Revista Agrogeoambiental Espécies do cerrado com potencial ...
Reproductive strategies in Cerrado plants: outcrossing vs ... · Reproductive strategies in Cerrado...
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Paulo Eugênio Oliveira
Universidade Federal de Uberlândia - MG
Reproductive strategies in
Cerrado plants: outcrossing vs.
apomixis and the organization of
plant-pollinator interaction webs
in Central Brazil
Felipe Ribeiro
Cerrado:Neotropical savanna
areas in Central Brazil
Cerrado Physiognomies
Gallery
forestCampos Cerrado Cerradão
Deforestation and
land use in Cerrado
Klink & Moreira 2002
Cerrado as a conservation hot spot
Native areas44,53%
Cultivatedpastureland41,56%
others
0,59%
Urban areas1,90%
Forestry
0,07%
Agriculture
11,35%
Mendonça e at (1998), Ratter et al (2003), Felfili et al (2001), Sano et al. (2008)
376sites
Ratter et al, 2003
Bridgewater et al. 2004
Floristic basis
• Woody species = 953 spp
• Herbs/woody ratio= 3-4/1
• No species occurred in all sites
• Gallery forest and riverine woody species = 1 900 spp
• Total = > 12000 spp
High β diversity and
species turn-over
Cerrado Determinants
Nutrient
poor soils
FireSeasonal
climate
Woody species with
deep roots
Herbaceous plants
with shallow root
system
Water use and
seasonality
Accumulated biomass and fire
Fire
Traditional ideas suggested fire and environmental
restrictions would render sexual reproduction of woody
species very difficult and would favor asexual reproduction
Gallery
forestCampos Cerrado Cerradão
Gallery
forestCampos Cerrado Cerradão
Malvaceae (Gottsberger 1986)
Malvaceae (Gottsberger 1986)
Woody hummingbirds and
bat pollinated species with
specialized flowersHerbaceous
bee pollinated
promiscuous groups
Diversity of pollination systems
Oliveira & Gibbs 2000
Cerrado Dry Secondary Cloudy Dry Lowland
forest forest forest forest rainforest
BSB Costa Rica Venezuela Venezuela Mexico Costa Rica
Dioecy 15 22 24 31 20 23
Monoecy 5 10 0 3 10 11
Self-incompat. 66 54 64 26 53 53
Autogamous 14 14 12 43 17 13
Outcrossing 81 76 88 57 73 75
Breeding systems of woody species in Netropical areas
Oliveira & Gibbs 2000
Polinizadores, vicariancia, fenologia e processo de especiaçãoV. pumila
V. pyramidalis
V. rufa
V. thyrsoidea
V. tucanorum
V. elliptica
Oliveira & Gibbs 1994
Polinizadores, vicariancia, fenologia e processo de especiaçãoV. pumila
V. pyramidalis
V. rufa
V. thyrsoidea
V. tucanorum
V. elliptica
Oliveira & Gibbs 1994
Distribution of Vochysia (Vochysiaceae)
in Cerrado plant formations.
- self-incompatible
- dependent on large bees for fruit set.
- guild of plants (including other groups) and large bees
Sphingids Plants
Guilds and interaction web
Amorim 2008
Average flower lenght (cm)
Av
era
ge
pro
bo
scis
len
gh
t (c
m)
Blind connections: association between
hawkmoths and plants in Cerrado.
Amorim 2008
1,25 cm
1,25 cm1,75cm
1,75cm
Xylocopa (Neoxylocopa) suspecta (Moure & Camargo)
Xylocopa (Neoxylocopa) frontalis (Olivier)
Xylocopa (Neoxylocopa) grisescens Lepeletier
Clá
udia
Inês
da
Silv
a
Cláudia Inês Silva
Pollen load on Xylocopa bees
Xylocopa frontalis
Xylocopa suspecta
Xylocopa grisescens
Xylocopa hirsutissima
30 plant spp. intensely used
Cláudia Inês Silva
Xylocopa frontalis
Xylocopa suspecta
Xylocopa grisecens
Xylocopa hirsutissima
Floral visitors on Matayba
guianensis (Sapindaceae)
in Cerrados of Uberlândia
Ana Coelho Carvalho,
Unpublished data
Wasps
50
Beetles
37
Bees
110
!
Guilds, redundancy and resilience
Obligatory outcrossing: dependency on pollinators.
Specialized relationships are
rare. Generalist guilds?
Redundancy and stability?
Guilds and resilience after
disturbance.
Adjust to high β-diversity?
Importance
of apomixisMelastomataceae
Miconieae39%
63%
Eriotheca pubescens
Goldenberg & Shepherd 1998Oliveira et al. 1992
Martins & Oliveira 2003
Mendes-Rodrigues et al. 2005
Grupos EspecíficosSpecific groups
Polyembryony
and apomixis
Salomão & Allem 2001
Allem 2003
Apomixis in Cerrado
Questions:
How common is polyembryony among cerrado plants?
Is polyembryony a diagnostic feature to estimate
importance of apomixis in Cerrado plants?
How widespread are polyembryony and apomixis in
Cerrado environment?
What is the origin of apomictic taxa?
Consequences and perspectives?
Apomixis in Cerrado plantsCNPq research grant (2005-2007)
Family Species Method Mean SD N SE Min. Max Mono Poly %M %P
Anacardiaceae Astronium fraxinifolium Embryos 1,00 0,00 253,00 0,00 1,00 1,00 253,00 0,00 100,00 0,00
Anacardiaceae Astronium fraxinifolium Seedlings 1,01 0,08 697,00 0,00 1,00 2,00 692,00 5,00 99,28 0,72
Anacardiaceae Lithraeae molleoides Seedlings 1,00 0,00 1.335,00 0,00 1,00 1,00 1.335,00 0,00 100,00 0,00
Anacardiaceae Myracrodruon urundeuva Embryos 1,01 0,11 250,00 0,01 1,00 2,00 247,00 3,00 98,80 1,20
Anacardiaceae Myracrodruon urundeuva Seedlings 1,01 0,10 486,00 0,00 1,00 2,00 481,00 5,00 98,97 1,03
Apocynaceae Aspidosperma cylindrocarpum Seedlings 1,00 0,00 221,00 0,00 1,00 1,00 221,00 0,00 100,00 0,00
Bignoniaceae Jacaranda cuspidifolia Seedlings 1,01 0,11 404,00 0,01 1,00 3,00 402,00 2,00 99,50 0,50
Bignoniaceae Tabebuia avellanedae Seedlings 1,00 0,00 346,00 0,00 1,00 1,00 346,00 0,00 100,00 0,00
Bignoniaceae Tabebuia caraiba Seedlings 1,00 0,00 132,00 0,00 1,00 1,00 132,00 0,00 100,00 0,00
Bignoniaceae Tabebuia chysotricha Embryos 2,18 1,12 606,00 0,05 1,00 7,00 192,00 414,00 31,68 68,32
Bignoniaceae Tabebuia chysotricha Seedlings 1,66 0,74 563,00 0,03 1,00 5,00 270,00 293,00 47,96 52,04
Bignoniaceae Tabebuia ochracea Seedlings 1,36 0,48 45,00 0,07 1,00 2,00 29,00 16,00 64,44 35,56
Bignoniaceae Tabebuia roseo-alba Seedlings 1,01 0,08 676,00 0,00 1,00 2,00 672,00 4,00 99,41 0,59
Bignoniaceae Tabebuia serratifolia Embryos 1,26 0,57 46,00 0,08 1,00 4,00 36,00 10,00 78,26 21,74
Bignoniaceae Tabebuia serratifolia Seedlings 1,18 0,38 200,00 0,03 1,00 2,00 165,00 35,00 82,50 17,50
Caesalpiniaceae Copaifera langsdorfii Seedlings 1,00 0,04 577,00 0,00 1,00 2,00 576,00 1,00 99,83 0,17
Caesalpiniaceae Hymenaea courbaril Seedlings 1,00 0,03 1.173,00 0,00 1,00 2,00 1.172,00 1,00 99,91 0,09
Caesalpiniaceae Senna sylvestris Seedlings 1,00 0,05 346,00 0,00 1,00 2,00 345,00 1,00 99,71 0,29
Clusiaceae Kielmeyera coriaceae Embryos 1,00 0,00 62,00 0,00 1,00 1,00 62,00 0,00 100,00 0,00
Clusiaceae Kielmeyera grandiflora Embryos 1,00 0,00 200,00 0,00 1,00 1,00 200,00 0,00 100,00 0,00
Clusiaceae Kielmeyera variabilis Embryos 1,00 0,00 50,00 0,00 1,00 1,00 50,00 0,00 100,00 0,00
Cocholospermaceae Cocholospermum regium Embryos 1,00 0,00 62,00 0,00 1,00 1,00 62,00 0,00 100,00 0,00
Connaraceae Rourea induta Embryos 1,03 0,16 40,00 0,03 1,00 2,00 39,00 1,00 97,50 2,50
Erythroxylaceae Erythroxylum daphnites Embryos 1,00 0,00 139,00 0,00 1,00 1,00 139,00 0,00 100,00 0,00
Euphorbiaceae Manihot anomala Embryos 1,00 0,00 60,00 0,00 1,00 1,00 60,00 0,00 100,00 0,00
Polyembryony analysis
Seeds from 10 different
plants for each species in
the triângulo mineiro region
Polyembryony and apomixis
Our survey: 70 species – 27 families
Total (+ Salomão & Allem): 125 species – 39 families
Apomictics?
Monoembryonic
Polyembryonic
57%
29%
71%
Monoembryonic
Polyembryonic43%
0,00
10,00
20,00
30,00
40,00
50,00
60,00
70,00
80,00
90,00
0 20 40 60 80 100 120 140
6%
23%71%
5%
Polyembryonic
and apomictic ?
Monoembryonic
Polyembryonic
< 5 % seeds
Polyembryonic
> 5 % seeds
Species ranked by
frequency of
apomictic seeds
Polyembryonic species and breeding systems
Family Species Polyembryony % Source BS
Anacardiaceae Astronium fraxinifolium Poli 0,72 Original data Dioecious
Anacardiaceae Myracrodruon urundeuva Poli 2,00 Original data Dioecious
Anacardiaceae Tapirira guianensis Poli 2,00 Salomão e Allem 2001 Dioecious
Apocynaceae Hancornia speciosa Poli 5,00 Salomão e Allem 2001 SI
Bignoniaceae Jacaranda cuspidifolia Poli 0,50 Original data ni
Bignoniaceae Tabebuia chysotricha Poli 68,32 Original data Apomictic
Bignoniaceae Tabebuia ochracea Poli 35,56 Original data SI/Apomictic
Bignoniaceae Tabebuia roseo-alba Poli 0,59 Original data SI
Bignoniaceae Tabebuia serratifolia Poli 21,74 Original data SI/Apomictic
Burseraceae Commiphora lepthophloeos Poli 2,00 Salomão e Allem 2001 ni
Caesalpiniaceae Copaifera langsdorfii Poli 0,17 Original data SI
Caesalpiniaceae Hymenaea courbaril Poli 0,09 Original data SI
Caesalpiniaceae Senna sylvestris Poli 0,29 Original data SI
Connaraceae Rourea induta Poli 2,50 Original data SI
Fabaceae Machaerium acutifolium Poli 2,22 Original data ni
Fabaceae Ormosia arborea Poli 0,25 Original data ni
Flaucortiaceae Casearia sylvestris Poli 0,89 Original data SC
Hippocrateaceae Salacia campestris Poli 2,08 Original data ni
Hippocrateaceae Salacia crassifolia Poli 1,94 Original data ni
Malpighiaceae Byrsonima basiloba Poli 3,00 Salomão e Allem 2001 ni
Malvaceae Eriotheca pubescens Poli 81,82 Original data Apomictic
Malvaceae Guazuma ulmifolia Poli 1,21 Original data SI
Melastomataceae Clidemia hirta Poli 12,53 Original data Apomictic
Melastomataceae Miconia albicans Poli 1,51 Original data Apomictic
Melastomataceae Miconia ferruginata Poli 24,74 Original data Apomictic
Mimosaceae Acacia polyphylla Poli 2,00 Original data
Mimosaceae Inga laurina Poli 41,33 Original data Apomictic?
Myrtaceae Campomanesia pubescens Poli 0,12 Original data SI/SC
Myrtaceae Eugenia calycina Poli 1,31 Original data ni
Myrtaceae Eugenia hegeliana Poli 3,40 Original data ni
Myrtaceae Eugenia punicifolia Poli 0,45 Original data SC
Rhamnaceae Zizyphus joazeiro Poli 2,00 Salomão e Allem 2001 SC
Rubiaceae Genipa americana Poli 3,00 Salomão e Allem 2001 Dioecious
Sapindaceae Magonia pubescens Poli 3,00 Salomão e Allem 2001 ni
Vochysiaceae Callisthene fasciculata Poli 0,22 Original data SI
Vochysiaceae Callisthene major Poli 0,57 Original data SI
36 species
17 families
At least 11 SI
Contrasting
breeding
system results
in some
putative
apomictics
Polyembryony
without apomixis
Callisthene species
(Vochysiaceae)
R. Carmo-Oliveira & L.N. Custódio
unpublished data
Callisthene fasciculata
Callisthene major
Callisthene major
Callisthene fasciculata
Qualea multiflora
Eriotheca pubescensMendes-Rodrigues et al. 2005 Plant biology 7:533-540
Polyembryony and
sporophitic apomixis
Eriotheca pubescensEriotheca gracilipes
Embryos and seedlings per seed of Tabebuia chrysotricha (embryos n =
606 seeds; seedlings n = 640 seeds).
0
5
10
15
20
25
30
35
40
45
1 2 3 4 5 6 7
Number
Fre
qu
ency
(%
)
Embryos
Seedlings
Costa et al. 2004
Sampaio et al. 2007
Handroanthus (syn. Tabebuia)
Bignoniaceae
Apomixis and Neopolyploidy
Oliveira et al. 1992 Sampaio et al. 2008
2n= 276
2n = 40
2n = 80
Eriotheca pubescens
Miconia albicans
Caetano 2007
Apomixis without polyembryony
Goldenberg & Shepherd 1998 Cortez et al. 2008
How widespread are
apomictic taxa?
Handroanthus ochraceus
Eriotheca pubescens
Miconia albicans
Between the 125 most
frequent woody species in
Cerrado (ca. 1000 spp.)
Bridgewaters et al. 2004
2n=40
2n=80
Polyembryonic
individuals
A
100 Km
BA
100 Km
B
Monoembryonic
individuals
Handroanthus ochraceus
(syn. Tabebuia ochracea)
Eriotheca pubescens
Breeding systems mosaics?
Mila et al. in preparation
Breeding system and
distribution in the
Melastomataceae
40%75%
84%
60%
25%16%
0
5
10
15
20
25
30
35
40
Apomictis Self-incompatible Self compatible
Breeding system
Sp
ec
ies
wide
restrict
Apomictics widespread
Questions:
How common is polyembryony among cerrado plants?
Is polyembryony a diagnostic feature to estimate
importance of apomixis in Cerrado plants?
How widespread are polyembryony and apomixis in
Cerrado environment?
What is the origin of apomictic taxa?
±Hybridization?
Polyploidy?
Widely distributed apomictics
Consequences: apomixis as a reproductive alternative ?
Foto: Rogério Gribel
Collevatti, R.G., Grattapaglia, D., & Hay, J.D. 2001. Population genetic structure of the endangered
tropical tree species Caryocar brasiliense, based on variability at microsatellite loci. Molecular
ecology 10: 349-356
Collevatti, R.G., Grattapaglia, D., & Hay, J.D. 2001. High resolution microsatellite based analysis of
the mating system allows the detection of significant biparental inbreeding in Caryocar brasiliense,
an endangered tropical tree species. Heredity 86 (1): 60-67
Disturbance and reduced
gene flow
Caryocar brasiliense
Miconia albicans
- Independent of pollinators
- Bird dispersed fruits and viable seeds
- But no pollen for bees
The future of Cerrado ?
Clesnan Mendes-Rodrigues
Diana Sampaio
Ana Paula Mila
Daniela Simão
Renata Carmo-Oliveira
Luciana N. Custódio
Rosana Romero
Pietro K. Maruyama
Felipe W. Amorim
Ana Paula Caetano
Peter E. Gibbs