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Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae)

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Abstract

Pericarp structure is investigated in 143 specimens belonging to 59 out of ca. 65 genera and 134 out of ca. 2,000 species of Lamiaceae subfamily Nepetoideae tribe Mentheae. Some of the data presented by earlier authors are believed to be incorrect. The variation in pericarp structure has been found to be strongly correlated to the variations in DNA and gross morphological characters. The groups of two-staminate genera (Salvia, etc.) and four-staminate genera of Salviinae (Lepechinia and Chaunostoma) can be distinguished from each other and from other labiates by differences in pericarp structure. The other two subtribes, Menthinae and Nepetinae, differ considerably, but their variations strongly overlap. The differences in pericarp structure also suggest that Menthinae can be divided into three main monophyletic groups based on other data: (1) the main bulk of the tribe, (2) a group of Prunella and Cleonia and perhaps also Horminum, and (3) the characteristic genus Lycopus. Pericarp data disagree with a suggestion that the genus Melissa should be included in the subtribe Salviinae. Modifications of the subtribal classification of Mentheae are discussed.

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Acknowledgments

I wish to thank the curators at LD and UPS for allowing me to sample nutlets from their specimens.

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Authors and Affiliations

  1. The Botanical Garden & Museum, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark

    O. Ryding

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  1. O. Ryding
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Correspondence to O. Ryding.

Appendix

Appendix

Voucher specimens. When two or more specimens have been studied in a species, they are enumerated in the same order as in Table 1. Acanthomintha ilicifolia A.Gray—USA, California, Lewis & Rose 32227 (UPS). Agastache foeniculum (Pursh) Kuntze—cult. Copenhagen Botanical Garden S1972-0260 (C). Agastache micrantha (A. Gray) Wooton & Standl.—Mexico, Bye 5719 (C). Agastache nepetoides (L.) Kuntze—USA, Vermont, Zika 6595 (C). Agastache rugosa (Fisch. & C.A. Mey.) Kuntze—Japan, Togasi 1583 (C). Agastache scrophularifolia (Willd.) Kuntze—USA, Illinois, Kline 22 (C); USA, Virginia, Fosberg 36019 (C); cult, garden in Copenhagen (C). Agastache urticifolia (Benth.) Kuntze—USA, Washington, Tureson s.n. (UPS). Blephilia ciliata (L.) Benth.—USA, Ohio, Herb. Tuckerm (UPS). Blephilia hirsuta (Pursh) Benth.—USA, Illinois, Underwood s.n. (IX.1882)(C). Bystropogon canariensis (L.) L’Hér.—Canary Islands, Teneriffa, Tullgren s.n. (UPS). Bystropogon maderensis Webb. & Berthel.—Madeira, Hansen s.n. (26.VIII.1970) (C). Cedronella canariensis (L.) Webb & Bertel.—Madeira, O.Dahlgren s.n. (UPS). Chaunostoma mecistandrum J.D. Sm.—Mexico, Matuda 3915 (GH). Cleonia lusitanica (L.) L.—Algeria, Faure 8 (UPS). Clinopodium acinos (L.) Kuntze—?, Ralska s.n. (9.VIII.1956)(C). Clinopodium alpinum (L.) Kuntze—Austria, Ryding 961 (UPS). Clinopodium bolivianum (Benth.) Kuntze—Bolivia, Steinbach 8458 (S). Clinopodium nepeta (L.) Kuntze—France, Rastetter 7751 (C). Clinopodium taxifolium (Kunth) Govaerts—Ecuador, Asplund 17961 (UPS). Clinopodium vulgare L.—Bulgaria, Frost-Olsen 1052 (C). Clinopodium. coccineum (Hook.) Kuntze—Brazil, Rio de Janeiro State, Glaziou 6663 (C). Conradina canescens A.Gray—USA, Moldenke 1133 (K). Conradina grandiflora Small—USA, Schallert 5349 (LD). Cuminia eriantha (Benth.) Benth. var. fernandezia (Colla) Harley—Chile, Juan Fernandez Islands, Bertero 1490 (S). Cunila fasciculata Benth.—Brazil, Sello s.n. (UPS). Cunila incisa Benth.—Brazil, Dusén 17818 (S). Cunila origanoides (L.) Britton—USA, New Jersey, A. Gray s.n. (UPS). Cyclotrichium origanifolium (Labill.) Manden & Scheng.—Turkey, Davis 14144 (C). Cyclotrichium stamineum (Boiss. & Hohen.) Manden. & Scheng.—Iraq, Kurdistan, Kotschy 311 (S). Dicerandra densiflora Benth.—USA, Florida, D’Arcy 2286 (C). Dicerandra frutescens Shinners—USA, Florida, Tehler et al. 110 (S). Dorystaechas hastata Benth.—Turkey, Davis 13948 (K). Dracocephalum moldavica L.—Russia, illegible (C Dracocephalum ruyschiana L.—Finland, Grøntved 50 (C). Drepanocaryum sewerzowii (Regel) Pojark—Uzbekistan, Knorring 99 (S). Glechoma hirsuta Waldst. & Kit.– Romania, Arvat et al. 2037 (C). Glechon ciliata Benth.—Brazil, Regnell s.n. (S). Glechon marifolia Benth—Brazil, Hatschbach 52353 (C). Gontscharovia popovii (B. Fedtsch. & Gontsch.) Boriss.—Tadshikistan, Vvedensky 6795 (C). Hedeoma dentata Torr.—USA, Arizona, Gooding 2408 (C). Hedeoma drummondii Benth.—Mexico, Pringle 13740 (C). Hedeoma mollis Torr.—USA, Texas, Havard s.n. (UPS). Hesperozygis myrtoides (Benth.) Epling—Brazil, Rio de Janeiro State, Dusén s.n. (S). Hesperozygis rhododon Epling—Brazil, Hatschbach 2405 (C Heterolamium debile (Hemsl.) C.Y. Wu—East Asia, Wilson 4348 (K). Hoehnea epilobioides (Epling) Epling—Paraguay, Hassler 6847 (S). Hoehnea parvula (Epling) Epling—Brazil, Paraná, Hatschbach 40439 (C). Horminum pyrenaicum L.—Italy, Friis s.n. (5.VIII.1968)(C); Pyrenean Mts., Böcher & Larsen 195 (C); Austria, St-Lager s.n. (13.VIII.1890)(C); Italy, Thornberg s.n. (15.VIII.1968)(C). Hymenocrater bituminosus Fisch. & C.A. Mey.—Iran, Rechinger 312 (S). Hyssopus officinalis L.—Romania, Topa 2891 (C). Kurzamra pulchella (D. Clos.) Kunze—Chile, Atacama Province, Werdermann 957 (S). Lallemantia iberica (M. Bieb.) Fisch. & C.A. Mey.—Turkey, Davis & Hedge 29183 (C). Lallemantia royleana (Benth.) Benth.—Turcmenistan, Frantskevich 723 (C). Lepechinia betonicifolia (Lam.) Epling—Ecuador, Asplund 6646 (UPS). Lepechinia calycina (Benth.) Epling—USA, California, Clokey 5044 (UPS). Lepechinia chamaedryoides (Balb.) Epling—Chile, Hohenacker 707 (UPS). Lepechinia floribunda (Benth.) Epling—Argentina, Krapovickas & Vanni 39499 (C). Lepechinia mutica (Benth.) Epling—Ecuador, Sparre 16111 (UPS). Lepechinia salviifolia (Kunth) Epling—Colombia, Alonso & Jaramillo 8209 (C). Lepechinia vesiculosa (Benth.) Epling—Argentina, Tucum n Prov., Meyer 13988 (UPS). Lophanthus tschimganicus Lipsky—Baranov & Raikova 6426 (C). Lycopus europaeus L.—Turkey, Davis 13913 (C). Marmoritis pharica (Prain) A.L. Budantzev—Nepal, Stainton & al. 7255 (E). Melissa axilaris (Benth.) Bakh. f.—India or Bangladesh, Herb. Griffith 3982 (C). Melissa officinalis L.—Bosnia-Herzegovina, Kaae s.n. (5.X.1967) (C). Mentha aquatica L.—Bulgaria, Vyhodcevski 714 (C). Mentha arvensis L.—? (UPS). Mentha cervina L.—?, Wiik s.n. (UPS); France, Avignon, collector unknown (LD). Mentha longifolia (L.) L.—Canary Islands, A. Hansen s.n. (22.X.1970) (C). Mentha spicata L.—Belgium, Lambinon 11565 (C). Meriandra dianthera (Roem. & Schult.) Briq.—Eritrea, Pappi 1538 (S). Micromeria imbricata (Forssk.) C. Chr.—Ethiopia, Friis et al. 486 (C). Micromeria juliana (L.) Benth. ex Rchb.—Greece, Corfu Island, Ryding 2297 (UPS). Minthostachys mollis (Benth.) Griseb.—Argentina, Cordoba Prov., Hieronymus 5189 (UPS). Minthostachys mollis (Benth.) Griseb.—Ecuador, Asplund 7200 (UPS). Minthostachys verticillata (Griseb.) Epling—Argentina, Tucuman Prov., Risso 581 (UPS). Monarda bradburiana Beck—USA, Pammel 17/88 (C). M. citriodora Cerv. ex Lag.—USA, Albert Ruth 1320 (C). M. clinopodia L.—USA, Schallert 372 (C). M. clinopodioides A. Gray—USA, Whitehouse 16224 (BRIT). M. didyma L.—Cult. in Copenhagen. M. fistulosa L.—USA, Bro Arsène 17679 (C). M. lindheimeri A. Gray—USA, Fryxell 1011 (BRIT). M. pectinata Nutt.—USA, Higgins 4375 (MICH). M. punctata L.—USA, Leonard & Leonard 5696 (C). M. russeliana Nutt. ex Sims—USA, Cory 23185 (MICH); USA, Demaree 35455 (BRIT). Monardella odoratissima Benth.—USA, Oregon, T.Howell s.n. (UPS). Nepeta azurea Benth.—Eritrea, Ryding 2054 (UPS). Nepeta cyanea Steven—Iran, Skofitz s.n. (C). Nepeta nepetella L.—Spain, Leadlay & Petty 86 (C). Nepeta nepetoides (Batt. ex Pit.) Harley—Morocco, Jahandiez 45 (C). Nepeta parnassica Heldr. & Sart.—Greece, Baden 999 (C). Nepeta tuberosa L.—Portugal, Kaae s.n. (27.IX.1968) (C). Origanum onites L.—Turkey, Heldreich 1465 (UPS). Origanum scaber Boiss. & Heldr.) Briq.—Greece, Orphanides 42 (C). Perovskia abrotanoides Kar.—India, Herb. India Orient., Hook.f. & Thomson s.n. (S). Perovskia scrophulariifolia Bunge—cult. in USA, Copenhagen Botanical Garden (C). Piloblephis rigida (Benth.) Raf.—USA, Florida, Divers 4 (K). Pogogyne douglasi Benth.—USA, California, Skottsberg s.n. (UPS). Pogogyne serpylloides (Torr.) A. Gray—USA, California, Gray s.n. (UPS). Pogogyne ziziphoroides Benth.—USA, California, Rose 33191 (UPS). Poliomintha incana (Torr.) A. Gray—USA, Utah, Holmgren et al. 2043 (S). Prunella vulgaris L. subsp. hispida (Benth.) Hultén—Himalaya, Duthie s.n. (16.VIII.1899) (C). Pycnanthemum flexuosum (Walter) Britton—Canada, Ontario, Breitung & Dore 9296 (UPS). Pycnanthemum incanum (L.) Michx.—USA, Washington D.C., illegible (UPS). Rhabdocaulon denudatum (Benth.) Epling—Brazil, Minas Gerais State, Widgren s.n. (UPS). Rhabdocaulon lavanduloides (Benth.) Epling—Brazil, Minas Gerais State, Regnell I/325 (UPS). Rhododon ciliatus (Benth.) Epling—USA, Hall 443 (K). Rosmarinus eriocalyx Jord. & Fourr.—Spain, Fernández Casas s.n. (12.III.1972)(C). R. officinalis L.—Portugal, Ryding 428 (UPS). Saccocalyx satureioides Coss. & Durand—Algeria, illegible (UPS). Salvia aegyptiaca L.—Morocco, Ryding 2404 (C). Salvia azurea Michx. ex Vahl—USA, Kansas, Rose 17159 (C). Salvia canariensis L.—Canary Islands, Anker Petersen s.n. (VI.1965)(C). Salvia coccinea Etl.—Argentina, Pedersen 7480 (C). Salvia farinacea Benth.—cult., native in USA, Texas, Schallert 563 (C). Salvia glutinosa L.—Austria, Vitek 98-1808 (C). Salvia officinalis L.—? (UPS). Salvia sonomensis Greene—USA, California, Hall & Essig 10184 (C). Salvia splendens Sellow—Cuba, Van Hermann 2682 (C). Salvia taraxacifolia Coss. & Balansa—Morocco, Lewalle 10583 (C). Salvia verbascifolia M. Bieb.—Turkey, Davis & Hedge 29498 (C). Satureja hortensis L.—Albania, Kit Tan 24620 (C). Satureja montana L.—Spain, Font Quer s.n. (24.IX.1949) (C). Satureja thymbra L.—Greece, Pedersen s.n. (IV.1979) (C). Schizonepeta tenuifolia (Benth.) Briq.—Japan, Togasi 1519 (UPS). Stachydeoma graveolens (A. Gray) Small—USA, Florida, Godfrey 62494 (UPS). Thymbra capitata (L.) Cav.—Greece, Kaae s.n. (26.IX.1973) (C). Thymbra spicata L.—Iran, Balansa 330 (C); Turkey, Wängsjö 2723 (LD). Thymus serpyllum L.—Sweden, Almquist s.n. (VII.1866) (C). Zhumeria majdae Rech.f. & Wendelbo—Iran, Wendelbo & Foroughi 15731 (E). Ziziphora capitata L.—Iran, Rechinger 1189 (C). Ziziphora hispanica L.—Spain, Bourgeau 1419/2 (C). Ziziphora taurica M. Bieb.—Greece, Kapetanidis s.n. (C). Ziziphora. tenior L.—Turkmenistan, Nikitin s.n. (C); Iran, Køie 278 (C).

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Ryding, O. Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae). Plant Syst Evol 285, 165–175 (2010). https://doi.org/10.1007/s00606-010-0270-9

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