Acta Botanica Brasilica 27(2): 436-444. 2013.
Pollen morphology of Menispermaceae in the state of Bahia, Brazil1
Michella Del Rei Teixeira2,5, André Márcio Amorim2,3,4 and Francisco de Assis Ribeiro dos Santos2
Submitted: 26 May, 2012. Accepted: 2 April, 2013
ABSTRACT
There have been few palynological studies of the family Menispermaceae. Here, we analyzed the pollen grains of
12 species from seven genera of Menispermaceae occurring in the state of Bahia, Brazil: Abuta selloana Eichl.; Borismene japurensis (Mart.) Eichl.; Chondrodendron microphyllum (Eichl.) Moldenke; C. platyphyllum (A.St.-Hil.) Miers;
Cissampelos andromorpha DC.; C. ovalifolia DC.; C. pareira L.; C. sympodialis Eichl.; Hyperbaena domingensis (DC.)
Benth.; Odontocarya duckei Barneby; Sciadotenia campestris Barneby; and S. pubistaminea (K. Schum.) Diels. The
pollen grains were acetolysed, measured, described, and illustrated using light and scanning electron microscopy. The
pollen grains were found to be small, oblate spheroidal or prolate spheroidal, isopolar, and tricolpate or tricolporate;
the exine was found to be reticulate. It was concluded that the various genera can be separated based on pollen characters. Within each of the genera, the species studied were stenopalynous. For five Menispermaceae species, pollen
descriptions are presented here for the first time.
Key words: Palynology, taxonomy, stenopalynous
Introduction
The family Menispermaceae Juss. comprises 71 genera
and approximately 500 species with largely pantropical
distributions (e.g., Judd et al. 2007; Barneby & White 2004;
Jacques et al. 2007). Only a few species occur in temperate
zones, between the eastern region of North America and
eastern Asia (Jacques et al. 2007). Sixteen genera and 110
species occur in Brazil, with most of the native species
occurring in the northern part of the country (Braga 2010).
Menispermaceae is widely known for its medicinal
and toxic components. The roots and bark of many plants
within this family (e.g., those within the genera Abuta,
Chondrodendron, Cissampelos and Sciadotenia) contain
alkaloids used as muscle relaxants in modern medicine
or as poisons on the tips of arrows utilized by indigenous
populations (Barneby 1975; Judd et al. 2007).
There have been few palynological studies of Menispermaceae. The revision published by Thanikaimoni (1968) presented a complete survey of the pollen morphology visible in
light microscopy of representatives of the tribes of this family.
Other authors have examined the pollen grains of the Menispermaceae and their relationships with the taxonomy of tribes
such as Coscinieae, Fibraureae, Menispermeae, and Triclisieae
(Ferguson 1975, 1978; Harley & Ferguson 1982; Harley 1985).
More recent palynological studies of Menispermaceae
have been published in pollen floras of specific geographical areas and have described the pollen grains of some
species of the tribes Anomospermeae, Menispermeae,
Tiliacoreae, and Tinosporeae. Salgado-Labouriau (1973)
described the pollen grains of Cissampelos ovalifolia DC.
as a contribution to the palynology of vegetation in the
Brazilian cerrado (savanna); Palacios-Chávez et al. (1991)
described the pollen grains of Cissampelos pareira L. from
the Sian Ka’an Biosphere Reserve in Mexico; Roubik &
Moreno (1991) provided palynological data concerning
four genera (Abuta Barrère ex Aubl., Chondrodendron
Ruiz & Pav., Cissampelos L., and Odontocarya Miers)
represented by seven species in Panama; and Carreira &
Barth (2003) studied the pollen flora of vegetation growing on iron-rich soils known as cangas (hardpan soils)
in Serra de Carajás, Brazil, describing the pollen grains
of Cissampelos andromorpha DC.
The present study had the principal objective of
analyzing the pollen grains of Menispermaceae species
occurring in the state of Bahia, Brazil, in order to describe
their morpho-palynological nature as well as to identify
characters that could have diagnostic value at the genus
or species level and contribute to taxonomic and related
studies of this family.
Based on the Master’s dissertation of the first Author
Universidade Estadual de Feira de Santana, Programa de Pós-Graduação em Botânica, Feira de Santana, Bahia, Brazil
3
Universidade Estadual de Santa Cruz, Departamento de Ciências Biológicas, Ilhéus, Bahia, Brazil
4
Centro de Pesquisas do Cacau, Comissão Executiva do Plano da Lavoura Cacaueira Herbário CEPEC, Ilhéus, Bahia, Brazil
5
Author for correspondence: michelladelrei@yahoo.com.br
1
2
Pollen morphology of Menispermaceae in the state of Bahia, Brazil
Materials and methods
We examined the pollen grains of 12 species of seven
genera of the family Menispermaceae occurring in the state
of Bahia, Brazil. Although an earlier survey had registered
15 species in the state (Barneby 1996), three species—Abuta
convexa (Vell.) Diels, Curarea crassa Barneby, and Orthomene schomburgkii (Miers) Barneby & Krukoff—could not
be included in the present study, either because of a lack of
fertile material or because of a limited number of staminate
inflorescences on the specimens in herbaria.
The study materials obtained from specimens deposited in the Herbarium of the Center for Cacao Research (code, CEPEC), Herbarium of the (Bahia) State
University of Feira de Santana (code, HUEFS), and the
Curitiba Municipal Botanical Museum (code, MBM) are
listed as follows: Abuta selloana Eichl. – BRAZIL. Bahia:
Valença, RPPN Fazenda Água Branca, P. Fiaschi et al.
2595, 30/X/ 2004 (CEPEC). Borismene japurensis (Mart.)
Barneby. – BRAZIL. Bahia: Belmonte, Estação experimental Gregório Bondar, T.S. Santos 4345, 29/XI/1987
(CEPEC). Chondrodendron microphyllum (Eichl.) Moldenke – BRAZIL. Bahia: Arataca, Fazenda IESB - Serra
das Lontras, M. Del-Rei et al. 44, 7/II/2009 (CEPEC);
Bahia: Arataca, Serra Peito-de-Moça, W.W. Thomas et al.
14530, 18/I/2006 (CEPEC); Bahia: Santa Terezinha, Serra
da Jibóia, M.L.C. Neves 39, 1/IV/2004 (HUEFS). Chondrodendron platiphyllum (A. St.-Hil.) Miers – BRAZIL.
Bahia: Porto Seguro, BR 101, Montinho, T.S. Santos 1646,
15/V/1971 (CEPEC); Bahia: São Sebastião do Passé, Mata
da Mariquita, M. Del-Rei et al. 61, 27/IX/2009 (HUEFS). Cissampelos andromorpha DC. – BRAZIL. Bahia:
Ilhéus, área do CEPEC, J. L. Hage et al. 1408, 6/X/1981
(CEPEC); Bahia: Itacaré, 15,5 km da BR-101 via Itacaré, J.G. Jardim et al. 1136, 17/X/1997 (CEPEC); Bahia:
Santa Cruz Cabrália, Reserva Biológica Pau-brasil, A.
Eupunino 15, 1/X/1971 (CEPEC). Cissampelos ovalifolia
DC. – BRAZIL. Bahia: Feira de Santana, 16/VI/2009,
M. Del-Rei 58 (HUEFS); Bahia: Piatã, 16/X/1992, W.
Ganev 1236 (HUEFS); Bahia: Barreiras, 2/XI/1987, L.P.
Queiroz et al. 2095 (HUEFS). Cissampelos pareira L. –
BRAZIL. Bahia: Coribe, 8/VI/2007, M.M.M. Lopes et al.
1374 (CEPEC); Bahia: Feira de Santana, 16/VII/2009, M.
Del-Rei 55 (HUEFS); Bahia: Feira de Santana, 9/XI/2004,
S.F. Conceição et al. 86 (HUEFS). Cissampelos sympodialis
Eichl. – BRAZIL. Bahia: Caldeirão Grande, 17/XI/1986,
L.P. Queiroz 1166 (HUEFS); Bahia: Jacobina, 12/X/2007,
J.L. Ferreira 124 (HUEFS); Bahia: Morro do Chapéu, 8/
VI/2001, E.R. Souza et al. 110 (HUEFS). Curarea crassa
Barneby – BRAZIL. Bahia: Jaguaquara, 24/IV/2002,
R.P. Oliveira et al. 786 (HUEFS); Bahia: Uruçuca, 12/
IV/1995, W.W. Thomas et al. 10900 (CEPEC). Hyperbaena domingensis (DC.) Benth. – BRAZIL. Bahia: sem
localidade, BR-101, Vale do Rio Mucuri, 2/VI/1971,
T.S. Santos 1538 (CEPEC); Bahia: Teixeira de Freitas,
Acta bot. bras. 27(2): 436-444. 2013.
11/X/1971, T.S. Santos et al. 2106 (CEPEC). Odontocarya
duckei Barneby – BRAZIL. Bahia: Catolés, 4/II/2003,
F. França et al. 4272 (HUEFS); Bahia: Rio de Contas,
3/XI/1988, R.M. Harley et al. 25898 (CEPEC). Sciadotenia campestris Barneby – BRAZIL. Bahia: Barreiras,
20/VIII/1986, G. Hatschbach & J.M. Silva 50530 (holótipo
MBM); Bahia: Correntina, 16/V/2000, G. Hatschbach et
al. 71238 (MBM). Sciadotenia pubistaminea (K. Schum.)
Diels – BRAZIL. Bahia: Encruzilhada, 17/VIII/2001,
A.M. Carvalho et al. 6981 (CEPEC); Bahia: Vitória da
Conquista, 22/III/1996, W.W. Thomas 11098 (CEPEC).
The pollen samples were prepared for light microscopy
using the acetolysis method devised by Erdtman (1960).
Slides were mounted using glycerin jelly. The measurements
of polar and equatorial diameters were taken from 25 randomly pollen grains (whenever possible) in equatorial view;
exine thicknesses were determined with 10 measurements.
The polar/equatorial diameter ratio was used to define the
shapes of the grains. The slides were subsequently incorporated into the palynological archives of the Plant Micromorphology Laboratory (LAMIV) at the State University
of Feira de Santana.
For scanning electron microscopy (SEM) analyses, non-acetolysed pollen grains were scattered directly onto metallic supports and then coated with a thin layer of palladium
gold (for approximately 3 min) in a sputter-coater (SCD050; Bal-Tec/Leica Microsystems, Bannockburn, IL, USA).
The palynological terminology adopted follows Punt
et al. (2007). For pollen size classes, we followed Erdtman
(1952).
Photomicrographs were taken using an Olympus digital
camera (Olympus, Tokyo, Brazil) mounted to a microscope
(Axiostar Plus; Carl Zeiss, Oberkochen, Germany) with a
100× objective. The electron micrographs were taken using
a scanning electron microscope (LEO 1430 VP; Carl Zeiss
do Brasil, São Paulo, Brazil).
Results
The descriptions of pollen of Menispermaceae genera
were based on species occurring in the state of Bahia, as
listed below and detailed in Tab. 1.
1. Abuta Barrère ex Aubl.
Fig. 1-3.
Pollen grains in monads, small, prolate spheroidal,
isopolar, amb circular, tricolpate, polar area large; colpus
approximately 2/3 the length of the polar axis, apertural
membrane ornamented with microspines; exine microreticulate to microechinate, irregular muri surface with sparse
microspines (Fig. 1 and 3) that are visible only by SEM and
difficult to measure, lumens poorly visible, heterogeneous;
sexine the same thickness as nexine. Species examined:
Abuta selloana Eichl.
437
Michella Del Rei Teixeira, André Márcio Amorim and Francisco de Assis Ribeiro dos Santos
Table 1. Measurements of the pollen grains of Menispermaceae species in the state of Bahia, Brazil.
Species/Collector (Herbarium)
Polar diameter
(μm)
Equatorial diameter
(μm)
Shape
P/E
Exine
Sexine/ AS
thickness
PAI
Nexine (μm)
(μm)
Min.
(x–)
Max.
Min.
(x–)
Max.
13.0
(13.5)
14.0
12.0
(13.0)
14.0
prolate spheroidal
1.04
1.5
S=N
7
0.54
13.0
(14.0)
15.0
13.0
(14.5)
16.0
oblate spheroidal
0.97
1
S>N
5
0.34
M. Del-Rei et al. 44 (CEPEC)
14.0
(15.0)
16.0
14.0
(15.0)
16.0
spheroidal
1
2
S>N
5
0.32
M.L.C. Neves et al. 39 (HUEFS)
16.0
(17.0)
18.0
15.0
(16.0)
17.0
prolate spheroidal
1.06
2
S>N
--
--
W.W. Thomas et al. 14530 (CEPEC)
15.0
(16.0)
17.0
14.0
(15.0)
16.0
prolate spheroidal
1.07
2
S>N
--
--
M. Del-Rei et al. 61 (HUEFS)
14.0
(16.0)
18.0
13.0
(14.5)
16.0
prolate spheroidal
1.1
2
S>N
5
0.36
T.S. dos Santos et al. 1646 (CEPEC)
14.0
(15.5)
17.0
12.0
(13.5)
15.0
prolate spheroidal
1.15
1
S>N
--
--
15.0
(16.0)
17.0
15.0
(16.0)
17.0
spheroidal
1
2
S=N
6
0.38
Abuta selloana (Velloso) Diels
P. Fiaschi et al. 2595 (CEPEC)
Borismene japurensis (Mart.) Eichler
T.S. Santos et al. 4345 (CEPEC)
Chondrodendron microphyllum (Eichler)
Moldenke
Chondrodendron platyphyllum (A. St.-Hil.) Miers
Cissampelos andromorpha DC.
A. Eupunino 15 (CEPEC)
J.G. Jardim et al. 1136 (CEPEC)
15.0
(15.5)
16.0
15.0
(16.0)
17.0
oblate spheroidal
0.97
2
S=N
--
--
J.L. Hage et al. 1408 (CEPEC)
15.0
(16.0)
17.0
15.0
(16.0)
17.0
oblate spheroidal
1
2
S=N
--
--
M. Del-Rei 58 (HUEFS)
16.0
(18.0)
20.0
18.0
(19.0)
20.0
oblate spheroidal
0.95
2
S=N
7
0.39
W. Ganev et al. 1236 (HUEFS)
16.0
(17.0)
18.0
16.0
(17.0)
18.0
spheroidal
1
2
S=N
--
--
L.P. Queiroz et al. 2095 (HUEFS)
16.0
(17.5)
19.0
16.0
(18.0)
20.0
oblate spheroidal
0.97
2
S=N
--
--
M.M.M. Lopes 1374 (CEPEC)
17.0
(18.0)
19.0
19.0
(20.0)
21.0
oblate spheroidal
0.9
2
S=N
5
0.25
M. Del-Rei 55 (HUEFS)
14.0
(15.0)
16.0
14.0
(15.5)
17.0
oblate spheroidal
0.97
2
S>N
5
0.31
S.F. Conceição et al. 86 (HUEFS)
14.0
(15.5)
17.0
15.0
(16.5)
18.0
oblate spheroidal
0.94
2
S>N
--
--
12.0
(13.5)
15.0
12.0
(13.5)
15.0
spheroidal
1
2
S>N
6
0.41
Cissampelos ovalifolia DC.
Cissampelos pareira L.
Cissampelos sympodialis Eichl.
J.L. Ferreira et al. 124 (HUEFS)
E.R. Souza et al. 110 (HUEFS)
14.0
(15.5)
17.0
14.0
(15.5)
17.0
spheroidal
1
2
S>N
--
--
L.P. Queiroz et al. 1166 (HUEFS)
12.0
(14.0)
16.0
12.0
(14.0)
16.0
spheroidal
1
2
S>N
--
--
T.S. Santos 1538 (CEPEC)
14.0
(15.0)
16.0
13.0
(13.5)
14.0
prolate spheroidal
1.11
1
S=N
6
0.41
T.S. Santos 2106 (CEPEC)
13.0
(15.0)
17.0
13.0
(14.5)
16.0
prolate spheroidal
1.03
1
S=N
--
--
2
0.15
Hyperbaena domingensis (DC.) Benth.
Odontocarya duckei Barneby
F. França et al. PCD 1296 (CEPEC)
13.0
(14.0)
15.0
13.0
(14.0)
15.0
spheroidal
1
1
S>N
R.M. Harley 25898 (HUEFS)
13.0
(14.5)
16.0
12.0
(13.5)
15.0
prolate spheroidal
1.07
1
S>N
Sciadotenia campestris Barneby
G.M. Hatschbach et al. 50530 (MBM)
12.0
(13.0)
14.0
12.0
(13.0)
14.0
spheroidal
1
1
S=N
6
0.43
G.M. Hatschbach et al. 71238 (MBM)
12.0
(14.0)
16.0
13.0
(14.5)
16.0
oblate spheroidal
0.97
2
S=N
--
--
W.W. Thomas et al. 11098 (CEPEC)
13.0
(14.0)
15.0
12.0
(13.0)
14.0
prolate spheroidal
1.08
1
S=N
7
0.5
A.M. Carvalho et al. 6981 (CEPEC)
14.0
(15.0)
16.0
13.0
(14.5)
16.0
prolate spheroidal
1.03
1
S=N
--
--
Sciadotenia pubistaminea (K. Schum.) Diels
Min. – minimum; x– – arithmetic mean; Max. – maximum; P/E – polar/equatorial (diameter) ratio; AS – apocolpium side; PAI – polar area index; CEPEC – Herbarium of the Center for Cacao Research; HUEFS – Herbarium of the (Bahia) State University of Feira de Santana; MBM – Curitiba Municipal Botanical Museum.
438
Acta bot. bras. 27(2): 436-444. 2013.
Pollen morphology of Menispermaceae in the state of Bahia, Brazil
2. Borismene Barneby
Fig. 4-6.
Pollen grains in monads, small, oblate spheroidal, isopolar, amb circular, tricolporate, polar area small; ectoaperture
approximately 2/3-3/4 the length of the polar axis, aperture
membrane strongly granulate (Fig. 6), sometimes covering
the circular endoaperture, making it difficult to observe; exine
microreticulate, muri surface smooth, lumen small, heterogeneous, smaller near the apertures; sexine thicker than nexine.
Species examined: Borismene japurensis (Mart.) Eichler.
3. Chondrodendron Ruiz & Pav.
Fig. 7-13
Pollen grains in monads, small, spheroidal to prolate
spheroidal in C. microphyllum and prolate spheroidal in
C. platyphyllum, isopolar, amb subcircular, tricolporate,
polar area small; ectoapertures approximately 2/3 the
length of the polar axis, narrow; endoapertures lolongate
and difficult to see; exine microreticulate, muri surface
irregular, with transversal constrictions (small folds) on
the surface and scabrae that are difficult to see (Fig. 10),
lumen small, polygonal, heterogeneous; sexine thicker
than nexine. Species examined: C. microphyllum (Eichl.)
Moldenke (Fig. 7-9) and C. platyphyllum (A. St.-Hil.)
Miers (Fig. 10-13).
4. Cissampelos L.
Fig. 14-25
Pollen grains in monads, small, oblate spheroidal in C.
pareira, oblate spheroidal to spheroidal in C. andromorpha
and C. ovalifolia, spheroidal in C. sympodialis, isopolar,
amb subcircular, tricolporate, polar area small; ectoapertures approximately 2/3-3/4 the length of the polar axis;
opercula narrow and long over the ectoapertures (Fig. 18
and 23), smaller in C. ovalifolia than in the other species;
endoapertures lalongate and difficult to see; exine reticulate,
muri surface smooth, lumen large, heterogeneous; sexine
twice as thick as nexine. Species examined: C. andromorpha
DC.(Fig. 14-16), C. ovalifolia DC. (Fig. 16-17), C. pareira L.
(Fig. 18-22), and C. sympodialis Eichl. (Fig. 23-25).
5. Hyperbaena Miers ex Benth.
Fig. 26-28
Pollen grains in monads, small, prolate-spheroidal, isopolar,
amb sub-circular, tricolporate, polar area small; ectoaperture
approximately 1/2-2/3 the length of the polar axis; endoaperture circular; exine with cerebroid reticulate ornamentation (Fig.
26), muri thick and sinuous, surface smooth, lumens very small
to large, heterogeneous; sexine the same thickness as nexine.
Species examined: Hyperbaena domingensis (DC.) Benth.
6. Odontocarya Miers
Fig. 29-31
Pollen grains in monads, small, spheroidal to prolate
spheroidal, isopolar, amb circular to subcircular, tricolporate, polar area very small; ectoaperture more than 3/4
the length of the polar axis, sometimes fused at both poles
(syncolporate, Fig. 29); opercula narrow and long over the
ectoapertures (Fig. 29 and 31); endoaperture lolongate,
diffuse, and difficult to see. Exine reticulate, with thin and
slightly sinuous muri but with variable thicknesses, surface
undulate, lumens heterogeneous; sexine twice as thick as
nexine. Species examined: Odontocarya duckei Barneby.
7. Sciadotenia Miers.
Fig. 32-35
Pollen grains in monads, small, spheroidal in S. campestris and prolate spheroidal in S. pubistaminea, isopolar, amb
circular, tricolpate, polar area small; colpi approximately
2/3-3/4 the length of the polar axis; exine microreticulate,
with thick and sinuous walls (Fig. 33), muri surface smooth, lumen of variable sizes, heterogeneous; sexine twice as
thick as nexine. Species examined: S. campestris Barneby
(Fig. 32-34) and S. pubistaminea (K.Schum.) Diels (Fig. 35).
Pollen key for separating the genera of Menispermaceae occurring in the state of Bahia:
1. Pollen grains operculate
2. Pollen grains ≥ 15 μm, exine thickness 2 μm .............................................................................................. 4. Cissampelos
2’. Pollen grains < 15 μm, exine thickness 1 μm ............................................................................................ 6. Odontocarya
1’. Pollen grains not operculate
3. Pollen grains colpate
4. Exine with microspines ....................................................................................................................................... 1. Abuta
4’. Exine without microspines ........................................................................................................................ 7. Sciadotenia
3’. Pollen grains colporate
5. Muri with transversal constrictions ............................................................................................... 3. Chondrodendron
5’. Muri without transversal constrictions, smooth
6. Exine with cerebroid reticulate ornamentation ............................................................................... 5. Hyperbaena
6’. Exine with microreticulate ornamentation ......................................................................................... 2. Borismene
Acta bot. bras. 27(2): 436-444. 2013.
439
Michella Del Rei Teixeira, André Márcio Amorim and Francisco de Assis Ribeiro dos Santos
Figures 1-13. Pollen grains of Menispermaceae in the state of Bahia, Brazil, under light microscopy and scanning electron microscopy (SEM). Fig. 1-3.
Abuta selloana Eichl.: 1. Polar view (SEM); 2. Polar view; 3. Equatorial view, colpus and surface (SEM)—Fig. 4-6. Borismene japurensis (Mart.) Eichl.: 4.
Polar view (SEM); 5. Polar view; 6. Equatorial view (SEM). 7-9. Chondrodendron microphyllum (Eichl.) Moldenke: 7. Equatorial view, colpus and surface
(SEM); 8, 9. Polar view—Fig. 10-13. Chondrodendron platyphyllum (A. St.-Hil.) Miers: 10. Equatorial view (SEM); 11. Polar view (SEM); 12. Polar view; 13.
Polar view, surface. (Scale bar = 10 μm, Fig. 2, 5, 8, 9, 12).
440
Acta bot. bras. 27(2): 436-444. 2013.
Pollen morphology of Menispermaceae in the state of Bahia, Brazil
Figures 14-25. Pollen grains of Menispermaceae in the state of Bahia, Brazil, under light microscopy and scanning electron microscopy (SEM). Fig. 14,
15. Cissampelos andromorpha DC.: 14. Polar view, surface (SEM); 15. Polar view—Fig. 16, 17. Cissampelos ovalifolia DC.: 16. Equatorial view, colpus and
surface (SEM); 17. Polar view—Fig. 18-22. Cissampelos pareira L.: 18. Equatorial view, colpus and surface (SEM); 19. Polar view, surface; 20. Polar view; 21.
Equatorial view, colpus and surface (SEM); 22. Polar view—Fig. 23-25. Cissampelos sympodialis Eichl.: 23. Equatorial view, colpus (SEM); 24. Polar view;
25. Polar view, surface. (Scale bar = 10 μm, Fig. 15, 17, 19, 20, 22-24).
Acta bot. bras. 27(2): 436-444. 2013.
441
Michella Del Rei Teixeira, André Márcio Amorim and Francisco de Assis Ribeiro dos Santos
Figures 26-35. Pollen grains de Menispermaceae in the state of Bahia, Brazil, under light microscopy and scanning electron microscopy (SEM). Fig. 26-28.
Hyperbaena domingensis (DC.) Benth.: 26. Equatorial view (SEM); 27. Polar view; 28. Equatorial view, surface (SEM)—Fig. 29-31. Odontocarya duckei Barneby:
29. Polar view (SEM). 30. Polar view. 31. Equatorial view, colpus and surface (SEM)—Fig. 32-34. Sciadotenia campestris Barneby: 32. Equatorial view (SEM);
33. Equatorial view, surface (SEM); 34. Polar view—Fig. 35. Sciadotenia pubistaminea (K. Schum.) Diels: Polar view. (Scale bar = 10 μm, Fig. 27, 30, 34, 35).
Discussion
The seven genera of Menispermaceae occurring in the
state of Bahia were studied and the characteristics of the 13
species analyzed corroborated the findings of classical studies (Erdtman 1952; Thanikaimoni 1968; Ferguson 1975).
For five species, the present study represents the first time
that their pollen grains have been described: Abuta selloana,
Chondrodendron microphyllum, Cissampelos sympodialis,
Odontocarya duckei, and Sciadotenia campestris.
442
The pollen grains of species within the same genus were
found to be stenopalynous, showing only small morphological differences. The genera, however, are more palynologically distinct, and it is possible to identify them according
to their pollen characters.
All of the pollen grains studied are dispersed in monads
and are isopolar, varying in shape from oblate to prolate-spheroidal. The Cissampelos species analyzed here vary
from oblate spheroidal to spheroidal. The Chondrodendron
species have pollen grains that are predominantly prolate
Acta bot. bras. 27(2): 436-444. 2013.
Pollen morphology of Menispermaceae in the state of Bahia, Brazil
spheroidal, except for one specimen of C. microphyllum
that presents spheroidal pollen grains. The pollen grains
of the Sciadotenia species vary from oblate-spheroidal (S.
campestris) to prolate-spheroidal (S. pubistaminea). All of
the species have small pollen grains, varying from 13 μm in
Sciadotenia campestris (G.M. Hatschbach et al. 50530 ) to 20
μm in Cissampelos pareira (M.M.M. Lopes 1374).
The pollen grains of the genera Abuta and Sciadotenia
are predominantly colpate, whereas those of Borismene,
Chondrodendron, Cissampelos, Hyperbaena, and Odontocarya are colporate. Colpi lengths vary from 2/3 to 3/4 of the
length of the polar axis. The pollen grains of Odontocarya
duckei have longer colpi, which were often fused at the
poles (syncolporate). Most of the colporate pollen grains,
mainly those with opercula, had endoapertures that were
quite difficult to see.
Most of the species have microreticulate sexine with
smooth surfaces; transversal constrictions on the muri
surface were observed only in Chondrodendron species.
Microspines were observed on the surface of the pollen
grains of Abuta selloana.
The proportions of sexine and nexine thicknesses in the
pollen wall varied among the genera studied, although the
sexine are generally observed to be thicker than the nexine.
Nexine and sexine thicknesses are similar only among the
genera Abuta, Hyperbaena, and Sciadotenia.
The prolate spheroidal shape attributed to the pollen
grains of Cissampelos ovalifolia by Salgado-Labouriau (1973)
diverged from the results of the present study, although
interpretations of pollen shapes are often influenced by the
laboratory techniques used.
For many years, a tricolpate aperture was thought to
represent a primitive characteristic in this family, with
tricolpate, triporate and inaperturate pollen grains being
considered derived characters (Thanikaimoni 1986). However, within the phylogenetic context of the molecular data
presented by Ortiz et al. (2007) and Hoot et al. (2009), the
pollen grains were found to be extremely homoplastic—the
basal condition of the family is probably triaperturate-tricolpate or triporate (Hoot et al. 2009).
The small size of the sample examined here prevented
us from establishing an evolutionary hypothesis related to
the pollen characters of the Menispermaceae. However,
our results, contextualized within those of the molecular
phylogenetic studies conducted by Ortiz et al. (2007) and
Hoot et al. (2009), indicate that the apomorphic condition
of the colporate pollen type with a circular endoaperture
(shared by Borismene and Hyperbaena) and the presence
of an operculum (shared by Odontocarya and Cissampelos)
are homoplastic aspects and have arisen several times in the
evolutionary history of this family.
Based on data presented here, we can conclude that
species within each genus of Menispermaceae demonstrate
significant morphological homogeneity in terms of their
pollen characters, principally in terms of the number, type,
Acta bot. bras. 27(2): 436-444. 2013.
and ornamentation of their apertures. However, major differences were noted between the genera in terms of characters
such as aperture type, size, and exine ornamentation as
observed under SEM.
Despite the similarity of the pollen grains (suggesting
close relationships) among the species of the various genera
analyzed here, these taxa are otherwise quite distinct in
terms of their vegetative and reproductive morphologies.
Although the analysis of pollen morphology can be useful
in taxonomic studies of this group, it should be considered
only as an additional tool for species classification.
Acknowledgments
The authors would like to thank the staff of the Plant
Micromorphology Laboratory at the Universidade Estadual de Feira de Santana (UEFS, [Bahia] State University of
Feira de Santana) especially Dr. Paulino Pereira Oliveira,
for providing access to the laboratory facilities; the staff
of the UEFS Electron Microscopy Laboratory, especially
the technician Gisele Rocha for her help in preparing the
electron micrographs; and the staff of the Animal Histology Laboratory at the Universidade Estadual de Santa Cruz
(UESC, Santa Cruz State University [of Bahia]), especially
Prof. Guisla Boehs for her help in preparing the light photomicrographs. This study received financial support in the
form of research grants from the Brazilian Coordenação
de Aperfeiçoamento de Pessoal de Nível Superior (CAPES,
Office for the Advancement of Higher Education; master’s
scholarship grant to MDRT) and Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq, National
Council for Scientific and Technological Development;
productivity grants to AMA and FARS).
References
Barneby, R.C. 1975. Menispermaceae. Pp. 3-40. In: Reitz R. (Ed.). Flora
Ilustrada Catarinense. Itajaí, Herbário Barbosa Rodrigues.
Barneby, R.C. 1996. Tiding of Menispermaceae from interior French
Guiana and from the Brazilian State of Bahia. Brittonia 48(1): 20-25.
Barneby, R.C.; White, P. 2004. Menispermaceae (Moonseed Family). Pp.
247-249. In: Smith, N.; Mori, S.A.; Henderson, A.; Stevenson, D.W.
& Heald, S.V. (Eds). Flowering Plants of the Neotropics. Princeton,
Princeton University Press.
Braga, J.M.A. 2010. Menispermaceae. In: Forzza, R.C.; Leitman, P.M.;
Costa, A.F.; Carvalho Jr., A.A.; Peixoto, A.L.; Walter, B.M.T.; Bicudo, C.;
Zappi, D.; Costa, D.P.; Lleras, E.; Martinelli, G.; Lima, H.C.; Prado, J.;
Stehmann, J.R.; Baumgratz, J.F.A.; Pirani, J.R.; Sylvestre, L.; Maia, L.C.;
Lohmann, L.G.; Queiroz, L.P.; Silveira, M.; Coelho, M.N.; Mamede,
M.C.; Bastos, M.N.C.; Morim, M.N.C.; Barbosa, M.R.; Menezes, M.;
Hopkins, M.; Secco, R.; Cavalcanti, T.B. & Souza, V.C. (Orgs). Lista
de Espécies da Flora do Brasil. http://floradobrasil.jbrj.gov.br/2010/
FB000163. Acesso em 22/09/2010.
Carreira, L.M.M. & Barth, O.M. 2003. Atlas de pólen da vegetação de
canga da Serra de Carajás. Belém, Museu Paraense Emílio Goeldi.
Erdtman, G. 1952. Pollen Morphology and Plant Taxonomy Angiosperms. Stockholm, Almquist & Wiksell.
Erdtman, G. 1960. The acetolysis method. A revised description. Svensk
Botanisk Tidskrft 39: 561-564.
443
Michella Del Rei Teixeira, André Márcio Amorim and Francisco de Assis Ribeiro dos Santos
Ferguson, I.K. 1975. Pollen Morphology of the Tribe Triclisieae of the Menispermaceae in relation to its taxonomy. Kew Bulletin 30(1): 49-75.
Ferguson, I.K. 1978. Pollen Morphology of the Tribe Coscinieae of the
Menispermaceae in relation to its taxonomy. Kew Bulletin 32(2):
339-346.
Harley, M.M. 1985. Pollen morphology and Taxonomy of the Tribe Fibraureae (Menispermaceae). Kew Bulletin 40(3): 553-565.
Harley, M.M. & Ferguson, I.K. 1982. Pollen Morphology and Taxonomy
of the Tribe Menispermeae (Menispermaceae). Kew Bulletin 37(3):
353-366.
Hoot, S.B.; Zautke, H.; Harris, D.J.; Crane P.R. & Neves S.S. 2009. Phylogenetic Patterns in Menispermaceae Based on Multiple Chloroplast
Sequence Data. Systematic Botany 34(1): 44-56.
Jacques, F.M.B.; Gallut, C.; Vignes-Lebbe, V. & Bagils, R.Z. 2007. Resolving
phylogenetic reconstruction in Menispermaceae (Ranunculales) using
fossils and a novel statistical test. Taxon 56(2): 379-392.
Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F. & Donoghue, M.J.
2007. Plant systematics: a phylogenetic approach. 2nd ed. Sunderland, Sinauer Associates.
Ortiz, R.; Kellogg, E.A. & van der Werff, H. 2007. Molecular phylogeny of
the moonseed family (Menispermaceae): implications for morphological diversification. American Journal of Botany 94: 1425-1438.
Palacios-Chávez, R.; Ludlow-Wiechers, B. & Villanueva, R.G. 1991. Flora
palinológica de la Reserva de La Biósfera de Sian Ka’an, Quintana
Roo, Mexico. Chetumal, Centro de Investigaciones de Quintana Roo.
Punt, W.; Hoen, P.P.; Blackmore, S.; Nilsson, S. & Le Thomas, A. 2007.
Glossary of pollen and spores terminology. Review of Paleobotany
and Palynology 143: 1-81.
Roubik, D.W. & Moreno P., J.E. 1991. Pollen and spores of Barro Colorado
Island. Monographs in Systematic Botany 36: 1-270.
Salgado-Labouriau, M.L. 1973. Contribuição à palinologia dos cerrados.
Rio de Janeiro, Academia Brasileria de Ciências.
Thanikaimoni, G. 1968. Morphologie des pollens des Menispermacées.
Institut Français de Pondichéry. Travaux de la Section Scientifique
et Technique 5: 1-56.
Thanikaimoni, G. 1986. Evolution of Menispermaceae. Canadian Journal
of Botany 64: 3130-3133.
Online version: www.scielo.br/abb and http://www.botanica.org.br/acta/ojs
444
Acta bot. bras. 27(2): 436-444. 2013.