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RESEARCH ARTICLE
Contents Vol 57(1)

Phylogenetic relationships among southern South American species of Camptosema, Galactia and Collaea (Diocleinae: Papilionoideae: Leguminosae) on the basis of molecular and morphological data

Silvana M. Sede A F , Daniela Tosto B , Paola Talia B , Melissa Luckow C , Lidia Poggio D and Renée Fortunato E
+ Author Affiliations
- Author Affiliations

A Instituto de Botánica Darwinion, Labardén 200, Casilla de Correo 22, B1642HYD, San Isidro, Buenos Aires, Argentina.

B Unidad Integrada Facultad de Ciencias Exactas y Naturales, UBA y Centro de Investigaciones en Ciencias Veterinarias y Agronómicas (CICVyA), INTA, Argentina.

C L. H. Bailey Hortorium, 462 Mann Library, Cornell University, Ithaca, New York 14853, USA.

D Laboratorio de Citogenética y Evolución, Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Argentina.

E CONICET, Instituto de Recursos Biológicos, Centro de Investigación en Recursos Naturales, INTA, Argentina.

F Corresponding autor. Email: ssede@darwin.edu.ar

Australian Journal of Botany 57(1) 76-86 https://doi.org/10.1071/BT08091
Submitted: 30 May 2008  Accepted: 19 January 2009   Published: 23 March 2009

Abstract

The neotropical genus Camptosema in its southern distribution is represented by the following four species: Ca. paraguariense, Ca. praeandinum, Ca. rubicundum and Ca. scarlatinum. Ca. rubicundum is the only species with all the diagnostic characters of the genus, i.e. tubular calyx. The other three taxa are related morphologically and cytologically to Galactia and Collaea, two closely allied genera in the subtribe Diocleinae. Individual and combined phylogenetic analyses of morphology, ITS and trnL-F were conducted to examine the position of these species as well as to explore their relationship with Galactia and Collaea species in southern South America. In none of the analyses does Camptosema species form a monophyletic group. Instead, they are scattered in different groups. These results together with cytological, morphological and AFLP data provide good support for re-evaluating the taxonomic position of these species within Camptosema.


Acknowledgements

This work was supported by UBA (Universidad de Buenos Aires) – Grant UBACyT EX317 ANPCyT (Agencia Nacional de Promoción Científica y Técnica, Argentina), Grant PICT 14119 CONICET (Consejo Nacional de Investigaciones Científicas y Técnicas) and Grant PIP 02296 and IAPT (International Association of Plant Taxonomy). Collections were supported by the Myndel Botanica Foundation. We thank INTA Cerro Azul, Misiones, Cecilia Laporta, Fernando Fernandez, Silvina Soto and Julián Greppi for assistance and help during the field trips to Misiones and Corrientes, Argentina.


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Appendix 1. List of specimens analysed in the molecular analyses, in alphabetical order by tribe and subtribe, and Genbank accession numbers

Previously published sequences are indicated by Genbank number (*) and literature citation

Outgroup

Tribe Abreae

Abrus precatorius L. ITS AF467015* (Hu et al. 2002)

Tribe Genisteae

Lupinus paraguariensis Chodat & Hassl. ITS AF007476* (Ainouche and Bayer 1999), trnL-F AF538709* (A Ainouche, MT Misset, RJ Bayer, unpubl.)

Tribe Millettieae

Austrosteenisia blackii (F.Muell.) Geesink. ITS AF467020* (Hu et al. 2002)

Lonchocarpus rugosus Benth. ITS AF467062* (Hu et al. 2002)

Lonchocarpus sp. ITS AF467064* (Hu et al. 2002)

Millettia grandis Skeels. ITS AF467474* (Hu et al. 2002)

Millettia sp. ITS AF467480* (Hu et al. 2002)

Tribe Phaseoleae – subtribe Diocleinae

Canavalia bonariensis Lindl. ITS AY293839* (Varela et al. 2004)

Can. brasiliensis Mart. ex Benth. ITS AF467034* (Hu et al. 2002)

Cleobulia multiflora Mart. ex Benth. ITS AY881100* (TB Grangeiro, DMM Jorge, ES Varela, JPMS Lima, WM Bezerra, EP Nunes, MAO Alves, unpubl.)

Cymbosema roseum Benth. ITS AY293836* (Varela et al. 2004)

Dioclea bicolor Benth. ITS AY293833* (Varela et al. 2004)

Dioclea virgata (Rich.) Amshoff. ITS AY293835* (Varela et al. 2004)

Tribe Phaseoleae – subtribe Phaseoliinae

Macroptilium psammodes (Lindm.) S.I.Drewes & R.A.Palacios. ITS1 DQ888774*, ITS2 DQ888786*, trnL-F DQ888796* (Espert et al. 2007)

Ingroup

Tribe Phaseoleae – subtribe Diocleinae

Camptosema coccineum (Mart. ex Benth.) Benth. Fortunato 8433 (BAB), ITS EU927595, trnL-F FJ613485

Ca. coriaceum (Nees & Mart.) Benth. Fortunato 8452 (BAB), ITS EU927596, trnL-F FJ613486

Ca. paraguariense (Chodat & Hassl.) Hassl. Fortunato 6805 (BAB), ITS EU927592, trnL-F FJ613481

Ca. praeandinum Burkart. Fortunato 6936 (BAB), ITS EU927593, trnL-F FJ613482

Ca. rubicundum Hook. & Arn. ITS EU499368* (Espert et al. 2008), trnL-F FJ613483

Ca. scarlatinum (Mart. ex Benth.) Burkart. Fortunato 8003 (BAB), ITS EU927594, trnL-F FJ613484

Collaea argentina Griseb. Fortunato 7673 (BAB), ITS EU927599, trnL-F FJ613490

Co. aschersoniana (Taub.) Burkart. Fortunato 8002 (BAB), ITS EU927597, trnL-F FJ613487

Co. cipoensis Fortunato. ITS EU499369* (Espert et al. 2008), trnL-F FJ613491

Co. paraguariensis (Hassl.) R. García. Fortunato 8043 (BAB), ITS EU927598, trnL-F FJ613489

Co. stenophylla (Hook. & Arn.) Benth. EU499370* (Espert et al. 2008), ITS trnL-F FJ613488

Galactia benthamiana Micheli. Fortunato 7886 (BAB), ITS EU927600, trnL-F FJ613492

G. boavista (Vell.) Burkart. Fortunato 7544 (BAB), ITS EU927601, trnL-F FJ613493

G. dubia DC. Fortunato 7170 (BAB), ITS EU927602, trnL-F FJ613494

G. fiebrigiana Burkart. Sede 12 (BAB), ITS EU927603, trnL-F FJ613495

G. latisiliqua Desv. Fortunato 6840 (BAB), ITS EU927604, trnL-F FJ613496

G. lindenii Burkart. ITS EU499371* (Espert et al. 2008), trnL-F FJ613497

G. longifolia (Jacq.) Benth. Fortunato 7907 (BAB), ITS EU927605, trnL-F FJ613498

G. marginalis Benth. Fortunato 7185 (BAB), ITS EU927606, trnL-F FJ613499

G. martii DC. Fortunato 8447 (BAB), ITS EU927607, trnL-F FJ613500

G. neesii DC. Fortunato 8060 (BAB), ITS EU927608, trnL-F FJ613501

G. striata (Jacq.) Urban. Sede 9 (BAB), ITS EU927609, trnL-F FJ613502

G. texana (Scheele) A.Gray. Fortunato 7577 (BAB), ITS EU927610, trnL-F FJ613503


Appendix 2. List of specimens analysed in the morphological analysis, in alphabetical order by tribe and subtribe

Outgroup

Tribe Millettieae

Lonchocarpus nitidus (Vogel) Benth., Argentina: Dematteis 265 (SI); Rodríguez 711 (SI). Brasil: Burkart 25131 (SI)

Tribe Phaseoleae – subtribe Diocleinae

Canavalia bonariensis Lindl. Argentina: Burkart 4886 (SI); Jozami 263 (SI)

Cleobulia multiflora Mart. ex Benth. Brasil: Williams and Assis 6803 (SI); Pabst s/n (SI); Pabst 5384 (SI)

Cratylia mollis Mart. ex Benth. Brasil: Hatschbach 1515 (SI)

Dioclea virgata (L.C. Rich) Amshoff. Bolivia: Serato 19 (BAB)

Ingroup

Tribe Phaseoleae – subtribe Diocleinae

Camptosema coccineum (Mart. ex Benth.) Benth. Brasil: Hatschbach 51096 (BAB); Fortunato 8452 (BAB); Fortunato 8433 (BAB)

Ca. coriaceum (Nees & Mart.) Benth. Brasil: Arbo 7540 (BAB); Hatschbach 59569 (MBM); Fortunato 8452 (BAB)

Ca. paraguariense (Chodat & Hassl.) Hassl. Argentina: Ragonese 7025 (BAB); Fortunato 2869 (BAB); Fortunato 6805 (BAB)

Ca. praeandinum Burkart. Argentina: Fortunato 6936 (BAB); Venturi 1379 (SI); Fortunato 7271 (BAB)

Ca. rubicundum Hook. & Arn. Argentina: Hurrell 3445 (BAB); Greppi 25 (BAB); Sede 17 (BAB)

Ca. scarlatinum (Mart. ex Benth.) Burkart. Argentina: Llamas 826 (BAB); Fortunato 8003 (BAB). Brasil: Barreto 5688 (SP)

Collaea argentina Griseb. Argentina: Fortunato 5333 (BAB); Krapovickas 4980 (BAB); Fortunato 7673 (BAB)

Co. aschersoniana (Taub.) Burkart. Argentina: Fortunato 8002 (BAB). Brasil: Jonsson 1072a (G) Co. cipoensis Fortunato. Brasil: Chukr et al. (BAB); Abbud (BAB). Fortunato 8411 (BAB)

Co. paraguariensis (Hassl.) R.García. Argentina: Fontana 5089 (BAB); Jorgensen 578 (BAB); Fortunato 8043 (BAB)

Co. stenophylla (Hook. & Arn.) Benth. Argentina: Crovetto 6301 (BAB); Sede and Fernández 6 (BAB); Sede 3 (BAB)

Galactia benthamiana Micheli. Argentina: Ragonese 6832 (BAB); Zamboni 34767 (SI); Fortunato 7886 (BAB)

G. boavista (Vell.) Burkart. Argentina: Fortunato 8072 (BAB); Sede and Fernández 4 (BAB); Fortunato 7544 (BAB)

G. dubia DC. Argentina: Kiesling 5714 (SI). Fortunato 7175 (BAB); Fortunato 7170 (BAB)

G. lindenii Burkart. Colombia: Haught 6182 (SI); Ruiz 2 (BAB); Ruiz 3 (BAB)

G. fiebrigiana Burkart. Argentina: Fortunato 8167 (BAB); Sede 12 (BAB)

G. latisiliqua Desv. Argentina: Fortunato 7916 (BAB); Bacigalupo 1345 (SI); Fortunato 6840 (BAB) G. longifolia (Jacq.) Benth. Argentina: Schulz 17745 (BAB); Fortunato 2797 (BAB); Fortunato 7907 (BAB)

G. marginalis Benth. Argentina: Crovetto 8649 (BAB); Fortunato 7908 (BAB); Fortunato 7185 (BAB)

G. martii DC. Brasil: Hatschbach 47543 (MBM); Macedo 2734 (SP); Fortunato 8447 (BAB)

G. neesii DC. Argentina. Fortunato 8060 (BAB). Brasil: Hatschbach 54517 (MBM)

G. striata (Jacq.) Urban. Argentina. Bacigalupo et al. 1302 (BAB); Fortunato et al. 8050 (BAB); Sede 9 (BAB)

G. texana (Scheele) A.Gray. Argentina: Fortunato 5997 (BAB); Fortunato 7577 (BAB)


Appendix 3. List of morphological characters used in the phylogenetic analysis

Vegetative character

1. Habit: trees (0); lianas, vines or shrubs (1); subffrutex or herbs (2). 2. Leaf rachis: absent (0); present (1). 3. Number of leaflets: one (0); three (1). 4. Petiole: absent (0); present (0). 5. Leaf texture: membranous, papyraceous or chartaceous (0); coriaceous (1). 6. Leaf shape: narrowly oblong (0); broadly elliptic-obovate (1). 7. Stipels: absent (0); semipersistent (caducous) (1); persistent (2).

Inflorescence

8. Length of inflorescence v. length of subtending leaf: same size or longer (0); shorter (1). 9. Brachyblast: absent (0); present (1). 10. Woodiness of the rachis: not woody (0); woody (1). 11. Inflorescence type: elongate raceme (0); umbelliform/corymbiform raceme (1). 12. Flower bracts: not strobiliform (0); strobiliform (1). 13. Bracteole persistence: caducous (0); persistent (1). 14. Length of the bracteole v. length of the calyx tube: shorter (0); of the same size or longer (1). 15. Bracteole position: at the base of the calyx (0); below the base of the calyx (1). 16. Bracteole shape: obovate to orbicular (0); subulate to triangular (1). 17. Flower bud shape: ovate to cylindrical (0); orbicular (1). 18. Length of the pedicels: short (0 to <7 mm) (0); median (7 to <15 mm) (1); large (15 to <30 mm) (2) (states were determined with ANOVA, followed by comparisons with Scheffée’s test (Sokal and Rohlf 1995)).

Flower

19. Calyx color: red (0); green (1). 20. Calyx shape: campanulate (0); tubular (1). 21. Calyx inner surface indument: glabrous (0); only at the lobes (1); all over the surface (2) (ordered). 22. Upper lobe shape: triangular to lanceolate (0); elliptic to ovate (1); broadly rectangular (2). 23. Petal color: red (0); blue lilac (1). 24. Standard shape: orbicular (0); obovate to elliptic (1); oblong (2). 25. Standard outer surface indument: glabrous (0); only at the upper portion (1); all over the surface (2) (ordered). 26. Callous areas: absent (0); present (1). 27. Callous areas indument: absent (0); present (1). 28. Standard reflexion: absent (0); present (1). 29. Standard auricles: vestigial or absent (0); present (1). 30. Wing shape: elliptic to oblong (0); broadly obovate (1). 31. Indument on the margins of the wings: absent (0); present (1). 32. Keel apex: straight (0); incurved (1). 33. Keel upper margin: entire (0); toothed (1). 34. Stamen connation: pseudomonadelphous (0); diadelphous (1). 35. Anther indument: absent (0); present (1); 36. Nectary disc shape: cylindrical (0); conic (1); truncate (2). 37. Nectary disc margin: irregular (0); lobulate (1). 38. Ovary: sessile to subsessile (0); short-stipitate (1); long-stipitate (1). 39. Style: geniculate at the base or the middle (0); straight (1).

Fruit

40. Fruit margin: straight (0); undulate (1). 41. Fruit shape: straight (0); curved (1). 42. Fruit indument: glabrous (0); pilose (1); sericeous (2). 43. Beak position: central (0); lateral (1).

Seed

44. Seed lateral compression: not compressed (0); compressed (1). 45. Strophiole: incomplete (0); complete (1). 46. Hilum shape: obovate to elliptic (0); linear (1).