New combinations in Luciliocline with notes on South American Gnaphalieae (Asteraceae)

16 Pages·2003·7.8 MB·English

by  Michael Dillon

45-60. 2003
10(1):
New
Combinations in Luciliocline with notes on South American
Gnaphalieae
(Asteraceae)
MICHAEL DILLON
O.
Botam
Department
of
Museum
The Field
USA
Chicago, IL 60605-2496,
a lichimuseum.org
i/ilion
There once was a farmer who could not between
distinguish
his two horses, so he cut the the white horse and
tail off left
on
the the black one. Turner
tail - Billie L.
"Habia una vez un campesino quien no podia entre
distinguir
sus dos por eso
caballos, corto cola del caballo bianco
la y
dejo cola del caballo negro
la
».
Resumen
Se proporciona la revision de los representantes sudamericanos de la tribu
&
Gnaphalieae Cass, ex Lecoq (Asteraceae), incluyendo una discusion de
Julliet. las
Una
posiciones de las subtribus. revision de morfologia en conjunto de generos
la los
asignados «grupo Lucilia» revelo necesidad de una nueva
al la circunscripcion para
el
grupo; estos cambios han requerido la transferencia de varias especies. Las nuevas
&
combinaciones
propuestas son: Luciliocline longifolia (Cuatrec. Aristeg.) M.O.Dillon
& &
comb,
Sagast.. nov., L. pickeringii (A.Gray) M.O.Dillon Sagast, comb, nov,
L.
& &
piptolepis (Wedd.) M.O.Dillon comb, nov,
Sagast., L. plicatifolia (Sagast.
& L &
M.O.Dillon) M.O.Dillon Sagast., comb, nov, radians (Benth.) M.O.Dillon
Sagast.,
&
comb, nov, (Wedd.). M.O.Dillon comb, nov,
L. schultzii Sagast., L. spathulifolia
&
& &
M.O.Dillon) M.O.Dillon comb, nov,
(Sagast. Sagast., y L. turneri (Sagast.
&
nov
M.O.Dillon) M.O.Dillon comb,
Sagast.,
A
review of the South American representatives of Gnaphalieae
tribe Cass, e*
&
Lecoq
Juillet (Asteraceae) is provided, including a discussion of subtribal positions.
A
review of morphology of
the overall the genera assigned to the "Lucilia group" revealed
new
need
the for a circumscription for the group; these changes required the transfer of
The new
several species. combinations proposed Luciliocline
are: longifolia (Cuatrec.
& &
&
Aristeg.) M.O.Dillon Sagast, comb, nov, pickeringii (A.Gray) M.O.Dillon
L.
&
Sagast, comb, nov, L. piptolepis (Wedd.) M.O.Dillon comb,
Sagast., nov, L.
& &
plicatifolia (Sagast. M.O.Dillon) M.O.Dillon comb, nov, radians
Sagast., L.
&
&
(Benth.) M.O.Dillon Sagast., comb, nov, L. schultdi (Wedd.). M.O.Dillon
Sagast.,
& &
comb, nov,
L. spathulifolia (Sagast. M.O.Dillon) M.O.Dillon comb, nov,
Sagast.,
& &
and turned
L. (Sagast. M.O.Dillon) M.O.Dillon comb.
Sagast., nov.
Introduction
This paper based upon an and Phytogeny
is oral presentation abstract entitled,
and
South American
Classification of the Inuleae
(sens, ampl.) delivered the
at
International Compositae Conference, Royal Botanical Gardens, Kew, 24 July 5 August
-
1994. That paper presented the relationships amongst genera based upon
detailed
&
observations (Dillon Sagastegui, 199 and morphological
la) cladistic analysis (Dillon,
2000). In October 2000, a modified version of this paper was posted on the ABIS website
and an abbreviated version presented here background new combinations.
as for
is
South American Gnaphalieae
The
Inuleae ampl. Asteraceae) included South American
traditionally the taxa
s. (
&
now
two
accepted in segregate tribes, Gnaphalieae Cass, ex Lecoq and
Juillet
Plucheeae Anderb. (Bremer, Worldwide, Gnaphalieae
1994). the contains 187 genera
and about 1250 species (Bayer South America, Gnaphalieae
et in press). In the
al.,
consists of perhaps 22 genera and over 00 species with highest diversity in the tropical
1
and Andean The
subtropical Cordillera. majority of genera are Neotropical endemics, but
some
are cosmopolitan or Pantropical, Achyrocline, Gamochaeta, and
e.g.,
Pseudognaphalium,
however,
their greatest species diversity is in South America.
Anderberg 99
provided a morphological of Gnaphalieae
cladistic analysis the
( 1
1 )
that included 72 genera, utilizing 82 characters to establish five subtribes and several
putative monophyletic groups. His treatment ultimately included 46 genera or over half
1
He
added
intuitively. placed South American genera into three subtribes: Cassiniinae
Gnaphaliinae
Anderb., (Cass.) Dumort, and Loricariinae Anderb.
Subtribe Gnaphaliinae South America
in
The
subtribe Gnaphaliinae (sensu Anderberg) consisted of a group of genera with
worldwide diagnosed
distribution as annual to perennial herbs (occasionally subshrubs).
I
)
capitula with marginal pistillate florets greatly outnumbering the central hermaphroditic or
staminate and oblong achenes with pubescence of clavate trichomes. The
florets, short,
subtribe encompassed 47 genera, with centers of diversity in Africa, Asia, and the
No
Neotropics. fewer than 14 genera were partially or wholly distributed in South America.
Anderberg proposed several informal groups, as suggested by his cladistic analysis and
group"
"Lucilia
Anderberg 99 established the "Lucilia group" diagnosed as herbs or subshrubs,
)
( 1
1
with polychromous phyllaries, pistillate florets with generally yellow corollas, hermaphroditic
florets with generally purple corollas, and the corolla veins ending below the apex of the
The
lobes. group initially contained Lucilia, Belloa, Chevreulia, Jalcophila,
Cuatrecasasiella, Berroa, and Facelis. Subsequently, Anderberg and Freire 99
( 1
1
Gamochaeta
considered taxon group" and Micropsis and
as the sister to the "Lucilia
were
Stuckertiella included subsequent to their cladistic analysis. Freire 986, 987)
( 1 1
published species-level cladistic analyses where both Belloa and Lucilia were combined
form an expanded Anderberg and 99
to Lucilia. Freire published another cladistic
( 1
)
1
analysis where the expanded Lucilia was dismantled and constituent taxa placed
in
its
new
four genera: a revised Belloa a revised Lucilia and two
(1) (9 spp.), (2) (8 spp.),
&
genera, Gamochaetopsis Anderb. and Anderb.
S.E.Freire Luciliocline
(3) (1 sp.), (4)
&S.E.Freire(5spp.).
To
test the various hypotheses of relationships presented by Anderberg 99 and
( 1 )
1
Anderberg and Freire 99 an independent morphological analysis ofthe "Lucilia group"
( 1 ),
1
was undertaken The
(Dillon, 2000). analysis utilized 35 discrete characters analyzed
PAUP
with (Swofford, The genus Chionolaena was
1991). (including Leucopholis)
The
used as the outgroup. analysis yielded a basal polytomy with Chionolaena,
Lucilia,
Gamochaetopsis, and two These
additional c lades. results suggested a close potential
between Chionolaena and members of Both
relationship the other the "Lucilia group".
Lucilia and Gamochaetopsis formed a basal grade with no unambiguous characters
to
distinguish then by Anderberg
l ered
member
99 to be a of the Cassiniinae, shares synapomorphies with Lucilia, including
1 )
(
1
biseriate trichomes (zwillingshaares; Hess, 1938) with enlarged adaxial basal In
cells.
Chionolaena, Gamochaetopsis, and Lucilia, the apical cells of achenial trichomes are
um
elongate and in excess of 50 in length, thick-walled, acute, and never rupturing
in
1
&
by Anderberg
Lucilia contains the species listed Freire (1991), plus a suite
of reduced, caespitose Andean species, included by them in Belloa or reduced synonymy.
to
L
These include araucana conoidea Wedd., kunthiana (DC.)
ucilia Phil., L. L. Zardini,
)
and nivea These
L. (Phil.) Cabrera. are all cushion-form species most closely related to
species found in eastern Argentina and Brazil. This trend in habit reduction also evident
is
recurvata Wedd. and Wedd. The
in L. L. flagelliformis superficial resemblance of
members some
reduced
these
to Luciliocline taxa Luciliocline
(e.g., longifolia, L.
radiata, and L. schultzii) is purely convergence, and has led some workers to propose
non-monophyletic
groupings.
Luciliocline, as interpreted here contain 13 species distributed in high-elevation
Andean
habitats throughout the Cordillera from Venezuela to Chile and Argentina. Greatest
&
diversity is found in Peru where eight species have been recorded (Dillon Sagastegui,
99
b). It is diagnosed as possessing heterogamous capitula, pappus bristles fused the
1 at
1
base, style branches of hermaphroditic florets rounded or obtuse, and achenes with
biseriate,
multicellular, capitate-glandular trichomes (Figs. The species composition of
1,2,
3).
&
expanded
Luciliocline over of Anderberg and
is that Freire (1991) several species
require transfers from Belloa and/or Lucilia (see below).
Belloa here considered monotypic and
is restricted to austral Chile and adjacent
&
The
Argentina. lectotype {Hooker Arnott 342, K) has been examined and,
just as
Hooker
and
Arnott
835) stated, their material contained only old capitula lacking
florets
( 1
The
or achenes. distinctness of species was recognized by DeCandolle when
this (1838)
&
he established Lucilia section Lucihodes based upon Lucilia chilensis Hook. Arn.
&
Remy
847), in establishing Belloa chilensis (Hook. Arn.) Remy, apparently did not
1
(
examine
type material, but stated that the primary difference between Belloa and
his
was new
Lucilia in the achenial trichomes, with his genus possessing achenes "papulosa,
non
villosa" (Fig. whereas, Lucilia possessed densely villous achenes. This difference
4),
achenial pubescence was by Cabrera
in also stressed
(1958).
Anderberg and Freire 99 based genus, Gamochaetopsis, upon Laennecia
their
( 1 )
1
&
alpina Poepp.
Endl., a species originally collected in austral Chile; however, has not
it
been established whether the type material was examined by the authors and whether
it
The
supports the diagnosis. genus said to be isolated and contain three autapomorphies
is
83): "lanate adaxial leaf surfaces, divided stereome [phyllaries], and cypselas with
(p.
1
&
The
short, clavate twin hairs." identity of Gamochaetopsis alpina (Poepp. Endl.)
&
Anderb. 199 was by
S.E.Freire problematic. type considered Cabrera 96
is Its
( 1)
( 1
1
to be congeneric with Lucilia and stated that could be confused vegetatively with
it
Gamochaeta
was
nivalis (Phil.) Cabrera, but easily distinguished from the taxon
latter
by
the sericeo-pubescent achenes elongate trichomes). Cabrera 97 considered
(i.e.,
)
( 1
1
Lucilia alpina distinct from both L. araucana and L. nivea. However, three taxa
all
were
described as possessing sericeo-pubescent achenes.
An
examination of
probable type material ofLaennecia alpina collected by Poeppig
at the type locality [Poeppig s.n., F 878548 ex B], shows to possess achenes with
it
elongate trichomes identical to those found in Lucilia. The achenial trichomes described
&
known
as "short, clavate twin hairs" are only from Belloa chilensis (Hook. Arn.)
&
Remy
(= Lucilia chilensis Hook. Arn.), and possible that material referable to
is
it
Belloa chilensis was taken for alpina. Whether Gamochaetopsis should continue
L.
may
to be recognized will await further study, but well prove congeneric with Lucilia.
it
Gamochaeta and and diagnosed by
Stuckertiella are sister taxa truncate style
branches and achenial pubescence of paired myxogenic Gamochaeta
sessile, cells.
New
warmer
contains approximately 80 species distributed primarily in the regions of the
World, but with several species adventive in the Old World. Cabrera 96 resurrected
)
( 1
1
Gamochaeta and described several new species from Argentina. Drury 970, 1971)
( 1
New
Gamochaeta
analyzed the gnaphaloid elements in Zealand and reduced
to sectional
Gamochaeta was by Holub Merxmuller
status. accepted 976) in Flora Europaea, but
( 1
et (1977) once again treated the genus as a section of Gnaphalium. Anderberg
al.
99 accepted Gamochaeta and had do with Gnaphalium
stated that to
) little (s.s.).
( 1 it
1
A
listing of South American species can be found in Anderberg (1991), Dillon and
&
99 and
Sagastegui Freire Iharlegui 997). Stuckertiella has achenes identical
a),
( 1 1 ( 1
to Gamochaeta, but diagnosed by several autapomorphies, including involute leafmargins,
is
4-merous florets, and clavate pappus apices. The 4-merous floret character has been
Gamochaeta
observed from
in as well (Diaz-Piedrahita, pers. com.). Preliminary data
molecular (Bayer between Gamochaeta and
studies et 2003) point to relationships
al.,
members
other of the "Lucilia group" as discussed here.
Facelis, Berroa, and Micropsis form a clade diagnosed as annual herbs with varia-
ble capitulescences and achenes with elongate (> 50 urn), myxogenic trichomes. In
all
1
when
these taxa, the achenial trichomes rupture through terminal pores in the apical cells
hydrated. Berroa and Micropsis are taxa and are distinguished from Facelis by
sister
achenial trichomes with twisted apical Micropsis further defined by
cells.
is
autapomorphies, including paleate receptacles where the outer phyllaries enclose the
pistillate florets, and achenial trichomes with unfused apical cells. Anderberg 1994)
(
suggested that the closest relative ofMicropsis was obscure, though was most frequently
it
associated with genera of the "Filago group." Overall morphology suggests that these
three genera are best placed in the "Lucilia group" and have more obscure relationships
with the "Filago group" (see below).
viviilia and uatrccasasiella are diagnosed as herbs with opposite, distichous
( 7/i
(
leaves, and a persistent pappus. Chevreulia contains six species primarily in austral
South America with three reaching the northern Andean Cordillera and diagnosed by
is
possessing fusiform achenes contracted into a filiform rostrum, barbellate pappus
bristles,
um
and myxogenic
biseriate, trichomes approximately 40 long, with bulbous
apical
cells.
Cuatrecasasiella diagnosed
is as dioecious herbs with glabrous achenes; two species
its
represent closely related Northern Southern Andean The
to disjuncts. relationships of
two
these genera are predicted to be with other Andean members of the "Lucilia group"
based only on morphology.
overall
The
three species of Jalcophila are recorded from the northern Andes,
i.e., /.
&
&
colombiana Diaz Velez
(Diaz-Piedrahita Velez-Nauer,
ecuadoriense
1999),
J.
&
&
&
M.O.Dillon
Sagast. and peruviana M.O.Dillon
J. Sagast. (Dillon Sagastegui,
&
These
1986). three species are very different from Jalcophila Anderb.
boliviensis
upon
S.E.Freire (based Lucilia hypoleuca Wedd. ex nom.
Schultz-Bip, taxon
nud.), a
&
described from southern Bolivia (Anderberg The
Freire, 990). a highly reduced
1 latter, is
though
species, not considered here belonging
as to Jalcophila, since lacks the
phyllaries,
it
pappus, achene and
shape, achenial trichomes members
that distinguish other of the
genus. Further, possesses a single, large capitulum on an elongate with over 40
it pedicle,
florets, a character quite aberrant in Jalcophila. This species most a Gamochaeta,
likely
is
A
a genus diagnosed by of
all the characters that distinguish this species. population of
MO)
this rare species has been discovered C. Solomon 4925, and should provide
(J. a
A
source of
material for molecular formal Gamochaeta
studies. transfer provided
to
is
below
allow monophyly
to for the reestablishment of
in Jalcophila.
"Helichrysum group"
The
"Helichrysum
group", with Pseudognaphalium and was
Achyrocline,
diagnosed by Anderberg 99 on
of
the basis phyllaries with divided stereomes, yellow
( )
1
1
and
florets, papillose achenial pubescence. Individual genera were defined by the
ratio
of
pistillate to hermaphroditic florets. Subsequently, Stenophalium was placed with
this
group. These genera exhibit a grade of floret ratios beginning with Stenophalium, where
number
the of pistillate florets is reduced to one or two per capitulum and the
fertile
hermaphroditic
florets are typically only Achyrocline has
five. - (-23) pistillate florets
1 1 1
and
-4(-6) fertile, hermaphroditic florets. Pseudognaphalium has (25-) 40- 30
1 pistillate
1
and 5-10
florets (-25) functionally staminate, hermaphroditic
florets. Finally, in
Helichrysum,
the central hermaphroditic number
florets far out the
pistillate florets.
Should of
the character the floral ratio of capitula be discarded, these weakly defined
genera could be combined, with Helichrysum name
as the oldest valid The
available.
from
molecular
initial result studies (Bayer et al., 2003) has identified a monophyletic
group
containing Anaphalis, Helichrysum, Pseudognaphalium and
several other species
Anderberg 99
that considered close Chionolaena
to in his C
( 1 )
1
))
.
group"
"Filago
The
"Filago group" an
essentially Nearctic clade with African, Eurasian, and
is
North American
elements. Psilocarphns a predominately North American genus
is
represented in South America by one Chilean endemic (Cronquist, Anderberg
1950).
99 and Morefield 992) related Psilocarphus Anderberg 99
to Stylocline.
( 1 ) ( 1
1 ( 1
1
suggested that Micropsis belong in this group, an alternative position for Micropsis in the
group"
"Lucilia discussed above.
is
Subtribe Cassiniinae Anderb. South America
in
Anderberg 99
established the subtribe Cassiniinae for a group of genera with
( 1 )
1
worldwide
and diagnosed
distribution as follows: often dioecious or subdioecious shrubs
phloem
or herbs, fibers absent, phyllaries with opaque laminae, hermaphroditic
florets
with truncate styles possessing trichomes on abaxial surfaces, achenes usually with two
vascular bundles, and pappus with
bristles clavate apical
cells.
Among
the 16 genera placed were Anaphalis, Antennaria,
there
Gnaphaliothamnus, and
Chionolaena. Anderberg monotypic
(1991) placed the
&
Colombian
genus, Pseudoligandra M.O. upon
Dillon based Oligandra
Sagast.,
&
chtysocoma Wedd.
(Dillon Sagastegui, 1990), under the synonymy of Chionolaena
comment
with a would
that recognition cause Chionolaena be Another
its to paraphyletic.
&
segregate, Parachionolaena M.O.Dillon was
Chionolaena
Sagast., established for
&
columbiana
Blake
S. F. (Dillon Sagastegui, 99 a). Freire 993) treated this genus
1 1 ( 1
as congeneric with Chionolaena, along with Gnaphaliothamnus
the several species of
Nesom
1990a,b) considered Gnaphaliothamnus
as distinct from, but with relationships
(
Nesom
Chionolaena.
to, Later, 994) provided a critique of the various classifications
( 1
of Gnaphaliothamnus
and stated that the Mexican and Central American taxa formed
a monophyletic group Nesom
potentially related to Chionolaena. Subsequently, (200
1
stated that he could not support Gnaphaliothamnus and sank remaining
as distinct
its
taxa into Chionolaena, transferring a suite of Mexican and Central American species.
If the segregate genera proposed by Dillon and Sagastegui 990, 99 a) are to be included
( 1 1 1
Chionolaena, must
in the diagnosis be modified morphological
to reflect the not
traits
by
shared the majority of
the species.
Analysis of both morphological and molecular data (Bayer 2003) suggest
et
al.,
that the Cassiniinae Anderb. polyphyletic and both Antennaria and Chionolaena
is
Gnaphaliothamnus) more
(including
are closely related to genera of the "Lucilia group"
defined
as here.
Subtribe Loricariinae Anderb. South America
in
Anderberg 99
established the subtribes Loricariinae and Relhaniinae
as
( 1 ) sister
1
made woody
taxa in a clade largely up of genera with leaves possessing involute margins
and pubescence on The woody
adaxial surfaces. Relhaniinae contained 19 African
genera with discolorous ray florets and rod-like achenes. The Loricariinae was diagnosed
as compact, often dioecious shrubs without phloem, crowded
fibers in the leaves with
adaxial pubescence, achenes with more than two vascular bundles, and dimorphic pappus
of
(i.e., apical cells bristles acute in the pistillate florets and clavate in the hermaphroditic
The
florets). subtribe included Pterygopappus (Tasmania), Psychrophyton (New
and two Andean
Zealand), endemics, Loricaria and Mniodes,
in the original cladistic
analysis. Later, Anderberg added Raouliopsis (Colombian endemic) and
Sinoleontopodium The
(C*hina) to his group, a posteriori. greatly condensed and
compacted
leafy stems of Mniodes and Raouliopsis are similar to highly reduced
Luciliocline species. Authentic material of Sinoleontopodium has not been examined,
common
but the generic description could refer to a shrubby species of Anaphalis, a genus
to the Himalayas. Preliminary results from molecular studies (Bayer 2003) suggest
et
al.,
may
that the Loricariinae be an
artificial c
New
coml
& &
Gamochaeta
bolivensis (Anderb. S.E.Freire) M.O.Dillon Sagast, comb.
nov.
&
Jalcophila bolivensis Anderb. S.E.Freire, Brittonia42: 139. 1990. TYPE: Bolivia,
Mandon
NY).
Larecaja, 79 (holotype, isotype,
1 S,
& &
Luciliocline longifolia (Cuatrec. Aristeg.) M.O.Dillon Sagast., comb. nov.
&
TYPE;
Lucilia longifolia Cuatrec. Aristeg., Venezuela 367. 1964. Vene-
Fl. 10:
km
zuela, Edo. Merida, camino a Pico Bolivar, 15 al sudeste de Merida, E.L.
VEN).
15725, (holotype,
Little
& &
Belloa
longifolia (Cuatrec. Aristeg.) Sagast. M.O.Dillon, Phytologia 58: 396.
1985.
&
Luciliocline pickeringii (A. Gray) M.O.Dillon Sagast., comb. nov.
TYPE:
Lucilia pickeringii A.Gray, Proc. Amer. Acad. Arts 138. 862. Peru, Dept.
5:
1
GH;
Junin, Prov. Yauli, Banos-Alpamarca, Capt. Wilkes (holotype, isotype:
s.n.
US)
&
Belloa pickeringii (A.Gray)
Sagast. M.O.Dillon, Phytologia 58: 396. 1985.
&
Luciliocline piptolepis Wedd.) M.O.Dillon Sagast, comb. nov.
(
SYNTYPES:
Meropepiptolepis Wedd., And.
Chlor. 162. 1856. Peru, Dept. Puno,
1:
HA.
4514 by
Maravillas, Weddell (lectotype, designated Cabrera,
1978).
P,
Belloa piptolepis (Wedd.) Cabr., Bol. Soc. Argent. Bot. 8 1958.
7:
1.
Gnaphalium
(Wedd.) Abh.
piptolepis Griseb., Konigl. Ges. Wiss. Gottingen.
24:
186. 1879.
& &
Luciliocline M.O.Dillon) M.O.Dillon comb.
plicatifolia (Sagast. Sagast., nov.
&
TYPE:
Belloa plicatifolia Sagast. M.O.Dillon, Phytologia 58: 394. 1985. Peru,
Dept. Cajamarca, Prov. Contumaza, Cascabamba, de Contumaza,
arriba A.
Sagdstegui Garcia Lopez M. &J. Mostacero 707/
A., E. A., L. 7 (holotype,
S.
HUT; HUT, MO).
isotypes, K,
F,
&
Lucilia plicatifolia (Sagast. M.O.Dillon) S.E.Freire, Darwiniana, 28: 41 987.
1 1
.
&
Luciliocline radians M.O.Dillon comb.
(Benth.) Sagast., nov.
Gnaphalium TYPE:
radians Benth., Hartweg.: 207, 35b. 845. Colombia, Prov.
PI.
t. 1
Popayan, prope Laguna de Guanacas, Hartweg 1146 K)
(holotype,
Lucilia radians (Benth.) Cuatrec, Trab. Mus. Cienc. Nat. Madrid, Ser. Bot. 33: 138.
1936.
&
Belloa radians (Benth.) Sagast. M.O.Dillon, Phytologia 58: 396. 1985.
&
Luciliocline (Wedd). M.O.Dillon comb.
schultzii Sagast., nov.
Merope TYPE:
Wedd., And.
schidtzii Chlor. 163. 1856. Peru, Dept. Puno, Prov.
1:
Carabaya, Ayapata, W. Lechler 1984 (holotype, F neg 37608).
P,
Belloa schultzii (Wedd.) Cabrera, Revista Invest. Agrlc. 404. 1957.
1 1
:
& &
Luciliocline M.O.Dillon) M.O.Dillon comb.
spathulifolia (Sagast. Sagast., nov.
&
TYPE:
Belloa spathulifolia Sagast. M.O.Dillon, Phytologia 58: 394. 1985. Peru,
La
Dept. Libertad, Prov. Santiago de Chuco, entre Chota Motil y Shorey, A.
M
HUT;
Sagdstegui Mostacero 11695
A., J. L., Diestra Q. (holotype, isotype,
F,MO,NY).
& &
Luciliocline turneri M.O.Dillon) M.O.Dillon comb.
(Sagast.
Sagast., nov.
&
Belloa turneri Sagast. M.O.Dillon, Phytologia 58: 392. 1985. TYPE: Peru, Dept.
Cajamarca, Prov. Contumaza, Pozo Kuan, Garcia
A. Sagdstegiii E.
A., A.,
S.
&
Lopez M. Mostacero 10087 HUT; HUT, MO,
J. L. (holotype, isotype,
F,
TEX).
Acknowledgements
thank Billie L. Turner for relating the fable quoted the beginning of paper,
I at this
my
has shaped do
for it in part thinking in this group; not overlook the obvious in classification.
thank Abundio Sagastegui Alva for companionship during providing much
I field studies,
and on Andean Mary
original material, discussions generic limits in genera. Reynolds,
SEM
Ron
Betty Strack and Wibel provided with The
assistance the original
studies.
American Taxonomy
Society of Plant thanked
is for partial financial support to attend the
Kew
1994 Compositae Nancy
Conference. Fred Barrie, Hensold, and Lucia Kawasaki
acknowledged
are for reading various drafts of this paper and making valuable comments.
Guy
Randall Bayer, Use Breitwieser, Nesom, and Josephine Ward have shared
all
unpublished data and their thoughts on various aspects Gnaphalieae classification and
phylogeny. thank the curators and collection managers of various herbaria where ma-
I
was CONC, CPUN, COL, GH, HUT, MO,
studied or borrowed, including K, NY,
terial
SGO, TEX, USM.
US, and Finally, Mario Zapata Cruz thanked for providing help with
is
Spanish
the
translation.
References
Anderberg, A. A. 1991. Taxonomy and phylogeny of Gnaphalieae
the tribe (Asteraceae).
Opera
104:1-195.
Bot.
1994. Tribe Gnaphalieae. In K. Bremer, Asteraceae, Cladistics and
.
Classification, 304-364. Timber Press, Portland, Oregon.
pp.
&
new
Freire. 990. Jalcophila of South American
S. boliviensis, a species
1
.
Asteraceae (Gnaphalieae). Brittonia42: 138-141.
A
-&
1991. and biogeographic of
cladistic analysis the Lucilia
.
group
(Asteraceae, Gnaphalieae). Linn. Soc. Bot. 106: 173-198.
J.
&
Bayer, M. M. Koekemoer, Ward.
R., Breitwieser, Dillon, 2003. Phylogeny
I. J.
cpDNA
of the Gnaphalieae based on three sequences (matK, trnL and
intron,
tmL/trnF miexgcmc Compositae
spacer). Newsletter 40.
&
Ward,
Bayer, R., Breitwieser, J. C. Puttock. xxxx. Gnaphalieae (Asteraceae)
I.
Pages xx-xx. In K. Kubitzki The Families and Genera of Vascular
(ed.), Plants.
Xx.
Vol. Springer Verlag, Berlin, in press.