Phytoneuron

Pruski, J.F. Lowryanthus rubens {Compositae: Athroismeae), a new genus and species from southeastern Madagascar.
Phytoneuron 2014-51: 1-11. Published 20 May 2014. ISSN 2153 733X

LOWRYANTHUS RUBENS (COMPO SITAE: ATHROISMEAE),
A NEW GENUS AND SPECIES FROM SOUTHEASTERN MADAGASCAR

John R Pruski

Missouri Botanical Garden

P.O. Box 299
St. Louis, Missouri 63166

ABSTRACT

The new genus and species Lowryanthus rubens Pruski (Compositae) from Madagascar is
described. By paleate elinanthia. tailed anthers. 2-banded styles branches, and especially by the
strange carbonized, obcompressed, geniculate-rostrate cypselae, Lowryanthus Pruski is placed in
African- centered tribe Athroismeae, which contains only 3 subtribes, 7 genera, and about 60 species.
Although the epappose long-ciliate cypselae and open paniculate capitulescence make confident
placement of Lowryanthus in either sub tribe Anisopappinae or Athroisminae difficult the carbonized
obcompressed cypselar character of Lowryanthus is used here to place the new genus in
Athroisminae. However, the three other genera of Athroisminae are totally different and
characterized by glomerate capitulescences and typically white corollas.

Exploration in Madagascar by a team of botanists from the Missouri Botanical Garden has
resulted in the discovery of a hitherto unknown, red-flowered, discoid-capitulate Compositae. This
striking plant (Fig. 1) was brought to my attention in 2007 by Porter P. (Pete) Lowry II (MO) and is
described here as monotypic Lowryanthus Pruski and placed in tribe Athroismeae, which is
characterized in part by its strange, carbonized, obcompressed, geniculate-rostrate cypselae (Figs. 2,
51)). An early draft of this manuscript was reviewed in 2007 by Henk Beentje (K) and Arne
Anderberg (S), but validation of the plant was withheld pending examination of possible newer
collections, the existence of which I was aler ted to in 2008 by Pete Lowry. In. 20 13 this second suite
of collections, made by J. Rabenantoandroin in 2008, were processed and mounted at MO. These
2008 specimens were seen recently during routine herbarium specimen filing, and their mature, black,
carbonized, geniculate-rostrate cypselae confirm mis important character as given in original draft
description of Lowryanthus. I do not anticipate that possible additional collections will change
drastically either the description or circumscription of the plant, and it now appears appropriate to
validate the new genus and species, apparently endemic to Madagascar.

LOWRYANTHUS RUBENS Pruski, gen. et sp. nov. TYPE: MADAGASCAR. Toliara. Anosy Region,
Bemangidy Forest, ca. 3 km W of Antsotso, along RN 12a, 65 km N of Ft. Dauphin, humid
evergreen forest on steep slope, E of Ivohibe peak, 24° 35' 02" S, 47° 12 44" E, 100-250 m,
7 Feb 2006, P.P. Lowry II, J. Rabenantoandro, F. Randriatafika, E. Lowry, E, Ramisy, & B.
Mara 6648 (holotype: MO; isotypes: P [photograph: MO], S, TAN). Figs. 1-5.

Plantae herbaceae perennes vel iruticosae 1-4 m altae; caules rubri subtereti hirsutuli; folia
simplicia alterna petiolata, lamina oblanceolata vel anguste obovata 5-19(-25) >< 2-7.5(-9.5) cm chartacea
pinnativenia eglandulosa hirsutula denticulata, trichomatibus patentibus uniseriatas multi eel hilar ibus,
petiolo 2-3.5(-5) cm longo; capitulescentia terminalis corymbiformis vel paniculata, pedunculis 7-20 mm
longis, capitula discoidea paleacea, involucrum turbinatum vel campanulatum 7-9 x 4.5-8.5 mm,
phyllaria 5-8 circiter 2-seriata oblanceolata vel obovata 6.5-8 x 2-5.5 mm, chartacea rubra; clinanthium
paleaceum; paleae late lanceolatae vel oblanceolatae 7-1 1 x 1.5-2.5 mm stramineae vel rubrae; flosculi
disci 1 0-15, corolla anguste campanulata quinquelobata 4-5.8 mm longa rubra vel rosea, tubo 1 .5-2. 1 mm
longo glanduloso necnon piloso, glandulis biseriatis brevistipitatis, Umbo anguste ampliato, lobis 1.2-1.5

Pruski: Lowryanthus rubens (Compositae: Attroismeae) 2

Pruski: Lowryanthus rubens (Compos\tae: Athroismeae) 3

Figure 2. SEM micrograph ol con ! e\ ahaual lace ot art uumatui e opsela of
owr) ! / rt he vn g m ulitt 1 ostium (v j p . n Jit) (Antilahii 1
4801, MO; sample preparation method is as given in Pruski 2012).

Perennial herbs to shrubs, 1-4 m tall; stems ascending to erect, few-branched distally,
striate, subterete to angled or even somewhat flattened in capitulescence, reddish, slightly hirsutulous
distally, leafy in the proximal half with somewhat congested internodes, these becoming well-spaced
distally, internodes 1^1(-12) cm long, pith solid. Leaves simple, alternate, petiolate; blade
progressively reduced in size distally, oblanceolate to narrowly obovate, 5— 19(— 25) x 2-7.5(-9.5) cm,
chartaceous to stiffly so, venation pinnate, secondary veins departing from midrib at angles of 45° or
greater, third order venation obscure, surfaces eglandular, hirsutulous. trichomes patent, uniseriate,
multicellular, without well-developed subsidiary cells, abaxial surface of immature leaves sometimes
densely hirsute, blade bise alttimate mn,> petiole, margins denticulate, the closely spaced callous-
tipped teeth usually ca. 0.5 mm long, apex obtuse to rounded; petiole narrowly winged to base, 2-
3.5(-5) cm long, canaliculate, rose-colored, broadened basally and subclasping (subamplexicaul) the
stem, f ipitnlisiiiMf m Mb turnup] plutn. ipilul it>. open, _ 01 vuhi form-paniculate, branching

Pruski: Lowryanthus rubens{Compos\t&t: Athroismeae) 4

alternate appearing more or less divaricate, branches reddish, often 1-bracteolate, slightly hirsutulous,
bracteoles typically positioned toward proximal half of branch, oblanceolate, 5-8 x 1-2 mm wide;
peduncles reddish, somewhat stout, 7-20 mm long, slightly hirsutulous, often l(-4)-bracteolate,
bracteoles resembling those of the capitulescence branches, often positioned {Q-)2-A mm proximal to
subtended capitulum. Capitula discoid, 10-15-flowered, 10-12 mm tall; involucre turbinate to
narrowly campanulate, 7-9 x 4.5-8.5 mm, sometimes appearing double by closely subtending
peduncular bracteoles; phyllaries 5-8, oblanceolate to obovate, 6.5-8 x 2-5.5 mm, subequal or
slightly graduated, loosely imbricate, ca. 2-seriate, erect, reddish, bracteoles and outer phyllaries not
articulated basally, stiffly chartaceous, obscurely ca. 5-7-nerved, glabrous or short-ciliate distally,
apex somewhat cucullate especially in bud, disk corollas partly exserted from top of involucre;
chnanihmm palealc hemi^pheijcal ro clavate, 1.5^.5 x 1-2.5 mm; paleae broadly lanceolate to
oblanceolate, 7-11 x 1.5-2.5 mm, nearly planar grading to the inner ones somewhat conduplicate,
chartaceous, stramineous with rose-colored apices, 3-5-nerved, glabrous or short-ciliate distally,
apically rounded to less commonly truncate. Ray florets none. Marginal pistillate florets
apparently none. Disk florets 10-15, bisexual, 5-merous, the outermost sometimes aborted; corolla
actinomorphic, narrowly campanulate, 5-lobed, 4-5.8 mm long, tissue moderately thick, tube 1.5-2.1
mm long, reddish with pinkish lobes, broadened basally around nectary, with short-stipitate biseriate-
glandular trichomes (similar to those in Eriksson 1995: 118, Fig. 4), also often pilose, the limb
moderately ampliate, throat glabrous or stipitate-glandular, lobes 5, 1.2-1.5 mm long, reflexed to
slightly so, subsessile-glandular, 2-nerved, nerves intramarginal; filaments 1.2-1.5 mm long,
glabrous, collar same diam. as filaments, shorter than the tails, collar cell walls evenly thick, anthers
often fully exserted, ca. 2.2 mm long, pinkish, caudate (sterile-tailed) or calcarate-caudate (tails partly
polleniferous), thecae ca. 1.5 mm long, endothecial tissue polarized, thecae and connectives
somewhat discolorous, tails ca. 0.7 mm long, distal tail surface short-papillose, papillae apices
rounded, tails of adjacent thecae appressed to slightly connate, apical anther appendage elliptic-ovate,
ca. 0.3 mm long, longer than wide, well-differentiated from thecae, slightly concave longitudinally,
non-sculptured; pollen markedly echinate, white; style 4.7-6 mm long, red, exappendiculate, trunk
3.5^1.2 mm long, basally glabrous, non-enlarged, branches 1.2-1.3 mm long, broadly linear,
tangentially spreading to recurved, each with a 2-banded stigmatic surface, apically acute to obtuse,
with short sweeping papillae, papillae apices rounded, style immersed in basal nectary, nectary ca, 0.6
mm long, commonly positioned asymmetrically atop rostrum. Cypselae obcompressed (tangentially
flattened), obovate in outline, 3.7-5 x to 2 mm, rostrate, epappose, body and rostrum black-
carbonized, eglandular, faces finely roughened, phytomelanin deposits first manifest in spots,
glabrous to sparsely pilose distally, abaxial face convex, adaxial face concave, body ultimately with 2
well-developed lateral shoulders, the rostrum thus appearing as arising from a slight recession on the
top of the fruit, margins to 0.4 mm broad, sometimes brownish when maturing, spreading-antrorse
long-ciliate, trichomes multicellular, biseriate-celled, to ca. 2.5 mm long, falsely resembling pappus
(nearly as to position on the technically epappose cypselae), but ontogenetically not homologous and
not borne upon the apical annulus, terminal trichome cells erect and non-diverging, rostrum broad, ca.
0.7 mm tall, strongly deflected (geniculate) abaxially, annulus a minute ring atop rostrum;
carpopodium nearly annular, ca. 0.3 mm long, somewhat broadly triangular in cross-section,
stramineous. Chromosome number unknown.

Paratypes. MADAGASCAR. Toliara. Bemangidy, Commune Iabakoko, Fokontany
Antsontso, Ivohibe forest, 24° 34' 13" S, 47° 12 01" E, 286 m, 23 May 2006, Antilahimena et al.
4801 (MO, P, S, TAN, TEF); Anosy, Fort Dauphin, Iabakoho, Antsotso, 24° 34' 15" S, 47° 12 05" E,
280 m, 2 Apr 2008, Rabenantoandro et al. 1895 (MO, P, TAN); Anosy, Tolagnaro, Iabakoho,
Antsotso avaratra. Foret dense humide de basse altitude dTvohibe. 24° 34' 03" S, 47° 12 09" E, 270
m, 3 Apr 2008, Rabenantoandro et al. 1903 (MO, P, TAN); Fort-Dauphin, Iabokoko, Antsotso, Foret
Ivohibe, 24° 33' 52" S, 47° 14' 25" E, 112 m, 26 Nov 2005, Razakamalala et al. 2369 (MO, P, S,
TAN).

Pruski: Lowryanthus rubers (Compositae: Atrroismeae) g

Etymology. I am both pleased and privileged to dedicate this new genus to Porter P. (Pete)
Lowry II (MO), director of MBG's Madagascar program and collector of the type material. Pete
kindly and enthusiastically showed me photograplis of his then-unrecognized, reel- flowered
Compositae collection and shortly thereafter routed the initial collections from 2005 and 2006 to me
for identification. The epithet refers to the crimson-red branchlets, phyllaries, and corollas.

4

▲ Lowryanthus rubens iW/ 1 "^^

A

I fryr- is /
r VsY \ 1

\ a * c /
\ T\ ) J

Figure 4. Distribution map (with the five major biozones of
Madagascar indicated) of Lowryanthus rubens.

Habitat and ecology. Lowryanthus rubens is known from only a handful of collections
made within a few kilometers of each other in the Bemangidy-Ivohibe forest, downhill from Ivohibe
peak and about 50 kilometers north of Fort Dauphin in southeastern Madagascar. Lowryanthus
rubens occurs between about 100-286 meters elevation in low-elevational humid evergreen for ests on
the eastern slope of the Tsitongambarika range, the easternmost mountain chain in southern
Madagascar and basically bordering the Indian Ocean The Bemangidy-Ivohibe forest is of special
interest as it is among the few remaining regional low-elevational wet forests. The species has been
collected in flower from November to May and is pollinated possibly by insects. I do not know the
soil type where the plants grow. Five of the seven known genera of Athroismeae occur in
Madagascar, but only Lowryanthus is endemic there.

Figure. 5. Microscopic features of Lowryanthus ruber'.!: A. Bisk floret showing corolla and immature eypsela.
The arrow points toward trichomes which in the close-up are seen as biseriate with erect non -diverging apical
cells. B. Two post-anthesis stamens showing tailed anthers (arrow points toward the end of tails). C. Bifid
style. D Close-up of ob compressed nearly mature eypsela shewing abaxial geniculate rostrum (arrow) (Lowry
etal. 6648, MO).

Pruski: Lowryanthus rubens{Compos\t&t: Athroismeae) 8

Malagasy Lowryanthus has paleate clinanthia, tailed anthers, style branches with 2-banded
stigmatic surfaces, and obcompressed carbonized genicu late-rostrate cypselae (Figs. 2, 5D) and is a
member of the Heliantheae Alliance (subfamily Asteroideae sensu Bremer 1994; Panero 2007a), but
upon first dissection its tribal placement was not at all immediately apparent to me. Within
Asteroideae, Inuleae (including Plucheeae) is the only traditionally recognized tribe characterized in
part by consistently tailed anthers. It was basically similar to Inuleae, from which African-centered
Athroismeae Panero, Cuban Feddeeae Pruski et al., and cosmopolitan Gnaphalieae Anderberg were
segregated (viz Anderberg 1991; Cariaga et al. 2008; Panero 2005, 2007a, 2007b), and that is where
the affinities of Lowryanthus were sought. Lowryanthus would be clearly anomalous, however, in
either Feddeeae or Gnaphalieae.

Although the character of tailed anthers only places Lowryanthus somewhere in the general
vicinity of Inuleae sensu lato (sensu Merxmiiller et al. 1977) and segregate tribes, it is the very
strange, obcompressed, carbonized, asymmetrically rostrate cypselae character (viz Eriksson, 1990,
1992, 1995; Figs. 2, 5D; described by Eriksson in 1990 as "abaxially curved") that specifically allows
Lowryanthus to be placed near the genera of the African-centered Blepharispermum group, treated
earlier by Merxmiiller et al. (1977) as Inuleae.

The three genera of the Blepharispermum group (Athroisma DC, Blepharispermum Wight ex
DC, Leucoblepharis Am.) since have been placed in Heliantheae subtribe Ecliptinae (Eriksson 1990,
1991) as "unassigned to a subtribe" within tribe Helenieae (Karis and Ryding 1994: 536) and based
on molecular studies were placed in the newly described Athroismeae (viz Panero 2005, 2007a,
2007b). Panero (2005) expanded Athroismeae from 3 to 5 genera by adding the new monogeneric
Centipedinae Panero and Anisopappinae Panero. Panero (2007b) added epaleate Welwitschiella O.
Hoffm. to Anisopappinae, but Brouillet et al. (2009) placed this genus in Astereae. Ortiz (2010)
described Cardosoa S. Ortiz, a sixth genus of Athroismeae. The present addition of Lowryanthus
raises to 7 the number of genera (each monographed) recognized in the relatively small African-
centered Athroismeae (Table 1), which contains 3 subtribes, 7 genera, and about 60 species.
Subtribes, genera, species numbers, distributions, primary references, and a key to subtribes and

'-I IllK Ulll.'IMlK.K l.'ll.'U

Table 1. The subtribes, genera, species numbers, and distributions of Athroismeae.

I. Athroisminae

1 . Athroisma DC [as Polycline
Oliv. in Humbert 1960-1963]

12 species; Africa, Madagascar, 1 sp.
■ I ..I ..„ 1 ■ ■■ ■

. . .- .- .■ .-■ 1 i.-l.i . I ■■

15 species; Africa, Madagascar, Asia.

■ I nl i---: .

3. Leucoblepharis Arn.

1 species; India.
■ I ,,l ! ■

4. Lowryanthus Pruski

1 species; Madagascar.

■ 1 i ii 1 i _■ ■ U | i . .1,1 |- ,|-.i ■

II. Anisopappinae

5, Amsopappus Hook. & Arn. [as
EpallageDC. in Humbert 1960-

17^40 species; Africa, Madagascar, 1

1963]

(Wild 1964; Eldenas and Anderberg
1996; Ortiz et al. 1996; Ortiz 2005;
Panero 2005).

6. Cardosoa S. Ortiz

1 species; Africa.
(Ortiz 2010).

III. Centipedinae

7. Centipeda Lour.

10 species; mostly circumaustral, also
Madagascar.

(Walsh 200 1 ; Nylmder et al. 20 1 3).

Pruski: Lowryanthus rubens (Compositae: Athroismeae) 9

Key to subtribes and genera of tribe Athroismeae (modified from Panero 2007b)

1. Cypselae brown to black, not carbonized, subterete to compressed.

2. Capitula disciform, qialeate; innermost disk florets 4-merous; cypselae epappose;

(Centipedinae, 1 genus) Centipeda

2. Capitula radiate, paleate; disk florets 5-merous; cypselae pappose; (Anisopappinae, 2 genera),

3. Capitulescence lax corymb if onn; marginal florets radiate; disk corollas strictly

actinomorphic; anther appendage acute apically; cypselae 10-20-costate Anisopappus

3. Capitulescence racemiform-cymose; marginal florets subbilabiate; disk corollas slightly
zygomorphic, anther appendage apieulate apically; cypselae < 10-costate Cardosoa

1. Cypselae black, carbonized, obcompressed; (Athroisminae, three core genera + Lowryanthus).

4. Capitulescence open; capitula discoid; corollas red Lowryanthus

4. Capitulescence of globose glomerules; capitula discoid or disciform; corollas mostly white.

5. Capitula 4-6-flowered; inner florets functionally stamina te; cypselae pappose

Blepharispermum

5. Capitula 4-47-flowered; disk florets bisexual; cypselae epappose.

6. Capitula short-bracteate; twin trichomes of cypselae with diverging or recurved apices

Athroisma

6. Capitula long-bracteate; twin trichomes of cypselae with erect apices Leucoblepharis

Within Athroisminae, the non-glomerate, red-flowered Lowryanthus is very different from
the three other genera, each of which lias densely glomerate, globose capitulescences and typically
white corollas. However, by long marginal and apical twin trichomes on the epappose cypselae
Lowryanthus is somewhat similar to Athroisma and Leucoblepharis. Additionally, Leucoblepharis
has trinervate (vs. pinnately veined) leaves; Blepharispermum has occasionally thorny (vs. unarmed)
stems, unisexual (vs. bisexual) disk florets, and cypselae with (vs. without) a true pappus of awns or
scales; and Athroisma has diverging to recurved (vs. erect) apical ceils of the twin trichomes of the
cypselae, further distinguishing these three genera from Lowryanthus. The two non-typical subtribes
have non-carbonized cypselae and either radiate capitula or innermost disk florets 4-merous, thereby
differing from Lowryanthus.

I should also note here that the combination of morphological characters diagnostic for
Lowryanthus is unmatched in any of the detailed regional treatments of Humbert (1923, 1960-1963),
Hind et al. (1993), Beentje (2000, 2002), Beentje et al. (2005), and nothing similar can be found in
the family overview of Bremer (1994). Thus, the striking Lowryanthus rubens is described as a new
genus and species and placed in tribe Athroismeae. Molecular support of the affinities of
Lowryanthus awaits study, which I suspect will show that a new subtribe is needed to accommodate
Lowryanthus.

ACKNOWLEDGEMENTS
I would like to thank Arne Anderberg (S) and Henk Beentje (K) for reviewing the manuscript;
Henk van der Werff (MO) for correcting the Latin description; Porter P. (Pete) Lowry II (MO) for use
of his field photographs; Pete Lowry, Guy Nesom, and Rosa Ortiz for helpful comments on the
manuscript; Frederic Tronchet (P) for sending me photographs of the specimens of Lowryanthus in P;
and Cynthia Hong-Wa (MO) for helpful discussion and for sending me the base map used here.

Pruski: Lowryanthus rubens (Compositae: Athroismeae) 10

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