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Seed morphology and variation in the genus Pachycereus (Cactaceae)

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Abstract

Seeds of 13 Pachycereus species and two Stenocereus species that have been suggested as closely related were examined with the scanning electron microscope. Quantitative features were evaluated using multivariate analysis in order to identify characters that distinguish them. Several species groups were recognized on the basis of 16 qualitative characters. All species studied are keeled. Stenocereus aragonii and S. eichlamii share with most Pachycereus species large size, glossy appearance, and a flat relief on periclinal cells in the lateral region. Pachycereus gatesii and P. schottii are unique in having the smallest seeds and a deeply impressed hilum-micropylar region. P. hollianus does not exhibit micro-relief on periclinal walls in the lateral region, and P. fulviceps has no expanded testa border. Multivariate analysis showed that four characters, length, breadth, hilum-micropylar region length, and angle, made the greatest contribution to distinguishing among species groups. More than 80% of P. fulviceps, P. hollianus, P. tepamo, P. weberi, and S. eichlamii seeds could be classified correctly using four seed features and the percentage was even higher using just two or three features for P. gatesii, P. grandis, P. militaris, P. pringlei, and P. schottii. Testa appearance, testa cell-pattern, and position relative to the rim of the hilum-micropylar region were found to be potentially informative and should be combined with other sources of data in future phylogenetic analyses.

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Fig. 1A, B
Fig. 2A–H
Fig. 3A–F
Fig. 4A–H
Fig. 5A–H
Fig. 6
Fig. 7A–F

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Acknowledgements

This study was supported by Consejo Nacional de Ciencia y Tecnología grant 33064-V (T.T.) and scholarship 127274 (S.A.). We thank the herbaria ariz, chapa, cicy, mexu, ny, and us for allowing us to sample seeds. We are also indebted to H. Arreola-Nava, U. Guzmán, and A. Orduño, who provided some samples from collection in the wild, to Silvia Espinosa for technical assistance in SEM, to Lauro López-Mata for helpful comments on the statistics, to Héctor Hernández for darkroom assistance, and to two reviewers for helpful comments on the manuscript.

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Correspondence to Teresa Terrazas.

Appendix

Appendix

Species seed descriptions are given below.

  • Pachycereus fulviceps (S. Arias 863) (Figs. 2A, 4A, 5D). Seed broadly oval, large, 2.5–2.95×1.6–2.45 mm, black-brown, glossy, periphery keeled; border not expanded around hilum; cells uniform, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices undifferentiated; relief flat; convexities none; micro-relief minutely rugose; HMR small, 0.64–1.16 mm, oblique, 29–49°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. gatesii (S. Arias 1343a, b, 1344a, b) (Figs. 2B, 4B). Seed broadly oval, medium-sized 1.59–2.58×1–1.79 mm, black to black-brown, glossy, periphery keeled; border highly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, uncommon on lateral region; relief flat on lateral region; slightly convex on dorsal and apical ridges, convexities low domes; micro-relief minutely rugose; HMR small, 0.16–1 mm, oblique, 34–57°, deeply impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. gaumeri (S. Arias 1087, 1359, 1360, 1366; Escalante 1023; U. Guzmán-Cruz 995) (Figs. 4D, 5A). Seed broadly oval, very large, 2.22–3.72×1.8–2.69 mm, black to black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, uncommon on lateral region; relief flat on lateral region; slightly convex on peripheral ridges; convexities low domes; micro-relief rugose; HMR medium-sized, 0.8–1.54 mm, oblique, 26–54°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. grandis (S. Arias 1383a, b, c, d; U. Guzmán-Cruz 1119) (Figs. 2C, 4C). Seed oval, extremely large, 3.84–6.15×2.44–3.68 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries slightly raised on lateral region, but inconspicuous around peripheral regions, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR medium-sized, 0.17–2.27 mm, oblique, 28–61°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. hollianus (S. Arias 830, 1331, s.n.; L. Scheinvar s.n.; J.N. Rose 10148) (Figs. 2G, 5C). Seed broadly oval, large, 2.23–3.15×1.6–2.45 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, sometimes inconspicuous on lateral region, straight; interstices minutely pitted, common on all regions; relief flat on lateral region, slightly convex on peripheral ridges; convexities low domes; micro-relief none; HMR medium size, 0.9–1.69 mm, oblique, 44–73°, finely impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. lepidanthus (F. Eichlam 23a, b) (Figs. 2F, 4E, 5B). Seed broadly oval, large, 2.45–3.49×1.69–2.36 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR small, 0.77–1.35 mm, oblique, 32–51°, finely impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. marginatus (S. Arias 1386; H. Bravo s.n. a, b; T. Terrazas 362a, b) (Fig. 3A). Seed broadly oval, extremely large, 3.25–5.45×2.15–4.52 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric and elongate-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, more common on peripheral region; relief flat on lateral region, slightly convex on peripheral ridges; convexities low domes; micro-relief minutely rugose; HMR medium size, 0.93–2.24 mm, oblique, 21–56°, finely impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. militaris (S. Arias 1332, 1336, 1338, 1339) (Fig. 3B). Seed broadly oval, extremely large, 3.81–5.13×2.57–3.8 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted; uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR medium size, 0.79–1.75 mm, oblique, 16–49°, finely impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. pecten-aboriginum (S. Arias 986; T. Terrazas 415, 419a, b, 478) (Figs. 2D, 5G). Seed oval, extremely large, 3.35–4.78×2.24–3.44 mm, black-brown, glossy, periphery slightly keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries inconspicuous, but slightly channelled around peripheral ridges, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR medium size, 1.34–2.27 mm, oblique, 31–48°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. pringlei (A. Orduño 3, 4; R. Felger 13337) (Fig. 3C). Seed broadly oval, very large, 2.69–3.84×1.86–2.94 mm, black, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region elongate-polygonal, anticlinal boundaries slightly raised, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR small, 0.78–1.6 mm, oblique, 30–69°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. schottii (S. Arias 1319, 1341, 1345, 1347, 1349a) (chapa). Seed broadly oval, large, 1.8–2.43×1.14–1.94 mm, black to black-brown, glossy, periphery keeled; border highly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted; uncommon on lateral region, relief flat on lateral region; slightly convex on dorsal and apical ridges; convexities low domes; micro-relief minutely rugose; HMR small, 0.49–1.12 mm, oblique, 38–68°, deeply impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. tepamo (S. Arias 1333; S. Gama 107a, b; U. Guzmán-Cruz 906a, b) (Figs. 2H, 4F, 5E). Seed broadly oval, very large, 3.27–4.13×2.24–3.07 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region elongate-polygonal, anticlinal boundaries raised, and incomplete channelled, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR medium-sized, 1.27–2.16 mm, oblique, 39–65°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • P. weberi (S. Gama 70, 100, 103; T. Terrazas 518, 519) (Figs. 2E, 5F). Seed oval, large, 2.44–3.55×1.58–2.37 mm, black, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric and elongate-polygonal, anticlinal boundaries slightly raised, straight; interstices minutely pitted, uncommon on lateral region; relief flat; convexities none; micro-relief minutely rugose; HMR small, 0.72–1.5 mm, oblique, 41–75°, finely impressed, micropyle and hilum conjunct but separated by sclerified tissue, oval.

  • Stenocereus aragonii (W.R. Maxon 7858) (Figs. 3E, 4G). Seed broadly oval, extremely large, 3.26–6.24×2.15–4.4 mm, black-brown to brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, uncommon on lateral region; relief par-convex on lateral region; convexities low domes; micro-relief none; HMR medium size, 1.18–2.53 mm, oblique, 28–60°, superficial, conjunct but separated by sclerified band, oval.

  • S. eichlamii (S. Arias 1329) (Figs. 3F, 4H, 5H). Seed broadly oval, very large, 3.52–4.2×2.27–3.04 mm, black-brown, glossy, periphery keeled; border slightly expanded around hilum; cells gradually smaller toward hilum, in lateral region isodiametric-polygonal, anticlinal boundaries channelled, straight; interstices minutely pitted, uncommon on lateral region; relief par-convex on lateral region; convexities low domes; micro-relief minutely rugose; HMR medium size, 1.22–1.78 mm, oblique, 24–56°, superficial, micropyle and hilum conjunct but separated by sclerified tissue, oval.

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Arias, S., Terrazas, T. Seed morphology and variation in the genus Pachycereus (Cactaceae). J Plant Res 117, 277–289 (2004). https://doi.org/10.1007/s10265-004-0156-4

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