Table Of Content!
AND
REEVALUATION OF AYLACOPHORA PALEAEPAPPUS
ASTEREAE)
(ASTERACEAE:
M.Bonifacino
Jose
Bosque 9G
°aseo del s/n, C.P.I
[email protected]
l.'iS.Moin^Kk'oJiRUGUA:
o/v.'os
Sancho
Gisela
,i;!u!nitniif\20560-0166,U.SA
ho.niwkmNMNH.SI.FDU
,an<
Permanent
address:
lenaasNaturalesyMuseo
Foe iltadde<
ARGENTINA
Bosque s/n,CP. 1900 La Plata,
ABSTRACT
Nesom and NanlophyJlum pdtd^onna nu.ihre,.
a (Cabreiw)
mean Patagonia that were originally desi
No
cacpappusm l°o3and
respectively.
1
/\\,'u'« >/>ll:H -),
1
. l 1
genus arguing
.-Vihaopiioo •ithin the rvi/rdnprivlhun.
i
etweeu the pah u o p ippu "1 h i\n lie u Pdidd, p mi ih p p'|"
, , ,'/ 1 .1 1 I | 11 i
11
ll
[ 1 I
I
I
Wl mi
II, II lll| 11 |> ll I. hill III u HI 1 1" /, / l| H , I Mil U I " '
1 [ I I I
unmlaand Hum
uhUmIh \unhplnlhu,nh \anh<
in mil pih u In
,P pi ol r ,i
,, 11
1
wide and enclose the lords, while Ndulophs species the paleae. present.;
re in tlu in if
I
RESUMEN
h
mi
mnnoiipii o \\ pt !,, 1, ' d, ,n poppio. 11 l i \ P> (1 > n [V , n
I 1 I ! ,
Nesom o op/iooi dolompMppio dent ro del genero \di\hph\
lu\
iih \v,',i, \
.
imiiiiuiiLulis cut papo palaeeo de MThophdv.i v PiiluipM
dis, re el
m, u toscomogi
Xli ; J,,, 1, (Pin, 110,11 niu on id, 1 n n, 10 111, , pi lulu 1
Aylacophora Paleaepappus, respectivamente.
y
Endemic
Argentinean agon Nardophyllu mdcscrlnola Nesom
to (Cabrera)
'at ia,
I
and Nesom
Naidophyllutn paiayouu
urn (Cabrera) are 2 of the species recog-
1.0
&
Nesom
nized inside Nardophylluin look. Arn. by
(1993).
I
Hum
Ncm/o/'/n de sen uola and paUigoiueum were
A. originally described
by Cabrera under the monotypic genera Aylacophora and
Paleaepappus.
Aylacophora (Cabrera b)"C>) was characterized by paleaceous receptacle, scaly
its
pappus,and compressed
cypsclae wit h 1( })ciliate ribs Paleaepappus (Cabrera
was
1969) defined by paleaceous receptacle and pappus 7-8
its of paleae. In
its
m N54
contrast, Nardophyllu sensu Cabrera has receptacles naked or with only
1 )
pappus and more
3-6C-13) paleae, a ol bristles, terete, or less pubescent cypselae.
According Nesom,
to the discontinuities between the paleaceous pappus
Aylacophora am]
Palcacp<ippus;uu\ pappus do
ol bristles Ntad.ojdiylluin
ilic oi
not support from Nardophyllum
their distinction because tendency
there a
is
m
pappus somewhat
the
for bristles to be flattened Nardophyllum.
However, according our observations and
to the interpretation of the data
gathered (see below), Ayhuophora and Palcacpitpus should be considered
as
independent genera from Nardophylluin
as follows:
Aylacophora Cabrera
Pol Sex Argent Hot 4:200 19=) n Aylacophora
Spkcies:
3
I
NSy
deserliuha Cabrera. Hoi. Soe. Argent. Rot, 4:208. \,udophxUum\lcscrtueld
u
Nesom. ARGENTINA.
abrera.) Phviologia 75:3o2. 1003. Fig. A-D.
Tu-'i-::
1
PkoviNOiA Np.uqufm
Plaza llunuul, Apr ahvni
12 1052. A.I. A /I252 U mi otyh-:
LSOTYPF:
LP!; US!).
cm
Shrub
50 high, densely branched; old branches aphyl Ions, bearing furrows;
new
branches with sparse nodes; leaves near; capit ula discoid, solitary ends
at
1 1
branches;
of involucresglobo.se: receptacles paleaceous, paleae wide, apically
pubescent, each palea enclosing cvpselae compressed,
a floret; 2(-3) nerved,
pubescent only on pappus
the ribs; of 10-11 oblong scales no longer than
1.2
Ecology- Ayhuophora
desertuola inhabits semidesert areas in Patagonia.
The
very few herbarium specimens
them
this species lead us regard
o\ to as
very narrowly distributed.,According SteibeKpers.comm.),
o
lug. A. deserticola
t
Cabrera grows on edaphic communities Monte
in the biogeographic province
&
(Cabrera Willink where
127 present on sane! dunes with very sparse
>), it is
shrub
with
cover, associated Larreadivaricata,Atriplexlampa,Prosopisflexuosa
A-D, Aylacophora desertia
Fig. 1
.
patagonicus. Capitulum. Receptacular
E. F.
bracteolatum.l. Capitulum.J. Receptacular palea. K. Cypsela with pappus.L. Pappus bristk. M-O.Nardophyllumbryoides.
M. Receptacular palea. N. Pappus bristle. 0. Cypsela with pappus. (A-D, based on Cabrera 11053, LP; E-H, based on Rio
Chico, 1900, Ameghino s.n., LP; l-L, based on Sena 77, LP; M-O, based on Ruiz Leal 27011, LP)
var.depressa,Scnnadph\llasuh>\\di\<niiitue(J}ui(uiraeacrinacea,Gutierrezia
patagouica Larrca Aylacophora deserticolahzs
solbrigii,Fabiana cuiu'ifolia.
i\ne\
been Argent Neuquen, Dptos. Conl uencia, Cutralco, Anelo,
collected na, Prov.
in
i I
Pehuenches, and Zapala. West Rio Covunco and south of Rio Neuquen in
ol
Dpto. Zapala, A. descrtuola the dominant species regarding surface cover.
is
Flowering the
in
fall.
Mnspn
Paleaepappus Cabrera, Bol.Soc. Argent. hot. 11:27 >.ls>o9. irn J'dcac/><i/>/nis
i
11*0
Nurdoph\Uumpata^onieum
/HJfdgcmcu.s Cabrera. Bol. See. Argent. Hot. 11:27
ARGENTINA.
K .ahivrajNesom.Plnaologia 75:302. 1001 iMg.l 10 n Chubut.
li 1.
1 :
Shrub densely branched, brandies sharp ended' leaves oblong to spatu-
lateral
ends branches; involucres
eonaceous; eapiiula discoid, solitary at the of
l.iie,
campanulate; receptacles paleaceous, paleae wide, apically pubescent, each
enclosing a floret; cypsclae terete, densely pubescent; pappi of 9-10 elliptic
nun
paleae long, in 2 series.
7
Ecology —Paleaepappus piitagonicus inhabits semidesert areas in
and
Knowledge ecology Paleaepappus scarce speculative
Patagonia. of the
ol is
and no
because the only record o\ this species he tvpe itsell there is ecologi-
is
t
.
cal information on the label. The conservation status of his species could aptly
I
be recorded as endangered.
Aylaeophoraand Pa Icac pappus are stronglv segregated from Nardophyllum
num-
elements pappus such number, shape, and
by characteristics of the of as
The pappus AyUuojmora and Paleaepappus con
paleaceous
ber of
series. o\
m
pappus
trast with the bristles Nardophyllum. Intermediate states of shape,
m
narrow ha\c nor been found Nardophyllum. but occasionallyflat
paleae.
like
have been found Nardophyllum.
bristles in
Nardophylhun circumscription
&
Nardophyllum Am. (1830) was described by Cabrera (1954) as shrubs
look.
I
end
densely branched, leaves alternate, small: heads solitary at the of the
campanulate; convex,
branches; discoid capitula, involucre receptacle small,
4-5 pappus com-
naked or with few paleae; cypselas turbinate, ribbed, hairy;
posed The circumscri prion ai Nardophyllum adopted here
of several brist is
les.
where he included
the presented by Cabrera (1954) in his revision o\ the genus,
Nardophyllum armatum (Wedd.) Reiche, N. hraeieolatum llauman,
7 species:
chilwtruhioidcs
N.genistioules (Phil.) Gray, N. hryoides (Lam.) Cabrera, N.
Hook.&
(Remy)A.Gray, /minima (MeyenjCabrera.and N.obtusi folium Arn.
N.
\anloph\
Palraepappus
,\ad lliun
pappus
Contrasting with the pappus Paleaepappus (big. the ol
o\ II),
1
mm
com 5-10 L),sometimes
Nardophvllum posed olea. 30 bristles long
(.Fig.
is 1
apex arranged 2-3(-5)
flattened, especially ar the (lag. N), in series.
1
Involucres and shape
o\ Palcticpiippus cvpselac arc similar those
to of
Nardophyllum. number
Differences and shape
in of receptacular paleae are also
found in these two genera. Palcacpappus has paleaceous with wide
receptacles
and
apically pubescent paleae enclosing each The
floret (Fig. receptacular
F).
1
paleae Nardophyllum do
of are narrow, not enclose the florets (Fig. J and M),
1
mid vary from armalum
absent cV. to Vo> (rarely more numerous, 9-13
in N.
)
bracteolatum).
Aylacophora and Nardophyllum
The pappus C
Aylacophora and
of (Fig. D) contrast highly with the pappus of
1 1
Nardophyllum
add
(see above). In lon.ot her characters distinguish Ay /acophora
it
from Nardophyllum:
the involucre Aylacophora globose
ol Fig. A); cypselae
is its
1
I
are compressed, 2(-3). and pubescent only on the ribs The involucre
(lag. C). of
I
Nardophyllum campanulate toobcomcal band
is (Fig. the cypselas are
terete,
1
(4-)5-7(-8) nerved and uniformly pubescent K and
(Fig. O).
1
1
Aylacophora has paleaceous with wide and
a receptacle apically pubes-
cent paleae enclosing each The number and shape
floret. of receptacular paleae
Aylacophora
of (Fig. B) contribute to set this (axon apart from Nardophyllum.
1
Aylacophora and Palcacpapus were placed Chihotrichum group
the
in
&
(Zhang Bremer Bremer
1993; 1994) that includes shrubs with mostly densely
ami
coriaceous, often abaxial tomentose and most
set. leaves, for of the genera,
\
paleate receptacles (Bremer Within group
1994). this there are intermediate
morphotypes
pappus elements range from
for that
terete bristles in
Chilliophyllum, through narrow paleae Pcpidophyllum (Cabrera
in 1971), to
paleae Aylacophora and number
in \i lee epop/arv In addition, the of receptacular
/
paleae vary within the Chiliad ichum gmup. defence ehacu
In to this tor the
i
when
variation observed wiihin Nardophvlluiu comparing
isalso present other
genera Chihotrichum
of group, such Lcpidophyllum without
as paleae,
Chiliotrichum with lew and and
paleae. '.hilioplwllum Chiliotrichiopsis with
(
fully paleate receptacles labrera L971, 978) Because the gradation in pappus
((
1
and
receptacular paleae inherent he Ch richum group
is to Hint itself, this varia-
t
tion cannot be an argument againsl the recognition of both Aylacophora and
Palcacpappus
as distinct from Nardophyllum.
Genera v
Chili rich tun roup ensu bremei F-W4) can
o\' ol CI identified as
i
follows:
appusofca.30ormor.>hriMlr'
Nardophylk
in 2-3C-5).
ippusoflOorfewrrpalu.iwu
,i,-,n n,;d to the Avlacophc
iiif ii ill, h, in ribs.
1 1 i . 1
?ceptacles epaleate.
Diplostephk
(Nesom
Nesom's reinstatement of Nardophyllum scoparium Philippi 1993) is
not accepted here. Presence oi pistillate ligulate florets in the periphery (5-6)
The
ionmgthisiavon \ doph
doubts km posh inside m. taxo-
casts serious at villi
n
;il
nomic placement Chilean species and possible status as an
of this its
Nesom
undescribed genus allied to some genus inside Chiliot richum group as
suggests comm.) being reviewed by one oi us ()MB) and will be pre-
(pers.
is
ruhum
sented as a more comprehensive study of the (dwliol group (in prep.).
W),
According Bremer Tvlm ophoi a and Pakacpapput.;are very similar
to
.1
(
was Amhcpliv! and
and possibly sister groups. Ayldcoplh'/u related to urn Crnliofri-
I
U
\m< and Avldou
>m w
ohiopsis by Cabrera : Palcitop>opj elated lo >pliyllum
C l a-, I.
\ l > > i
morphology
p ho (Cabrera 190Q) Ouropinion, based on observations of evpsclar
ra
number of series of the pappus elements, in vol uc re shape, leaves shape and plant
between Palcacpappus and
relationship N'mt/onliyllimi.
habit, favors a closer
Nesom tomentosecaulmesulcae
(1993)suggest the narrow, internally
that
homologous somespecicsol Nardo-
Aylacophora could be Inwo
oi ith hu.incl in
\\
t
am and so denoting a closer rclationshipol Avlmuplamc! to Nardophyllum.
/dn'l
I
both Aylacophora
To complicate the matters further, the paleate receptacle
i^i
and Palcapapus, would favor a closer relationship o( these two genera with
hum which Nesom
and nr
Chi Hot richiopsisX'.hi iophyllum Cli iliot instead, are, as
I
them
.VmlnplivlImM and have
(1993) points out the closest relatives to all oi a
m
Ch paleated,Chi/iotnehum
paleate receptacle X'hi Hot rich topsis, '/it>p/n hi ul
\
i I I I
15-21 paleae). Evidently, as Bremer U994) states, the elucidation of the phylo-
among genera demands more comprehensive study
genetic relationships these a
oi the Chiliotrichum group that beyond the scope ol this paper.
is
Adem
and Irom Nmdn/'liyllum on the
Segregation ol Ay/a. nia'i.M a />np/)m
pappus shape supported by other characters such as the shape
basis of ol
is
both cypsela and capitulum Aylacophora, and the shape and quantity ol
in
both ophoro and Palcacpappus.
reeeptacular paleae A\
in lu<
Although Aylacophora and I'alcacpappas are closely rehired to the rest of
Nardophyllum species, the discontinuities basically observed in the shape ol
the pappus elements, with no defined intermediate states, justify the consider-
wo
ation oi both species as distinct genera.
t
Nesom These based
Otirconclusion issupported b\ al.Un press). mil hors.
el
and mainly on pappus morphology have included
on features of the involucre
S
1
new Wi On
Peruvian A.ki.x uu cu
a spec a so! inside /ml n«;hiopsis Cabrera
&
Nesom
(OnliotrichiopsLS Nesorn, Robinson Granda).
pcrin'iciiirt et (in
al.
found pappus moi
press) that the phologj good harai ter to separate gen-
i <
era inside Cihiliotrichum group and concluded that Cabrera's narrow generic
concept Nardophyllum
of w d< iption of the diversity inside
b<
tt<
\ <
i
Chiliotrichum group then fore ha\ ing incf pi ndcnth an ived at the same con-
we
clusion have, regarding the consideration Aylacophora and Paleaepappus
of
from
Nardophyllum.
as distinct
APPENDIX
1
Additional specimens examined of Aylacophora deserticola Cabrera. ARGENTINA. Neuquen, Plaza
19*
Dec L
Huincul, 11 hi II, H ][ h H Troiani etal 12504
il ,
1
i
5h
(SRFA);PasodekoslndiosaCutrako
ft RFA), Paso de Bardas,
, n las
[ 1 < - i )
-e,
II
Feb
2 ]999, H.Troiani\ " I, Indtos, 5 Feb 1999, P.Steibely H. Troiani
HP"
I
oIihImh
H
/4069(SRFA) Paso del o
liulio Troiani 14073 (SRFA)
it ,1 '
; \
Nardophyllum. Nardophyllum armatum ARGENTINA. San
(Wedd.) Reiche. Juan:
Iglesia,
Cam
camino h h Pampa
a el Fierro d _ >i P) de Pauacan,
I i it \ 1 I
entrelasAguaditasyChepical, D
12 La Rioja: Gral.Sarmiento.Rio del
- LP).
, ,
1-jn d
i { ii
I i I t , i
n Wde
San Antonio delosCobres, 2^ km
Salta: P).Jujuy: Quiaca,
l.n 'I al la
I 1 1
&
Feb Meyer
960,
1
H
Esquinas Blancas, 22 Ene 1966,Gv/vm Pampa, Feb 1937, Cas tellanos 20229
If
< - . i ,
(LP).
Nardophyllum bracteolatum Hauman. ARGENTINA. Mendoza:
San Carlos, Pedernal, 25
El
Mar 1916, Sanzin 1810 (LP);Tunuyan, Paso del Portillo, Cuesta de los Afligidos, 29 Jan 934, Ruiz Leal
1
2052 (LP); San Carlos Pnu >nd L n 41 ~ an Carlos, Arroyo de
it ' (I r la
I > I I I i
Qda/'Casa de Jan 77
Piedra," 7 952, Serra
(LP).
1 1
Nardophyllum bryoides (Lam) Cabrera CHILE. Magallanes: Naciomll
III
|u
ii ti
I i
Lago Paine, 7 Jan At uLNTINA. Santa Cruz: alrededores de
El
1
I
Chalten,10Feb2000,P<»/i a m. rlinero.Cabo Virgenes,
L
i il
i i i i
Mi l'4ho//n r
Riod h
Vizcachas,Cerro de
PI illt-go H E le las
(I r
1 , i i
Ml
-crmW.25864
(LP);LagoArgentino,ParqueNacionalh<
J hi (LP).Tierra
i t I i 1 I
^
delFuego. Est.Cullen,52°44'5,08° Bahia Lee.52°
in -, P) 52'
> i , - ,> il
70° 6' W, 6 Nov 1971, Moore 2339
(LP).
1
Nardophyllum ARGENTINA.San
chiliotrirl- Juan:Calingasta,Qda.Los
I-.-
ii
m
avestruces (oeste de Cerro Castanet Rio Mauriquc
F .4 (LP), a
I I i • >
Pmt
1" Ine
-II, |, I, ' tl tilt. . Iltl 1 • • hi. , I hi, lIF ,1 II I, I, h 1
I
1947, Soriano 2786 (LP); Ravs. m, in I, 40, astruvie,o & Lopez 23 13
' il f ft, 1 (SI).
1 I I 1 i
hi
huh
CHILE. Santiago: Cotdlhtad, UCr/ermcmn
621
Ft u> in (LP).
it it it,
I i I
Nardophyllum lanatum ARGENTINA. Mendoza: km
(Meyen) Cabrera Malargue, 40,20
ruta
- h NPi.qi.on:
alf,u u
L i-
I,
> 1 1 ' I I , I
Calmuco,15 Feb 1942 n lm Pio Barranca Buta Ranquil,8 Feb
Ft y
j
I i
950, Boelcke 4235 (LP). CHILE. Colchagua fen 9 Feb 96i <ner833 (LP);Vegas del
1 1
III
Flaco, de Qucl>t hl<h II s Valparaiso:
al E la It L >•" (LP).
1 I I
i I t
n n
Cerro Roble, Cordtlk h Ovalle: Ge/sses.n (LP 60269).
I,
I l t I I I
Nardophyllum obtusifolium Hook & Am. CHILE. Magallanes: an Gn 76 Oct
gorio, 968,
1
ARGENTINA.
Cekalovic 33903 (kP). Santa Cruz: Pto.Sai luliai M n (LP) Corpen Aike 20
i i I i
.
l/l^t^fi >M
hnHIVHidRiK'm.Kut.i o . I" < L),L,.in|cnL b astillo.11 Feb 1936,5coff
'
\ ! >< ',
^
V
Nicora3638 (LP); Pueito nio in A/bo// gu. ' Apt 946, Scotnik 306 (LP);
>. il
[ < I 1 I ) I I 1
Leleque, 13 Jan 1947,5ooono ?3A Pcpita, Alto Rio Viuior, Fob 947, Soriano 2585 (LP).
P); hst. i
I (I 1 1
lm
Neuquen:Charahui!h A,, inn Q Jan 1966;camino
a
, I ,, , I I I 1 I I i I i 1 1
!•-
aNinhuau, 16Jan Santa Cruz: olt Dec
.-. . P alios Pellegrini, Est. La Flora,
. ' L
" " inqen 2
(LP).
I ' >
ACKNOWLEDGMENTS
It is a pleasure to acknowledge Lihana Katinas and Jorge V. Crisci lor helplul
comments Nesom
critical on the final manuscript; tiny lor his helpful critical
new
comments making
on this paper, available lo usihe inanuscript "A species
from (m\ Nesom, Harold Rob-
Chiliotrichiopsisi teraceai tereae Peru.
ol '
\ \
)
m
m
i.mpn
in on no! u ro unl.i Patu ai Bi ilioni In en< mi n nit
• t ,i ' i i
1 1 I
towards publication; John Strother for his corrections on the final manuscript;
lng. Agr. Pablo E. Steil x for he n formation regarding the ecology of Aylacophora
I t i
and geographic on me the< on .ejoNai uMialif InvestigacionesCientificasy
its
Tecnicas de Argentina (CONIC.LT) and . '.omision Sectorial de lnvesfigacton
(
(dentil ica(CSIC), Uruguay, the financial support on tins investigation, and
ioi
LPAG, MVF, SRFA and US loans herbarium specimens.
LP, SI, for of
RHHHRHNC.liS
Br[mer,K. 1994.Asteraceae bit Portland, Oregon.
Cldcli th hi
it ti
I 1 il i I i
Un nuevo genero de astemn bepubl
Carrfra, A.L. 1953. dt a a Argentina. Rol 5t> Ai
,
4:266-271.
gent. Bot.
Cabrera, A.L. 1954. Las especies del genero Nardophyllum. Notas Mus. La Plata, Bot. 83,
17:55-66.
CABRbRA, A.L. 1969. Compuestas nuevas de Patagonia. Bol.Soi Anient. Bot. 1:273-275.
1
Cabrfra, A.L. 1971. Compositae. En M. N. Correa (cd.), LI Pataqonica, Colecc. Ci. Inst. Nac.
Tecnol. Agropecu. 45
8(
7):
1 1
1978 on In' Cabreia
Cabrfra,A.L. uju\ Ci.lnst.Nac.Tecnol.
it). LI Pi le
( I.
if
Agropecu. -726.
13(10):!
no m
p
and Wn oi ili'io a bene de
Cabrera, A.L. A. E Biologi'a,
1 ' I i i ill i It ' t '
Monografia Washington, D.C
13.
P
toman m: M mine, and
GrauJ. 1977.Asteie.it l In IL *>, Lirl B.L.Turner, eds.
\ > \
,
I | I
The biology and bemisliy of the Compositae At aclemic Press. London. Pp. 539-576.
).
t
NtsoM,G.L 1993 laM in mi inn v eae Astereae) with
i i ' i i
observations on the definition of Nardophyllum. bbytologia 75:358-365.
Astereae) from Peru. Bnttonia press).
(in
