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J. Bot. Res. Inst. Texas 9(1): 63 – 73. 2015
GNAPHALIOTHAMNUS NESOMII (ASTERACEAE: GNAPHALIEAE),
A NEW SPECIES FROM GUATEMALA AND NOMENCLATORIAL CHANGES
Michael O. Dillon Federico Luebert
Botany Department Universität Bonn
The Field Museum Nees-Institut für Biodiversität der Pflanzen
1400 South Lake Shore Drive Meckenheimer Allee 170
Chicago, IL 60605, U.S.A. D-53115 Bonn, GERMANY
mdillon@fieldmuseum.org
abstr act
Gnaphaliothamnus nesomii M.O. Dillon & L uebert (Asteraceae: Gnaphalieae) is a new species fr om the Sierra de los Cuchumat anes, De-
partment Huehuetenango, Guatemala. The generic boundaries w ithin the Gnaph alieae have been controversial and the genus Gnaphalio-
thamnus has not been universally accepted. New molecular phylogenetic studies support the acceptance of Gnaphaliothamnus as distinct
from Chionolaena, which i s congruent w ith cypsela tric home morphology. Two Gnaphalium species ar e transfer red as Pseudognaphalium
stolonatum (S.F. Blake) M.O. Dillo n and P. paramorum (S.F. Blake) M.O. Dillon.
key words: Chionolaena, Gamochaeta, Gnaphaliothamnus, Gnaphalium, Pseudognaphalium, Asteraceae, Gnaphal ieae, Gnaphalii nae, Guate-
mala, Lucilia-group, c omb. nov., sp. nov., Sierra d e los Cuchumata nes, taxonomy
resum en
Gnaphaliothamnus nesomii M.O. Dillon & Luebert (Asterace ae: Gnaphalie ae), es una nueva especie proveniente de Sierra de los Cuchu-
mata nes, Depart amento Huehuetenango, Guatem ala. Los límites genéricos e n Gnaphalieae son controvertido s y el género Gnaphaliotham-
nus no ha sido universalmente aceptado. Nuevos estudios filogenéticos molecul ares apoyan la aceptación de Gnaphaliothamnus como un
género diferente de Chionolaena, lo que es congruente con la morfología de los tricom as de las cipsel as. Dos e species de Gnaphalium son
tran sferidas c omo Pseudognaphalium stolonatum (S.F. Blake) M.O. Di llon y P. paramorum (S.F. Blake) M.O. Dillon.
palabras clave: Chionolaena, Gamochaeta, Gnaphaliothamnus, Gnaphalium, Pseudognaphalium, Asteraceae, Gnaphalieae, Gnaphaliinae,
Guatemala, Lucilia- group, comb. nov., sp. nov., Sierra de lo s Cuchumatane s, taxonomy
int roduc tio n
Gnaphaliothamnus Kirp. (Asteraceae, Gnaphalieae) comprises around 11 species distributed from Mexico to
Costa Rica. It has been accepted (Nesom 1990a,b; 1994) or subsumed in the South American Chionolaena DC.
(Anderberg & Freire 1989, 1991; Freire 1993; Nesom 2001; Freire et al. 2015). Based upon differences i n cypse-
lar (achenial) trichome s, Dillon (2003) argued that Gnaphaliothamnus was a monophyletic g roup not necessar-
ily close to Chionolaena. The cypselar trichomes in Gnaphaliothamnus are described (Hess 1938) as short cla-
vate (zwillingshaares) with an enlarged adaxi al basal cel l (schwellpolster) and myxogenic apica l cells 65–125 µm
long. The two apical cells in the cypselar trichomes in Chionolaena are much longer (250–850 µm long) and
not myxogenic (Dillon & Sagástegui 1991; Loeuille et al. 2011).
Pruski (2012) transferred a Guatem alan species, Gnaphalium stolonatum S.F.Blake, to Chionolaena, with the
comment th at it resembled other Mex ican and Central American members. He apparently did not examine the
type material of G. stolonatum, providing a new description drawn from a herbarium collection that he termed
a “topotype” (A. Molina et al. 16441, NY). From this voucher, he described the plants as reduced subshrubs to
30 cm tal l with clusters of capitula each havi ng 30–70+ pistillate florets and 11–25 hermaph roditic disc floret s.
He described the cypselas as setose with elongate trichomes, a condition very different from that described for
the type by Blake, i.e., minutely hispidulous with conical, bluntish, few-celled, dark-based hairs (Blake 1937).
An examination of the holotype of Gnaphalium stolonatum (A.F. Skutch 1098, GH) has shown it to be distinct
from other collections from the same general region annotated a s G. stolonatum. Gnaphalium stolonatum is here
treated as a Pseudognaphalium Kirp. and transferred to that genus. The other morphologically different collec-
This article has been licensed for personal, non-commercial use by the author(s) and publishers.
64 Journal of the Botanical Research Institute of Texas 9(1)
tions (e.g., A. Molina et al. 16441) are descr ibed here as a new species of Gnaphaliothamnus from t he Sierra de los
Cuchumatanes.
In the ultimate offering from H. Robinson (2015), he relegates Pseudoligandra M.O. Dillon & Sagást. (1990)
and Gnaphaliothamnus to the synonymy of Chionolaena. He states that the publications by Freire (1993) and
Nesom (2001) have “totally resolved that problem by reducing all three genera to synonymy under the name
Chionolaena.” While we have been un successful i n extracting DNA from the sa mples representi ng Pseudoligan-
dra at our disposal, we are confident that our nrDNA results have more definitively resolved the problem of the
recognition of Gnaphaliothamnus as distinct from Chionolaena.
taxonomy
Gnaphaliothamnus nesomii M.O. Dillon & Luebert, sp. nov. (Fig. 1). type. GUATEMAL A. Huehuetena ngo: common on
moist bank along road to San Juan Ixcoy, Sierra Cuchumatanes, 12–23 Jan 1966, A. Molina R., W.C. Burger, and B. Wallenta 16441
(holotype: F1637117; isotype: NY, n.v.)
Similar to Gnaphaliothamnus salicifolius (Bertol.) G.L. Nesom but habit considerably smaller (to 15 cm ta ll), capitulescences glomer ulate,
obscu red in dense a rachnoid-tome ntose indument um, cyps elar trich omes ca. 100 µm long
Subshrubs, 5–10(–15) cm tall; stems ascending, basally-branched from lignified base. Leaves sessile, oblan-
ceolate to spatulate, 10–42 mm long, 2– 4 mm wide, abaxial sur faces white-tomentose, adaxi al surfaces darker,
weakly arachnoid-tomentose, apices acute, apiculate, proximal leaves marcescent. Capitulescences densely
glomerulate with 7–11(–20) tightly grouped capitula, pedicles obscured by dense arachnoid-tomentose indu-
mentum. Capitula 5–7 mm long, 2–3 mm diam.; involucres narrowly campanulate, graduated, submerged in
dense arachnoid-tomentose indumentum; phyllaries 4–6-seriate, the outer with arachnoid-tomentose bases;
the inne r with white-opaque, oblong apical lamina, 5 –7 mm long, ca. 2 mm wide; outer floret s 20–32, pistill ate,
fertile; central disc florets 13–15, hermaphroditic, ovary sterile. Cypselas oblong, ca. 1 mm long, trichomes
4-celled, clavate, ca. 100 µm long, the apical cells myxogenic, not rupturing (Fig. 2A); pappus of ca. 20, setose
bristles, 2.8–3.2 mm long, apical cells of bristles of hermaphroditic florets expanded, apical cells of pistillate
florets obtuse, not expanded.
Etymology.—Gnaphaliothamnus nesomii is dedicated to Dr. Guy L. Nesom, noted synanthrologist and
systematist, one of the first botanists to accept Gnaphaliothamnus in modern usage, and first to provide a de-
tailed revision for Mexican and Central American taxa. Over the years, Dr. Nesom has been generous with his
detailed knowledge of Mexican and Central American Asteraceae and particularly the Gnaphalieae. He also
commented on the uniqueness of some of the sheet s examined within his loan of material from Field Museum
in 1990.
Distribution and Ecology.—The type locality is within the Sierra de los Cuchumatanes in the Department
of Huehuetenango, Guatemala [15°31'S, 91°32'W]. This is the highest non-volcanic mountain range in Central
America and has the most extensive highlands above 3000 m. The region is home to a variety of different bi-
omes, including montane pine-oak forest, intermittent shr ublands, and grasslands. Gnaphaliothamnus nesomii
is found in alpine habitats at ca. 3700 m, notably different from the environments associated with the “llanos”
or plains where Gnaphalium stolonatum occurs (see below).
Conservation status.— Gnaphaliothamnus nesomii deserves a preliminary status of Critically Endangered
(CR) because total area of its known distribution is less than 100 km² and only three populations are known
(I UCN 20 01).
Discussion.— Recent molecular phylogenetic studie s (nrDNA) have provided re sults that sug gest the great
majority of New World genera of Gnaphalieae are associated with the Lucilia-group within the FLAG clade
(Freire et al. 2015; Luebert et al., unpubl.). The Lucilia-group has been expanded to include Gnaphaliothamnus,
Chionolaena, and Antennaria Gaertn. from the subtribe Cassiniinae Anderb. (Anderberg 1991). These results
support the hypothe sis that the gross morphological similar ity between Gnaphaliothamnus and Chionolaena is
convergence; they are found in distant well-supported clades w ithin the Lucilia-group (Fig. 3). Gnaphaliotham-
nus is phylogenetically related to Antennaria, while Chionolaena appears to be sister to a group including the
type species of Lucilia Cass., which is in agreement with their similarity in cypselar trichomes.
Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala 65
Fig. 1. Gnaphaliothamnus nesomii. Photograph of holotype collection, A. Molina R. et al. 16441 (F1637117, imgV0093984F).
66 Journal of the Botanical Research Institute of Texas 9(1)
Fig. 2. Cypselar trichomes, white scale bar = 100 µm. A. Gnaphaliothamnus nesomii (F1637117), B. G. salicifolius (F1639348), C. Pseudognaphalium
stolonatum (GH00008351), D. P. antennarioides (F1148157).
Nesom (1990a) provided a detailed taxonomic history for Gnaphaliothamnus and his rationales for its ac-
ceptance (Nesom 1990b, 1994). Interestingly, Nesom also discussed the putative relationships for Gnaphalium
stolonatum as outside of Gnaphaliothamnus (Nesom 1990a, p. 367). He commented that this taxon did have
white-tipped phyllaries, but it had more florets per capitulum in general (+100) and fertile central disc florets,
Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala 67
Fig. 3. Simplified cladogram of the Lucilia-group derived from preliminary result of a Bayesian analysis of Gnaphalieae with nrDNA sequence data
(ITS+ETS) carried out in MrBayes v.3.1.2 (Ronquist & Huelsenbeck 2003). The numbers above branches are Bayesian posterior probabilities. The results
are congruent with those reported by Freire et al. (2015). Chionolaena and Gnaphaliothamnus do not form a clade.
68 Journal of the Botanical Research Institute of Texas 9(1)
Table 1. Comparison of salient morphological characters in the species of Gnaphaliothamnus and Pseudognaphalium discussed here. Cypselar trichomes are
illustrated in Figure 2.
Character G. nesomii G. salicifolia P. stolonatum P. antennarioides
Habit subshrubs to 15 cm subshrubs to 100 cm perennial herbs perennial herbs
Stolons none none present present
Capitula height 5–7 mm 6–7 4.5–5.5 mm 6–7 mm
Pistillate florets 20–32 (22–)34–55 44–92 (50–)75–190
Hermaphroditic florets 13–15 3–4(–7) 7–14 7–16
Cypselar trichomes ca. 100 µm long, Fig. 2A 62–78 µm long, Fig. 2B 38–40 µm long, Fig. 2C ca. 50 µm long, Fig. 2D
and suggested it was probably best maintained in Gnaphalium L. s.s. There has been acceptance of Pseudog-
naphalium and the majority of New World species previously classified as Gnaphalium were transferred there
(Anderberg 1991).
Gnaphaliothamnus nesomii was originally treated as Gnaphalium stolonatum. In a note from Guy Nesom
(16 Feb 1990) accompanying the return of his Field Museum loan, he mentioned the presence of some un-
usual or atypical specimens with fewer pistillate florets. When Pruski (2012) encountered what he considered
conspecific material (A. Molina 16441, NY), he transferred the species to Chionolaena and described its floral
morphology exactly as it is in Gnaphaliothamnus. He related it to other Mexican and Central American species,
including C. eleagnoides Klatt, C. lavandulifolia (Kunth) Benth & Hook. f., C. concinna (A. Gray) Anderb. & S.
E. Freire (as C. mexicana S. E. Freire), and C. salicifolia (Bertol.) G.L. Nesom, all species considered here to be-
long within Gnaphaliothamnus.
Gnaphaliothamnus nesomii is distinctive among members of Gnaphaliothamnus. It has a dwarf woody veg-
etative habit with stems only 5–10(–15) cm long, the basal leaves are marcescent and cloaking the lower por-
tions of the stems, and the glomerulate capitulescences with the heads immersed in dense, white, arachnoid-
tomentose indumentum, is a combination of characteristics unmatched in the genus. It has oblanceolate to
spathulate leaves to 42 mm long, white-opaque phyllary apices, and capitula with 20–32 pistillate florets, and
13–15 hermaphroditic florets, Nesom’s (1990a) key would lead to G. salicifolius; which is the only other mem-
ber of Gnaphaliothamnus to be recorded from Guatemala, and it has a widespread distr ibution extending nor th
from Guatemala to central Mexico.
Gnaphaliothamnus salicifolius has a distribution quite unlike all other species, which tend to be narrowly
endemic and geographically restricted (Nesom 1990a,b, 1994). In Guatemala, it inhabits the edge of pine-oak
forests at about 3000 m and is also found in the Sierra de los Cuchumatanes, recorded on the road between
Paquix and San Juan Ixcoy (A. Molina R. 21293, F1661761; A. Molina R. et al. 30031, F1734420; A. Molina R. et al.
16550, F1639348) and N of Santa Eulalia (F. Almeda and J.L. Luteyn 1686, F17 33 978).
Table 1 allows for comparison of the salient morphological characteristics of three of the Gnaphalieae
taxa known to occur in region around the Sierra de los Cuchumatanes.
Additional material examined: GUATEMALA. Huehuetenango: Sierra Cuchumatanes between Paqui x and San Juan Ixcoy, 8 Jan 1974,
3000 –3350 m, A. Molina R., A.R. Mo lina, and J.A. Mol ina 30055 (F1734422); between Tojquiá a nd Caxín blu ff, summit o f Sierra de los Cuch u-
mata nes, 3700 m, 6 Aug 1942, J.A. Ste yermark 50159 (F114815 0).
key to g naph al ioth am nus spec ies
The following annotated key will allow for identification of Gnaphaliothamnus species (adapted from Nesom
1990a):
1. Inner phyllaries lacking prominent, white lamina; pappus bristles monomorphic (Chiapas - Mexico) _________ G. cryptocephalus
G.L. Nesom
1. Inner phyllaries with prominent, white lamina.
2. Adaxial leaf surfaces with stipitate glandular trichomes beneath the arachnoid-tomentum.
3. Pistillate florets 5–10; pappus bristles strongly dimorphic, caducous; cypselas glabrous (Costa Rica) _________ G. costaricensis
G.L. Nesom
Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala 69
3. Pistillate florets 12–24; pappus bristles monomorphic to weakly or strongly dimorphic, basally persistent; cypselas
with trichomes.
4. Pistillate florets 12–18, usually equal the number of hermaphroditic; pappus bristles strongly dimorphic (Oaxaca
- Mexico) _______________________________________________________________________ G. macdonaldii G.L. Nesom
4. Pistillate florets 21–24, usually about twice as many as the hermaphroditic; pappus bristles monomorphic to very
weakly dimorphic (Veracruz, Puebla, Tlaxcala, Morelos - Mexico) _________________ G. lavandulifolius (Kunth) G.L. Nesom
2. Adaxial leaf surfaces arachnoid-tomentose to glabrate, eglandular.
5. Leaves 7–8 mm long; phyllaries subequal (Oaxaca - Mexico) _______________________________ G. sartorii (Klatt) G.L. Nesom
5. Leaves shorter than 5 mm or longer than 10 mm long; phyllaries strongly graduated.
6. Heads few in tight clusters at tips of leafy stems.
7. Plants dioecious; leaves 2.5–5 mm long; phyllaries with reddish midregion (Oaxaca - Mexico) _______ G. aecidiocephalus
(Grierson) G.L. Nesom
7. Plants polygamodioecious; leaves 10–20 mm long; phyllaries without a red midregion (San Luis Potosí -
Mexico) _________________________________________________________________ G. concinnus (A. Gray) G.L. Nesom
6. Heads numerous in corymbs above the leaves.
8. Leaves elliptic to elliptic-oblanceolate; pappus bristles weakly dimorphic.
9. Leaves elliptic to elliptic-oblanceolate, 15–42 mm long, 4–8 mm wide (Hidalgo, Oaxaca - Mexico)
_______________________________________________________________________G. eleagnoides (Klatt) G.L. Nesom
9. Leaves elliptic-obovate, 10–20 mm long, 3–5 mm wide (Durango - Mexico) _____________ G. durangensis G.L. Nesom
8. Leaves linear to oblanceolate or spathulate; pappus bristles strongly dimorphic.
10. Leaves 20–80 mm long, 1–3(–5) mm wide; pistillate florets (22–)34–55; hermaphroditic florets 3–4(–7)
(Guatemala, Mexico) ___________________________________________________ G. salicifolius (Bertol.) G.L. Nesom
10. Leaves 10–42 mm long, 2–4 mm wide; pistillate florets 20–32; hermaphroditic florets 13–15 (Guatemala)
__________________________________________________________G. nesomii M.O. Dillon & Luebert
new com binations in p seud ogna ph ali um
Pseudognaphalium stolonatum (S.F. Blake) M.O. Dillon, comb. nov. (Fig. 4). Gnaphalium stolonatum S.F. Blake, Brit-
tonia 2:341. 1937. type: GUATEMALA. HueHuetenango: llanos of the Sier ra Cuchumatanes, 10,500 ft [3200 m], 24 Aug 1934, A. F.
Skutch 1098 (Holotype: GH00008351; isotype: TEX-L L003737 32, n.v.).
Chionolaena stolonata (S.F. Blake) Pr uski, Phytoneu ron 2012-1:4. 2012.
Perennial herbs 10–25 mm tall, roots fibrous; stolons from the base, filiform; stems erect, usually simple,
rarely branched, thinly arachnoid-tomentose, purplish. Leaves oblanceolate to linear, 2–4.5 mm long, 0.5–1
mm wide, abaxial surfaces gray with dense tomentose indumentum, adaxial surfaces obscurely stipitate-
glandular under thinly arachnoid-tomentose indumentum, apices apiculate, bases slightly decurrent. Capit-
ulescences glomerulate, crowded at apic es. Capitula campanulate, ses sile, 4.5–5.5 mm tall, 4.5–5.5 mm d iam.;
phyllaries ca. 4-seriate, graduated, thinly arachnoid-tomentose; outer ovate to oblong-lanceolate, obtuse to
acute, inner narrowly lanceolate, narrowed to an obtuse apex; pistillate florets 85–92, the corollas filiform,
white, ca. 2.5 mm long, pappus bristles ca. 3 mm long, apices acute; hermaphroditic florets (7–)11(–14), the
corollas cylindrical, white with rose-color lobes, the styles apically subtruncate, ovary sterile; pappus bristles
ca. 3 mm long, apically obtuse, slightly expanded. Cypselas ca. 1 mm long, minutely hispidulous, the tri-
chomes clavate, 38–40 µm long (Fig. 2C).
Pseudognaphalium stolonatum has been collected within the Sierra de los Cuchumatanes from alpine
meadows on rock outcrops, 3200–3750 m. It was included in Gnaphalium s.s. by Anderberg (1991). Inspection
of the type has shown it to have floral morphology consistent with Pseudognaphalium (Anderberg 1991).
Blake (1937) specifically related Pseudognaphalium stolonatum to P. antennarioides (DC.) Anderb., a rosu-
late, stoloniferous herb distributed from Colombia to Bolivia with capitula having (50–)75–190 pistillate and
7–16 hermaphroditic florets. Phylogenetic studies have shown that Pseudognaphalium is a member of the HAP
clade (Smissen et al. 2011; Nie et al. 2013; Galbany-Casals et al. 2014), along with Achyrocline (Less.) DC., He-
lichrysum Mill., and Anaphalis DC .; Gnaphaliothamnus and Chionolaena are members of the FL AG clade, which
is only distantly related (Smissen et al. 2011).
Pseudognaphalium subsericeum (S.F. Blake) Anderb. (= Gnaphalium subsericeum S.F. Blake, 1927), a Costa
Rican endemic, was also mentioned as potentially related to Gnaphaliothamnus salicifolius by Blake (p. 62). An
examination of conspecific material from Costa Rica (F1692005, F1692006) shows P. subsericeum more simi-
70 Journal of the Botanical Research Institute of Texas 9(1)
Fig. 4. Pseudognaphalium stolonatum. Photograph of holotype collection of Gnaphalium stolonatum S.F.Blake, A.F. Skutch 1098 (GH000008351).
Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala 71
Fig. 5. Pseudognaphalium paramorum. Photograph of holotype collection of Gnaphalium paramorum S.F.Blake, A. Jahn 883 (US1186590, img00129548).
72 Journal of the Botanical Research Institute of Texas 9(1)
lar to P. stolonatum or P. antennarioides (DC.) Anderb., the latter an Andean species that has similar overall
morphology and cypselar trichomes (Fig. 2D). All these species have capitula in dense cymes or glomerules
and phyllaries with showy, white apices.
Additional specimens examined: GUATEMALA. Huehuetenango: vicinit y of Chemal, sum mit of Sierra de los Cuchumat anes, 3700–3750
m, 8 Aug 1942, J. A. Ste yermark 50253 (F1148157), 50273 (F114818 0), 50275 (F114817 9).
Pseudognaphalium paramorum (S.F. Blake) M.O. Dillon, comb. nov. (Fig. 5). basionym: Gnaphalium paramorum S.F.
Blake, J. Wash. Acad. Sci. 21:328. 1931. type: VENEZUELA. Estado Mérida: Páramo Quirorá, 2900 m, 24 Feb 1922, A. Jahn 883
(Holotype: US118 6590).
Gamochaeta paramora (S.F. Blake) Anderb., Ope ra Bot. 104:157. 1991.
Blake (1931) commented that Pseudognaphalium paramorum had the appearance of Gnaphalium antennarioides
DC. (= P. antennarioides (DC.) Anderb.), and he related it to that taxon, which is a rosulate, stoloniferous herb
distributed from Colombia to Bolivia.
Anderberg (1991) transferred Gnaphalium paramorum to Gamochaeta Wedd.; an examination of the type
collection shows it to only superficially resemble Gamochaeta. It has a basal rosette of subspathulate leaves and
unbranched, ascending stems. The terminal capitulescences are not spicate; they are densely glomerulate cor-
ymbs. The t ype collection has capitular and flora l morphology consistent with Pseudognaphalium; the cy pselas
are papillose and lack the 2-celled, sessile trichomes diagnostic for Gamochaeta.
acknowled gments
We thank the curators and staff of the herbaria at F, GH, MO, NY, TEX, and US for permitting access to their
material. Especially, we wish to acknowledge Ms. Emily Wood, Collections Manager at the Gray Herbarium,
who expedited loan of type material from that institution. We thank John Strother for many constructive sug-
gestions in the review process. Andrés Moreira-Muñoz and Miguel Álvarez provided plant material for phylo-
genetic studies. Digital images were obtained via the Internet in Figure 4 from GH and Figure 5 from US. We
than k Field Museum’s Christine Niezgoda, Anna Balla, Daniel Le, and Allie Stone for hand ling various aspects
of loans and digitizing collections.
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