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Crystals in calyces of Lamiaceae and their phylogenetic and adaptive significance

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Abstract

The diversity of crystal types is studied in fruiting calyces of 435 out of the c. 7,200 species, and 184 out of the c. 236 genera of Lamiaceae, mainly in the subfamilies Nepetoideae and Lamioideae. Calcium oxalate crystals are found in about half of the studied calyces, and belong to the following main types: prismatics in the inner epidermis, prismatics in fibres, prismatics in sclereids, prismatics in the mesophyll and druses in the mesophyll. Presence of epidermal prismatics may constitute a synapomorphy of the subfamily Nepetoideae or of a subclade within this subfamily consisting of the two largest tribes Mentheae and Ocimeae. The presence of narrow mesophyll prismatics seems to support a large clade of Lamioideae genera in a recent molecular phylogeny of the subfamily. Epidermal prismatics are found to be particularly frequent in closed calyces, calyces with a narrow mouth and internally hairy calyces. These findings and the position of the crystals in the calyces are regarded as supporting the hypothesis that calcium oxalate crystals protect the fruit against insect predators.

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Acknowledgments

I thank the directors, curators and other staff of the herbaria of E, K and UPS for allowing me to sample or sending me calyces of some species. I am also indebted to Dr. Ruth Nielsen and Prof. Øjvind Moestrup for helping me in the use of the digital microscope camera, and Christian Lange for the photo of the calyx of Salvia verticillata.

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Correspondence to O. Ryding.

Appendix

Appendix

The species identifications and vouchers of some of the studied species are given in this list. The data on the other species are given in Ryding (2007). Order of the data: species name, author of the species, voucher specimen, herbarium abbreviation within brackets when the herbarium is another than C. Acanthomintha ilicifolia A. Gray; Elmer 4538. Achyrospermum carvalhi Gürke, J. Hansen 123. Aeollanthus rehmannii Gürke; Ryding 2377. A. repens Oliv.; Saintenoy 20. A. subacaulis (Baker) Hua & Briq.; Malaisse & Goetghebeur 106. A. ukamensis Gürke; Vollesen MRC3844. Asterohyptis stellulata (Benth.) Epling; Pringle 11677. Betonica officinalis L.; Gerrans 1607. Blephilia hirsuta (Pursh) Benth.; Bryant 120. Bystropogon origanifolius L’Her.; Hansen 88. Capitanopsis albida (Baker) Hedge; Humbert 13152. Clinopodium coccineum (Nutt. ex Hook.) Kuntze; Lehmann 3628 (or 3618?). C. dalmaticum (Benth.) Bräuchler & Heubl; Papanicolaou 117. C. darwinii (Benth.) Kuntze; O’Donell 3718. C. gilliesii (Benth.) Kuntze; Poulsen s.n. (29.XII.1951). C. gracile (Benth.) Kuntze; Tanaka & Shimada 17775. C. kilimandschari (Gürke) Ryding; Hansen 229. C. menthifolium (Host) Stace; Krendl s.n. (15.VII.1979). C. myrianthum (Baker) Ryding; Blackmore et al. 1419. C. nepeta (L.) Kuntze; Auquier 7751. C. nubigenum (Benth.) Kuntze; Holm-Nielsen & Jeppesen 1462. C. paradoxum (Vatke) Ryding; de Wilde 8220. C. revolutum (Ruiz. & Pav.) Govaerts; Sørensen & Holm 212. C. sericeum (C. Presl ex Benth.) Govaerts; Sørensen & Holm 67. C. serpyllifolium (M. Bieb.) Kuntze; Naftolsky s.n. (1925). C. simense (Benth.) Kuntze; Ryding 2418. C. suaveolens (Sm.) Kuntze; Strid & Papanicolaou 15793. C. uhligii (Gürke) Ryding; Friis et al. 1065. C. umbrosum (Bieb.) Kuntze; Rechinger 43400. C. vulgare L.; Gerrans 1679. C. vulgare; Ryding 2440. Conradina canescens A. Gray; Kral 29951. Cunila angustifolia Benth.; Smith & Reitz 14255. C. spicata Benth.; Hatschbach et al. 72656. Dicerandra densiflora Benth.; D’Arcy 2286. Dracocephalum nutans L.; Bowes Lyon 196. Elsholtzia winitiana Craib; Wu 2746. Endostemon camporus (Gürke) M.R. Ashby; Fries & Fries 38 (UPS). E. gracilis (Benth.) M.R. Ashby; Harvey et al. 48. E. kelleri (Briq.) Ryding ex A.J. Paton et al.; Friis et al. 8773. E. obtusifolius (E. Mey.) N.E. Br.; Vollesen MRC4918. Glechon marifolia Benth.; Hatschbach 52353. G. spathulata Benth.; Pedersen 11954. Haumaniastrum katangense (S. Moore) J. Duvign. & Plancke; Bingham 9587. Hedeoma hispida Pursh; Ahles 75616. Hesperozygis rhododon Epling; Silva & Cruz 2405. Hymenocrater bituminosus Fisch. & C.A. Mey; Belianina & Proskuriakova 189. Hypenia salzmannii (Benth.) Harley; Hatschbach et al. 67698. Hyptidendron asperrimum (Spreng.) Harley; Hatschbach 26975. Hyptis floribunda Briq.; Schinini & Mroginski 4647. H. macrostachys Benth.; Hatschbach 47495. Hyssopus seravschanicus (Dubj.) Pasij; Kudrjaschev 6796a. Killickia pilosa (Benth.) Brächler et al.; Schlechter 6808. Lallemantia peltata (L.) Fisch. & Mey; Bot. Have 1980-07-30. Lavandula coronopifolia Poir.; Gilbert et al. 7390. L. dentata L.; Pedersen s.n. (IV.1968). L. lanata Boiss.; St.-Lager s.n. (23.VI.1908). L. latifolia Medik.; Lange s.n. L. pubescens Decne; Hein 96-143. L. setifera T. Anderson; Schweinfurth 135. L. stoechas L.; Hansen s.n. Leocus membranaceus J.K. Morton; Jacques-Georges 12174. Lophanthus chinensis Benth.; Saposchnibov s.n. (1914). Lycopus uniflorus Michx.; Schaeffer 62471. Marsypianthes foliolosa Benth.; Hatschbach & Barbosa 59563. Melissa axillaris (Benth.) Bakh. f.; Herb. Griffith 3982. M. officinalis L.; wild plant in the Botanical Garden in Copenhagen. Mentha arvensis L.; Grøntved 875. M. suaveolens Erhr.; Hansen s.n. Micromeria pineolens Svent.; Hansen s.n. Moluccella aucheri (Boiss.) A.-C. Scheen; Popov 123. Monarda fistulosa L.; Fork 3133. M. punctata L.; Leonard 5696. Nepeta racemosa Lam.; cultivated in Botanical garden in Copenhagen s.n. N. spruneri Boiss.; Baden 1078. Ocimum americanum L.; Friis et al. 9480. O. carnosum (Spreng.) Benth.; Sehnem 3910. O. filamentosum Forssk.; Kuchar 17578. O. gratissimum L.; Segawa 15. O. jamesii Sebald; Hansen et al. 6462. O. lamiifolium Benth.; Friis et al. 8819. O. obovatum Benth.; Schrire 57. O. spectabile (Gürke) A.J. Paton; Gilbert et al. 8015. O. spicatum Deflers; Friis et al. 8594. Origanum majorana L.; Jansen 2439. O. sipyleum L.; Davis 18360. Orthosiphon aristatus (Blume) Miq.; Vesterdal 4E. Otostegia modesta S. Moore; Hansen & Heemstra 6425. Peltodon tomentosum Pohl; Lund s.n. Perillula reptans Maxim.; Taquet 1498. Platostoma cororatum (D. Don) A.J. Paton; Bunchuai 1424. P. denticulatum Robyns; Bridson 168. P. hildebrandtii (Vatke) A.J. Paton; Abdallah & Vollesen 96/236. P. intermedium A.J. Paton; Geesink et al. 7070. P. madascariensis (Benth.) A.J. Paton & Hedge; Decary 61/7. P. menthoides (L.) A.J. Paton; Børgesen 275. Plectranthus guerkei Briq.; Jacques-Georges 17563. P. lanuginosus (Benth.) Agnew; Friis et al. 1330. P. laxiflorus Benth.; Friis et al. 1505. P. longipes Baker; Kalema 1752. P. montanus Benth.; Phillipson 4841. P. otostegioides (Gürke) Ryding; Friis & al. 9583. P. prostratus Gürke; Ryding 2095. P. orthodonta Gürke; Kaunda & Balaka 31. Pogogyne serpylloides (Torr.) A. Gray; Abrams 5100. Poliomintha longifolia A. Gray; Palmer 203. Pseudocaryopteris bicolor (Roxb. ex Hardw.) P.D. Cantino; Wallich s.n. Rosmarinus eriocalyx Jord. ex Fourr.; Fernandez Casas s.n. (12.III.1972). Rydingia integrifolia (Benth.) A.-C. Scheen; Gilbert & Getachew 2309. R. limbata (Benth.) Scheen & V.A. Albert; Falconer s.n. Salvia amplexicaulis Lam.; Strid 36567. S. azurea Mich. ex Vahl; Rose 17159. S. ballotiflora Benth.; Lindheimer 1087. S. clevelandii (A. Gray) Greene; Schallert 28850. S. coccinea Etl.; Pedersen 7480. S. columbariae Benth.; Bot. Garden Copenhagen s.n. S. farinacea Benth.; Schallert 563. S. fruticosa Mill.; Kaae s.n. S. glutinosa L.; cultivated in Copenhagen Bot. Gard., from A. Strid. S. greggii A. Gray; Sörensen 7348. S. hirsuta Jacq.; Pringle 8456. S. involucrata Cav.; Moore 5057. S. lavandulifolia Vahl; Thornberg s.n. S. merjamie Forssk.; Ryding 2053. S. microphylla Kunth; cultivated in Copenhagen Bot. Gard. S. officinalis L.; cult in Copenhagen Bot. Garden s.n. S. pachyphylla Epling ex Muny; Wiggins & Demaree 4892. S. subincisa Benth.; Pringle 13603. S. taraxacifolia Coss. ex Balansa; Lewalle 10543. S. verticillata L.; Strid 977. Satureja montana L.; Borza 596. Schizonepeta annua (Pall.) Schischk.; Kridov s.n. (1910). Siphocranion macranthum (Hook. f.) C.Y. Wu; Liljestrand 137 (UPS). Syncolostemon parviflorus Benth.; van Hoepen 1998. Tetradenia urticifolia (Baker) Phillipson; Phillipson 5017. Thymus vulgaris L.; Lopez & Morales 2633. Ziziphora clinopodioides Lam.; Klements s.n. Z. hispanica L.; Reading Univ./BM Exped. 509.

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Ryding, O. Crystals in calyces of Lamiaceae and their phylogenetic and adaptive significance. Plant Syst Evol 290, 201–215 (2010). https://doi.org/10.1007/s00606-010-0361-7

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