Intraspecific Variation in Four Distinct Populations of Anemia villosa Humb. & Bonpl. ex Willd. (Anemiaceae) Occurring in Rio de Janeiro, Brazil

2011·5.5 MB·English

by  Maria Luiza Ribeiro Da Costa Ribeiro

Four
Intraspecific Variation
in Distinct Populations of
Anemia
Humb. &
villosa Bonpl. ex
Willd.
(Anemiaceae) Occurring
Rio de
in Janeiro, Brazil
Maria
Luiza da Costa
Ribeiro Ribeiro*
UNIGRANRIO
Escola de Ciencias da Saude,
Marcelo
Guerra Santos
Departamento de Ciencias, Faculdade de Formagao de Professores, Universidade do Estado do Rio
de
Janeiro (Brazil)
Claudla Franca Barros
Laboratorio de Botanica Estrutural, Instituto Jardim Botanico do Rio de Janeiro (Brazil)
Costa
Cecilia G.
The
diversity of plant communities comprise
that the Brazilian Atlantic rain
complex prompted number
forest has a of studies on functional variation of
species
that are vi^idespread in various habitats within this complex. All of
these studies have indicated a large degree of intraspecific variation between
distinct populations at different habitats, regarding both anatomical and
physiological traits (Scarano et al., 2002; Scarano et 2005; Mantuano et al,
al.,
2006).
Anatomical and may
morphological
characters serve as reliable indicators in
and
the study understanding
of ecological adaptations of living organisms
(Fahn, 1964]. Environmental conditions, such and
as light intensity nutrient
availability, can influence the size and shape of leaves as well as anatomical
many
aspects. For example, high
in light habitats, leaves of plants tend be
to
and more
smaller deeply incised than under shadier conditions (MacLellan,
known
2000). In addition, there are other well shifts in leaf traits such as
* Corresponding author. Escola de Ciencias da Saude, UNIGRANRIO, Rua Professor de
Jose
Souza Herdy Duque
1160, 25 de agosto, de Caxias, Brazil, [email protected]
RIBEIRO ET AL.: INTRASPECIFIC VARIATION IN ANEMIA VILLOSA
cuticle thickness, epidermal stomatal frequency and length, and
cell size,
trichome frequency and length that are often related environmental
to
conditions (Olson and Rogas Hlwatika and
Carlquist, 2000; et al, 2001; Bhat,
2002].
common
on morphology and anatomy
Studies fern are (Kraus et al, 1993,
Gragano et al, 2001; Chaerle and Viane, 2004; Pita et al, 2006a, however
b),
on such
studies intraspecific variation in traits in ferns are not usual (Liu et al,
Boeger al According Mickel hundred
2006; et 2007). to (1962), in the last
Anemia
years, has been an explored subject for morphological studies that
have been largely concerned to the taxonomy. Little work, other than that of
how
Ribeiro al has been done on understanding environmental
et (2007),
conditions influence morphology genus Anemia.
in the
Anemia Humb. &
villosa Bonpl. ex Willd. widely distributed in eastern
is
Brazil, from Santa Catarina to Ceara, and also in northern South America from
commonly
Peru Surinam. Rio de populations
to In Janeiro State, large are
with on and
associated vegetation islands rocky outcrops (Santos Sylvestre,
According Mickel
2006). to (1962) this species has a considerable variation in
size and form that can lead to confusion with other species. In Rio de Janeiro
State, A. villosa populations exhibit two different specimen sizes. The aim of
study was analyze anatomical and environmental parameters
this to variation
in A. villosa growing at four different sites in Rio de Janeiro State, Brazil to
generate a better understanding of the degree of plasticity exhibited by this
species across a variety of habitats.
Material and Methods
Anemia
Leaf samples of villosa were collected in four distinct sites in Rio de
Janeiro State, Southeastern Brazil: Pedra de Itacoatiara (Serra da Tiririca State
Park) and Forte Imbuhy, on the coast of Niteroi; Pedra Dubois at municipality
of Santa Maria Madalena and Lumiar, municipality of Nova Friburgo. All
grew which was
populations in inselbergs, except for Lumiar, collected at the
The two
border of the Atlantic rain forest (Table populations in
1).
km
municipality of Niteroi are approximately 14 apart from each other.
each were randomly
In collection site, five plants chosen, except for the site
where From
Pedra Dubois, only were each
of three plants selected. plant, five
were and immediately
leaves collected fixed in formalin-acetic acid-alcohol
70%
and
(FAA70) (Berlyn Miksche, Fragments each pinna
1976). of last
cm^) were gradually dehydrated in ethanol and embedded in glycol
(0.5
methacrylate resin (Feder and O'Brien, Five micrometer thick transverse
1968).
made
sections were with a rotary microtome, and were subsequently stained
O
with 0.1% toluidine blue (O'Brien and McCully, 1981). The tracheary
elements were observed in material dissociated using the Franklin method
and by
(1945) stained hydroalcoholic safranin (Johansen, 1940). Identification
of vascular bundle types followed Ogura (1972) and stomata Sen and De
(1992).
AMERICAN FERN VOLUME NUMBER
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Forte Pedra de
Pedra Dubois Lumiar Imbuhy
Itacoatiara
Anatomical observations and measurements were made Olympus
using an
BX50
microscope with
light the aid of Image-Pro Plus version 4 software. The
images were
acquired by a video camera Cool SNAP-Pro.
The
following leaf anatomical parameters were measured each
for sample:
thickness
of leaf blade, mesophyll, adaxial and abaxial epidermis; trichoma
and
length density; stomata length and and
density; lignified petiole
area.
Means
and
standard were
deviations calculated from a sample size of 25 fields
A ANOVA
each measurement
for (N-25). nested was used study
to the
association between the outcome and Component
variable group. Principal
Analysis (PCA) was used
to find the higher variance components. To
the
test
hypothesis that the analyzed sample was composed
morphologically
of
different discrete groups, a Discriminant Function Analysis (DFA) was
performed
(Schlichting, 1986; Zar, 1996). Overall differences between the
compared
groups are presented by Mahalanobis The
distance.
statistical
analyses were performed with
Statistica 6.0 (Zar, 1996].
v.
As
described by Mikel populations Anemia show
(1962), different of
villosa
differences in leaf morphology. The specimens from Lumiar and Pedra Dubois
exhibit the typical oblong blade with pinnae
leaf fertile close to the
sterile
pinnae, and the specimens from have
Niterdi triangular leaf blade with
fertile
pinnae
that vary in their localization from remote to close to the sterile pinnae
(Fig.
1).
The
four populations analyzed
are similar in qualitative characters such
as
type and The
of stomata, trichomes, stele. petiole has a uniseriated epidermis
with
lignified walls along with uniseriated trichomes. The stomata
are parallel
in the adaxial epidermis, making up two and
supported above
lines, are the
common
epidermal by with Below
cells cells lignified walls. the epidermis
there are four to five layers of sclerenchyma followed by parenchyma with
some
The
amyloplasts. V-shaped and endodermis
stele the
is clearly visible
is
and shows
evident Casparian two
were
strips; layers of pericycle observed.

AMERICAN FERN VOLUME NUMBER
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_
©
o
.(D
e 0
e
Figs. 3-14. Sections of leaves of four populations Anemia Humb. &
of villosa Bonpl. ex Willd.
occurring in Rio de Janeiro state, Brazil. 3-5. Leaf of Itacoatira populations. 3) Cross section of leaf
blades. 4) Adaxial epidermis surface. 5) Abaxial epidermis surface. 6-8. Leaf of Imbuhy
populations. 6) Cross sections of leaf blades. 7) Adaxial epidermis surface. 8) Abaxial epidermis
surface. 9-11. Leaf of Lumiar populations. 9) Cross sections of leaf blades. 10) Adaxial epidermis
surface. 11) Abaxial epidermis surface. 12-14. Leaf of Pedra Dubois populations. 12) Cross sections
^"
nm ^
Figs. 200
3. 5, 6, 8. 9, 11, 12, 14; in Figs. 4, 7, 10, 13.
All populations have hypostomatic The
leaves. epidermis comprised
of
is
one
layer of cells with sinuous anticlinal walls and convex
periclinal walls, in
both and The
abaxial adaxial surfaces. stomata are desmocytic and
polocytic,
and
the guard cells of the stomata protrude a from the surface of the
little
epidermis. Pluricellular trichomes are present in both adaxial and abaxial
The
surfaces. mesophyll shows 3-4 which more
layers of arm-cells, are
compactly
arranged near the adaxial epidermis
3-14).
(Figs.
RIBEIRO ET INTRASPECIFIC VARIATION ANEMIA VILLOSA
AL.: IN 75
^
ex Willd (Mean ± standard deviation) and F values (NESTED-GLM).
ANOVA
The among
results of the nested were significant sites (F -115.67)
ANOVA
and among =
significant individuals (F 5.66). Nested results for the
among
quantitative variables groups are presented in Table All populations
2.
were
similar for one out of 10 variables (epidermis thickness of adaxial
surface). All populations were different in three variables (trichome density on
adaxial surface, leaf thickness and mesophyll thickness). The inland mountain
(Pedra Dubois and Lumiar) and coastal land (Pedra de Itacoatiara and Forte
Imbuhy) were
populations statistically different in three variables (trichomes
lengh of adaxial surface, trichomes length of abaxial surface and stomata
length).
and two components and
Factor loadings eigenvalues the (PCsl
for
first 2)
PCA
extracted in the are shown in Table These accounted for 57.71% of the
3.
total variance. Anatomical characters such as trichome density of abaxial
surface, uniseriate trichome length of adaxial surface, uniseriate trichome
and showed
length of abaxial surface, trichome density of adaxial surface the
highest (either positive or negative) correlations with PCI, and epidermis
and showed
thickness of adaxial abaxial surface the highest correlations with
PCA
PC2. Figure 15 shows that separated the populations analyzed in two
groups: one formed by the coastal land populations of Imbuhy and Itacoatiara,
and other formed by the inland mountain populations of Parque Estadual do
Pedra Dubois and Lumiar.
Factor structure for the 10 variables and eigenvalues for the two factors
first
DFA
(DFl and shown DFl was
extracted in the are in Table positively
2) 3.
76 AMERICAN FERN JOURNAL: VOLUME NUMBER
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2 (2011)
correlated with trichome lengh of abaxial surface and negatively correlated
with trichome
density DF2
of abaxial surface, whereas was
positively
DFA
with
correlated stomata
density. was 98.27%
classification individ-
for
uals from groups and
(Fig.
I, II III 16).
Discussion
In this study all populations of A. villosa showed similar qualitative
characters but variation was demonstrated
in the quantitative characters. This
quantitative variation occurred between specimens
at different latitudes,
temperatures and
altitudes in the Rio de Janeiro State showing two distinct
groups: the coastal land populations and the inland mountain
populations.
These two groups were by
described Mickel
Rio de
(1962) in Janeiro state as
specimens
with oblong and
leaf blades pinnae approximate
fertile to the sterile
pinnae (inland mountain population) and those with broader leaf blades with
pinnae
fertile varying in their origin from remote approximate
to the
to sterile
pinnae
land
(coastal populations).
Other have
studies
verified quantitative variation populations
in of
angiosperm
species that occur in different habitats or micro-habitats
(e.g.,
Rogas
et al, 1997, 2001; Hlwatika and Bhat, 2002; Mantuano The
et 2006).
al.,
anatomical Anemia
differences
in villosa are evident in traits such as trichome
density of adaxial and abaxial surfaces, leaf thickness and mesophyll thickness
between
populations and in trichome length of adaxial and abaxial surfaces
and
stomata between
length the inland mountain and
coastal land popula-
The
inland mountain
populations, submitted lower temperatures and
to
higher humidity, presented thicker leaves with longer stomata and
trichomes,
and low when
stomata frequency compared
to coastal land populations. The
difference in thickness of the leaf and mesophyll was expressed in length but
RIBEIRO ET INTRASPECIFIC VARIATION ANEMIA VILLOSA
AL.: IN
mesdens,ty
Tricho abaxlal surface
in
OOaOo'
°
Trichomeslerghonadaxiplsu^c^
°
i
v.*.
-2,0 -1,5 -1,0 -0,5 0,0 0,5 1,0 1,5
not in number of cell layers. According to some studies {Woodward, 1979;
and
Willians Black, 1993; Korner, 1999; Loeys et al, 2002) biomass allocation
is temperature sensitive, and exposure to low temperatures results in plants
that exhibit reduced investment in the shoot and leaves that are thicker than
their warm-growth counterparts. Differences in stomata frequency were also
observed by Hlwatika and Bhat studying Rapanea melanophloes
(2002)
(L.)
Mez
and Cunonia These
capensis authors
in distinct ecological
L. sites.
may
suggest that the higher stomatal frequencies in sclerophyllous vegetation
be a reaction to the favorable photosynthetic conditions.
Rogas studying Alcbornea
et (1997, 2001), intraspecific variation in
al.
Mantuano
triplinervia (Spreng.) Miill. Arg., et (2006) verifying intraspecific
al.
variation in Erythroxylum ovalifolium Peyr., and Pereira et al. (2009) studying
variation of Andira legalis Toledo leaves, observed differences in
(Veil.)
epidermis thickness between the populations and related this to changes in
light regimes. In A. villosa populations analyzed are fully exposed to
all
which may why
sunlight, explain there no difference in their epidermis
is
thickness.
Although intraspecific variation in ferns has been poorly explored,
it is
crucial to understand the distribution of this group. According to Liu et
al.
(2006), the leaf anatomy of Isoetes shows less environmentally induced
variation than the external leaf morphology. In this present study, however,
it
AMERICAN FERN VOLUME NUMBER
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Root Root 2
vs.
1
was
under
verified that different environmental conditions Anemia
villosa
populations shov^ed both anatomical and morphological
modifications.
We
thank the Institute de Pesquisas Jardim Botanico do Rio de CAPES, CNPq
Janeiro, (Conselho
Nacional de Desenvolvimento and Museu
Cienti'fico e Tecnologico) Nacional-UFRJ
for fellowships,
We
funding and MSc
research
grants. are grateful to the Gabriel U. C. Araiijo dos Santos for the
English revision and to Carlos Eduardo Jascone for the aid in collecting plants. This paper was
derived from
the dissertation of the author.
first
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RIBEIRO ET INTRASPECIFIC VARIATION IN ANEMIA VILLOSA
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