Table Of ContentAND
PHYLOGENY, BIOGEOGRAPHY
SYSTEMATICS,
OF (MELASTOMATACEAE)
CHAETOLEPIS
Grimm and Almeda
Daniel Frank
Department of Botany
Academy
of Sciences
California
55 Music Concourse Golden Gate Park
Drive,
dgrimmO @yahoo.com; [email protected]
I
RESUMEN
INTRODUCTION
montane shmbs and subshrubs
genus of
neotmpical
mostly
small
Mi,. (Melastomalaceae) a
is
charac,enaed^4.merous,d.^^^
readily
thatare
species
fatelyannuaU). consismofll
As
interpreted here,
it
welWeveloped pedoconnecuves^
lack
stamens that
«emonous isomVic inappendrcula.e
mostly
flowers,
compre^flatened
mostly or
withatestathatconsists
seeds
MocuUr.aptcallyse, and cochleate
use ovaries,
Andean
hasindudedspecresof South
inthe past,
circumscribed
i»h"ligi.adngcefc. The genus,asithasbeen
Cuba Recently the motrotypic genus
and
America,
America, Central
the Guayana Highlands, southern
''-ophlr.NaudimLmweltropicalAfricawas.ramferredtoCIwtdephO-que^Felix
monophyly of the geographically dispa-
Because the
Study
has attempted Chaetolepis or Nerophila.
evaluate
to
seemed ripe for testing.
been evaluated, it
rate had never
assemblage
Chaetolepis
of^taxa refel^ed to
218
The
primary goals of this study were to determine if Chaetolepis is monophyletic, assess the
phyloge-
netic relationships of the c species based on morphology, and evaluate character evolution by
using
A
cladistic techniques. L ttaa: xonomic revision is provided that includes detailed descriptions of all recognized
n microscope images of seeds summary known
for selected species, a of chromosome
numoers, miormation on and
habitat geographic distributions, flowering phenology,
distribution maps,
il-
lustrations of all recognized species, citations of specimens examined, and a list of excluded taxa. This was
supplemented by
limited field study of four species in Costa Rica, Colombia, and Venezuela.
—
Tribal and Generic
Affinities. Chaetolepis belongs to the tribe Melastomeae, the core mbersof^^
i
are characterized by
cochleate seeds with
a testa that is tuberculate or papillate, a crown inenomes
oi ur ap-
pendages on
the ovary apex, and the presence of staminal pedoconnectives (Renner and Meyer
2001;
Freire-
Fierro 2002; Michelangeli Based on
et al. 2013). overall morphological Wurdack
similarities, (1973) hypoth-
esized that Chaetolepis most No
is closely related to Tibouchina Aubl. cladistic analyses or molecular
data,
however, had been used
to assess the possible
sister relationships of Chaetolepis. Other neotropical
relatives
of Chaetolepis include Don, &
Aciotis D. Arthrostemma Pav. ex D. Don, Heterocentron Hook Am.,
Microlicia
D.
Don,
Microlepts (DC.) Monochaetum
Miq.,
(DC.) Naudin,
Nepsera Naudin, and
Pterolepis (DC.) Miq., Rhexia
based on
L. a study using ndhF and cpDNA
rpll6 sequences 24 members
of of Melastomeae and outgroups
(Renner and Meyer
2001). Based on an expanded DNA
molecular sampling
using sequence
data, Bucquetia
DC.,
Castratella Naudin, and Monochaetum now
are considered
the
closest sister taxa of Chaetolepis (Peimeys
etal. 2010; Michelangehetal.
2013). Like
Chaetolepis, the majority of
species in thesegenera are also restricted
to higher elevation montane
environments
Andes
in the
of northwestern South America and
have
tetramer-
all
ousn^ers,
an
apicallysetulose ovary
thatUfourlocuUrandfree
tarn hypamhium,
the and
capsularfrulB.
Morphology
(Characters Taxonomic
of Value).—The
species of Chaetolepis are easily dis-
tinguished from
each
other by
characters
involving
the indument,
anther
connective, seed surface, petal
coloration, and
petal margin. The indument
found on
leaves, intemodes, and
nodes, organs includes
floral
eglandular tnchomes,
glandular
trichomes,
barbellate/dendritic
trichomes, and The
scales. distribution of the
structures vanes
greatly. ranges from
It absent to densely covering the
surfaces^IfTeeeLtS^lnTfl^^l
organs. The
anther
connective
1^ prolongation ranges from absent to minutely
nrolnno^ft
t 1 i T
dwTrwtb
1^^^^
prevailingly smooth with compressed/flattened
imerdmatim'^ZttZ
t
Lk interdigitating The
cells. petals vary in color fromyel-
llooww iTalel
ttoo pink oorr magenta anHd
their margins
are either
glabrous
or
ciliate
ainoris.ics.udi«(Su„dCl9MTribJuW^™'’'lTo^^^
Wurdack
Almeda
1973; some
' Over of
2001). the past 80 years
the soecies have been k a
•
.he placemen,
of
Leon and
Alain MamnsT989)
1974;
ciesa„d.h.eevade.ies.
Because
a,
^
identify symapomorphies
and
only cumorv w,
aupmlon
“
'
vegetative and ”
reproductive llco^tunWiSr
chamctem.
monograph
example
of
some Melastoma.aceae,
of the °f for
chatacters used
to characieri
varyingin
color from rose .op„rple;4-Cw^t.ts“f"''“^^^^
smZr
connectives; and
ovoid
cochleateLds
With
ofasmallersamplingofspecielTstiWbuX^^^^^^^
T
and
the seed
surface appeam
-smooth
or dimta
fit^ly and
No
any
^ ^uriculate.” work mentioned
sculptural prior
details of the seed
SFM
testa because n
^
,1
and
consistency
detail relating of
to the seed and
other
mavlT'^
structures
perhaps
iolepis, biased by
the
four-merous
flowers and
‘U greltirreduc^
greatly reduced some
pedoconnective. Furthermore,
219
monophyly
doubt about the of the genus has persisted because no cladistic analysis of the group has ever been
and
undertaken to evaluate character states their evolution.
—
Ingroup. ^All sixteen taxa thought to comprise Chaetolepis form the ingroup for this study. Chaetolepis alpina
var. alpina, C. alpina var. latijolia, C. anisandra, C. cubensis, C. ci^odontisii, C. gentianoides, C. lindeniana, C. lori-
carella, C. microphylla, C. perijensis var. perijensis, C. perijensis var. glandulosa, C. phelpsiae var. phelpsiae, C.
C
andC.
phelpsiaevar. chimantensis, C. santamartensis,
sessilis, thymifolia.
Outgroup.—To monophyly among
assess the of Chaetolepis, seven species the genera Tibouchina, Mi-
and Monochaetum were used as the outgroup. The exemplars used were gleasoniana Wurdack,
crolicia, T. T.
Wurdack, Almeda and Naudin, M. Cogn.
narinoensis wurdackii Todzia, Microlicia canastrensis obtusifolia
T.
ex Romero, Monochaetum amabile Almeda, and M. compactum Almeda. Tibouchina was chosen as part of
R.
common
Wurdack most
outgroup because hypothesized shares a recent ancestor
the (1973) that likely
it
with Chaetolepis based on many shared characteristics, such as perigynous flowers, campanulate or urceolate
and
hypanthia, superior ovaries that are apically pubescent, capsular fruits, cochleate seeds. In addition, T.
^easoniana, narinoensis, and wurdackii were chosen as part of the outgroup because the tuberculate seed
T. T.
The
testa and foliar scales (see Todzia and Almeda 1991) are identical to those found in C. cubensis. species of
Microlicia and Monochaetum were chosen other members of the outgroup because of their close phylogenetic
as
proximity to Tibouchina based on molecular studies by Renner and Meyer (2001).
Material Examined.— Over 400 herbarium specimens, including types, were used in the phylogenetic
analysis and taxonomic Specimens from the following herbaria with significant holdings of Central
revision.
DUKE,
and on BM, BR, CAS, COL, DS,
South American Melastomataceae were borrowed or studied site: F,
UTMC images
FMB, G, GH, HECASA, M, MO, NY, and (acronyms according to Thiers 2012). Internet
K, US,
P,
on
of type specimens OSTOR Plant Science) were examined for those taxa that were not readUy available loan.
and The
Characters.— and characters were used in the analysis (Tables 1 2).
Forty-one vegetative floral
was informative in this study.
type of indument and structures particularly
covering both vegetative floral
Data Analysis.—The program PAUP* 4.0bl0 (Swofford 2002) was used to generate phylogenetic
software
matrix
estimates. Maximum parsimony with branch and bound search was selected to evaluate the data that
a
Two were conducted. Analysis
was generated in Maclade (Maddison and Maddison 2000). analyses 1
4.03
ingroup
was provided insight into relation-
performed and analysis 2
monophyly Chaetolepis
to assess the of
was prompted by the results of the
ships cubensis)
after the exclusion of one of the species of Chaetolepis (C.
formed
and 23 Sixteen species the
taxa.
first analysis. The analysis used 41 morphological characters
first
and formed
Monochaetum, Microlica the
presumed
ingroup and seven species from the genera Tibouchina,
was appUed
backbone constraint that
a
outgroup. analysis
Default options were used run the analysis. In 2,
to
unique and/or seed characters
grouped because of their floral
outgroup members together
all plus C. cubensis
&
Mad-
MacClade (Maddison
grouped was done by constructing a tree in
ingroup This
all taxa together.
loading the
dison PAUP (Swofford 2002), derooting the tree,
2000) with opening in
the data matrix, the file
the search.
constraint in
the
as a backbone and enforcing
finally
constraint in the analysis,
generated, a bootstrap analysis
phylogenies
To various
evaluate clades in the
the strength of the different
emp
branch swapping was oyed; ot er-
employing
Stepwise-addition
PAUP.
a heuristic search was utilized in
default
options were
used.
.
The
among the lineages of Chaetolepis. “l
MacClade
was icters
used
to infer the evolutic
ges,” and were chosen.
“all possible changes” options
mo
conducted^
a™lys« were
O
Chartolepis,
gain — biogeography of
insight
into the historical
witnm
taxa
patterns of the diilerent
peerrffoormr^mserdi j_ r^rioin aanndd ddiissopeerrssaall patl
.. origin
to provi; de insight into the
MacClade, a data mai
^«nca. opir
used Fitch
Central America, and This analysis
Africa.
'°"P«ratingallChaetolepi
glands
sessile (2).
5 {1);
and Almeda, Systematics of Chaetolepis
Grimm
PAUP
with
generated in
in which they are found (Table and one of the 26 most parsimonious tree topologies
3),
may
how have
Chaetolepis taxa
was performed to infer
polytomies resolved The second analysis
arbitrarily.
number
only in the of area states (five)
migrated within the Andes. This analysis differed from the first analysis
were examined.
(MPR’s)
reconstructions
used. In the analysis, equally most parsimonious
all
Chromosome
H.
Cytology
Camoy’s (Bradley 1948) solution (4 chloro-
modified
Horal bud material was collected in the field, fixed in
under
70% ethanol and stored refrig-
form, 3 24 hours, transferred to
ethanol, glacial aceitic acid, v/v/v) for
1
v
Chaetolepis cufodontisii
eration The new chromosome count reported here for
until studied.
andalOOxoil
with phase contrast
microscope
acetocarmine smears mother using a Zeiss light
of pollen cells
The
2100x.
magnification of
made by camera lucida at a
immersion were
Drawings meiotic figures
objective. of
among specimens cited.
voucher new marked with an asterisk
collection count
for this is
on
based dried her-
characters
and reproductive
vegetative
all
:e
an
mature open flower in the center of
one
from
either
taken
e
numerous
being recorded,
were
ot
were characteristics r
scored from matur^branches. To eLure that aberrant
were
specimens used.
were examined and with typical traits
those
“actoljs
morphobgy of . i
A seed
study the
scanning efecmn (SEM) was used io
microscope
t«th gold-palladium
and sputter-coated
tape
double-stick
Seeds were aluminum stubs with
affixed
to
laboramryatthe^^^^
microscope
^l»y.San,ples scanning electron
were photographed the
in
SEM
Voucher collectton data for the
20kV,
microscope at
“(Sciences using Leo 14M BP scanning electron
a
‘“'“ges are provided legends.
in the relevant figure
RESULTS
^fogeny.-in
22 n
the analysis,
first cladistic
was and the reter
0.66
224
which
Cl, of nine are unique,
three are homoplasious
and
one
include
ambiguous. The
is unique characters
^te,
appresse
flattened
scales, basally
attached with
a
raised thickening near the point where the scale is at-
’
tached to the leal surface and an erose margin on
the hypanthium
base, hypanthium surface,
apex, adaxial leaf
abaxial primacy
leaf surface
veins, branch
surface
intemodes,
branch surface nodes Lercalydne lobe sinus,
alyxvesiuure
excluding
the midrib, and
calyx midrib
(characters
3, 6, 11, 17, 20 23 25 28 31 respectively).
(char-
m homoplasious
""oo*
‘>"= egUndular smooth glandular tri-
trichomes,
MUK/d
S^mes
or b
Systematics of Chaetolepis
Almeda,
and
Grimm
226
ffi
