Phaseolus (Papilionoideae - Phaseoleae)

V International Legumes Conference

in the Neotropics before Man

D. G. Debouck

Buenos Aires, 12 August 2010

1/27 setting the scene . . .

Tropical Legumes

formation of Phaseoleae by 36 My ago

formation of Papilionoideae by 60‐56 My ago

origin in west Gondwana

South America

•Andean mountain building initiated by 95‐85 My ago

•united with North America by closure of Isthmus of Panama by late Pliocene 3 My ago

• separated from the rest of Gondwana (Laurasia, Africa, Australia) by late Jurassic 144 My ago

map: Taylor 1995 (South America about 80 My ago) 2/27 The Americans cross the Straight of Behring 12,000 – 70,000 years ago

(0.06-3.8 % of the duration of Quaternary [1.8 MyA] !)

source: Zimmer 2005 source: Crawford 1998 3/27 what is wild? what is weedy?

wild; 6 g/ 100 seed

escape; 11 g/ 100 seed 4/27 what is wild? what is weedy?

•growing wild in climax vegetations

•reproduce itself for generationS

•wild attributes in reproduction

(pod dehiscence, dormancy, . . .)

•presence of antinutritional factors (trypsin inhibitor, HCN, . . .)

•higher level of polymorphisms in

proteins, enzymes, DNA fragments

5/27 Phaseolus (Papilionoideae - Phaseoleae) in the Neotropics before Man

1. How many beans, and which were/ are they?

2. How old were/ are these beans ?

How do they relate to each other ?

3. How people have affected the bean potential ?

definition of actions and time set for conservation

6/27 Nature has found different ways at twisting a keel !

“Carina … spiraliter contra solem revoluta … ” Linnaeus 1754 “Phaseolus carina spiralis Vigna omnia Phaseoli nisi carina erostris vel rostro obliquo valde incurvo sed spiram perfectam non efficiente ” Bentham 1865 7/27 1790-1890 The century of botanical explorations Sessé & Mociño, 1787‐1803: Mexico

Lewis & Clark, 1804‐06: W USA

Humboldt & Bonpland, 1815‐25: Mexico

Wilhem Karwinsky, 1826‐31: Mexico

Karl Hartweg, 1836‐39: Mexico

Thomas Coulter, 1825‐32: Mexico

David Douglas, 1826‐32: W coast USA over 400 species described ! Charles Darwin, 1835: Galapagos Is.

Henri Galeotti, 1837‐58: Mexico

Eugène Bourgeau, 1865‐66: Mexico

Edward Palmer, 1865‐85: Mexico

8/27 Towards a better definition . . .

Linnaeus (1753) : 11 sp.

DeCandolle (1825) : 57 sp.

Bentham & Hooker (1865) : ca. 60 sp.

Piper (1926) : 92 sp.

Verdcourt (1971) : circa 50 sp.

I International Legumes Conference Maréchal et al. (1978) : 31 sp.

Lackey (1983) : circa 50 sp.

9/27 Phaseolus rotundatus Freytag & Debouck what is a bean ?

1. all Neotropical

2. trifoliolate leaves with stipules and stipels

3. style with inner brush

4. uncinate hairs on all aerial parts (Ù Vigna, Macroptilium)

5. stipules without extension (Ù Vigna Catiang)

6. length of pedicel = or > calyx (Ù Vigna, Macroptilium)

7. primary bracts permanent (Ù Vigna, Macroptilium)

8. lack of pedicelar glands (Ù Vigna Catiang) (present on stipels and bracteoles)

9. style with 1.5‐2 closed coils (Ù Vigna, Macroptilium)

10. style without extension (Ù Vigna)

11. terminal part of style caducous (Ù Lablab)

12. pods without internal septa (Ù Strophostyles)

after the works by Maréchal et al. 1978; Lackey 1983 drawing from Salcedo et al. 2009 10/27 The last reviews : Freytag & Debouck 2002 : 76 sp. 15 sections Delgado‐Salinas 1985 : 36 sp. 4 sections 1. Acutifolii : 2 sp. 2. Bracteati : 2 sp. 3. Brevilegumeni : 3 sp. Chiapasana (1 species). 4. Chiapasana : 1 sp. 5. Coccinei : 1 sp. 6. Coriacei : 4 sp. 7. Digitati : 5 sp. 8. Falcati : 5 sp. Minkelersia (8 sp.). 9. Minkelersia : 10 sp. 10. Paniculati : 18 sp. Phaseolus (25 sp.). 11. Pedicellati : 11 sp. 12. Phaseoli : 4 sp. 13. Revoluti : 1 sp. 14. Rugosi : 3 sp. Xanthotricha (2 sp.). 15. Xanthotricha : 6 sp. 11/27 Minkelersia Revoluti ! • monophyly Pedicellati Digitati ! • 6-7 My old! • 2 clades Bracteati ! • mcp included! Brevilegumeni Xanthotricha ! Chiapasana

Rugosi 9

Acutifolii Phaseoli ! Coccinei

Falcati 9 8 groups 15 sections

Paniculati Bayesian consensus phylogeny from trnK locus (cpDNA) Coriacei ! source: Delgado-Salinas et al. 2006 source: Freytag & Debouck 2002 12/27 Phaseolus albiflorus Freytag & Debouck

Phaseolus grayanus Wooton & Standley

Section Digitati does not match morphologically with Section Pedicellati

13/27 source: Freytag & Debouck 2002 Minkelersia : 10 Minkelersia : 10 Revoluti : 1 Revoluti : 1

Pedicellati : 11 Pedicellati : 12 Digitati : 5 Digitati : 4 10 Bracteati : 2 Bracteati : 2 9 Brevilegumeni : 3 Brevilegumeni : 5 Xanthotricha : 6 Xanthotricha : 6 Chiapasana : 1 Chiapasana : 1 9

3 Rugosi : 3 Rugosi : 3

Acutifolii : 2 Acutifolii : 2 Phaseoli : 4 Phaseoli : 6 8 Coccinei: 1

Falcati : 5 Falcati : 3 3

71 sp. 7 76 sp. 79 sp.

Paniculati : 18 Paniculati : 19 Bayesian consensus phylogeny from trnK locus (cpDNA) 17 Coriacei : 4 Coriacei : 5 source: Delgado-Salinas et al. 2006 source: Debouck 2010 14/27 How robust are these sections ? Are they valid ? (1) vulg x angu embryo did not grow Belivanis & Doré 1986 vulg x angu 12,72 I + 3,45 II + 0,45 III + 0.18 IV Balasubramanian et al. 2005 vulg x filif 5,69 I + 8,08 II + 0,04 III Maréchal & Baudoin 1978 vulg x filif 12,14 I + 4,71 II + 0,14 III Balasubramanian et al. 2005 vulg x acut 6,0 I + 8,0 II Katanga & Baudoin 1990 vulg x dumo 0,88 I + 10,56 II Schmit et al. 1992 vulg x cocci 0,32 I + 10,84 II Maréchal 1971

Rugosi

Acutifolii

Phaseoli

triporate pollen ! Coccinei parsimony tree from combined analysis of trnK and ITS sequences: Delgado‐Salinas et al. 2006 15/27 g. h. The Rugosi have unique microscopic features in their seed testa

Phaseolus filiformis Bentham

Phaseolus microcarpus Martens after Salcedo‐Castaño et al. 2009 16/27 How robust are these sections ? Are they valid ? (2)

vulg x luna embryo did not grow Kuboyama et al. 1991 luna x jali 2,06 I + 9,97 II Katanga & Baudoin 1990 luna x polys 10,0 I + 6,0 II Dhaliwal et al. 1962 luna x macu 2,31 I + 9,70 II + 0,03 III + 0,06 IV Le Marchand et al. 1976 luna x rite 1,95 I + 10,02 II Katanga & Baudoin 1990 luna x sali 3,61 I + 9,19 II Katanga & Baudoin 1990

Paniculati

Coriacei

Paniculati tricolporate pollen ! Coriacei parsimony tree from combined analysis of trnK and ITS sequences: Delgado‐Salinas et al. 2006 17/27 Pollen types found in Phaseolus species

3-Colporate 3-Colporate 3-Colporate 3-Porate

Pseudocolpus present Pseudocolpus: trace No pseudocolpus No pseudocolpus

lunatus polystachyus coccineus vulgaris maculatus angustissimus albescens polyanthus jaliscanus novoleonensis acutifolius coccineus pachyrrhizoides marechalii carteri costaricensis rotundatus salicifolius filiformis texensis juquilensis xolocotzii microcarpus tuerckheimii neglectus

Sources: Le Marchand & Maréchal 1977; Maréchal et al. 1978; Delgado 1985; Solarte 2000 Ramírez & Delgado 1999; Delgado 2000; Salcedo et al. 2006, 2009; Delgado & Carr 2007 18/27 Time . . .

would some of our molecular clocks be like the ones by Dali ?

“The Disintegration of the Persistence of Memory” Salvador Dali 1952‐54 19/27 ITS major consensus tree from a Bayesian approach

Estimation of divergence times on 200 ITS trees

Average age SD Node MRCA (My) (My)

A Xeroderris – Phaseolus 45.2 1.7

B Wajira – Phaseolus 10.7 0.9

C Vigna ‐ Phaseolus 8.0 0.8

1 rest Mesoamerican – allied Andean 2.2 0.2

2 P. lunatus – allied Andean species 1.0 0.08

3 P. lunatus: pools Andean vs 0.5 0.02 ‘Mesoamericans’

Ages for the Most Recent Common Ancestor for nodes A, B, C are according to Lavin et al. 2005

closure of Isthmus of Panama: 3 mybp !

secondary genepool

tertiary genepool 3

source: Serrano-Serrano et al. 2010 20/27 3 Impact of people: introduction of cattle

21/27 Impact of people: deforestation

22/27 Impact of people: deforestation

San Sebastián de Yali ‐ San Rafael del Norte, western Jinotega, Nicaragua, 13 Dec 2007 23/27 Costa Rica, Alajuela, Zarcero

Four years and nine months later . . .

Impact of people: expansion of road system 8 February 1998

the plant # 3121 was here

. . . the site is gone ! 24/27 Conclusions (1)

the definition of the genus as stated in 1978 holds on (e.g. monophyly, new species)

the number of species is NOT final yet ! up to 80, to 90 sp. (endemisms !) ?!

formation of the genus in extreme SW corner of Laurasia 6‐8 Mya ?! ; speciation thus initiated in that very corner, where logically today highest no. of taxa

formation happened before the closure of the Isthmus of Panama (about 3 Mya), and

speciation went on with most volcanism events in Mexico and Central America

the current marker studies told us the species are of the same age (about 1 My): is that sure ? or is it because we do not have the appropriate marker yet ?

expansion towards the N, the Andes, the Caribbean, thanks to climatic oscillations

25/27 Conclusions (2)

during the SE expansions, three geological breakages acted as barriers;

these were: Isthmus of Tehuantepec, Lake Nicaragua, and Isthmus of Panama

the numbers of species go further down E of Tehuantepec (19), Lake Nicaragua (8) and Isthmus of Panama (5); went SE only vulgaris (twice) and the root of lunatus

defining sections is still worth the effort, because of practical implications

sections that hold true: Chiapasana, Falcati, Minkelersia, Rugosi; more work: Paniculati

monospecific sections? what if there are/ were several sp. after all? extinct lineages?

negative impact by people on the long‐living late species of Coriacei, Paniculati

limited expansion in man‐made habitats (of the species candidate for domestication with epigeal germination, fibrous root, annual, monocarpic)

26/27 Thank you ! Ana L Caicedo Jorge Acosta AGCD Genis Castillo Rodolfo Araya Banco de la República César Azurdia Maria I Chacón BMZ Raúl Castillo Antonio Hernández CIAT Hipólito de la Cruz Jorge Hernández Paul Gepts EU Celia Lima Rogelio Lépiz IBPGR Jenny Motta Luís López IUCN Jorge Liñan Martha Serrano Swiss Aid José Muruaga Alba M Torres USAID Raúl Ríos Joe Tohme María C Sevillano USDA Orlando Toro Luís Valera World Bank

in the lab in the countries for the funding 27/27