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A Revision of Erato (Compositae: Liabeae)

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Systematic Botany (2006), 31(3): pp. 597–609
䉷 Copyright 2006 by the American Society of Plant Taxonomists

A Revision of Erato (Compositae: Liabeae)

EMILY MORAN1,3 and V. A. FUNK2

1
Duke University, Durham, North Carolina 27708-0338 U.S.A.;
2
U.S. National Herbarium, National Museum of Natural History, Smithsonian Institution,
Washington, DC 20013-7012 U.S.A.
3
Author for correspondence (emily.moran@duke.edu)

Communicating Editor: Thomas G. Lammers

ABSTRACT. Erato DC. contains five species, distributed from Costa Rica to Bolivia, with its main center of diversity in
Ecuador. The revision includes a new species endemic to Costa Rica and Panama, Erato costaricensis E. Moran & V. A.
Funk. Morphological and molecular data support Erato as a monophyletic group, sister to Philoglossa. The phylogenetic
analysis based on morphology used Munnozia Ruiz & Pavon, Chrysactinium (H.B.K.) Wedd., and Philoglossa DC. as outgroups.
The phylogeny supports the monophyly of Erato, but the relationships among the species within Erato have only weak
support. The genus is believed to be a recent radiation because of the morphological similarity among the taxa and their
location in some of the youngest areas of the Andes.
RESUMEN. El genero Erato contiene 5 especies, distribuidas desde Costa Rica hasta Bolivia, con su centro de distribución
en Ecuador. Este revista incluye una nueva especie que es endémica de Costa Rica y Panamá, Erato costaricensis E. Moran &
V. A. Funk. Datos morfológicos y genéticos confirman la hipótesis que Erato es un grupo monofilético hermano a Philoglossa.
El análisis filogenético utilizó Munnozia Ruiz & Pavón, Chrysactinium (HBK) Wedd. y Philoglossa DC. como grupos externos.
La filogenia confirma Erato como un grupo monofilético, pero los relaciones dentro de Erato solo tienen soportes débiles.
Créen que el genero es una radiación reciente a causa de la semejanzas morfológicas entre de las especies y porque occurien
en unos de los áreas mas jóvenes de los Ándes.
KEYWORDS: Asteraceae, biogeography, Compositae, endemic, phylogeny, taxonomy.

Erato DC. (Compositae), which has not previously which it shares stiff, thick-based hairs on the stems
been revised, contains five species. It is a member of and leaves, irregularly dispersed pollen spines, and a
the tribe Liabeae, which has approximately 180 species reduced number of achene ribs, two in Philoglossa and
in 15 genera, all confined to the Neotropics. Most of four in Erato.
the Liabeae are perennial herbs or shrubs; some are Characteristics that distinguish Erato from other
annuals, small trees, or climbers. Characteristics of the members of the Liabeae include ovate leaves that are
tribe include milky sap, opposite leaves, and arachnoid bright green above and paler below, blades that are
tomentum, but not all genera possess all three of these palmately veined with 5–9 main veins and dentate
characteristics. The complex history of the classification margins, and petioles that are often reddish. Arach-
of the Liabeae reflects the difficulty of both determin- noid tomentum, characteristic of most of the Liabeae,
ing the genera to be included in this tribe and inferring is almost totally lacking in Erato except for tufts on the
the relationships among them (Kim et al. 2003). apices of the involucral bracts in two species. In Erato
Erato is distributed from Costa Rica to Bolivia; three the indument usually consists of stiff, thick-based hairs
of the five species are native to Ecuador. Erato was orig- and the achenes are usually four sided. Members of
inally described by Candolle (1836) and was placed in Erato are coarse, upright herbs to shrubs; Munnozia
a position remote from other Liabeae. It was later species are lax shrubs with many-sided achenes, found
placed within Liabum by Bentham in Bentham and in open often sloped areas, Philoglossa species are small
Hooker (1873). In the generic revision of the Liabeae, herbs found in wet areas, with single heads arising
Robinson and Brettell (1974) placed it within a broad from the leaf axils, and the species of Chrysactinium
concept of Munnozia. It was restored to separate ge- are small acaulus herbs covered in tomentum and hav-
neric status during a study of the members of the tribe ing solitary heads.
in Ecuador (Robinson 1976, 1978) and since then has Our goals in this study were to use morphological
remained as such. characters, combined with some genetic data, to deter-
Currently, Erato is recognized as part of the subtribe mine the phylogenetic relationships among the species
Munnoziinae, which also includes the genera Munnozia of Erato and to identify the closest relative of Erato
Ruiz & Pavon, Chrysactinium (Kunth in H.B.K.) Wedd., within the Munnoziinae. Close examination of herbar-
and Philoglossa DC. This subtribe is distinguished by ium collections revealed that populations of Erato in
the presence of black or dark anther thecae. Philoglossa Costa Rica and northern Panama, previously identified
is usually identified as the sister group of Erato, with as E. vulcanica, were a distinct species.
597
598 SYSTEMATIC BOTANY [Volume 31

TABLE 1. Character List for Erato and outgroups.

1. Habit. Small herb (0), large herb to shrub (1)

2. Milky sap. Absent (0), present (1)
3. Petioles. Present (0), absent (0)
4. Number of main veins in leaves. 3 (0), 1 (1), 5–7 (2)
5. Pattern of leaf venation. Tri-nervate (0), pinnate (1), palmate (2)
6. Leaf dentation-1. Entire (0), irregular small teeth (1)
7. Leaf dentation-2. Entire (0), large regular teeth (1)
8. Base of hairs on leaves. Gradually tapering (0), bulbose (1)
9. Inflorescence form. Branched (0), single solitary (1), multiple solitary (2)
10. Inflorescence location. Terminal (0), axils of leaves (1), solitary (2)
11. Peduncle pubescens. Arachnoid and purplish hairs (0), stiff, erect, white hairs (1), appressed white hairs (2), long bulbose
based hairs (3), tomentose and glandular (4)
12. Tufts of arachnoid tomentum on bracts. Absent (0), few, scattered (1), many, dense (2)
13. Outer involucral bracts (both surfaces): without stiff hairs (0), with stiff hairs (1)
14. Number of main veins in involucral bracts. 3 (0), 5 (1), 7 (2)
15. Inner involucral bract l/w ratio. 5 or less (0), 7 or more (1)
16. Pales. Present (0), absent (1)
17. Number of ray florets. Less than 70 (0), 75–120 (1), 120–225 (2)
18. Ray floret length. 12.5–26.5 mm (0), 7–11 mm (1)
19. Number of disc florets. 30–100 (0), 25–33 (1), ⬍ 16 (2), ⬎ 100 (3)
20. Style length. 4.5–7 mm (0), 8–13 mm (1)
21. Pappus type. Long bristles (0), small squamell or awns in one series (1), absent (2), short, multiseriate awns (3)
22. Pappus persistence. Persistant (0), deciduous or possibly absent (1)
23. Achene indument. Pubulent (0), glabrous (1)
24. Achene keels. Absent (0), present (1)
25. Achene shape. Prismatic; 6–10 ribs (0), compressed; two ribbed (1), 4-sided (2)
26. Pollen spines. Regularly dispersed (0), irregularly dispersed (1)
27. Molecular data-1. Absent (0), several site mutations supporting the Munnozia-Chrysactinium clade as monophyletic (1)
28. Molecular data-2. Absent (0), several site mutations supporting the Erato-Philoglossa clade (1)

MATERIALS AND METHODS Outgroups. The monophyly of the four genera of the subtribe
Munnoziinae was demonstrated using ITS sequence data (Kim et
al. 2002); 13 base pair changes defined the node supporting the
Characters. The morphological study was based on specimens monophyly of the subtribe (98% bootstrap value). Therefore, Mun-
in AAU, MO, NY, and US, all of which have extensive plant col- nozia, Chrysactinium, and Philoglossa were included as outgroups.
lections from Ecuador and Peru. The data derived from the spec- It should be noted that, according this same molecular analysis,
imens were supplemented by information from the literature. For Chrysactinium is nested within Munnozia. However, since not all
microscopic examination, floral parts were rehydrated and mount- species of Munnozia were sampled and because of the disparity in
ed on microscope slides using Hoyer’s mounting solution. A total
morphology between the two, they are maintained here as sepa-
of 26 morphological characters were assessed, as well as two ge-
rate genera.
netic characters from a previous study by Kim et al. (2003; Tables
Data Analysis. Maximum parsimony analysis and parsimony
1, 2). The Kim et al. study used ITS sequence data to evaluate the
bootstrap analysis (with 1000 replicate runs, each with 10 random
monophyly of the subtribe Munnoziinae and to separate the four
taxon additions, TBR branch swapping, and MULPARS in effect)
genera and some of the species of the subtribe; two species of Erato
of the data matrix were preformed using full heuristic searches
were included in the study. Each genetic ‘‘character’’ in this study
with PAUP* (Swofford 2002). No weighting was used. Maximum
is derived from a well supported node on the molecular clado-
parsimony analysis (with ACTRAN) using a branch-and-bound
gram of Kim et al. (2003): ‘‘character 27’’ represents 19 base pair
search was also performed. The bootstrap runs employed 1000
changes (the node had a bootstrap value of 99%) and ‘‘character
replicates with branch-and-bound searches.
28’’ represents 16 base pair changes (the node had a bootstrap
value of 98%).
Most characters are self-explanatory (Table 1), but there are sev-
RESULTS
eral that may seem similar and therefore need some discussion.
Characters 4 and 5 may appear to be the same, but they are dis-
tinct in that one is the overall pattern of venation and the other is Because of morphological differences, specimens
the number of main veins. The removal of either of these charac- from Costa Rica previously identified as Erato vulcanica
ters from the analysis does not change the results. In characters 6 were described as a new species, E. costaricensis.
and 7, the small teeth are independent of the larger regular teeth,
and in characters 24 and 25, the compression and number of ribs
Maximum parsimony analysis yielded one most
are believed to be independent from the keels. Although there parsimonious tree for relationships within Erato; Fig. 1
were 28 characters in total, only five were informative within the is a phylogram of that tree with the branch lengths
genus Erato. representing the number of characters (L⫽51, ci⫽.804,
Not all specimens studied are listed in this paper; however, the
label information from all specimens used in this project has been ri⫽.655). Figure 2 is the bootstrap consensus tree. Erato
sent to MO to be deposited in their online database, TROPICOS. is monophyletic and sister to Philoglossa; E. polymnioides
2006] MORAN & FUNK: ERATO 599

is always the sister species to the rest of the genus.

28

0
0
1
1
1
1
1
1
Erato costaricensis, E. vulcanica, and E. sodiroi form a

27

1
1
0
0
0
0
0
0
monophyletic group and E. stenolepis, the Peruvian
species with the wide involucral bracts, is sister to that
26

0
0
1
1
1
1
1
1
clade. Bootstrap support for the monophyly of Erato
25

0
0
1
2
2
2
2
2
was 95% and 92% for the Erato/Philoglossa clade. The
relationships within Erato have short branches and
24

0
0
0
0
0
0
0
1
weaker bootstrap support with four of the taxa col-
23

0
0
1
1
1
1
1
0
lapsing into a polytomy and only 68% support for the
clade including all species except E. polymnioides. The
22

0
0
1
0
0
0
0
1
fact that the species of Erato are not sorting out in a
morphological analysis is not surprising, for although
0&1

1&2
21

0
0
0
0
3
the genus is unique in the family, and each species
within the genus has several apomorphies, there are
0&1
20

few synapomorphies among the taxa.

0
0
0
1
1
1
?

DISCUSSION
0&1
19

0
0
0
1
3
2

The species of Erato have a narrow range of mor-

phological and ecological diversity and this similarity
18

1
0
1
1
0
0
0
0

could be interpreted as the result of a recent radiation.

17

0
0
0
1
1
1
2
1

This scenario fits with the geologic history of the area.

The Andean Cordillera is thought to be of recent ori-
16

0
0
1
1
1
1
1
1

gin; it was (and continues to be) formed by the Nazca

15

0
0
0
0
1
0
0
0

plate colliding with the South American plate along

the Peru-Chile trench (James 1973; Jordan et al. 1983).
14

0
0
0
1
1
1
2
2

About three million years ago the Isthmus of Panama

13

0
0
1
0
1
0
0
1

first connected North and South America and there is

evidence of faunal movement across the isthmus at 2.8
12

0
0
0
0
0
1
2
2

MYA (Knowlton et al. 1991; Bermingham pers.

11

comm.). Sea level fluctuated several times; as little as

4
0
3
1
1
2
1
1

12,000 years ago it was lowered, exposing the isthmus.

10

0
2
1
0
0
0
0
0

At the same time the climatic zones in the Andes were

lowered (B. S. Vuilleumier 1975; Gentry 1982). It may
0
1
2
0
0
0
0
0
9

have been at this later date that Erato, along with some
0
0
1
1
1
1
1
1
8

other members of the tribe, Liabum, Munnozia, and Oli-

gactis, managed to colonize Central America and
0
0
0
1
1
1
1
1
7

southern Mexico. Molecular and morphological studies

1
0
1
1
1
1
1
1
6

(Funk et al. 1996; Kim et al. 2003) indicate that an an-

cestor of the extant members of the Munnoziinae was
0&1
0
0
2
2
2
2
2
5

most likely a beautiful herb from Ecuador and north-

ern Peru.
0&1
0
0
2
2
2
2
2
4

TAXONOMIC TREATMENT
0
1
0
0
0
0
0
0
3

ERATO Candolle, Prodr. 5:318, 1836.—Munnozia subg.

Erato (Candolle) H. Robinson & Brettell, Phytolo-
1
0
0
1
1
1
1
1
2

gia 28:56, 1974.—TYPE: Erato polymnioides Can-

TABLE 2. Data matrix for Erato.

dolle.
1
0
0
1
1
1
1
1
1

Perennial herbs to large shrubs, occasionally climb-

ers, 1–5 m tall, sap milky. Stems sparsely to densely
Erato polymnioides

Erato costaricensis

pubescent, hairs stiff, white; lengths of internodes

Erato stenolepis

Erato vulcanica
Chrysactinium

variable, usually 3–16 cm. Leaves opposite, ligh-

Erato sodiroi
Philoglossa

ter green abaxially and lacking tomentum; stipules

Taxa

Munnozia

0.5–5.0 cm long, broadly oblong, usually emarginate;

petioles 1–25 cm long, unwinged, often reddish in col-
or; blades ovate to broadly ovate; 3–27 ⫻ 3–27 cm, pal-
600 SYSTEMATIC BOTANY [Volume 31

FIG. 2. Bootstrap tree for the five species of Erato plus out-
groups; branches show bootstrap values.
FIG. 1. The most parsimonious tree for the five species of
Erato and its three outgroups. Munnozia and Chrysactinia are
shown as sister taxa because of information from a previous
publication (Kim et al. 2003; see text for details).

cells of anther collars not or weakly annulated on

walls, thecae 2.5–3.0 mm long, black, not digitate at
bases, apices 0.2–0.5 mm, acute; nectaries elongate,
mately veined, main veins 5–7 (–9), bases truncate, cor- slightly lobed; style branches short. Achenes four-sided,
date, rounded, attenuate, or slightly indented, some- one species with keel-like ridges on two sides, glabrous
times asymmetrical; margins usually irregularly den- or puberulous, 1.2–2.0 ⫻ 0.5–1.0 mm, light to dark
tate; apices acute to shortly and sharply acuminate; brown. Pappus of either 20–48 persistent pale bristles,
both surfaces strigose or pubescent, hairs short, ap-
4–8 mm long, or ca. 20, short, broad, fragile, pale,
pressed. Inflorescence terminal, loosely cymiform to
straw-colored scales (awns) 0.5–1.5 mm long. Pollen
densely subumbelliform; peduncles 1–13 cm long,
grains 30–40 ␮m in diameter, spines unevenly dis-
lightly to densely hispid, hairs stiff, white. Heads
persed, distinct internal columellae grouped under
broadly campanulate, usually 0.6–1.8 ⫻ 0.6–2.7 cm. In-
spines.
volucral bracts (phyllaries) 40–100 in 4–6 series, trian-
Distribution and Habitat. Erato is native to Costa
gular to lanceolate, sometimes with tufts of arachnoid
Rica, Panama, Venezuela, Colombia, Ecuador, Peru,
tomentum at apices, inner and outer series distinct;
outer bracts 3.0–10.5 ⫻ 1.7–3.0 mm, triangular to lan- and Bolivia at 360–3800 meters. Its members often
ceolate, main veins 5 or 7 main, margins ciliate, apices grow in open forest, pastureland, and forest edges, or
acute with herbaceous tips; inner bracts 7–12 ⫻ 1.5– along roadsides and streams, in part shade to full sun.
2.5 mm, lanceolate to oblong, usually lighter in color, Chromosomes. Erato generally has numbers of n
sometimes with ciliate margins, apices acute to round- ⫽7 or 9, in contrast to n ⫽18 in Philoglossa and n ⫽
ed. Ray florets 75–225, fertile; corollas yellow, 7.0–26.5 10, 11, 12, c. 13, c. 24 in Munnozia/Chrysactinium (Rob-
mm long, tubes 2.5–5.5 ⫻ 0.25–0.50 mm; laminae 5.0– inson et al. 1985).
22.5 ⫻ 0.5–1.0 mm, 2–3 apical teeth 0.3–0.5 mm long; Notes. Erato is easily recognizable because of its
styles 5.9–10.0 mm long, style branches 1–3 mm long. unusual leaves, which are opposite, shiny green, and
Disc florets 11–150, bisexual; corollas 6.5–9.0 mm long, palmately veined, with reddish petioles and margins
tubes 2.5–5.0 ⫻ 0.5 mm, throats 3.2–5.5 ⫻ 2.0–3.5 mm, that are irregularly dentate or with two levels of den-
lobes 2.5–3.0 mm long with stomates near margin and tation. Production of latex varies over time; sometimes
apices strongly spiculiferous; stamens 6–8 mm long, plants might seem to lack latex.
2006] MORAN & FUNK: ERATO 601

Key to Species of Erato (English)

1. Involucral bracts with 7 main veins; arachnoid tomentum at apices of involucral bracts in prominent or sparse tufts . . . . . . 2
2. Achenes puberulous with 2 prominent keel-shaped ridges; pappus of ca 20 short, broad, deciduous scales, 0.5–1.5 mm; ray
florets 90–120; disc florets 110–150; tufts of arachnoid tomentum mainly on the inside of involucral bracts; Ecuador . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. E. sodiroi
2. Achenes glabrous, without keels; pappus of 27–48 persistent bristles 4–7 mm long; 120–225 ray florets; 25–100 disc florets;
arachnoid tomentum sparse on outside of involucral bracts; Venezuela, Colombia, and Ecuador . . . . . . . . 5. E. vulcanica
1. Involucral bracts with 5 main veins; arachnoid tomentum at apices of involucral bracts either absent or, rarely, in small tufts . . . . . 3
3. Heads large, ca 120 disc florets; involucral bracts without tufts of arachnoid tomentum; surfaces of involucral bracts with
hairs; inner involucral bracts long, length/width ratio ⬎ 7; ray floret tubes glabrous; Peru . . . . . . . . . . . 3. E. stenolepis
3. Heads small, ⬍ 40 disc florets; involucral bracts rarely or never with tufts of arachnoid tomentum; surfaces of involucral
bracts with few to no hairs; inner involucral bracts short, length/width ratio ⬍ 5; ray floret tubes hairy . . . . . . . . . . 4
4. Heads small, 23–33 disc florets; ray corollas 7.0–11.5 mm long; involucral bracts never with tufts of arachnoid tomentum;
plants 1–5 m tall; peduncles with white, stiff, erect hairs; elevation 360–3050 m; Colombia, Ecuador, Peru, and Bolivia
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. E. polymnioides
4. Heads very small, 11–16 disc florets; ray corolla 13.0–16.0 mm long; involucral bracts occasionally with small tufts of
arachnoid tomentum; plant 0.5–3.0 m tall; peduncles covered with white, appressed hairs; elevation 1200–1700 m; Costa
Rica and Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. E. costaricensis

Key To Species of Erato (Spanish)

1. Brácteas involucrales con 7 nervios principale; tomento aracnoide en el ápice de las brácteas involucrales . . . . . . . . . . . . . 2
2. Aquenios puberulentos, con dos quillas prominentes; pappus de ca. 20 escamas cortas, anchas, y deciduas, 0.5–1.5 mm; flores
radiadas 90–120; flores del disco110–150; tomento principalmente en el reverso de las brácteas involucrales; Ecuador . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. E. sodiroi
2. Aquenios glabros, sin quillas; pappus de 27–48 cerdas persistentes, 4–7 mm de largo; flores radiadas 120–225; flores del disco
25–100; presente en Venezuela, Colombia y Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. E. vulcanica
1. Brácteas involucrales con 5 nervios principales; tomento aracnoide ausente o raro . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Capı́tulos largos, con ca. 120 flores del disco; brácteas involucrales sin tomento aracnoide; brácteas involucrales pubescentes en ambas
superficies, y brácteas interiores largas, largo/anchura ⬎7; tubo de flores radiadas glabros; Perú . . . . . . . . . . . 3. E. stenolepis
3. Capı́tulos pequeños, con ⬍ 40 flores del disco; brácteas involucrales raramente o nunca con tomento aracnoide; brácteas
involucrales glabras o con pocos pelos, y brácteas interiores cortas, largo/anchura ⬍5; tubo de las flores radiadas pubes-
cente . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Capı́tulos pequeños, con 23–33 flores del disco; flores radiatas 7.0–11.5 mm; brácteas involucrales sin tomento aracnoide;
arbusto de 1–5 m de altura; pedunculos 1.0–7.5 con pelos blancos cano-hirsutos; rango altitudinal 360–3050 m; presente
en Colombia, Ecuador, Perú y Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. E. polymnioides
4. Capı́tulos muy pequeños, con 11–16 flores del disco; flores radiadas 13.0–16.0 mm de largo; brácteas involucrales a veces
con un poco de tomento aracnoide; arbusto 0.5–3.0 m de altura; pedunculos cano serı́ceos; rango altitudinal 1200–
1700; Costa Rica y Panamá . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. E. costaricensis

1. Erato costaricensis E. Moran & V. A. Funk, sp. sometimes truncate or slightly indented, margins ir-
nov.,—TYPE: COSTA RICA. Cartago: Refugio Na- regularly dentate; apices shortly to sharply acuminate;
cional de Vida Silvestre Tapantı́, 14 Feb 1992, F. both surfaces with scattered, slender, short, appressed
Almeda 7001 (Holotype: US!; isotypes NY! CA). hairs. Inflorescence terminal, cymiform; peduncles 1.5–
Fig. 3. 8.2 cm long, densely pubescent, hairs appressed. Heads
broadly campanulate, usually 0.8–1.3 ⫻ 0.8–1.9 cm. In-
Similis Erato polymnioides sed: Frutex vel herba volucral bracts 50–70 in 5–6 series, oblong to lanceolate,
grossa 0.5 ad 3.0 m altus, aliquando scandens, pedun- usually without tufts of arachnoid tomentum; outer
culi cum pilis albis appressis, capitulum 0.8 ad 1.3 cm bracts 5–6 ⫻ 1.5–2.0 mm, triangular to oblong, main
altum, 0.8 ad 1.9 cm latum, flores disci 11–16, bracteae veins 5, margins lightly ciliate, apices acute; inner
involucralae apice persaepe cum caespibus tomenti ar- bracts lighter in color, oblong with hyaline margins,
achnoidei, bracteae externae 5 ad 6 mm ⫻ 1.5 ad 2.0 5.5–8.5 ⫻ 1.2–2.0 mm, apices rounded to acute. Ray
mm latae, flores radii corolla longiora, 13 ad 16 mm, florets 80–113; corollas yellow, 13–16 mm long, tubes
flores disci pauci 11 ad 16. 4.0–5.5 ⫻ 0.25 mm, sparsely puberulous distally; lam-
Perennial herbs to shrubs, 0.5–3.0 m tall, sometimes ina 9.0–10.5 ⫻ 0.3–0.5 mm, apical teeth 3, 0.3 mm long;
vine-like, sap milky. Stems terete, hairs scattered, stiff, styles ca 8.0 mm, style branches 1.5–2.5 mm. Disk flo-
white; internodes variable in length, usually 3.5–13.0 rets 11–16; corollas yellow, 7–9 mm long, tubes 2.5–3.5
cm; stipules 1.1–1.8 cm long, hairs sparse. Leaves darker ⫻ 0.5 mm, throats 4.5–5.5 ⫻ 2.5 mm, sparsely puber-
green adaxially, lighter abaxially; petioles 2–15 cm ulous distally, lobes 2–3 mm long; stamens 6–8 mm
long, reddish; blades ovate to broadly ovate, 10.5–22.5 long, thecae 2.2–2.5 mm long, black, apical append-
⫻ 4–20 cm, main veins 5–7, base usually rounded, ages 0.2 mm, acute; styles 7–10 mm, style branches 0.6
602 SYSTEMATIC BOTANY [Volume 31

FIG. 3. Erato coastaricensis E. Moran & V. A. Funk. A. habit, B. head, C. disc corolla, stamens and style, D. ray corolla and
style, E. disc style, F. achene and pappus. Illustration by Alice Tangerini (US).

mm, apices acute. Achenes 1.5 ⫻ 0.4–0.6 mm, brown, from Panama (Fig. 4). It is usually found in wet forest,
glabrous. Pappus of ca 30 setae, 4.0–6.5 mm long, white on forested hillsides, or in cut-over areas and along
to straw-colored. roadsides. It grows in part shade to full sun at 1,200–
Distribution and Habit. Erato costaricensis is 1,700 meters.
known mostly from Costa Rica with one collection Phenology. This species has been collected in flow-
2006] MORAN & FUNK: ERATO 603

FIG. 4. Map showing the distribution of Erato costaricensis

FIG. 5. Map showing the distribution of Erato sodiroi (Hi-
E. Moran & V. A. Funk.
eron.) H. Rob.

er in December, February, March, May, June, and Au- usually 3.5–8.0 cm; stipules 0.5–1.2 cm long, puberu-
gust. lous. Leaves darker green adaxially, lighter abaxially;
Notes. Erato costaricensis can be distinguished from petioles 1–10 cm long; blades ovate to very broadly
other species in the genus by its relataively small heads ovate, 4–16 ⫻ 2–14 cm, main veins 5–7, base truncate,
(disc florets 11–16), long ray florets (13–16 mm), ap- in older leaves slightly indented, in younger leaves of-
pressed, rather than bristly, hairs on the peduncles, ten attenuate, margins irregularly dentate, teeth coarse;
and a pappus of ca. 30 setae. apices shortly and sharply acuminate; adaxial surface
Representative Specimens Examined. COSTA RICA. Alajuela:
Vera Blanca - San Miguel rd, 17 Aug 1994, Kress 4810 (US); Vara- sparsely to densely strigose, abaxial surface with
blanca intersection on the rd to Puerto Viejo, 28 Feb 1986, Almeda dense, short, slender, appressed hairs. Inflorescence ter-
5172 (US). Cartago: Tapantı́ Hydroelectric Project, 25 Jun 1976, minal, loosely cymiform with few branches; peduncles
Utley 5173 (US); 25 Feb 1990, King 10001 (MO, US); S of Tapantı́, 3.0–12.0 cm long, densely hispid, hairs stiff, white.
12 Dec 1969 Burger 6790 (MO); 9 km NW of Tapantı́ Dam, 10 Aug
1980 Wilbur 30775 (MO). Heredia: 15 km NE of Santa Domingo, Heads broadly campanulate, usually 1.0–1.7 ⫻ 1.3–2.5
31 Dec 1974, Taylor 17868 (US); 35km. NE of Alajuela, 18 Aug 1967, cm. Involucral bracts 60–100 in 4–5 series, apices with
Taylor 4539 (MO, NY, US). San Jose: La Hondura, 2–4 Mar 1924, tufts of arachnoid tomentum; outer bracts triangular
Standley 36589 (US); La Palma, 17–18 Jul 1923, Maxon 8030 (US); to lanceolate, ca 7.0 ⫻ 3.0 mm, main veins 7, both sur-
Nubes, 13 Jun 1974, King 6782 (US); ca. 5 km N of Tunel Zarqui,
faces with stiff hairs, margins ciliate, apices acute with
5 Dec 1995 Hammel 20005 (INB, MO). Puntarenas: Monteverde Re-
serve, 10 Jan 1980, Funk 3063 (US); Monteverde, Mirador La Ven- herbaceous tip, 3–7 mm long; inner bracts lanceolate
tana, 9 Feb 1994, Lépiz 144 (CR, NY); Guanacaste, rd at Continental to oblong, ca 9.0 ⫻ 2.5 mm, lighter in color with hya-
Divide, 1 Nov 1977, Dryer 1111 (MO). line margins, apices rounded to acute. Ray florets 90–
PANAMA. Bocas del Toro: 5 km ENE of Cerro Pate Macho, 120; corollas yellow, 18.5–26.5 mm long, tubes 3.0–4.0
headwaters of Rio Culebra, 11 Feb 1979, Hammel 6146 (MO, PMA).
⫻ 0.25 mm, distally densely hirsute, hairs long; lamina
15.5–22.5 ⫻1.0–1.5 mm, apical teeth 3, 0.5 mm long;
2. ERATO SODIROI (Hieronymus) H. Robinson, Phyto-
styles 10 mm, style branches 3 mm. Disk florets 110–
logia 34:379, 1976.—Liabum sodiroi Hieron., Bot.
150; corollas yellow, 8 mm long, tubes ca. 4.0 ⫻ 0.5
Jahrb. 29:61, 1900. Munnozia sodiroi (Hieron.) H.
mm, throat 4.0 ⫻ 2.5 mm sparsely puberulous to hir-
Robinson & Brettell, Phytologia 28:57, 1974.—
sute distally, lobes 2.5 mm long; stamens 6 mm long,
TYPE: ECUADOR. Near San Florencio, Pallatanga
thecae 3 mm long, dark brown to black, apices 0.25
etc., growing in subtropical regions, s.d., Rev. P.(A.)
mm, acute; styles 10.5 mm, style branches 0.7 mm, api-
Sodiro S.J., s.n. (55/11) (B [destroyed]; Lectotype
ces acute. Achenes with prominent keel-shaped ridges
here designated: QPLS, photos of lectotype have
on two edges, ca 2 ⫻ 1 mm, brown, puberulous. Pappus
been examined).
of ca. 20 scales, 0.5–1.5 mm long, broad, fragile, pale,
Coarse herbs, occasionally scrambling or shrub-like, straw colored.
usually 1–3(–5) m tall, sap milky. Stems terete, some- Distribution and Habitat. Erato sodiroi is known
times hexagonal when dry, brownish, pubescence from Ecuador, in disturbed cloud forest, on steep road-
dense, hairs stiff, white; internodes variable in length, sides, and along creeks in wet forests (Fig. 5). It is
604 SYSTEMATIC BOTANY [Volume 31

sometimes described as climbing over shrubs. It grows

at 1780–2769 meters.
Phenology. This species blooms between late May
and early September, most commonly in July.
Notes. Erato sodiroi can be distinguished from oth-
er species in the genus by its short, easily deciduous
pappus scales and its puberulous, double-keeled
achenes.
Conservation Status. Vulnerable.
Specimens Examined. ECUADOR. Azuay: Cuenca, 3 Aug 1996,
Palacios 13812 (MO). Bolivar: Chillanes-Bucay rd, 1 Sep 1987, Zak
2680 (AAU, MO, NY, US); E of Chillanes on rd to Pallatanga, 23
May 1990, King 10202 (MO, US); Chillanes-Yaquibusu Rd, 20 Jul
1991, Van der Werff 12528 (NY, US); Chillanes-Tambillo-Trigoloma
rd, 5 Sep 1987, Zak 2756 (AAU, MO, NY, US); Guaranda-San Pablo
rd, 28 Aug 1987, Zak 2548 (AAU, MO, NY, US). Cañar-Chimborazo
border: Between Sta. Rosa and Joyagshi, 6–9 Jul 1945, Camp, E-
4034 (NY, US). Chimborazo: Chunchı́, 27 Jul 1959, Barclay 8314
(MO, US); Huigra 28 Aug 1918, Rose 22413 (NY, US); Cañon of the
Rio Chanchan, ca. 5 km N of Huigra 19–28 May 1945, Camp E-
3263 (MO, NY); N of the intersection of the southernmost entrance FIG. 6. Map showing the distribution of Erato stenolepis (S.
to Huigra and the Pan American Hwy on rd to Riobamba, 5 Jul F. Blake) H. Rob.
1992, Panero 2930 (US); Valle de Pallatanga, s.d., Spruce s.n. (NY);
Valle Pallatango and M. Chimborazo, s.d., Sodiro s.n. (BAF, pres-
ence confirmed but specimen not seen). Cotopaxi: E of Macuchi, es 1.5 mm, apices acute. Achenes four-sided, 1.25–1.5 ⫻
1–24 Jul 1982, Dodson 13440 (MO, US); Pilaloa-Macuchi rd, 17 Nov
1939, Haught 2959 (NY, US). 0.5–0.7 mm, brown, glabrous. Pappus of 24–45 setae, 5–
8 mm long, straw colored.
3. ERATO STENOLEPIS (S. F. Blake) H. Robinson, Phy- Distribution. Erato stenolepis is endemic to Peru,
tologia 28:43–63, 1974.—Liabum stenolepis S. F. found at elevations of around 2000 meters (Fig. 6).
Blake, J. Wash. Acad. Sci. 17:302, 1927.—TYPE: Notes. This species can be distinguished from oth-
PERU. Huanuco: Muña, trail to Tambo De Vaca, er species in the genus by its large heads and extreme-
2440 m, 27 Jun 1923, J.F. Macbride 4338 (Holotype: ly long and narrow inner involucral bracts. In his orig-
US!; Isotype: F, presence of isotype at F was con- inal description Blake quoted from the label that the
firmed, specimen was not seen). plant was a ‘‘Liana, flowers lemon-yellow.’’ (Blake
1927).
Large perennial herbs, occasional climber, sap Specimens Examined. PERU. Amazonas: Florida, 18 Jan 1983,
milky. Stems terete, hairs scattered to moderately King 9237 (US). Junin: La Merced, 15 Aug 1957, Hutchison 1190
(NY, US).
dense, stiff, white; stipules 0.6–1.5 cm long, sparsely
hairy. Leaves darker green adaxially, lighter abaxially;
4. ERATO POLYMNIOIDES deCandolle, Prodr. 5:318,
petioles 1.5–7.0 cm long; blades ovate, 6.5–13.0 ⫻ 3.5–
1836.—TYPE: PERU. ‘‘Peruvian mountains’’, 1791,
11.0 cm, main veins 5–9, base acute, margins dentate;
T. Haenke, 8161 (G-DC! Image of holotype seen).
apices acuminate; leaf surfaces densely strigose or pu-
Munnozia polymnioides (DC.) H. Robinson & Bret-
bescent, hairs short, appressed. Inflorescence terminal,
tell, Phytologia 28:56, 1974.
loosely cymiform with few branches; peduncles 1.0–
Liabum pallatangense Hieron. Bot. Jahrb. 29:60, 1900.
11.5 cm long, densely hispid, hairs stiff, white. Heads
TYPE: ECUADOR. Pallatanga valley and Piloton,
broadly campanulate, usually 1.2–1.8 ⫻ 1.1–2.7 cm. In-
s.d., Rev. P. Sodiro s.n. 55/12 (Holotype: B [de-
volucral bracts 72–88 in 4–5 series, margins ciliate; outer
stroyed]). Fig. 7.
bracts ca 10.5 ⫻ 2.0 mm, oblong to lanceolate, main
veins 5, both surfaces with stiff hairs, apices acute; in- Shrubs to coarse herbs, 1–5 m tall, sap milky. Stems
ner bracts ca 12.0 ⫻ 1.0–1.5 mm, lighter in color, elon- hexagonal when dry, older stems terete, brownish to
gate, margins sometimes darker in color, apices acute. reddish, usually pubescent, hairs stiff, white; inter-
Ray florets ca 100; corollas lemon yellow to greenish nodes variable in length, usually 4.0–16.0 cm; stipules
yellow, 16–20 mm long, tubes 6–8 ⫻ 0.25 mm, gla- 0.9–5.0 cm long, emarginate, hairs few. Leaves deep to
brous; lamina 10–12 ⫻ 1 mm, apical teeth 3, 0.5 mm bright green adaxially, lighter abaxially; petioles 2.0–
long; styles 9 mm, style branches 2.5 mm. Disk florets 25.0 cm long; blades ovate to very broadly ovate; 9–26
ca 120; corollas yellow, 8.0 mm long, tubes ca 4.0 ⫻ ⫻ 3–27 cm, main veins 5–7(–9), base usually cordate,
0.5 mm, throat 4.0 ⫻ 2.5 mm, hirsute at base, lobes 3 rounded, or attenuate, sometimes truncate or slightly
mm long; stamens 8.0 mm long, thecae 2.7 mm long, indented, margins singly or doubly dentate, teeth
black, apices 0.5 mm, acute; styles 6 mm, style branch- coarse; apices shortly and sharply acuminate; both
2006] MORAN & FUNK: ERATO 605

FIG. 7. Erato polymnioides DC. A. habit, B. hairs from disc corolla, C. head, D. ray corolla and style, E. disc corolla, F. disc
style, G. achene with pappus. Illustration by Alice Tangerini (US).

surfaces sparsely to moderately densely strigose or pu- mm, triangular to lanceolate, main veins 5, hairs ab-
bescent, hairs short, appressed. Inflorescence terminal, sent or few, margins ciliate, apices acute with short
usually strongly cymose to densely subumbellate; pe- herbaceous tip; inner bracts ca 7.0 ⫻ 1.5 mm, lanceo-
duncles 1.0–7.5 cm long, lightly to densely hispid, late to oblong, apices acute to slightly rounded. Ray
hairs stiff, white. Heads broadly campanulate, often florets 75–100; corollas yellow, 7.0–11.5 mm long, tubes
densely clustered, usually 0.6–1.8 ⫻ 0.6–2.4 cm. Invo- 2.5–4.5 ⫻ 0.25 mm, puberulous, lamina 5.0–7.5 ⫻ 0.5
lucral bracts 40–70 in 4–5 series, triangular to oblong, mm, 2–3 apical teeth 0.5 mm long; styles 5.9–7.0 mm,
no arachnoid tomentum; outer bracts 3.0–5.5 ⫻ 1.7–2.0 style branches 1.0–1.4 mm. Disk florets 23–33; corollas
606 SYSTEMATIC BOTANY [Volume 31

FIG. 8. Map showing the distribution of Erato polymoniodes FIG. 9. Map showing the distribution of Erato vulcanica
DC. (Klatt) H. Rob.

yellow, 6.5–7.0 mm long, tubes 3.0–3.5 ⫻ 0.5 mm, hir- 1943, Steyermark 54229 (NY); W of Piñas, 4 Feb 1979, King 7971
(US). Guayas: 21 km E of El Triunfo, 25 Jan 1979, King 7799 (US).
sute, throat 3.5 ⫻ 2.5–3.5 mm, glabrous except at base, Imbabura: Cotacachi, Hda. La Florida, 28 Aug 1992, Alvarez 630
lobes ca. 2.5 mm long; stamens 6.0 mm long, thecae (MO, US); Cotachi-Nangulbı́ and Apuela rd, 11–14 Aug 1990, Rubio
ca. 2.5 mm long, black, apices 0.25 mm, acute; styles 536 (AAU, MO, US). Loja: Loja-Zamora rd, km 15, 23–24 Apr 1988,
4.0–5.5 mm, style branches 1mm, apices acute. Achenes Madsen 74012 (AAU). Morona-Santiago: Sigsig-Chiquinda rd, 26
Oct 1995, Funk 11455 (US); Guarumales (Cola de San Pablo), 19–
four-sided, 1.2–1.5 ⫻ 0.5 mm, light to dark brown, gla-
20 Sep 1983, Larsen 45309 (AAU); 9–10 km SE of San Juan Bosco,
brous. Pappus of 20–40 setae, 4–6 mm long, persistent, 27 Jan 1981, Gentry 30845 (MO). Napo: Baeza, 28 Jul 1974, Plowman
pale. 3902 (US); Cerro Huacamayos, 9–10 Aug 1980, Øllgaard 35901
Distribution and Habitat. Erato polymnioides is na- (AAU, US); Cordillera del Guacamayo, 3 Aug 1984, Dodson 14850
tive to Ecuador, Peru, Colombia, and Bolivia. It is (US); Cosanga, 21 Feb 1978, Kirkbride 4271 (US); E of Papallacta,
26 Mar 1972, MacBryde 1265 (US); Salcedo-Napo rd, 4 Feb 1977,
found in primary, secondary, or disturbed moist for- Boeke 900 (AAU, NY, US); San Ramón, 8–12 Jun 1929, Killip 24529
ests, pastureland, scrub chaparral, steep rocky slopes (US); Yanayacu Biological Station, 28 Dec 2000, Clark 5784 (US);
and is often along roads or riverbanks or in pasture- Parque Nacional Llanganates, Salcedo-Tena rd, km 74–75, 14 Sep
land, usually in bright to partial sun. It is often locally 1998, Vargas 2349 (MO, QCNE); Cotundo-Coca, new rd km 2, 5
Aug 1984, Dodson 15025 (MO). Napo-Pastaza: Mera, in rastrojo, 4
abundant at elevations from 360–3050 meters. Fig. 8.
Apr 1956, Asplund 20135 (NY). Pastaza: Mera, 27 Mar 1968, Harling
Phenology. This species apparently blooms 7880 (US); 30 Jul 1980, Øllgaard 35575 (AAU, US); 5 km NE of Mera
throughout the year. Carretera al Rio Ansu, 3 Mar 1985, Neill 5941 (AAU, MO, NY);
Notes. Erato polymnioides can be distinguished Puyo 18–20 Feb 1935, Mexia 6957 (US); Veracruz, 24 Jun 1968, Lugo
from other species in the genus by its small heads with 36 (NY, US); 2 km N of Shell-Mera, 6 Jun 1968, Holm-Nielsen 350
(AAU, NY). Pichincha: Bosque Protector Maquipucuna 1 Sep 1993,
involucral bracts that are completely lacking tufts of Webster 30106 (US); Old rd to Chiriboga km 39, 23 Jul 1984, Dodson
arachnoid tomentum and by the bristly pubescence on 14348 (AAU, US); Cornejo Astorga (Tandapi), 7–10 May 1968, Har-
the peduncle. ling 9390 (US); Quito Cantón, 3 Dec 1996, Clark 3536 (MO, US);
Representative Specimens Examined. BOLIVIA. Cochabamba: Reserva Florı́stica-Ecológica ‘‘Rio Guajalito’’, 18 Aug 1985, Zak 566
Colomi- Tunari rd, 8 Feb 1978, King 7702 (MO, US). (AAU, MO, NY, PMA, US); Saloya, 28 Jun 1939, Asplund 7300 (US);
COLOMBIA. Caqueta: Florencia-Suaza rd, km 47, 20 Nov 1993 San Juan to Chiriboga, 23 Feb 1993, Funk 11044 (US); Lloa-Mindo
Ramirez 5329 (JAUM, MO). Valle: Buga-El Placer rd, 12 Sep 1991, rd, btw km 30–34, 4 Jul 1987, Zak 2114 (AAU, MO, NY); 13 kms
Silverstone-Sopkin 6369 (US). N of Nono, 21 Jul 1977, Stuessy 4861 (MO); 11 km W of Tandapi,
ECUADOR. Bolivar: Balsapampa, 19 May 1968, Harling 9645 26 Oct 1974, Gentry 12086 (AAU, MO); Chillogallo-Santo Domingo
(US). Cañar: 78 km W of Canar, 23 Jan 1979, King 7797 (US); Rd rd below Chiriboga, 13 Aug 1980, Holm-Nielsen 24850 (AAU); Par-
El Triunfo-Cañar, 5 Feb 1976, King 7002 (NY, US); 40km E of oquia Nanegal, Reserva Maquipucuna, 12 Jul 1990, Webster 28177
bridge at Guayaquil, 21 Jan 1979, King 7734 (US). Carchi: near (MO). Sucumbios: Reserva Ecológica Cayambe-Coca, 12 Aug
Maldonado, 31 May 1978, Madison 4823 (AAU). Chimborazo: Rd 1999, Vargas 3958 (MO, US). Tungurahua: Baños, 25 Sep 1923,
to Chillanes, 22 May 1990, King 10190 (US); Near Huigra, 7 Sep Hitchcock 21793 (NY, US); Btw Ambato & Baños, 10 Jan 1981,
1918, Rose 22583 (NY, US); 7–14 May 1945, Camp, E-3152 (MO, NY, D’Arcy 13998 (MO); NW of Puyo, 23 Jan 1974, King 6575 (NY, US);
US); Pallatanga, Jul 1903, Sodiro s.n. (US); s.d., Sodiro s.n. (BAF, Baños-Mera rd ca. 30 from Mera, 27 Mar 1983, Lawesson 43287
location confirmed but specimen not seen, MO); Jul 1903, Mille s.n. (AAU). Zamora-Chinchipe: Cordillera del Cóndor, 15 Dec 2000,
(NY). Cotopaxi: Macuchi, 1 May 1968, Harling 8872 (NY, US). El Montenegro 163 (MO, US); Loja-Zamora rd, 30 Dec 1980, Balslev
Oro: Trail from Sambotambo along hwy to Portovelo, 29 Aug 1264 (AAU, NY, US); Loja-Zamora rd, Km 24.5, 9 Aug 1997, Lewis
2006] MORAN & FUNK: ERATO 607

3423 (US); Nangaritza Canton Pachicutza, 18 Oct 1991, Palacios adaxially, paler abaxially; petioles 1.0–17.0 cm long, of-
8278 (MO, US); Tapichalaca Reserve, 29 May 2003, Clark 8084 (US);
ten reddish to purplish; blades ovate to very broadly
Rı́o Tundayme, 12 Dec 2000, Cerna 405 (MO, US); 4 km E of Pa-
quishsa, 6 Feb 89, Øllgaard 90432 (AAU, NY); Trail at Romerillo ovate, mostly 7–27 ⫻ 3–26 cm, main veins 5–7(–10),
towards Parque Nacional Podocarpus, 6 Dec 1988, Madsen 75860 base rounded, truncate, attenuate, or cordate, occasion-
(AAU). ally indented or asymmetrical, margins usually slight-
PERU. Amazonas: 17 kms N of Pedro Ruiz, 14 Jul 1995, Sanchez ly to strongly dentate; apices acute to shortly and
Vega 8015 (MO, US); Trail from La Peca to Serrania de Bagua, 13
Jun 78, Gentry 22847 (MO); Btw El Soro and Playa Granda, 4 Nov
sharply acuminate; both surfaces sparsely to densely
1991, Sanchez Vega 6027 (MO). Ayacucho: San Miguel, 7 Jun 1915, strigose or pubescent, hairs short, appressed. Inflores-
Cook 1111 (US). Cajamarca: Paraguay, 7 Aug 1994, Leiva 1372 (MO, cence terminal, cymose to subumbellate; peduncles 1–
US); San Ignacio-El Chaupe rd, 4 Jan 1995, Leiva 1567 (MO, NY, 13 cm long, densely hispid, hairs stiff, white to light
US); San José de Lourdes, 28 Oct 1995, Rodrı́guez 670 (MO, US); 4
brown. Heads broadly campanulate, usually 0.8–1.7 ⫻
Apr 1997, Campos 3750 (US); San Martı́n, 4 Aug 1994, Leiva 1337
(US). Cuzco: Ceja de Selva, Kosñipata, San Pedro, 28 Mar 1991, 1.3–2.5 cm. Involucral bracts 50–80 in 4–6 series, trian-
Nuñez 13364 (MO, US); Llactahuaman, N of Rı́o Apuramic, 14 Jul gular to oblong with tufts of arachnoid tomentum at
1998, Baldeon 3058 (US); Cord. Verónica, 30 Apr 1957, Raupt-Hirsch the apices; outer bracts 7.0–11.⫻ 2.5–3.5 mm, trian-
P1063 (NY); Machu Pichu, 21 Apr 1957, Raupt-Hirsch P8321 (NY). gular to lanceolate, main veins 7, margins ciliate, api-
Huánuco: Carpish, 3 Oct 1950, Ferreyra 8168 (MO, US); 16 Nov
1979, Jones 9234 (US); 15 Jun 1958, Humbert 30946 (NY); Rd along
ces acute, herbaceous to 4.0 mm; inner bracts 9.0–10.0
summit of Cerro Carpish, 28 Feb 1978, Luteyn 5481 (NY); Carpish ⫻ 1.5–2.5 mm, oblong to lanceolate with hyaline mar-
- Tingo Marı́a rd, 10 Jan 1986, Diaz 1989 (MO, US); Muña, 23 May- gins, apices rounded. Ray florets 120–225; corollas yel-
4 Jun 1923, Macbride 4054 (US). Junin: Huacapistana, 6 Jun 1929, low, 12.5–14.0 mm long, tubes 4.0–5.0 ⫻ 0.25 mm,
Killip 24282 (NY, US); Oxapampa, Aug 1944, Soukup 2421 (US);
sparsely puberulous above; lamina 7.5–10.0 ⫻ 0.5 mm,
Between Tarma and La Merced, 9 Sept 1972, Canne 268 (US); 51
km NE of Tarma, rd to San Ramon, 19 Dec 1978, Dillon 1426 (MO). apical teeth 3, ca 0.3 mm long; styles 7.0–7.5 mm, style
Loreto: 20 km NNE of Tingo Maria, rd to Pucallpa, 16 Jul 1981, branches 1.5–2.5 mm. Disk florets 25–100; corollas yel-
Dillon 2643 (MO). Pasco: Mallampampa, 22 Jan 1984, Smith 5842 low, 7.0–8.0 mm long, tubes 3.0–4.8 ⫻ 0.5 mm, sparsely
(MO, US); Oxapampa, 27 Aug 1968, Riccio 5701 (US); 30 May 1979,
puberulous distally, throat 3.2–4.0 ⫻ 2.0–3.0 mm, lobes
Teppner 18 (US). San Martin: La Divisoria, 14 Aug 1946, Ferreyra
1647 (MO, US); Pedro Ruiz-Moyobamba rd, 29 Jul 1983, Smith 4497 2.0 mm long; stamens 7.0–8.5 mm long, thecae 2.5 mm
(MO, US); Tingo Marı́a-Pucallpa rd, 17 Nov 1949 to15 Jan 1950, long, black, apices 0.4 mm, acute; styles 8.5 mm, style
Allard 21316 (US); 1 Nov 1949–5 Jan 1950, Allard 22063 (US); Plan- branches 1 mm, apices acute. Achenes four-sided, 1.2–
tación Margarita, 14 Aug 1946, Ferreyra 1056 (MO, NY, US). 1.8 ⫻ 0.5 mm, dark brown, glabrous. Pappus of 27–48
setae, 4–7 mm long, slender, persistent.
5. ERATO VULCANICA (Klatt) H. Robinson, Phytologia Distribution and Habitat. Erato vulcanica is native
34:379, 1976.—Liabum vulcanicum Klatt, Bot. Jahrb. to Colombia, Ecuador, and Venezuela where it is found
8:47, 1887.—TYPE: COLOMBIA. Cauca: Near Pu- in open, disturbed, or secondary forest, along forest
racé, western slopes of mountains, 2600–3200 m, edges and slopes, near streams, or along the roadside,
2 Jan 1884, F.C. Lehmann 3504 (Holotype: B [de- often in wet environments in open sun to part shade.
stroyed]; lectotype here designated: US!; isolec- This species is often locally abundant and is found at
totype: GH!, digital image seen). Munnozia vul- elevations from 1100–3800 meters, rarely as low as 255
canica (Klatt) H. Robinson & Brettell, Phytologia meters. Fig. 9.
28:57, 1974. Phenology. This species has been collected in flow-
Liabum anatinum Benoist, Bull. Soc. Bot., France 84:633, er during every month of the year, but in Colombia
1938.—TYPE: ECUADOR. Pichincha: San José de and Venezuela it is most commonly collected in June
Minas, flowers yellow, 3 Mar 1931, R. Benoist 3962 and July.
(Holotype: P; isotypes: PC). Munnozia anatina (Be- Notes. Erato vulcanica can be distinguished from
noist) H. Robinson & Brettell, Phytologia 28:56, other species in the genus by its largish heads bearing
1974.
involucral bracts with prominent tufts of arachnoid to-
Liabum insigne Badillo, Bol. Soc. Venez. Cienc. Nat. 10:
mentum, but without stiff hairs on both surfaces of the
313, 1946.—TYPE: VENEZUELA. Merida:
bracts. Like E. sodiroi it has seven main veins in the
Bosques húmedos, Los quebraditos, arriba de Jajı́,
involucral bracts but it has a glabrous achene while E.
2590 m, 21 Apr 1944, J. Steyermark 55981 (Holo-
sodiroi has a puberulous one.
type: VEN; isotypes: NY(2)!) [type from VEN was
This species is variable in the length of its peduncle.
seen as digital image].
In Ecuador in parts of Napo, Sucumbios, and Carchi
Coarse herbs to shrubs, 1–4(—5)m tall, sap milky. the peduncles are much longer than the usual 3–7 cm,
Stems terete, sometimes hexagonal when dry, brownish regularly reaching 8–10 and sometimes up to 13 cm
to reddish, hairs scattered to dense, stiff, white; inter- long. Some might chose to recognize these populations
nodes variable in length usually 3.0–15.0 cm; stipules as a separate species, but it seems more like a local
0.6–2.0 cm long, scarcely emarginate, hairs scattered, variant. Also, two or three specimens in this same area
particularly along margins. Leaves dark to light green have involucral bracts that are glabrous.
608 SYSTEMATIC BOTANY [Volume 31

Representative Specimens Examined. COLOMBIA. Antioquia: Pittier 733 (US); Rio Guadalajara, 27 Feb 1969, Cuatrecasas 27594
Páramo de Sonsón, 8 Jan 1956, Garganta Fabrega 2116 (US); de La (US); Rd to Cerro del Inglés, 22 Jul 2004, Pedraza 1115 (NY).
Ceja a Sonsón, 12 Apr 1951, Romero-Castañeda 2380 (AAU); Sonsón- ECUADOR. Carchi: El Carmen to Chical rd, Agua Amarilla, 8
Nariño rd km 11, 1 Apr 1987, Zarucchi 5210 (MO, NY, US); Gra- Jul 2003, Clark 8538 (US); NW side of Rı́o Gualpi Chico, 25 Jan
nada-San Carlos rd, 21 Feb 1989, MacDougal 4121 (MO, US); Las 1988, Hoover 3059 (MO, US); Páramo del Angel, Tulcán to Chicai
Palmas, 27 May 1948, Gutierrez 171 (US); La Planada Reserve, 25 Rd, 26 Feb 1995, Ortiz 430 (NY); Julio Andrade-El Carmelo rd, turn
Jul 1986, Gentry 55129 (MO, US); Rio Medellı́n, 27 Jan 1984, Juncosa off towards El Ajún, 10 Aug 1990, 92372 Jørgensen (AAU, US);
1966 (MO, US); 10 km NE Sonsón, 27 Mar 1979, Luteyn 7148 (NY, Above San Marcos de los Coaiqueres, 7 Feb 1985, Øllgaard 57457
US); Nutibara-Murri rd km 15.5, 22 Sep 1987, Zarucchi 5605 (NY, (AAU); 19 Jan 1983, Barfod 41505 (AAU). Imbabura: Pimampiro,
US); paraje Paramitos, Jul 1951, Uribe 2140 (US); San Jose de San 17 Sept 1988, Zak 3816 (AAU, MO, US). Napo: Baeza-Lago Agrio
Andreas, 1 May 1948, Correa 30 (US); Vereda La Corrala, 27 Apr Rd, 7 Aug 1980, Øllgaard 35625 (AAU, US); Hollı́n-Loreto rd, km
1987, Escobar 7590 (US); between Yarumal and the Llanos de Cui- 40–50, 10–22 Oct 1988, Hurtado 658 (AAU, MO, US); Reserva Ecol-
ba, 20 Feb 1942, Metcalf 30139 (MO, US). Caldas: Las Brisas to La ógica Antisana, 2 Jul 1995, Tirado 1583 (MO, QCNE, US); Btw Tena
Carbonera, 16 Oct 1946, Cuatrecasas 22182 (US); La Selva, 13 Jan and Papallacta, 12 Jan 1981, D’Arcy 14111 (MO, US); Slopes of Gua-
1946, Von Sneidern 5480 (US); 21 km ESE of Manizales, 16 Jul 1965, gua Urdu above Rı́o Borja, 25 Sep 1980, Holm-Nielsen 27025 (AAU,
King 5976 (NY, US); Salento, 27 Jul 1922, Killip 8825 (NY, US); US); Rı́o Panteor SW of Borja, 22 Sep 1980, Holm-Nielsen 26688
Pinares above Salento, 2–10 Aug 1922, Pennell 9402 (NY, US); Nev- (AAU). Pichincha: Near Quito, Valle Chillos, 1906, Mille 591 (NY,
ado del Ruiz, 15–16 Jul 1965, King 5966 (NY). Cali: Jugar, near La US). Sucumbios: Julio Andrada-La Bonita rd, 6 Mar 1992, Funk
Habana, 27 Feb 1969, Ramos 3674 (MO). Caqueta: 29 km SE of 11085 (US); El Salado, Rı́o Quijos, 27 May 1990, Cerón 10010 (MO,
US); San Rafael, NE of cascada, 11 Oct 1990, Jaramillo 13201 (NY);
Guadalupe on rd to Florencia, 9 Jan 1974, Davidse 5607 (MO, NY);
Playón de San Francisco, rd to Santa Bárbara, 30 Dec 1980, Jar-
30 km SE of Guadelupe, 9 Jan 1974, Gentry (AAU). Carpinterias:
amillo 3960 (AAU).
Forest between the cerros de Muchique y Altamira, 15 Jul 1939,
VENEZUELA. Mérida: La Carbonera, Jul 1957, Aristeguieta
Arbelaez 6132 (US). Cauca: Coconuco – Popayan rd, 19 Jun 1922,
2830 (NY, US); Btw Merida and La Azulita, 15 km SE of La Azul-
Pennell 6899 (US); ‘‘La Gallera’’ Micay valley, 1 Jul 1922, Killip 7965
ita, 7 Aug 1982, Croat 54824 (MO, US). Táchira: Btw Alcitran and
(NY, US); Cerro Munchique, 5 Nov 1968, Espinal 3194 (CUVC, MO,
San Vincente de la Revancha on rd to Las Copas, 3 Jan 1989, Hahn
US); ‘‘San Jose’’ San Antonio, 30 Jun 1922, Pennell 7563 (NY, US);
4951 (NY, US); Btw Bramón y Las Delicias, 15–16 May 1967, Stey-
Tambo, 13 Nov 1946, Haught 5246 (US); 22 Km NW of El Tambo,
ermark 98292 (NY); Mata Mula, N of Delicias, 26 Jul 1979, Steyer-
3 Jun 1977, Olsen 516 (NY); Parque Nacional Munchique, 25 Apr mark 118738 (MO, US); 20–25 KM.
1979, Luteyn 7437 (NY); Rio Palo, 15 Dec 1944, Cuatrecasas 19324
(MO). Choco: Cerro del Torra, 8 Aug 1988, Silverstone-Sopkin 4190
(MO, NY, US); un lugar denomina do La Mansa, 21 Jan 1949, Mo- ACKNOWLEDGEMENTS. We thank the following herbaria for the
lina 19Ch-31 (US); Rd Ansermanuevo-San José del Palmer, 18 Feb use of specimens and for information and digital images of types
1977, Forero 2891 (MO, NY). Choco-Antioquia: headwaters of Rio (AAU, BAF, BP, F, G, GH, K, MO, NY, P, PMA, QAP, QPLS, US,
Duro, 4 Mar 1944, Fosberg 21534 (US). Cundinamarca: El Colegio, VEN). Without such inter-institutional cooperation, revisions
would be impossible. We also thank Tom Hollowell, for assistance
13 Jul 1979, Stuessy 5529 (US); 21 km NW of Facatativa, 14 Jul 1965,
with graphics, and Pedro Acevedo for his help with the Spanish
Kinget al. 5934 (NY, US); 24 km NE of Fusagasuga, 19 Jun 1965,
translations. The Latin diagnosis was written with the help of Eliz-
King 5673 (NY, US); Salto de Tequendama, 1–3 Oct 1938, Cuatre-
abeth Wolfram and Harold Robinson and the illustrations were
casas 256 (type of fma. macrolepis Cuatre., holotype: US, photo of
prepared by Alice Tangerini. We thank Dr. Carols E. Ceron M.,
holotype: NY); 20 Aug 1942, Schultes 4032 (US); Apr 1972, Barclay
Director adhonorem del Herbario ‘‘Alfredo Paredes’’ (QAP) for
3309 (US); Tequendama Falls, near Bogota, 27 Feb 1945, Schiefer taking the time to make a special trip to visit QPLS and find the
469 (NY); Sibate, 23 Jun 1944, St. John 20527 (US); San Francisco, Sodiro material; John Clark (US) helped facilitate that search. We
7 Mar 1985, Sanabria 75 (US); 15 km NW of Medina, Gazaunta thank a number of individuals who spent time searching their
Valley, 24 Sep 1944, Grant 10275 (NY, US). Huila: Guadalupe en herbaria for types: Zoltán Barina (BP), Mireya Correa (PMA), B.
Resina, 20 Mar 1940, Arbelaez 8377 (US); E of San Antonio, 26 May C. Giberti (BAF), Nicholas Hind (K), Fernand Jacquemoud (G),
1944, Little 7950 (US); E of Neiva, 1–8 Aug 1917, Rugby 557 (NY); Christine Niezgoda (F), John Pruski (MO), Leyda Rodriguez
Around Mehrenberg rd from Popoyan, 6 Jul 1984, D’Arcy 15615 (VEN), Amber Swanson (GH), Anne-Elizabeth Wolf (P), and Em-
(MO). Medellin: Santa Elena, 20 Nov 1946, O’Donoghue 50 (US). ily Wood (GH). This study was undertaken as part of the Research
Meta: Rio Grande (?) S of Cordillera de las Cruces, 21 Aug 1943, Training Program at the National Museum of Natural History,
Fosberg 20880 (US). Nariño: Ipiales, 5 Aug 1939, Barriga 7781 (US); funded by the National Science Foundation; unfortunately this pro-
Pasto-Sandona Rd, 31 May, Vogelmann 2018 (US); Rd Tuquerres- gram has now been discontinued at NMNH.
Tumaco, 3 Apr 1989, Smith 1535 (US); Victoria, 23 Sep 1944, Ewan
16185 (US); La Planada Biological Reserve, 7 Km S of Chucunez,
10 Aug 1990, Luteyn 13990 (NY). Putumayo: between Sachamate
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BENTHAM, G. 1873. Compositae. Pp. 165–533 in Genera Plantarum,
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dean orogeny? Annals of the Missouri Botanical Garden 69: 557– ROBINSON, H. 1976. Studies in the Liabeae (Asteraceae), VI: Notes
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