)!; K! (ex Herbarium
Hookerianum); K! (ex Herbarium Benthamianum), US!,
NY-2 (digital JSTOR-image)!, GH (digital JSTOR-image)!,
Photos: RB!, SPF!].
Trichogonia pseudocampestris R. M. King & H. Rob., Phytologia
45(2): 108. 1980.—TYPE: BRAZIL. Bahia, “ca. 2 km SW
of the town of Morro do Chapeú, on the Utinga road,”
3 Mar 1977, R. M. Harley, S. J. Mayo, R. M. Storr, T. S.
Santos & R. S. Pinheiro 19320 (holotype: CEPEC!;
isotypes: IPA!, K!, NY!, RB!, US!; Photo: SPF!), syn. nov.
Trichogonia santosii R. M. King & H. Rob., Phytologia 45(2):
110. 1980.—TYPE: BRAZIL. Bahia, “1.5 km S of
Santo Inácio on Gentio do Ouro road,” 24 Feb 1977, R.
M. Harley, S. J. Mayo, R. M. Storr, T. S. Santos & R. S.
Pinheiro 18987 (holotype: CEPEC!; isotypes: IPA!, NY!,
RB!, US!), syn. nov.
Erect subshrubs 0.3–0.6 m tall; branches pubescent to
tomentose, glandular trichomes sessile and stipitate. Leaves
alternate, spirally arranged, chartaceous, discolorous, petiole
1–1.5 cm long; laminae 1.5–2.5(–3)
0.2–0.5 cm, narrowelliptic to elliptic, obtuse at apex, crenate to serrate on
+
530
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
531
Fig. 4. Distribution of T. attenuata, T. campestris and T. chodatii.
+
+
+
margins, attenuate at apex, tomentose, glandular sessile trichomes, venation trinervate. Involucres 0.8–1.2 0.6–0.8 cm;
receptacles flat; phyllaries 15–20, obtuse at apex, 2 series, the
outer 5–6
1.5 mm, oblanceolate, the inner 6–7
0.5 mm,
linear, tomentose on upper third, puberulous below, glandular trichomes sessile, sparse. Florets 30–40, 0.8–1.1 cm long,
corolla 4–5 mm long, corolla lobes 0.3 mm long, inconspicuous, corolla tube 1.5–2.5 mm long, glabrous; anther connective appendages retuse, wider than long; styles 7–8 mm long.
Cypselae 3.5–5 mm long, stipitate (0.5–1 mm long), villose,
glandular trichomes sessile; carpopodium annular; pappus
2.5–3 mm long, regular. Figure 5A–B.
Distribution and Habitat—Trichogonia campestris occurs in
the Brazilian states of Tocantins, Piauı́, and Bahia, on sandy
soils in open savanna vegetation (campos gerais, cerrado)
and less commonly in campo rupestres, caatinga or riparian
forests. Figure 4.
Representative Specimens Examined—BRAZIL. Bahia: Abaı́ra, 13 150 S,
41 450 W, 31 Jan 1992, Hind et al. H51418 (ALCB); Alagoinhas, Calu, 7 Oct
2002, Jesus 1486 (ALCB); Barra da Estiva, 19 Jul 1981, Giulietti et al. CFCR
1369 (SPF, UEC); Boquira, Torre Telebahia, 19 Jan 1997, Hatschbach et al.
66004 (CEPEC, US); Canudos, 9 550 1500 S, 39 10 5500 W, 26 Jun 2002, Queiroz
et al. 7151 (HUEFS); Gentio do Ouro, 11 020 1800 S, 42 420 4800 W, 24 Jun 1996,
Guedes et al. PCD 3040 (ALCB, CEPEC, HRB, SPF); Ibiquara, 23 Jan 1984,
Hatschbach 47538 (CEPEC, MBM, US); Irará, 8 May 1957, Lordelo 57–537
(ALCB); Jacobina, 11 050 1300 S, 40 400 2100 W, 2 Jul 1996, Hind et al. PCD 3342
(ALCB, CEPEC, HRB, SPF); João Correia, 17 Dec 2009, Guedes et al. 16887
(ALCB); Lençóis, 14 Nov 1983, Lima et al. 263 (HRB); Mirangaba, 10 460 S,
40 38 0 W, 23 Apr 1981, Fonseca 411 (HRB); Morro do Chapéu, 11 40 0 22 00 S,
41 00 0 39 00 W, trail to cidade das Pedras, 24 May 2008, Roque & Liro 1843
(ALCB, US); Mucugê, Serra do Sincorá, 13 35‘S, 41 28 0 22 00 W, 4 Dec 2003,
Roque 785 et al. (ALCB); Piatã, 13 04 0 19 00 S, 41 47 0 33 00 W, 11 Nov 1996,
Hind & Bautista PCD 4181 (ALCB, CEPEC); Remanso, 9 320 0200 S, 41 58 0 48 00
W, 23 Sep 2009, Guedes et al. 16046 (ALCB; HUEFS); Santa Brı́gida, 9 44 00 9 00
S, 38 18 0 3 00 W, 29 Nov 2009, Melo et al. 7302 (HUEFS); Seabra, 3 Feb
1981, King & Bishop 8788 (UB, US); Serrinha, road to Biritinga a Sátiro
Dias, 11 42 0 36 00 S, 38 51 0 W, 3 Sep 2006, Amorim et al. 6247 (CEPEC). Piauı́,
Caracol, PARNA Serra das Confusões, 9 13 0 23 00 S, 43 29 0 367 00 W, 22 Mar
2006, Barros et al. 2720 (HUEFS). Tocantins, estrada de São Félix do Tocantins
a Novo Acordo, 28 Mar 2011, Bringel & Moreira 740 (ALCB, UB).
Notes—Trichogonia campestris is defined by its shrubby
habit reaching 60 cm tall, lamina narrow-elliptic to oblanceolate with base attenuate and petiolate leaves (petiole 1–1.5 cm
long). Trichogonia campestris reseambles Trichogonia arguta but
can be told apart by the petiolate leaves (1–1.5 cm vs. 3–5 mm
long) and more numerous florets per head (30–40 vs 20–25).
Trichogonia campestris as well as T. arguta, may have purple
phyllaries (Gomes et al. 461) and pappus (Rodrigues 22;
Freitas 25).
Herein we are proposing two new synonyms for T.
campestris. Trichogonia pseudocampestris, based on the apically
obtuse and tomentose phyllaries, and by its short, stipitate
glandular trichomes and T. santosii based on bigger heads
and more numerous phyllaries and florets. These differences,
532
SYSTEMATIC BOTANY
[Volume 37
Fig. 5. Trichogonia campestris Gardner. A. Floriferous branch. B. Floret showing cypsela stipitate. (Guedes 9803, ALCB). T. capitata (Rusby) B. L. Rob.
C. Floriferous branch showing asymmetric leaves (Bang 2114, US). D. Cypsela (Jardim & Rosas-Hurtado 1512, US).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
533
Fig. 6. Trichogonia chodatii (Hassler) R. M. King & H. Rob. A. Floriferous branch showing obovate leaves (Drawn from the holotype). T. cinerea (Gardner)
R. M. King & H. Rob. B. Floriferous branch. C. Floret epappose. D. Receptacle convex. E. Ribs decurrent onto the carpopodium (Anderson 9997, MBM).
SYSTEMATIC BOTANY
Trichogonia capitata (Rusby) B. L. Rob. Proc. Am. Ac. 47:
193. 1911. Eupatorium capitatum Rusby, Bull. New York
Bot. Gard. 4(14): 380. 1907.—TYPE: BOLIVIA. Coripata,
Yungas, 25 Mar 1894, M. Bang 2114 (holotype: NY!;
isotypes: GH (digital JSTOR-image)!, K!, LD (digital
JSTOR-image)!, US!, MICH!, Photo: SPF!).
Trichogonia bishopii R. M. King & H. Rob., Phytologia 39: 334.
1978.—TYPE: BOLIVIA. Santa Cruz. One Km from
Comarapa on road to Cochabamba, 5 Feb 1978, R. M.
King & E. Bishop 7626 (holotype: US!; isotypes: K!, MO!,
US!; Photo: SPF!), syn. nov.
+
Trichogonia cinerea (Gardner) R. M. King & H. Rob.,
Phytologia 24: 178. 1972. Piqueria cinerea Gardner, London
J. Bot. 6: 432. 1847.—TYPE: BRAZIL. Goiás Apr 1840,
G. Gardner 3810 [lectotype, here designated: BM! (Photo:
SPF!); isolectotypes: K! (ex Herbarium Hookerianum,
Photo: SPF!); K! (ex Herbarium Benthamianum, Photo:
SPF!), NY-4 (digital JSTOR-image!).
Notes—Trichogonia capitata is easily recognized by leaves
with ovate lamina, with crenate-serrulate margins, truncate
and slightly subcordate and asymmetrical base, grayishgreen abaxial surface and petioles (1.5–)3–5(–7) cm long.
The high number of florets per head (60–100) and the
glabrescent, long-stipitate cypselae, are also diagnostic characters for the species.
Trichogonia bishopii is here proposed as a synonym of
T. capitata as they are morphological identical.
Trichogonia chodatii (Hassler) R. M. King & H. Rob.,
Phytologia 24: 178. 1972. Eupatorium chodatii Hassler,
Repert. Spec. Nov. Regni Veg. 11: 169. 1912.—TYPE:
PARAGUAY. Caaguazú, Mar 1905, E. Hassler 9087 (holotype: G; isotype: K!; BM (digital JSTOR-image)!, Photos:
SPF!, US!).
+
Erect subshrubs 0.3–0.5 m tall; branches puberulous. Leaves
alternate, spirally arranged, usually lower leaves opposite
and similar to upper leaves, chartaceous, discolorous, sub3–4.5 cm,
sessile, petiole ca. 0.5 cm long; laminae 3.5–6(–7)
+
+
Erect subshrubs 0.5–1.2 m tall; branches tomentose, glandular trichomes stipitate and sessile, sticky. Leaves alternate,
spirally arranged, usually lower leaves opposite and similar
to upper leaves, chartaceous, discolorous, petiole 0.5–1 cm
0.6–1(–1.5) cm, oblong to elliptic,
long; laminae (2–)3–4.5
obtuse at apex, crenate on margins, decurrent at base, adaxial
surface pubescent to glabrescent, abaxial surface tomentose
to puberulous, venation trinervate. Involucres 6–9 6–7 mm;
receptacles convex; phyllaries 23–32, 2-seriate, the outer ca.
4 1.5 mm, elliptic, apex acute, the inner ca. 5 1 mm, linear,
apex acute, margin hyaline, tomentose to puberulous, glandular trichomes stipitate and sessile. Florets 60–100, 9–10 mm
long, corolla 3–4 mm long, corolla lobes ca. 0.3 mm long,
inconspicuous, corolla tube 1.5–2 mm long, pilose and with
glandular trichomes stipitate; anther connective appendages
obtuse to retuse, as long as wide, styles ca. 5–6 mm long.
Cypselae 2–3 mm long, short-stipitate (ca. 0.3 mm long), glabrous; carpopodium annular or the ribs decurrent into it;
pappus absent. Figure 6B-E.
Distribution and Habitat—Trichogonia cinerea is endemic
to Brazil (Goiás, Bahia, and Distrito Federal); it has been
collected in areas of cerrado (savanna woodland) and campo
rupestre. Figure 7.
+
All Specimens Examined—BOLIVIA. Cochabamba: Prov. Campero,
9 Jan 1994, Saraiva & López ES 906 (US). Santa Cruz, 1 km of Comarapa, 5 Feb
1978, King & Bishop 7630 (US), Prov. Cordillera, 29 Dec 1994, Jardim & RosasHurtado 1512 (US), Prov. Florida, 8 Mar 1988, Solomon & Nee 18034 (US),
18 Apr 2002, Nee et al. 52199 (US). La Paz, Yungas, Dec 1917, Buchtien 4761 (US).
Trichogonia dubia (B. L. Rob.) R. M. King & H. Rob.,
Phytologia 24: 178. 1972. Alomia dubia B. L. Rob., Proc.
Amer. Acad. Arts 42: 33. 1907.—TYPE: BRAZIL. Goiás,
1896, A. Glaziou 21579 (holotype: K!, Photos: F!, SPF!,
US!), syn. nov.
+
+
+
+
+
Erect subshrubs 0.8–2.5 m tall; branches aromatic, pubescent, glandular trichomes stipitate, sticky. Leaves alternate,
spirally arranged, usually lower leaves opposite and similar
to upper leaves, chartaceous, discolorous, petioles (1.5–)3–5
(–7) cm long; laminae (3–)6–10(–11) (1.5–)3–6(–7) cm, ovate,
acute to obtuse at apex, crenate-serrulate on margins, truncate to slightly subcordate and asymmetric at base, adaxial
surface puberulous, abaxial surface grayish-pubescent, with
glandular sessile trichomes, venation trinervate. Involucres
0.8–1 1–1.4 cm, receptacles flat; phyllaries 20–30, 2–3 seriate, the outer ca. 7 1.2 mm, spatulate, acute to acuminate
at apex, tomentose, the inner linear, acuminate at apex, 6–
7 1 mm, pubescent. Florets 60–100, 0.9–1.2 cm long, corolla
5–6 mm long, corolla lobes 0.5 mm long, conspicuous,
corolla tube 2.5–3 mm long, glabrous; anther connective
appendages obtuse to retuse, as long as wide, styles 4–6 mm
long. Cypselae 5–6 mm long, puberulous to glabrescent on
the ribs, long-stipitate (2–3 mm long); carpopodium annular;
pappus 3.5–4 mm long. Figure 5C-D.
Distribution and Habitat—Trichogonia capitata is known
only from Bolivia in xerophytic vegetations, mostly on rocky
outcrops and rocky soils. Figure 3.
obovate, obtuse at apex, upper half crenate-dentate on
margins, attenuate at base, scabrous in the veins, venation
trinervate. Involucres 1–1.5 1.5–2 cm, receptacles flat; phyllaries ca. 18, 2-seriate, acute at apex, the outer 1–1.2 0.8 cm,
lanceolate, the inner ca. 1 0.5 mm, linear, margin hyaline
and ciliate, glandular trichomes sessile. Florets ca. 30, 6–7 mm
long; corolla 4–6 mm long, corolla lobes 0.5 mm long, conspicuous, corolla tube 2–2.5 mm, glabrous; anther connective
appendages obtuse, as long as wide, styles ca. 5 mm long.
Cypselae 4–6 mm long, puberulous, especially along ribs,
glandular trichomes sessile; carpopodium annular; pappus
3–4 mm long, regular. Figure 6A.
Distribution and Habitat—Trichogonia chodatii is endemic
to the highlands of Amambay and Caaguazú, Paraguay. The
species is known to flower from February to April (Cabrera
1996). Figure 4.
Notes—Trichogonia chodatii is distinguished from other
species of Trichogonia by its subsessile, obovate leaves with
obtuse apex and crenate-dentate margins on distal half.
+
however, fall within the range of a continuous variation
observed and therefore cannot be used to recognize species in Trichogonia. Thus, T. pseudocampestris and T. santosii
are included in the same specific concept of T. campestris.
[Volume 37
+
534
Representative Specimens Examined—BRAZIL. Bahia: Riachão das
Neves: 13 460 4900 S, 44 540 3900 W, 600 m, 6 Apr 2005, Carvalho-Sobrinho et al.
449 (ALCB, UEFS). Brası́lia: Barracão, Árie do Capetinga, 8 Jun 1988, Silva
694 (IBGE); Chapada da Contagem, 18 Mar 1998, Calago 92 (CEN). Goiás:
Alto Paraı́so, Chapada dos Veadeiros, 7 Feb 1981, King & Bishop 8857 (UB,
US); Cavalcante, caminho para Vão do Moleque, 21 Mai 2011, Bringel &
Pastore 772 (ALCB, UB); Corumbá de Goiás, 15 May 1973, Anderson 10333
(INPA, MG, US); Cristalina, 15 Apr 1965, Heringer 10139 (UB); Formosa,
29 Apr 1966, Irwin et al. 15483 (US); Goiás Velho, Serra Dourada, 10 May
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
535
Fig. 7. Distribution of T. cinerea, T. eupatorioides, T. grazielae and T. harleyi.
1973, Anderson 9997 (INPA, MBM, R, US); Niquelândia, 14 090 S, 48 130 W,
14 Apr 1992, Walter et al. 1313 (SPF); Pirenópolis, s/data, Irnaldo et al. s. n.
(HB 45096); São João da Aliança, 22 Mar 1973, Anderson 7640 (UB, US).
Notes—Trichogonia cinerea is characterized by oblong to
elliptic and puberulous leaves, with crenate margins and
attenuate base, high number of florets per head (60–100),
glabrous cypsela and epappose florets.
Trichogonia cinerea belongs to the Trichogonia group which
has convex receptacles and corolla tubes with glandular
stipitate trichomes. Trichogonia cinerea seems to be close to
T. eupatorioides as they both have a defective pappus or lack
them altogether. Trichogonia cinerea can be distinguished
from T. eupatorioides by the oblong to elliptic (vs. lanceolate),
60–100 florest per head (vs. 40) and by pappus absent in all
florets (vs. absent at least at marginal florets).
King and Robinson (1972) distinguished Trichogonia dubia
by the epappose florets and the high number of florets per
head (ca. 65), characters that also are found in T. cinerea.
Thus, we are proposing T. dubia as a synonym of T. cinerea.
Trichogonia eupatorioides (Gardner) R. M. King & H.
Rob., Phytologia 45: 106. 1980. Isocarpha eupatorioides
Gardner, London J. Bot. 5: 456. 1846. Piqueria eupatorioides
(Gardner) Gardner, London J. Bot. 6: 431. 1847.
Trichogonia salviifolia Gardner var. calva Baker, in C. F. P.
von Martius, Fl. Bras. 6(2): 217. 1876, nom. illeg..
Trichogonia menthifolia Gardner var. calva (Baker) Mattf.,
Notizbl. Berlin 8: 448. 1923, nom. illeg.—TYPE: BRAZIL.
Minas Gerais, in moist open places, near San Romão, Jul
1840, G. Gardner 4839 [lectotype, here designated: BM!
(Photo: SPF!); isolectotypes: K! (ex Herbarium
Hookerianum, NY-3!, RB!, Photos SPF!, US!].
Trichogonia menthifolia Gardner, London J. Bot. 6: 434. 1847.
Trichogonia menthifolia Gardner var. genuina Hassler,
Repert. Spec. Nov. Regni Veg. 14: 291. 1916.—TYPE:
BRAZIL. Goiás, campos near Capella da Passe, May 1840,
Gardner 4226. [lectotype, here designated: BM! (Photo:
SPF!); isolectotypes: K-2! (ex Herbarium Hookerianum,
Photos: SPF-2!, US!), K! (ex Herbarium Benthamianum,
Photo: SPF!), NY-3!; Photo: RB!], syn. nov.
Trichogonia martii Baker, in C. F. P. von Martius, Fl. Bras. 6(2):
216. 1876. Trichogonia menthifolia Gardner var. martii
(Baker) Hassler. Repert. Spec. Nov. Regni Veg. 14: 291.
1916.—TYPE: BRAZIL. Minas Gerais, Ouro Preto,
Martius s. n. (holotype: M; Photos: RB!, US!), syn. nov.
Trichogonia zehntneri Mattf., Notizbl. Bot. Gart. Berlin-Dahlem
8: 446. 1924.—TYPE: BRAZIL. Bahia, Sento Sé, Ago 1912,
L. Zehntner 1914 (holotype: RB!; Photo: RB!), syn. nov.
Erect herbs to subshrubs to 0.5 m tall; branches densetomentose to puberulous, with glandular trichomes stipitate.
Leaves alternate, spirally arranged, usually lower leaves opposite and similar to upper leaves, chartaceous, discolorous,
536
SYSTEMATIC BOTANY
[Volume 37
Fig. 8. Trichogonia eupatorioides (Gardner) R. M. King & H. Rob. A. Floriferous branch. B. Corolla and style branches. C. Cypsela of marginal floret,
epappose. D. Cypsela of inner florets (Anderson et al. 37154, R). Trichogonia grazielae R. M. King & H. Rob. E. Floriferous branch. F. Floret. G. Stamens
(Anderson 12490, MBM).
ROQUE ET AL.: REVISION OF TRICHOGONIA
Trichogonia grazielae R. M. King & H. Rob., Phytologia
45: 106. 1980.—TYPE: BRAZIL. Goiás, Chapada dos
Veadeiros, 6–7 Km E of Alto Paraı́so on road to Nova
Roma, 7 Mar 1973, W. R. Anderson, M. T. Catlin Arroyo,
S. R. Hill, R. Reis dos Santos & R. Souza 6584 (holotype:
UB!, Photos: K!, SPF!; isotypes: MG!, US!).
Erect subshrubs 0.5–1.5 m tall; branches puberulous with
glandular trichomes stipitate. Leaves alternate, spirally
arranged, usually lower leaves opposite and similar to upper
+
+
+
+
All Specimens Examined—BRAZIL. Goiás: Alto Paraı́so, Chapada
dos Veadeiros, 13 Jun 1993, Hatschbach et al. 59492 (MBM, US); São
João da Aliança, 23 Jan 1980, King & Almeda 8271 (K, Photos SPF, UB,
US). Minas Gerais: Carrancas, 9 Dec 1983, Leitão Filho et al. 15384
(UEC); Ouro Preto, 12 Jan 1972, Macedo 5129 (MBM); Poços de Caldas,
26 Jan 1919, Hoehne 2917 (SP); Rio Bonito, Braçanã, 8 Dec 1974, Laclette
894 (R).
Notes—Trichogonia grazielae is characterized by large (5–
9 cm long), lanceolate leaves with serrate margins and attenuate bases. Trichogonia grazielae looks superficially similar to
T. salviifolia, a widely distributed species, but differs from the
latter by the leaves with attenuate base (vs. truncate), shorter
petioles 1–1.5 cm long (vs. 2–4 cm), 37–50 florets per head
(vs. 20–25) and pappus 2–2.5 mm long (vs. 3–4 mm long).
Trichogonia harleyi R. M. King & H. Rob., Phytologia 39:
335. 1978.—TYPE: BRAZIL. Bahia, Eduardo Magalhães,
16 km NW of Lagoinha on side road to Minas do
Mimoso, 8 Mar 1974, R. M. Harley, S. A. Renvoize, C. M.
Erskine, C. A. Brighton & R. Pinheiro 16977 (holotype: US!;
isotypes: CEPEC!, K!, Photo: SPF!).
Erect subshrubs ca. 1.5 m tall; branches tomentose, glandular trichomes sessile. Leaves alternate, fasciculate, congested,
subcoriaceous, concolorous, petiole 2–4 mm long; laminae
0.2–0.3 cm, orbicular, obtuse at apex, crenate on
0.3–0.4
margins, rounded at base, tomentose, with glandular sessile
trichomes. Involucres 0.7–1 0.7–1 cm, receptacles flat; phyllaries ca. 14, 2-seriate, the outer ca. 4
1 mm, elliptic, the
inner ca. 5 0.5 mm, linear, pubescent on the 1/3 upper and
puberulous below, with sessile glandular trichomes. Florets
20–23, ca. 8 mm long; corolla 3–3.5 mm long, corolla lobes
0.2 mm long, inconspicuous, corolla tube 1.5–2.5 mm long;
anther connective appendages obtuse, longer than wide;
style 5–6 mm long. Cypselae 4.5–5 mm long, short-stipitate
(base 0.5–1 mm long), setose, denser on the ribs, glandular
trichomes sessile; carpopodium annular, ribs decurrent on to
it; pappus 3–4 mm long, regular. Figure 9A–B.
Distribution and Habitat—Trichogonia harleyi is known
only from the type collection in cerrado area in the state of
Bahia, Brazil. Figure 7.
Notes—Trichogonia harleyi and T. tombadorensis, both endemic
to Bahia, are the only species in the genus with alternate, fasciculate, and congested leaves with laminas 3–9 mm long. However,
T. harleyi differs from T. tombadorensis by the smaller (3–4 mm vs.
4–9 mm) orbicular, and crenate leaves (vs. obovate and entire).
+
Notes—Gardner (1846) described Isocarpha eupatorioides as
having a paleaceous receptacle, a mistake he corrected in
1847, by transferring the species to Piqueria and by characterizing the receptacle as glabrous (nudum). Baker (1876)
described Trichogonia salviifolia Gardner var. calva based on
the same type. This variety was later transferred by Mattfeld
as a variety of Trichogonia menthifolia. King and Robinson
(1980) transferred Isocarpha eupatorioides into Trichogonia, but
later in 1987, they placed T. eupatorioides as a synonym of
T. menthifolia. Trichogonia eupatorioides however, is based on
an older epithet and should have priority over T. menthifolia.
In the present work, T. martii Baker and T. zehntneri Mattf.,
are reduced to synonymy of T. eupatorioides, as they all have
the diagnostic character of a defective pappus, meaning that
the pappus is absent at least in the outer florets.
Trichogonia eupatorioides and T. cinerea share the presence of
corolla tube with glandular stipitate trichomes and defective
and epappose pappus. However, Trichogonia eupatorioides can
be distinguished from the latter by its lanceolate glabrescent
lamina with serrate margins (vs. oblong or elliptic, pubescent, with crenate margins), and by the inner florets with
pappus (vs. lacking pappus).
leaves, chartaceous, discolorous, petiole 1–1.5(–2) cm long;
laminae 5–9(–11)
(0.6–)1–2 cm, lanceolate, acute at apex,
serrate on margins, attenuate at base, adaxial surface
glabrescent, abaxial surface puberulous, with glandular sessile trichomes, venation pinnate. Involucres 0.7–0.8 0.8 cm;
receptacles flat; phyllaries ca. 16, 2-seriate, the outer ca. 4
1 mm, lanceolate, the inner ca. 4 0.5 mm, linear, acuminate
to acute and tomentose at the apex, glandular trichomes
sessile and stipitate. Florets 37–50, 6–8 mm long; corolla
3–3.5 mm long, corolla lobes 0.3 mm long, inconspicuous,
corolla tube 1.5–2 mm long; anther connective appendages
obtuse to retuse, longer than wide, style 5–6 mm long.
Cypselae 2–3 mm long, puberulous, setose along ribs, shortstipitate (base 0.2–0.5 mm long); carpopodium annular; pappus
2–2.5 mm long, regular. Figure 8E–G.
Distribution and Habitat—Trichogonia grazielae is endemic
to Brazil and has been collected only in the states of Goiás
and Minas Gerais, mostly in cerrado (savanna woodland)
and campos rupestres areas at 1,100 m elevation. Figure 7.
+
Representative Specimens Examined—BRAZIL. Bahia: Bahianópolis,
24 Jul 1963, Costa s. n. (ALCB 4420); Cocos, Espigão Mestre, 15 Mar
1972, Anderson et al. 37010 (UB, US); Correntina, 13 20 0 S, 44 38 0 W,
23 Apr 1980, Harley et al. 21625 (CEPEC); Feira da Mata, Road
Cocos/Feira da Mata, 14 15 0 400 S, 44 22 0 10 00 W, 17 Apr 2001, Jardim
et al. 3594 (ALCB, CEPEC); Formosa do Rio Preto, 23 Feb 2005, Xavier
& Guedes 46 (ALCB); Eduardo Magalhães, Serra do Mimoso, 8 Jun
1912, Lutz 168 (R); São Desidério, 12 360 S, 44 51 0 W, 1 Jul 2001, Alves
et al. 254 (ALCB); Tabocas do Brejo Velho, Serra Dourada, 12 39 0 S,
44 02 0 W, 1 May 1980, Harley et al. 21995 (CEPEC, IPA, US). Ceará:
Baturité, Serra do Baturité, Aug 1937, Eugênio 12 (RB). Goiás: Apr 1840,
Gardner 3818 (K). Minas Gerais: Januária, distrito de Fabião, 15 070 8500 S,
44 150 1700 W, 25 May 1997, Lombardi & Salino 1792 (US); Montalvânia,
18 Mar 1972, Anderson et al. 37154 (INPA, R, US).
537
+
+
+
+
+
petiole 0.7–1 cm long; laminae (2–)2.5–6(–7) (0.5–)1–2.5
(–3) cm, lanceolate, acute at apex, serrate on margins, attenuate at base, adaxial surface glabrescent, abaxial surface
puberulous, venation trinervate. Involucres ca. 0.6 0.5 cm,
receptacles convex; phyllaries ca. 17, 2-seriate, acute at
apex, the outer ca. 4 1 mm, oblong to elliptic, the inner ca.
4 1 mm, linear to spatulate, pubescent, ciliate, with glandular sessile trichomes. Florets 40–57, 5 –7 mm long; corolla
3– 4 mm long, corolla lobes 0.2–0.5 mm long, inconspicuous, corolla tube 1–1.5 mm long, with glandular stipitate
trichomes; anther connective appendages acute, longer than
wide, styles 4–5 mm long. Cypselae 1–2 mm long, setose on
the inner florets, and cypselae glabrous, not or short-stipitate
(ca. 0.1 mm long) on the outer florets; carpopodium annular,
the ribs decurrent into it; pappus 2–2.5 mm long, defective,
absent on the outer florets. Figure 8A–D.
Distribution and Habitat—Trichogonia eupatorioides is
known from the Brazilian states of Minas Gerais, Goiás,
Bahia, and Ceará in cerrado (savanna woodland), caatinga
and open disturbed vegetation. Figure 7.
+
2012]
538
SYSTEMATIC BOTANY
[Volume 37
Fig. 9. Trichogonia harleyi R. M. King & H. Rob. A. Floriferous branch. B. Cypsela (Drawn from the holotype). Trichogonia hassleri Mattf. C.
Floriferous branch. D. Floret with cypsela cylindric. E. Corolla with trichomes on the limb and tube. F. Detail of the pedicellate trichomes on the
corolla tube (T. Rojas 9893, G).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
539
Fig. 10. Trichogonia heringeri R. M. King & H. Rob. A. Floriferous branch. B. Floret. C. Cypsela stipitade (Moreira s. n., ALCB 82230). Trichogonia laxa
Gardner. D. Floriferous branch. E. Corolla and style branches. F. Cypsela (Rezende et al. 105, UB).
SYSTEMATIC BOTANY
Trichogonia menthifolia Gardner var. tomentosula Hassler f.
cordata Hassler, Repert. Spec. Nov. Regni Veg. 14: 291.
1916.—TYPE: PARAGUAY. Amambay: “Sierra de
Amambay in campis siccis Punta Porá,” Dec 1907–1908,
E. Hassler & T. Rojas 9893 (holotype: G [00306013]!;
isotypes: BAF, G [0306014]!, G [00306015]!, K!, LIL,
Photo: SPF!, US!).
[synonyms proposed by Mattfeld 1923]
+
+
+
+
Erect subshrubs ca. 1 m tall; branches griseo-puberulous.
Leaves alternate, spirally arranged, usually lower leaves opposite and similar to upper leaves, chartaceous, discolorous, petiole 0.5–1 cm long; laminae 2.5–4.5(–5) 1–2.5 cm long, ovate,
acute to obtuse at apex, dentate on margins, cordate at base,
adaxial surface puberulous, abaxial surface griseo-tomentose
with glandular sessile trichomes, venation trinervate. Involucres 5–7
6–8 cm; phyllaries 16–20, spatulate, rounded at
1.5–2 cm, tomentose, the
apex, 2-seriate, the outer 4.5–5
inner 5.5–6
1 cm, tomentose on upper third; receptacles
convex. Florets 25–30, 6–8 mm long, corolla 4–5 mm long,
corolla lobes 0.5 mm long, conspicuous, corolla tube 1.5–3 mm
long with stipitate glandular trichomes; anther connective
appendages longer than wide; style ca. 6 mm long. Cypselae
2–3 mm long, cylindrical, setose; carpopodium annular; pappus 3–4 mm long, regular. Figure 9C–F.
Distribution and Habit—The species is known only from
the state of Mato Grosso do Sul state (Brazil) and Paraguay,
occurring mostly in cerrado areas. According to Cabrera
(1996), T. hassleri blooms in spring and summer. Figure 3.
All Specimens Examined—BRAZIL. Mato Grosso do Sul, Estação
de Bodoquena, Dec 1941, s/ collector (R 40.867); Bela Vista, Road
MS-472, Rod. BR-267 to Bela Vista, 11 Mar 2003, Hatschbach et al.
74603 (HUEFS, US); Miranda, Fazenda Lagoão Fechado, 18 Mar 2003,
Hatschbach et al. 74897 (US).
PARAGUAY. Alto Paraná: Estancia Bello Horizonte, 17 Sep 1993, Zardini &
Guerrero 37281 (US); Caaguazú, Dec 1898–1899, Hassler 5780 (G–3, NY);
Concepción, Estancia Santa Marı́a, 16 Mar 1994, Zardini & Guerrero
38960 (US).
Notes—Mattfeld (1923) considered Trichogonia menthifolia
var. tomentosula Hassler and T. menthifolia var. martii Hassler
to be the same taxon and decided to elevate it to the species
level, providing the name Trichogonia hassleri. As a result,
T. hassleri is based on several syntypes (Hassler 8739, 9893,
and 11023), and therefore a lectotype was selected by Roque
et al. (2011).
Trichogonia heringeri R. M. King & H. Rob., Phytologia
45: 107. 1980.—TYPE: BRAZIL. Pernambuco, between
Petrolina and Afrânio, 19 Apr 1971, E. P. Heringer, D. A.
Lima, J. P. L. Sobrinho, A. C. Sarmento 165 (holotype: UB!;
isotype: RB!, Photos: SPF!, US!).
Erect herbs 0.3–0.6 m tall; branches pubescent. Leaves
alternate, spirally arranged, membranaceous, hyaline,
discolorous, petiole 1.5–2(–2.5) cm long; laminae 3–4.5(6–)
(–0.5)1–2 cm, elliptic, obtuse at apex, irregularly deeply serrate on margins, attenuate at base, puberulous, glabrescent
with glandular sessile trichomes. Involucres ca. 0.5 0.9 cm;
phyllaries 17, acuminate to acute at apex, 2-seriate, the outer
4–5 1 mm, oblanceolate to linear, the inner 4–5 0.5 mm,
linear, pubescent, with glandular sessile trichomes; receptacles flat. Florets 20–32, 9–10 mm long; corolla 3–3.5 mm long,
corolla lobes 0.2 mm long, inconspicuous, corolla tube 2–3 mm
long, with glandular sessile trichomes; anther connective
appendages obtuse to retuse, wider than long or as long
as wide; styles ca. 6 mm long. Cypselae 4–4.5 mm long,
stipitate (base 1–1.5 mm long), setose, especially on the ribs;
carpopodium annular; pappus 2–2.5 mm long, regular.
Figure 10A–C.
Distribution and Habitat—Trichogonia heringeri is endemic
to the Brazilian states of Pernambuco and Bahia, occurring
mostly on sandy soils in caatinga. Figure 11.
+
Trichogonia menthifolia Gardner var. tomentosula Hassler f.
atenuatta Hassler, Repert. Spec. Nov. Regni Veg. 14: 291.
1916.—TYPE: PARAGUAY. Guairá, “Prope Villarica in
campis humidis,” Jan.1905, Hassler 8739 (lectotype by
Roque et al. 2011: G [00306019]!; isolectotypes: G
[00306020]!, G [00306021]!, K!, Photo: SPF!).
Hassler (1916) described Trichogonia menthifolia var.
tomentosula f. atenuatta based on two syntypes, (Hassler 8739
and Hassler 5780), thus making it necessary to select a lectotype for this name. As Matffeld (1923) didn’t see ‘Hassler
5780’, ‘Hassler 8739’ was choosen as the lectotype of this
infraspecific name (Roque et al. 2011).
Trichogonia hassleri belongs to a small group in Trichogonia
(including T. cinerea, T. eupatorioides, and T. prancei) characterized by convex receptacle and corolla tube with glandular
stipitate trichomes. Of all these species, T. hassleri and
T. prancei are the only one with regular pappus. Trichogonia
hassleri is distinctive from the latter mainly by leaves ovate
(vs. elliptic), florets 25–30 (vs. 60–80), cypsela cylindrical
(vs. short-stipitate) and pappus 3–4 mm long (vs. 0.6 mm long).
+
Trichogonia menthifolia Gardner var. tomentosula Hassler in
Repert. Spec. Nov. Regni. Veg. 14: 291. 1916.—TYPE:
PARAGUAY. Amambay: “Sierra de Amambay in
campis siccis Punta Porá, Dec 1907–1908, E. Hassler & T.
Rojas 9893 (lectotype by Roque et al. 2011: G [00306013]!;
isolectotypes: BAF, G [00306014]!, G [00306015]!, K!, LIL,
Photo: SPF!,US!).
[Volume 37
+
Trichogonia hassleri Mattf., Notizbl. Bot. Gart. BerlinDahlem 8: 449. 1923.—TYPE: PARAGUAY. Amambay:
“Sierra de Amambay in colle Cerro Margarita”, Mar
1907–1908, E. Hassler & T. Rojas 11023 (lectotype by
Roque et al. 2011: G [003006009]!; isolectotypes: BAF, K!,
Photo: SPF!).
+
540
Representative Specimens Examined—BRAZIL. Bahia: Canudos,
9 56 0 3400 S, 38 59 0 17 00 W, 27 Apr 2007, Ferreira et al. 240 (HUEFS);
Jeremoabo, 14 May 1981, Gonçalves 53 (ALCB, HRB); Livramento de
Brumado, 20 Mar 1980, Mori & Benton 13495 (CEPEC, US); Maniçoba,
Serra da Tiririca, s. d., s. collector (R 48840); Mundo Novo, 8 Feb
1970, Costa et al. (ALCB 04421); Pilão Arcado, 10 000 S, 42 300 W, Apr 2007,
Moreira s. n. (ALCB-82230); Remanso, 9 380 4400 S, 42 140 500 W, 27 Feb 2000,
Fonseca 1289 (ALCB, HRB, SPF); Vitória da Conquista, May 1954, Vidal IV1057 (R). Pernambuco: Afrânio, 08 490 S, 40 490 W, 1 Jun 1984, Salgado et al.
393 (ALCB, CEPEC, HRB-2, MBM, US); Ibimirim, 2 Jun 1982, Ataı́de et al.
13 (SPF); Parnamirim, 21 Jun 1984, Araújo 172 (SPF); Petrolina, Apr 1954,
Vidal 922 (R), 18 Apr 1971, Heringer et al. 123 (UB, Photo US).
Notes—Trichogonia heringeri is easily recognized by its
elliptic, membranaceous, hyaline, glabrescent leaves with
deeply irregularly serrate margins. In dried collections,
T. heringeri is distinctive by its green leaves and light
brown stems.
Trichogonia hirtiflora (DC.) Sch. Bip. ex Baker in C. F. P.
von Martius, Fl. Bras. 6(2): 214. 1876. Kuhnia hirtiflora
DC., Prodr. 5: 127. 1836.—TYPE: BRAZIL. Minas Gerais.
s. d., Gaudichaud (holotype: P; Photos: RB!, US!).
Trichogonia apparicioi G. M. Barroso, Arch. Jard. Bot. Rio de
Janeiro 11: 14. 1951.—TYPE: BRAZIL. Serra do
Cipó, 8 Dec 1949, A. P. Duarte 2308 (holotype: RB!).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
541
Fig. 11. Distribution of T. heringeri, T. hirtiflora, T. laxa, T. phlebodes and T. prancei.
[synonym proposed by Hind (1993) to T. crenulata (Gardner)
D. J. N. Hind].
Trichogonia margarethiae Soares Nunes, Bradea 3(18): 131.
1981.—TYPE: BRAZIL. Minas Gerais, Grão Mogol, Serra
do Jambreiro, 10 May 1979, H. C. de Lima et al. 992 (holotype: RB!; isotypes: RB-2!). [synonym proposed by Hind
(1993) to T. crenulata (Gardner) D. J. N. Hind].
+
+
+
+
Trichogonia crenulata (Gardner) D. J. N. Hind, Kew Bulletin
48(2): 411. 1993. Eupatorium crenulatum Gardner, London J.
Bot. 5: 474. 1846.—TYPE: BRAZIL. Minas Gerais, Elevated
rocky places in the Diamond District, Jul 1840, G. Gardner
4850 (lectotype, here designated: BM!, Photo: SPF!;
isolectotypes: K–2!, Photo: SPF!), syn. nov.
Erect subshrubs 0.5–1 m tall; branches tomentose to
puberulous, glandular trichomes stipitate or sessile. Leaves
alternate, spirally arranged, subcoriaceous, discolorous, petiole 1–4 mm long; laminae 0.5–2(–2.5) 0.5–1.5(–2) cm, ovate
to triangular, acute at apex, crenate on margins, cordate to
truncate at base, adaxial surface bullate (sunken veins),
glabrescent, abaxial surface with prominent veins, velutinous,
with glandular sessile trichomes, venation trinervate. Involu1–1.5 cm; receptacles flat; phyllaries 13–16,
cres 1–1.2
2 mm, oblanceolate, acute at apex,
2-seriate, the outer 5–6
1 mm, spatulate to oblanceolate, acute to
the inner ca. 6
obtuse at apex, pubescent, ciliate on apex. Florets 30–55,
0.9–1.2 cm long; corolla 4–5 mm long, corolla lobes 0.2 mm
long, conspicuous, corolla tube 2–3 mm long, glabrous or
with sessile glandular trichomes; anther connective appendages obtuse to retuse, longer than wide or as long as wide;
styles 6–7 mm long. Cypselae 4–5 mm long, short to
stipitate (base 0.5–1 mm long), setose, especially on the
ribs, glandular trichomes sessile; carpopodium annular;
pappus 3–4 mm long, regular. Figure 12A–F.
Distribution and Habitat—Trichogonia hirtiflora is
restricted to the state of Minas Gerais, Brazil, occurring in
cerrado and campos rupestres vegetation, mainly on sandyrocky soils. The species was cited by Baker (1876) as occurring in Espı́rito Santo and Rio de Janeiro in addition to
Minas Gerais, based on misidentifications of Trichogoniopsis
podocarpa (DC.) R. M. King & H. Rob.). In addition, there is a
collection without date or collector at R (R-48827) that refers
to T. hirtiflora and whose origin is indicated as from São
Paulo without specific locality. We view this record as a
locality mistake since the species is not otherwise known
outside the state of Minas Gerais. Figure 11.
Representative Specimens Examined—BRAZIL. Minas Gerais: Água
Limpa, Serra do Curral, 27 Jul 1956, Barroso 1646 (SPF); Belo
Horizonte, Serra do Caraça, 18 Nov 1978, Cruz et al. 6415 (UEC);
Botumirim, Serra da Canastra, road to Rio do Peixe, 16 53 0 42 00 S,
42 590 3000 W, 18 Nov 2007, Saavedra et al. 554, (SPF); Brumadinho, Serra da
Calçada, 27 Oct 2009, Cardoso et al. 2727 (HUEFS); Caeté, Serra da
Piedade, 19 Jan 1971, Irwin et al. 28719 (US); Buenópolis, Serra da Cabral,
542
SYSTEMATIC BOTANY
[Volume 37
Fig. 12. Trichogonia hirtiflora (DC.) Sch. Bip. ex Baker. A. Floriferous branch showing the smaller leaves (Hatchbach et al. 41697, MBM). B. Floriferous
branch showing the bigger leaves (Semir & Sazima 608, UEC). C. Corolla with style branches. D. Floret. E-F. Cypselae with pappus (Semir & Sazima 608, UEC).
ROQUE ET AL.: REVISION OF TRICHOGONIA
Trichogonia laxa Gardner, London J. Bot. 6: 435. 1847.—
TYPE: BRAZIL. Goiás, near Capella da Passe, May 1840,
G. Gardner 4225 [lectotype, here designated: BM!
(Photos: SPF!, US!); isolectotypes: K! (ex Herbarium
Hookerianum, Photo: SPF!), K! (ex Herbarium
Benthamianum, Photo: SPF!), RB!, NY (digital JSTORimage)!].
Notes—Trichogonia laxa resembles T. spathulifolia but differs by its oblong (vs. oblanceolate), crenate and slightly
revolute margins (vs. plane, not revolute margins) and
tomentose (vs. glabrous) lamina.
Trichogonia phlebodes (B. L. Rob.) R. M. King & H. Rob.,
Phytologia 24: 178. 1972. Eupatorium phlebodes B. L. Rob.,
Contr. Gray Herb. 100: 16. 1932.—TYPE: PARAGUAY.
Cordillera del Amambay, Sierra de Amambay, Feb 1908,
E. Hassler & T. Rojas 10178c (pro parte) (holotype: BM;
isotype: G, GH (digital JSTOR-image)!, Photo: US!).
Erect subshrubs 0.7–1 m tall; branches puberulous and
setose with sessile glandular trichomes. Leaves alternate, spirally arranged, usually lower leaves opposite and similar to
upper leaves, coriaceous, discolorous, sessile; laminae 6–
7.5 3–5.5 cm, broadly ovate, acute at apex, crenate-serrate
on margins, subcordate to rounded at base, scabrous, venation 5-nerved. Involucres 1–1.5 0.8–1.5 cm; phyllaries ca. 20,
2-seriate, subequal, 10–12 2–3 mm, oblanceolate, acute, base
glandular-puberulous. Florets ca. 40, corolla 5 mm long,
corolla lobes 0.1 mm long, conspicuous, corolla tube 2–
2.5 mm long, glabrous; styles 9–10 mm long. Cypselae 5.5–
6 mm long, cylindrical, pubescent; carpopodium annular;
pappus 5–6 mm long, regular. Figure 13A–C.
We have not been able to study the type of this species,
therefore we follow the descriptions of Robinson (1932) and
Cabrera (1996).
Distribution and Habitat—Trichogonia phlebodes is known
only from the type collection. The species blooms in February
and occurs in high fields in the Cordillera of Amambay,
Paraguay. Figure 11.
Notes—Trichogonia phlebodes differs from the other species of Trichogonia by its sessile, elliptic to broadly ovate,
5-nervate leaves with crenate-serrate margins, and deltoid
to rounded base.
Trichogonia prancei G. M. Barroso, Loefgrenia 36: 3. 1969.—
TYPE: BRAZIL. Goiás, Chapada dos Veadeiros, 2 km
from Veadeiros, 18 Jul 1964, G. T. Prance & N. T. Silva
58211 (holotype: RB!; isotypes: UB!, NY!).
Trichogonia munhozii H. Rob., Phytologia 88: 150. 2006.—
TYPE: BRAZIL. Goiás, Alto Paraı́so, road to Colinas do
Sul, Rio das Cobras, 14 Jun 1993, G. & M. Hatschbach 59528
& E. Barbosa (holotype: MBM; isotype: US!), syn. nov.
Erect subshrubs 0.7–1 m tall, branches tomentose, commonly lanate, silvery. Leaves alternate, spirally arranged,
usually lower leaves opposite, decussate and similar to
upper leaves, chartaceous, discolorous, petiole 0.5–1 cm long;
lamina 2.5–7(–15) 0.5–3(–5) cm, elliptic, obtuse at apex, crenate on margins, cuneate at base, adaxial surface tomentose,
+
+
+
+
+
Erect subshrubs 0.4–1 m tall; branches densely tomentose
with glandular stipitate or sessile trichomes. Leaves alternate, spirally arranged, usually lower leaves opposite and
similar to upper leaves, chartaceous, discolorous, petiole 1–2
(–2.5) cm long; laminae 1.5–3.5(–5) 0.6–1.5(–2) cm, oblong,
obtuse at apex, crenate on margins, slightly revolute, base
decurrent onto the petiole, tomentose, with glandular
stipitate trichomes, venation trinervate, inconspicuous. Involucres 0.9–1
0.8–1 cm, receptacles flat; phyllaries ca. 16,
acute at apex, 2-seriate, the outer ca. 7
2 mm, elliptic, the
inner ca. 7
1 mm, linear, pubescent on the apex and
puberulous below, with glandular sessile trichomes. Florets
30–50, 1–1.2 cm long; corolla 4–5 mm long, corolla lobes
0.3 mm long, inconspicuous, corolla tube 2–3 mm long,
glabrous; anther connective appendages obtuse, longer than
All Specimens Examined—BRAZIL, Bahia: Barreiras, 8 Mar 1972,
Anderson et al. 36861 (MG, US-2); Correntina, 13 -14 S, 45 45 0 46 15 0 W, 9 Jan 1991, Rezende et al. 105 (SPF, UB); Brası́lia, Road to
Roda Velha, 4 Feb 1981, King & Bishop 8792 (UB, US). Goiás: Alto
Paraı́so de Goiás, 21 Dec 1968, Barroso et al. 546 (US-2); Posse, 6 Apr
1966, Irwin et al. 14461 (SP, UB, US); São Domingos, 15 May 2000,
Hatschbach et al. 71143 (US).
+
+
Notes—Trichogonia hirtiflora is distinguished by its alter0.5–2 cm), triangular to
nate leaves, with small (0.5–2.5
cordate lamina and abaxially prominent veins. Collections
from Serro County (Franco et al. 211, 324 and Quarema et al.
69, 78) all have green-purple phyllaries.
King and Robinson (1987) and Hind (1993) proposed
Trichogonia apparicioi G. M. Barroso and Trichogonia margarethiae
Soares Nunes respectively to be synonyms of T. hirtiflora, a
conclusion accepted in this study.
We consider T. crenulata a synonym of T. hirtiflora based
on the study of their types. Baker (1876) synonymized
Eupatorium crenulatum Gardner into Trichogonia hirtiflora
(DC.) Sch. Bip. ex Baker, an opinion shared by Barroso
(1951) and King and Robinson (1987). However, Hind
(1993), based on foliar characters, considered E. crenulatum a
distinct species, thus proposing the new combination
Trichogonia crenulata (Gardner) D. J. N. Hind. According to
Hind, T. crenulata differs from T. hirtiflora by its commonly
narrower leaves (0.8 cm wide vs. two cm), abaxial surface
with few (vs. several) glandular stipitate trichomes, margin
serrate-crenate (vs. serrate to deeply crenate), and cypsela ca.
4.5 mm long (vs. three mm long), respectively. Our studies,
however, shows these characters to be part of a continium,
therefore, we only recognize T. hirtiflora.
wide or as long as wide, styles 6–7 mm long. Cypselae 3–
5 mm long, stipitate (base 0.5–1 mm long), setose; carpopodium
annular, decurrent on the ribs; pappus 3–4 mm long, regular.
Figure 10D–F.
Distribution and Habitat—Trichogonia laxa is endemic to
the Brazilian states of Bahia and Goiás, mainly in cerrado
areas. Figure 11.
+
9 Dec 2009, Quarema et al. 28 (ALCB); Congonhas do Norte, Serra da
Carapina, 3 Mar 1998, Pirani et al. 4178 (SPF), Serra Talhada, 19 Jan 2004,
Mello-Silva et al. 2384 (SPF); Cristália, Morro do Chapéu, 6 Jan 1986, Pirani
et al. CFCR 8892 (ALCB, SPF); Datas, 20 Nov 1997, Hatschbach et al. 67327
(INPA, US); Diamantina, 18 160 57.800 S, 43 430 14.300 W, 1,371 m, 26 Jun
2011, Roque et al. 3102 (ALCB); Gouveia, 3 Jun 1985, Semir et al. 17247
(UEC); Grão Mogol, trail of Tropa, 11 Dec 1989, Pirani et al. CFCR 12490
(SPF); Itacambira, 14 Dec 1989, Pirani et al. CFCR 12756 (ALCB, SPF);
Itamarandiba, Santa Joana, 13 Jan 1998, Pirani et al. 3951 (SPF);
Jacoticatubas, Serra do Cipó, 12 Feb 1996, Roque et al. 118 (SPF); Joaquim
Felı́cio, Serra do Cabral, 20 Oct 1999, Hatschbach et al. 69376 (US); Moeda,
Serra da Moeda, 25 Jan 1965, Duarte 8770 (SPF, UB); Ouro Preto, Parque
Estadual de Itacolomi, Trilha da Serrinha, 27 Jan 2006, Almeida et al. 264
(ALCB, VIC); Monte Boa Vista, Serra da Conceição, 6 Mar 1982, Hensold
et al. CFCR 2899 (ALCB, SPF); Rio Vermelho, Serra do Ambrósio, 31 Jul
1985, Pirani et al. CFCR 7823 (ALCB,SPF); Santana do Riacho, Serra do
Cipó, 29 Sep 1998, Roque & Hervêncio 477 (SPF, US); Santo Antônio do
Itambé, Pico do Itambé, 23 Nov 1997, Hatschbach et al. 67443 (US); Serro,
estrada para Capivari, Parque Estadual do Pico do Itambé, 21 Apr 2010,
Franco et al. 324 (ALCB).
543
+
2012]
544
SYSTEMATIC BOTANY
[Volume 37
Fig. 13. Trichogonia phlebodes (B. L. Rob.) R. M. King & H. Rob. A. Floriferous branch. B. Head. C. Floret [Drawn from Cabrera (1996)]. Trichogonia
prancei G. M. Barroso. D. Floriferous branch. E. Receptacle. F. Floret. G. Detail of the pedicellate trichomes in the corolla. H. Stamen with retuse anther
connective appendage. E. Receptacle convex. I. Cypsela with the smaller pappus (Hatschbach et al. 59340, MBM).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
545
Fig. 14. Trichogonia rhodotricha Malme. A. (Drawn from the holotype). Trichogonia salviifolia Gardner. B. Floriferous branch showing the truncate
leaves. C. Corolla and style branches. D. Cypsela (Pires 153, ALCB).
546
SYSTEMATIC BOTANY
+
+
+
abaxial surface conspicuously silvery lanate, glandular
stipitate or sessile trichomes, venation pinnate. Involucres
0.8–1 1–1.2 cm, receptacles convex; phyllaries ca. 34, acute
at apex, 2-seriate, the outer ca. 5 1–2 mm, elliptic, the inner
5–6
0.5 mm, narrowly elliptic to oblanceolate, lanate with
sessile glandular trichomes. Florets 60–80, 6–7 mm long;
corolla 3–3.5 mm long, corolla lobes 0.2 mm long, inconspicuous, corolla tube ca. 2 mm long with stipitate glandular trichomes; anther connective appendages obtuse to retuse, as
long as wide; styles 5–6 mm long. Cypselae ca. 3 mm long,
setose, short-stipitate (base ca. 0.5 mm long); carpopodium
anuliform; pappus 0.2–0.5 mm long, regular. Figure 13D–I.
Distributions and Habitat—Trichogonia prancei is endemic
to Brazil and is restricted to Goiás and the Distrito Federal,
occurring among rocky outcrops in cerrado and campo
rupestre vegetation. Figure 11.
Representative Specimens Examined—BRAZIL. Distrito Federal:
Brası́lia, Fazenda Água Limpa, divisa com o Cristo Redentor (Jardim
Botânico de Brası́lia), 22 Apr 2000, Munhoz et al. 1158 (US). Goiás:
Alto Paraı́so de Goiás, Chapada dos Veadeiros, Água Fria, 10 Mar
2001, Munhoz et al. 2599 (US); Cavalcante, Ponte da Pedra, 13 53 0 7 00 S,
47 31 0 12 00 W, 5 Apr 2007, Pastore & Suganuma 1903 (ALCB, HUEFS);
Pouso Alto, 21 Dec 1968, Barroso et al. 521 (RB, UB).
Notes—Trichogonia prancei belongs to the group of species
defined by the convex receptacle and corolla tube with glandular stipitate trichomes. It is distinguished from other
species in this group by the regular and reduced pappus
(0.2–0.5 mm long) instead of defective pappus as in T. cinerea
and absent in T. eupatorioides. Trichogonia prancei and T. hassleri
have regular regular pappus but differ by several characters.
Trichogonia prancei has elliptic leaves (vs. ovate), florets 60–
80 per head (vs. 25–30) and pappus smaller than 0.6 mm long
(vs. 3–4 mm long).
Trichogonia munhozii H. Rob. was described as having crenate, cuneate lamina, reduced petiole (0.5–1 cm long), large
number of florets per head (60–80), convex receptacle, corolla
tube with glandular stipitate trichomes and minute pappus.
These characters however, are also present in T. prancei,
therefore, we consider T. munhozii a synonym of T. prancei.
Trichogonia rhodotricha Malme, Kongl. Svenska
Vetenskapsakad. Handl. 12(2): 30. 1933.—TYPE: BRAZIL.
Paraná, Capão Grande, Jaguariahyba, 14 Mar 1914, G. O.
A. Malme 4167 (holotype: R!; isotype US!).
+
+
+
+
Erect subshrubs 50–70 cm tall; branches puberulous, with
glandular trichomes. Leaves alternate, spirally arranged,
usually lower opposite, decussate and larger than upper
leaves, chartaceous, discolorous, petiole 1–1.5 cm long; lami2–4 cm, ovate, obtuse at apex, broadly crenate on
nae 4–6
margins, attenuate at apex, glabrescent, venation trinervate.
Involucres ca. 0.7 0.7 cm, receptacles flat; phyllaries ca. 12,
2 mm, elliptic, the
acute at apex, 2-seriate, the outer ca. 4
1 mm, linear, with hyaline, ciliate margins,.
inner ca. 5
Florets ca. 20, 7–8 mm long; corolla 3.5–4 mm long, corolla
lobes ca. 0.3 mm long, triangular, conspicuous; corolla tube
2–2.5 mm, glabrous; anther connective appendages obtuse, as
long as wide, styles ca. 7 mm long. Cypselae ca. 5 mm long,
cylindrical, short-stipitate (base 0.3 mm long), puberulous,
glandular trichomes sessile; carpopodium anuliform; pappus
3–4 mm long, regular. Figure 14A.
Distribution and Habitat—Trichogonia rhodotricha is
endemic to southern Brazil (Paraná and Rio Grande do Sul)
and occurs in open fields. Figure 15.
All Specimens Examined—BRAZIL. Paraná: Ponta Grossa, 10 Mar
1969, Hatschbach & Koczicki 21228 (HUEFS, US). Rio Grande do Sul:
Without specific locality, 1816–1821, Saint-Hilaire 2688 (Photo US).
[Volume 37
Notes—Trichogonia rhodotricha is distinguished from other
species of Trichogonia by its ovate leaves with broadlycrenate margins.
Trichogonia salviifolia Gardner, London J. Bot. 6: 460.
1846.—TYPE: BRAZIL. Minas Gerais: “in a dry campo,
near Morro Velho,” Sep 1840, G. Gardner 4839/1 (pro
parte) [lectotype, here designated: BM!, Photo SPF!].
Eupatorium conoclinioides Gardner, London J. Bot. 5: 474. 1846.—
TYPE: BRAZIL. Ceará, Feb 1839, G. Gardner 2419 [lectotype, here designated: BM!, Photo: SPF!; isolectotypes:
K! (ex Herbarium Hookerianum, Photo: SPF!), K!
(ex Herbarium Benthamianun, Photo: SPF!)]. [synonym
proposed by King and Robinson 1987].
Trichogonia rhadinocarpa B. L. Rob., Proc. Amer. Acad. Arts.
42: 36. 1906.—TYPE: VENEZUELA. Near Tovar, 1854–
1855, A. Fendler 651 (holotype: G; isotype: NY!, Photo:
US!), syn. nov.
Trichogonia alba V. M. Badillo, Bol. Soc. Venez. Ci. Nat. 10:
295. 1946.—TYPE: VENEZUELA. Aragua, Guamitas.
Parque Nacional Aragua, Jun 1945, V. M. Badillo 1302
(holotype: VEN (digital JSTOR-image)!; Photo: US!).
(synonym proposed by King and Robinson 1987).
Trichogonia fiebrigii Mattf., Notizbl. Bot. Berlin-Dahlem 8: 450.
1923.—TYPE: PARAGUAY. Concepción: “N. Paraguay:
zwischen Rio Apa und Rio Aquidaban,” 1908–1909,
K. Fiebrig 4289 (lectotype by Roque et al. 2011: M [0029560]!;
isolectotypes: G [00306026]!, G [00306027]!, K!, Photo:
SPF!), syn nov.
Trichogonia salviifolia Gardner ex Hassler, Repert. Spec. Nov.
Regni Veg. 14: 291. 1916, nom. lleg.; non Trichogonia
salviifolia Gardner, London J. Bot. 5: 460. 1846.—TYPE:
BRAZIL. Minas Gerais: “in a dry campo, near Morro
Velho”, Sep 1840, G. Gardner 4839/1 (pro parte)].
Trichogonia salviifolia Gardner f. macrophylla Hassler,
Repert. Spec. Nov. Regni Veg. 14: 291. 1916.—TYPE:
PARAGUAY. Concepción: “Nördl. Paraguay (22-23o
lat.) zwischen Rio Apa u. Aquidaban. San Luis,” Dec
1908, K. Fiebrig 4441 (holotype: G [00306024]!; isotype: M
[0029558]!, Photo: US!).
Trichogonia salviifolia Gardner f. linearifolia Hassler, Repert.
Spec. Nov. Regni Veg. 14: 291. 1916.—TYPE:
PARAGUAY. Concepción: “N. Paraguay: zwischen
Rio Apa und Rio Aquidaban. Centurion,” Nov 1908–
1909, K. Fiebrig 4271 (holotype: G [00306022]!; isotypes:
G [00306023]!, K!, MO, Photo: SPF!). [synonyms proposed by Mattfeld 1923].
Trichogonia scottmorii R. M. King & H. Rob. Phytologia 45:
110. 1980.—TYPE: BRAZIL. Bahia, Maracás, road BA
250, 40 Km east of Maracás, 13 Jul 1979, R. M. King &
S. A. Mori 8018 (holotype: CEPEC!), syn. nov.
Trichogoniopsis grazielae Soares Nunes. Bradea 3(18): 132.
1981.—TYPE: BRAZIL. Pernambuco, between Triunfo
and Flores, 25 May 1971, E. P. Heringer, D. de Andrade
Lima, J. P. Lanna Sobrinho & A. C. Sarmento 908 (holotype:
RB!), syn. nov.
Erect subshrubs 0.3–1.5 m tall; branches aromatic,
sticky, puberulous with sessile and stipitate glandular trichomes. Leaves alternate, spirally arranged, usually lower
leaves opposite and similar to upper leaves, chartaceous,
slightly discolorous, aromatic, sticky, petiole 2–4 cm long;
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
547
Fig. 15. Distribution of T. rhodotricha, T. spathufilofia, T. tombadorensis and T. villosa.
+
+
+
+
laminae (2–)3–7 1–3 cm, lanceolate to triangular, acute at
apex, crenate to dentate on margins, truncate at base, adaxial surface glabrescent, abaxial surface puberulous, with
glandular sessile trichomes especially along veins, venation
inconspicuous or 3–4-nerved. Involucres (0.7–)0.9–1 0.6–
0.7 cm, receptacles flat; phyllaries ca. 15, ca. 2-seriate,
the outer 4–5 1 mm, elliptic to oblanceolate, the inner 5–
6 0.5–1 mm, linear to oblanceolate, pubescent, with glandular sessile trichomes and ciliate at apex. Florets 18–25,
0.8–1 cm long; corolla 5–6 mm long, corolla lobes 0.1 mm
long, inconspicuous, corolla tube 4–5 mm with glandular
sessile trichomes; anther connective appendages obtuse,
longer than wide; style ca. 6 mm long. Cypselae 3–4 mm
long, stipitate to long-stipitate (0.5–2 mm long), setose,
glandular trichomes on apex; carpopodium annular; pappus 3–4 mm long, regular. anuliform; pappus 0.2–0.5 mm
long, regular. Figure 14B–D.
Distribution and Habitat—Trichogonia salviifolia is the
most widely distributed species in the genus, occurring
in Venezuela, Colombia, Paraguay, Bolivia and Brazil. In
Brazil, it is found in the states of Ceará, Pernambuco, Piauı́,
Bahia, Goiás, Distrito Federal, Mato Grosso do Sul, Minas
Gerais, São Paulo, Rio de Janeiro, and Espı́rito Santo. The
species occurs in cerrado, caatinga, campo rupestres, margins
of riparian forests, and in disturbed areas. Figure 16.
Representative Specimens Examined—VENEZUELA. Distrito Federal, Caracas: Mission station, 9 Sep 1925, Zehntner 89 (US); Parque
Nacional Henry Pittier, 21 Jun 1967, Trujillo 8465 US). Falcón, Dtto.
Mauroa, 23 May 1980, Bunting & Stoddart 9356 (US). Maracay, without
specific locality, 1934, Vogel 768 (US). Mérida, Carretera BarrancasBarinas, 23 Jul 1966, Marcano Berti 1015 (US). Miranda, 11 Dec 1938,
Alston 5479 (US). Sucre, valley of Cocollar, 28 Apr 1945, Steyermark
62379 (US). Trujillo, Carache, Jul 1995, Licata et al. 398 (US). Yaracuy,
Mendoza, La Cuchilla, 16 Jan 1928, Pittier 12620 (US).
BOLIVIA. Santa Cruz: Chiquitos, 18 190 1000 ,500 S, 59 340 46.800 W, 31 Jan
1995, R. R. Abbott & B. Mostacedo 15952 (US).
BRAZIL. Bahia: Abaı́ra, 15 170 0000 S, 41 510 0000 W, 19 Dec 1991, Hind et al.
H50018 (ALCB2x, US); Barra da Estiva, 13 290 1200 S, 41 130 2300 W, 7 Mar
1996, Roque et al. PCD 2182 (ALCB-3, CEPEC); Bom Jesus da Lapa,
14 Mar 2002, Roque et al. 660 (SPF); Bonito, 20 May 2001. Alves et al. 125
(ALCB); Caetité, Brejinho das Ametistas, 28 Apr 2003, Roque et al. 624
(ALCB); Campo Alegre de Lourdes, Morro da Carlota, 9 290 3300 S,
43 50 3300 W, 21 May 2000, Queiroz et al. 6219 (CEPEC); Campo Formoso,
Serra dos Morgados, 14 Apr 2006, Santos 538 (ALCB, HUEFS); Catolés, road
to Piatã, 26 Mar 2005, Guedes et al. 11881 (CEPEC); Conceição de Feira, Aug
1974, Pinto 42356 (ALCB-2, HRB 16480); Itaberaba, 30 Jan 1981, King &
Bishop 8703 (UB, US); Itagibá, Mata do Cacau, 27 Oct 2008, Branches et al.
440 (ALCB); Itiruçu, 13 Feb 1979, Santos et al. 3442 (US); Jacobina,
11 200 4000 S, 40 290 5500 W, 7 Apr 2001, Bautista et al. 3067 (ALCB, CEPEC);
Maracás, 25 Mar 2008, Ferreira et al. 381 (HUEFS); Miguel Calmon, vicinity
Parque Sete Passagens, 22 Dec 2006, Guedes et al. 13140 (ALCB); Mirangaba,
23 Apr 1981, Fonseca 411 (ALCB, HRB, RB, UB); Morro do Chapéu, trail to
Ventura, Cidade das Pedras, 24 May 2008, Roque & Liro 1824 (ALCB, US);
Mundo Novo, Feb 1968, A. L. Costa s. n. (ALCB 4324); Palmeiras, idem,
23 May 1980, Harley 22422 (CEPEC); Piatã, 13 160 0100 S, 41 440 19.800 W,
548
[Volume 37
Distribution of Trichogonia salviifolia.
Del Rei, São Tomé das Letras, 5 Feb 1973, Hatschbach & Ahumada 31273 (US).
Pernambuco: Buique, 21 Jun 1975, Lima 75–8068 (IPA); Gravatá, 6 Mar 1966,
Lima 66–4458 (IPA). Piauı́: Guaribas, Serra das Confusões, 28 Mar 2007,
Barrot et al. 2906 (HUEFS). Rio de Janeiro: Paineiras, 17 Jan 1943, Bueno 6
(R); Petrópolis, 02 Jan 1971, Barra 237 (R). São Paulo: Cabreúva, 9 Mar 1994,
Barreto et al. 2136 (ESA, SPF); João Ramalho, 13 Feb 1996, V. C. Souza & J.
Souza 10833 (ESA); Luiz Antônio, 4 Feb 1987, Leitão Filho et al. 18886 (UEC);
Rancharia, 14 Feb 1996, V. C. Souza & J. Souza 10932 (ESA, SP, SPF); Santa
Rita do Passa Quatro, ARIE, Cerrado Pé de Gigante, 16 Jan 1998, Godoy et al.
1265 (SPF); Paraguaçu Paulista, 8 Feb 1965, Eiten et al. 5927 (US); Rancharia,
14 Feb 1996, V. C .Souza & J. Souza 10932 (ESA,SP); São Carlos, 17 Apr 1994,
V. C. Souza et al. 5839 (ESA).
COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, 25 Sep 1959,
Cuatrecasas & Romero Castañeda 24422 (US). Santander, Road from Abrego
to Sadinanta, 27 May 1969, Garcı́a-Barriga & Jaramillo Mejia 19953 (US),
Norte de Santander, 26 Sep 1969, Cuatrecasas & Rodrı́guez 27958 (US).
PARAQUAY. Amambay: 11 Dec 1997, Schinini & Dematteis 33737 (SPF).
Conceptión, Zwischen Rio Apa und Rio Aquidaban, Jan 1908/1909,
K. Fiebrig 4289 (K-Paratype). Paraguari - Compañia Costa Segunda,
2 Jun 1988, Zardini 4555 (US).
Notes—Trichogonia salviifolia is distinguished from other species of Trichogonia by its long petiolate (2–4 cm long), large [(2–)
3–7 1–3 cm], triangular leaves with truncate base, 20–25 florets
per head and pappus 3–4 cm long, a combination of characters
not found in other species of Trichogonia. The species has
aromatic stems and leaves and its vernacular name in Bahia
(Brazil) is ‘barba-de-velho’(old beard) or ‘amargoso’ (bitter),
which is not apparently related to its aromatic vegetative parts.
Robinson (2006) considered T. rhadinocarpa as closely
related to T. campestris but distinguished the first by wider
leaves and larger stipitate cypsela. However, the characters
1.3–2.5 cm, with crenate-serrate
(lanceolate lamina, 3–5.5
+
9 Nov 1996, Hind et al. PCD 4164 (ALCB, CEPEC, SPF); Pindobaçu,
10 560 3600 S, 40 240 2800 W, 10 Mar 2001, Jesus et al. 1365 (ALCB, CEPEC);
Rio de Contas, 13 320 0000 S, 41 560 0000 W, 27 Dec 1988, Harley et al. 27378
(ALCB, CEPEC, SPF, US); Saúde, 10 560 3700 S, 40 240 2900 W, 9 Apr 2001, Jesus
et al.1352 (ALCB, CEPEC); Seabra, Serra do Bebedor, 15 Nov 1983, Bautista
1340 (CEPEC); Ubaı́ra, 13 150 3400 S, 39 500 900 W, 10 Mar 2008, Oliveira et al.
1502 (HUEFS); Utinga, 12 280 0000 S, 41 270 0000 W, 10 Mar 1996, Roque et al.
PCD 2211 (ALCB). Ceará: base da serra de Maranguape, 24 Nov 1955, Lima
55–2415 (IPA); Alcântera, Serra da Meruoca, 4 Apr 2001, Souza 560
(HUEFS); Fortaleza, 23 Mar 1966, Matos & Barros, Barros s. n. (EAC 5423);
Monguba, 1839, Gardner 2419 (RB); Novo Oriente, 15 Feb 1991, Araújo 283
(EAC, IPA). Distrito Federal: Brası́lia, Reserva Ecológica do IBGE, 23 Jan
1997, Roque 302 (ALCB, SPF). Espı́rito Santo: Santa Tereza, 12 Nov 1985,
Fernandes 1642 (SPF); Vitória, 13 Apr 1871, Sello 266 (RB, US). Goiás: Caldas
Novas, Termas do Rio Quente, 5 Jan 1977, Heringer 16649 (US); Cristalina,
10 km de Cristalina, 5 Mar 1966, Irwin et al. 13554 (US); Nova Roma, Alto
Paraı́so, 5 Feb 1981, King & Bishop 8804 (UB, US); Posse 9 Jan 1965, Belém &
Mendes 147 (CEPEC, MG, UB, US). Maranhão: Carolina, 18 Mar 1934,
Swallen 3893 (US). Mato Grosso do Sul: Porto Murtinho, Road MS 382,
Bonito – Campo dos Índios, 12 Oct 2003, Hatschbach et al. 76222 (SPF, US).
Minas Gerais: Belo Horizonte, Serra do Curral, Jan 1958, Vidal V.190/222
(R); Brumadinho, Serra da Moeda, 10 Jan 1999, Silveira 152 (US); Campanha,
25 Feb 1976, Davidse & Ramamoorthy 10616 (US); Diamantina, 3 Mar 1996,
Roque et al. 186 (ALCB, SPF); Grão Mogol, 4 Feb 2007, Santos et al. 997
(HUEFS); Januaria, 14 Feb 1998, Salino & Gotschalg 4021 (US); Joaquim
Felı́cio, Parque Estadual da Serra do Cabral, 10 Dec 2009, Quaresma et al. 36
(ALCB); Lavras, Lagoa Santa, 2 Jan 1977, Gibbs 4109 (UEC); Mariana,
Parque Estadual do Itacolomi, 16 Mar 2006, Almeida & Custódio 351 (ALCB);
Montes Claros, 21 Jan 1981, King & Bishop 8578 (UB, US); Montezuma,
toward Espinosa, 15 Mar 1994, Souza et al. 5512 (SPF); Ouro Preto, Parque
Estadual de Itacolomi, trail to Morro do Cachorro, 26 Sep 2005, Almeida et al.
105 (ALCB, VIC); Paracatu, 3 Feb 1970, Irwin et al. 25932 (SP, UB, US);
Piumbi, 21 Feb 1978, Shepherd et al. 7124 (UEC); Poços de Caldas, 21 Mar
1965, Emmerich 2447 (RB); Sabará, Jan 1916, Hoehne 6780, 6855 (R); São João
+
Fig. 16.
SYSTEMATIC BOTANY
ROQUE ET AL.: REVISION OF TRICHOGONIA
Trichogonia tombadorensis R. M. King & H. Rob.,
Phytologia 45: 111. 1980.—TYPE: BRAZIL. Bahia. Morro
do Chapéu, Serra do Tombador, 20 Feb 1971, H. S. Irwin,
+
+
+
+
Erect subshrubs 0.5–1 m tall; branches tomentose with
glandular sessile trichomes. Leaves alternate, fasciculate,
congested, subcoriaceous, concolorous, petiole 2–6 mm long;
laminae 0.4–0.9 0.3–0.7 cm, obovate, obtuse at apex, entire
to serrulate on margins, attenuate at base, tomentose, with
glandular sessile trichomes, venation pinnate. Involucres 0.7–
1 0.7–1 cm; receptacles flat; phyllaries 12–15, 2-seriate,
rounded at apex, the outer ca. 4 1 mm, elliptic to spatulate,
the inner ca. 5 0.5–1 mm, linear, pubescent on upper third
and puberulous below, glandular trichomes on abaxial surface. Florets 20–35, ca. 9 mm long, corolla 3–4 mm long,
corolla lobes 0.3 mm long, triangular, conspicuous, corolla
tube 2–3 mm long, glabrous; anther connective appendages
obtuse to retuse, wider than long, style 6–7 mm long.
Cypselae 3–4 mm long, stipitate (base 0.5–1 mm long), setose,
with glandular sessile trichomes; carpopodium inconspicuous; pappus ca. 3 mm long, regular. Figure 17F-H.
Distribution and Habitat—Trichogonia tombadorensis is
endemic to the state of Bahia, Brazil, and is found in campo
rupestre vegetation. Figure 15.
Representative Specimens Examined—BRAZIL. Bahia: Boqueirão dos
Lopes, 18 May 1975, Costa & Barroso s. n. (ALCB 7260); Morro do Chapéu,
11 370 7000 S, 41 000 0700 W, 21 Jan 2006, Roque et al. 1318 (ALCB), trail to
Cidade das Pedras, 24 May 2008, Roque & Liro 1819 (ALCB, US).
Notes—Trichogonia tombadorensis is similar to T. harleyi, as
they both have small, fasciculate, congested leaves. However,
T. tombadorensis is distinguished from T. harleyi by the longer
(0.4–0.9 vs. 0.3–0.4 cm), obovate (vs. orbiculate) lamina with
entire to serrulate (vs. crenate) margins.
Trichogonia villosa (Spreng.) Sch. Bip. ex Baker in C. F. P.
von Martius, Fl. Bras. 6 (2): 213. 1876. Kuhnia villosa
Spreng. Syst. Veg. 3: 439. 1826.—TYPE: BRAZIL. Brasilia,
Sellow 763 [lectotype, here designated: P!; isolectotypes
p. p.: K! (Photo: SPF!), P!; Photo: F!].
Trichogonia multiflora Gardner, London J. Bot. 5: 460. 1846.
Trichogonia villosa (Spreng.) Sch. Bip. ex Baker var.
multiflora (Gardner) Baker in C. F. P. von Martius,
Fl. Bras. 6(2): 213. 1876.—TYPE: BRAZIL. Minas Gerais
“Elevated Campos. Diamond District,” Jul 1840, G.
Gardner 4851 [lectotype, here designated: BM! (Photo:
SPF!); isolectotypes: K! (ex Herbarium Benthamianum,
Photo: SPF!), P-2!, K! (ex Herbarium Hookerianum,
Photo: SPF!), R!, NY-2!, US!, syn. nov.
Erect subshrubs 0.5–1.5 m tall; branches puberulous, glandular trichomes stipitate, sticky. Leaves alternate, spirally
arranged to fasciculate, chartaceous, discolorous, sessile;
laminae 2–5(–6) 0.2–1 cm, linear to elliptic, obtuse to acute
at apex, entire, slightly dentate on the upper third to serrate
on margins, revolute, cuneate at base, puberulous, glandular
stipitate or sessile trichomes, sticky, venation pinnate. Involucres 0.8–1.2 0.7–1.1 cm; receptacles flat; phyllaries 16–30,
2–3-seriate, the outer 5–8
1.5–2 mm, elliptic, the inner 6–
8 1 mm, linear, puberulous, abaxial surface with glandular
stipitate or sessile trichomes, setose, ciliate at margin petiole
2–6 mm long. Florets 45–75, 1–1.2 cm long, corolla 5–6 mm
long, corolla lobes 0.3 mm long, conspicuous, corolla tube
3–4 mm, pubescent on apex; anther connective appendages
obtuse to retuse at apex, longer than wide; styles 8 mm long.
Cypselae 4–6 mm long, stipitate (0.5–1 mm long), setose;
carpopodium anuliform; pappus 4–5 mm long, regular.
Figure 18A–E.
+
+
+
+
Prostrate subshrubs ca. 0.5 m tall; branches griseopuberulous. Leaves alternate, spirally arranged, usually lower
leaves opposite and similar to upper leaves, subcoriaceous,
discolorous, petiole 0.8–1 cm long; lamina ca. 2.5 0.5 cm,
oblanceolate, rounded at apex, entire on margins, cuneate at
base, adaxial surface puberulous, abaxial surface incanotomentose, with glandular sessile trichomes, venation pinnate. Involucres 1.2–1.5 cm diam., receptacles flat; phyllaries
ca. 13, 2-seriate, the outer 6.5–7 1.5–1.8 mm, oblong-spatulate,
obtuse, the inner 6.5–7 0.8 mm, oblong-rhombic, obtuse,
incano-tomentose, with sessile glandular trichomes. Florets
ca. 30, 9–10 mm long; corolla ca. 5 mm long, corolla lobes
0.8 mm long, conspicuous, corolla tube 2.5 mm, with glandular sessile trichomes; anther connective appendages obtuse,
as long as wide; style ca. 1 cm long. Cypselae 5–5.5 mm long,
stipitate to long-stipitate (1.5–2 mm long), setose; carpopodium
annular; pappus ca. 4 mm long, regular. Figure 17A–E.
This species is known only from the picture of the type,
therefore, we follow Mattfeld (1923).
Distribution and Habitat—Trichogonia spathulifolia is
known only from the type specimen collected in Rio de
Contas Municipality, Bahia state, Brazil. Figure 15.
Notes—Trichogonia spathulifolia is distinguished by its
griseo-puberulous branches, oblanceolate, abaxially whitetomentose lamina, rounded at apex, with entire margins.
R. M. Harley & G. L. Smith 30671 (holotype: US!; isotype:
US!, NY).
+
TRICHOGONIA SPATHULIFOLIA Mattf., Notizbl. Bot. Gart. Berlin 8:
445. 1923.—TYPE: BRAZIL. Bahia, Rio de Contas,
Boqueirão Calderão, 13 Aug 1913, Lützelburg 412 (holotype: M; Photos: RB!, US!).
549
+
+
margins, capitula with 18 florets, long-stipitate cypsela, and
4 mm long pappus) used by him to support T. rhadinocarpa
are also considered diagnostic for T. salviifolia. Thus, we
propose T. rhadinocarpa as a synonym of T. salviifolia.
Trichogonia alba Badillo, from Venezuela was considered
to be a synonym of T. rhadinocarpa B. L. Rob. by King and
Robinson (1987), a decision that is accepted by us. Therefore,
we also consider it to be a synonym of T. salviifolia.
From specimens collected in Paraguay, Hassler (1916)
described three new forms in Trichogonia salviifolia Gardner
(T. salviifolia f. typica, T. salviifolia f. microphylla and T. salviifolia
f. linearifolia) based on leaf shape and size. Mattfeld (1923)
considered these forms as a new species, which he named
Trichogonia fiebrigii. After a detailed study of the type specimens and protologues, we are proposing T. fiebrigii as synonym of T. salviifolia because the characters used to delineate
the former fall within the variation found in T. salviifolia.
Trichogonia scottmori R. M. King & H. Rob. is considered to
be a synonym of T. salviifolia because the characters utilized
to recognize it (i.e. anther connective appendages as long as
wide, deltoid or ovate, 3-nerved leaves with crenate-serrulate
margins) are included within the morphological variation of
T. salviifolia.
Finally, Trichogoniopsis grazielae Soares Nunes is proposed
as a synonym of T. salviifolia based on corolla lobes pubescence, lanceolate-ovate (3 1.7 cm) leaves with acute apices,
and obtuse-truncate bases, and a larger pappus (4 mm long).
The corolla lobe pubescence and plumose pappus are diagnostic of Trichogonia while the remaining cited characters fit
under the specific concept of T. salviifolia.
+
2012]
550
SYSTEMATIC BOTANY
[Volume 37
Fig. 17. Trichogonia spathulifolia Mattf. A. Floriferous branch. B. Floret. C. Corolla with two types of trichomes. D. Tector and glandular trichomes on
the corolla. E. Corolla opened showing the lobes (Drawn from Lützelburg 412). Trichogonia tombadorensis R. M. King & H. Rob. F. Floriferous branch. G.
Corolla and style branches. H. Cypsela (Hatschbach 39570, MBM).
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
551
Fig. 18. Trichogonia villosa. A. Specimen showing wider leaves (Semir et al. CFCR 223, ALCB). B. Specimen showing the thinner leaves. C. Corolla and
style branches. D. Receptacle plane. E. Cypsela (B–E Roque et al. 189, ALCB).
552
SYSTEMATIC BOTANY
Distribution and Habitat—Trichogonia villosa is endemic to
Brazil where it is found in the states of Bahia and Minas
Gerais. This species occurs in campos rupestres vegetation,
and it is common in Santana do Riacho (Serra do Cipó) and
Diamantina (Planalto de Diamantina) municipalities of
Minas Gerais. Figure 15.
Representative Specimens Examined—BRAZIL. Bahia: Abaı́ra, road
to Marques- Boa Vista, 27 Apr 1994, Ganev 3147 (SPF, US); Mucugê,
4 Jul 2009, Roque et al. 2068 (ALCB, US); Rio de Contas, road to Pico
das Almas, 28 Jan 2000, Jardim et al. 2576 (CEPEC). Minas Gerais:
Augusto de Lima, Serra do Cabral, 20 Mar 1994, Roque et al. CFCR
15303 (SPF); Belo Horzionte, Serra do Itabarito, 7 Feb 1968, Irwin et al.
19525 (US); Buenopolis, 15 Dec 1991, Harley et al. 49950 (US);
Carrancas, 9 Dec 1983, Leitão Filho et al. 15376 (UEC); Cristália, Morro
do Chapéu, 6 Jan 1986, Cordeiro CFCR 8928 (ALCB, SPF); Datas,
Morro do Coco, 18 26 0 S, 43 41 0 W, 8 Jan 1988, Simão et al. CFCR 11668
(ALCB, SPF); Diamantina, 18 16 0 57.8 00 S, 43 43 0 14.3 00 W, 1371m, 26 Jun
2011, Roque et al. 3103, 3104 (ALCB); Gouveia, road para Barão de
Guacui, 24 Jul 1998, Hatschbach et al. 68181 (US); Grão Mogol, 16 Apr
1981, Rossi et al. CFCR 1027 (UEC); Itabirito, Pico da Cata Branca,
1 Jun 1945, Williams & Assis 7368 (SPF); Jaboticatubas, Serra do Cipó,
9 Mar 1985, Lewinsohn & Martins 18093 (UB, UEC); Joaquim Felicio, 17 Nov
1997, Hatschbach et al. 67193 (US); Lavras, Reserva Biológica Poço-Bonito,
11 Dec 1980, Leitão Filho et al. 12043 (UEC); Montes Claros, 62 Km NE of
Montes Claros to Salinas, 21 Jan 981, King & Bishop 8579 (UB); Ouro
Branco, old road Ouro Branco-Ouro Preto, 8 Mar 1995, Souza et al. 7872
(SPF); Ouro Preto, Parque estadual de Itacolomi, trail to Morro do
Cachoro, 18 Apr 2006, Almeida et al. 380 (ALCB, VIC); Rio Vermelho, Pedra
Menina, Morro do Ambrósio, 9 Sep 1986, Mello-Silva et al. CFCR 10246
(ALCB, SPF); Santa Bárbara, Serra do Caraça, 14 Dec 1978, Leitão Filho
et al. 9767 (UEC); Santana do Pirapama, Serra do Cipó, Trilha do João
Carrinho, 18 Feb 2007, Souza et al. 32653 (SPF); Santana do Riacho, Capão
Redondo, 4 Km from bifurcation to Morro do Pilar, 13 Feb 1996, Roque et al.
162 (ALCB, SPF); São João Del Rei, Serra do Lenheiro, 4 Dec 1998, Roque &
Faria 507 (ALCB, SPF); São João da Chapada, 24 Mar 1970, Irwin et al. 28215
(SP, US); Turvolândia, 24 Apr 1926, Hoehne & Gehrt 17486 (SP); Tiradentes,
Serra de São José, 16 Jan 1994, Giulietti et al. CFCR 13704 (SPF); Várzea de
Palma, Serra do Cabral, 16 Jan 1996, Hatschbach et al. 64185 (US).
Notes—Baker (1876) transferred T. multiflora Gardner as a
variety of T. villosa based on the large morphological variation of leaf shape and number of florets found in T. villosa.
However, after analyzing populations in the field, several
specimens and the type of T. villosa concept, we have concluded that the species has high morphological plasticity. For
instance, the number of florets per head and leaf shape
(linear to elliptic), are variable within individuals belonging
to the same population. Thus, we are proposing to place
T. villosa var. multiflora under T. villosa since there are no
morphological, biogeographical, or ecological characters to
distinguish any infra-specific taxa.
Excluded Names (listed by King and Robinson 1987)—
Trichogonia barrosoana G. M. Barroso, Arch. Jard. Bot.
Rio de Janeiro 11: 13. 1951. = Campuloclinium
campuloclinioides (Baker) R. M. King & H. Rob.
Trichogonia gardneri A. Gray, Hooker’s J. Bot. Kew Gard.
Misc. 3: 224. 1851. = Trichogoniopsis adenantha
(DC.) R. M. King & H. Rob.
Trichogonia leiantha Sch. Bip. ex Baker, Fl. Bras. 6(2): 215. 1876,
nom. nud. in syn. = Trichogoniopsis adenantha (DC.)
R. M. King & H. Rob.
Trichogonia macrolepis Baker, Fl. Bras. 6(2): 215. 1876. =
Trichogoniopsis podocarpa (DC.) R. M. King & H. Rob.
Trichogonia melissaefolia (DC) Mattf, Notizbl. Bot. Gart. BerlinDahlem 8: 450. 1923. = Platypodanthera melissaefolia
(DC.) R. M. King & H. Rob.
Trichogonia podocarpa (DC.) Shultz-Bip. ex Baker, Fl. Bras.
6(2): 216. 1876. = Trichogoniopsis podocarpa (DC.)
R. M. King & H. Rob.
[Volume 37
Trichogonia scabra Klatt, Arbeiten Bot. Mus. Hamb. 3. 1890. =
Campuloclinium tubaraoense (Hieron) R. M. King &
H. Rob.
Trichogonia selloi Sch. Bip. ex Baker, Fl. Bras. 6(2): 215. 1876.
nom. nud. in syn.= Trichogoniopsis adenantha (DC.)
R. M. King & H. Rob.
Trichogonia viscosa Sch. Bip. ex Kraschen, Bot. Mater. Gerb.
Glavn. Bot. Sada 3: 161. 1922. nom. nud. in syn. =
Acritopappus longifolius (Gardner) R. M. King & H. Rob.
Acknowledgments. The authors are grateful to the curators of the
herbaria that granted loans of the studied material. The senior author is
grateful to Prof. José Rubens Pirani, then curator of the herbarium SPF,
for his support on this revision. Special thanks go to Pedro Acevedo
Rodrı́guez and the reviewers of the manuscript for valuable comments
and suggestions. We also thank to Lorenzo Ramella for his helpful
advices in the typification of names from Paraguay. Finally, we thank
Natanael Nascimento dos Santos for drawing the fine illustrations of the
species, and Luciano Pataro for creating the distribution maps.
Literature Cited
Baker, J. G. 1876. Compositae III–Eupatorieae. Pp. 212–217 in Flora
Brasiliensis 6, eds. C. F. P. Martius and A. W. Eichler. Munchen:
Monachii, Typografia Regia.
Barroso, G. M. 1951. Estudos das espécies Brazileiras de Trichogonia
Gardner. Arquivos do Jardim Botânico do Rio de Janeiro 11: 7–18.
Bremer, K. 1994. Asteraceae–cladistics and classification. Portland: Timber Press.
Cabrera, A. L. 1996. Compositae III. Asteroideae, Eupatorieae.
Adenostemma, Ageratum, Eupatorium, Gymnocoronis, Stevia, Trichogonia.
Pp. 1–349 in Flora del Paraguay 25, eds. Spichiger R. and L. Ramella.
Genève: Conservatoire et Jardin botaniques de la Ville de Genève.
De Candolle, A. P. 1836. Compositae, Eupatoriaceae. Pp. 103–211 in
Prodromus Systematis Naturalis Regni Vegetabilis 5, ed. A. P. de Candolle.
Paris: Treuttel and Wurtz.
Ferreira, S. C. 2010. Filogenia molecular da subtribo Gyptidinae R. M. King &
H. Rob. (Eupatorieae– Asteraceae) e revisão taxonômica do gênero
Lasiolaena R. M. King & H. Rob. Ph. D. thesis, Feira de Santana, Bahia:
Universidade Estadual de Feira de Santana.
Funk, V. A., A. Susanna, T. F. Stuessy, and R. J. Bayer. 2009a. Systematics,
evolution and biogeography of the Compositae. Vienna: IAPT.
Funk, V. A., A. Susanna, T. F. Stuessy, and H. Robinson. 2009b. Classification of Compositae. Pp. 171–189 in Systematics, evolution and biogeography of the Compositae, eds. V. A. Funk, A. Susanna, T. F. Stuessy,
and R. J. Bayer. Vienna: IAPT.
Gardner, G. 1846. Contributions towards a Flora of Brazil, being the
characters of several new species of Compositae, belonging to the
tribe Eupatoriaceae. London Journal of Botany 5: 455–491.
Hassler, E. 1916. Ex herbario Hassleriano: Novitates paraguarienses. XXI.
Repertorium Specierum Novarum Regni Vegetabilis 14: 263–298.
Hind, D. J. N. 1993. A new combination in Trichogonia (Compositae). Kew
Bulletin 48: 411–412.
Hind, D. J. N. 1999. A new genus, Semiria (Compositae: Eupatorieae:
Gyptidinae), and a discussion of its affinities within the Gyptidinae
of Bahia, Brazil. Kew Bulletin 54: 425–432.
Hind, D. J. N. 2000. A new genus, Catolesia (Compositae: Eupatorieae),
and a discussion of its affinities within the subtribe Gyptidinae of
Bahia, Brazil. Kew Bulletin 55: 941–948.
Hind, D. J. N. and E. B. Miranda. 2008. Lista preliminar da famı́lia Compositae
na região Nordeste do Brazil. Kew: Royal Botanic Gardens.
Hind, D. J. N. and H. Robinson. 2007. Tribe Eupatorieae. Pp. 510–573 in
The families and genera of vascular plants vol. 8, ed. K. Kubitzki. Berlin:
Springer Verlag.
Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index
herbariorum. New York: New York Botanical Garden.
King, R. M. and H. Robinson. 1972. Studies in the Eupatorieae
(Asteraceae). CXII. The genus Trichogonia. Phytologia 24: 173–179.
King, R. M. and H. Robinson. 1977. Studies in the Eupatorieae
(Asteraceae). CLXI. A new species of Stevia from Costa Rica.
Phytologia 35: 229–232.
King, R. M. and H. Robinson. 1978. Studies in the Eupatorieae (Asteraceae).
CLXXXI. Two new species of Trichogonia. Phytologia 39: 334–338.
King, R. M. and H. Robinson. 1980. Studies in the Eupatorieae (Asteraceae).
CLXXXVII. Additions to Trichogonia and Trichogoniopsis. Phytologia
45: 105–123.
2012]
ROQUE ET AL.: REVISION OF TRICHOGONIA
King, R. M. and H. Robinson. 1987. The genera of the Eupatorieae
(Asteraceae). Monographs in Systematic Botany from the Missouri Botanical Garden 22: 1–581.
Mattfeld, J. 1923. Compositae - Plantae Lützelburgianae brasiliensis I. Notizblatt
des Botanischen Gartens und Museums zu Berlin-Dahlem 8: 428–451.
Panero, J. L. and V. A. Funk. 2008. The value of sampling anomalous taxa
in phylogenetic studies: Major clades of the Compositae revealed.
Molecular Phylogenetics and Evolution 47: 757–782.
Radford, A. E., W. C. Dickison, J. R. Massey, and C. R. Bell. 1974. Vascular
plant systematics. New York: Harper and Row Publishers.
Robinson, B. L. 1913. A key to the genera of the Asteraceae – Eupatorieae.
Proceedings American Academy Arts 49: 429–437.
Robinson, B. L. 1932. Records preliminary to a general treatment
of the Eupatorieae, X. Contributions from the Asa Gray Herbarium
100: 3–19.
Robinson, H. 2006. New species and new combinations in Brazilian
Eupatorieae (Asteraceae). Phytologia 88: 136–153.
553
Robinson, H., A. M. Powell, G. D. Carr, R. M. King, and J. F. Weedin.
1989. Chromosome numbers in Compositae, XVI: Eupatorieae II.
Annals of the Missouri Botanical Garden 76: 1004–1011.
Robinson, H., E. Schilling, and J. L. Panero. 2009. Eupatorieae. Pp. 731–744
in Systematics, evolution and biogeography of Compositae, eds. V. A. Funk,
A. Susanna, T. F. Stuessy, and R. J. Bayer. Vienna: IAPT.
Roque, N. and H. P. Bautista. 2008. Asteraceae: Caracterização e Morfologia
Floral. Salvador: Edufba.
Roque, N., D. J. Keil, and A. Susanna. 2009. Illustrated Glossary of
Compositae. Pp: 781–806 in Systematics, evolution and biogeography of
the Compositae, eds. V. A. Funk, A. Susanna, T. F. Stuessy, and R. J.
Bayer. Vienna: IAPT.
Roque, N., L. Ramella, and P. Perret. 2011. Tipificaciones en el género
Trichogonia (DC.) Gardner (Compositae – Eupatorieae) de la Flora
del Paraguay. Candollea 66: 221–222.
Veloso, H. P., A. L. R. Rangel Filho, and J. C. A. Lima. 1991. Classificação da
vegetação Brasileira, adaptada a um sistema universal. Rio de Janeiro: IBGE.