Volumes published in this series:
Volume I Pteridophytes and Gymnosperms The Families Edited by K.U. Kramer and P.S. Green (1990) Date of publication: 28.9.1990
Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families and Genera Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993) Date of publication: 28.7.1993 of Vascular Plants Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae) Edited by K. Kubitzki (1998) Date of publication: 27.8.1998 Edited by K. Kubitzki
Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae (except Gramineae) Edited by K. Kubitzki (1998) Date of publication: 27.8.1998 Flowering Plants • Dicotyledons Volume V Flowering Plants. Dicotyledons: Malvales, Capparales and Non-betalain Caryophyllales VII Lamiales (except Acanthaceae including Avicenniaceae) Edited by K. Kubitzki and C. Bayer (2003) Date of publication: 12.9.2002 Volume Editor: Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales Edited by K. Kubitzki (2004) J.W. Kadereit Date of publication: 21.1.2004
Volume VIIFlowering Plants. Dicotyledons: Lamiales (except Acanthaceae including Avicenniaceae) Edited by J.W. Kadereit (2004) With 60 Figures, 8 in Color
Springer Professor Dr. Klaus Kubitzki Preface Universitat Hamburg Biozentrum Klein-Flottbek und Botanischer Garten Ohnhorststrafie 18 22609 Hamburg Germany
Professor Dr. Joachim W. Kadereit Johannes Gutenberg-Universitat Mainz Institut fur Spezielle Botanik und Botanischer Garten This volume, the seventh in this series, deals with the families of the Lamiales, a group of 55099 Mainz the asterids or Sympetalae, which was recognised by early workers such as Bartling and Germany Lindley under the name Labiatiflorae or Personatae, and which has only moderately been modified in circumscription over time. Nevertheless, our considerably increased knowl edge of the group and the results of molecular studies provide a more detailed basis for understanding relationships among and evolution within the families of the Lamiales, although, as J.W. Kadereit emphasises inhis introduction, in this connection and also in relation to family delimitation, many questions remain currently unanswered. My sincere thanks go to all authors for their scholarly treatment of families in this volume. In particular, the authors/organisers who handled the four large families with more than 100 genera, Bignoniaceae, Gesneriaceae, Labiatae and Scrophulariaceae, deserve special mention for their herculean work. The authors also emphasised the various themes of diversification, differing from group to group, that are so important for the understanding and interpretation of taxonomic diversity, such as phytochemical diversification in Labiatae and pollinator interactions in Gesneriaceae, to cite only two salient examples. In the circumscription of Scrophulariaceae, a pragmatic position has been taken; since this series aims primarily at providing baseline information for fami lies and genera, the problem of a phylogenetically satisfactory classification of the Scro phulariaceae complex that avoids paraphyly at our present state of knowledge must be left undecided, and indeed is not the primary focus of this volume. Sincere thanks are also extended to Professor J.W. Kadereit for patiently bringing his work on the present ISBN 3-540-40593-3 Springer-Verlag Berlin Heidelberg New York volume to an end, in spite of all discouraging vicissitudes. I am most grateful to all copyright holders for so generously authorising the use of library of Congress Cataloging-in-Publication Data their precious illustrative material. Finally, it is a pleasure to thank, also in the name of Professor Kadereit, the staff of Springer-Verlag and ProEdit, Heidelberg, for their dedi Flowering plants, dicotyledons : Lamiales (except Acanthaceae including Avicenniaceae) / volume editor, J.W. Kadereit. cated work on the present volume. I would like to express special thanks to the copy editor, p. cm. - (The families and genera of vascular plants ; 7) Includes bibliographical references. Monique Delafontaine, for the many improvements she made to the manuscript. ISBN 3-540-40593-3 (alk. paper) 1. Lamiales - Classification. I. Kadereit, J.W. (Joachim W.), 1956- II. Series. Hamburg, December 2003 K. Kubitzki QK495.A12F57 2004 583'.96 - dc22 2003065479
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Lamiales: Introduction and Conspectus J.W.Kadereit 1
Bignoniaceae E. Fischer, I. Theisen and L.G. Lohmann 9 Buddlejaceae B. Oxelman, P. Kornhall and E.M. Norman 39 Byblidaceae J.G. Conran and R. Carolin 45
Callitrichaceae C. Erbar and P. Leins 50 Carlemanniaceae M.Thiv 57 Cyclocheilaceae Sebsebe Demissew 60
Gesneriaceae A. Weber 63 Globulariaceae G.Wagenitz 159
Hippuridaceae P. Leins and C. Erbar 163
Labiatae R.M. Harley, S. Atkins, A.L. Budantsev, P.D. Cantino, B.J. Conn, R. Grayer, M.M. Harley, R. de Kok, T. Krestovskaja, R. Morales, A.J. Paton, O. Ryding and T. Upson 167 Lentibulariaceae E. Fischer, W. Barthlott, R. Seine and I. Theisen 276
Martyniaceae H.-D. Ihlenfeldt 283 Myoporaceae I. Theisen and E. Fischer 289
Nesogenaceae R.M. Harley 293
Oleaceae P.S. Green 296
Pedaliaceae H.-D. Ihlenfeldt 307 Phrymaceae P.D. Cantino 323 Plantaginaceae A.E. Schwarzbach 327 Plocospermataceae L. Struwe and S.R. Jensen 330
Scrophulariaceae E. Fischer 333 Stilbaceae H.P. Linder 433
Tetrachondraceae S.J.Wagstaff 441 Trapellaceae H.-D. Ihlenfeldt 445
Verbenaceae S.Atkins 449
Index to Scientific Names 469 List of Contributors IX
List ofContributors Leins, P. Heidelberger Institut fur Pflanzenwissenschaften (HIP) - Biodiversitat und Pflanzensystematik, Im Neuenheimer Feld 345,69120 Heidelberg, Germany
LlNDER, H.P. Institute for Systematic Botany, University of Zurich, Zollikerstrasse 107,8008 Zurich, Switzerland
LOHMANN, L.G. Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 64507, USA
Atkins, S. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK Morales, R. Herbario, Real Jardin Botanico, Plaza de Murillo 2, Madrid 28014, Spain Barthlott, W. Botanisches Institut und Botanischer Garten der Rheinischen Friedrich-Wilhelms-Universitat, Meckenheimer Allee 170, Norman, E.M. Department of Biology, Stetson University, Deland, 53115 Bonn, Germany FL 32723, USA
Budantsev, A.L. Komarov Botanical Institute, Herbarium, Prof. Popov Street, Oxelman, B. Department ofSystematic Botany, Evolutionary Biology St. Petersburg 197376, Russia Centre Uppsala University, Norbyv. 18D, 75236 Uppsala, Sweden Cantino, P.D. Department of Environmental and Plant Biology, Ohio University, Athens, OH 45701-2979, USA Paton, A.J. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK
Carolin, R. Pulman's Cottage, 30 Pulman Street, Berry, N.S.W. 2535, Ryding, O. Botanical Museum, University of Copenhagen, Australia Gothersgade 130,1123 Copenhagen K, Denmark Department of Biological Sciences, Kent State University, Conn, BJ. Royal Botanic Garden, Sydney, Australia schwarzbach, a.e. Kent, OH 44242, USA Conran, J.G. Environmental Biology, School of Earth and Environmental The National Herbarium, Science Faculty, Addis Ababa Sciences, The University of Adelaide, SA 5005,Australia Sebsebe Demissew University, P.O. Box 3434, Addis Ababa, Ethiopia Erbar, C. Heidelberger Institut fur Pflanzenwissenschaften (HIP) - Seine, R. European Astronaut Centre, Linder Hohe, Biodiversitat und Pflanzensystematik, 51147 Koln, Germany Im Neuenheimer Feld 345,69120 Heidelberg, Germany Struwe, L. Department of Ecology, Evolution, and Natural Resources, Fischer, E. Institut fur Biologie, Universitat Koblenz/Landau, Rutgers University - Cook College, 237 Foran Hall, 59 Dudley Universitatsstr. 1,56070 Koblenz, Germany Road, New Brunswick, NJ 08901-8551, USA Grayer, R. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK Theisen, I. Botanisches Institut und Botanischer Garten der Rheinischen Green, P.S. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK Friedrich-Wilhelms-Universitat, Meckenheimer Allee 170, 53115 Bonn, Germany Ihlenfeldt, H.-D. Ostseestr. 40,24369 Waabs, Germany Thiv,M. Staatl. Museum fur Naturkunde, Rosenstein 1, Harley, M.M. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK 70191 Stuttgart, Germany
Harley, R.M. Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK Upson, T. Cambridge University Botanic Garden, Cory Lodge, Bateman Jensen, S.R. Department of Chemistry, The Technical University of Street, Cambridge CB2 1JF, UK Denmark, 2800 Lyngby, Denmark Wagenitz, G. A. v. Haller-Institut, Abteilung Systematische Botanik, Kadereit, J.W. Institut fur Spezielle Botanik und Botanischer Garten, Untere Karspule 2,37073 Gottingen, Germany Johannes Gutenberg-Universitat Mainz, 55099 Mainz, Wagstaff, S.J. Allan Herbarium, Landcare Research, P.O. Box 69, Germany Lincoln 8152, New Zealand
Kok, R. de Royal Botanic Gardens Kew, Richmond, Surrey TW9 3AB, UK Weber, A. Institut fur Botanik, Universitat Wien, Rennweg 14, 1030 Wien, Austria KORNHALL, P. Department of Systematic Botany, Evolutionary Biology Centre Uppsala University, Norbyv. 18D, 752 36 Uppsala, Sweden
Krestovskja, T. Komarov Botanical Institute, Herbarium, Prof. Popov Street, St. Petersburg 197376, Russia
KUBITZKI, K. Institut fur Allgemeine Botanik, Universitat Hamburg, Ohnhorststr. 18,22609 Hamburg, Germany Labiatae 167 166 P. Leins and C. Erbar
Selected Bibliography Metcalfe, C.R., Chalk, L. 1957. Anatomy of the dicotyledons II. Labiatae Oxford: Clarendon Press. Moore, R.J. (ed.) 1973. Index to plant chromosome numbers R.M. Harley,1 S. Atkins, A.L. Budantsev, P.D. Cantino, B.J. Conn, R. Grayer, M.M. Harley, Behnke, H.D. 1991. Distribution and evolution of forms and 1967-1971. Regnum Vegetabile 90. types of sieve-element plastids in the dicotyledons. Aliso 13: R. de Kok, T. Krestovskaja, R. Morales, A.J. Paton, O. Ryding and T. Upson Olmstead, R.G., Reeves, P.A. 1995. Evidence for the polyphyly 167-182. of the Scrophulariaceae based on chloroplast rbcL and ndhF Cronquist, A. 1981. An integrated system of classification of sequences. Ann. Missouri Bot. Gard. 82:176-193. flowering plants. New York: Columbia University Press. Olmstead, R.G., DePamphilis, C.W.,Wolfe, A.D., Young, N.D., Davis, G.L. 1966. Systematic embryology of the angiosperms. Elisons, W.J., Reeves, P.A. 2001. Disintegration of the Scro New York: Wiley. phulariaceae. Am. J. Bot. 88: 348-361. De Candolle, A.P. 1828. Prodromus Systematis Naturalis Regni Reeves, P.A., Olmstead, R.G. 1998. Evolution of novel morpho Vegetabilis III. Paris: Treuttel & Wiirtz, pp. 65-72. Labiatae Adans., Fam. Pi. 2: 180 (1763). annulus of hairs or appendaged within, rarely logical and reproductive traits in a clade containing Antir Erdtman, G. 1966. Pollen morphology and plant taxonomy. Lamiaceae Martynov (1820), nom. alt. corolla resupinate. Stamens attached within rhinum majus (Scrophulariaceae). Am. J. Bot. 85:1047-1056. New York: Hafher Publ. Co. Reveal, J.L., Judd, W.S., Olmstead, R.G. 1999. Proposal to con corolla-tube, usually 4 or 2 by abortion and then Federov, ALA. (ed.) 1969. Chromosome numbers of flowering serve the name Antirrhinaceae against Plantaginaceae Trees, shrubs, subshrubs or perennial or annual staminodes often present, or stamens 5-8(-16), plants. Leningrad: Nauka. (Magnoliophyta). Taxon 48:182. herbs, rarely climbers, aromatic or not. Roots Harada, 1.1952. Chromosome studies of some dicotyledonous when 4 often didynamous (rarely a fifth, posterior Rothmaler, W. 1987. Exkursionsflora. Band 3: Atlas der water plants. Jap. J. Genet. 27:117-120. rarely tuberous. Stems often quadrangular, erect to vestigial staminode present), free or rarely mon- Gefafipflanzen, 6. Aufl. Berlin: Volk und Wissen. Hegi, G. 1926: Illustrierte Flora von Mitteleuropa vol. 2. prostrate, sometimes forming stolons or large or odelphic, filaments short or often elongate, usually Savolainen, V., Fay, M.F., Albach, D.C., Backlund, A., van der Miinchen: Lehmann. Bank, M., Cameron, K.M., Johnson, S.A., Lledo, M.D., slender rhizomes. Indumentum usually present, of exserted from corolla-tube and sometimes long- Hegnauer, R. 1967. Chemical characters in plant taxonomy: Pintaud, J.-C, Powell, M., Sheahan, M.C., Soltis, D.E., Soltis, glandular and non-glandular trichomes, often exserted from corolla; parallel, divergent or some possibilities and limitations. Pure Appl. Chem. 14: P.S., Weston, P., Whitten, W.M., Wurdack, K.J., Chase, M.W. hair-like, rarely scale-like, usually multicellular- 173-187. ascending and sometimes included within or lying 2000. Phylogeny of the eudicots: a nearly complete familial Hegnauer, R. 1989. Chemotaxonomie der Pflanzen VIII. Basel: uniseriate, simple, branched, dendroid or stellate, under the posterior corolla-lip, or dedinate and analysis based on rbcL gene sequences. Kew Bull. 55: Birkhauser. sometimes gland-tipped, large-headed subsessile 257-309. then sometimes included within the anterior Juel, 0.1910. Cynomorium und Hippuris. Svensk Bot. Tidskr. 4: Schindler, A.K. 1904. Die Abtrennung der Hippuridaceen von glands rarely absent. corolla-lip, anthers usually dithecous, tetrasporan- 151-159. den Halorrhagaceen. Bot. Jahrb. Syst. 34, Beibl. 77:1-77. Leaves opposite, often decussate, sometimes giate or monothecous by abortion, thecae parallel Juel, 0.1911. Studien iiber die Entwicklung von Hippuris vul- Solereder, H. 1899. Systematische Anatomie der Dicotyledo- whorled, very rarely alternate, simple, entire, garis. Nova Acta Regiae Soc. Sci. Upsal. IV, 2, N. 11: 1-26. or divergent, occasionally widely separated by an nen. Stuttgart: Enke. Leins, P. 2000. Bltite und Frucht. Aspekte der Morphologic toothed or lobed, sometimes compound and then elongate connective, or apically confluent or syn- Takhtajan, A. 1987. Systema Magnoliophytorum (in Russian). Entwicklungsgeschichte, Phylogenie, Funktion und digitate or pinnate, petiolate or sessile, rarely Leningrad: Nauka. thecous, opening by longitudinal slits or rarely by Okologie. Stuttgart: Schweizerbart'sche Verlagsbuchhand- Thorne, R.F. 1968. Synopsisof a putatively phylogenetic clas forming a basal rosette, estipulate. Inflorescence pores. Disc at base of ovary often present, usually lung. sification of the flowering plants. Aliso 6: 57-66. often bracteate, bracts sometimes conspicuous, fleshy, entire or irregularly or often 4-lobed, ante Leins, P., Erbar, C. 1988. Einige Bemerkungen zur Blutenent- Thorne, R.F. 1992. Classification and geography of the wicklung und systematische Stellung der Wasserpflanzen persistent or deciduous, rarely (Lavandula and rior lobe sometimes longer than others, nectari flowering plants. Bot Rev. 58: 225-348. Callitriche, Hippuris und Hydrostachys. Beitr. Biol. Pflanzen Scutellaria) spirally arranged, composed of cymes, ferous. Gynoecium 2-carpellate, often 4-locular by Uphof,J.CTh. 1962. Plant hairs. In: Zimmermann, W, Ozenda, 63: 157-178. bracteolate or not, and often arranged in a termi P.G. (eds.) Encyclopedia of plant anatomy IV, 5, 2nd edn. intrusion of carpel wall forming "false septum", or Leins, P., Erbar, C. 1990. The possible relationship of Berlin-Nikolassee: Borntrager. nal, lax or congested, indeterminate thyrse which rarely imperfectly 2-locular and free towards apex, Hydrostachyaceae based on comparative ontogenetical Wieffering, J.H. 1966. Aucubinartige Glucoside may be paniculate, raceme-like with cymes often flower studies. Mitt. Inst. Allg. Bot Hamburg 23b: 723- ovary usually 4-ovuled, 2-locular ovaries generally (Pseudoindikane) und verwandte Heteroside als systema 729. 1-flowered, or spike-like, or rarely congested into with loculi 2-ovuled, and 4-locular ovaries with 1 tische Merkmale. Phytochemistry 5:1053-1064. a capitulum, with or without a distinct involucre ovule per loculus, ovary entire or lobed, with ter of bracteoles. Flowers hypogynous, usually bisex minal style, or more often deeply 4-lobed, the ual, or less often unisexual due to gynodioecy or loculi often separated and with style gynobasic, gynomonoecy, very rarely due to dioecy. Perianth style usually with 2 equal or unequal stigma-lobes, biseriate, sepals 4-5(-9), connate, actinomorphic rarely entire with 1 stigma-lobe vestigial, or to zygomorphic, sometimes 2-lipped, lobes stigma capitate or very rarely 4-lobed. Ovules 2-many, often 5, equalor unequal, rarely obsolete, anatropous to hemianatropous, usually basal or some lobes often fused, or lips entire, calyx-tube sub-basal, erect, rarely orthotropous, apical, (5-)10-15-nerved, straight or curved, throat hairy pendulous, borne laterally or submarginally on the or glabrous, calyx often accrescent, rarely inflated placenta, unitegmic, tenuinucellate. Fruit drupa or fleshy in fruit. Petals (4-)5(-16) connate, actin ceous, often with pyrenes, or dry, indehiscent, or omorphic to more often slightly to strongly zygo separating into two 2-seeded or frequently, four 1- morphic, often 2-lipped, rarely 1-lipped, lobes seeded mericarps, sometimes fewer by abortion. (2-)4-5(-16), equal or unequal, porrect to patent, Mericarps (nutlets) often with sculptured, tuber- one or other lip often concave to galeate, corolla- culate, hairyor rarely winged pericarp, mucilage tube short to elongate, rarely spurred, often with cells often present. Seeds albuminous or exalbu- minous, epigeal. Embryo straight or bent, invest 1 For details of authorship see p. 275. ing or spatulate. 2 Inthe generic descriptions, calyx and corolla-lobe numbers The family contains 236 genera and about 7173 of zygomorphic flowers are often represented by a formula, species, almost cosmopolitan, but absent from the with the number of lobes in the posterior lip followed by that of the anterior lip, e.g. 3/2. coldest regions of high latitude or altitude. 224 R.M.Harley etal. Labiatae 225
cymes;calyx actinomorphic, campanulate, 32,34, 36,38,40,42,44,46, 50, About 56. 140 spp., 106.Stenogyne Benth., nom. cons., nonCass. mucronateorrounded lobes; inflorescence thyr
5-lobed,lobesequal, subtriangular, notspinescent, various habitats, Macaronesia, Mediterranean soidor racemoid, withup to 5-flowered cymes, StenogyneBenth., Edward's Bot.Reg. ad. 1.1292 (1830); Weller stronglyaccrescent infruit; corolla weakly 2- regiontoRussia, Tibet and W China.Mendoza- bracteolesspinescent; calyxslightly zygomorphic, andSakai inWagner etal. (eds.) Manual ofthe Flowering Heuer(1977) recognises twosubgenera: subg. (4-)5-lobed,lobessubequal or unequal (3/2), lipped,5-lobed (2/3), roseor white, lobes scarcely PlantsofHawaii 1:831-843 (1990), rev. differentiated,rathershort; stamens included in Sideritis andsubg. Marrubiastrumy thelatter posterioroftenlonger, lobes spreading orerect, corolla-tube;filaments short, thecae distinct; endemic toMacaronesia. Vinesorrarely erect, perennial herbs,with simple oblong tobroadly ovate, often spinose-mucronate, stigma-lobesequal;nutlet usually only one per hairs;leaves sometimes palmately lobedin young equalling orlonger than tube; corolla strongly 2-
lipped,5-lobed (2/3), white, yellowish-white, rarely fruit,broadly obovoid, rounded atapex. One plants;inflorescence thyrsoidor racemoid, not 104.Haplostachys (A.Gray.) W.F.Hillebr. pink,posterior liplong, straight, shallowly species,P.parviflora (Benth.) Vved.Rocky moun terminal;calyx5-lobed, becoming wider, often
hooded,denselyvilous outside, 2-lobed,lobes tainslopes, NE Iran, Afghanistan, Uzbekistan, Haplostachys (A. Gray.) W.F.Hillebr., Fl.Hawaii IsL:346 (1888); broadlycampanulate infruit, lobes equal or
sometimes2-toothed, anterior lipwith lateral Pakistan,NW India (W Himalayas). Someauthors Wagner et al., Manual ofthe Flowering Plants ofHawaii 1: unequal(3/2), sometimes irregularly so;corolla 2-
798-801(1990), rev. lobesacute oremarginate, medianlobe obcordate, includePhlomidoschema inStachys. lipped,4-lobed (1/3), redto pink, purple, white or PhyllostegiaBenth.sect. Haplostachys A.Gray tubedensely pilose-annulate inside;stamens not green,posterior lipmoderately long, usually
orshortly exserted from corolla-tube, filaments longerthan lower lip, shallowly hooded oralmost 102.Prasium L. Perennialwoody herbs, with simple hairs; leaves complanate,anthers glabrous orhairy, thecae flat,entire to2-lobed, corolla-tube oftenlong and coriaceous;inflorescence racemoid;bracts mostly distinct;stigma-lobes equal;nutlets trigonous, PrasiumL,Sp. PI. 601 2:(1753). falcate,oftenstrongly dilated distally; stamens distinctlysmaller than the leaves; bracteoles obovoid,apex truncate orrounded, glabrous, equalorposterior pairslightly longer, exserted or presentatbase of pedicels; calyx5-lobed, lobes piloseorglandular. In= 22,34. About 40 species. Smallshrub, usually glabrous orvery sparsely not,thecae weakly distinct; stigma-lobes subequal equal,short, throat ± closed infruit; corolla 2- Dryslopes, valleys, deserts ofIran to Mongolia, pubescentwithsimple hairs; leaves toothed, peti- orslightly unequal; nutlets obovoid, often very lipped,4-lobed (1/3), white tosometimes purple, Russia(S.Siberia), NW China and N. Pakistan. olate;inflorescence lax,racemoid or rarely thyr- large,fleshy, margins slightly winged, obliquely posteriorlipmoderately long,undulate, crisped, Somemedicinal, ornamental. soid,cymes up to 2-flowered; bracteoles atbase attachedatbase, rounded atapex, glabrous. In= entire;stamens equal or posterior pairslightly ofpedicels; calyxbroadly campanulate infruit, 32,(34), 64.About 20 spp., inforests andsubalpine longer,notexserted fromcorolla, thecae weakly slightly2-lipped, 5-lobed (3/2), lobes equal or shrubland,Hawaii. Some species apparently polli distinct;styleincluded in corolla-tube, with 109.Roylea Wall, exBenth. subequal,shortly spinose; corolla strongly 2- natedby honeycreepers. stigma-lobesshortlyclavate; nutlets dry,hairy at lipped,4-lobed (1/3), white topale cream, poste RoyleaWall, exBenth. inEdward's Bot.Reg. 15:1.1289 (1829). apex.Five spp. (four ofthese probably extinct), in riorlip long, hooded; stamens notexserted from dryforests and shrubland atlow elevations, 107.Chamaesphacos Schrenk exFisch. & corolla,thecae weakly distinct, divergent; stigma- Shrubwith simple hairs; leaves petiolate, blade Hawaii. C.A.Mey. lobesequal or subequal;nutlets drupaceous, ovate,deeply crenate; inflorescence lax,thyrsoid, roundedatapex, black, glabrous. In= 34,36. One cymesup to6-flowered; calyxtubular, 5-lobed, ChamaesphacosSchrenk exFisch. & C.A. Mey., Enum. PI. lobesequal orsubequal, oblong-elliptic, narrowed sp.,P.majus L., dry places, mainly coastal, Canary Schrenk.1:27 (1841). 105.Phyllostegia Benth.
Islands,Iberian Peninsula, Morocco and Mediter atbase; corolla strongly 2-lipped, 4-lobed (1/3),
PhyllostegiaBenth.,Edward's Bot. Reg. 15: t. 1292 (1830); raneanregion. Annualherb, subglabrous orvery sparsely pubes pinkto white, posteriorliplong, hooded, densely
Wagneretal., Manual ofthe Flowering Plantsof Hawaii 1: centwith simple hairs; leaves petiolate, spines- pubescentoutside; stamens not exserted from 810-826(1990), reg.rev.; Wagner, Novon 9:265-297 (1999), cent-aristate;inflorescence thyrsoid, withup to corolla,thecae weakly distinct; stigma-lobes nomend. 103.Sideritis L. Fig.21G 3-floweredcymes; calyx weakly 2-lipped, 5-lobed shortly subequal; nutlets rounded atapex, smooth.
In= 34.One sp., R.cinerea (D.Don) BailL, NW L,Sp. PL 2: 574 (1753); Pe*rez dePaz & Negrin, (3/2),lobes spinose; corolla 2-lipped, 5-lobed Sideritis Perennialherbs,lianas orsubshrubs, withsimple
Phanerog.Monogr. 20:1-327, plates 1-22 + 48 figs. (1992), (1-2/3),purple, posterior liplong, flator shallowly India, Nepal. hairs;leaves simple or sometimes irregularly rev.ofsubg.; Obon de Castro & Rivera, Phanerog. Monogr. hooded,anterior lipdeflexed, tubevery long, pinnatisect;inflorescence thyrsoidor racemoid; 21:1-640 (1994), rev.of section. straight,floccose-villous withinatthroat; stamens LeucophaeWebb & Berthel.(1845). calyx5-lobed, becoming broader, often broadly 110.Stachyopsis Popov& Vved. notexserted, filaments pubescent, thecaeweakly campanulateinfruit, lobesequal orsubequal (3/2)
distinct;stigma-lobes subequal; nutlets oblong, Stachyopsis Popov& Vved., Trudy Turkest. Nauch. Obsch. 1: Annualor perennial,sometimes rhizomatous sometimes irregularly so,occasionally dentate- 120(1923). roundedatapex, smooth, narrowly membranous- herbsor shrubs, with simple and sometimes serrate; corolla 2-lipped, 4-lobed (1/3), white to
winged.Onesp., C.ilicifolius Schrenk.Sandy areas, branchedhairs, usually aromatic; leaves often purple, posterior lipmoderately long,shorter than Perennialherbs with short simple hairs; leaves Iranto Kazakhstan andW China(Xinjiang). narrow,usually toothed, sessile to petiolate; lowerlip, shallowly hooded oralmost flat, entire or toothed;inflorescence thyrsoid with many- inflorescencethyrsoidor sometimes racemoid, emarginate, corolla-tube usuallycurved; stamens floweredcymes; calyx actinomorphic, obconical- oftendense; bracteoles present orabsent; calyx equal or posterior pairslightly longer, notor 108.Lagochilus Bungeex Benth. campanulate,5-lobed, lobes equal, triangular, zygomorphicoralmost regular, 5-lobed, lobes only shortly exserted, thecae weakly distinct; spinescent;corollastrongly 2-lipped, 5-lobed LagochilusBungeex Benth., Lab.Gen. et Sp. 640 (1834). subequalorunequal (1/4), mostly triangular, oftenstigma-lobes short,± clavate, subequal oranterior (2/3),purplish tolilac-pink, posterior lipdeeply ChlainanthusBriq.inEngler & Prantl (1896). spinose,tubeoften barbate inthroat; corolla broader; nutlets spreading andexserted from LagochilopsisKnorring(1966). hooded,erect, entire, densely villous, anterior lip 2-lipped,4-5-lobed (1-2/3), usually yellow or calyxatmaturity, obliquely attached atbase, fleshy, withmedian lobe orbicular toobovate, reflexed, white,sometimes purple, posterior lipflat or withslightly winged margins, rounded atapex, Shrubs,subshrubs orperennial herbswith woody emarginate,muchlarger than lateral lobes, tube almostso, deeply emarginate toentire, tube glabrous. In= 64, 66. About 34 spp., inmesic to rootstocks,withor without spines inleaf-axils, ratherbroad, slightly curved, annulate; thecae shorterthan calyx; stamens included incorolla- moistforest, Hawaii, withone species (P.tahiten- glabrousorsparsely hirsute with simple hairs; distinct; stigma-lobes subequal; nutlets oblong, tube,thecae confluent; stigma-lobes unequal; sisNadeaud) inTahiti andone (P. tongaensis H.St. leavesentire ormostly palmatifid withspinose- triquetrous,apexobliquely truncate, glabrous. nutletsrounded atapex. In = 20,22,24,26,28,30,lohn)inTonga Is. Labiatae 237 236 R.M. Harley et al. accrescent; corolla 2-lipped, 5-lobed (4/1), ?red, obovoid-tetrahedral, often with conspicuously 24,26,28,30,32,34,36,38,42,44,46,48,60,66,84, 133. Lepechinia Willd. posterior lip with 2 median lobes, separated by toothed to tuberculate apical crest, glabrous, inner 86, 240. At least 900 species, open to forest and Lepechinia Willd, Hort. Berol 1: 21, t. 21 (1803); Epling, short sinus, lateral lobes rounded,almost obsolete, and upper faces often densely glandular. In = 22. montane habitats, almost cosmopolitan with main Brittonia 6: 352-364 (1948); Hart, Ph.D. Diss., Harvard, anterior lip rounded, tube short, exannulate; About 14 species of wet habitats, mostly in the centres of diversity in SW Asia and Central and Cambridge, Mass., U.S.A. (1983), rev. stamens 4, didynamous, ascending, long-exserted, north-temperate zone (one species in Australia). South America. Especially the Mediterranean S. SphaceleBenth. (1829). filaments curved, thecae 2, strongly divergent, officinalis L. widely used as potherb, also many confluent, elongate; stigma-lobes short, subequal; ornamentals, such as S. microphylla Kunth and Perennial herbs to shrubs, rarely gynodioecious, 136. HyssopusL. fofad\e± disc fleshy, scarcely lobed; nutlets ovoid, smooth, S. splendens Sellow & Nees from the New World. or dioecious (sect. Parviflorae Epling; see Hart black, with minute basal scar. One species: the Hyssopus L., Sp. PL 2: 569 (1753). The infrageneric classification proposed, espe 1985); often aromatic; leaves entire to toothed, rarely collected C. mecistandrum Donn. Sm. is cially by Briquet (1897), largely based on staminal often rugose; inflorescence terminal and often known only from S Mexico (Chiapas) and Perennial subshrub, aromatic; leaves simple, structure, is outdated, although the neotropical axillary, a paniculate thyrse of lax, (l-)3- to many- entire, usually glabrous; inflorescence of 6-many- Guatemala. subgenera Calosphace Benth. and Audibertia flowered cymes, or spiciform of ± congested, flowered verticillasters merging to form a termi usually sessile, 6-to many-flowered verticillasters; (Benth.) Epling appear to be monophyletic. nal, usually ± secund, spiciform thyrse; bracts bracts usually small, bracteoles inconspicuous; Pobedimova (1972) recognised the segregate leaf-like; flowers very shortly pedicellate, bracteo- flowers largeto very small; calyx ± actinomorphic VII.2.B. Subtribe Menthinae (Dumort.) Endl. genus Arischrada Pobed. for species with two late; calyx 2-lipped (3/2), ± tubular, 15-nerved, (1838). fertile anther-thecae, but the separation seems to weakly 2-lipped, 5-lobed (3/2), lobes deltoid to lobes equal, deltoid, acuminate or aristate, ciliate, unjustified. Hedge (1974, 1982) has informally subulate or aristate, tube campanulate, 10-15- Perennial or annual herbs or shrubs. Hairs simple with thickened fold at each sinus; corolla 2-lipped, recognised various smaller groups of closely nerved, often accrescent and inflated, lobes 5-lobed (2/3), violet to whitish, posterior lip deeply spreading to connivent, throat exannulate; corolla or rarely branched. Leaves various, rarely hastate, related species. emarginate, concave, patent, anterior lip with weakly 2-lipped, 5-lobed (2/3), white, yellowish, petiole unwinged, lamina rarely finely rugose. Flowers with calyx-tube 5- to 15-nerved; stamens median lobe ± suborbicular, emarginate, curved purplish to red or blue, lobes often subequal, or downwards, lateral lobes rounded and smaller, 132. RosmarinusL. f hc^\zc, 4, anterior pair usually longer than posterior, or anterior lobe larger, tube cylindrical to campanu tube cylindrical; stamens 4, inserted in middle of late, sometimes shorter than calyx, usually annu stamens divergent, subequal, or stamens 2 with Rosmarinus L, Sp. PL 1: 23 (1753). tube, curved, exserted, thecae divergent, confluent, late within; stamens 4, didynamous, included posterior pair reduced to staminodes or absent; stamens usually 2-thecous, thecae usually ellip connective ± triangular; stigma-lobes equal, ± Shrubs, sometimes gynodioecious, strongly aro to long-exserted, anterior pair usually longer, soid, distinct to confluent, parallel to divaricate, acicular; nutlets ± ovoid, smooth to papillose, matic, with simple and dendroid hairs; leaves ascending, thecae 2, divergent to divaricate, elon connective often conspicuous; nectary-disc entire puberulent, with mucilaginous cells. In = 12. Two linear to lanceolate, revolute, entire, sessile; inflo gate to oblong, sometimes confluent; stigma-lobes species in Europe, NW Africa, Central and SW short, subequal; disc 4-lobed, nutlets ovoid, or often 4-lobed, with lobes usually subequal. rescences as short, 5-15-flowered racemes in axils Asia, including China. Nutlets often areolate. of upper leaves; bracts inconspicuous, bracteoles smooth, brown or black, abscission-scar minute, absent; flowers pedicellate; calyx 2-lipped, 3 or not mucilaginous. In - 32, 66. About 40 species, 137. Mentha L. 5-lobed (1-3/2), campanulate to infundibuliform, often montane, Andes from Argentina and Chile 135. LycopusL. widening above, posterior lip curved upwards, northwards to Mexico and U.S.A. (California), Mentha L., Sp. PL 2: 576 (1753); Harley, Flora Europaea 3: Lycopus L, Sp. PL 1: 21 (1753); L, Gen. PL ed. 5: 12 (1754); triangular, tube 11-12-nerved, accrescent; corolla with outlying species in Brazil and Hawaii, the 183-186 (1972), reg. rev.; Harley, in P.H. Davis (ed.) Flora of Henderson, Am. Midi. Nat. 68: 95-138 (1962), rev. TUrkey 7: 384-394 (1982), reg. rev. strongly 2-lipped, 5-lobed (2/3), violet to white, Chilean L. chamaedryoides (Balbis) Epling natu ralized in Reunion Is. Some species used medici Pulegium Mill. (1753). posterior lip with lobes spreading, anterior lip Perennial herbs, with horizontal rhizomes or Preslia Opiz (1824). nally or as ornamentals. Epling (1948) recognised with lateral lobes, shorter, median lobe rounded, runners, sometimes tuberiferous, weakly or not eight sections, based mainly on inflorescence concave, toothed, tube short, exannulate; stamens aromatic, with simple hairs; leaves opposite, petio- Perennial, rarely annual herbs, subshrubs or structure and floral morphology. Hart (1983) 2, posterior pair absent or rudimentary, anterior late or sessile, ovate to lanceolate or linear, vari shrubs, usually gynodioecious, strongly aromatic; recognised two: sect. Lepechinia and sect. pairlong-exserted, ascending, much shorter in ously toothed or pinnatifid; inflorescence of tight, stems often rhizomatous; inflorescence of few- to Parviflorae, considering the former to be male-sterile flowers, anthers elongate, with single axillary verticillasters; flowers sessile to subsessile; many-flowered axillary cymes, pedunculate or not, polyphyletic. fertile theca separated by an elongate, downward- calyx actinomorphic to weakly zygomorphic, not often forming either dense verticillasters along curving connective from the sterile theca repre accrescent, tubular-campanulate to campanulate, stem, in axils of leaf-like bracts, or congested into sented by small, tooth-like appendage, connective 134. Chaunostoma Donn. Sm. 4-5-nerved, 4-5-lobed, lobes ± equal, ovate to terminal spiciform thyrses in axils of reduced not articulating with filament and appearing lanceolate-subulate or aristate, throatglabrous; bracts, bracteoles inconspicuous, rarely broader or Chaunostoma Donn. Sm., Bot. Gaz. 20: 9, t. 3 (1895). continuous; style curved, exserted, stigma-lobes corolla white, usually weakly zygomorphic, occa digitately lobed; calyx ± actinomorphic, 5-(rarely short, unequal; disc 4-lobed, anterior lobe largest; sionally ± actinomorphic or 2-lipped, lobes usually 4-)lobed, lobes subequal, triangular to subulate or Shrub, cauliflorous, densely hairy, with dendroid nutlets ovoid, smooth, abscission-scar large with 4, posterior and sometimes anterior lobes larger acicular, or slightly 2-lipped, (3/2), anterior lobes and simple hairs; leaves bullate, serrate, petioles central pore, not mucilaginous, pseudostrophio- than lateral ones, all entire or posterior lobe bifid, often longer, tube cylindrical to infundibuliform, unwinged; inflorescences axillary, borne on bare late. In = 24. Three species, with natural hybrids, tube straight; stamens 2 (posterior pair reduced 10-15-nerved, throat hairy or glabrous; corolla stems below leaves, forming short, racemoid in the Mediterranean region and Caucasus. Ros to staminodes or absent), exserted or included, ± actinomorphic, 4-lobed, lilac, pink or white, lobes thyrses, cymes single-flowered; bracts caducous, marinus officinalis L. has been widely cultivated filaments glabrous, thecae parallel to divergent, spreading, subequal, or posterior lobe slightly bracteoles inconspicuous; flowers medium-sized, since antiquity as an ornamental. The genus is separate at dehiscence; style often expanded near broader, often emarginate, corolla-tube ± cylindri pedicellate; calyx actinomorphic to weakly 2- scarcely separable from Salvia. base to form flange covering top of ovary, stigma- cal or rarely gibbous on anterior side, usually lipped, 5-lobed, bright blue, lobes subequal, lobes equalor unequal; disc symmetrical; nutlets included within calyx-tube; stamens 4, subequal, deltoid, tube broadly campanulate, membranous, R.M. Harley et al. 238 Labiatae 239 exserted, divergent (included in male-sterile ± quadrangular, hairy all round, on two opposite cal to campanulate, 10-13-nerved, pilose in throat; axillary cymes, shortly pedunculate or sessile, plants), anthers ellipsoidal, thecae parallel,distinct; sides or only on angles; leavessmall, usually entire, corolla 2-lipped, usually 5-lobed (2/3), white, pink forming dense verticillasters on upper part of stigma-lobes subequal, shortly spreading; disc ± frequently revolute; inflorescence of verticillasters or purple, lobes subequal, tube straight or curved, stem, in axils of leaf-like bracts, often somewhat symmetrical; nutlets ovoid, smooth to rugose,pale forming a terminal, condensed, often spiciform sometimes ventrally saccate, short or to 3 cm in O. congested; bracteoles inconspicuous; flowers to dark brown, not mucilaginous; In = 18,20,24,36, or interrupted thyrse; bracts leaf-like or not, amanum Post; stamens 4, sometimes reduced or subsessile; calyx long-cylindrical, actinomorphic, 40,42,48,54,60,66,72,78,84,90,96,108,120,132. sometimes coloured, bracteoles inconspicuous; absent, usually unequal, ascending to divergent, equally 5-lobed, lobes subulate, tube 10-nerved, 5- About 20 species in damp, open habitats, often by calyx 2-lipped, 5-lobed (3/2), sometimes nearly anterior longer, usually exserted, posterior angled, 5 main nerves along crest of narrow wings, water, especially Mediterranean and Australasian actinomorphic, ± campanulate or cylindrical, 10- exserted to included, thecae divergent to divari alternating with 5 weaker nerves in lower half of (one species in New Zealand), butextending nerved, lobes triangular or setaceous (in T. mas- cate, distinct; stigma-lobes subequal; disc ± membranous intercostal furrows, throat hairy through Europe, Asia and Africa to the Cape, and tichina L.), rarely posterior lip entire, anterior lip symmetrical; nutlets ovoid toapiculate, smooth. within; corolla ± actinomorphic to weakly 2- with one species in North America. Some species upcurved or spreading, throat bearded; corolla 2- In = 30 (32). About 40 species, especially Mediter lipped, 5-lobed (2/3), pink, lobes subequal, corolla- cultivated since antiquity as potherbs and now lipped, or rarely nearly actinomorphic, 4-lobed ranean, extending through temperate Eurasia and tube cylindrical, very narrow, slightly longer than introduced worldwide. Hybrids are of frequent (1/3), white, cream, pink or violet, posterior lip N Africa, from Azores to Taiwan. Cultivated widely calyx, exannulate; stamens 4, subequal, included in occurrence in sect. Mentha (Harley and Brighton ± rounded, emarginate, straight, anterior rectan as culinary and medicinal herbs, especially O. corolla, filaments straight, anterior slightly longer, 1977). These can persist and spread widely by veg gular to suborbicular, rounded, spreading, tube vulgare L, O. onites L. and O. majorana L., some anthers ellipsoidal, thecae ± parallel, distinct; etative means. Mentha spicata L. and its hybrids, ± cylindrical, sometimes very long (to 20 mm); ornamental. Many hybrids recorded. Ietswaart stigma-lobes subequal; nutlets oblong, glabrous. including M. xpiperita L. and, in the subtropics,M. stamens 4, exserted or not, thecae parallel,distinct; (1980) recognises 10 sections. One sp., P. subulifera Hand.-Mazz., dry limestone canadensis L., are of commercial importance. No stigma-lobes ± equal; nutlets ovoid, smooth. 2« = slopes, 400-1000 m, N Iraq and SE Turkey. modern infrageneric classification has yet been 24,26,28,30,32,42,48,50,52,54,56,58,60,84,90. 141. Zataria Boiss. published, while previous systems such as those by About 220 species, open areas, Europe, Mediter Briquet (1897) are unsatisfactory. ranean region and Macaronesia, Asia, N Africa and Zataria Boiss., Diagn. PL Or. Nov. ser. 1,5:18 (1884). 143. Satureja L. % titles mountains of Ethiopia, S Greenland. Aromatic, Satureja L, Sp. PL 2: 567 (1753); Doroszenko, Doct. Thesis, ornamental and medicinal plants used as flavour Subshrub, apparently gynodioecious, aromatic, Edinburgh University: 175-205 (1985), rev. 138. ThymbraL.
Low subshrub, aromatic; leaves elliptic to linear- 146. Micromeria Benth. £ conspicuous, lowest sometimes leaf-like; calyx 149. Minthostachys (Benth.) Spach oblong, flat, entire, gland-dotted especially on cylindrical-campanulate, ± actinomorphic, equally Micromeria Benth., Edwards's Bot. Reg. 15:1.1282 (1829); Perez Minthostachys (Benth.) Spach, Hist. Veg. Phan. 9: 164 (1840); 5-lobed, lobes triangular-subulate, sometimes abaxial surface, subsessile, palisade layer isolat- de Paz, Monograffas Secc. 4, Ctencias Naturales 16: 1-306 Epling, Feddes Rep. Spec. Nov. Beih. 85:162-168 (1936), rev. eral; inflorescence of verticillasters, cymes shortly (1978), reg. rev.; Morales, Bot. Complutensis 18: 157-168 plumose, tube 10-nerved, often long-pilose, throat Bystropogon sect. Minthostachys Benth. (1834). pedunculate, 1-6-flowered, in axils of reduced (1993), reg. rev. hairy; corolla white, pink or lilac, weakly 2-lipped, bracts and forming a lax, terminal, spiciform 5-lobed (2/3), posterior lip emarginate to 2-fid, Weak-stemmed, often semi-scandent shrubs, aro thyrse; bracteoles inconspicuous; calyx ± actin Perennialor annual (M. cymuligera Boiss. & anterior lip with median lobe broader, tube in matic, gynodioecious; leaves shallowly toothed to omorphic, 5-lobed, lobes equal, triangular-lanceo Haussk.) herbs, subshrubs or shrubs, often aro cluded in calyx; stamens 4, shorter than corolla, subentire; inflorescence of compound, congested, late, calyx-tube cylindrical-campanulate, straight, matic; leaves entire or toothed, linear to ovate, flat didynamous, anterior pair longer, spreading, sometimes 2-3, often pedunculate cymes, in 15-nerved, glabrous in throat; corolla pale pink, or revolute; inflorescence in axillary verticillasters, filaments glabrous, thecae parallel or divergent, axils of leaf-like bracts, these becoming reduced 2-lipped, 5-lobed (2/3), posterior lip straight, 2-20-flowered, or sometimes flowers solitary in distinct, connective slightly dilated; stigma-lobes towards stem-apex, upper cymes often forming a flat, anterior lip spreading, 3-lobed, equalling or leaf-axils, bracts ± similar to leaves, bracteoles unequal to subequal, long-subulate, anterior lobe terminal, spiciform thyrse; bracteoles inconspicu slightly shorter than posterior, corolla-tube exan- often present; flowers (possibly resupinate in M. broader; disc symmetrical, reddish-orange; nutlets ous; calyx cylindrical, ± actinomorphic, 5-lobed, nulate; stamens 4, didynamous, filaments straight, cymuligera) pedicellate or not, pedicels sometimes oblong-ovoid, smooth or finely tuberculate. In = lobes subulate to narrowly triangular, calyx-tube rather short, anterior pair longer and slightly very long; calyx actinomorphic to weakly 2-lipped, 42,48. Seven species, endemic to Canary Is. and often externally long-pilose, 13(-12)-nerved, exserted from tube, ascending under posterior campanulate or cylindrical, scarcely accrescent, Madeira. Mountain slopes and clearings. Hybrids throat hairy; corolla white, or pale lilac, 2-lipped, lip of corolla, thecae slightly divergent, separate; 13-15-nerved, 5-lobed, lobes ± equal or (3/2), ante have been reported. La Serna (1984) recognised 5-lobed (2/3), anterior lip with median lobe stigma-lobes equal, subulate, nutlets oblong, apic- rior often longer, sometimes curved, spreading, two sections. Sect. Bystropogon^ flowers in dichasial larger, corolla-tube hairy within, longer to shorter ulate, glabrous. One species, G. popovii (Fedsch. 8c throat bearded or not; corolla usually small, white cymes, calyces with lobes narrow, subulate, than calyx; stamens 4, didynamous, shorter than Gontsch.) Boriss. Pamir-Alai: Uzbekistan, Tajik to yellowish, mauve, pink or purple, 2-lipped, 4- plumose, nutlets smooth, 4 spp. Canary Is. Sect. corolla, slightly exserted from tube, thecae slightly istan, Kazakhstan, Afghanistan, Pakistan (Chitral), lobed (1/3), posterior lip emarginate, ± straight, Canariense La-Serna, flowers in congested divergent, distinct, connective slightly dilated; India: Kashmir. Dry limestone slopes. anterior lip spreading, tube cylindrical, sometimes glomerules, calyces with lobes short, triangular- stigma-lobes long, unequal, anterior longer; hairy in throat; stamens 4, didynamous, included acuminate, not plumose, nutlets somewhat tuber disc symmetrical reddish orange; nutlets ovoid, or exserted from tube, filaments straight or con- culate, 3 spp. Madeira and Canary Is. finely reticulate. In = 46. Probably less than 145. Saccocalyx Coss. & Durieu nivent, thecae divaricate, ellipsoidal, confluent; twelve spp., montane, Andes from Venezuela Saccocalyx Coss. & Durieu, Ann. Sci. Nat. Bot. Ser. 3, 20: 80 stigma-lobes subequal, subulate; nutlets ovoid, to Argentina. Species limits are very poorly 148. Cuminia Colla (1853), nom. cons. sometimes apiculate, ± smooth, sometimes hairy, understood.
mucilage cells present. In = 20, 22, 30, 50,60. Cuminia Colla, Mem. Acad. Sci. Torino 38,6:139 (1835); Harley, Small shrub, strongly aromatic; older stems often About 70 species, Europe, Mediterranean region, Kew Mag. 3,4: 151-156 (1986), rev. 150. Clinopodium L. Fig. 28 decumbent and rooting at base, with longitudi Macaronesia, N Africa, Asia, mountains of E and Skottsbergiella Epling (1937), non Petrak (1927). Johowia Epling & Looser (1937). nally fissured bark; leaves small, remote, ± spathu- S Africa, and W tropical Africa, Madagascar. Clinopodium L, Sp. PL 2: 587 (1753); L, Gen. PL ed. 5: 256 late, sessile, entire, canaliculate, conspicuously Aromatic and medicinal plants used as flavouring (1754); Cantino & Wagstaff, Brittonia 50: 63-70 (1998), reg. gland-dotted, margin ciliate with long, white hairs and source of essential oils. Of the six sections Small tree or shrub, aromatic, gynodioecious consp.; Harley & Granda Paucar, Kew Bull. 55: 917-927 in lower half, cauUne leaves with axillary fascicles recognised by Morales (1993), only four are or gynomonoecious; leaves petiolate, sharply and (2000), reg. consp. irregularly serrate; inflorescence of compound, Calamintha Mill. (1754). of smaller leaves; inflorescence of verticillasters, included in Micromeria as delimited here: Acinos Mill. (1754). dichasial, pedunculate cymes in axils of, and cymes sessile, 1-3-flowered in axils of leaf-like Pseudomelissa Benth., Micromeria^ Pineolentia P. Gardoquia Ruiz et Pav. (1794). ■ - 1 < i s - 'V < bracts; bracteoles inconspicuous; flowers minute; P^rez, Cymularia Boiss. The supposedly resupinate shorter than, large, leaf-like bracts; bracteoles Xenopoma wmd. (1811).j ^Pl fe"i'^ "'^'7 " calyx urceolate, actinomorphic, 5-lobed, lobes natureof the flower of M. cymuligera has been inconspicuous; calyx tubular-campanulate, rather Antonina Vved. (1961). "~ broadly deltate, slightly spreading, calyx-tube ca. questioned. J Itk fleshy, actinomorphic, 5-lobed, lobes triangular- 13-nerved with inconspicuous nerves, densely lanceolate, ciliate, often slightly spreading, Perennial, rarely annual herbs and shrubs, aro spreading-hairy outside, annulate at throat within, calyx-tube obscurely 12-14-nerved, exannulate, matic, with simple or branched hairs; leaves peti 147. Bystropogon L'He*r. tube strongly accrescent in fruit, becoming becoming distended at base to accommodate olate to subsessile, simple, variously toothed or globose-urceolate, inflated; corolla purplish-pink, Bystropogon L'He*r., Sert. Angl. 19, t. 22,23 (1788), nom. cons.; nutlets, throat glabrous; corolla white to lavender entire, margin sometimes revolute; inflorescence ± actinomorphic, very small, 5-lobed, lobes sube- La-Serna, Revisidn del ge"nero Bystropogon, J. Cramer, Vaduz or dark purple, ± actinomorphic, 5-lobed, lobes of axillary cymes, these sometimes reduced to qual, erect, corolla-tube shortly campanulate- (1984), rev. rounded, posterior pair erect, tube hairy outside, solitary flowers or merging to form a terminal, cylindrical, glabrous within; stamens 4, subequal, longer than calyx, annulate within; stamens 4, paniculiform to spiciform, verticillastrate, or sub- ± included within corolla, filaments glabrous, very Shrubs, strongly aromatic, frequently gynodioe- subequal, included in corolla-tube, filaments capitate thyrse; calyx actinomorphic to 2-lipped, cious; leaves petiolate, crenate, dentate to sub- short, thecae weakly divergent, separate, connec short, thecae divergent, distinct; stigma-lobes not accrescent, usually cylindrical to tubular-cam tive small; stigma-lobes slightly unequal, posterior entire; inflorescence in compound, terminal or slightly unequal, posterior shorter; disc sym panulate, straightto sigmoid, sometimes gibbous, lobe shorter; disc with elongate anterior lobe; axillary, dichasial, pedunculate, few to many- metrical, fleshy; nutlets ovoid-trigonous, fleshy usually 13- (11-, 15-)nerved, 5-lobed, lobes often nutlets oblong, apiculate, smooth, mucilaginous flowered, often corymbiform cymes, lax or con with thickened angles, rather large. In - 46, 44. upcurved, posterior 3 often partially fused to form when wetted. One species, S. satureioides Coss. & gested, in axils of leaf-like bracts, often forming One species, C. eriantha (Benth.) Benth., in a lip, anterior 2 usually free above tube, throat Durieu, Algeria. Sandy margins of salt lakes, in spherical glomerules, often with solitary, terminal lower montane forest, 200-450 m, Chile: hairy or not; corolla white to blue, lavender, red, or flower at lowest cyme-branch; bracteoles in- desert. Masatierra Is., Robinson Crusoe (Juan Fernandez) orange (rarely yellow), 2-lipped (2/3), posterior lip Archipelago. usually notched, tube straight or curved; stamens Labiatae 243 242 R.M. Harley et al.
sessile or pedunculate cymes in axils of leaf-like usually 4 (rarely the posterior pair reduced to campanulate, downcurved, 11-13-nerved, 5-lobed 156. Hedeoma Pers. bracts, forming a terminal, spiciform or capituli- staminodes), didynamous, included or exserted, (3/2), lobes usually slightly upcurved, posterior Hedeoma Pers., Syn. PI. 131 (1807); Irving, Sida 8(3): 218-295 form thyrse; bracts patent or imbricate, ± conspic filaments usually glabrous, thecae parallel to lobes deltoid-cuspidate, fused to form a lip, ante (1980), rev. uous, bracteoles inconspicuous; flowers shortly divaricate, separate at dehiscence; stigma-lobes rior lobes lanceolate-subulate, free above tube, Pseudocunila Brade (1944). pedicellate, pedicels sometimes flattened; calyx ± equal or unequal; disc usually symmetrical; throat densely villous; corolla usually blue to pink, actinomorphic,5-lobed (±3/2), lobes triangular, nutlets ellipsoid to ovoid, obloid, obovoid, or 2-lipped, posterior lip deeply bifid, anterior lip Annual or perennial herbs or subshrubs, strongly posterior lobes shorter than straight or curved subglobose, often somewhat trigonal, smooth or 0.8x to as long as tube, 3-lobed, median lobe aromatic; leaves entire or toothed; inflorescence of anterior lobes, calyx-tube cylindrical, sometimes minutely sculptured, glabrous or puberulent. In = largest, notched, tube ± straight; stamens 4, didy few- to many-flowered, pedunculate to subsessile curved, gibbous below, 13-nerved, accrescent, 10-72 (most frequently 18 or 20). About 100 spp., namous, included, filaments glabrous, thecae cymes, or flowers rarely solitary, in axils of often throat bearded, closed; corolla 2-lipped, ±4-lobed mostly in the New World (both temperate and divaricate, separate at dehiscence; stigma-lobes leaf-like bracts, upper cymes often congested in a (1/3), white, rose-red or purple, posterior lip tropical) and temperate Eurasia, but a few in unequal; disc symmetrical; nutlets ellipsoid, terminal spiciform thyrse; bracteoles inconspicu rounded, sometimes emarginate, straight, anterior Africa, tropical Asia and Indomalaysia. As delim smooth, glabrous. One species, 0. caerulescens ous; flowers sometimes deistogamous; calyx lobes ± rounded, corolla-tube cylindrical, short, ited here, Clinopodium includes most of the native (Bentham) Doroszenko & P.D. Cantino, N ± actinomorphic, subequally 5-lobed, or 2-lipped sometimes scarcely exserted from calyx, exannu- New World species of Satureja sensu Epling and Colombia. (3/2) with lobes unequal, posterior somewhat late; stamens 2 (posterior pair staminodal or Jativa (1964,1966), see Cantino and Wagstaff connate, deltoid to subulate, anterior lobes usually absent), anterior pair ascending under posterior (1998). longer, tube 13-nerved, cylindrical to infundibular, 153. Kurzamra Kuntze corolla-lip, exserted from tube, anthers often often gibbous at base, throat densely hairy within; cohering at margins, with fertile theca ellipsoidal, corolla white, lilac, pink or orange-red, 2-lipped Kurzamra Kuntze, Rev. Gen.: 520 (1891). 151. Cyclotrichium (Boiss.) Manden. & Scheng. divaricate, becoming confluent, or often dimidiate, Soliera Clos (1849) non J.G. Agardh (1842). (2/3), posterior lip sometimes subentire, concave lower theca sterile and forming a small Cyclotrichium (Boiss.) Manden. & Scheng., Not. Syst. to flat, erect or spreading, median anterior lobe appendage; stigma-lobes unequal; disc symmetri Leningrad 15: 332-337 (1953). Dwarf, mat-forming perennial herb, aromatic; usually larger, often emarginate, tube short to Calamintha sect. Cyclotrichium Boiss. (1879). stems procumbent, often rhizomatous, subter cal; nutlets ovoid to trigonous, smooth or granu elongate, widening upwards, annulate or not ranean at base, with reduced scale-leaves; leaves, late. In = 12,16,18,24,32,36. About 20 species in within; stamens 2, posterior pair staminodal, ante Perennial subshrubs, hairs simple or dendroid; small, entire, densely hairy; inflorescence of l(-2)- S Europe, NW Africa and Asia to Himalayas and rior pair ascending under posterior corolla-lip, leaves entire or toothed; inflorescence of axillary flowered cymes, sessile, subtended by leaf-like Altai mountains. Open, usually xeric habitats. included to exserted, thecae ± parallel to widely cymes, sometimes very shortly pedunculate, bracts and crowded towards shoot-apex; bracte Aromatic and medicinal plants used as flavouring divaricate, distinct, connective well developed; forming terminal thyrse of 6-many-flowered, oles paired, inconspicuous; calyx ± actinomor- and as source of essential oils. stigma-lobes unequal, ± subulate, posterior very usually remote verticillasters; bracts, at least phic, 5-lobed, lobes equal, triangular, with apex small; disc symmetrical; nutlets ovoid to oblong, the lowest, leaf-like, upper small, inconspicuous, long-subulate, plumose, curved, more than 2x as 155. Stachydeoma Small usually 1.5 mm long or less, areolate to foveate, bracteoles ± inconspicuous, smaller than to long as tube, spreading in fruit, calyx-tube shortly sometimes glaucous, often mucilaginous. In = 34, equalling calyx; calyx weakly 2-lipped, 5-lobed cylindrical- infundibuliform, 15-nerved, externally Stachydeoma Small, Fl. S. E. United States 1040,1337 (1903). 36,44, 72, 76,144. About 42 species, from Canada (3/2), lobes triangular-subulate, posterior lobes patent-hairy, throat annulate within; corolla lilac, southwards to Mexico, Central and Southern usually shorter than anterior, calyx-tube cylindri 2-lipped, 5-lobed (2/3), posterior lip shallowly Low shrubs or subshrubs, aromatic, conspicuously S America, from Peru and Bolivia to S Brazil, cal, straight or shortly curved, 13-nerved, throat lobed, shorter than anterior lip, corolla-tube long- glandular and setose; leaves opposite, subsessile, Uruguay and Argentina. Open montane or sub hairy; corolla resupinate, 2-lipped, 4-lobed (3/1), exserted from calyx, narrow at base, gradually ovate to elliptical, entire or few-toothed, usually montane, sometimes xeric habitats. Irving (1980) white or lilac, posterior (lower) lip entire or emar- widening towards throat, straight, glabrous within, revolute; inflorescence a terminal thyrse or recognises four subgenera, based mainly on habit ginate, anterior (upper) lip 3-lobed, slightly stamens 4, didynamous, exserted from corolla- raceme; calyx zygomorphic, not accrescent, cam- and reproductive characters. deflexed, median lobe larger, corolla-tube twisted, tube, divergent, anterior pair longer, thecae weakly panulate, 13-nerved, glandular-punctate, 5-lobed annulate near middle; stamens 4, ± equal, long- (3/2), setose, posterior lobes triangular, largely divergent, separate, connective small; stigma- 157. Rhododon Epling exserted, anthers ellipsoid, thecae 2, parallel, dis lobes flattened, slightly unequal, posterior shorter; fused to form a shallowly lobed lip, anterior lobes tinct with short connective; stigma-lobes unequal, disc ± symmetrical; nutlets oblong, mid-brown, subulate, slightly upcurved, free above tube, throat Rhododon Epling, Rep. Spec. Nov. Regn. Veg. Beih. 115: 14 ± subulate, posterior lobe longer; disc weakly 4- smooth, not mucilaginous. One species, K. hairy; corolla purplish, 2-lipped (1/3), posterior lip (1939). lobed; nutlets ovoid, smooth, glabrous. About eight pulchella (Clos) Kuntze, Chile: Atacama and entire, anterior lip with median lobe notched, tube species in SW Asia. Coquimbo and adjacent Argentina. straight; fertile stamens 2 (posterior pair reduced Annual herbs; leaves subsessile, entire with to staminodes), included to slightly exserted, margins long-ciliate; inflorescence of verticil filaments glabrous, thecae nearly parallel, asym lasters, cymes 1-4-flowered, in axils of leaf-like 152. Obtegomeria P.D. Cantino & Doroszenko 154. ZiziphoraL. £ her&les metrical (one theca attached higher than other), bracts, forming terminal spiciform, congested or Obtegomeria P.D. Cantino & Doroszenko, Novon 8:1-3 (1998). separate at dehiscence; stigma-lobes unequal; interrupted thyrse; bracteoles long-ciliate, curved, Ziziphora L, Sp. PL 1: 21 (1753); Hedge, Notes Roy. Bot. Gard. Edinburgh 23: 209-221 (1961); Rechinger f., Fl. Iranica 150: disc symmetrical; nutlets subglobose, smooth, purple-tinged; calyx often purple-tinged, 2-lipped, Low, ericoid shrub with simple hairs; stems trans 480 (1982), reg. rev. glabrous. In = 18. One species, Stachydeoma grave- 5-lobed (3/2), lobes elongate, long-ciliate, anterior versely ridged at nodes; leaves subsessile, stiffly Faldermannia Trautv.(1839). olens (A. Gray) Small, Florida. Differs from narrower with deeper sinus, tube 15-nerved, con ascending, linear-oblong, entire, replicate (abaxial Hedeoma in chromosome number and asymmet spicuously ribbed, cylindrical, weakly gibbous, surface hidden); flowers axillary, solitary, with 2 Annual or perennial herbs or subshrubs, aromatic; rical anthers. slightly curved, annulate at throat; corolla lilac to lanceolate bracteoles near base of pedicel; calyx leaves ± entire, subsessile to shortly petiolate; purple, 2-lipped, 5-lobed (2/3), posterior lip flat, strongly zygomorphic, not accrescent, tubular- inflorescence of 2-12-flowered verticillasters, of anterior longer and broader, tube narrowly cylin- Labiatae 249 248 R.M.Harley etal.
posteriorliplonger than anterior withshallower stronglyarcuate; stamens usually included; tap- subulateappendage, thecae 2,divaricate, distinct, montaneforests. Manyspecies haverestricted
sinusbetween teeth, tube13-nerved, cylindrical- rootedannuals, biennials, short-lived perennials, ± parallel tofilament; stigma-lobes shortlysubu rangeswith specialised soilconditions. Epling
campanulate,exannulate within;corolla whiteor orlow shrubs: subgenus Cheilyctis (Raf.)E.M. late; disc unlobed; nutlets ovoid, apiculate or (1925)divided thegenus into two subgenera: subg.
pink,2-lipped, 4-5-lobed (1-2/3), posterior lip McClint.andEpling. rounded,± angulate, mucilaginous. In= 28,30,32. Monardella(assubg. Pycnanthae) andsubg.
concave,erect,anterior lipusually longer, broad, Aboutseven species inEurope, NW Africaand MacranthaeBriq.,based on relative length of
spreading,medianlobe longer thanlaterals, often Asia.Prunella vulgaris L.widely naturalized in corollalobesand tube and on calyx length. 174.Blephilia Raf. emarginate,tubenarrowly cylindrical, equallingto temperateand montane regions ofN andS
long-exsertedfromcalyx; stamens 4,didynamous America,Africa and Australasia. BlephiliaRaf.,Am. Monthly Mag. & Crit.Rev. 4: 190 (1819); 171.Pycnanthemum Michx. withposterior pairshorter, or2 and then poste Simmers& Krai, Rhodora 94:1-14 (1992), newsp.
riorpair staminodal, anterior pairascending, PycnanthemumMichx., Fl.Bor.-Am. 2:7 (1803), nom.cons; 176.CleoniaL. g_ t Grant& Epling, Univ.Calif. Publ.Bot. 20: 195-240 (1943), included under posterior corolla-lip, filaments Rhizomatousorstoloniferous perennial herbs,
rev. glabrous,thecae divergent, distinct, sometimes weaklytostrongly aromatic; leavespetiolate, ovateCleonia L.,Sp. PL 2 ed. 2: 837 (1763).
KoelliaMoench (1794). unequal, glabrousorhairy, connective apparent; tolanceolate or elliptic, serrate serrulate; to
discsymmetrical; stylehairy toglabrous, stigma- inflorescencea verticillastrate thyrseofterminal Annual herb, weakly aromatic; leaves simple, Rhizomatousperennial herbs,strongly aromatic, lobesunequal, subulate, posterior shorter; nutlets andsubterminal glomerules, eachsubtended by crenate todentate orpinnately lobed; inflores withsimple hairs; leaves sessile orshort-petiolate, obovoid-oblong,smooth,mucilaginous. In- 19. aninvolucre ofovate tolanceolate bracts;flowers cence of 6-flowered verticillasters, mergingto ovatetolanceolate, ellipticalor linear, variously Fourspecies, U.S.A.(California), oneextending subsessile,ebracteolate; calyxzygomorphic, not form a terminal,subcapitate thyrse;bracts con toothedorentire; inflorescence ofverticillasters, intoMexico (Baja California). Openareas in rocky accrescent,cylindrical, 13-nerved, 5-lobed (3/2), spicuous, ciliate, pectinate-pinnatifid, segments or capitate,or corymbiform, composed of orclay soils. lobeslanceolate-attenuate tosubulate, setose, pos aristate;calyx2-lipped (3/2), campanulate, 10- compactorloose, terminal andupper-axillary teriorlobes partially fusedto form long upper lip, nerved; posterior lobes deltoid, anterior lobes glomerules(paired cymes); calyx actinomorphic anteriorlobes shorter, freeabove tube, throat longer, aUstraight; corolla 2-lipped (1/3), violet- tozygomorphic, notaccrescent, cylindrical to 173.Monarda L. sparselyhairy; corolla purple towhite, 2-lipped purple to whitish, posterior lobeemarginate, tubular-campanulate,10-13-nerved, 5-lobed, (1/3),posterior lipentire, anterior lip3-lobed, vertical, anterior lipwith median lobe whitish, MonardaL,Sp. Pi. 1:22 (1753); Gen.PL ed. 5:14 (1754); Scora, lobestriangular toovate, lanceolate oraristate, all ± suborbicular,emarginate, horizontal, lateral Univ.Calif. Publ.Bot. 41:1-59 (1967), rev. medianlobe narrower than other 2,tube curved, similaroranterior 2more deeply cleft than poste corollainbud falcate; fertilestamens 2 (posterior lobes suborbicular, truncate,tube ± infundibular; rior3,throat glabrous; corollausually white to Rhizomatousperennial herbs or taprooted pairreduced tostaminodes), exserted, filaments stamens 4,didynamous, posterior shorter, anterior lavenderwithpurple spots, 2-lipped, posterior lip annualsorbiennials (rarelylow shrubs), aromatic, glabrous,thecaedivaricate, separateat dehiscence; pairexserted fromtube, filaments glabrous, thecae entireoremarginate, anteriorlip 3-lobed, tube withsimple hairs; leaves petiolate tosubsessile, stigma-lobesunequal; discsymmetrical; nutlets ellipsoidal, confluent, connective appendaged; ± straight;stamens 4,didynamous orsubequal, ovatetolanceolate, ellipticorlinear, serrateto den ellipsoidtoovoid, obscurely trigonal, smooth, stigma-lobes 4,short, subequal; nutlets ovoid, exsertedorincluded, filaments glabrous, thecae ticulate(rarelyentire); inflorescence terminalor glabrous.Threespecies, eastern North America. smooth. In= 20.One species: C.lusitanica L.,fre parallel,separate atdehiscence; stigma-lobes inverticillasters, ofinvolucrate glomerules; bracts quentlyongypsiferous soils,Iberian peninsula, unequaltosubequal (rarely equal); discsymmet sometimesbrightly coloured; flowers subsessile, NW Africa. rical;nutlets ovoidto obloid, ±trigonal, smoothor 175.Prunella L. ebracteolate;calyxactinomorphic, notaccrescent, rugose,glabrous orpubescent. In= 36,38,40,72, cylindrical,13-15-nerved, 5-lobed,lobes lanceo PrunellaL,Sp. PL 2: 600 (1753). 177.HorminumL. orcurved, corolla inbud falcate; fertilestamens 2 ofusually 6-flowered verticillasters, ina terminal, inflorescence of2-6-flowered verticillasters form condensed,spiciform thyrse, pedunculate ornot; ing a lax,secund, spiciform thyrse, scapose, or 172.Acanthomintha (A.Gray) Benth. & Hook. f. (posteriorpairreduced tostaminodes orabsent), includedorslightly exserted, filaments usually bractsimbricate, broadlyovate-acuminate, bractesometimes afew pairs ofleaves present onflower Acanthomintha(A.Gray) Benth. & Hook, f.,Gen. PL 2,2:1192 pubescent,anthersusually lightly fused(coherent olespresent or not;calyx 2-lipped, tubular- ingstem; bracts not conspicuous, ovate,acute, (1876);Gray,Syn. Fl.N. Am. 2, 1: 365 (1878), rev.;Jokerst, buteasily separated), thecaedivaricate, separateat campanulate,accrescent, 10-nerved, sometimes as long as or shorter than calyx; ebracteolate; Madrono38: 278-286 (1991), rev. dehiscence;stigma-lobes usuallyunequal; disc with2 lateralkeels, 5-lobed (3/2), posterior lip± calyx2-lipped (3/2), campanulate, 13-nerved, symmetrical;nutletsovoid toobloid, obscurely truncate,lobesshort, mucronate, anterior lipwith posterior lobesdeltoid, upcurved, anterior lobes Annualherbs, aromatic; leavespetiolate, coarsely trigonal,smooth,glabrous. In= 18,22,36 (rarely lobestriangular, upcurved, throat notbarbate, ± long-deltoid, straight,throat glabrous; corolla 2- toothedtosubentire; inflorescence ofverticil 24,32,34).About20species, SCanada toS Mexico; closedinfruit; corolla 2-lipped, 4-lobed (1/3), lipped, 4-lobed (1/3), violet, posterior lipsemi- lasterswithcymes 3-5-flowered, inthe axils of somewidely grown asornamentals. Twosubgen purple,blue,cream orwhite, posterior liphooded, orbicular, emarginate, anterior lipwith lobes ± leaf-likebracts,toothed andoften apically spinose; concave,± rounded, anterior lipwith median lobe rounded, truncate, reflexed, tube± cylindrical, ina terminal, congested orinterrupted spiciform era recognised are (McClintock andEpling 1942; Scora1967). Glomerules usually solitary (rarely concave,usually dentate, lateral lobes obliquely hairy inthroat; stamens 4,didynamous, posterior thyrse;bracteoles conspicuous, 2 per cyme, two),terminal; upperlip of corolla straight toonly truncate,tube± infundibular,annulatewithin, pair shorter, inserted inmiddle oftube, exserted, broadlyovateto rotund, spinose-dentate, subses- slightlyarcuate; stamens exserted; rhizomatous contractedatthroat; stamens 4,didynamous, anthers cohering in pairs, thecae confluent; sile,green or straw-coloured, equallingor over perennialherbs:subgenus Monarda, Glomerules exsertedfromtube, ascending underposterior lip,stigma-lobes shortlysubulate; nutlets ovoid, ± toppingcalyx; calyx 2-lipped, 5-lobed(3/2), lobes filamentscurved, bifurcating atapex to form a papillate.In= 12. One species: H.pyrenaicum L., subequal,triangular-lanceolate, spinose-tipped,several, in verticillasters; upperlip of corolla Labiatae 251 250 R.M. Harley et al. thyrsoid, spike-like or interrupted, cymes long- Perennial herbs, aromatic; leaves simple, dentate; calcicole in mountains of Europe, Alps, Pyrenees, 179. Drepanocaryum Pojark. to short-pedunculate or sessile; bracts leaf-like, inflorescence thyrsoid, terminal, spike-like or Cantabrian mountains. Drepanocaryum Pojark., Fl. U.R.S.S. 20:516 (1954); Budantsev, oblong-elliptic to ± ovate, bracteoles inconspicu panicle-like, cymes pedunculate or sessile; bracts Bot. Zhurn. 77,12:118-128 (1992), rev. ous; flowers sometimes resupinate; calyx tubular, reduced, lower often leaf-like, bracteoles subulate; campanulate, straight or curved, annulate within, calyx tubular or campanulate, straight, indistinctly VIL2.C. Subtribe Nepetinae Coss. & Germ. Annual herbs, not aromatic; leaves simple, petio- actinomorphic or indistinctly 2-lipped, 5-lobed 2-lipped, 5-lobed (3/2), lobes equal, throat open; (1845). late; inflorescence thyrsoid, cymes axillary, long- (3/2), lobes subequal, broadly ovate, becoming corolla strongly 2-lipped, 5-lobed (2/3), pink, blue pedunculate, distant; bracts similar to the leaves, patent, accrescent in fruit, membranous-scarious, to violet or white, posterior lip straight, anterior Calyx-tube 15-nerved. Corolla strongly 2-lipped bracteoles subulate to acerose, inconspicuous; often pale green, rose or purplish, tube funnel- lip with median lobe widest, spreading, entire or (2/3), often bluish. Stamens 4 (rarely anteriorpair calyx markedly oblique-gibbous at base, weakly 2- shaped, distinctly ribbed; corolla 2-lipped (2/3), undulate, lateral lobes straight, tube gradually absent in some species of Nepeta), posterior pair lipped, 5-lobed (3/2), lobes subequal or anterior blue-violet, posterior lip shortly 2-lobed, anterior dilating to throat; stamens much exserted or longer than anterior, both usually ascending under lobes slightly longer, triangular, throat ± straight; lip with median lobe concave or flat, margins included, filaments parallel or posterior pair posterior corolla-lip, sometimes exserted. Stigma- corolla slightly exceeding calyx, 2-lipped, 5-lobed undulate or entire, tube elongate, straight, narrow, antrorsely bent, thecae parallel; stigma-lobes lobes subulate. Nectary disc 4-lobed or 4-horned. (2/3), blue-lilac, anterior lip with median lobe gradually widening above; stamens exserted or equal, spreading; disc 4-lobed; nutlets ellipsoid to Nutlets elliptic to oblong, rarely curved and concave, margin undulate, tube gradually widen included, filaments parallel, thecae divergent at obovoid, smooth with apex hairy, areole oblong or globose, abscission-scar usually with well-defined ing above; stamen-filaments parallel, thecae 180°; stigma-lobes slightly unequal; disc 4-lobed; orbicular. In = 18. About 22 species. Mountains or areole. glabrous, divergent at 180°; stigma-lobes sube nutlets ovoid or elliptic, smooth or tuberculate. deserts of U.S.A, southern Canada and Mexico, qual; disc 4-horned; nutlets globose, tuberculate, In = 18. About ten species, Iran and Afghanistan. one species in W Asia. Sectional treatment after mucilaginous, areole large, curved. One species, 178. Nepeta L. Some medicinal, ornamental. Sanders (1987). Sect. Agastache: stamens long- D. sewerzowii (Regel)Pojark. Mountains, W and exserted, filaments of posterior stamens antrorsely Nepeta L, Sp. Pi.2: 570 (1753); Budantsev, Bot. Zhurn. 78,1: CAsia. bent, and crossing anterior pair. Seven species in 91-105 (1993), rev. 182. Marmoritis Benth. Kudrjaschevia Pojark (1953). U.S.A. and Canada, one species in Asia. Sect. Brit- Pitardia Batt. ex Pit. (1918). 180. Lophanthus Adans. Marmoritis Benth., Hooker's Bot. Misc. 3: 377 (1833); Budant tonastrum (Briq.) Lint & Epling: stamens included sev, Bot. Zhurn. 77,12:118-128 (1992), rev. or shortly exserted under posterior lip of corolla, Lophanthus Adans., Fam. PI. 2: 194 (1763); Budantsev, Bot. Subshrubs or annual or perennial herbs, aromatic, Phyllophyton Kudo (1929). filaments of stamens all parallel. 14 species. SW Zhurn. 77,9: 69-77 (1992), rev. Pseudolophanthus Levin (1941). sometimes gynodioecious; leaves simple; inflores United States and Mexico. cences thyrsoid, cymes lax to congested, pedun Perennial herbs or subshrubs, aromatic; leaves Perennial herbs, aromatic; flowering stems erect or culate or sessile, axillary, distant, or crowded simple; inflorescence thyrsoid, terminal, spike-like ascending, densely leafy above and bare below 184. Meehania Britton into spike-like or ovoid heads; bracts leaf-like or or in ovoid heads or paniculate, cymes often long- with small scale-leaves and long rhizomes; leaves reduced, sometimes longer than calyx, bracteoles Meehania Britton, Bull. Torrey Bot. Club. 21:32 (1894); Budant pedunculate or shortly pedunculate in axils of simple, sessile or subsessile, densely imbricate, inconspicuous; calyx tubular to campanulate, sev, Bot. Zhurn. 77,12:118-128 (1992), rev. leaf-like bracts, bracteoles subulate, inconspicu ± orbicular to reniform; inflorescence thyrsoid, actinomorphic to strongly 2-lipped, 5-lobed (3/2), ous; flowers resupinate; calyx tubular or tubular- terminal, cymes few-flowered, sessile, partially or rarely with lips entire, lobes subequal to Annual or perennial, stoloniferous herbs, not campanulate, indistinctly 2-lipped, 5-lobed, (3/2), obscured in axils of upper leaf-like bracts; bracte unequal, throat straight to strongly oblique; aromatic; leaves simple, dentate; inflorescence lobes unequal, triangular, throat open, calyx- oles entire, linear, shorter than calyces; flower corolla 2-lipped, 5-lobed (2/3), blue, violet, pink, thyrsoid, terminal, cymes pedunculate or sessile, tube annulate within; corolla strongly 2-lipped, resupinate or not; calyx tubular or campanulate, yellow or white, posterior lip straight or curved, few-flowered or sometimes solitary, lax, in axils of 5-lobed, (2/3), blue-violet or pink, anterior lip curved or ± straight, indistinctly 2-lipped, 5-lobed anterior lip with median lobe concave or ± flat, leaf-like bracts, bracteoles subulate, inconspicu 3-lobed, median lobe concave or flat, margin (3/2), lobes ovate to linear, posterior lobes longer, undulate or entire, tube included in or exserted ous; flowers usually large; calyx tubular to cam undulate or entire,lateral lobes deltoid to ovate, calyx-tube annulate or glabrous within; corolla from calyx, straight or curved, gradually or ± panulate, accrescent in fruit, 2-lipped, 5-lobed posterior lip 2-lobed, tube gradually widening strongly 2-lipped, 5-lobed (2/3), white or purplish, abruptly dilating upwards; stamens of hermaph (3/2), lobes triangular to lanceolate, acute, those of above, exserted orincluded in calyx; stamens posterior lip short, straight, bifid, anterior lip rodite flowers 4, rarely 2 with anterior pair absent, anterior lip shorter; corolla strongly 2-lipped, 5- exserted or included, filaments parallel, thecae with median lobe oblong or orbicular, lateral lobes usually shorter than corolla, filaments parallel, lobed (2/3), blue-violet to purplish, posterior lip parallel ordivergent at acute angle; stigma-lobes oblong-ovate, tube gradually dilated towards apex, thecae divergent at 18Q° (stamens of male-sterile straight, apex emarginate or 2-lobed, anterior lip subequal; disc 4-lobed; nutlets trigonous to ellip exserted; stamens included or exserted, filaments flowers rudimentary); stigma-lobes subequal; disc with median lobe widely obovate-ovoid, concave, soid, smooth or tuberculate. In = 16. Circa 20 parallel, thecae parallel or divergent at 90°; 4-lobed; nutlets ellipsoid to obovoid, rounded to entire or slightly undulate, tube long-exserted, species. Mountains of Afghanistan, C Asia, Mon stigma-lobes subequal; nutlets oblong, smooth, acuminate, smooth to tuberculate, glabrousor widely and abruptly dilated at throat; stamens golia and China. Some species used medicinally. glabrous. Four or five species. High alpine screes hairy at apex, mucilaginous or not. 2« = 14,16, shorter than corolla, filaments parallel, thecae par of the Himalaya and China. 18, 30, 32, 34, 36, 54. Over 200 species. Mainly in allel, papillose; stigma-lobes subequal; disc 4- mountains, semi-deserts and steppes of Eurasia, 181. Hymenocrater Fisch. & C.A. Mey. lobed; nutlets orbicular to elliptic, hairy, areole extending into North Africa. Some cultivated 183. Agastache J. Clayton ex Gronov. orbicular. In = 18. About six species. Temperate or Hymenocrater Fisch. & C.A. Mey., Index Sem. Hort. Bot. subtropical eastern Asia, one species in N America. for medicinaluse or ornament. For a synopsis Petropol. 2:39 (1835); Budantsev, Bot. Zhurn. 77,12:118-128 of infrageneric classification, see Budantsev Agastache J. Claytonex Gronov., Fl. Virgin.: 88 (1762); Lint & (1992), rev. Epling, Am. Midi. Nat. 33:207-230 (1945), rev.; Sanders, Syst. (1993). Bot. Mon. 15: 1-92 (1987), rev. section; Budantsev, Bot. Shrubs, subshrubs or perennial herbs, aromatic; Zhurn. 78,2:106-115 (1993), rev. leaves simple, strongly toothed; inflorescence Dekinia M. Martens & Galeotti (1844). Labiatae 252 R.M.Harley etal. 253 185.Glechoma L. nousor not; areole arcuate. In= 10,12,14,18,20, abruptlydilated towards throat; stamens attached dilated atthroat; stamens 4,didynamous, exserted 36,72.About 70 species. Usually montane, Eurasia, nearthroat, included under posterior lipof fromtube, anterior pair longer, filaments ± GlechomaL,Sp. PL 2: 578 (1753); Budantsev, Bot.Zhurn. 77, N Africa(one species), N America (one species). corolla,filaments parallel, thecaedistinct, parallel; straight, thecae divergent, confluent atbase; 12:118-128(1992), rev. Somespecies medicinal, ornamental orbee plants. stigma-lobes disc subequal;4-lobed; nutlets ellip stigma-lobes subulate, subequal; nutlets ovoid, MeehaniopsisKudo(1929). Threesubgenera after Budantsev (1987): Draco tic,smooth, glandular-pubescent, areoleorbicular. smooth, mucilaginous. In= 32,34,64. Fourspecies, cephalum(stamens included, anthers glabrous - 2n= 10.One species, Cedronella canariensis (L.)usually inforests, Europe, N Africa, Macaronesia Stoloniferousperennialherbs, not aromatic, often sevensections), Fedtschenkiella (Kudr.)Schischk. Webb& Berth.,Canary Is.and Madeira. andAsia. Plants used as a source ofessential oils, gynodioecious;leavessimple; inflorescence of (stamensexserted, anthers glabrous), Ruy- bothmedicinally andas flavouring. distant,few-flowered cymes,shortly pedunculate schiana(Mill.) Briq.(stamens included, anthers tosessile, inaxils ofleaf-like bracts,bracteoles 189.Schizonepeta (Benth.) Briq. pubescent). subulate,inconspicuous; calyxtubular tocampan- 191.Heterolamium C.Y.Wu Schizonepeta(Benth.) Briq.in Engler & Prantl, Nat.Pflanzen- ulate,slightly curved near throat, indistinctly fam.IV, 3a: 235 (1896); Budantsev, Bot.Zhurn. 78,2:106-115Heterolamium C.Y.Wu, Acta Phytotax. Sin.10: 254 (1965). 2-lipped,5-lobed (3/2), lobes equal, broadly 187.Lallemantia Fisch.& C.A. Mey. (1993),rev. triangulartolinear; corolla strongly 2-lipped Slenderperennial herb;leaves cordate, toothed, LallemantiaFisch.& C.A. Mey., Index Sem. Hort. Bot.Petropol. (2/3),blue-violet, lilacorpink, posterior lip Annualorperennial herbs, aromatic; leaves entire long-petiolate; inflorescence secund, of 1-3- 6:52 (1839). straight,2-lobed,anterior lipwith median lobe flat tobipinnate; inflorescence thyrsoid,cymes sessile, flowered cymes, shortly pedunculate tosessile, in orconcave, flabellate orreniform, marginundulate terminal,crowded into spike-like, usually Annualorbiennial herbs; leaves simple; inflores axilsofreduced bracts, forming anelongate, ter orentire, tubelonger than calyx, gradually or unbranchedheads; bracts, except sometimes cencethyrsoid, terminal, spike-like oroblong, minalthyrse; bracteoles inconspicuous; calyx abruptlydilated towards throat; stamens ofher thelowest, reduced, sometimes broadly ovate, ofteninterrupted, cymesfew-flowered, sessileor oftenpurple-tinged, cylindrical tocampanulate, maphroditeflowers exserted fromtube, included coloured,orsubulate, inconspicuous, bracteoles rarelyonvery short peduncles inaxils ofleaf-like strongly2-lipped, 5-lobed, (3/2),posterior lipwith underlipof corolla, inserted nearthroat, filaments subulate,inconspicuous; calyxstraight orweakly bracts,bracts ovate tolinear, weakly toothed to medianlobe broadly ovate, rounded at apex, parallel,thecae divergentat90°; nutlets oblong- oblique,actinomorphic or2-lipped, 5-lobed, lobes entire,rarely aristate above, bracteoles ovateto slightlyspreading andweakly decurrent ontube at ovoid,smooth, areole basal, orbicular orelliptic. ovate-oblongtolanceolate, acuteorattenuate into rotundor sometimeslinear, aristate-toothed; base,lateral lobes shortly triangular-acuminate, In= 18,24, 36. Four to eight species. Mostly in spinuloseawns, sinuses between anterior teeth pedicelsflattened, erect;calyx tubular, straight or anteriorliplonger, with sinus between lobes grasslands,forestmargins and stream sides of withswollen folds at base; corolla strongly 2- curved,2-lipped, 5-lobed (3/2), lobes unequal, tri shorter,tubeslightly gibbous atbase, 15-nerved, temperateEurasia; cultivated inN and S America. lipped,5-lobed (2/3), white or blue-violet, poste angular,aristate, posterior lipwith median lobe annulatewithin, below throat; corolla white, anteriorlipwith median lobe broadest,sinusbetween calyx-lobes withathick riorlip straight, strongly2-lipped, 5-lobed (4/1), posterior lipof 4 186.Dracocephalum L. flator scarcely concave, entire to crenate,tube enedfold ornot, throat closed byconnivent lipsin short,rounded, subequal lobes, anterior liplonger, includedorscarcely exserted fromcalyx, gradually fruit;corolla strongly 2-lipped (2/3),posterior lip withlobe slightly concave, rounded, tubenarrowly DracocephalumL,Sp. PL 2: 594 (1753); Budantsev, Bot.Zhurn. wideningabove; stamens with anterior pair cylindrical, straight, exserted from calyx, and 72,2:260-267 (1987), rev. with2longitudinal foldsinside, anterior lipwith antrorselybent,posterior pairarcuate-ascending MoldavicaFabr.(1759). laterallobesdirected downwards, tubewidening abruptlyexpanded above, exannulate; stamens underupper lip, thecae divergent at90° or at an above;stamens included under corolla-lip, 4,didynamous, long-exserted, ascending under acuteangle; stigma-lobes subequal; disc4-lobed; Herbs,annual orperennial, orsubshrubs, aro filaments parallel, thecaeglabrous, divergent at upperlip, later becoming slightly divergent, poste nutletsoblong toelliptic, smooth,mucilaginous or matic;leaves entire, dentate or pinnatisect;180°;stigma-lobes subequal; disc4-lobed; nutlets riorpair longer atearly anthesis, anterior pair not.In = 12.Three species. Steppes ofsouthern inflorescencethyrsoid, usually terminal,con oblong, trigonous, smooth, mucilaginous. In= 14. apparentlyextending later, anthers 2-thecous, Siberia,Mongolia and China. densed,spike-like orcapitate, cymessessile or Five species. Mediterranean, SW andC Asia. Some thecaeelliptic, widelydivaricate, confluentat apex; styleexserted, stigma-lobes subequal, subulate, pedunclesveryshort; bracts entire orlobed, medicinal, seedoils. usuallyaristate, sometimes coloured orleaf-like, spreading;disc4-lobed, anterior lobe elongate; bracteolesoftensimilar tobracts butsmaller, or VII.2.D.Tribe Mentheae: Incertae Sedis nutletsoblong-ovoid, smoothto finely striate, 188.Cedronella Moench subulate,oftenaristately toothed;calyx tubular to withsmall, elliptic areole. Onespecies, H.debile 190. campanulate,straightor slightly curved, 2-lipped, Cedronella Moench, Meth. PL: 411 (1794); Budantsev, Bot. (Hemsl.)C.Y.Wu, with several varieties, China. 5-lobed(3/2), lobes aristate, oftenupper lip with Zhurn.78,2:106-115 (1993),rev. MelissaL.,Sp. PL 2: 592 (1753). lobesequal ormedian lobe clearly largerthan lat VII.3.Tribe Ocimeae Dumort. (1829). erals,lower lipwith lobes shorter, sinuses between Perennial herbs, aromatic; leaves 3(-5)-foliolate, Perennialherbs, ± rhizomatous, aromatic; leaves lobeswith swollen folds atbase, throat glabrous dentate; inflorescence thyrsoid,terminal, spike- simple,toothed; inflorescence ofaxillary verticil- Aromaticherbs, shrubs orrarely trees. Stamens within;corolla 2-lipped, 5-lobed (2/3), blue to likeorovoid, cymes shortly pedunculate tosessile; lasters,widelyspaced, with2-16 flowers perverti- declinatetoporrect (Fig. 29), anthers synthecous, violet,pinkor white, posterior lipstraight or bracts usually small, linear, coloured, bracteoles cillaster;bractsleaf-like, bracteoles small,patent; dorsifixed.Discwith lobes alternate toor opposite slightlyconcave, subentire orbifid, anterior lipinconspicuous; calyxtubular, slightly curved and flowerswith curved pedicels; calyx2-lipped, 5- ovary-lobes. withmedian lobe largest, tubegradually or dilated towards throat, actinomorphic, campanu lobed(3/2), ±campanulate, lobesupcurved,ciliate, abruptlywidening frombase; stamens included latein fruit, 5-lobed, lobesequal, narrowly trian posteriorlipwith lobes very short, lower lipwith 1.Inflorescence congested,spiciform, terminal atthe end of underposterior corolla-lip, rarelyexserted, paralgular, acute, ± spine-tipped, throatopen, slightly lobeslong-triangular, tube13-nerved; corolla 2- a longaxis (but see Hyptis). Posterior calyx-lobe often oblique;corolla strongly 2-lipped, 5-lobed (2/3), lel,thecae pilose orglabrous, divergent at180°; appendaged.Stamens included incorolla-tube. Disc-lobes lipped,hairy,white, cream orreddish, posterior lip stigma-lobessubequal; disc4-lobed; nutlets pinkish, posterior lip±flattened, emarginate, ante oppositeovary-lobes a.Lavandulinae: 192.Lavandula shorter,withlobe rounded and emarginate, ante - Inflorescencethyrsoid.Posterior lobesometimes larger ovoid-oblong,weaklytrigonous, apicallytruncate- riorlip with median lobe larger, obovate, margin riorlipwith median lobe semi-orbicular, deflexed, thanothers, sometimes decurrent on tube, but not roundedorslightly flattened, smooth,mucilagi undulate, concave, tube exserted from calyx, laterallobesvery small, ± triangular, corolla-tube appendaged. Stamens exserted fromtube (except Endoste- R.M. Harley et al. 274 Labiatae 275 Ryding, O. 2002. Myxocarpy in the Nepetoideae (Lamiaceae) Trudel, M.C.G., Morton, J.K. 1992. 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