Abstract
The genus Peniocereus contains distinctive cacti with slender stems and thickened roots. The phylogenetic relationship of Peniocereus to other members of the subtribe Pachycereinae is unresolved, and a sister group for the genus has not been suggested in previous molecular phylogenies. Therefore, this study aims to corroborate the monophyly of Peniocereus, to determine the relationships within the genus, to clarify their relationship with Nyctocereus, and to reconstruct the ancestral states for eight morphological characters and the areas of distribution. Five chloroplast markers (rpl16, petL-psbE, psbA-trnH, trnL-trnF, and rpl32-trnL) and a broad outgroup represented by all genera of the subtribes Pachycereinae and Echinocereinae were used in phylogenetic analyses. The results strongly supported the monophyly of Peniocereus, which comprises P. greggii, P. johnstonii, P. lazaro-cardenasii, P. marianus, P. striatus, P. viperinus, and P. zopilotensis. The genus was sister to the Pachycereus group, which comprises Carnegiea, Lophocereus, Marshallocereus, Nyctocereus, and Pachycereus (including Backebergia and Pterocereus). The scandent branches, slender stems, and papillose epidermis are ancestral characters to Peniocereus. A complex region including the Pacific Lowlands, the Balsas Basin, and the Sonoran Desert was inferred as the ancestral area for the genus. This work provides the most comprehensive phylogeny of the subtribe Pachycereinae.
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Acknowledgements
The first author thanks the Posgrado en Ciencias Biológicas UNAM for providing the facilities for this project and Consejo Nacional de Ciencia y Tecnología for the scholarship (385257). We thank the UNAM-DGAPA-PAPIIT program <IN208619> for SA, which supported this work. DS thanks to the program “Investigadoras e investigadores por México” (CONACYT, project 985). We are grateful to Lidia Cabrera (Instituto de Biología, UNAM) for their kind help and Yolanda Morales, Isaura Rosas, Miguel Á. Cruz, David Aquino, Gabriel Olalde, and Héctor J. Tapia (Jardín Botánico, IB-UNAM) for supporting the laboratory and field work. We thank Berenice García Martínez for the illustration of Peniocereus. We thank the reviewers and editors for their valuable comments and suggestions, which have improved our manuscript.
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Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (Grant No. IN208619).
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All authors contributed to the study conception and design. S.A. conducted funding acquisition. Material preparation, data collection, and analysis were performed by D.F.E., D.S., and S.A. The first draft of the manuscript was written by D.F.E. and D.S. All authors commented on previous versions of the manuscript and approved the final manuscript.
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Appendix 1
Appendix 1
List of taxa sampled in alphabetical order, provenance, voucher (HERBARIUM ACRONYM), and GenBank numbers in the following order: trnL-trnF/psbA-trnH/petL-psbE/rpl32-trnL/rpl16. Country codes: GT Guatemala, MX Mexico, PR Puerto Rico, SV El Salvador, and USA United States of America. The dash (–) indicates missing sequence, only in Lophocereus schottii for rpl32-trnL.
1. Acanthocereus chiapensis Bravo, Chiapas, MX, Guzmán 999 (MEXU): KU598057/KU597952/OK302947/OK302980/KU598005. 2. Bergerocactus emoryi (Engelm.) Britton & Rose, Baja Cal., MX, Arias 1307 (MEXU): DQ099925/KF783478/KY624800/KY624730/DQ099994. 3. Cephalocereus apicicephalium E. Y. Dawson, Oaxaca, MX, Arias 1257 (MEXU): DQ099927/KY624684/KY624787/KY624717/DQ099996. 4. C. fulviceps (F. A. C. Weber ex K. Schum.) H. E. Moore, Puebla, MX, Tapia 29 (MEXU): AY181621/KY624683/KY624786/KY624716/AY181602. 5. C. mezcalaensis Bravo, Guerrero, MX, Terrazas 533 (CHAPA): AY181645/KY624671/KY624774/KY624704/AY181600. 6. C. polylophus (DC.) Britton & Rose, Hidalgo, MX, Terrazas 530 (CHAPA): AY181644/KY624673/KY624776/KY624706/AY181597. 7. C. senilis (Haw.) Pfeiff., Hidalgo, MX, Terrazas 529 (CHAPA): AY181638/KY624687/KY624790/KY624720/AY181616. 8. C. tetetzo (F. A. C. Weber ex J. M. Coult.) Diguet., Oaxaca, MX, Tapia 41 (MEXU): AY181632/KY624682/KY624785/KY624715/AY181592. 9. Carnegiea gigantea (Engelm.) Britton & Rose, Arizona, USA, NYBG-sn. cult.: AY181619/KY624692/KY624795/KY624725/AY181591. 10. Deamia chontalensis (Alexander) Doweld., Oaxaca, MX, Yañez 05 (MEXU): MH107805/OK302930/OK302949/OK302982/MH107790. 11. D. montalvoae J. Menjívar & S. Arias, Chiapas, MX, Arias & Cruz 1640 (MEXU): MH107806/MH107794/OK302950/OK302983/MH107791. 12. D. testudo (Karw. ex Zucc.) Britton & Rose, Izabal, GT, Véliz 19628 (BIGU): KY624663/KY624696/KY624802/KY624732/KY624649. 13. Echinocereus pentalophus (DC.) Lem., Querétaro, MX, Arias 1737 (MEXU): KF783628/KF783509/OK302951/OK302984/KF783558. 14. E. polyacanthus Engelm., Chihuahua, MX, Sánchez 24 (MEXU): KF783655/KF783510/OK302952/OK302985/KF783585. 15. Escontria chiotilla (F. A. C. Weber ex K. Schum.) Rose., Oaxaca, MX, Arias 891 (MEXU): KF783680/KF783542/KY624796/KY624726/KF783610. 16. Harrisia earlei Britton & Rose., Saint Croix Bot. Garden, USA, Fleming 118 cult.: DQ099939/KY624665/KY624768/KY624698/DQ100008. 17. Lemaireocereus hollianus (F. A. C. Weber ex J. M. Coult.) Britton & Rose, Puebla, MX, Arias 1373 (CHAPA): AY181623/KF783544/KY624805/KY624735/AY181603. 18. L. lepidanthus (Eichlam) S. Arias & Terrazas, Guatemala, GT, Cseh s.n. cult.: AY181639/OK302932/OK302953/OK302986/AY181598. 19. Lophocereus gatesii M. E. Jones, Baja Cal. Sur, MX, Hamann s.n. cult.: AY181637/OK302934/OK302954/OK302987/AY181601. 20. L. marginatus (DC.) S. Arias & Terrazas, Puebla, MX, Arias 1372 (CHAPA): AY181627/OK302935/OK302955/OK302988/AY181618. 21. L. schottii (Engelm.) Britton & Rose, Baja Cal. Sur, MX, Terrazas 474 (CHAPA): AY181620/MH107796/OK302956/–/AY181613. 22. Leptocereus quadricostatus (Bello) Britton & Rose, Cabo Rojo, PR, Arias 1464 (MEXU): KF783690/OK302933/KY624807/KY624737/KF783620. 23. Marshallocereus aragonii (F. A. C. Weber) Backeb. subsp. aragonii, Departamento Chalatenango, SV, Menjívar 2996 (MHES): AY181630/OK302936/OK302957/OK302989/AY181611. 24. M. aragonii (F. A. C. Weber) Backeb. subsp. eichlamii (Britton & Rose) Guiggi, Yucatán, MX, Arias 1363 (MEXU): AY181629/KY624690/KY624793/KY624723/AY181610. 25. Myrtillocactus geometrizans (Mart. ex Pfeiff.) Console, Querétaro, MX, Terrazas 557 (CHAPA): DQ099943/KF783546/KY624798/KY624728/DQ100012. 26. M. schenckii (J. A. Purpus) Britton & Rose, Puebla, MX, Terrazas 500 (CHAPA): AY181633/KF783547/OK302959/OK302991/AY181607. 27. Morangaya pensilis (K. Brandegee) G. D. Rowley, Baja Cal. Sur, MX, Arias 1295 (MEXU): KF783679/KF783538/OK302958/OK302990/KF783609. 28. Nyctocereus serpentinus (Lag. & Rodr.) Britton & Rose, Michoacán (MICH1), MX, Arias 2229 (MEXU): OK040809/OK302937/OK302960/OK302992/OK302927. 29. N. serpentinus (Lag. & Rodr.) Britton & Rose, Michoacán (MICH2), MX, Sánchez-Mejorada 4249 (MEXU): DQ099959/MH107797/OK302961/OK302993/DQ100028. 30. N. serpentinus (Lag. & Rodr.) Britton & Rose, Oaxaca, MX, Guzmán 1285 (MEXU): DQ099960/OK302938/OK302962/OK302994/DQ100029. 31. Peniocereus greggii (Engelm.) Britton & Rose subsp. greggii, Durango, MX, Arias 2054 (MEXU): DQ099948/OK302941/OK302967/OK302999/DQ100017. 32. P. greggii (Engelm.) Britton & Rose subsp. transmontanus (Engelm.) U. Guzmán, Arizona, USA, Quirk s.n. (DES-1960.6704): DQ099949/OK302942/OK302968/OK303000/DQ100018. 33. P. johnstonii Britton & Rose, Baja Cal. Sur, MX, Arias 1296A (MEXU): DQ099951/OK302943/OK302969/OK303001/DQ100020. 34. P. lazaro-cardenasii (Contreras et al.) D. R. Hunt, Michoacán, MX, Contreras 20 (MEXU): DQ099952/KF783550/OK302970/OK303002/DQ100021. 35. P. marianus (Gentry) Sánchez-Mej., Sinaloa, MX, Guzmán 1038 (MEXU): DQ099955/MH107800/OK302971/OK303003/DQ100024. 36. P. pecten-aboriginum (Engelm. ex S. Watson) Britton & Rose, Guerrero, MX, Terrazas 535 (CHAPA): AY181624/OK302939/OK302964/OK302996/AY181615. 37. P. pringlei (S. Watson) Britton & Rose, Baja Cal. Sur, MX, Arias 1348 (CHAPA): AY181642/KF783549/KY624799/KY624729/AY181589. 38. P. striatus (T. Brandegee) Buxb., Sonora, MX, McGill s.n. (DES-1962.7225): DQ099962/MH107801/OK302972/OK303004/DQ100031. 39. P. tepamo Gama-López & S. Arias, Michoacán, MX, Gama 107 (MEXU): AY181647/MH107799/OK302965/OK302997/AY181593. 40. P. viperinus (F. A. C. Weber) Kreuz., Morelos, MX, Gold s.n. (MEXU): OK040810/OK302944/OK302973/OK303005/OK302928. 41. P. viperinus (F. A. C. Weber) Kreuz., Puebla, MX, Gama 37 (MEXU): DQ099964/MH107802/OK302974/OK303006/DQ100033. 42. P. weberi (J. M. Coult.) Backeb., Puebla, MX, Arias 747 (MEXU): AY181631/OK302940/OK302966/OK302998/AY181614. 43. P. zopilotensis (J. Meyrán) Buxb., Guerrero, MX, Scheinvar s.n. (MEXU): DQ099965/OK302945/OK302975/OK303007/DQ100034. 44. Pachycereus gaumeri Britton & Rose, Yucatán, MX, Arias 1360 (MEXU): AY181626/KY624691/KY624794/KY624724/AY181606. 45. P. grandis Rose, Puebla, MX, DFE 31 (MEXU): AY181646/MH107798/OK302963/OK302995/AY181605. 46. P. militaris (Audot) D. R. Hunt, Michoacán, MX, Arias 1339 (CHAPA): AY181628/OK302946/OK302948/OK302981/AY181609. 47. Polaskia chende (Rol.-Goss.) A. C. Gibson & K. E. Horak., Puebla, MX, Arias 1528 (MEXU): OK040808/OK302931/OK302976/OK303008/OK302929. 48. P. chichipe (Rol.-Goss.) Backeb., Puebla, MX, Arias 1529 (MEXU): KF783684/KF783551/OK302977/OK303009/KF783614. 49. Stenocereus pruinosus (Otto ex Pfeiff.) Buxb., Puebla, MX, Arias 750 (MEXU): KF783688/KF783554/OK302979/OK303011/KF783618. 50. S. stellatus (Pfeiff.) Riccob., Puebla, MX, Arias 1375 (MEXU): AY181640/KF783555/KY624797/KY624727/AY181590. 51. Selenicereus atropilosus Kimnach, Jalisco, MX, Arreola 1473 (MEXU): KU598082/KU597977/OK302978/OK303010/KU598029. 52. Strophocactus wittii (K. Schum.) Britton & Rose, Hannover, Herrenhäuser Gärten B05X138 cult.: KY624664/KY624697/KY624803/KY624733/KY624650.
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Franco-Estrada, D., Sánchez, D., Terrazas, T. et al. Phylogenetic analysis of Peniocereus (Cactaceae, Echinocereeae) based on five chloroplast DNA markers. Braz. J. Bot 44, 903–916 (2021). https://doi.org/10.1007/s40415-021-00774-0
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DOI: https://doi.org/10.1007/s40415-021-00774-0