Abstract
Allometric relationships in organisms are considered a universal phenomenon. A positive scaling has been reported between stem size and cellular size of tracheary elements in wood of different vascular plants, but few studies have been carried out in slow-growing succulent plants. The aim of this study was to evaluate if a relationship exists between size, growth form and wood cell size among individual species of Cacteae. Forty-four species belonging to 16 genera of the tribe Cacteae with differing growth forms and sizes were studied. When analyzing plant size, we found a positive allometric scaling and the larger-sized species showing a higher percentage of succulent tissue and less accumulation of wood tissue. The positive scaling found between plant size (height and diameter) and vessel elements and fiber length support the universality of the allometric relationship proposed for other vascular plants with non-succulent stems. Notably, wide-band tracheids do not scale with plant size or growth form. Succulence associated with narrow vessel elements with distinctive helical secondary walls and wide-band tracheids suggest they are the key adaptations to tolerate drought and provide support to the stems of most taxa in Cacteae. Fibers do not have the primary role of giving mechanical support; therefore, we assume the scarce fibers in clusters represent reaction wood that, along with the fundamental tissue, maintains the vertical position and shape of those species growing in rocky cracks. Our results with species having short succulent stems support the universal theory of positive allometric scaling of vascular plants.



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Acknowledgments
Thanks to Posgrado en Ciencias Biológicas, UNAM and Consejo Nacional de Ciencia y Tecnología (CONACyT) for the PhD scholarship to M.V.S. (41991). Funding was provided by the Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica, DGAPA, UNAM IN224307 to T.T. and by Instituto de Biología. Lauro López-Mata gave valuable comments.
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Appendix
Appendix
Species studied, collector number, and state in Mexico in which the plant was collected. All herbarium vouchers were deposited at Herbario Nacional, Universidad Nacional Autónoma de México (MEXU).
Acharagma roseana (Boed.) E.F.Anderson, C.Glass6443, Coahuila; Astrophytum asterias (Zucc.) Lem., TT846, Tamaulipas; Astrophytum myriostigma Lem., SA1730, San Luis Potosí; Astrophytum ornatum (DC.) F.A.C.Weber ex Britton & Rose, SA1699, Querétaro; Aztekium ritterii (Boed.) Boed. ex Berger, SA1868, Nuevo León; Coryphantha clavata (Scheidw.) Backeb., SA1705, Querétaro; Coryphantha cornifera (DC.) Lem., SA1700, Querétaro; Coryphantha erecta (Lem.) Lem., SA1684, Querétaro; Coryphantha macromeris (Engelm.) Lem., SA1775, Nuevo León; Echinocactus horizonthalonius Lem., SA1691, Querétaro; Echinocactus parryi Engelm., SA1791, Chihuahua; Echinocactus platyacanthus Link & Otto, SA1850, San Luis Potosí; Echinocactus texensis Hopffer, TT851, Tamaulipas; Ferocactus chrysacanthus (Orcutt) Britton & Rose, SA1815, Baja California; Ferocactus fordii (Orcutt) Britton & Rose, SA1809, Baja California; Ferocactus glaucescens (DC.) Britton & Rose, SA1701, Querétaro; Ferocactus haematacanthus (Salm-Dyck) Bravo, SA1796, Puebla; Ferocactus hamatacanthus (Muehlenpf.) Britton & Rose, TT839, Nuevo León; Ferocactus herrerae J.G. Ortega, SA1833, Sonora; Ferocactus macrodiscus (Mart.) Britton & Rose, SA1798, Oaxaca; Ferocactus pilosus (Galeotti ex Salm-Dyck) Werderm., SA890, San Luis Potosí; Ferocactus recurvus (Mill.) Y. Ito ex G.E. Linds., SA1794, Puebla; Ferocactus reppenhagenii G. Unger, HJA1179, Jalisco; Ferocactus robustus (Pfeiff.) Britton & Rose, SA1795, Puebla; Glandulicactus crassihamathus (F.A.C. Weber) Backeb., SA1688, Querétaro; Glandulicactus uncinatus (Galeotti ex Pfeiff. & Otto) Backeb., SA1899, Durango; Leuchtenbergia principis Hook., HSM3826, San Luis Potosí; Lophophora diffusa (Croizat) Bravo, SA1698, Querétaro; Mammillaria albilanata Backeb., TT816, Oaxaca; Mammillaria elongata DC., SA1697, Querétaro; Mammillaria heyderi Muehlenpf., TT829, San Luis Potosí; Mammillaria winterae Boed., SA1870, Nuevo León; Neolloydia conoidea (DC.) Britton & Rose, SA1695, Querétaro; Pelecyphora aselliformis C. Ehrenb., UG2907, San Luis Potosí; Stenocactus dichroacanthus (Mart. ex Pfeiff.) A. Berger ex Backeb. & F.M. Knuth., SA1758, Aguascalientes; Strombocactus disciformis (DC.) Britton & Rose, SA1738, Querétaro; Thelocactus bicolor (Galeotti ex Pfeiff.) Britton & Rose, TT895, Coahuila; Thelocactus conothelos (Regel & Klein) Backeb. & F.M. Knuth, TT844, Nuevo León; Thelocactus heterochromus (F.A.C.Weber) van Oosten, SA1898A, Querétaro; Thelocactus hexaedrophorus (Lem.) Britton & Rose, TT838, San Luis Potosí; Thelocactus leucacanthus (Zucc. ex Pfeiff.) Britton & Rose, SA1681, Querétaro; Thelocactus setispinus (Engelm.) E.F.Anderson, SA1856, Tamaulipas; Turbinicarpus beguinii (N.P. Taylor) Mosco & Zanovello, SA1852, Nuevo León; Turbinicarpus schmiedickeanus (Boed.) Buxb. & Backeb., TT820, San Luis Potosí.
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Vázquez-Sánchez, M., Terrazas, T. Stem and wood allometric relationships in Cacteae (Cactaceae). Trees 25, 755–767 (2011). https://doi.org/10.1007/s00468-011-0553-y
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DOI: https://doi.org/10.1007/s00468-011-0553-y


