Table Of Content2
TWO NEW
FROM THE BREVANTHERUM CLADE
SPECIES
OF SOLANUM (SOLANACEAE) FROM EASTERN BRAZIL
Leandro Giacomin Lynn Bohs
L.
Department ofBiology
University of Utah
257 South 1400 East
Av. Antonio Carlos, 6627, 31270-901 Salt Lake City, Utah 841 12-0840, U.SA.
MG,
BRAZIL
Belo Horizonte,
[email protected]
Stehmann
Joao
R.
ABSTRACT
RESUMO
INTRODUCTION
Howering plants and contains approximately 1500 spe-
among genera of
Solarium L. (Solanaceae) the largest
is
comprehensive taxonomic
has been focus of a species-level
cies with a worldwide distribution. Since 2004, it
Knapp
species (the FBI Solanum project; et al. 2004;
inventory aiming provide online descriptions of all its
to
new
from
has led to the discovery of several taxa
This collaborative effort
httpy/www.solanaceaesource.org).
&
&
Knapp Knapp Nee
Orozco
2006; 2007;
Granados-Tochoy
different 2005;
countries Peralta et al.
(e.g.,
& &
&
& Knapp 2010a Farmggia Bohs 2011; Gia-
Bohs 2009; b;
2009; Nee 2006- Stem Bohs 2009; Tepe
et al
&
comin & Knapp 2012). The project was coincidentally developed together with
Stehmann Vorontsova
2011-
i
8ot hKt -
Res. Teas
7(1): 95 107.
Journal of the Botanical Research
Institute of Texas
7(1)
a large-scale initiative in Brazil to achieve the Target 1 of the Global Strategy for Plant Conservation adopted by
the parties to the Convention on Biological Diversity (CBD) in 2002: the construction of a working of
list the
plant species for the country (Forzza The
et al. 2010). construction of the Brazilian of Solanaceae (Stehm-
list
ann
et al. 2010) revealed several poorly known or unknown and two
taxa, of these are described
here.
Previous molecular
phylogenetic have
studies Brevantherum
identified the clade as one of the 12
to 15
&
main
lineages of Solarium (Bohs 2005; Weese Bohs
2007). This clade has been
target of intensive studies by
the authors in order to better understand the evolution of the group and the relationships among
species.
its It
New
has a World
distribution, occurring from the southern United States through Argentina, with a center
of
diversity in Brazil. It comprises unarmed herbs to shrubs with unbranched to variously branched and
stellate
trichomes encompassing
at least six sections treated in past
classification systems. Molecular data have
identi-
&
two
fied subclades within the Brevantherum Weese
clade Bohs 2007 more
(see The
for subclade
details). first
composes members
of Gonatotrichum which
S. sect. Bitter, are herbs to shrubs, mainly with an indument
of
unbranched
trichomes
(except for Ugnescens Femald), and
S. are easily characterized by
the explosive dehis-
&
cence of their fruits (Stem Bohs Stem
2012; The
et al. 2013). other subclade contains from
species
five sec-
&
tions treated in past classifications Brevantherum Cemuum
(sects. Seithe, Carvalho Sheph., Extensum
D’Arcy,
Lepidotum and
Seithe
Stellatigeminatum and
Child), share branched
all to stellate or peltate hairs as a main
character. Molecular phylogenies based on
three gene
regions (nuclear ITS and GBSSI and
plastid tmT-F)
indi-
cate that Solanum Brevantherum
sect. is largely monophyletic Giacomin
(L. et al., unpub. data), but the mono-
phyly of
the other four sections has not been established and among
the relationships them
are uncertain. The
limits of these sections are not well demarcated and
they are in dire need of a revision, as pointed out by other
authors Nee
(Child
1998;
1999).
Although most
recent
classification systems
place species with lepidote scales in Solanum
Lepidotum
sect.
and
species with large leaves and usually
bristly-pubescent or scaly inflorescence axes Cemuum,
in
sect. there
no
is similar consensus about the characters defining Extensum and
S. sects. Stellatigeminatum
(Child
1998;
Nee
1999). Child (1998) proposed that the concept
of S. sect. Extensum should be applied only
climbing
to or
sprawling from
species Central America with somewhat
foliaceous calyces and
sympodia
plurifoliate even
though
the protologue
for Extensum
S. sect. did not the sympodial stmcture
list as diagnostic (DArcy
1972).
The
species from South America
with
foliaceous calyces
(a diagnostic character cited by DArcy,
but with
1972),
difohate-geminate sympodial
units were placed
in
S. sect. Stellatigeminatum by
Child Nee
(1998)
(1999) dis-
agreed with
this proposition and, DArcy
like included
(1972), the species with
foliaceous
accrescent calyces
and
fasciculate axillary inllorescences
in Extensum. The
sect.
entire Brevantherum
clade currently under
is
by
investigation the authors
in order to better understand
its infrageneric groupings
While
sampling
the species of the Brevanthemm
clade from
Brazil
for phylogenetic studies an unde-
scnbed was
species revealed from
Santo
Espi'rito
state. Also, while
reevaluating
the
species limits of Sofonum
megalochiton Mart, from new
the northeast, a segregate was
species
discovered. The new
species do not have
lepidote scales or scaly
inflorescence and
axes,
are expected
belong
to
to the undefined mixed
which
clade
to
the type
species of sections Extensum and
Stellatigeminatum
are assigned.
MaKrial fram
he following herbaria was
, reviewed
(acronyms
from
Index
Herbariorum.
hupV/sweelgum.
PEUFR, RB
and
UFP,
as
well as the non-indexed
HST R
1
nes and Crilma (lUCN
2010) and
for the extent of occurrence
mtalmum
calculation convex^^lJ^n^CB
a
b^d
was
on
Roe
(1971) and Mentz
et The
al. (2000).
delimitation new
of the taxa was on morpW-
based
a
1995).
:t
Taxonomic Treatment
nov 2A-F, 3A-D).
(Figs. T
1,
BHCB[BHCB0027301;ls
Shnibs to small trees up to 5 m, usually with a prominent trunk, branching in the upper stems, these spread-
ing. Young stems moderately to densely hirsute to hispid with few-rayed (4-6) porrect-stellate trichomes, these
mm
mm,
sometimes multangulate, on multiseriate stalks 0.3-1. 2 long, the rays 0.2-0.8 usually with a
2-3-celled central ray longer than the 1-celled lateral ones; old growth glabrescent to pubescent, usually with
broken trichomes comprising a hispid vestiture of multiseriate stalk remnants. Bark of older stems yellowish-
white, usually shiny; new growth whitish-green to dark purple near the insertion of the inflorescence, always
shiny. Sympodial units usually rarely di- or plurifoliate, often with solitary leaves; if geminate, leaves
trifoliate,
up
paired only branching points and differing mainly in size, the smaller to the size of the larger ones.
at
cm cm membra-
narrowly
Leaves simple, the blades 1.7-11.8(-13) long, 1.2-4.8 wide, elliptic to ovate-elliptic,
when
naceous, weakly discolorous (dark shiny green above, pale green beneath fresh), the abaxial surfaces
on
moderately to densely hirsute, the lamina always visible, with short- (more often the lamina) to long-stalked
(more often on primary and secondary veins) porrect-stellate trichomes with 3-5 lateral rays supported by a
mm
and a 2-3 celled central ray somewhat longer than the lateral ones (Fig.
multiseriate stalk 0.3-0.9(-1.4)
mm
moderately densely hirsute, the lamina always visible, with
2B), 0.8-1.2 long, the adaxial surfaces to
mm
up and (more often on the lamina)
mixed hyaline unbranched trichomes (Fig. 2D) to 1.8 long, sessile to
am
on primary
short-stalked (more often
few (1-4; Fig. 2C) lateral rays and a 2-3-celled c
mm
camptodromous, impressed or not above
up venation (barely
supported by a multiseriate stalk to 0.4 long;
5-9 base rounded cuneate, asymmetric;
secondary veins pairs; to
visible in dried material), raised beneath, the
on
margins apex acute to long attenuate-caudate (only acute the smaller geminate
entire, slightly revolute;
mm
with the same trichomes as those of the abaxial midrib. In-
leaves); petioles 1.8-5.5 long, densely hirsute,
when
an unbranched cyme, terminal, lateral, intemodal, or subopposite the leaves, in-
florescence elongated
cm
comments; 3A-B), the axes (0.35-)0.9-4.3
branch Figs.
temodal, changing the orientation of the (see 1,
cm
moderately
peduncles 0.1-1.47 long, hirsute
pedunculate, the
long, with (3-)6-15 subsessile to
flowers,
with those of the stems, the rachis pale green, densely pubescent with three
porrect-stellate trichomes like
eglandular or glandular-capitate trichomes, sessile to
types unbranched, 2-4-celled,
of trichomes: hyaline,
with few (1-4) lateral rays or only the 2-4-ceIled, sometimes
trichomes
short-stalked multangulate-stellate
much than the^oAers, and supported by a inultismate
longer
glandular-capitate central ray developed, this
mm cm
cm 1.9-2.5 in fruit, pale green to dark pink to purplish at
to 0.9 pedicek 0.3-L2 long in flower,
long;
mm same unbranched
pubescent with the trichomes
densely
base, articulated at base, 1 in diameter at base,
mm
mixed with
up 2 long, sessile to short-stalked
found on with glandular head, to
the rachis, almost all a
Buds elongated with developed
multangulate trichomes with 1-4 lateral rays and a longer central ray. slightly
when developed. Flowers perfect, 5-merous. Calyx tube 1-2
calyx included even fully
lobes, the corolla nearly
mm
mm
mm
6-8 3-4 wide in flower, the apices acuminate,
long,
long, with narrow-ovate lobes
lanceolate to
mm
mm
8-10
8.6-12 long, wide, the apices
broadly ovate to cordate,
lobes
ehairswith
acute, densely covered abaxi
= 5(»|im;E-J=100|ini.
Journal of the
Botanical Research
Institute of Texas
7(1)
New Solanum from
Giacomin et species of Brazil
al.,
unbranched
with
and the central ray always longer than the others and often glandular, adaxially
shorter stalks
mm
up
trichomes 2 long
(sometimes with a poorly developed secondary ray) glandular capitate to
2-4-celled
commonly found also on the margin, associated with sessile to subsesstle
gland these
2E), the ellipsoidal,
(Fig.
on midveins and near the base. Corolla
concentrated the
globose dark glandular trichomes (Fig. 2F), these
mm
mm
4-6 4.5-6.5 wide,
1.3-1.7 cm in diameter, white, stellate, lobed up to to the base, the lobes long,
moderately pubescent
spreading anthesis, sparsely to
with involute apices, at
deltoid to ovate, acute, slightly
unbranched
with the same trichome types found adaxially on the calyx and with a tuft of small eg-
abaxially
mm
Filament tube up long, with a minute free portion less
landular trichomes at apex, glabrous adaxially. to 1
mm
mm
mm
anthers 2.8-4 long, 1.3-1.5 wide, sometimes slightly unequal, elliptic to
than long, glabrous;
1
opening
oblong, connivent, yellow, glabrous on both faces, poricidal at apex, with the pores directed introrsely,
mm
6-7 stigma clavate, with a papillose
Ovary long, cylindrical;
with glabrous; style
longitudinal age.
into slits
when
cm green when immature, purple to black
white pale
Fruit a globose berry 1-1.4 in diameter, to
surface.
which spreads open
brown completely covered by the accrescent calyx
drying dark,
mature, shiny, glabrous, to
mm
mm
golden
1.7-2.3 wide, flattened, reniform, to
2.3-3.4 long,
maturity. Seeds 9-31(-35) per fruit,
at
with
brown, the surfaces irregular pits.
W.W nomas (BHCB, NY. RB). Bahia: Mun. Ibirapitanga, Far. Paineiras.
May 12441
550-600 m, 14 2001 et al.
9»13.53'S, 35“52.776'W, (fl, fr),
do Urubu, 20 Apr 1994 A.M. Miran-
Pernambnco: Mun. Maraial^agoa dos Gatos, Serra (fi, fr),
Feb 1998 ConceifOo 213 (MG).
17 A.S.
(fl, fr),
&
da 1628 (EAC, HST).
L.P. Felix
m
up with
5
forming small trees to tall
with a robust habit,
Discussion—Solanum anisocladum a species
is
Solanum meg-
and drying dark. similar to
dark green, It is
upper surface shiny,
prominent trunks, with the leaf
m
covered
up 3 with the surface matte, often
much branched shrubs lo taU.
aloAitm which nonnally fonns
when Both species share an inflated calyx that
green dried.
and normally pale
by indument,
a whitish-yellow
was previously used to separate S. tnegulocliilOT
This feature
maturity (Fig 3D).
is strongly accrescent at fruit
other characters that
presents several
However, utiisoclodum
didymum Dunal. 5.
from such
similar species as
S.
and unbranched hairs reaching
indument of very long stellate
readily dislinguUh from megalochiton. Its
S.
it
naked The old growth
mm the eye.
2A-B) visible to
easily
more than 2 long and drying ferruginous (Figs. 1, is
The
trichomes remain.
stalks of the stellate
where only the multiseriate
becomes somewhat hispid
hirsute to
mm
normally transparem to yellowish
much reaches up to 1.5 long, is
indument in S mcgulocbiton is softer,
unbranched
robust
when become hispid. The inflorescence is also a distinctively
and stems never
dried, the
more than 4 cm long whereas the inflorescences of S.
cyme sometimes with an axis teaching
3A-B),
(Fig.
composed of eglandular porrect-
indument
adaxial leaf is
than 2.5 cm. Its
normally shorter
megulocliilon are
represem-
apparently
with muUicellular bases,
unbranched hairs, the latter
trichomes assocUted with
stellate
composed
only
adaxialleafindumentofS. megulocliilon is
The
ing without developed lateral rays.
hairs
stellate
branching pattern not
has a distinctive
also
Solumim utiisoclodum
of sometimes glandular.
stelUte hairs these
the^s,
Aejnflon
When from between
arising
observed in megulochiton (Fig. 3A).
S.
herbarium sheets, I
on small brar
1
always visible
Strong change growth direction (not
in
of megalochiton are
flowering brai s S.
the
contrast,
in every specimen seen the In
at field).
straight.
was occurring
megulochiton cited as tn
(Stehmannetal.2010),SoIunum
In the Brazilian plan, species list
Pernambuco and Bahia no
Duni rbaria in
and Bahia.
Alagoas
northeastern Brazil from the states of
seenco-occutnngwithS.umsocl4dum.AsheetfromAlagoas(71iomusl244I;BHCB.
specimens matching were
it
umm^
confo^
our concept of
acmally to S.
experra
Solunum
by other
NY,
RB) megulochiton
determined as
S.
isknowntooccuronb-mafewoc^onsmthe^^^^^^
megulochiton
cludum. Thus,innor.heas.ernBraz.l,S.
m
oun
ttoncana
ofthesouthempartofBahiaandnotfurthernorthward.Solonummegooc
aKsoud,eas.ernandsouUremregionsofBrazil.wi.halewaddi.ionalcollecnonsm.hesrateofGo^andMa.o
butth.scanno.be
sympaurtc,
S.megukxhit™ are not
and
Sunisocludum
Grosso.
This suggests that
(e.g.,JPB).
group
because of the
not important collections
all
tog
pan
when
ingiin unequal branches the infloresce from between the nodes.
i
—
and Solanum
Distribution habitat. anis >ccurs in forest edges of the Atlantic coastal rainforest
of
northeastern Brazil in the states of Alagoas, Bahia, and Pernambuco, in partial to complete shade, at elevations
m
of 500-700 (Fig. 4). Primary wet forest fragments at this elevation are quite rare in northeastern Brazil today,
and the type collection, despite the fact that was found in a forest edge, was from a well preserved fragment
it
—
Phenology ^The species was collected with flowers and fruits from February through May, but a flower-
.
ing peak was observed in February and March, and developed were found from March through
fully fruits
May.
MCP
only three localities and its calculated extent of occurrence using the is 834 km^. Despite the fact that the
MCP
Bahia from one Pernambuco narrow
registry is far the in state, the reflects the range of coastal rainforeS
where the species is expected to occur. Although the type locality is now part of a private reserve owned by a
sugar company (RPPN Frei Caneca), consists of a small group of forest fragments in a severely damaged
it
landscape mainly modified by based on monoculture
extensive agriculture (plantations). Generally, the Atlan-
damaged
tic coastal rainforest has been severely in northeastern Brazil in the past decades due to urban expan-
sion and extensive farming.
much
Shrubs, up to 3 m, sometimes scandent, branched, the apical branches scandent, held downwards,
arch-
ing. Young stems densely pubescent with sessile to short-stalked porrect-stellate trichomes normally with 8
mm
lateral rays, rarely fewer, commonly of unequal length, up to 0.3 long, with a central ray much shorter than
growth pubescent
the lateral ones (Fig. 2G); old sparsely to glabrescent. Bark of older stems bright white to
yellowish, shiny; new growth gray due to dense vestiture, with a white to dark purple lamina Sympo-
beneath.
dial units 2-3 foliate, geminate, with paired leaves differing in size and shape, the smaller ones than of
less
cm
cm
the size of the larger. Leaves simple, the blades 2.2-7 long, 1-3 wide, elliptic to broadly (the
elliptic
sometimes
smaller geminate leaves broadly elliptic to ovate, slightly obovate), membranaceous
to chartaceous,
slightly discolorous, drying dark adaxially, the abaxial surfaces sparsely to moderately
pubescent, pubes-
the
cence denser on veins, with sessile to short-stalked porrect-stellate trichomes with 4-9
up
lateral rays 0.3
to
mm
long, frequently unequal in length, and a minute central ray (Fig. 2H), always shorter than up
the
laterals,
mm
mm
to 0.1 long, the stalks multiseriate, reaching 0.1 long, the adaxial surfaces glabrescent,
with only
hairs
mm
more up
frequently rayed), to 0.3 long, frequently
unequal,
; with
a slightly
mm
the abaxial surface (Fig. reaching 0.2 long, but
21),
still shorter
than
the
lateral
m
camptodromous, slightly to not raised above, raised beneath, the secondary
veins 4-6
pairs;
base cuneate to obtuse (rounded on the smaller geminate leaves), asymmetric; margins
entire, apex
plane-
densely pubescent with the same trichomes as the stem and abaxial midrib.
Inflorescence
a
congested un-
branched cyme, intemodal, or subopposite the leaves, rarely terminal, the
lateral, axis
sessile to up
subsessile
nm
or absent, green, moderately to densely covered
with
the same
porrect
very
closely
spaced;
pedicels
Journal of the Botanical Research Institute of Texas
7(1)
Fk. 5. botype of Sokmum aektolm Oiacomin & Stehmann. (K Dtmmer et aL 3273, BHCB [BHCB 002734]). <
