A REVISION OF CORDIA SECTION GERASCANTHUS (BORAGINALES: CORDIACEAE)

2013·17.2 MB·English

by  James S Miller

GERASCANTHUS
SECTION
OF CORDIA
A REVISION
CORDIACEAE)
(BORAGINALES;
James
Miller
S.
New Garden
The York Botanical
New
York 10458, U.S.A.
Bronx,
ABSTRACT
RESUMEN
INTRODUCTION
with estimates lot the
2001)
Millet
Johnston 1930;
broad sense (e.g.
been defined in a
Cordia has historically
two decade^
However, the
in last
Shaw 350 (Miller 2001).
1973) to
from 230 (Airy
number ranging
of species
withm
relationships
genera,
with other related
relationships
understanding of
number advances in our its
a of
we unde«a.rf.
what
have added considerably to
species
constituent
and studies of the
the genus, floristic
com-
been considered to
which have histoncaUy
shown Boraginaceae,
^ have that
studies
phylogenetic
Several
HeltorioptoA-
Ehrettotdeae,
Cordioideae,
subfamUies:
ddferent
and morphologically
distinct
prise four
&
Hil^r 2001; Gottschltng et al.
(Gottschling
more complicated
were
L, Boraginolae (lohnston 1931),
other famthes, or
Several
& Weigend submitted).
et
Hufford 2012; al.
Nazaire Sr
Moore 2006;
2005- Janse
and
Codon Hoplesligmu,
genera
and African
Lennoaceae, the
Hydrophyllaceae,
including
enigmatic genera,
WeLdtiawereallembeddedwithintheBoraginaceae,butalloftheoriginalfoursublamil«wereprovento
(Gottschltng et
al.
been placed in the
had
few genera historically
be monophyletic, altbough a
more
with Ehreha, are closely
both long considered allied
and
Coldcnin.
now
Sacccllitm
clear that
2005).
It is
(Gmtschhng ab
and Vorronm et
Cordiu
within and the later sister to
included
former it
Cordio, the
related to
mcludmg
2W5),
Boraginales (Gottschltng et al.
defined
broadly
recognhion of a
2005). This has lead to the
Hehoriopuceae,
Cordiaceae Ehretiaceae,
elevated to familial status,
subfamilies
each of the four tmditional
Wellstedttace-
Hoplestigmataceae,
Lennoaceae,
Hydrophyllaceae,
of the
and inclusion
and Boraginaceae, also
&Hilger
(Weigend 2010). w
Codonaceae
and newly described M
ae.
prccumto
.
. Old World
the
elucidated that
(2005)
Gottschling et
al.
studies of
the
For the Cordiaceae,
CorAu
and then
woody members of Cordiaceae
other
was of the
procumbens sister to all
annual herb Coldeniu
had been recognized be-
small genera that
Cordia and the three
of
was of the rest
section Varronia sister to all
(Auxe^ L and
Putag^ulu
Miers,
calyces
accrescent
which have large ,
unusual all
cause of unique, fruits,
on
Based
section Cordia.
and with the species of Coirfio
embedded in Coniia allied
were
SuccelliuM Bonpl.), all
membem
and Cordia to the of sec-
restricted
Varronia
segregated
(2007)
and Gottschling
this evidence. Miller
show Old
the
that
though molecular data clearly
the
Myxa, and Superbiflorae,
Gerascanthus,
lions Cordia,
morphologically
World and New World species of section Myxa are distinct clades, even though they appear
be approximately
Now that checklists are being compiled for the Online World Flora, there appear to
similar.
dozens
of
230 known species of Cordia and an additional 124 species of Varronia, though there are still clearly
major herbarium col-
from undetermined folders in
both genera awaiting description
undescribed species of
lections.
Wheeler Johnston
Gerascanthus has always been a difficult group (see quotes in 1942;
Cordia section
by
primarily
and recognizable as a group of species, characterized
morphologically distinct easily
1950).
It is
and
surrounded
and most
distinctively
unusual fruits, which are ellipsoidal, single-seeded, fibrous-walled,
its
brown
turning
and marsescent with the corolla drying,
completely enclosed in the persistent calyx corolla,
wind mature But while the section
and apparently assisting like a small parachute with dispersal of the fruit.
prompting Chodat and Vischer
may be easily recognized, the species have long been difficult to tell apart,
and
(1920) to write “We are, in fact, deahng with species which are but feebly defined morphologically the
The
number.”
taxonomy of which will require revision from time to time as observations in the field increase in
species of section Gerascanthus are not distributed evenly, and though they occur throughout the Neotropics,
most of the South American species were known from few collections or were discovered relatively recently,
and the greatest concentration of species in western Mexico, a group provided with a relatively solid revision
is
(Johnston 1950), though Johnston knew eight of his twelve species from fewer than 3 collections, four from the
type
only.
Most species of Cordia sect. Gerascanthus are small to medium-sized trees. Only a few species are truly large.
m
Cordia megalontha the largest, occasionally reaching 60 and C. alliodora and C. trichotoma often reach
is tall,
30 m. Cordia insignis a shrub of Brazilian cerrado that apparently grows from an underground xylopodium
is
m
and flowering individuals are generally only 1-5 tall and at least in some situations, above ground stems
its
appear be
to annual.
Descriptions of bark are generally not included with most specimen
label data, though bark often a
is
characteristic feature. Most species have smooth pale gray bark. Cordia morelosana has dark rough and
gray,
deeply fissured bark, which is one character that distinguishes from
C. sonorae, a
it species
i
and
cally similar difficult to distinguish from herbarium specimens
but has smooth,
pale gray
bark.
Species of Cordia are not infrequently associated with ants but
the association generally with
is casual,
ants living under patches of bark or in hollowed twigs. There are, however, two true myrmecophilous
species
which
of Cordia, are unrelated and
independently
derived
(Gottschling,
2005). Cordia nodosa Myxa,
of sect
hasdomatia
that are swollen, hollow
petioles. Cordia
alliodora, of
sect. Gerascanthus, has domatia
that are hol-
ell^td
low, sweUings
at the ends of branches and
the bases
of
inflorescences
that are generally inhabited by
bttmg
small, ants of more than dozen
a genera (Wheeler
1942). These
swellings
are filled with a loose pithy
material that presumably
is easily removed by the The
ants. domatia
are
characteristic of the plants and not
the
.s„UdWesta.ion^theamsasdo™a.Ufon„nonnally
on grownm,
plants
he gree^
absence of ams.Anj5
the have been
associated wi,hall of the treesihave
observed
in Mexico, but they have
frequently been
absent from
trees in Central
America.
Tnchomes
of various types
are
often associated with E
in the genus
n
vanous
species of
section
Gerascanthus;
the
fifth type, echii
Gerascanthus
Myxa. The four types of hairs found in section
known only in Cordia cymosa of section
are
hairs,
include:
found in the majority of the species and are variable in
hairs that are
hairs.—These are unicellular
Simple
1)
from a swollen,
unicellular hairs that arise
are thick-walled,
SimplThl^lT/rom
2)
and the hairs are mostly ap-
usually postulate
The shaft of these hairs is
pedestal or cystolith.
basal
and
on length
depending the
induments,
and scabrous
pressed giving rise to strigose, strigillose,
In some cases, the shaft
appears multicellular.
generally
The pedestal or cystolith
thicknes of the hairs.
rough
which result in a sur-
the persistent cystoliths,
and with time, leaving
weathers lost
of the hairs is
common
Gerascanthus.
common in the genus, but less in section
from pedestals are
Hairs
face.
and
of Cordia alliodora
star-shaped hairs that are characteristic
multiply-branched,
hairs.-These are
Stellate
3)
numerous branches
with
have hairs
Both these species stellate
of
Gerascanthus.
trichotoma in section
C.
which
genus Varronia, are
These from the stellate hairs in the related
and the hairs are sessile. differ
and obviously have arisen independently.
stalked
They
andamedianattachment.
with two arms
T-shapedhairs
are unicellular,
Malpighiaceoushairs.-These
4)
with
each time a slight-
times in the family,
independently at least four
but have arisen
are rare in Cordia
among section Ger-
the species of
known only from Cordia glabrata,
They
appearance. are
different
ly
undersurface of the leaves.
the
whitish cast to
where they give rise to a
ascanthus,
and are presumed to be de-
tL Gerascanthus are simple, alternate, entire,
leaves of species of Cordia section
some
Many flower without their leaves, so it is in
species
confirmed in species.
not all
ciduous, although this is
Leaves^ryconsiderablemsize
specimens with leaves.
with
paiinoweringspedmens sterile
cases difficuU to
distingnishingspecies-Allspecieshavepenolate
and form and shape and texture are useful characters for
leaf
only
thoughafewspeciesare flattened
on the adaxial surface,
canaliculate
generally
leaves,and the petioles are
and nrominent groove.
lack a
condtttonBap-
that
by helicoid or scorpioid inflorescences,
thoughtofascharacterized
While Boraginales are
and
only in the ultimate branches of inflorescences, tt ts
Conhu and then generally
parent only occasionally in
cymes
expanded
from
vary
in the section
Inflorescences
Gerascanthus.
section
seldom apparent in species of
but they vary
cymose-paniculate,
compressed and nearly umbellate. All of these are basically
and
panicles to
between three different
continuum of variatton
a
branches giving rise to
the
in the degree of elongation of
suchasm
cymes,
axisthatisapanicleof small
withadistmct central
expanded
imLulateinforescencesare
macrantha.
Cordia
l j
r •
and broad.er t. n form, such as ts found
expanded, but lack a central axis
Cymose mnotescences are also
2)
CordiacolimensisandC.elaeagnoides.
with none of the branches elongating and, gi• ving rise .to a
ccoommpprreessssed
very
are
Umbellate inflorescences
3)
umbeU ratheramere con-
but
glohuh/em these are not true
Cordia
in
of flowers,
umbellatecluster
nearly
umbel-
umheltea they appear to be truly
but in C.
branched inflorescence,
a
branches of
of the
traction
from a single point.
with flowers arising
late all
i^orescences vary
has that
Cordia sonoroe
each species,
consistent lor
generally
While type
inflorescence is
S,^^olCordiasect.Getnscanthusaretypicallydistyh,usandmdividualpUntscanread%bese^ratedmm
between
varyingsignificanUy the
stamenand heights
style
with respective
morphs,
and
long-style short-style
When
known
two. species are from adequate numbers of collections, floral measurements are provided sepa-
rately for the two morphs, though this unfortunately is not the case for the majority of species. In two species,
has broken down.
distyly In Cordia morph
alliodora, the long-style has been and have
lost all individuals ex-
serted stamens and
a short style. In Cordia trichotoma an unusual variation occurs, with both long-style and
short-style individuals being
present in populations but with morph
a third with equal stamen and
style
&
heights (Gibbs Taroda
1983).
The
species of Cordia
sect. Gerascanthus have a tubular calyx
that is usually distinctly 10-ribbed. In a few
such
species, as Cordia morelosana, the indument on
the calyx dense and long enough
is to obscure the ribs. In
other such
species, as Cordia
thaisiana, the calyx merely and
is striate lacks the clearly defined ribs characteris-
of most The
tic species. calyx
persists in and
all species encloses the mature
fruit
Section Gerascanthus
characterized by
is
corollas that are marcescent and
persist, enclosing the mature
As
runs. f^its mature, the dry,tum
corollas brown, and
act as small
parachutes wind
that aid in dispersal of
ta^Corolteofspedesof
.he
the section are her tubuUrwith
ei, spreading
lobes
^.al
5
corolla
y lobes, tftough c^ollas with 4 6
or lobes are not nncomnton
some
in spedes. Whether
the
l,^s
corolla are oblong, wtth
parallel sides, or ovate debate
to is a feature useful in distinguishing The
species
^
»>• ^nd
only
in Cordia
a^ ntegalantha,
C. latiloba,
and C
C. umbellifera are the corolla lobes drawn an
to acute, sharp apex
characurfordistinguislungspeciesandvar^bleonlyinCurdiagerascanthLandCm^^^^^
The
gynoecturn Cordw
of
consists of a
superior, U
bicaipellate ovary
that no. divided
or lobed
Earlv in
development
tt IS btlocular, but becomes
later
it falsely
4.|ocular as a septum forms *"*"*'"*
dividi
t.
two
separate chambers (Uunence 1987^1 fTT
1937-
Khaleel 1975
,
T
thotropus
ovule but onlyonedevelopsmmaml
fXonTa
aid
?
A"
species of the section have an annuL needle
H-
compared
with the mto ‘“^8'
nectaries of 0^7
species of cto"
le n
grocilipesischaracterizedbyhavingadiscthatisciliatealoneth
““Sin
The
and “PP*"^ style in Cor(ka
twice terminal
bifid with 4 is
separate
stigma
lobes ^
that vary from
fil f
«‘l«oid
but
in section are always
Cerascantims. clavate
Fruits
The
fruits in
species of
section
Gerascanthus
are thp a-
thin,
fibrous
wall, lacking
a mesocarp, "
and
are capped
bv
slightly during
development
and
remains 7'™i
borne
within
the ners
™|.dispe.sedwi.h.hemarcesce„.corona,aidinginwinr:;^^^^
PollenoftheBoiaginalesisknown.obediver!e(Avetissianl95fil
m
7
thrm
i,
types
of
polfen
grains
in the genus Cmdia <*’73) initm
in a broad
and
Gottschling
Q007),
Varronia and
^mn Cordia
section
Cordta
7
C^o^ontlms 7““'
’”'‘™
shares ^0.^12'
pollen gmins
that
are
^
Tarodaand
Gibbs(l986)groupedthese
section7nrr7
three
taxonomic
treatment
Browne)
G. Don,
•
Gen.
Hist.
380.
1837; A. DC,
Prodr.
9:471
If
Geraxanthus &
Schlect
Cham.,
Linna
Revision of Cordia section Gerascanthus
Miller,
Cerdan
Cordia
section
a{Ruiz&Pavon)Oken.
Small to large trees. Leaves deciduous, alternate, petiolate. Infloresce
much
sometimes with the axes reduced and nearly umbellate. Flowers
dia alliodora: calyx tubular, usually distinctly 10-ribbed or occasionally merely striate; corolla whii
cent, tubular with spreading lobes or funnelform, usually 5-merous, the lobes usually spreading; si
same number as the corolla lobes, the filaments pubesc
an evident, usually annular, disc. Fruits borne within
soid, the walls thin and fibrous, capped by the discoid base of the style.
Cordia section Gerascanthus consists of 23 species, all restricted to the Neotropics. Thirteen species occur
in Mexico, and though the species are spread throughout Central America, the West Indies, and South Ameri-
there no other substantial center of diversity.
ca, is
Journal of the Botanical Research
Institute of Texas
7(1)
^^3-4):396.
1988. Type; PERU. HuAnuco: Pn^nr^ p.h, .
. ,
&
.r«« ccraam
Koem.
Schult., Syst. Veg.
4:467. 1819.
'"*'
non. August, Hassler 7155 in campis siccis, prope Concep-
(holotype:
G).
Cordia
alliodora
var. glabra A. DC..
Prodr.
9:472.
11 S.TYPEiPERUrPovOns.)
m
Tree
to 20(-25)
tall, slender
cr
b
attenuate ^^(-12)-ftbbed,
f white, mm teeth,
these
(8.5-)9.5-12(-141 0.5-K-l
i ,
*”
mm ^ mm marcescent,
(3.5-)4.5-6(-8 ^bese
5-7(_8
5) 5)
1 1
Gerascmtims
rang-
»tn.lAmencaa«l,heWes.lndies.,„B,BzU,
Miller, Revision of Cordia section Gerascanthus 61
by
Bolivia, and Argentina. replaced in Eastern Brazil its closest relative Cordia trichotoma. occurs across
It is It
m
a wide vareity of habitats, from dry to wet forests and ranges from sea level to 1,500 or rarely as high as 1,900
Orange Walk:
30 Whitfoord2866 (MO). 17557 (MO).
lev. ft, 1
MO, BOLIVIA. V
5407 (BRH, SEL). Beni: Prov.
samo
1224 (MO).
Y
Cochabamba, Bang 1178 (MO, NY, US). La Paz. S.
i:
W
km N
Krufeoiifs.n.(MO.NY).Pai bank of Rio Madeira, 2 of Abuna, Prance et al. 6114 (MO, NY, I
lo.
km NE
Hierba Buena, 1,300 Steinbach 270 (MO). Tarija: 0.2-0.4 of the bridge over Rio Pilcoi
elev.
,
W
road. km Nee 53257 (MO. NY). BRAZIL. Acre: near mouth of Rio Embira (Tribuui
5.5 (by road) of ce
,
uth of Rio Embira (Tributary of Rio Tarauaca), Krukojf 4788 (MO, NY, U5
n 22 da Rodovia llheus^Ttabuna (BR Jardim 1102 (NY). Ceara: without
415), et al. localit
km
Heringer et al. 7183 (NY). Goias: Serra do Caiapd, ca. 5 (straight line) S
I,
km
5 de Imperatriz, Fires 16117 (MO). Mato Gross
a,
16668 (NY, US). Para: Upper Cupory River, plateau betwet
iL
&
km NE Gentry Renteria 23690 (MO). Bolivan Norosi-Tiquisio Lands of Lova, 1500-600 m.
;a Valley, 15 of Bolivar, trail. elev.
Journal of the
Botanical Research Institute of Texas
7{1)
«
n,Stehle603I(
600 MEXICO. A km
ft, Proctor 18900 (US). St: Vincent: Smith 1249 (GH, NY). Campeche: 18 S de Santa Maria Xcabak aproximadamente
al
&
MO
road to Mai Palo, elev. 800 m, Breedlove Thome 21019 (CHAPA, DS, F, LL, MEXU, MICH, NY, RSA). CoUma: 20-30 mi NE of Colima
MO,
11734 (ARIZ, GH, K, N
F,
1.
MEXU,
de Chamela, (ENCB,
iologia Perez 767 ^
Rzedowski 17976 (ENCB, MEXU). Oaxaca: Mpio. of Chiltep
i.
Quinta
brera 1688 (MO). San Luis Potosi: Mpio. de Xilitla, El Jobo, elev. 200 m. Rzedowski 10151 (DS, ENCB, MEXU, SLPM, TEX, WIS). Sinaloa:
MEXU,
Imala, Gentry 4946 (ARIZ.
F, t
W
Mpio. Gomez Farias, 4-5 km al N de Gomez Farias,
MO). Yucatan: Lolon, Flores 4 NICARAGUA. Atlantico Norte: E of Finca Waylawas, 2 km from base of Cerro Waylawas, 90 m.
(F). elev.
km NE
Pipoly 4346 (MO). Atlantico Sur: El Zapote, 40 de Nueva Guinea, elev. 130-150 m, Sandino 4834 (MO). Boaco: Camino a Mom-
N
etepe, al lado del Volcan Concepcidn, cafetales do las Angeles y sus alrededores, elev. 250-350 m, Sandino 536 (MO). Zelaya: vicinity of La
&
PANAMA.
Luz-Siuna, elev. 150-200 m. Bunting Ucht 657 (F, UC, WIS). Bocas del Toro: vicinity of Chiriqul Lagoon, Old Bank Island,
Wedel 1877 (MO, US). Canal Area: Barro Colorado Island, Croat 7694 (MO). Chiriqui: vicinity of San Felix, elev. 0-120 m. Pittier5279 (US).
Gentry 4349 (MO). Herrera:
vi
i.
Panama:
San Jo Island,Johnston 570 (GH). S
etal. 154 (AAU,
km SW PARAGUAY. &
6 of Aguadulce, elev. 0-50 m, Knapp et al. 3348 (MO). Alto Parana: Estancial Rio Bonito, Zardini
Smith 1682 (MO). P
etal.
i.
km N
Kayap 806 {MO). Cajam a, Soukup 3839 (US). C »: 1 of Pilcopata, edge of Rio Madre de Dios, e
PUERTO
Tlooking Rio Huallaga, 350 m, Gentry 37716 (MO). RICO.
elev. et al.
On
Adjnntas: Base of Pico Guilarte, 100 m, Liogier 10015 (NY, US). Agnada: road from Rincon to Aguada, Vives 2999 (UPR). Aguadillas:
Aguadillas, Sargent 573 (US). Aibonita: Inter Aibonita et Cari-Blanco, Sintenis 2946 (US). Cabo Rojo: Cabo Rojo in Monte Grande, Sintenis
&
709 (MO. US). Caye; Caye, Hess Stevens 2924 (NY). Ceiba: near Ceiba, Wagner 692 (A). Fajardo: near westernmost radio tower just S of the
Route3/Route 985 comer, 10 m, Taylor 9133 (UPRRP). Isabela: Bosque Insular de Guajataca, Little 13516 (A, NY, UPR, US). Jayna: Ja-
elev.
Km
)^a. Sargent 3050 (US). Juncos: Juncos, in Monte Goyo, sintenis 2038 (US). Manati: 26, Manati-Villalba highway. Little 13712 (US); Na-
U
V 465-720 m, Shafer 3476 (US). Penuelas: Penuelas, Sintenis 4883 (MO,
N
Mameyes U
Coastal hill 2 mi. of (Palmar), elev. 50 m. Hartley 13376 (A,
;
outes 162 and 717, Taylor 6432 (UPRRP). San Juan:
I
B
Gregory 527 (UPR). Vega
e,
1
Journal of the Botanical Research Institute of Texas
7(1)
16342 (MO). Dis-
lidadde
SW
de
il los tanque de la Electricidad de Caracas (Cocuizal), elev. 900-1,100 m, Steyermark 91829 Guarico:
(US).
km SW
15 of Tamaco along main highway between
Altagracia de Orituco and Chaguaramus,
elev. 150 m, Davidse 4218 (MO). Maracaibo:
SW
uatro Bocas, km
Aristeguieta 6824 Merida: 30
et al. (US). of Mriida, along road to El
irais the most widespread
species of section Gerascanthus, and
very
it is distinctive in evident
its
Metepubescenceontostems,
leaves, inBorescem^ebranches,
and
calyxsott
slenlellhaslhe
smallest Dowers mm
ot the
seclion with
corollas than
less 14 long. most
Itis closely related to
an most
easi y confused with
Coidlu trtchotmm, mm
hut differs in having smaller Dowers,
(8.5-)9 5-12(-14)
>“
rT”
4nt domatia, which
C. Iricholoma does not
i """I'
Q^ndla^
?
S^uuT:^
inn. Missouri
Bot. Gard.
74(3):672-673,
f. 2. 7. (Fig. 2). Type: GUATEMALA,
Contreras 6835 i
(holotype: LL; LL
isotypes: DS, US)
, F.
1.3-
1.4-
ersdlstylotm, on
short spurs L5
rmTongXTubuLT5‘!^t‘'T‘"T^r''^'^^^
to
evenly
lobed and 7“^'
tearing upon 7”
dehMng
dehiscence
or
circumscis^h
o
sfriate
Hremlent; T
corolla white,
tubular with somewhat
sprLing
m
mm lobes 14 2 16 8
oblong, mm
5.2-6.2
long, 4.8-5.9 mm
wide
the tube 4 7-5
mm 1 I mm
long, the upper
7-7.6
9.8-11.5
free,
glabrous,
the
anthers oblnna 9 t
mm ^
2.5 mm
broad; mm
disc
anular,
1.4
3 k
tall 1 hrni,4
mm ^
1.6 long,
the stigma
lobes
clavate. Fruits unknown^
occurrs^a7;d'::!:re:;™^=’^“"^"'-"»”'^'^^
ntlated and
to. most
easily"i^nhld
iving
a calyx
that shorter
is
A
31:186. (F»
1950.
3),<
«(l.M.Johnst.)Borhidi.
nal,
compressed
cymose-panicu-