Bulletin of The Natural History Museum Botany Series BRITISH MU8&M (NAWttL PRiOTWTBD GENERAL LIBRARY THE NATURAL HISTORY MUSEUM VOLUME 23 NUMBER 2 25 NOVEMBER 1993 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Dr R. Huxley Assistant Editor: Mrs M.J. West Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum's resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (0264) 334748 Fax: (0264) 334058 Wo rid List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) The Natural History Museum, 1993 Botany Series ISSN 0968-0446 Vol. 23, No. 2, pp. 55-177 The Natural History Museum Cromwell Road London SW7 5BD Issued 25 November 1993 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford Bull. not. Hist. Mus. Land. (Bot.) 23(2): 55-59 Issued 25 November 1993 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region BRITISH 3 DEC 1993 TING-NUNG HO AND SHANG-WU LIU Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China PRESENTED GENERAL LIBRAR SYNOPSIS. Eleven new species* and two new varieties from Western China and the Himalayan region are described. They are G. leucantha H. Smith, G. chateril. N. Ho, G. hicksii H. Smith, G. masonill. N. Ho, G. zekuensisT. N. Ho et S. W. Liu, G. laxiflora T. N. Ho, G. depressa var. stenophylla T. N. Ho, G. bryophylla H. Smith, G. micans var. latifolia T. N. Ho, G. glabriuscula H. Smith, G. winchuanensis T. N. Ho, G. shaanxiensis T. N. Ho and G. subuliformis S. W. Liu. The following three new names are also made: G. himalayaensis T. N. Ho, G. alata T. N. Ho and G. taiwanica T. N. Ho. *Among them three species were diagnosed, described and typified and one species labelled by Harry Smith but were left unpublished at his death in 1971. NEW TAXA 1. Gentiana leucantha H. Smith, sp. nov. (Sect. Otophora Kusn.) Species Gentiana decor atae Diels et Gentiana infelici C. B. Clarke affinis, sed a prima tubo corollae lobis multo longiore (nee subaequilongo), a secunda flore duplo majore, ab ambo- bus flore albo (in sicco lutescenti) bene differt. Planta perennis, omnino glabra, e collo radicem principalem carnosulam, fusiformem et caules floriferos numerosos edens. Caules floriferi perennes, assurgentes, breves, dense parvifoliati. Folia rosularia decunt, caulina sessilia, interno- diis multo longiora, ovata, 3.5-4 mm longa, 1.8-2 mm lata, apice acuta, submucronata, margine cartilagineo- microscabridula. Flores solitarii, terminales, erecti, sessiles; tubus calycis campanulatus, 5-6 mm longus, lobis ellipticis, 3-3.5 mm longis, c. 2 mm latis, apice acuminatis, mucronatis; corolla alba (in sicco lutescens), calyce triple vel ultra lon- gior, tubo subcylindrico, 13-15 mm longo, lobis ovatis, 4-5 mm longis, fere 4 mm latis, obtusis, plicae lobis later- aliter adnatae, c. 2.5 mm longae et 1.5 mm latae, bifidae, laciniatae vel fere integrae subtruncataeque; stamina tubo corollae c. 1 mm alte affixa, ex ore corollae exserta, filamen- tis filiformibus, 6-8 mm longis, antheris linearibus, vix 2 mm longis; ovarium ellipticum, 8-10 mm longum, apice in stylum c. 1 mm longum attenuatum, stipite c. 2 mm longo. Capsula inclusa, c. 11 mm longa; semina brunnea, oblonga, 1.1 X 0.4 mm magna, tenuiter reiculata. China: S Tibet: Tsari, Karkyn la, on open grassy hillside, 4420m, 24 June 1936, Ludlow & Sherriff 2209 (BM) ; ibid, on mossy open hillside, 4CKXM270 m, 11 October 1938, Ludlow, Sheriff & Taylor 6592 (BM); Chayul chu, Kashong la, 4420 m, 14 July 1936, Ludlow & Sherriff 2356 (BM); SE Tibet: Langgong, 2845' N 94E, on open grassy hillside, 4420 m, 5 June 1938, Ludlow, Sherriff& Taylor 3963 (BM, E); Kongbo, Kusha la near Paka, 2915' N 942' E, 4270-4570 m, 25 July 1938, Ludlow, Sherriff& Taylor 5934 (BM-holotype); ibid, on open grassy meadows, 4270 m,' 27 July 1938, Ludlow, Sherriff & Taylor 5951 (BM, E); ibid. Deyang la, on stony grassy slopes, 4115 m, 8 August 1947, Ludlow, Sherriff & Elliot 14273 (BM, E). NE Bhutan: Me la, 4420 m, 5 August 1933 Ludlow & Sherriff 415 (BM); ibid. 4270 m, 26 August 1949, Ludlow, Sherriff & Hicks 21104 (BM, E, K); Ju la, Mangde chu, 19 July 1949, Ludlow, Sherriff & Hicks 16886 (BM,E). 2. Gentiana chateri T. N. Ho, sp. nov. (Sect. Otophora Kusn.) Species ob folia rosularia linearia et corollam albam usque flavescentem Gentiana damyonensi Marquand et Gentiana hicksii H. Smith similis, sed a prima corolla epunctata, lobis tubo brevioribus, foliis caulinis ovato-lanceolatis, a secunda corolla minore, 18-20 mm longa, ab ambobus floribus 3-4 in cymam laxam dispositis, lobis calycis irregularibus, oblan- ceolatis et spathulatis, basi contractis recedit. Herba perennis, 3-4 cm alta, e collo radicem principalem, caules floriferos et rosulem basalem emittens. Radix carno- sula, verticalis, fusiformis, 4-6 mm diam. Caules floriferi 1^4, ascendenti-erecti, glabri. Folia rosularia linearia vel lineari- oblonga, 8-22 mm longa et 2-4 mm lata, apice acuta vel obtusa, basi attenuata, nervis 1-3, subtus prominentibus, petiolis c. 2 mm longis; ilia caulina 2-3-juga, remota, ovato- lanceolata, 7-18 mm longa et 3-7 mm lata, apice acuta, basi breviter petiolata. Flores saepe 3^ in cymam laxam dispositi, raro solitarii; pedicelli 4-5 mm longi; calyx 7-9 mm longus, tubo tubuloso, lobis irregularibus, obovatis et lineari- spathulatis, 3-4 mm longis et 0.5-1.5 mm latis, apice acutis vel obtusis, basi contractis; corolla alba (flavescens in sicco), atrocaeruleo-striata, tubulosa, 18-20 mm longa, lobis tubo brevioribus, oblongis, 6-7 mm longis, acutis, plicae obliquae, auriculatae, lobis lateraliter adnatae; stamina tubo corollae inferioris inserta, filamentis subulatis, c. 10 mm longis, ad basin breviter ampliatis, antheris luteis, c. 1 mm longis; ovarium lineare, c. 15 mm longum, stylo c. 5 mm longo, stigmatibus linearibus, recurvatis. Nepal: Iswa Khola, 4000 m, 8 August 1971, L. W. Beer, C. R. Lancaster & D. Morris 9545 (BM), Kasuwa khola, 4000 m, 23 August 1975, L. W. Beer 25363 (BM-holotype). We take particular pleasure in naming this species after Mr The Natural History Museum. 1993 56 Arthur O. Chater, former curator of the flowering plant herbarium, The Natural History Museum, who worked for some years on the Nepalese Flora. 3. Gentiana hicksii H. Smith, sp. nov. (Sect. Otophora Kusn.) Species Gentiana damyonensi Marquand affinis, sed corolla majore, 25-35 mm longa, tubo lobis longiore (nee breviore) differt. Herba perennis, c. 8 cm alta, e collo radicem principalem, caules floriferos et rosulam basalem emittens. Radix carno- sula, verticalis, fusiformis, 3-5 mm diam. Caules floriferi 1-3, ascendenti-erecti, glabri. Folia rosularia linearia vel lanceolato-linearia, 3-4 mm longa et 2-3.5 mm lata; ilia caulina 3-4-juga, internodiis breviora vel interdum sub- aequilonga, lanceolato-linearia, 0.7-1.2 cm longa, acuta, ses- silia. Flores saepe solitarii, terminales; calyx 7-9 mm longus, tubo obconico, lobis subregularibus, linearibus; corolla alba, coeruleo-striata, cylindrico-obconica, 25-35 mm longa, tubo c. 19 mm longo, lobis ovatis, 8-9 mm longis et 5 mm latis, obtusis, plicae dentiformes, latere lobi adnata; stamina tubo c. 6 mm alte affixa, filamentis 13-15 mm longis, antheris angustis, vix 2 mm longis; ovarium lanceolatum, c. 15 mm longum, apice in stylum c. 4 mm longum attenuatum, stipite 2-3 mm longo. Capsula et semina non visa. Bhutan: Pang la, on open wet slopes, 4000 m, 21 September 1949, Ludlow, Sherriff& Hicks 21456 (BM - holotype; UPS - isotype). 4. Gentiana musonii T. N. Ho, sp. nov. (Sect. Otophora Kusn.) Fig. 1. Species Gentiana otophorae Franchet ex Forbes & Hemsley affinis, sed corolla omnino coerulea, epunctata bene differt. Herba perennis, omnino glabra, 14-18 cm alta, e collo radi- cem principalem, caules floriferos et rosulam basalem emit- tens. Radix carnosula, cylindrica, 4-7 mm diam. Caules floriferi 1-3, ascendenti erecti. Folia rosularia oblongo- spathulata vel oblonga, 3-7.5 cm longa et 0.9-1.9 cm lata, apice acuta vel obtusa, basic in petiolos 2.5-6 cm longos attenuata, nervis 3-5, subtus prominentibus, ilia caulina 2-3-juga, remota, oblonga vel ovato-elliptica, 1.4-2.6 cm longa et 0.7-1.2 cm lata, apice obtusa vel acuta, basi in petiolos 3-7 mm longos attenuata. Flores 3-4 in cymam laxam dispositi; calyx 6-8 mm longus, lobis irregularibus, spathulatis, 1.5-2.5 mm longis apice rotundatis vel acutis, basi contractis; corolla caerulea, epunctata, cylindrica, 16-20 mm longa, lobis tubo longioribus, oblongis, 13-16 mm longis et c. 5 mm latis, apice acutis, plicae minimae, c. 1 mm longae, auriculatae, lobis lateraliter adnatae; stamina medio corollae inserta, filamentis subulatis, c. 10 mm longis, anth- eris caeruleis, c. 1.5 mm longis; ovarium cylindricum, 11-13 mm longum, breviter stipitatum, stylo breve, stigmate breve. Upper Burma: N'Maikha-Salwin divide, 2628' N 9848' E, September 1924, Forrest 24944 (K); ibid. 2650' N 9848' E, on alpine meadows, 4270 m, September 1925, Forrest 27222 (BM - holotype; K - isotype). This new species is dedicated to my friend, Prof. Howard S. Mason, a member of American Rock Garden Society. He has been devoting himself to cultivated Gentians of different Gentian seedlings. T.-N. HO AND S.-W. LIU 5. Gentiana zekuensis T. N. Ho & S. W. Liu, sp. nov. (Sect. Cruciata Gaudin) Fig. 2. Species habitu Gentiana officinali H. Smith valde similis, sed tubo calycis obtuso (nee truncate), lobis filiformibus, usque ad 5 mm longis (nee dentiformibus, 1-1.5 mm longis), corolla flavida, caeruleo-punctata, versus limbum caerulea, raro corolla omnino caerulea (nee omnino flavida) differt. Herba perennis, 15-40 cm alta. Radices subcarnosae, pau- cae, in radicem verticalem, cylindricam conniventes. Caules floriferi 3-10, ascendentes, caudice reliquis petiolorum bru- neis, fibrosis obtecto. Folia rosularia anguste elliptica vel lineari-lanceolata, 8-22.5 cm longa et 0.5-2.5 cm lata, apice acuta, basi attenuata, utrinque glabra, nervis 3-5, subtus prominentibus; ilia caulina 2-3-juga, elliptica vel lanceolata, 1.8-5.5 cm longa et 0.6-1.2 cm lata, apice acuta, basi in vaginam 0.8-3 cm longam connata, eis summis bractescenti- bus, lanceolatis usque lineari-lanceolatis, 1.2-5 cm longis et 0.5-1 cm latis, acuminatis. Flores sessiles, numerosi, termina- les axillaresque; calyx fissus fere ad basin, 6-13 mm longus, tubo spathaceo, apice obtuso, lobis filiformibus, usque 5 mm longis; corolla flavida, caeruleo-punctata, versus limbum caerulea, raro corollla omnino caerulea, tubulosa, 18-20 mm longa, lobis ovatis, 2.5-3 mm longis, acutis, plicae obliquo- triangulatae, c. 1 mm longae, acuminatae; stamina tubo corollae inferioris inserta, filamentis linearibus, 7-8 mm longis, antheris flavidis, linearibus, c. 1.5 mm longis; ovarium lanceolatum, 11-13 mm longum, stylo cum stigmatibus c. 2 mm longo. The corollas of G. zekuensis are yellowish, bluish towards the limb, seldom whole corolla bluish. G. zekuensis is somewhat intermediate between G. macrophylla and G.officinalis but closer to the latter. It differs from both mainly by the distinct and filiform calyx lobes. China: Qinghai. Zeku, among shrubs, 3600 m, 24 July 1967, Y. C. Young 1822 (HNWP); ibid, on grassy hillside, 3200 m, 27 August 1970, L. H. Zhou & L. N. Sun 1958 (HNWP - holotype); ibid, among shrubs, 3300 m, 14 August 1975, Z. B. Wang 349 (HNWP); Tongren among shrubs, 3400 m, 26 July 1970, 5. W. Liu 1474 (HNWP). 6. Gentiana laxiflora T. N. Ho, sp. nov. (Sect. Frigida Kusn.) Species G. trichotomae Kusn. et G. erecto-sepalo T. N. Ho affinis, ab hac corolla 22-25 mm longa, lobis calycis angustis, linearibus et subulatis, sinubus inter lobos calycis rotundatis, ab ilia corolla caerulea, versus basin flavescente differt. Herba perennis, 10-14 cm alta. Rhizoma breve, horizontale vel verticale, radices adventitias, caules floriferos et rosulam basalem emittens. Caules 1-3, erecti, subglabri, basi vaginis veteribus membranaceis obtecti. Folia plerumque basilaria, petiolata, laminae lineari-oblongae, 2.5-8 cm longae et 3-6 mm latae, apice acutae vel obtusae, basi attenuatae; ilia caulina 1-2 juga, minora, sessilia. Flores 5-7, terminales et axillares, in florescentiam laxam dispositi; pedicelli graciles, 5-21 mm longi; calyx tubulosus, 8-11 mm longus, lobis angustis, linearibus et subulatis, 3-6 mm longis, sinubus inter lobos calycis rotundatis; corolla caerulea, versus basin flave- scens, infundibuliformis, 2-25 mm longa, lobis ovatis, 1.5-3 mm longis, acutis, plicae truncatae, lobis breviores; stamina tubo corollae inferioris inserta, filamenti subulatis, c. 10 mm longis, antheris luteis, c. 2 mm longi. Capsula breviter NEW TAX A OF GENTIAN A 57 cm cm cm Fig. 1 Gentiana masonii T. N. Ho: a, habit ; b, cauline leaves ; c, calyx opened out; d, corolla opened showing stamens. stipitata, ex corolla paullo exserta. China. SE Tibet. Nam la, 2959' N 9417' E, 4115 m, 30 August 1938, Ludlow, Sherriff & Taylor 6944 (BM), Kongbo, 30N 9420' E, 4115 m, 20 July 1947, Ludlow, Sherriff & Elliot 15492 (BM - holotype; E - isotype). 7. Gentiana himalayaensis T. N. Ho, stat. et nom. nov. Gentiana nubigena var. parviflora C. B. Clarke in Hook, f., Fl. Brit. India 4: 117 (1883). Type: Sikkim, Kinchinjhow, 4880-5200 m, /. D. Hooker (K - holotype!; BM - iso- type!) Gentiana algida var. parviflora (C. B. Clarke) Kusn. in Acta Horti Petrop. 15: 266 (1898). DISTRIBUTION. C. & E. Himalayan region. China (S. & SE Tibet), Nepal, Sikkim, Bhutan. 8. Gentiana depressa D. Don var. stenophylla T. N. Ho, var. nov. (Sect. Isomeria Kusn.) A var. typo recedit foliis angustis, lineari-oblongis vel lan- ceolatis, 1-3.5 cm longis et 2.5-4(-8) mm latis, acuminatis. Leaves linear-oblong or lanceolate; calyx-lobes narrowly tri- angular or linear-elliptical. Nepal: East of Chalike Pahar, 4000 m, Stainton, Sykes & Williams 4593 (BM); Moktintan, on grassy slopes near Pass, 4000 m, 1 October 1954, Stainton, Sykes & Williams 8060 (BM); Phagune Dhuri, 4000 m, 13 October 1954, Stainton, Sykes & Williams 9007 (BM); Kali Gandaki, Tasang/ESE of Tukche, 3840 m, 4 November 1976, G. Miehe 194 (BM); S. side Deorali ridge, 2900 m, 11 November 1979, A. D. Schilling 2445 (BM - holotype). 9. Gentiana alata T. N. Ho, nom. nov. Gentiana kusnezowii Franchet in Bull. Soc. hot. Fr. 43: 492 (Nov. 1896), non Gilg May 1896 (in Engler, Bot. Jahrb. 22: 325). Type: China, Yunnan, Yunnansen (Kunming), Delavay s.n. (P - holotype!). DISTRIBUTION. China (C. Yunnan). 10. Gentiana bryophylla H. Smith, sp. nov. (Sect. Chondro- phylla Bunge) Species habitu, forma folii et colore magnitudineque floris Gentiana burmensi Marquand persimilis, sed plicis corollae margine crenulatis, efimbriatis valde differt. Herba perennis, c. 5 cm alta, stolone breve praedita. Caulis ascendens vel erectus, uniflorus, subglaber. Folia basilaria minuta, 2-3 mm longa, 1-1.5 mm lata; ilia caulina 6-7-juga, inferiora minora, superiora ad 5 mm longa et 2 mm lata, lanceolata, recurvato-aristata, margine cartilaginea, subgla- braque, versus basin gradatim dense ciliolata. Tubus calycis 58 T.-N. HO AND S.-W. LIU cm cm Fig. 2 Gentiana zekuensis S. W. Liu: a, habit ; b, flower; c, calyx opened out; d, corolla opened inside with stamens and gynoecium. 6-7 mm longus, lobis 3.5 mm longis, acicularibus, subarista- tis; corolla purpurea, c. 17 mm longa, tubo c. 15 mm longo, lobis paullo obliquis, rotundato-triangularibus, acutis, apicu- latis, 2.5 mm longis et latis, plicae lobis paullo breviores, subrotundatae, margine crenulatae; stamina tubo corollae 6 mm alte adnata, filamentis filiformibus, 4.5 mm longis, antheris c. 1 mm longis; ovarium stipitatum, obovato- ellipsoideum, c. 5 mm longum, stylo fere 2 mm longo. Cap- sula amplanato-obovata, c. 5 mm longa, alata; semina brunnea, subangulariter ellipsoidea, 0.7 X 0.3 mm magna, reticulata. Burma: Nam Tamai valley, 28 N 9745' E. , on steep slopes in the mossy carpet underneath Arundinaria and Rhododen- dron, 3000 m, 8 September 1937, Kingdon Ward 13184 (BM - holotype); ibid. 9 Septemper 1937, Kingdon Ward 13208 (BM). 11. Gentiana in leans C. B. Clarke var. latifolia T. N. Ho, var. nov. (Sect. Chondrophylla Bunge) Gentiana melaensis H. Smith in herb. A var. typo recedit foliis caulinis latioribus, ovato-lanceolatis usque lineari-lanceolatis, patentibus (nee acicularibus, caule fere adpressis), lobis calycis brevioribus, latioribus, breviter acutis (nee longe acuminatis), plicis bifidis (nee integris minute laciniatis). Bhutan: Liaru Thang, Ringchen chu, 3750 m, 8 June 1937, Ludlow & Sheiriff 3531 (BM); Thampe la, on bare hillside, 4570 m, 14 August 1949, Ludlow, Sherriff & Hicks 17134 (BM), Shingbe (Me la), 4115 m, 8 September 1949, Ludlow, Sherriff & Hicks 21166 (BM - holotype; UPS - isotype). . 12. Gentiana glabriuscula H. Smith ex T. N. Ho, sp. nov. (Sect. Chondrophylla Bunge). Species Gentiana pedicellatae (G. Don) Griseb. similis, sed planta omnino glabra, foliis basilaribus majoribus, involucri- formibus, orbicularibus vel late oblongis, illis caulinis minori- bus, ovatis valde differt. Planta annua vel biennis, 2-5 cm alta, omnino glabra. Caulis gracilis, ascendens, e basi ramosus quasi caespitosus. Folia basilaria majora, involucriformia, orbicularia vel late oblonga, 9-26 mm longa, 7-15 mm lata, apice rotundata, mucronata, basi in petiolos 3-5 mm longos abrupte contracta, nervis 1-3, utrinque prominentibus; ilia caulina minora, ovata, 3-5 mm longa et 2-3.5 mm lata, apice acuta mucronataque, basi leviter attenuata, subsessilia. Flores sessiles, solitarii ad apices ramu- lorum siti; calyx campanulatus, 3.5-5 mm longis, lobis ovatis, 2-3 mm longis et 1.5-2 mm latis, apice acutis mucronatisque, basi contractis, recurvatis; corolla caeruleo-grisea, extra atro- caerulea, fauce nigro-punctata, campanulata, 5-7 mm longa, calycem paullo superans, lobis ovatis, c. 2 mm longis, apice NEW TAXA OF GENTIAN A 59 acutis mucronatisque, plicae late ovatae, c. 1 mm longae, lobis breviores, acutae, integrae vel bilobatae; stamina medio corol- lae inserta, filamentis c. 1 mm longis; ovarium obovato- oblongum cum stipite c. 4 mm longum, stylo breviore, stigmatibus oblongis. Bhutan: Trashi Yanasi chu, on mossy rocks and banks in mixed forest, 2590 m, 5 September 1949, Ludlow, Sherriff & Hicks 20608 (BM - holotype). Sikkim: Gangtok-Karponang road, F. H. Lister 9 (K). China: S Tibet: Monyul, 2 April 1936, Ludlow & Sherriff 1264 (BM). 13. Gentiana taiwanica T. N. Ho, stat. et nom. nov. Gentiana scabrida var. angusta Masam. in Trans, nat. Hist. Soc. Formos. 29: 64 (1939). Type: China, Taiwan, Nan- hutashan, Taihoku-syu, 15 July 1931, Masamune & K. Mori s.n. (herb. Taihoku Imperial Univ. Taiwan - holo- type, not seen). Gentiana angusta (Masam.) Liu et Kuo in Bull. Exp. Forest Natn. Taiwan Univ. 114: 176, pi. 4 (1974), non M. E. Jones 1908 (in Contrib. Western. Bot. 12: 52). DISTRIBUTION. China (Taiwan). 14. Gentiana winchuanensis T. N. Ho, sp. nov. (Sect. Chon- drophylla Bunge). Species Gentiana piasezkii Maxim, capsula anguste oblonga, calyce carinato-alato af finis, sed foliis ovatis, lobis calycis triangularibus, multo brevioribus bene recedit. Herba annua, 8-10 cm alta. Caulis ascendens, ramosus. Folia basilaria ignoti; ilia caulina remota, internodiis breviora, ovata, 4-6 mm longa et 2-4 mm lata, apice acuta, basi in petiolos 2-3 mm longos abrupte contracta, margine carti- laginea ciliolataque, utrinque glabra. Flores subsessiles, soli- tarii ad apices ramulorum siti, calyx tubuloso- infundibuliformis, 13-17 mm longus, lobis triangularibus, 1.5-2 mm longis, basi c. I mm latis, acuminatis, margine cartilagineis, sinubus rotundatis; corolla caeruleo-purpurea, fauce flavida et nigro punctata, anguste infundibuliformis, 25-30 mm longa, lobis ovatis, 6-7 mm longis, apice acutis mucronatisque, plicae ovatae, lobis paullo breviores, denticu- latae, stamina medio corollae inserta, filamentis filiformibus, c. 5 mm longis, antheris c. 1 mm longis. Capsula inclusa, fusiformis, c. 10 mm longa, alata, stipite c. 5 mm longo; semina brunnea, oblonga, c. 1 mm longa, tenuiter reticulata. China: N Sichuan. Winchuan, 2400 m, in forest, 30 July 1975, Sichuan Veget. Exped. 8522 (HNWP - holotype). 15. Gentiana shaanxiensis T. N. Ho., sp. nov. (Sect. Chon- drophylla Bunge). Species habitu Gentiana piasezkii Maxim, similis, sed foliis caulinis linearibis lobis calycis linearibus, usque ad fauce corollae, lobis corollae acuminato-caudatis differt. Herba annua, 7-12 cm alta. Caulis purpureus, ascendens, dense papillosus, e basi ramosissimus. Folia apice acuta vel acuminata, margine dense papillosa, utrinque glabra, costis subtus prominentibus; ilia basilaria majora, anguste oblonga, 25-42 mm longa et 3.5-8 mm lata, sessilia vel subsessilia; ilia caulina linearia, 9-22 mm longa et 1.5-3 mm lata, sessilia. Flores solitarii ad apices ramulorum siti; pedicelli 2-10 mm longi, dense purpureo-papillosi, calyx tubuloso infundibuli- formis, 20-27 mm longus, usque ad faucem corollae, tubo carinato alato, lobis linearibus, 6-8 mm longis; corolla pur- purea, hypocrateriformis, 25-35 mm longa, lobis anguste lanceolatis, 5-9 mm longis, basi 2-3 mm latis, apice acuminato-caudatis, plicae ovatae, 4-5 mm longae et c. 3 mm latae, acutae, denticulatae; stamina medio corollae inserta, filamentis filiformibus, 50-55 mm longis, antheris linearibus, c. 1 mm longis; ovarium ellipticum, 7-8 mm longum, stylo cum stigmatibus 4-5 mm longo. Capsula e corolla exserta, spathulato-oblonga, 9-11 mm longa, alata, stipite corollam aequante vel superante; semina brunnea, oblonga, 1-1.2 mm longa, tenuiter reticulata. China: Shaanxi: Ningqiang, 11 July 1957, Northwest Univ. Biol. Exped. 57 (HNWP - holotype). 16. Gentiana subuliformis S. W. Liu, sp. nov. (Sect. Frigida Kusn.) Species Gentiana wilsonii Marquand similis, sed foliis basi- laribus latis, anguste ellipticis vel oblongis, corolla multo minore, 23-25 mm longa differt. Herba perennis, 12-20 cm alta. Rhizoma breve, horizontale vel verticale, radices adventitias carnosulas, caules floriferos et rosulam basalem emittens. Caules floriferi erecti, basi c. 2 mm diam., vaginis veteribus membranaceis obtecti. Folia plerumque basilaria, petiolata, petiolis 0.5-2 cm longis, lami- nis anguste ellipticis vel oblongis, raro oblongo-spathulatis, 1.5-3.5 cm longis et 0.5-0.9 cm latis, obtusis vel subrotunda- tis, basi attenuatis, nervis 3, subtus prominentibus; ilia caulina 1-2-juga, eis basilaribus similaria, 2-3.5 cm longa et ad 1.2 cm lata, sessilia vel breviter petiolata. Flores numerosi, terminales, aggregati in inflorescentiam subcapi- tatam; calyx tubulosus, c. 10 mm longus, saepe fissus, lobis recurvatis vel patentibus, subulatis, 1-2 mm longis; corolla atrocaerulea, epunctata, tubulosa, 23-25 mm longa, lobis ovatis, 1.5-2 mm longis, obtusis, plicae lobis breviores, trun- catae, integrae vel denticulatae; stamina tubo corollae inferi- oris inserta, filamentis subulatis, c. 9 mm longis, antheris luteis, linearibus, c. 1.5 mm longis; ovarium stipitatum, lin- eare, c. 20 mm longum, stylo breve, stigmatibus linearibus. China: SE Tibet: Bai ma, on alpine meadows, 4800 m, 10 August 1973, Northwest Inst. Biol. Tibet Exped. 1367 (HNWP - holotype); Bamda-Zogong, 4000 m, 5 September 1977, P. C. Kuo & W. Y. Wang 23411 (HNWP). ACKNOWLEDGEMENTS. We are particularly indebted to the directors and staff of the herbaria of The Natural History Museum, the Royal Botanical Gardens, Kew, and the Royal Botanical Garden at Edin- burgh for permission to consult their collections, for facilities pro- vided and for the loan of specimens. We must also thank Prof. Tang Yan-cheng, Mr A O Chater and Dr Mike Gilbert for useful discus- sions in the preparation of this publication. REFERENCES Ho, T.-N. & Liu, S.-W. & Wu, C.J. 1988, Gentianaceae. Flora Reipublicae Popularis Sinicae 62. Beijing. Kusnezow, N. I. 1894. Subgenus Eugentiana of genus Gentiana Tourn. Trudy imp. S.-Peterb. Obshch. Estest. 24(2): 1-531. [Latin translation in Trudy imp. S. -Peterb. hot. Sada 15: 1-507 (1896-1904)]. Naithani, H. B. 1990. Gentiana. In Flowering plants of India, Nepal and Bhutan: 291-295. Bull. not. Hist. Mus. Land. (Bot.) 23(2): 61-65 Issued 25 November 1993 New combinations, names and taxonomic notes on Gentianella (Gentianaceae) from South America and New Zealand TING-NUNG HO AND SHANG-WU LIU Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China SYNOPSIS. Gentianella is a large genus of about 200 species. It was segregated from the genus Gentiana in 1794. Since then the study histories of the two genera have been interwoven. During study on Gentiana and Gentianella it was realized that 104 new combinations and two names needed to be made from South America and New Zealand. INTRODUCTION The genus Gentianella was segregated from the genus Genti- ana L. by Moench in 1794. Since then the study histories of the two genera have been closely interwoven. The majority of species of Gentianella were originally described within Genti- ana. Since the last half century, Fabris (1953-1981), Gillet (1957), Holub (1967-1983) and Pringle (1981-1987) etc. have studied Gentianella, and made a number of nomenclatural transfers, but these studies are insufficient for the whole genus and there are still too many species under Gentiana. During study on Gentiana and Gentianella in Eurasian her- baria, it was realized that 104 new combinations and two names of Gentianella needed to be made from South America and New Zealand. Gentianella Moench is a large genus of about 200 species. It has an almost global distribution chiefly centred on South America and New Zealand. NEW COMBINATIONS AND TAXONOMIC NOTES Gentianella achalensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana achalensis Hieron., Bot. Acad. Nac. Cordova, 4: 373 (1881), nom. nud; Gilg, Bot. Jahrb., 22: 322 (1896). Type: Argentina, Hieronymus 526 (K!). Gentianella albido-caerulea (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana albido-caerulea Gilg, Bot. Jahrb. ,22: 323 (1896). Syntypes: Bolivia, M. Bang 1132 (n.v.), O. Kuntze (n.v.). Gentianella andreae-mathewsii (Briquet) T. N. Ho & S. W. Liu, comb. nov. Gentiana andreae-mathewsii Briquet, Candollea, 4: 326 (1931), based on Gentiana mathewsii Gilg, Bot. Jahrb., 54 (Beibl. 118): 64 (1917), non Petrie (1911) (Trans. Proc. N. Zeal. Inst., 44: 183). Type: Peru, Mathews (n.v.). Gentianella antarctica (Kirk) T. N. Ho & S. W. Liu, comb. nov. Gentiana concinna var. robusta Hook, f., Fl. Antarct., 1: 53 (1844). Gentiana antarctica Kirk, Trans. Proc. N. Zeal. Inst. 27: 339, 1894 (1895). Type: New Zealand, T. Kirk, W. 4729 (n.v.). Gentianella antipoda (Kirk) T. N. Ho & S. W. Liu, comb. nov. Gentiana antipoda Kirk, Trans. Proc. N. Zeal. Inst., 23: 440. 1890 (1891), nom. nud. et 27: 340. 1894 (1895). Type: New Zealand, T. Kirk (K! - isotype). Gentianella armerioides (Griseb.) T. N. Ho & S. W. Liu, comb. nov. Gentiana armerioides Griseb. in Lechler, Berb. Am. Austr., 58 (1857). Type: Peru, Lechler 2000a (P! - isotype). Gentianella astonii (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana astonii Petrie, Trans. Proc. N. Zeal. Inst., 48: 187. 1915 (1916). Type: New Zealand, Valley of Ure river, B. C. Aston (WELT! - holotype). Gentianella atroviolacea (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb. nov. Gentiana atroviolacea Gilg, Bot. Jahrb., 54(Beibl. 118): 53 (1917). Type: Columbia, Kalbreyer 1200 (K! - isotype). Gentiana solidagoides Reim., Bot. Jahrb., 62: 329 (1929). Type: Columbia, (K! no. 10574), syn. nov. Gentianella bangii (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb. nov. Gentiana bangii Gilg, Bot. Jahrb., 22: 324 (May 1896), nee Rusby (end 1896) (Mem. Torr. Bot. Club , 6: 79), Type: Bolivia, M. Bang 1 153 (K!- isotype). Gentianella bellatula (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana bellatula Gilg, Bot. Jahrb., 50(Beibl. Ill): 49 (1913). Syntypes: Bolivia, Hauthal 201, 218 (all n.v.). Gentianella bellidifolia (Hook, f.) Holub var. divisa (Kirk) T. N. Ho & S. W. Liu, comb. nov. Gentiana bellidifolia var. divisa Kirk, Trans. Proc. N. Zeal. Inst., 27: 337. 1894 (1895). Type: New Zealand, Ashburton Mountains, T. H. Potts, W. 4714 (n.v.). Gentiana divisa (Kirk) Cheeseman., Man. N. Zeal FL, 453 (1906). Gentianella bockii (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana bockii Gilg, Bot. Jahrb., 54(Beibl. 118): 34 (1917). Syn- types: Bolivia, Bock in herb. Herzog 2480e (n.v.), Buchtien 1481 (n.v.). Gentianella bridgesii (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, nov. Gentiana bridgesii Gilg, Bot. Jahrb., 22: 316 (1896). Syntypes: Bolivia, Bridges a. 1850 (n.v.), O. Kuntze (n.v.). Gentianella briquetiana (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana briquetiana Gilg, Bot. Jahrb., 54 (Beibl. 118): 32 (1917). Type: Bolivia, Larecaja, Mandon 361 (P! - holotype, BM! K! - isotypes). Gentianella bromifolia (Griseb.) T. N. Ho & S. W. Liu, comb. nov. Gentiana bromifolia Griseb., Glen. Abh., 19: 208 (1874). Syntypes: Argentia, Tucuman, Lorrentz 775 (K!), 303 (n.v.). Gentianella brunneo-tincta (Gilg) T. N. Ho & S. W. Liu, comb. nov. The Natural History Museum, 1993 62 T.-N. HO ANDS.-W. LIU Gentiana brunneo-tincta Gilg, Fedde, Rep. Nov. Sp., 2: 37 (1906). Type: Peru, Weberbauei 3092 (n.v.). Gentianella buchtienii (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana buchtienii Gilg, Bot. Jahrb., 54(Beibl. 118): 66 (1917). Type: Bolivia, Buchtien 48 (K! - lectotype designated here, BM! - isolectotype). Gentianella calcarea (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana calcarea Gilg, Fedde, Rep. Nov. Sp., 2: 42 (1906). Type: Peru, Weberbauer 2539 (n.v.). Gentianella centamalensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana centamalensis Gilg, Bot. Jahrb. ,22: 334 (1896). Type: Peru, Stuebel4\ (n.v.). Gentianella cerina (Hook, f.) T. N. Ho & S. W. Liu, comb. nov. Gentiana cerina Hook, f., Fl. Antarct., 1: 54, t. 36 (1844). Type: New Zealand, Ins. Auckland, J. D. Hooker (K! - holotype). Gentianella chathamica (Cheesman) T. N. Ho & S. W. Liu, comb, nov. Gentiana chathamica Cheeseman, Man. N. Zeal. Fl., 449 (1906). Type: New Zealand, F. A. D. Cox (BM! K! - isotypes). Gentianella chrysantha (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana chrysantha Gilg, Bot. Jahrb., 54(Beibl. 118): 81 (1917). Type: Bolivia, Herzog 2044 (n.v.). Gentianella chrysosphaera (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana chrysosphaera Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917). Type: Peru, Weberbauer 6521 (n.v.). Gentianella chrysotaenia (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana chrysotaenia Gilg, Bot. Jahrb., 54(Beibl. 118): 39 (1917). Syntypes: Peru, Weberbauer 6528, 6914 (all n.v.). Gentianella claytonioides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana claytonioides Gilg, Bot. Jahrb., 22: 318 (1896). Type: Argentina, Rioja, Hieronymous & Niederlein (n.v.). Gentianella coccinea (G. Don) T. N. Ho & S. W. Liu, comb. nov. Gentiana coccinea G. Don, Gen. Syst., 4: 196 (1838). Type: Peru, Pavon (BM! - isotype). Gentianella concinna (Hook, f.) T. N. Ho & S. W. Liu, comb. nov. Gentiana concinna Hook, f., Fl. Antarct., 53, t. 35 (1844). Type: New Zealand, Ins. Auckland, J. D. Hooker (K! - holotype). Gentianella coquimbensis (Briquet) T. N. Ho & S. W. Liu, comb, nov. Gentiana coquimbensis Briquet, Candollea, 4: 328 (1931). Type: Chile, Coquimbo, M. Gay (P! - isotype) Gentianella corallina (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana corallina Gilg, Fedde, Rep. Nov. Sp., 2: 48 (1906). Type: Peru, Weberbauer 4288 (n.v.). Gentianella crassiuscula T. N. Ho & S. W. Liu, nom. nov. Gentiana crassicaulis Gilg, Bot. Jahrb., 54(Beibl. 118): 60 (1917), non Duthie ex Burkill (1906) (/. As. Soc. Beng. n. s. 2: 316). Type: Peru, Lobb (n.v.). Gentianella crassulifolia Fabris, Bol. Soc. Argent. Bot., 8: 25 (1959). Gentiana crassulifolia Griseb., Gen. Sp. Gent., 227 (1838). Type: Columbia, Jameson 455 (K!). G. crassulifolia is a subshrub, characterised by crowded, imbricate, rigid and subcoriaceous leaves, very short pedicels and flowers crowded in a head at the top of the stem. Its calyx-lobes are as long as to slightly longer than the tube. Leaf shape shows some variation. They are oblong, elliptic, lanceolate to ovate. Two varieties can be recognised. var. crassulifolia Gentiana selaginifolia Griseb., Linnaea, 22: 42 (1849). Type: Colum- bia, Hartweg 1252 (P! - isotype). Gentiana engleri Gilg, Bot. Jahrb., 22: 314 (1896). Type: S. Colum- bia, Lehman 666 (BM!, K!), syn. nov. Gentiana dacrydioides Gilg, Bot. Jahrb., 22: 312 (1896). Type: S. Columbia, Lehman 2682 (BM! - isotype), syn. nov. Gentianella selaginifolia (Griseb.) Fabris, Bol. Soc. Argent. Bot., 8: 25 (1959). Gentianella dacrydioides (Gilg) Weaver & Rudenberg, J. Arnold. Arbor., 56(2): 215 (1975). Calyx-lobes lanceolate; corolla c. 15 mm. var. hypericoides (Gilg.) T. N. Ho, stat. et comb. nov. Gentiana hypericoides Gilg, Bot. Jahrb., 22: 312 (1896). Type: Ecuador, Lehman 6521( K!). Gentianella hypericoides (Gilg) Fabris, Bol. Soc. Argent. Bot., 8: 184 (1960). Calyx-lobes oblong-lanceolate; corolla c. 20-25 mm. Gentianella crossolaema (Wedd.) T. N. Ho & S. W. Liu, comb. nov. Gentiana crossolaema Wedd., Chlor. And., 2: 55 (1859). Type: Peru, Weddel(P\- holotype). Gentianella cupiformis T. N. Ho & S. W. Liu, nom. nov. Gentiana campanuloides Gilg, Bot. Jahrb., 22: 320 (1916), non Willd. ex Roem. & Schult. (1820) (Roem. & Schult., Syst., 6: 184). Type: Argentina, Lorentz & Hieronymus (n.v.). Gentianella dasythamna (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana dasythamna Gilg, Bot. Jahrb., 54 (Beibl. 118): 63 (1917). Type: Bolivia, Larecaja, Mandon 363 ( P! - holotype, BM! K! - isotypes). Gentianella dissitifolia (Griseb.) T. N. Ho & S. W. Liu, comb. nov. Gentiana dissitifolia Griseb., Gen. Sp. Gent., 229 (1839). Type: Peru, Dombey 394 (P! - lectotype designated here). Gentianella dombeyana (Wedd.) T. N. Ho & S. W. Liu, comb. nov. Gentiana dombeyana Wedd., Chlor. And., 2: 62 (1845). Type: Peru, Dombey (P! - holotype). Gentianella ericothamna (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana ericothamna Gilg, Fedde, Rep. Nov. Sp., 2: 50 (1906). Type: Peru, Weberbauer 3381 (n.v.). Gentianella eurysepala (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, nov. Gentiana eurysepala Gilg, Bot. Jahrb., 50 (Beibl. Ill): 50 (1913). Type: Peru, Weberbauer 5676 (n.v.). Gentianella fiebrigii (Gilg) Holub, Folia Geobot. & Phytotax., Praha, 2: 117(1967). Gentiana fiebrigii Gilg, Fedde, Rep. Nov. Sp. 2: 45 (1906). Type: Bolivia, K. Fiebrig 3187 (BM! K! - isotypes). Gentiana gynophora Gilg, Bot. Jahrb. ,22: 305 (1896). Type: Bolivia, M. Bang 1231 (BM! K! - isotypes), syn. nov. There is no clear dividing line between G. gynophora with long- petiolate basal leaves and scapose stem and G. fiebrigii with less long-petiolate basal leaves and scapose or foliate stem. G. gynophora is therefore not maintained as a distinct species. Gentianella filipes (Cheeseman) T. N. Ho & S. W. Liu, comb. nov. Gentiana filipes Cheeseman, Trans. Proc. N. Zeal. Inst., 28: 536 (1896). Type: New Zealand, Mount Arthur, T. F. Cheeseman (BM! K! - isotypes). Gentianella florida (Griseb.) Holub, Folia Geobot. Phytotax., Praha, 2: 117 ( 1967). Type: Argentina, Tucuman, Lorentz 310 (K!). Gentiana totorensis Gilg, Bot. Jahrb., 54(Beibl. 118): 84 (1917). Type: Bolivia, Herzog 2032 (n.v.), syn. nov. Gentianella fruticulosa (Domb. ex Wedd.) T. N. Ho & S. W. Liu, comb. nov. Gentiana fruticulosa Domb. ex Wedd., Chlor. And., 2: 71 (1839). Type: Peru, Dombey (P! - holotype). Gentianella gibbsii (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana gibbsii Petrie, Trans. Proc. N. Zeal. Inst., 49: 52. 1916 (1917). Type: New Zealand, Stavart Island, F. G. Gibbs in herb. D. Petrie, W. 4709 (WELT! - holotype). Gentianella gracilifolia (Cheeseman) T. N. Ho & S. W. Liu, comb, nov. Gentiana gracilifolia Cheeseman, Man. N. Zeal. Fl. 1144 (1906). Type: New Zealand, Mount Arthur Plateau, F. G. Gibbs in herb. T. F. Cheeseman 137 (BM! K! - isotypes). NEW COMBINATIONS, NAMES AND NOTES ON GENTIANELLA 63 Gentianella graebneriana (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana graebneriana Gilg, Bot. Jahrb., 54(Beibl. 118): 24 (1917). Type: Peru, Weberbauer 605 1 (n.v.). Gentianella grisebachii (Hook, f.) T. N. Ho, comb. nov. Gentiana grisebachii Hook, f., Ic. PL, t. 636 (1844). Type: New Zealand, Rotuari-Tongariro, J. C. Bidwill (K! - holotype). Gentianella ignea (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana ignea Gilg, Fedde, Rep. Nov. Sp., 2: 49 (1906). Type: Peru, Weberbauer 746 (n.v.). Gentianella krauseana (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, nov. Gentiana krauseana Gilg, Fedde, Rep. Nov. Sp., 2: 45 (1906). Type: Bolivia, K. Fiebrig 3187c (n.v.). Gentianella kuntzei (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana kuntzei Gilg, Bot. Jahrb., 22: 326 (May 1896). Type: Bolivia, O. Kuntze (n.v.). Gentiana cochabambensis Rusby, Mem. Torr. Bot. Club, 6: 76 (End 1896). Type: Bolivia, M. Bang 1232 (BM! K! - isotypes). Gentianella kusnezowii (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana kusnezowii Gilg, Bot. Jahrb., 22: 325 (1896). Type: Bolivia, M. Bang 1230 (BM! K! - isotypes). Gentianella larecajensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana larecajensis Gilg, Bot. Jahrb., 54(Beibl. 118): 31 (1917). Types: Bolivia, Larecaja, Mandon 362 (P! - lectotype designated here, BM! K! - isolectotypes), 363 P. P. (BM! K!). Gentianella lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana lilacina Gilg, Fedde, Rep. Nov. Sp., 2: 40 (1906). Type: Peru, Weberbauer 3223 (n.v.). Gentianella lilacino-flavescens (Gilg) T. N. Ho & S. W. Liu, comb, nov. Gentiana lilacino-flavescens Gilg, Bot. Jahrb., 54(Beibl. 118): 38 (1917). Type: Bolivia, Herzog 2114 (n.v.). Gentianella limoselloides ( Kunth ) Fabris, Bol. Soc. Argent. Bot., 8: 166(1960). Gentianaana limoselloides Kunth, Nov. Gen., 3: 130 (1818). Types: Ecuador, Humboldt & Bonpland (P! - lectotype designated here), Bonpland 2266 (PI). Gentiana paludicola Gilg, Fedde, Rep. Nov. Sp., 2: 42 (1906). Type: Peru, Weberbauer 2694 (n.v.), syn. nov. G. limoselloides is a widespread and variable species. It shows considerable variation especially in petiole length of the basal leaves and in cauline leaf shapes which are spathalate, oblong to lanceolate. Clearly, G. paludicola with lanceolate cauline leaves and long- petiolate basal leaves can not be maintained at any rank. Gentianella lineata (Kirk) T. N. Ho & S. W. Liu, comb. nov. Gentiana lineata Kirk, Trans. Proc. N. Zeal. Inst. , 27: 334, t. 27. 1894 (1895). Type: New Zealand, Hollows on the crest of the Longwool Range, T. Kirk (K! - isotype). Gentianella liniflora (Kunth) T. N. Ho & S. W. Liu, comb. nov. Gentiana liniflora Kunth, Nov. Gen. Sp., 3: 171 (1818). Type: Peru, Humboldt & Bonpland ( P! - holotype). Gentianella lithophila (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana lithophila Gilg, Bot. Jahrb., 54(Beibl. 118): 39 (1917). Type: Bolivia, Herzog 2115 (n.v.) Gentianella lobbii (Gilg) T. N. Ho & S. W. Li u, comb. nov. Gentiana lobbii Gilg, Bot. Jahrb., 54 (Beibl. 118): 60 (1917). Type: Peru, Lobb (n.v.). Gentianella lurido-violacea (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb. nov. Gentiana lurido-violacea Gilg, Fedde, Rep. Nov. Sp., 2: 37 (1906). Type: Peru, Weberbauer 3759 (n.v.). Gentianella lythroides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana lythroides Gilg, Bot. Jahrb., 54(Beibl. 118): 24 (1917). Type: Bolivia, Herzog 2229 (n.v.). Gentianella macrorriza (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, nov. Gentiana macrorriza Gilg, Bot. Jahrb., 54(Beibl. 118): 40 (1917). Type: Bolivia, Herzog 2271 (n.v.). Gentianella magellanica (Gaudich.) Fabris in D. M. Moore, Vase., Fl. Falkland. Isl. 103 (1968). Gentiana magellanica Gaudich. in Freyc. Voy. Bot. 449 (1826). Type: lies Malouines, Gaudichaud (K! P! - isotypes). Gentiana pearcei Philippi, Anal. Univ. Santiago, 18: 65 (1861). Type: Chile, Philippi (K! - isotype), syn. nov. Gentiana valdiviana Philippi, Anal. Univ. Chil., 40: 206 (1895). Type: Chile, Philippi (K! - isotype), syn. nov. G. magellanica is a variable species in leaf, calyx-lobe shapes and corolla size. Leaves are oblong, ovate or lanceolate. Calyx-lobes are usually oblong, rarely ovate or lanceolate and all longer than the tube. The range of corolla size is (10-)13-18(-20) mm. The differ- ences in calyx-lobe shape and corolla size used to separate G. valdiviana and G. pearcei from G. magellanica are not of taxonomic significance. Gentianella mathewsii (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana mathewsii Petrie, Trans. Proc. N. Zeal. Inst., 44: 183. 1911 (1912). Type: New Zealand, near Lake Harris, B. Petrie W. 4710 (WELT! -holotype). Gentianella mesembrianthemoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana mesembrianthemoides Gilg, Fedde, Rep. Nov. Sp., 2: 41 (1906). Type: Peru, Weberbauer 3303 (n.v.). Gentianella muscoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana muscoides Gilg, Fedde, Rep. Nov. Sp., 2: 35 (1906). Type: Peru, Weberbauer no. Ph. 96 (n.v.). Gentianella narcissoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana narcissoides Gilg, Bot. Jahrb., 54(Beibl. 118): 65 (1917). Type: Bolivia, Herzog 2414 (n.v.). Gentianella neomandonii (R. C. Foster) T. N. Ho & S. W. Liu, comb, nov. Gentiana neomandonii R. C. Foster, Rhodora, 56: 103 (1954), based on Gentiana mandonii Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917), non Rusby (1896) (Mem. Torr. Bot. Club, 6: 80). Type: Bolivia, Mandon 363a (P! - holotype, BM! K! - isotypes). Gentianella odontosepala (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana odontosepala Gilg, Fedde, Rep. Nov. Sp., 2: 48 (1906). Type: Bolivia, M. Bang 2671 (n.v.). Gentianella oreosilena (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana oreosilena Gilg, Fedde, Rep. Nov. Sp., 2: 40 (1906). Type: Peru, Weberbauer 4288 (n.v.). Gentianella orobanchoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana orobanchoides Gilg, Bot. Jahrb., 22: 333 (1896). Type: Bolivia, Mandon 366 (P! - lectotype designated here, BM! K! - isolectotypes). Gentianella pachystemon (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana pachystemon Gilg, Bot. Jahrb., 54(Beibl. 118): 46 (1917). Syntypes: Peru, Stuebel 51, 52 (all n.v.). Gentianella palcana (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana palcana Gilg, Bot. Jahrb., 54(Beibl. 118): 47 (1917). Syntypes: Bolivia, Herzog 2176 (n.v.), Stuebel 46c (n.v.). Gentianella pallido-lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana pallido-lilacina Gilg, Bot. Jahrb., 54( Beibl. 118): 58 (1917). Type: Bolivia, Herzog 2028 (n.v.). Gentianella parviflora (Griseb. ) T. N. Ho, stat. et comb. nov. Gentiana coerulescens Gill, ex Wedd. var. parviflora Griseb., Symb. Fl. Argent., 237 (1879). Type: Argentina, Hieronymus 464 (K!). Gentianella peruviana (Griseb.) Fabris, Bol. Soc. Argent. Bot., 7: 93 (1958). Gentiana limoselloides Kunth var. peruviana Griseb., Gen. Sp. Gent., 215 (1838). Type: Peru, Weddell 4445 (P! - Type). Gentiana peruviana (Griseb.) Gilg, Bot. Jahrb. 22: 304 (1896), non 64 Lamarck (1786) (Lamarck, Encycl. Meth. 2: 642). G. peruviana is a perennial dwarf herb characterized by a rather fleshy tap-root, developed, oblong to oblanceolate basal leaves and oblong, ovate-oblong to obovate calyx-lobes which are as long as, slightly shorter or longer than the tube. It includes four species (G. peruviana, G. hieronymii, G. boliviano and G. lobelioides) which are better treated as the following two varieties. The difference between var. peruviana and var. boliviano is only of corolla size: larger flowers (16-17 mm) of the former and smaller flowers (11-13 mm) of the latter. var. peruviana Gentiana hieronymii Gilg, Bot. Jahrb., 22: 305 (1896). Type: Argen- tina, Lorentz & Hieronymus 15 (K! - isotype), syn. nov. Gentianella hieronymii (Gilg) Fabris, Rev. Invest. Agric., Buenos Aires, 11: 396 (1958), in adnot. var. boliviana (Pax) T. N. Ho, stat. et comb. nov. Gentiana boliviana Pax, Fedde, Rep. Nov. Sp., 7: 243 (1909). Type: Bolivia, Buchtien 1482 (n.v.). Gentiana lobelioides Gilg, Bot. Jahrb., 54(Beibl. 118): 30 (1917). Type:Peru, Weberbauer 955 (n.v.), syn. nov. Gentianella petrophila (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana petrophila Gilg, Fedde, Rep. Nov. Sp.,2: 42 (1906). Type: Peru, Weberbauer 2562 (n.v.). Gentianella pilgeriana (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana pilgeriana Gilg, Bot. Jahrb., 54(Beibl.ll8): 42 (1917). Type: Bolivia, Herzog 2410 (n.v.) Gentianella poculifera (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana poculifera Gilg, Bot. Jahrb., 50(Beibl. 118): 48 (1913). Type: Peru, Weberbauer (n.v.) Gentianella porphyrantha (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb. nov. Gentiana porphyrantha Gilg, Fedde, Rep. Nov. Sp., 2: 39 (1906). Type: Peru, Weberbauer 2803 (n.v.). Gentianella potamophila (Gilg) Fabris ex T. N. Ho , comb. nov. Gentiana potamophila Gilg, Bot. Jahrb., 54(Beibl. 118): 74 (1917). Type: Peru, Weberbauer 6907 (n.v.). Gentianella primuloides (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana vaginalis Griseb. ex Wedd. Chlor. And., 2: 53 (1859), non Griseb. (1838) (Gen. Sp. Gent., 215). Gentiana primuloides Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917). Types: Peru, Lechler 2002 (P! - lectotype designated here); Bolivia, M. Bang 1888 (BM! K!), 1889 (BM! K!), O. Kuntze (n.v.), Bock in herb. Herzog 2480c (n.v.), Herzog 2081 (n.v.). Gentianella pseudolycopodium (Gilg) T. N. Ho & S. W. Liu, comb, nov. Gentiana pseudolycopodium Gilg, Fedde, Rep. Nov. Sp., 2: 38 (1906). Type: Peru, Huamalies, Weberbauer 3353 (n.v.). Gentianella pulla (Griseb.) T. N. Ho & S. W. Liu, comb. nov. Gentiana pulla Griseb., Goett. Abh., 19: 209 (1874). Type: Argen- tina, Lorentz 773 (K! - isotype). Gentianella punicea (Wedd.) Holub, Folia. Geobot. Phytotax., Praha, 2: 118(1967). Gentiana punicea Wedd., Chlor. And., 2: 70 (1859). Types: Bolivia, Mandon 364 (K! BM!); Peru, Weddell 4741 (P! - lectotype designated here). Gentiana dolichantha Gilg, Torrey, 5: 109 (1909). Syntype: Bolivia, Lobb (n.v.), R. S. Wiliams 2489 (K! BM!), syn. nov. Gentiana purpureiflora Gilg, Bot. Jahrb. 54 (Beibl. 118): 65 (1917). Type: Bolivia, Herzog 2168 (n.v.), syn. nov. The authors could find no distinction between G. dolichantha and G. punicea in any respect. G. purpureiflora is also identical to G. punicea in habit, leaf shape, corolla size as well as the character of the calyx-lobes. It seems that G. purpureiflora is supposed to differ from G. punicea in its corolla-lobes which are as long as the tube. However, there is a complete range of intermediates from short to long corolla-lobes in G. punicea. Therefore, G. dolichantha and G. purpureiflora are all reduced to synonyms. T.-N. HO AND S.-W. LIU Gentianella raimondiana (Wedd.) T. N. Ho & S. W. Liu, comb. nov. Gentiana raimondiana Wedd., Chlor. And., 2: 310 (1861). Type: Peru, Raimondi (n.v.). Gentianella roseo-lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana roseo-lilacina Gilg, Fedde, Rep. Nov. Sp., 2: 35 (1906). Type: Peru, Weberbauer 2952 (n.v.). Gentianella sancti-mathacii (R. C. Foster) T. N. Ho & S. W. Liu, comb. nov. Gentiana sancti-mathacii R. C. Foster, Rhodora, 56: 103 (1954), based on Gentiana praticola Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917), non Franchet (1896). (Bull. Soc. Bot. Fr., 43: 489). Type: Bolivia, Herzog 1977 (n.v.). Gentianella sandiensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana sandiensis Gilg, Fedde, Rep. Nov. Sp., 2: 36 (1906). Syntypes: Bolivia, Weberbauer 1016 (n.v.); Peru, Weberbauer 352, 445, 917a, 2593 (all n.v.), Poeppig (n.v.). Gentianella saxicola (Griseb.) T. N. Ho & S. W. Liu, comb. nov. Gentiana saxicola Griseb., Gen. Sp. Gent., 216 (1838). Type: Peru, Andes, Mathews (K! - isotype). Gentiana vaginalia Griseb., Gen. Sp. Gent., 215 (1838). Type: Peru, Pasco, Cruikshanks (K!), syn. nov. The type of G. vaginalia is identical to the type of G. saxicola in all respects, e. g. dwarf herb, crowded leaves forming a rosette, leaves ovate or triangular, calyx-lobes all papillose on upper surfaces and distinctly ciliate on margins. G. vaginalia is therefore not maintained here. Gentianella scarlatiflora (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana scarlatiflora Gilg, Bot. Jahrb., 50 (Beibl. Ill): 49 (1913). Syntypes: Peru, Lobb (n.v.), Weberbauer 5836 (n.v.). Gentianella scarlatina (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana scarlatina Gilg, Fedde, Rep. Nov. Sp., 2: 36 (1906). Type: Peru, Sandia, Weberbauer 1047 (n.v.). Gentianella scarlatino-striata (Gilg) T. N. Ho & S. W. Liu, comb, nov. Gentiana scarlatino-striata Gilg, Bot. Jahrb., 54(Beibl. 118): 67 (1917). Type: Peru, Weberbauer 6621 (n.v.). Gentianella scopulorum (Wedd.) Fabris ex T. N. Ho & S. W. Liu, comb. nov. Gentiana scopulorum Wedd., Chlor. And., 32: 67 (1859), nee Tide- str. (1925) (Contr. U.S. Nat. Herb., 26: 416). Type: Bolivia, Weddell 3928 (P! - holotype). Gentianella silenoides (Gilg) Fabris in A. L. Cabrera, Fl. Prov. Jujuy (Inst. Nac. Teen. Agropec., 13), 8: 68 (1983). Gentiana silenoides Gilg, Bot. Jahrb., 22: 319 (1896). Type: Bolivia, Lorentz & Hieronymous 878 P. P. (n.v.). var. silenoides Flowers in a lax cyme; corolla 16-18 mm. var. striaticalyx (Gilg) Ho, stat. et comb. nov. Gentiana striaticalyx Gilg, Bot. Jahrb., 54(Beibl. 118): 56 (1917). Type: Bolivia, //erzog2046b. (n.v.). Gentiana anthosphaera Gilg, Fedde, Rep. Nov. Sp., 2: 46 (1906). Type: Bolivia, Fiebrig 2246 (K! P! - isotypes), syn. nov. Gentiana herzogii Gilg, Bot. Jahrb. ,54 (Beibl. 118): 57 (1917). Type: Bolivia, Herzog 2018 (n.v.), syn. nov. Gentianella anthosphaera (Gilg) Holub, Folia Geobot. & Phytotax., Praha, 2: 116 (1967); Fabris in A. L. Cabrera, Fl. Prov. Jujuy (Inst. Nac. Teen. Agropec., 13), 8: 77 (1983), comb, superfl. Flowers in a lax cyme; corolla 20-30 mm. var. inaequicalyx (Gilg) Ho, stat. et comb. nov. Gentiana inaequicalyx Gilg, Bot. Jahrb., 22: 324 (1896). Syntypes: Bolivia, Mandon 365 (K!), M. Bang 1143 (K!), O. Kuntze (n.v.). Flowers in a dense umbel or subcapitulum; corolla 20-30(-35) mm. Gentianella serotina (Cockayne) T. N. Ho & S. W. Liu, comb. nov. Gentiana serotina Cockayne, Trans. Proc. N. Zeal. Inst., 47: 113. 1914 (1915). Type: New Zealand, W. 4724 (n.v.). Gentianella setipes (Gilg) T. N. Ho & S. W. Liu, comb. nov. NEW COMBINATIONS, NAMES AND NOTES ON GENTIANELLA 65 Gentiana setipes Gilg, Bot. Jahrb., 54(Beibl. 118): 43 (1917). Type: Peru, Weberbauer 6322 (n.v.). Gentianella soratensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana soratensis Gilg, Bot. Jahrb., 22: 332 (1896). Type: Bolivia, Rusby 675 (P! - holotype, BM! K! - isotypes). Gentianella speciosissima (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana speciosissima Gilg, Bot. Jahrb., 22: 325 (1896). Type: Peru, Stuebel24b,25(al\n.v.). Gentianella spedenii (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana spedenii Petrie, Trans. Proc. N. Zeal. Inst., 56: 14 (1926). Type: New Zealand, Princess Range, James Speden (WELT! - holotype). Gentianella spenceri (Kirk) T. N. Ho & S. W. Liu, comb. nov. Gentiana spenceri T. Kirk, Trans. Proc. N. Zeal. Inst., 27: 335, t. 27 A, B. 1894 (1895). Type: New Zealand, Mount Rochfort, F. H. Spencer, W. 4711 (n.v.). Gentianella stellarioides (Gilg) Fabris, Bol. Soc. Argent. Bot. , 8: 180 (1960). Gentiana stellarioides Gilg, Bot. Jahrb., 54(Beibl. 118): 79 (1917). Types: Ecuador, Spruce (K! - lectotype designated here), Jameson (K!). var. stellarioides var. androtricha ( Gilg) T. N. Ho, stat. et comb. nov. Gentiana androtricha Gilg, Bot. Jahrb., 54(Beibl. 118): 79 (1917). Type: Ecuador, Spruce 6050 (K!). This variety is very similar to var. stellarioides in having linear to linear-lanceolate calyx-lobes, very long pedicels and large flowers (17-20 x 15-20 mm), but is distinguished by basal leaves with long petioles and elliptic cauline leaves. Gentianella stenosepala (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana stenosepala Gilg, Bot. Jahrb., 22: 331 (1896). Type: Bolivia, O. Kuntze (n.v.). Gentianeila stricticaulis (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana stricticaulis Gilg, Bot. Jahrb., 54(Beibl. 118): 62 (1917). Type: Peru, Weberbauer 6097 (n.v.). Gentianella stuebelii (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana stuebelii Gilg, Bot. Jahrb., 22: 317 (1916). Type: Peru, Stuebel35f (n.v.). Gentianella tarapacana (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana tarapacana Gilg, Bot. Jahrb., 22: 305 (1896). Type: Chile, Philippi (K! - isotype). Gentianella tenuifolia (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana tenuifolia Petrie, Trans. Proc. N. Zeal. Inst., 45: 270. 1912 (1913). Type: New Zealand, near Lyell, 5. W. Nelson in herb. B. Petrie, W. Townson, W. 4721 (WELT! - holotype). Gentianella tereticaulis (Petrie) T. N. Ho & S. W. Liu, comb. nov. Gentiana tereticaulis Petrie, Trans. Proc. N. Zeal. Inst., 49: 51. 1916 (1917). Type: New Zealand, Lake Harries, Routeburn Valley, W. Petrie (WELT! - holotype). Gentianella thiosphaera (Gilg) Holub, Folia Geobot. Phytotax., Praha, 2: 118 (1967). Gentiana thiosphaera Gilg, Fedde, Rep. Nov. Sp.,2: 46 (1906). Type: S. Bolivia, K. Fiebrig 3156 (P! - holotype, BM! K! - isotypes). var. thiosphaera Gentiana comarapana Gilg, Bot. Jahrb., 54(Beibl. 118): 82 (1917). Type: Bolivia, Herzog 1914 (n.v.), syn. nov. G. comarapana was originally described as having lanceolate calyx- lobes which are as long as the tube, and thus differs from G. thiosphaera. The calyx-lobes of both species are lanceolate, linear- lanceolate to linear and as long as to much longer than the tube. G. comarapana is therefore not maintained here. var. macroclada (Gilg) Ho, stat. et comb. nov. Gentiana macroclada Gilg, Fedde, Rep. Nov. Sp., 2: 47 (1906). Type: Bolivia, K. Fiebrig 2654a. (n. v.). This variety is very similar to var. thiosphaera and is distinguished only by smaller flowers (usually 12-14 mm). Gentianella townsonii (Cheeseman) T. N. Ho & S. W. Liu, comb, nov. Gentiana townsonii Cheeseman, Man. N. Zeal. Fl., 450 (1906). Syntypes: New Zealand, Mount Rochfort, W. Townson 288 (BM! K!), 389 (BM! K!). Gentianella tristicha (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, nov. Gentiana tristicha Gilg, Fedde, Rep. Nov. Sp., 2: 39 (1906). Type: Peru, Weberbauer 2933 (n.v.). Gentianella vernicosa (Cheeseman) T. N. Ho & S. W. Liu, comb, nov. Gentiana vernicosa Cheeseman, Man. N. Zeal. Fl., 1145 (1906). Type: New Zealand, Mount Lockett, F. G. Gibbs (n.v.). ACKNOWLEDGEMENTS. We are grateful to the directors and staff of the herbaria of The Natural History Museum, Royal Botanic Gar- dens, Kew, Museum National d'Histore Naturelle, Paris and the National Museum of New Zealand, for permission to consult their collections and for the loan of specimens. We are much indebted to Mr Roy Vickery, Miss Sally Bidgood and Mr Thierry Deroin for their help in providing most of the herbarium material. Thanks are also due to Dr Mike Gilbert for useful suggestions in the preparation of this paper. REFERENCES Allan, H. H., 1961. Gentiana. In Flora of New Zealand 1: 764-779. Wellington. Fabris, H. A., 1953. Sinopsis preliminar de las Gentianaceae Argentinas, Bol. Soc. Argent. Bot., 4: 232-259. , 1955. Nuevas especies de 'Gentianella' del Peru, Bol. Soc. Argent. Bot., 6(1): 45-50. , 1958. Notas Sobre Gentianella del Peru. Bol. Soc. Argent. Bot., 7(2): 86-93. , 1959. Sobre la identid de dos species Sudamericanae de Gentianella, Bol. Soc. Argent. Bot., 8(1): 24-25. , 1960. El Genero Gentianella en Ecuador, Bol. Soc. Argent. Bot., 8(3-4): 160-191. Gilg, E., 1896. Beitrage zur Kenntnis der Gentianaceae I. Bot. Jahrb., 22: 301-347. , 1906. Beitrage zur Kenntnis der Gentianaceae HI. Gentianaceae, Andinae, Fedde, Rep. Nov. Sp., 2: 34-52. , 1917. Monographische Zusammenstellung der Gentiana-Arten Sud- Amerikas, Bot. Jahrb., 54(Beib. 118): 4-89. Gillet, J. N., 1957. A. revision of the north American species of Gentianella Moench,.4nn. Missouri Bot. Card., 44(3): 195-262. Grisebach, A. H. R., 1838 (1839). Genera et species Gentianearum. Stuttgart. Holub, J., 1967. Neue namen innerhalb der Gattungen Gentianella Moench, Gentianopsis Ma and Comas toma (Wettst.) Toyokuni, Folia Geobot, Phyto- tax. Praha, 2: 115-120. , 1968. Einige neue nomenklatorische Kombinationen innerhalb der Gentianinae, Folia Geobot. Phytotax. Praha, 3: 217-218. , 1983. A brief note on Slovak taxa of Gentianella, Preslia, 55(4): 371-373. Pringle, J. S., 1981, Nomenclatural transfers and taxonomic notes on some South American Gentianaceae, Phytologia, 48(4): 281-285. , 1987. A new species and taxonomic notes on Gentianella (Gentianaceae) in South America, Sida, 11(4): 357-369. Bull. not. Hist. Mus. Lond. (Bot.) 23(2): 67-70 Issued 25 November 1993 Studies in Hypericum: validation of new names NORMAN K.B. ROBSON do Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Introduction 67 Section 20. Myriandra (Spach) R. Keller 67 Section 23. Triadenioides Jaub.& Spach 68 Section 25. Adenotrias (Jaub.& Spach) R. Keller 68 Section 27. Adenosepalum Spach 69 References .. ..70 SYNOPSIS. The new names in Part 6 of 'Studies in the genus Hypericum L. (Guttiferae)' are validated in advance of publication of the main work. They are:- new taxa (tax. nov.): H. hypericoides (L.) Crantz subsp. prostratum N. Robson (Sect. 20. Myriandra), H. fieriense N. Robson (Sect. 23. Triadenioides), subsects. Aethiopica N. Robson, Pubescentes N. Robson and H. collenettiae N. Robson (Sect. 27. Adenosepalum); new combinations (comb, et stat. nov.): H.nitidum subsp. cubense (Turcz.) N. Robson, H. nitidum subsp. exile (P. Adams) N. Robson,//. aegypticum L. subsp. maroccanum (Pau) N. Robson, H. annulatum Moris subsp. intermedium (Steudel ex A. Rich.) N. Robson, H. annulatum subsp. afromontanum (Bullock) N. Robson. INTRODUCTION Part 6 of 'Studies in the genus Hypericum L. (Guttiferae)', a monographic series that is intended to cover the whole genus (Robson 1977, 1981, 1985, 1987, 1990), will contain accounts and analyses of Sections 20-28. As this part will not be published before 1994, it is necessary to validate in advance of publication the new names that will appear in it. Section 20. MYRIANDRA (Spach) R. Keller Hypericum nitidum Lam., Encycl. 4:160 (1797). When the populations from Cuba and Belize in the H. nitidum group are considered along with those in the U.S.A., they can be divided into three subspecies: a) Cuba and Belize (subsp. cubense), which is related to the Cuban H. limosum Griseb.; b) south-eastern U.S.A. (subsp. nitidum); and c) western Cuba and north-western Florida (subsp. exile), which is related to H. brachyphyllum (Spach) Steudel from south- eastern U.S.A. H. nitidum subsp. nitidum Leaves subcoriaceous, apex obtuse to rounded-apiculate, margin loosely inrolled leaving lower lamina partly exposed. Sepals obtuse to shortly apiculate. Capsule cylindric. U.S.A. (south-eastern Alabama to southern N. Carolina). The Natural History Museum, 1993 H. nitidum subsp. cubense (Turcz.) N. Robson, comb, et stat. nov. H. cubense Turcz. in Bull. Soc. Nat. Moscou 31(1): 384 (1858). H. fasciculatum sensu Alain in Leon & Alain, Fl. Cuba 3:317 (1953) pro parte. Leaves coriaceous, apex rounded-apiculate to rounded, mar- gin tightly inrolled leaving only midrib exposed. Sepals obtuse to rounded-apiculate. Capsule cylindric to rarely ovoid-conic. Cuba: (Oriente, Las Villas, Isla de Pinos), Belize (El Cayo). Hypericum nitidum Lam. subsp. exile (P. Adams) N. Robson, comb, et stat. nov. H. galioides var. cubense Griseb., Cat. PI. Cuba: 39 (1866), non H. cubense Turcz. (1858). H. galioides var. axillare sensu Griseb., loc. cit., pro parte (1866). H. galioides sensu Sauvalle, Fl. Cubana: 8 (1868). H. fasciculatum sensu Alain in Leon & Alain, Fl. Cuba 3: 317 (1953) pro parte excl. typum. H. exile P. Adams in Contr. Gray Herb. Harv. no. 189: 33 (1962). Leaves chartaceous, apex acute to long-acuminate, margin tightly inrolled leaving only midrib exposed. Sepals acute to long- acuminate. Capsule cylindric to narrowly conic. U.S.A. (north-western Florida), Cuba (Pinar del Rio, Isla de Pinos). 68 N.K.B. ROBSON Hypericum hypericoides (L.) Crantz, Inst. rei herb. 2:520 (1776). Ascyrum hypericoides L., Sp. pi. :789 (1753) excl. syn. Hort. Cliff, et Plukenet., 2nd ed.: 1108 (1763) excl. syn. Plukenet. Type: Hispaniola, Hypericoides frutescens erecta, flore luteo Plumier, Nov. pi. amer.: t.7 (1703), lectotype (Robson, 1980:272). Plumier (1703) distinguished two species from Hispaniola in his new genus Hypericoides: H. frutescens, erecta, flore luteo and H. frutescens, humi-fusa, flore luteo. The erect species is not distinguishable taxonomically from Hypericum hyperi- coides subsp. hypericoides as it is known in the other islands of the Greater Antilles, the Bahamas, Bermuda and the mainland (eastern N. America, eastern Mexico to Honduras Republic); but the other represents a taxon that has appar- ently evolved within Hispaniola, occurring at high altitudes in the Dominican Republic. The occurrence of a few somewhat intermediate specimens in the region between 1600 and 2000 m, where it co-exists with the typical form, indicates that the appropriate rank for this taxon is subspecies. Hypericum hypericoides subsp. prostratum N. Robson, subsp. nov. Hypericoides frutescens, humi-fusa, flore luteo Plumier, Nov. pl.amer.:52(17Q3). Ascyrum foliis lanceolato-linearibus, biglandulosis, ramidiffu- sis Burman, PI. amer.: 146, t.152 f.2 (1758). A subsp. hypericoides habitu prostrate, folium minoribus anguste oblongis vel oblongo-spathulatis, differt. Type: Dominican Republic, San Juan, Sabana Nueva, Cor- dillera Central N. of Rio Arriba del Norte, 1950 m, 17-20.ix.1944, R.A. & E.S. Howard 9080 (BM!, holotype; GH!, MICH!, NY!, US!, isotypes). Plant prostrate, with stems ? numerous, radiating and branching, forming mats. Leaves 3-8(-10) X 1-2.5 mm, narrowly oblong to oblong-spathulate. Inflorescence- branching pseudo-dichotomous. Open Pinus forest, grassland and open slopes, (1600-)1800-2900 m. Dominican Republic (La Vega, Santiago, San Juan, Peravia). Section 23. TRIADENIOIDES Jaub. & Spach Socotra, an island of endemics, is critical for the understand- ing of the evolution of Hypericum. It is already known to contain four endemic species, H. balfourii N. Robson and H. socotranum Good (Sect. 1. Campy losporus), the latter with two subspecies (Robson 1985), H. scopulorum Balf. f. and H. tortuosum Balf. f. (Sect. 23. Triadenioides). To these must be added a new species based on one collection from the Hagghiher Mountains by Smith and Lavranos. It is clearly near H. scopulorum but is more woody and larger in all its parts and has petiolate leaves, and the lower leaf surface, petiole and young stems are covered with a puberulous indumentum. Hypericum fieriense N. Robson, sp. nov. H. scopulorum Balf. f. affinis, sed caulibus crassioribus lignosioribus, foliorum lamina ovata subtus cum petiolo et ramis junioribus puberula, inflorescentia 3-5-florata, floribus maioribus sepalis crassioribus, capsulis coriaceis valvis leviter angustissime vittatis fere laevibus, inter alia differt. Type: Socotra, Hagghiher Mountains(1235'N, 5403'E), below Fieri peaks, 1350 m, 21. iv. 1967, Smith & Lavranos 475 (K!, holotype & isotype). 'Low scrub among Dracaena cinnabari trees', 1350 m. Socotra (Hagghiher Mts). Section 25. ADENOTRIAS (Jaub. & Spach) R. Keller Hypericum aegypticum L., Sp. PL: 784 (1753). H.aegypticum comprises a series of disjunct populations forming a morphological reduction trend from south Morocco to Crete and Cyrenaica (not Egypt). This trend is almost continuous, but it is possible to recognize three, rather poorly differentiated subspecies. Only essential synonymy is given here. Hypericum aegypticum L. subsp. maroccanum (Pau) N. Robson, stat. nov. H. aegypicum var. maroccanum Pau in Cavanillesia 4: 157 (1932) ['maroccana'] in Spanish; Maire in Bull. Soc. Hist, nat. Afr. N. 24: 206 (1933), in Latin. Plant erect, (0.15-)0.3-2 m tall, with branches erect to ascending. Leaves sessile, plane; lamina (7-)9-18 X (2-)3-4 mm, narrowly elliptic or narrowly oblong-elliptic, acute. Sepals 5-6 mm long. Petals 10-12(14?) mm long. Morocco (south-west), Algeria (southern Atlas Mts). H. aegypticum L. subsp. webbii (Spach) N. Robson, comb. et. stat. nov. Triadenia webbii Spach in Annls Sci. nat. (Bot.) II, 5: 174, t. 5A (1836), Hist. nat. veg. Phan. 5: 372 (1836). Plant erect to loosely spreading, 0.04-0.4 m tall, with branches erect or usually spreading and often tortuous, forming bushes up to 1 m across. Leaves subsessile to shortly (c. 0.3 mm) petiolate, plane or subcucullate; lamina 4-10 X 1.5-3 mm, narrowly oblong to broadly elliptic; acute to obtuse. Sepals 5-6 mm long. Petals 8-14 mm long. Lampedusa, Malta, Sardinia, Greece (Ionian Islands, west- ern Peloponnisos), Crete. H. aegypticum L. subsp. aegypticum Plant spreading 0.05-0.18 m tall, with branches spreading and tortuous, forming low bushes. Leaves shortly (0.4-0.5 mm) petiolate, always (?) incurved-cucullate; lamina 3-6 X 1-2 mm, narrowly oblong to broadly elliptic, acute. Sepals 3.5-5 mm long. Petals 6.5-9 mm long. Libya (Cyrenaica-Jebel el Akhdar). STUDIES IN HYPERICUM 69 Section 27. ADENOSEPALUM Spach When Sect. Adenosepalum (sensu Robson, 1977) has been 'purified' by the removal of the tropical Asian species(//. elodeoides group) to Sect. 9. Hypericum sensu lato and three Turkish species (H. huber-morathii N. Robson, H. minutum P.H. Davis & Poulter, H. formosissimum Takht.) to Sect. 12. Origanifolia, the remaining species form a natural group distributed over most of Africa, Macaronesia, Europe, Medi- terranean Asia and western Arabia. It can be divided into four subsections, as follows: 1. Subsect. Aethiopica N. Robson, subsect. nov. Planta omnino glabra. Folia libera. Bracteae bracteolaeque haud glanduloso-auriculatae. Typus: H. aethiopicum Thunb. 2. Subsect. Pubescentes N. Robson, subsect. nov. Planta usque ad sepala vel ad partem inferiorem inflorescentiae vel rare ad basin inflorescentiae indumentum ferens. Folia libera. Bracteae bracteolaeque haud glanduloso- auriculatae. Typus: H. pubescens Boiss. 3. Subsect. Caprifolia N. Robson, subsect. nov. Planta usque ad basin inflorescentiae indumentum ferens. Folia inter- dum inferiora excepta binatim conjuncta. Bracteae bracteolaeque interdum glanduloso-auriculatae. Typus: H. caprifolium Boiss. 4. Subsect. Adenosepalum. Planta usque ad basin inflores- centiae indumentum ferens vel rare caulibus vel foliis vel omnino glabra. Folia libera. Bracteae bracteolaeque per- saepe glanduloso-auriculatae. Typus: H. montanum L. Hypericum collenettiae N. Robson, sp. nov. H. sp. aff. sinaicum sensu Collenette, ///. Guide Fls Saudi Arabia: 262 + photos (1985). H. somalensi N. Robson affinis, sed indumento breviori, caulibus internodiis plerumque foliis brevioribus, foliis angus- tioribus, inflorescentia paucioriflora laxiore, floribus maiori- bus, sepalis petalisque glandulis nigris laminaribus ornatis, inter alia differt. Type: Saudi Arabia, Asir, Taif-Abha road 82 km S. of Baljurshi, Wadi Mahra, c.1800 m, 5.viii.l982 (fr), Collenette 3752 (BM!, holotype; K!, isotype). Collenette 1401 (K) is another collection (16. iv. 1979) from the same population. Shady rock crevices, c.1800 m. Saudi Arabia (Asir). This apparently solitary population is intermediate in mor- phology and distribution between H. somaliense N. Robson (N. Somalia) and H. sinaicum Hochst. ex Boiss. (Sinai and adjacent Saudi Arabia) but distinct from both these species. According to Mrs. Collenette, its habitat is under some threat. An earlier specimen from the same area (between Baljurshi and Abha) has recently come to light and may represent a second population:- Bashwat, 9.viii. 1975 (fl. & fr.), A. El-Sheikh in Herb. KSUH 1067 (KSUH). Hypericum annulatum Moris, Stirp. sard, elench.: 9 (1827). Although originally regarded as endemic to Monte Santa Vittoria esterzili in Sardinia, this species has subsequently been found in another locality in that island (Nodu 'e Littipori) (Arrigoni et al. 1973). Meanwhile it had been treated as conspecific with the central Balkan H. degenii Bornm. and the Ethiopian and East African H. intermedium Steudel ex A. Rich. (Milne-Redhead, 1953 a,b\ Robson, 1958, 1968; etc.). Further study of this variable species and its unusual discon- tinuous distribution has shown a) that the widely separate populations can be differentiated as three subspecies and b) that the Mt. Elgon endemic H. afromontanum Bullock is no more than a high-altitude form of the East African subspe- cies. The Ethiopian/Arabian subsp. intermedium is morpho- logically nearest to H. montanum L., the sister species of H. annulatum. H. annulatum Moris subsp. annulatum H. perfoliatum var. annulatum (Moris) Fieri in Fiori & Paoletti, Fl. Anal. It. 1: 389 (1898), Nuovo Fl. Anal. It. 1: 524 (1924). H. degenii Bornm. in Magyar Bot. Lap. 9:90 (1910). Stem without red or black glands, densely shortly pubescent. Leaves without laminar black glands, densely shortly pubes- cent. Sepals long- to short-glandular-ciliate (cilia shorter than to two or more times as long as glands), with laminar glands all pale. Petals without punctiform laminar black glands, not tinged red in bud. Sardinia, [Yugoslavia] (southern Serbia, Macedonia), north- ern Albania, Bulgaria, northern Greece. H. annulatum Moris subsp. intermedium (Steudel ex A. Rich.) N. Robson, comb, et stat. nov. H. intermedium Steudel ex A. Rich., Tent. Fl. Abyss. 1: 95 (1847). H. intermedium forma obtusifolium R. Keller ex Moggi & Pisacchi in Webbia 22: 272 (1967) in synon. H. annulatum sensu Cufod. in Bull. Jard. hot. Etat Brux. 29, Suppl.: 588 (1959); Moggi & Pisacchi in Webbia 22: 272 (1967) pro parte; Collenette, ///. Guide Fls Saudi Arabia: 261 + photo (1985), pro parte excl. typum et spec. cit. ex Harar. Stem without or rarely with few black glands, sparsely puberulous to glabrous. Leaves without laminar black glands, sparsely and very shortly pubescent to puberulous or glabrous. Sepals short- to long-glandular-ciliate (cilia shorter than to two or more times as long as glands), occasionally with some laminar glands black. Petals with few (rarely more numerous) punctiform laminar black glands, rarely red- veined in bud. Saudi Arabia (Asir), Sudan Republic (southern Red Sea Hills), northern Ethiopia (Eritraea to L. Tana and northern Shoa). H. annulatum Moris subsp. afromontanum (Bullock) N. Robson, comb, et stat. nov. H. afromontanum Bullock in Kew Bull. 1932: 492 (1932). H. annulatum sensu Milne-Redh. in Kew Bull. 8: 435 (1953), Fl. Trop. E. Afr., Hypericac.:6 (1953); Moggi & Pisacchi in Webbia 22: 272 (1967) pro parte; Agnew, Upland Kenya Wild Fls:186 (1974); N. Robson in Bamps, Robson & Verde., Fl. Trop. E. Afr. Guttif.:30 (1978). 70 N.K.B. ROBSON Stem usually with numerous black (very rarely red) glands, densely to sparsely puberulous or rarely glabrous. Leaves sometimes with few to numerous laminar black glands, puberulous above and densely pubescent beneath or very rarely wholly glabrous. Sepals long-glandular-ciliate (more than twice as long as glands), usually with some or all laminar glands black. Petals with few distal or numerous scattered punctiform laminar black glands, always (?) red-tinged in bud. Southeastern Ethiopia (Harar), East Africa (eastern Uganda, southwestern Kenya, northern Tanzania). The Harar population is somewhat intermediate between subsp. afromontanum and subsp. intermedium, but is more similar to the former than the latter. REFERENCES Arrigoni, P.V., Corrias, S.D., Nardi, E. & Valsecchi, F. 1973. Nuovo stazioni di Hypericum annulatum Moris e Ribes multiflorum Kit. ssp. sandalioticum Arrig. in Sardegna. Webbia 28:423-425. Milne-Redhead, E. 1953a. Tropical African plants: XXIV, Hypericaceae. Kew Bull. 8:434. 19536. Hypericaceae. In W.B. Turrill & E. Milne-Redhead (Eds), Flora of Tropical East Africa. London. Plumier, C. 1703. Nova plantarum americanum genera: 51. Paris. Robson, N.K.B. 1958. The genus Hypericum in Africa south of the Sahara, Madagascar and the Mascarenes. Kew Bull. 12:443-446. 1968. Guttiferae (Clusiaceae). In T.G. Tutin et al. (Eds), Flora Europaea 2:261-269. 1977. Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. Bull. Br. Mus. nat. Hist. (Bot.) 5:291-355. 1981. Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bull. Br. Mus. nat. Hist. (Bot.) 8:55-226. 1985. Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1. Campylosporus to 6a. Umbraculoides. Bull. Br. Mus. nat. Hist. (Bot) 12:163-325. 1987. Studies in the genus Hypericum L. (Guttiferae). 7. Section 29. Brathys (part 1). Bull. Br. Mus. nat. Hist. (Bot.) 16:1-106. 1990. Studies in the genus Hypericum L. (Guttiferae). 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. Bull. Br. Mus. nat. Hist. (Bot.) 20:1-151. Bull. not. Hist. Mus. Lond. (Bot.) 23(2): 71-177 Issued 25 November 1993 Generic monograph of the Asteraceae-Anthemideae KARE BREMER Department of Systematic Botany, Uppsala University, Villavdgen6, 5-752 36 Uppsala, Sweden CHRISTOPHER JOHN HUMPHRIES Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS Introduction 73 Materials and methods 73 Descriptive terminology 74 Taxonomic concepts 74 Cladistics 74 Tribal and subtribal divisions 74 Classification 76 Subtribal classification 76 Generic classification 77 Characters of the Anthemideae 78 Character scoring 78 Distribution 83 Key to genera 84 Anthemideae Cass 90 1 . Ursiniinae Bremer & Humphries 91 1 . Ursinia Gaertner 93 2. Lasiospermum Lagasca 94 3. Phymaspermum Less 94 4. EumorphiaDC 94 5. Gymnopentzia Benth. in Benth. & Hook, f 95 6. Hymenolepis Cass 95 7. Athanasia L 95 8. Asaemia (Harvey) Harvey ex Benth. in Benth. & Hook, f 96 2. Cancriniinae Bremer & Humphries 96 9. Trichanthemis Regel & Schmalh 97 10. Ugamia Pavlov 98 11. Richteria Karelin & Kir 98 12. Allardia Decne 98 13. Cancrinia Karelin & Kir 99 14. Cancriniella Tzvelev 9 3. Tanacetinae' Bremer & Humphries 99 15. Tanacetum L 100 16. Opisthopappus Shih 104 17. Tanacetopsis (Tzvelev) Kovalevsk 104 18. Xylanthemum Tzvelev 105 19. LepidolophaWmk\ 105 20. Hippolytia Polj 1 5 21 . Heliocauta Humphries 106 4. Gonosperminae Bremer & Humphries 106 22. LugoaDC 1 23. Gonospermum Less 107 24. Inulanthera Kallersjo I 1 5. Handeliinae Bremer & Humphries 1 25. Lepidolopsis Polj 1 9 26. Polychrysum (Tzvelev) Kovalevsk 1' 27. Pseudohandelia Tzvelev 1 9 28. Handelia Heimerl ! 29. Sclerorhachis (Rech. f.) Rech. f 11 The Natural History Museum, 1993 72 K. BREMER AND C. J. HUMPHRIES 6. Artemisiinae Less, emend. Bremer & Humphries 110 30. Brachanthemum DC 113 31. Dendranthema (DC.) Des Moul 113 32. Arctanthemum (Tzvelev) Tzvelev 114 33. Tridactylina (DC.) Sch. Bip 115 34. Ajania Polj 115 35. Phaeostigma Muld 115 36. Stilpnolepis H. Kraschen 1 16 37. Ajaniopsis Shin 116 38. Filifolium Kitam 116 39. Sphaeromeria Nutt 117 40. Kaschgaria Polj 117 41. Seriphidium (Besser ex Hooker) Four 117 42. Crossostephium Less 120 43. Artemisia L 120 44. Neopallasia Polj 125 45. Turaniphytum Polj 125 46. Mausolea Polj 125 47. Picrothamnus Nutt 125 7. Achilleinae Bremer & Humphries 126 48. SantolinaL 128 49. Otanthus Hoffsgg & Link 128 50. Achillea L 128 5 1 . Anacyclus L 129 52. Leucocyclus Boiss 130 53. Mecomischus Cosson ex Benth. in Benth. & Hook 130 54. Chamaemelum Miller 130 55. Rhetinolepis Cosson 130 56. Cladanthus Cass 131 8. Anthemidinae Dumort. emend. Bremer & Humphries 131 57. Anthemis L 132 58. NanantheaDC 134 9. Chrysantheminae Less, emend. Bremer & Humphries 134 59. Chrysanthemum L 135 60. Heteranthemis Schott 135 61 . Ismelia Cass 135 62. Argyranthemum Webb ex Schultz-Bip 136 10. Leucantheminae Bremer & Humphries 136 63. Lepidophorum Necker ex Cass 139 64. Nipponanthemum Kitam 139 65 . Leucanthemella Tzvelev 139 66. Nivellea Wilcox, Bremer & Humphries 140 67. Phalacrocarpum (DC.) Willk 140 68. Leucanthemopsis (Giroux) Heyw 140 69. Hymenostemma ( Kunze) Willk 140 70. Prolongoa Boiss 141 71. Leucanthemum Miller . 141 72. Rhodanthemum (Vogt) Bremer & Humphries 141 73. Leucoglossum Wilcox, Bremer & Humphries 142 74. Chlamydophora Ehrenb. ex Less 142 75. Chrysanthoglossum Wilcox, Bremer & Humphries 143 76. Glossopappus Kunze 143 77. Coleostephus Cass 143 78. Ptagius L'Herit. ex DC 143 1 1 . Thaminophyllinae Bremer & Humphries 144 79. Osmitopsis Cass 145 80. Adenanthellum B. Nord 145 81 . Inezia E . Phillips 146 82. Lidbeckia P. J. Bergius 146 83. Thaminophyllum Harvey . 146 12. Matricariinae Bremer & Humphries 146 84. Cymbopappus B. Nord 151 85. PentziaThunb 151 86. Marasmodes DC 152 87. Rennera Merxm 152 88. Oncosiphon Kallersjo 152 89. Otospermum Willk 153 GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 73 90. Heteromera Pomel 153 9 1 . Daveaua Willk . ex Mariz 153 92 . Matricaria L 153 93. Microcephala Pobed 154 94. Endopappus Schultz-Bip 154 95 . Myxopappus Kallersjo 155 96. Foveolina Kallersjo 155 97. Lonas Adans 155 98. Tripleurospermum Schultz-Bip 155 99. Aaronsohnia Warb. & Eig 156 100. Leucoptera B . Nord 157 Wl.AdenoglossaB. Nord 157 W2.HilliardiaE. Nord 157 103. CotulaL 157 104. Leptinella Cass 159 105. Soliva Ruiz Lopez & Pavon 159 106. Schistostephium Less 160 107. HippiaL 160 108. Eriocephalus L 160 Excluded genera 161 References 161 Index . . 165 SYNOPSIS. The Asteraceae tribe Anthemideae is revised. In all, 12 subtribes, 108 genera and 1741 species are recognized; nine tribes are described as new and three, the Anthemidinae, the Artemiisinae and the Chrysanthem- ineae are emended in circumscription. Four new genera are described and many have been revised in circumscrip- tion. The definitions of the tribe, subtribes and genera are expressed in terms of sister group relations and the most robust hypotheses of character distribution utilizing the principles of synapomorphy, parsimony and character congruence. Phylogenetic relationships were determined using the tree-building computer program, HENNIG86. Cladograms of the subtribes and genera are provided together with a synoptic character analysis of each individual clade. A key to all genera is provided, each genus is described, all species currently recognized are listed with a brief synonomy and the relevant nomenclature and taxonomic changes are discussed in detail. Summaries of distributions are given in tables and within the generic accounts. The account ends with a list of excluded taxa and a taxonomic index. INTRODUCTION MATERIALS AND METHODS The Asteraceae tribe Anthemideae is one of the largest tribes of the family with 1741 species predominantly distributed in Eurasia, North and South Africa, with fewer species in North America and Australasia. The circumscription of the tribe is new but generally follows that outlined by Bentham in Bentham & Hooker (18730). The principal taxonomic problems within the tribe are almost entirely relationships between genera but also circum- scription of genera especially within subtribal groups such as the Artemiisinae, the Chrysantheminae, and the 'Tanaceti- nae'. This revision deals with the systematics and generic circumscription of the Anthemideae utilizing modern cladistic methods. The nomenclature and descriptive taxonomy have been fully revised in line with our own analyses and the literature scanned up to the present time. In addition to the formal taxonomic treatment a complete description of charac- ter definitions, character distributions and morphologies has been provided. Phytogeography and phytogeny have been included to give the correct context for the novel classifica- tions and a brief historical review of tribal and subtribal groupings is also presented. The revision is based on herbarium specimens and living collections in botanic gardens. The rationale was to study as many of the recognized species as possible during the course of the work. Those species which we have been unable to locate are indicated with an asterisk (*) in the individual lists at the end of each generic account. We have studied speci- mens from the following herbaria; (Index Herbariorum', abbreviations as in Holmgren et al, 1981) BM, BOL, BRY, E, F, GH, K, LE, M, MO, NBG, NY, PRE, S, US. Several species have been transferred during the course of the revision and new combinations are provided. For some North African species new combinations based on names in the unpublished thesis of Helen Wilcox are validated here. The generic descriptions and the character matrices are based on dried herbarium specimens and living specimens when these were available. Cross-sections of cypselas were made from reconstituted collections, softened in water, embedded in paraffin and ceresin wax, cut by microtome and stained with safranin and light green. The material examined has not been listed but all specimens examined were of known provenance. 74 K. BREMER AND C. J. HUMPHRIES Descriptive terminology The descriptions and terms used throughout this work follow those in Featherly (1954) and Stearn (1966). The terminology for outlines and plane shapes adopted is that of the Systemat- ics Association Committee for Descriptive Biological Termi- nology (1962). Descriptions of variation in corolla morphology generally follow those of Jeffrey (1977). Taxonomic concepts The taxonomic concepts in this revision are based on mor- phological studies. In most generic revisions, particularly in large families such as the Asteraceae, genera and higher taxa are frequently circumscribed on uncritical character analyses. This approach is considered by us to be unacceptable since in no way does it reflect the principles of cladistics. Generic names, as with all taxa of any rank, can only be applied to monophyletic groups. To recognize monophyletic groups which can be designated as genera (or species, or tribes) the concept of resemblance has to be resolved so that hypotheses of characters can be used to generate hypotheses about groups. Evolutionary novelties or synapomorphies, are those characters which diagnose monophyletic groups and have been utilized here accordingly (see Humphries & Funk, 1984). Cladistics The technique we have used to diagnose monophyletic groups is the Wagner algorithm as implemented in the computer program Hennig86 (Farris, 1988). Hennig86 is an interactive program for cladistic analysis which can obtain most parsimonious trees by either exact or heuristic calcula- tions. Characters were scored from the raw character matri- ces into discrete binary codes. Many of the characters we used were simply presence or absence characters and thus all characters were discretely coded in either a or 1 state. However, a considerable number of the characters were polymorphic in their distribution especially in the larger genera. For precise details of the conventions adopted refer to the section on character coding. The character scores are presented in twelve matrices (Tables 4, 6, 8, 10, 12, 13, 15, 17, 19, 21, 23, and 25; see taxonomic accounts below); eleven are for individual subtribes (Anthemidineae is combined with the Chrysantheminae) and one for subtribes represented as individual taxa (Table 4). For the analysis of subtribes the matrix (Table 4) was analysed as presented. For the subtribe matrices there are three blocks of scores within each matrix; the first represents character distributions at a level higher than subtribe, the second is characters varying within the subtribe and the third provides a summary of ambiguous information for a given subtribe. Cladistic analysis for the subtribes is based on the central block of characters in each of the data matrices. Trees were calculated using a variety of options in Hen- nig86. All four commands, and five of the nine tree building options available in Hennig86 were utilized depending upon the size and complexity of each data matrix. The options were as follows; hennig (h), which constructs one tree with one pass through the data; mhennig* (mh*) which constructs several trees, each by a single pass, adding the terminal taxa in several different sequences. Branch swapping was applied to each of the initial trees and one tree from each initial one was retained in the memory; branch-breaker* (bb*) a similar option to mhennig* but all of the obtained equally parsimoni- ous trees were retained, as was allowed in the assigned memory of 128k in the computer; implicit enumeration (ie*) which generated all possible shortest trees. The results guar- anteed shortest trees and all equal length trees were retained in the available tree space; ie- which found one guaranteed shortest tree by implicit enumeration. Exact analyses (implicit enumeration) were applied to the Achilteinae, Anthemidinae, Cancriniinae, Chrysantheminae, Gonosperminae, Handeliinae, Tanacetinae, Thaminophylli- nae and the Ursiniinae. The hennig, mhennig* and bb* options were applied to the larger groups including the Anthemideae tribal analysis and the analyses of the Arte- misiinae, Leucantheminae and Matricariinae subtribes. The trees were all rooted utilizing the outgroup - a taxon scored from a North American group of genera within the Heliantheae tribe. All scored characters in each of the matrices were given equal weight. Unknown scores were left in since Hennig86 has the facility for coping with them. In the cases where more than one equally parsimonious tree was obtained the nelsen command was activated to produce a strict consensus tree (Farris, 1988). However, for the purpose of the classification only one of the final trees is figured in the text (see Figs 1, 2, 7, 11, 17, 20, 23, 31, 36, 41, 47 and 51; see taxonomic text below). Each tree was diagnosed using the xsteps command in order to determine the states of the hypothetical ancestors on each tree (i.e. the node states) and to generate statistics on length and fit. Precise details of character changes along each clade are provided in the subtribe accounts. Each cladogram is pro- vided with a unique letter and number code and the length and consistency of each tree is provided in the figure caption. The cladograms should be used in conjunction with the descriptive accounts of the character changes. Outgroup Bremer (1987) has put forward evidence for the hypothesis that Anthemideae and Heliantheae sensu lato are sister groups within a larger clade consisting of the Calenduleae, the Senecioneae, the Anthemideae and the Heliantheae. Bremer also showed that the Anthemideae and its postulated relatives in the Heliantheae share similar ray floret epidermis as elucidated in the work of Baag0e (1977). The two tribes also have similar habit and foliage with trinerved and dis- sected leaves especially in the subtribe Helenieae, and thus the North American species of this group were selected as the outgroup. It has been pointed out to us by Charles Jeffrey (in litt.) that the results published by Palmer et al. (1988) from an analysis of chloroplast DNA would suggest that we might consider a different outgroup. The consensus tree of Palmer et al. using 926 restriction site mutations places Astereae and Anthemideae as sister groups and the Eupatorieae as an ingroup of the Heliantheae. However, we consider that their sample of taxa is so preliminary that for the time being we remain with Bremer's (1987) morphological hypothesis. TRIBAL AND SUBTRIBAL DIVISIONS The first coherent tribal classification of the Asteraceae was that of Cassini (1817, 1826) who recognized 19 tribes in the GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 75 family. The precise systematic position of the Anthemideae within the Asteraceae is shown in a diagram of the family when Cassini used a map with an oval arrangement of abutting circles to express the relationships of the tribes (King & Dawson, 1975). The Anthemideae were sandwiched between the Inuleae and the Ambrosineae. Cassini expressed the opinion that the Anthemideae are most closely related to the Heliantheae although in style structure they were very similar to the Inuleae, the Senecioneae and the Nassauvi- ineae. Cassini (1826) divided the tribe into two major groups: Anthemidees - Prototypes (Anthemideae-Archetypae) and Anthemidees - Prototypes vraies, based on the prescence or absence of receptacular scales respectively. For many genera the division is quite workable and many of them can be placed in one group or the other. However, the artificiality of this character can clearly be demonstrated in several taxa especially the genus Anthemis (Ammanthus group) in which paleate and non-paleate forms can occur within one species (Heywood & Humphries, 1977). Lessing (1832) had a quite different treatment from that of Cassini, and although his classification was avowedly artifi- cial, since it was written with identification in mind, it clearly influenced the later major treatments of de Candolle (1837), Bentham (1873a, b), and Hoffmann (1894) (see Heywood & Humphries, 1977). Lessing recognized eight tribes and the genera of the Anthemideae were included in tribe VII Senecionideae subtribes Chrysanthemeae and Artemisieae. The Chrysanthemeae were further subdivided into two taxa of indefinite rank, the Anthemideae and Chrysanthemeae, and the Artemisieae were divided up into six taxa of indefi- nite rank. De Candolle (1837) recognized Cassini's tribe Anthemideae as a subtribe and more or less followed Lessing's indefinite ranks which he categorized as divi- sions. However, he synonymised the Santolineae with the Euanthemideae. Bentham (1873a) recognized the Anthemideae as a tribe and divided it into six informal taxa. Four of these were common to the Lessing and de Candolle treatments but he also recognized the Lidbeck- ieae and the Tanaceteae as separate taxa. Of other 19th century works, that of Hoffmann (1894) is the only one of significance and he simply recognized the Cassini classifica- tion, dividing the tribe into two subtribes. Baillon (1886) provided an end of century classification of the angiosperms in his classic Histoire des plantes. For the Asteraceae he recognized eight tribes and four hundred and three genera. The sub-series Anthemidees was included in an enlarged tribe Helianthees together with the Heleniees, Senecionees and the true Euhelianthees. Most 20th century contributions have been concerned with the classification and delimitation of genera but rather silent about suprageneric classification and the relationship of the tribe to other tribes. However, Bessey (1915) provided perhaps the most novel treatment of the Asteraceae when he raised the family to the rank of order and raised fourteen tribes to family status. Family 296, the Anthemidaceae was sandwiched between families 295 and 297, the Eupatoriaceae and the Senecionidaceae respectively. However, most treat- ments have simply tended to follow the tribal groupings outlined by Bentham (1873a) with no real modifications to the status of infra-tribal groups (e.g. Carlquist, 1976; Cron- quist, 1955, 1977; Wagenitz, 1976). Poljakov (1967) recognized six subtribes in the Anthemid- eae in a treatment that to some extent resembled that of Bentham but with a more detailed consideration of the Artemisiinae. One of the most recent attempts at creating subgroups within the tribe is that of Reitbrecht (1974) who considers that the Anthemideae consists of seven provisional groups (Humphries, 1979). These include the Ursinia group, to include Ursinia and its allies, the Lasiospermum group to include the majority of the African genera, especially Lasiospermum and Eriocephalus , the Chrysanthemum group for some members of the Tanacetinae, Chrysanthemineae and Leucanthemineae as recognized by us, the Matricaria group to include the Anthemidineae and some members of the Matricariinae, and the Cotula group for various other members of our Matricariinae. Bremer (1987) undertook the first cladistic analysis of the tribes of the Asteraceae. The cladogram he obtained sug- gested a number of novel groupings. The Anthemideae were clearly placed in the monophyletic subfamily, Asteroideae, and appeared as the sister group of the Heliantheae in a subgroup consisting of the Calenduleae, the Senecioneae, the Anthemideae and the Heliantheae, with the Astereae and the Eupatorieae as sister groups. Bremer obtained his result by considering that the Anthemideae and the Heliantheae share similar ray floret cells (Baag0e, 1977), and similar foliage with trinerved and dissected leaves which are interpreted as synapomorphies for the two tribes. Some of the older groups recognized by the 19th century authors have caused problems as a result of different inter- pretations of characters considered of importance in classifi- cation. One of the most problematic groups within the Anthemideae is the Ursinia group, and Ursinia in particular. Ursinia resembles the Anthemideae in habit and the Arc- totideae in terms of superficial features of the capitulum. The interpretation as to its exact position has varied, depending mostly on the weight of individual characters rather than any form of character analysis. Bentham (1873a) placed Ursinia in the tribe Arctotideae (sensu Norlindh, 1977) mainly because of its well developed pappus scales. Cassini (1816) followed by Beauverd (1915), Merxmuller (1954), Prassler (1967), and Reitbrecht (1974) considered it to be a member of Anthemideae. Robinson & Brettell (1973) argued that the inclusion in the Anthemideae of an anomalous genus like Ursinia with its conspicuous pappus scales, widely ovate apical anther appendages, and different pollen morphology (exine without columnar structure) stretched the workable tribal concept. They proposed a new tribe, Ursinieae, which included only Ursinia. However, they were not working within a cladistic framework and have used in our opinion autapomorphic characters to define their new tribe. Accep- tance of their solution does nothing to actually clarify the position of the Ursinieae in terms of its sister group relations and would render the Anthemideae as paraphyletic if the Ursinieae were excluded from it. As described in the generic account, we consider the large pappus scales and the shape of the apical anther appendage plesiomorphic within the Anthemideae, since similar structures occur in the outgroup. The pollen was investigated by Stix (1960) and she concluded that Ursinia belongs to the Anthemideae. The presence of unique furanosesquiterpenes which must be considered as apomorphic for Ursinia and other South African genera of the subtribe Ursiniineae, in our view diagnoses the Ursinieae as a natural group and corroborates its tribal position as a member group of the Anthemideae. Bentham's informal group, the Cotuleae, has also been 76 K. BREMER AND C. J. HUMPHRIES scrutinized by a variety of workers in recent years. Bentham (18736) recognized the 'Cotuleae' mostly by 'the mutual possession of characters representing loss or reduction in the habit and various parts - involucre, paleae, ray florets, pappus, number of corolla teeth and stamens (to four) and seed-sterility of disk florets' (Lloyd, 1972a). It is a classic polyphyletic group and all of its constituent members (12 genera; Table 1) have been analysed in terms of gross morphology (Heywood & Humphries, 1977), flower and fruit structure (Bruhl & Quinn, 1990, 1991), and pollen morphol- ogy (Jarvis, 1976; Bruhl & Quinn, 1991). Most of the taxa of the Cotuleae are very small plants occurring in the southern hemisphere with structurally reduced parts. The detailed morphological analyses have shown that most taxa are mis- placed within the Anthemideae and these are marked with asterisks in Table 1. Several of these have already been transferred to other tribes (see list of excluded genera, Heywood & Humphries, 1977). The remainder are all mem- bers of our revised group, the Matricariinae, in which Cotula, Leptinella, and Soliva are all closely related genera. In addition to those studies listed by Heywood & Humphries (1977), in recent years there have been several studies of higher classification in the Anthemideae, particularly by Swedish work- ers within the Museum of Natural History, but these have concentrated on taxa selected within the tribe. For example, Nordenstam (1976a) reclassified a number of genera masquerad- ing under the name of Chrysanthemum in South Africa and Kallersjo (1988) examined the patterns of relationship in the South African Pentzia group of genera. CLASSIFICATION Subtribal classification In this study an entirely new subtribal classification is pro- posed. We have grouped genera into larger monophyletic entities, subtribes, rather than by a more traditional method of dividing the tribe using particular characters. We are fairly confident that we have identified a number of monophyletic entities, and produced a number of well supported cla- dograms. At this time we find it appropriate to recognize 12 subtribes, most of which represent generic groups that are fairly well understood as monophyletic. Several peculiar genera of uncertain position have been more or less provi- sionally accommodated in otherwise homogenous subtribes. Table 1 The 'Cotuleae' group as recognized by Bruhl & Quinn (1990, 1991). 1 Abrotanella * 2 Centipeda * 3 Ceratogyne * 4 Cotula 5 Dimorphocoma * 6 Elachanthus * 1 Isoetopsis * 8 Leptinella 9 Nananthea 10 Plagiocheilus * 1 1 Soliva 12 Tripleurospermum = excluded genera We think it is better to present a hypothesis on the relation- ships of such genera of uncertain position rather than simply listing them at the end of the tribe. However, genera excluded from the Anthemideae are simply listed at the end of the revision. The predominantly Asian subtribe 'Tanacetinae' is only a provisional taxon in this treatment. It is at best paraphyletic and will probably turn out to be polyphyletic in later studies. It contains the large genus Tanacetum, a dubious taxon of critical importance in future studies of the Anthemideae. In our study we have grouped together a number of genera that appear to have their sister groups within Tanacetum. Further- more, it is possible that several other subtribes also have their sister groups within Tanacetum. It is clear to us that the subtribe Tanacetinae', and most particularly Tanacetum itself, are unresolved taxa. Ursiniinae, Gonosperminae, Handeliinae, Chrysanthemi- nae and Thaminophyllinae are homogeneous and monophyl- etic subtribes without inclusion of problematic genera. Some generic affinities within these subtribes have been understood by earlier authors, but in general they are emended here. For example, the Canary Island Lugoa and Gonospermum, a well known generic pair, are associated with the South African Inulanthera, and the three genera form our subtribe Gono- sperminae. Ursiniinae is the South African group containing fura- nosesquiterpenes (Greger, 1977, and references therein) and the precise delimitation of the Ursiniinae was undertaken by Kallersjo (1986). Handeliinae contains a number of small Asian genera. The association of Sclerorhachis with the other genera of the Handeliinae is new. The affinities between the other genera are described already by Tzvelev (in Komarov, 1961) although he did not recognize the subtribe and he placed them in widely different positions in his floristic classification. Chrysantheminae as we understand it is a small group that has been identified by earlier authors (see Humphries, 1976; Heywood & Humphries, 1977, and references therein). Thaminophyllinae is a South African subtribe that hitherto has hardly been considered, although affinities of some genera have been discussed by recent authors (Nordenstam, 1976; Bond, 1980). Cancriniinae, Artemisiinae, Achilleinae, Anthemidinae, Leucantheminae and Matricariinae are subtribes that we also consider monophyletic with the character information used in this study. Further investigations may result in changes in their delimitation, however, since a number of problematic genera are involved and the defining characters are rather few in some subtribes. Affinities between several of the genera within each of these subtribes have been indicated by earlier authors but their precise delimitations here are entirely new. The Asian Cancriniinae seems homogeneous but may eventually become a larger group, as its relatives probably occur within the undefined subtribe Tanacetinae'. The affini- ties between some of the genera of the Cancriniinae have also been described by Tzvelev (in Komarov, 1961). The mainly Asian and large subtribe Artemisiinae contains the Artemisia group of genera, an assemblage that has been recognized for some time (e.g. Besser, 1829). We have added their plesiomorphic relatives to this group. The relationships of some of these genera to Artemisia and its relatives have been indicated by Poljakov (1955) and Tzvelev in Komarov (1961). GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 77 The Eurasian/North African subtribe Achilleinae is a generic group hitherto unrecognized. Interrelationships between a few of the genera have been understood (e.g. Achillea, Anacydus, and Leucocydus, Humphries, 1979; Greger, 1977; Valant-Vetschera, 1981) but the subtribe is a new generic grouping. Santolina is a plesiomorphic genus of uncertain position. It may have close relatives within the 'Tanacetinae'. In this treatment the Anthemidinae contains only two genera: Anthemis and Nananthea, the latter a specialized genus of uncertain position, and discussed at greater length in the generic account. The predominantly Mediterranean Leucantheminae includes a large subgroup, the Leucanthemum group, which has been largely elucidated by the studies of Wilcox (1977) and is considered by us to be a well supported monophyletic unit. Leucanthemopsis is generally associated with Leucan- themum (Hey wood, 1954, 1976) and to it we add Phalacro- carpum, Hymenostemma and Prolonged. There are other small genera of more uncertain position, such as Lepidopho- rum, Nipponanthemum, Leucanthemella and Nivellea. Pend- ing further studies, we have considered it worthwhile to include them in the Leucantheminae. The large cosmopolitan subtribe, Matricariinae, represents one of the more interesting generic groupings presented in this study. We consider it monophyletic with reservations for a few genera of possible uncertain position such as Rennera and Oncoslphon. Within the Matricariinae there is the south- ern hemisphere Cotula group, an homogeneous, monophyl- etic unit, quite distinct from the generic assemblages introduced by Bentham (1873a) and discussed by more recent authors (see e.g. Heywood & Humphries, 1977; Bruhl & Quinn, 1990, 1991). We have refrained from recognizing the Cotula group as a subtribe, since it would probably make the remaining Matricariinae paraphyletic. Related to the Cotula group are several predominantly Mediterranean and South African genera, the interrelationships of which have hitherto been unprescribed. Although we feel that the majority of our subtribes are well supported and easily distinguished, subtribal interrelation- ships have been difficult to assess and the main cladogram (Fig. 1) is largely unresolved at the subtribal level. The cladogram as presented is one of several equally parsimoni- ous hypotheses. We suggest that the Ursiniinae and Cancri- niinae are plesiomorphic particularly in terms of the scale-like pappus structures as compared to other tribes. The genus Ursinia within the subtribe Ursiniinae has always been con- sidered to occupy an anomalous position in the Anthemid- eae. In our opinion, this is partly due to the presence of plesiomorphic features. Some of these, particularly basal arrangements of leaves and basal woodiness and the internal anatomy of cypselas in some members of the Ursiniinae and other relatively basal taxa such as the Cancriniinae, are remarkably similar to those found in members of the out- group which we interpret as examples of symplesiomorphy. Within the remaining group of ten subtribes it is reasonable to suggest that the predominantly Mediterranean and South African subtribes, such as the Achilleinae, Anthemidinae, Chrysantheminae, Leucantheminae, Thaminophyllinae and Matricariinae are closely related to each other because of shared possession of floral resin canals as compared to the mainly Asian Tanacetinae', Gonosperminae (African but apparently directly related to Tanacetum), Handeliinae and Artemisiinae. We admit that such a character is vague in circumscription but we offer this classification as a basis for further investigation. The Thaminophyllinae and Leucan- theminae may be closely related subtribes because of similari- ties in foliage characters. Generic classification At the generic level a number of changes should be noted. The status of particular genera as being monophyletic or non-monophyletic will be discussed briefly. We have adopted a conservative style and undertaken generic redelimitations only whenever necessary and wherever possible. Our aim has been to reclassify para- or polyphyletic genera and maximize the number of monophyletic genera within the Anthemideae. In many cases we have been content in discussing the non-monophyletic status of existing genera, to leave redelimi- tations of ambiguous taxa to future systematists. Many of the new and redefined genera have been analysed in more detail and considered by authors working in close connection with us (Kallersjo, 1986, 1988, 1991; Bremer & Kallersjo, 1986; Nordenstam, 1976; Ling, 1980a, b, 1991a, b; Wilcox, 1977). The definitions and dispositions of these new genera as determined by these authors include Inulanthera (Gonosperminae), Rhodanthemum (Leucantheminae), Oncosiphon, Myxopappus, Foveolina and Hilliardia (Matri- cariinae) and Seriphidium (Artemisiinae). Redefined genera include Phymaspermum, Hymenolepis, Athanasia (Ursinii- nae), and Pentzia and Rennera (Matricariinae). New genera described within this monograph include Chry- santhoglossum, Nivellea, Leucoglossum, and Rhodanthe- mum, taxa all within the Leucantheminae. Richteria (Cancriniinae) and Ismelia (Chrysantheminae) are two small genera originally described during the 19th century, later reduced into synonomy and then re-established here. Typi- cally misplaced species have been transferred to a variety of genera such as Microcephala and Aaronsohnia (Matricarii- nae). Also, it should be noted that taxonomic changes established here have dramatically improved the delimitation and definitions of well-known genera such as Artemisia, Chrysanthemum (s.s.), Leucanthemum and Tripleurosper- mum. By removing a number of species from these formerly non-monophyletic genera and assigning the misplaced species to new genera, the streamlined versions of the new taxa are now more likely to be monophyletic. The majority of genera within the Anthemideae are mono- phyletic taxa but there are several exceptions which will obviously require further taxonomic work, as indicated in further detail in the generic accounts. Of the relatively smaller taxa, genera without autapomorphies include Eumor- phia (Ursiniinae), Richteria (Cancriniinae), Tanacetopsis (Tanacetinae'), Gonospermum (Gonosperminae), Sphaer- omeria (Artemisiinae), Mecomischus and Chamaemelum (Achilleinae), Leucanthemopsis and Coleostephus (Leucan- theminae), and Cymbopappus and presumably Matricaria (Matricariinae). Future work should reveal whether they are monophyletic or non-monophyletic taxa. Amongst the larger genera, Achillea is one of similar uncertain stature. Tanacetum (Tanacetinae'), Dendranthema, Ajania and Artemisia (Artemisiinae), and Cotula (Matricariinae) are shown to be non-monophyletic. Other genera and even groups of genera have their sister groups within these taxa and eventually they will have to be split into smaller mono- phyletic units. In the next few years we envisage further changes of considerable magnitude which will necessitate 78 taxonomic and nomenclatural rearrangements quite different from those traditionally recognized. CHARACTERS OF THE ANTHEMIDEAE The characters used in the cladogram and for delimitation of genera are discussed below (see Table 2). The terms charac- ter and apomorphy are used synonymously, since any charac- ter is apomorphic at its universal level within the taxonomic hierarchy. The corresponding plesiomorphic condition is omitted from the character table. In obvious cases, such as perennial being considered as plesiomorphic as compared to annual, the perennial condition is not mentioned. In other characters the corresponding plesiomorphic conditions are explained elsewhere. The identification of characters and character states was undertaken by outgroup comparison with parts of the Asteraceae-Heliantheae. Character scoring There have been problems in scoring character states. For several character states, particular variables are expressed only within some species of a particular genus. Furthermore, the problem of unknown or inapplicable character states has caused conflicts during analysis. As far as this study is concerned each character may be scored in one of five different ways for a particular genus: 1: The character is present in all species of the genus. a: The character is polymorphic and present in only some but not necessarily all species of the genus. We take the line that all characters of this type are originally present within a genus but secondarily lost in some species. This interpreta- tion is based on comparison with the immediate relatives of the genus, identified during or after cladistic analysis. If the character is present in the sister taxa or close relatives, it is also considered originally present within the genus; 'a' stands for apomorphic. This method is simply outgroup comparison with the ingroup restricted to a particular genus being scored for the character. 0: The character is absent from all species of the genus. p: The character is absent from some but not necessarily most species of the genus. It is interpreted as originally absent within the genus; 'p' stands for plesiomorphic but indepen- dently derived in other species. The reason for this interpre- tation is similar to 'a' above. If the character is absent in the relatives, it is also considered originally absent within the genus. ?: The character is unknown or inapplicable to the genus. Characters relating to chemistry, embryology and chromo- some number are sporadic and unknown for many genera. Other characters are inapplicable to some genera. For example, characters of receptacular paleae and ray florets are inapplicable in epaleate and non-radiate genera, respectively. In the cladograms (Figs 1-12; see taxonomic text below) unknown or inapplicable characters are assumed to be present or absent following the principle of parsimony. If the character is known to be present in some genera of a particular group, it is also considered present in all genera of that group, and vice versa. K. BREMER AND C. J. HUMPHRIES Table 2 Characters used in the cladograms. 1 Plants annual. 2 Plants shrubby. 3 Plants spiny. 4 Plants compact and more or less scaphoid. 5 Plants with one or few sparsely branched stems arising from a woody villous caudex. 6 Plants rhizomatous with rosulate, spathulate-obovate-linear leaves. 7 Plants with branches in whorls below the first capitula. 8 Plants basally villous-tomentose with rather thick stems and a soft pith. 9 Plants covered with a dense greyish-white indumentum. 10 Plants covered with viscid hairs. 11 Plants with dolabriform hairs. 12 Plants with stellate hairs. 13 Plants with interxylary cork. 14 Leaves opposite. 15 Leaves variously deeply lobed or divided. 16 Leaves much pinnatisect with filiform lobes. 17 Leaves pectinate-pinnatisect with filiform, apically somewhat swollen and mucronulate lobes. 18 Leaves with few, oblong to rounded, apically mucronate lobes. 19 Leaves large with many rounded lobes. 20 Leaves spathulate in outline, ternate to ternately pinnate. 21 Leaves serrate-dentate. 22 Leaves entire or apically tridentate. 23 Leaves entire, ericoid. 24 Leaves rather fleshy, few-lobed or entire. 25 Leaves rather fleshy, entire, linear. 26 Leaves closely set, lanceolate to linear. 27 Leaves vermiform. 28 Leaves with secretory cavities. 29 Capitula densely corymbose. 30 Capitula very small and numerous in a large, dense, semiglobose corymb. 31 Capitula in a long narrow panicle or raceme. 32 Capitula in glomerules arranged in long spikes. 33 Capitula on short and nodding peduncles. 34 Capitula sessile along the stems. 35 Capitula discoid. 36 Capitula disciform. 37 Involucre rather narrowly urceolate. 38 Involucral bracts in 4-7 rows. 39 Involucral bracts in 1-2 rows, rather wide. 40 Involucral bracts in 2 unequal series. 41 Involucral bracts wide, flabelliform. 42 Involucral bracts subulate. 43 Involucral bracts with scarious margins. 44 Involucral bracts with dark brown margins. 45 Receptacle paleate. 46 Receptacle narrowly conical to subulate. 47 Receptacle hollow. 48 Receptacle pilose. 49 Receptacle densely hirsute. 50 Receptacular paleae pilose. 51 Floral parts with resin canals. 52 Ray floret limb white. 53 Ray floret limb golden yellow. 54 Ray floret limb bluish violet. 55 Ray floret limb deeply emarginate. 56 Ray floret limb epidermis cells tabular (senecioid or mutisioid type). 57 Ray floret tube sinus extending to the base. 58 Ray and disc floret tube dorsiventrally flattened. 59 Ray floret tube and cypsela pilose laterally; ray floret limb pilose abaxially. 60 Ray floret tube confluent with the cypsela. 61 Ray floret tube persistent on the cypsela. 62 Outer florets stalked. 63 Outer female florets in several rows. 64 Outer female florets subtended by scaphoid bracts. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 79 Table 2 cont 65 Outer female floret corollas 'flask-shaped', tapering above or narrowly cylindrical. 66 Outer female floret corollas without teeth. 67 Outer female florets without corollas. 68 Outer female floret style-branches lanceolate, flat, acute. 69 Corolla gradually expanded, rather thin and funnel-shaped. 70 Corolla inflated with a hollow space between outer surface and inner layer. 71 Corolla apically contracted. 72 Disc corolla 4-lobed. 73 Disc corolla lobes with dorsal appendages. 74 Corollas with continuous veins extending into the lobes. 75 Disc corolla lobes with central resin sacs. 76 Disc corolla zygomorphic with 2 smaller adaxial lobes with marginal resin canals extending from the base of the corolla and with 3 larger abaxial lobes. 77 Disc corolla red. 78 Corolla apically with erect, straight hairs. 79 Corolla apically with long, reddish hairs. 80 Corolla apically with stellate hairs. 81 Corolla cobwebby pilose. 82 Disc corolla tube thickened in fruit. 83 Disc corolla tube very thick and brittle. 84 Disc corolla tube basally saccate at least adaxially. 85 Disc corolla tube deeply and equally saccate both abaxially and adaxially. 86 Corolla tube basally copiously swollen and spongy, almost enclosing the cypsela especially laterally. 87 Disc corolla tube confluent with the cypsela. 88 Disc corolla tube pilose. 89 Corolla tube with long-stalked glands; stalk cells elongated. 90 Disc corolla tube and cypsela ribs with thick vascular strands. 91 Central florets of two kinds; outer perfect, inner completely sterile with reduced ovaries. 92 Anthers caudate. 93 Anthers with triangular-linear-lanceolate apical appendages, of rather thick-walled cells. 94 Anthers with an apical resin sac. 95 Anthers with endothecial tissue partly or wholly polarized. 96 Pollen grains with short or without spines. 97 Pollen grains without spines. 98 Pollen grains hexa-panto-colporate. 99 Style slender, parallel-sided at base. 100 Style immersed in a lobed nectary. 101 Style persistent and spinescent in fruit. 102 Style-branches brownish. 103 Disc floret style-branches long-penicillate. 104 Disc floret style-branches fused. 105 Stylopodium large and persistent in fruit. 106 Central floret ovaries reduced; florets functionally male. 107 Cypselas terete to weakly angled, or flattened. 108 Cypselas turbinate. 109 Cypselas arcuate. 110 Cypselas ellipsoid, small, c. 1 mm long. 1 1 1 Cypselas subglobose, with 2-3 very thin lateral-adaxial ribs. 1 12 Cypselas large, with 3 thick protruding sclerenchymatous ribs, somewhat winged in ray cypselas. 113 Cypselas slender and tuberculate with numerous obtuse excrescences. 114 Cypselas dorsiventrally flattened. 115 Ray cypselas dorsiventrally flattened with 3 adaxial ribs. 116 Disc cypselas laterally flattened. 117 Cypselas laterally winged. 1 18 Ray cypselas laterally winged; wings projected to apical teeth. 1 19 Cypselas with sclerenchymatic lateral wings. 120 Cypselas heteromorphic; ray cypselas triquetrous, winged; disc cypselas terete to prismatic to laterally flattened. 121 Disc cypselas abaxially and adaxially winged. 122 Cypsela wings as apical spines. 123 Cypselas with a mainly abaxial entire or toothed rim. 124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 125 Cypsela ribs basally fused into a more or less well developed foot callus. 126 Cypsela ribs protruding, narrow and somewhat wing-like. Table 2 cont 127 Cypselas with 5 mainly adaxially arranged ribs. 128 Cypselas with 1 abaxial and 2 lateral thick ribs and 2-3 adaxial ribs. 129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. 130 Cypselas with 2 lateral vascular strands, sometimes also with 1 adaxial strand. 131 Cypselas 10-ribbed with costal veins and resin canals. 132 Cypsela ribs with ellipsoid secretory cavities forming longitudinal ducts. 133 Cypselas with costal resin canals or sacs. 134 Cypselas with vallecular secretory canals. 135 Cypselas with vallecular vascular strands. 136 Cypselas abaxially and apically with 2 distinct, occasionally 1 or 3-5, resin sacs. 137 Cypselas with a single resin sac apically in the adaxial rib. 138 Cypselas with scattered elongated resin sacs. 139 Cypselas completely covered with rows of myxogenic cells. 140 Cypselas with myxogenic cells on abaxial surface and on the ribs of the adaxial surface. 141 Cypselas with dense rows of myxogenic cells also on the corona. 142 Cypselas with large myxogenic cells in rounded, scattered groups. 143 Cypselas with myxogenic cells in 2 distinct adaxial-lateral rows. 144 Cypselas with scattered, ovoid, myxogenic trichomes. 145 Cypselas papillose. 146 Cypselas long-papillose, seemingly pubescent. 147 Cypselas densely pilose; hairs subulate, with a few basal cells and one long apical cell. 148 Cypselas cobwebby pilose. 149 Cypselas with rather stiff unbranched hairs of 3-8 cells with spiral wall thickenings. 150 Cypselas copiously villous, seemingly covered in 'cotton wool'. 151 Cypsela wall without carbonized layer. 152 Cypsela wall several cell layers thick, partially or completely sclerified. 153 Cypsela thick-walled and conspicuously rugose. 154 Cypselas thin-walled, obovoid to oblanceolate, devoid of ribs. 155 Cypsela wall very thin, translucent and showing brownish black, rounded, very thick-walled testa epidermis cells. 156 Cypsela wall white and spongy; pericarpic cells isodiametric with thin spiral wall thickenings. 157 Cypsela wall with rod-shaped crystals in small packets. 158 Cypsela wall with numerous druses in the pericarp. 159 Cypsela wall with a continuous ring of fibre-like cells. 160 Cypsela wall with a continuous ring of sclerified isodiametric cells. 161 Pappus of short (not large and obovate or bristle-like) scales, an auricle, a corona, or absent. 162 Pappus of scales or teeth projected from the ribs. 163 Pappus of separate, mainly abaxial, subulate scales. 164 Pappus of 5 convolute-contorted scales. 165 Pappus a scarious, flimsy corona. 166 Pappus adaxially long. 167 Pappus a stiff adaxial auricle. 168 Pappus a large, scarious, adaxial but basally coroniform auricle, as long as the corolla or longer. 169 Pappus a large, scarious, adaxial, flabelliform auricle, as long as the corolla or longer. 170 Pappus scales brownish. 171 Pappus absent in disc cypselas, but present in ray cypselas. 172 Pappus absent in ray and disc cypselas. 173 Testa epidermis cells spirally arranged around the embryo. 174 Testa epidermis cells thick-walled and dark reddish. 175 Embryo sac tetrasporic. 176 Embryo sac disporic. 177 Chromosome number x=10. 178 'Irregular' monoterpenes in high concentrations present. 179 Furanosesquiterpenes present. 180 Particular thiophene derivatives present. 181 Amides present. 182 Flavonol 5-glucosides present. 183 Dehydrofalcarinone and dehydrofalcarinol present. 184 Anthocyanin present in root tips. 80 K. BREMER AND C. J. HUMPHRIES Habit The outgroup comprises perennial herbs or half-shrubs, as in many Anthemideae. Throughout the tribe there are scattered annual genera, groups of genera or odd annual species or groups of species within otherwise perennial genera. The annual habit (character 1) is independently derived several times within the Anthemideae. The same holds for the shrubby habit (2), although this condition is less common within the tribe. No species of the Anthemideae grows into a tree. Other uncommon habit characters absent in the out- group include characters 3-8. Indumentum The indumentum of the frequently pubescent Anthemideae commonly consists of unbranched hairs with a few basal stalk cells and a long apical cell. The same type is found in the outgroup and in many other Asteraceae. Occasionally in Anthemideae the hairs form a dense, tomentose to villous, greyish-white indumentum (character 9). In several Anthemideae dolabriform hairs are present (11) comprised of a stalk with few cells and transversely arranged apical cells, so that the hairs are T-shaped or Y-shaped. A distinction between T-shaped and Y-shaped hairs was made by Napp- Zinn & Eble (1980) but we find it difficult to maintain this distinction. Both types may be present, sometimes mixed together with intermediates. Dolabriform hairs are absent from the outgroup. In a few genera stellate hairs are present (12), possibly derived from the dolabriform hairs. The stellate hairs also have a stalk of a few cells, but a stellate apical cell. The type is absent from the outgroup. Glands composed of two parallel rows of cells with the apical pair enlarged are frequent in the Anthemideae, the outgroup and other Asteraceae. The number of cell pairs in the stalk varies, and at first sight there seem to be two types: one with several cells forming a stalk and one with a basal cell pair and an apical enlarged pair forming a sessile gland (cf. for example, Nordenstam, 1976). There are intermediates with two or more basal cell pairs, and hence we find it difficult to separate the two types into separate character states. Thus, we have avoided using any of them as a character. Occasionally, the viscid glands form a dense cover on the plant (10). Wood anatomy In Anthemideae-Artemisiinae a number of genera are pro- vided with interxylary cork (character 13; Holmgren et al., 1976; Moss, 1940). Many genera have not been investigated. As far as we know it does not occur in the outgroup. Foliage In the majority of the Heliantheae the leaves are opposite. The outgroup of the Anthemideae within Heliantheae and the majority of Anthemideae have alternate leaves. This condition then seems best interpreted as a synapomorphy for the Anthemideae and part of Heliantheae, whereas the secondarily opposite leaf arrangement (character 14) is a character within the Anthemideae. Variously dissected leaves are characteristic of most Anthemideae and entire leaves are rather rare exceptions. Dissected leaves also characterize the outgroup, and appear to be a synapomorphy (character 15) comprising the Anthemideae and the outgroup. Entire leaves then represent a reversal in this character. Entire and variously characteristic leaves have been scored as independent characters, particu- larly 22-26. The mode of leaf dissection is highly variable and difficult to apply at the generic level. Various types may be present within individual genera. We have designated only a few different types as characters which are absent in the outgroup, particularly 16-21. In Ursiniinae a number of genera have leaves with secre- tory cavities, present in floral structures but not in the leaves of many other Anthemideae. Secretory cavities are absent in the outgroup. Inflorescence The capitula (inflorescences) are usually terminal either singly at the end of a peduncle or aggregated into a variety of corymbose, spike-like or clustered synflorescences. Some- times the peduncles are structurally reduced so that the capitula are aggregated into central sessile clusters (an aggre- gated synflorescence) or, on rare occasions, into a single head (a syncephalum). The capitula are solitary or arranged in lax corymbs in the outgroup and commonly in the Anthemideae. They are also frequently arranged in dense corymbs within certain genera (character 29, further modified as 30). Rarely, the capitula are sessile (34) or solitary on nodding peduncles (33). Within the Artemisiinae the corymbose capitulum arrangement is further modified into a long narrow panicle- like or raceme-like synflorescence (31) generally with numer- ous capitula. The paniculate-like capitulum arrangement is variable and apparently contains a number of different types. The interpretation of these requires a detailed study, and we have only listed one distinct type as a character, the arrange- ment of the capitula in glomerules on long spikes (32). Floral and sex arrangement The array of sex expressions and floral morphology in most Anthemideae were surveyed by Heywood & Humphries (1977). Radiate, heterogamous capitula with hermaphroditic, perfect disc florets and female ray florets are the plesiomor- phic condition, being present in the outgroup. Sometimes the ray florets are neutral but we found this condition difficult to apply as a character at the generic level. The reduction of the ray florets resulting in discoid, homogamous capitula is common, and has occurred several times within genera. This character (35) has often been used for delimitation of genera, despite its homoplasious nature. Disciform, heterogamous capitula (36) with central, her- maphroditic florets and outer, female, non-radiate florets occur in several Anthemideae. It is possible to hypothesize two interpretations for discoid and disciform florets. Outer tubular female florets may either be modified central her- maphroditic florets or modified female ray florets. The first hypothesis implies that disciform capitula are derived modifi- cations of discoid capitula. The second hypothesis implies that disciform and discoid capitula may be independently derived from radiate capitula but it is possible also that discoid capitula are derived from disciform capitula by reduc- tion of the outer female florets. For analysis it seemed best to consider discoid and disciform capitula as independent non- additive characters. After construction of the cladogram, a particular interpretation may be considered the most parsi- GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 81 monious, as in the case of the Artemisiinae where discoid capitula are considered derived from disciform capitula. In other subtribes discoid capitula are best interpreted as derived directly from radiate capitula. It was eventually seen that there was no support for the hypothesis that disciform capitula are derived from discoid capitula in any group of Anthemideae. Within Artemisiinae the central florets may have reduced fertility. In Neopallasia the central florets are heteromorphic. The florets towards the periphery are perfect and the central ones completely sterile with reduced ovaries (91). In the Dracunculus group of the genus Artemisia, central floret ovaries are reduced and the florets are functionally male (106). Involucre Involucral shape is variable from widely campanulate to narrowly urceolate. In the outgroup the narrow types are absent, so that narrowly urceolate involucres are derived characters (37) within the Anthemideae. The involucral bracts are arranged in several imbricate series as in the outgroup and many other Asteraceae. Rarely there are 4-7 rows (38), more than in the outgroup, or 1-2 rows (39). In one genus (Eriocephalus) the involucral bracts are arranged in two distinctly unequal rows without intermediates (40). In almost all Anthemideae as well as in the outgroup the involucral bracts have scarious margins, otherwise they are generally absent in the Heliantheae. The chaffy bract is one of the main characters (43) which we used to identify the outgroup of the Anthemideae. Sometimes the involucral bracts are wide and flabelliform (41), or have dark brown margins (44), or they are subulate (42), all characters absent in the outgroup. Receptacle The receptacle is either paleate or epaleate. Within the Anthemideae the paleate receptacle is plesiomorphic. For- merly the tribe Helenieae was distinguished from Heliantheae by having epaleate receptacles rather than being paleate as in the Heliantheae s.s. Furthermore, the Anthemideae was divided into two subtribes based on recep- tacular paleae, the paleate Anthemidinae s.l. and the epale- ate Chrysantheminae s.l. The homoplasious nature of this character has since long been recognized and the distinction of the Helenieae and the two subtribes of Anthemideae has now been abandoned (see Bremer, 1987). It seems possible that the epaleate receptacle is a character for a portion of the Heliantheae (s.l.) and Anthemideae. The outgroup of the Anthemideae has epaleate receptacles. Within Anthemideae the presence of a paleate receptacle is hence considered a character (45) despite the fact that it might be under single gene control within certain genera (see Mitsuoka & Ehrendorfer, 1972). The shape of the receptacle varies from being flat to convex to conical and ultimately subulate. The latter conditions are absent from the outgroup. Receptacle shape is a potential source of more character information but we have found it difficult to interpret at this stage. It is used as a character (46) for a few genera. Rarely the receptacle is hollow (47), pilose (48) or densely hirsute (49), characters absent in the out- group. The hairs are scattered over the receptacle and similar to hairs on other parts, so that they cannot be hypothesized as being modified paleae. Occasionally, the paleae are also pilose (50) and thus the two characters cannot be considered as homologous (failure of the conjunction test, Patterson, 1982; Humphries & Funk, 1984). Resin canals In many genera of the Anthemideae floral parts are fre- quently invested with resin canals (character 51). They occur in a variety of forms, in the corolla lobes, in style-branches, in cypselas, sometimes in anther tips and often also in the receptacular paleae and involucral bracts. We admit that this generalized character is rather vaguely formulated but offer it as one hypothesis for suggesting the interrelationships of the various subtribes (see discussion below). Resin canals are absent in the outgroup. Ray florets The ray corolla limb is yellow in the outgroup and mostly white (character 52) but sometimes yellow in Anthemideae. Glossy golden yellow limbs (53) occur in Leucantheminae and bluish violet limbs (54) in one genus of the Cancriniinae (Allardia), although other colours do occur including pink, orange, and red. Occasionally, the limb is deeply emarginate (55). Baag0e (1977) distinguished a number of epidermal cell types on the ray floret limbs. Within the Anthemideae she recognized two groups, with Helianthoid and Senecionoid/ Mutisioid epidermal cells (56). The former type occurs in the outgroup. The ray floret tube furnishes a number of rather rare characters not present in the outgroup. The tube is sometimes dorsiventrally flattened (58), laterally pilose (also abaxially on the limb; 59), confluent with the cypsela (60), and persistent on the cypsela (61). Occasionally the ray floret tube sinus extends to the base (57), so that logically the tube is absent. Outer florets in disciform capitula Outer female florets and disciform capitula are not present in the outgroup. The characters of the outer female florets listed here mostly represent unique specializations within Arte- misiinae where the plesiomorphic and disciform genus Ajania serves as an outgroup. Within the Artemisiinae the outer female florets are occasionally subtended by scaphoid bracts (character 64), often their corollas are 'flask-shaped' and tapering above to narrowly cylindrical (65), and rarely the corollas are without teeth (66) or are totally absent (67). In disciform Anthemideae the outer female florets are sessile and arranged in one row. Rarely in the Matricariinae (Cotula group of genera) they are stalked and arranged in several rows (Solivd), both very peculiar conditions within the family as a whole. The pluriseriate arrangement is com- mon in other tribes, but is most parsimoniously interpreted as an independent character within the Anthemideae. Disc florets The corolla of the disc florets provides several characters. In the outgroup and in many Anthemideae the plesiomorphic condition is seen: yellow, glabrous and actinomorphic, with a more or less distinct, unswollen and non-saccate tube, and a 5-lobed limb without veins or resin sacs in the lobes. Within the Ursiniinae the corolla is often gradually 82 K. BREMER AND C. J. HUMPHRIES expanded and funnel-shaped without distinct tubes and limbs (character 69). In Sclerorhachis the limb is contracted at the apex (71). In some genera the corolla is 4-lobed (72). The lobes may be provided with dorsal appendages (73), continu- ous veins along the margins (74), with central resin sacs (75) or rarely (Adenoglossa) with marginal resin canals in two of the lobes which are smaller than the other three (76). Rarely the corolla is red (77) or provided with various types of hairs (78-81, 88, 89). The corolla tube is sometimes thickened in fruit (82), and in one extreme case (Oncosiphon) very thick and brittle (83). It is sometimes saccate basally, adaxially (84) or both abaxi- ally and adaxially (85). In an extreme case (Otanthus) the tube is copiously swollen at the base and spongy, and almost enclosing the cypsela (86). Rarely, the tube may be confluent with the cypsela (87, cf. character 60). Other unusual corolla types are those with long-stalked glands as found in the Ursiniinae (89), and those with very thick vascular strands (90). Anthers In the outgroup and in most Anthemideae the anthers are not caudate at the base. This was formerly considered an impor- tant character for defining the tribe but anther tails (character 92) are present in a few genera. The apical anther appendages are obtuse to rounded in the outgroup and in most Anthemid- eae. In the Artemisiinae they are triangular-linear-lanceolate and composed of rather thick-walled cells (93). Sometimes the appendages have an apical resin sac (94). The endothecial tissue in the outgroup and many Anthemideae and other Asteraceae is 'non-polarized', i.e. with the endothecial cell wall thickenings arranged evenly. However, in some Anthemideae (and other Asteraceae) the endothecial tissue is 'polarized' (character 95), with the thickenings arranged apically and basally on the cell (Dormer, 1962). Pollen Spiny pollen is the plesiomorphic condition of the Anthemid- eae occurring also in the outgroup. Within the Artemisiinae there is a gradual reduction of the spines (characters 96 and 97) until a rugose appearance is observed on the exine surface. Such a condition is considered a modification associ- ated with wind-pollination (Wodehouse, 1938). In Adenan- thellum, the pollen is hexa-panto-colpororate (98) rather than tricolporate as found generally within the Asteraceae. Styles The disc floret style-branches are of the common so-called Senecionoid type: apically penicillate, truncate with parallel stigmatic surfaces. In Prolongoa the apical hairs are rather long (character 103), and Phaeostigma is named after its brownish style-branches (102). In Mausolea (Artemisiinae) the style-branches are not linear as in other disciform Anthemideae but lanceolate, flat and acute (68). Sometimes the style-branches are fused (104) and the style is undivided, a condition associated with functionally male florets. The style is somewhat bulbous and often situated on a more or less developed stylopodium, as in the outgroup and many Asteraceae. Within Artemisiinae the style base may be slender and parallel-sided (99). Occasionally the stylopodium is large and persistent in fruit (105) and in Heliocauta it is modified into a lobed nectary (100). In Sollva the style is persistent and spinescent in the mature fruit (101). Cypselas The mature and developing cypselas provide a fruitful supply of characters within the Anthemideae (Bruhl & Quinn, 1990; Giroux, 1933; Humphries, 1976, 1977; Khandzhyan, 1983; Kneisl, 1981; Kynclova, 1970; Wilcox, 1977). In the outgroup the cypselas are sharply angled and provided with a heavily carbonized layer. This layer is never present in the Anthemideae but it is widespread throughout the Heliantheae. The reduction of the carbonized layer can be interpreted as a synapomorphy (character 151) for the Anthemideae as a whole. The cypselas of Anthemideae can also be terete to weakly angled, or flattened (107) and modified in various ways. The characters 108-113 relate to different cypsela shapes unique to the ingroup. The cypselas are sometimes much flattened, dorsiventrally (characters 114 and 115) or laterally (116), and the flattened cypselas are often winged (117-122). Cypselas without a pappus are apically rounded or truncate as in the outgroup, or sometimes provided with an apical rim. It is difficult to say whether this rim is homologous with a short coroniform pappus or is an independent outgrowth of the cypsela wall. We have avoided coding it as a character. In Oncosiphon the cypsela rim is distinctly abaxial (123) and seems to be a unique structure. The presence, number and arrangement of cypsela ribs is very variable. Cypselas with five or four evenly arranged ribs are common and may be plesiomorphic within the tribe. The cypsela ribs are difficult to interpret since similar ribs are absent from the outgroup. Characters 124-129 represent specialized rib structures within Anthemideae. The cypsela ribs are sometimes furnished with resin canals or sacs (131-133, 136, 137), not present in the outgroup. Rarely resin sacs are scattered over the cypsela (138). Within the Leucan- theminae, the Leucanthemum group is characterized by a unique cypsela type with vallecular resin canals (134) of a particular flattened type and vallecular vascular strands (135), situated in the valleys between the ribs, not inside the ribs (costal) as in other Anthemideae (Briquet, 1916; Briquet & Cavillier, 1916; Giroux, 1933). Frequently the cypselas are furnished with myxogenic cells within the Anthemideae but the character never occurs in the outgroup. However, it is difficult to apply the presence of myxogenic cells as a simple presence or absence character as it is homoplasious within the tribe as a whole. The myxogenic cells are commonly situated along the ribs and this appears to be the plesiomorphic arrangement. Other particular distribu- tions of the myxogenic cells may serve as characters (139-141 , 143). Characters 142 and 144 represent unique myxogenic cell shapes, quite different from the common type which consists of transversely compressed cells in elongated rows. The cypselas of the Anthemideae are generally smooth and glabrous or sometimes covered with scattered glands. Gla- brous and sparsely to rather densely hirsute cypselas are found in the outgroup. Papillose cypselas (145) and cypselas with other types of indumentum not present in the outgroup are those represented by characters 146-150. The cypsela wall is sometimes several cell layers thick and partially or completely sclerified (152). In other cases it is very thin (characters 154 and 155). The intermediate condi- tion is common and present in the outgroup. The anatomy of GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE the cypsela wall is a potential source of more character information but detailed investigations of the appropriate genera are necessary. The investigations by Reitbrecht (1974) need to be expanded and performed in more detail (see Bruhl & Quinn, 1990 on the 'Cotuleae'). Characters 156, 159 and 160 represent unique specializations within the Anthemideae (Kallersjo, 1986, 1988). The cells of the cypsela wall usually contain isodiametric druses as in many Asteraceae. Rarely they are numerous (158) and in Prolonged they are replaced by rod-shaped crystals in small packets (157). Pappus In the Anthemideae the cypselas are generally without a pappus or provided with a coroniform or scaly pappus. True bristles, which are never flattened and occur in other tribes, are never present. Such bristles are absent also from the outgroup, where the pappus, if present, consists of large whitish scales or numerous whitish, long, flat bristles (more properly bristle-like linear scales). Similar pappus types are present in Ursiniinae (Ursinia) and Cancriniinae and we assume that the presence of large or long-whitish pappus scales are plesiomorphic within the tribe. The five convolute- contorted pappus scales (character 164) of Ursinia seem to be unique for that genus, although the large size and the whitish colour appear to be plesiomorphic. A reduced or lost pappus is represented by characters 161, 171 and 172 within the Anthemideae. The coroniform pappus is either derived from fusion of the individual pappus scales or is an independent character unique to the Anthemideae. If we adopt the latter hypothesis, logically the coroniform pappus seems to be a plesiomorphic condition within the Anthemideae, excluding Ursiniinae and Cancriniinae, whereas specialized corona types and auricles are apomorphic, particularly as seen with characters 162, 163 and 165-170. 83 number of x=7, 8, and Artemisia a series based on x=5, 6, 7, 8 and 9. The highest values have been recorded in Leptinella with 2n=312 (Lloyd, 1972a). Most series and most variants are aneuploid or polyploid series within genera and variation is of little use for classification at the generic level. However, it appears that the base number of x=10 (character 177) as opposed to x=9 is plesiomorphic to a number of genera in the Thaminophyllinae and Matricariinae (Cotula group). Chemistry Chemical characters are difficult to utilize because they are mostly collected on a sporadic basis for one or two species within a genus. Furthermore, it is rare to find phytochemical papers that report absence traits as well as presence of a chemical. However, the Anthemideae are well-known for accumulating herbal and insecticidal chemicals and relative to many angiosperms have been richly studied (Greger, 1975, 1977; Bohlmann et al., 1973). For systematic purposes avail- able chemical evidence suggests that sesquiterpene lactones (character 179), flavonoids (182), particularly C-glycosyl fla- vones (Valant-Vetschera, 1981, 1982, 1985) and polyacety- lenes (183) are of most systematic value (Greger, 1977; Tetenyi, 1986). Presence of a chemical character has been scored at generic rank even though it is acknowledged that investigation has been sporadic, particularly with the larger genera, such as Artemisia. The scores thus represent hypothetical presence for all the species. Different compounds within the same class of compounds have been combined into a single score for polyacetylenes (183) and furanosesquiterpenes (179) when the class compound is considered apomorphic for a particular group. Red coloured root tips, due to the presence of anthocyanin (184) has been recorded in Leucanthemum (Favarger, 1966). Testa Testa epidermal types were investigated by Kneisl (1981), but the results are difficult to interpret. The testa epidermis cells are normally well developed with sinuose cells. Sometimes the testa collapses during maturation of the fruit. Rarely the testa epidermis cells are uniquely specialized, much elon- gated and spirally arranged around the embryo (character 173) or thick-walled and dark reddish (174). Embryology The Anthemideae have been fairly thoroughly investigated compared to many other tribes, by Hading (1950, 1951, 1960). Tetrasporic embryo sacs (character 175) unambigu- ously characterize a number of genera, and disporic embryo sacs (176) are reported as unique within Argyranthemum (Borgen, 1972). Harling also described development of the embryo sacs in detail but the variation is difficult to formulate into characters. Chromosome numbers The Anthemideae have been investigated based on the surveys of Federov (1969), Moore (1972, 1973, 1977), and Goldblatt (1980, 1981, 1984). X=9 is the base number for the tribe. Most genera have either a diploid number of 2n=18 or 2n=36. Cancrinia has a base number of x=7, Ursinia a base DISTRIBUTION The approximate total range of the Anthemideae is summa- rized in Table 3 and the general distribution of subtribes and genera given in Tables 3, 5, 7, 9, 11, 14, 16, 18, 20, 22 and 24; see taxonomic text below. The Anthemideae tribe has a worldwide distribution but with main concentrations of taxa in Central Asia, the Mediterranean region and South Africa. Some members of the subtribes Ursiniinae, Artemisiinae, Chrysantheminae, Leucantheminae, Anthemidinae, and Matricariinae are pernicious weeds, such as some of the Ursinia species introduced to Australia and New Zealand, and species of Chrysanthemum, Anthemis, Artemisia, Achil- lea, Leucanthemum, Tripleurospermum and Matricaria as more widespread weeds in both the northern and southern hemispheres. However, most taxa have discrete ranges and very obvious areas of endemism. The Thaminophyllinae, for example, occur only in South Africa. Similarly most Ursinii- nae are almost entirely restricted to South Africa, although one species of Ursinia occurs in Ethiopia and another species of Lasiospermum is found in the Sinai region of Egypt. The Gonosperminae is a small subtribe of three genera and 15 species which displays a presumably relictual distribution pattern. Gonospermum and Lugoa are endemic to the Canary Islands but have a curious disjunct distribution pat- tern with their sister genus, Inulanthera, which has ten species 84 K. BREMER AND C. J. HUMPHRIES Table 3 Summary of geography and natural distribution of the Anthemideae. N.Am. Number of genera 8 Ursiniinae Cancriniinae Tanacetinae 1 Gonosperminae Handeliinae Artemisiinae 4 Achilleinae Anthemidinae Chrysantheminae Leucantheminae Thaminophyllinae Matricariinae 3 Eur- Asia 25 C.& E. SW Asia Asia 38 13 6 6 4 15 2 S.Eur 28 2 5 1 2 12 N.Afr 38 1 2 1 7 1 4 11 10 S.Afr 29 8 1 1 5 14 Austr. S.Am. N.Zeal. 3 4 in southern Africa, mainly in Natal, but with one species also Capitula pedunculate 5 5 Pappus of 5 (rarely 8-10) large obovate scales and sometimes 5 The majority of the remainder of the tribe occur in the additional subulate scales (pappus rarely absent in Ursinia northern hemisphere. For example, the Chrysantheminae are trifida, a South African shrub with linear, entire or apically most prominent in North Africa and Macaronesia , although few-lobed leaves) 1 . Ursinia the two well-known species of Chrysanthemum, C. cow- Pappus a shallow corona, an auricle or rarely of few small narium and C. segetum are widespread throughout the north- scales, or absent 6 ern hemisphere Indeed C. coronarium is cultivated widely 6 Disc s 5 _ ribbed ^^ flat and as a salad vegetable in China. By contrast, the Leucanthemi- sterile with a few pappus scales; , eaves sermte nae are found in Eurasia and North Africa, and are particu- 53 Lepidophorum lady concentrated in the Mediterranean region. The Cypselas equal, without or with a coroniform or auriculiform Achilleinae is more of a Eurasian group with some of the pappus; leaves various 7 more distinctive taxa endemic to North Africa, southern Europe, the Mediterranean, and South-West Asia, although 7 Cypselas obovoid wuhout distinct ribs or wings 8 ... XT IL A T-L Cypselas angled or ribbed or prismatic, often turbmate, some- there are some outlying taxa oocunng in North America. The ^ tuber( f ulate; or las J attened and with 2 latera , ribs or Tanacetinae are well-represented in Eurasia and especially wings 9 in central Asia but there are some interesting endemics which occur also in North America and North Africa. The Cancri- 8 Leaves few-lobed or entire 53. Mecomischus niinae and Handeliinae are comprised of eleven genera and Leaves P innatifid to pinnatisect 54. Chamaemelum thirty-four species restricted entirely to the central steppes of 9 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad Asia. The Artemisiinae and Matricariinae are the most wings 10 widespread subtribes. The Artemisiinae which comprise Cypselas various, sometimes dorsiventrally flattened but then about a third of the tribe, with 18 genera and more than six rhombic in cross-section and not or only narrowly winged hundred species currently recognized, have a worldwide 57. Anthemis distribution but occur mainly in the northern hemisphere and , n /- u i r j . ... A . _ m , . ." 10 Cypselas with 2 lateral more or less distinct ribs but no especially central and eastern Asia. The Matricariinae, com- wings 50 Achillea prised of 25 genera and about 250 species, also has a Cypselas with 2 lateral wings ........................ 51. Anacyclus worldwide distribution but most genera occur in the Mediter- ranean region and South Africa. n Pa PP us of 5 (rarely 8-10) large obovate scales and sometimes 5 additional subulate scales 1 . Ursinia Pappus a shallow corona, an auricle, or of small scales, or absent 12 KEY TO GENERA 12 Cypselas copiously villous 2. Lasiospermum Cypselas glabrous, sometimes glandular or papillose 13 1 Receptacle distinctly paleate with paleae subtending florets (in Anthemis rarely with the basal part of the conical receptacle 13 Involucral bracts in 2 unequal series; one outer row of pubes- epaleate) .2 cent to glabrescent bracts and one inner row of generally Receptacle completely epaleate (though sometimes pilose or densely villous bracts 108. Eriocephalus hirsute) or capitula few-flowered and presence of paleae Involucral bracts imbricate and subequal, not in 2 unequal unclear . .. 41 series 14 Capitula radiate; rays present 3 Capitula discoid or disciform; rays absent 22 Rays yellow, rarely abaxially reddish 4 Rays white or rarely pink to reddish on both sides 11 14 Shrubs with opposite leaves (leaves alternate in Eumorphia davyi, a South African shrub with linear, closely set leaves); cypselas with 10 or more ribs 4. Eumorphia Herbs or rarely shrubs with alternate leaves (rarely with some leaves opposite basally on the stems); cypselas various 15 4 Capitula sessile 56. Cladanthus 15 Suffruticose perennial with large leaves with rounded lobes and GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 85 corymbose capitula; cypselas 5-ribbed with pappus of small scales projected from the ribs 22. Lugoa Habit and cypselas various but leaves not large and with rounded lobes and pappus if present not of scales projected mainly from the ribs 16 1 6 Cypselas obovoid , without distinct ribs or wings 17 Cypselas angled or ribbed or prismatic, often turbinate, some- times tuberculate; or cypselas flattened and with 2 lateral ribs or wings 18 17 Leaves few-lobed or entire 53. Mecomischus Leaves pinnatifid to pinnatisect 54. Chamaemelum 18 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad wings 19 Cypselas various, sometimes dorsiventrally flattened but then rhombic in cross-section and without 2 lateral ribs and not or only narrowly winged 21 19 Cypselas with 2 lateral more or less distinct ribs but no wings 50. Achillea Cypselas with 2 lateral wings 20 20 Leaves vermiform; disc corolla tube deeply and equally saccate both adaxially and abaxially 52 . Leucocyclus Leaves pinnatisect; disc corolla tube adaxially slightly sac- cate 51. Anacyclus 21 Leaves serrate-dentate, rarely pinnatifid or entire; anthers caudate 79. Osmitopsis Leaves pinnatisect to variously lobed; anthers not caudate 57 . Anthemis 22 Pappus present, of scales, an auricle or a corona 23 Pappus absent, but cypselas sometimes apically with an obtuse rim (rarely with a pseudopappus of bristle-like stalked glands in Athanasia) 28 23 Pappus of 5 (rarely 8-10) large obovate scales and sometimes 5 additional subulate scales 1 . Ursinia Pappus a shallow corona, an auricle or of small scales 24 24 Capitula solitary or laxly corymbose 57 . Anthemis Capitula densely corymbose 25 25 Glabrous annual herb; cypselas with 1 adaxial and 2 lateral ribs and with a single secretory cavity apically in the adaxial rib 97. Lonas More or less pubescent shrubs; cypselas 5-10-ribbed with or without several secretory cavities in the ribs 26 26 Capitula narrowly oblong-obconical, slender, and few-flowered; indumentum of stellate hairs 6. Hymenolepis Capitula rather widely urceolate to cyathiform-campanulate; indumentum of simple or bifid hairs 27 27 Cypselas 5-ribbed; anthers not caudate 23. Gonospermum Cypselas 8-10-ribbed; anthers caudate 24. Inulanthera 28 Cypselas copiously villous 2 . Lasiospermum Cypselas glabrous, sometimes glandular 29 29 Involucral bracts in 2 unequal series; one outer row of pubes- cent to glabrescent bracts and one inner row of generally densely villous bracts 108. Eriocephalus Involucral bracts imbricate and subequal, not in 2 unequal series 30 30 Leaves entire or crenate, covered with a dense greyish-white indumentum; corolla tube basally copiously swollen and spongy , almost enclosing the cypsela especially laterally 49. Otanthus Leaves various; corolla tube not or somewhat swollen and spongy but not enclosing the cypsela 31 3 1 Cypselas obovoid without distinct ribs or wings 32 Cypselas angled or ribbed or prismatic, often turbinate, some- times tuberculate; or cypselas flattened and with 2 lateral ribs or wings 34 32 Capitula numerous in a long panicle 41 . Seriphidium Capitula solitary or laxly corymbose or few closely together 33 33 Capitula almost sessile along the stems 55. Rhetinolepis Capitula pedunculate 54. Chamaemelum 34 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad wings 35 Cypselas various, sometimes dorsiventrally flattened but then rhombic in cross-section and not or only narrowly winged . 37 35 Cypselas with 2 lateral more or less distinct ribs but no wings 36 Cypselas with 2 lateral wings 51. Anacyclus 36 Leaves mainly rosulate and capitula solitary; cypselas actually 4-5-ribbed though with 2 major lateral ribs 21. Heliocauta Leaves alternate and capitula often corymbose, rarely solitary; cypselas with 2 lateral ribs only 50. Achillea 37 Shrublets; corolla basally saccate around the cypsela especially adaxially 48. Santolina Herbs or shrubs ; corolla not saccate basally 38 38 Shrubs with stellate hairs if present; corolla gradually expanded and funnel-shaped 7. Athanasia Herbs or half-shrubs with simple or bifid hairs; corolla more or less distinctly divided into tube and limb 39 39 Basally villous perennials with much pinnatisect, mainly basal leaves; cypselas 4-5-ribbed 40 Indumentum various but not mainly basal and villous; annuals or perennials; leaves and cypselas various 57. Anthemis 40 Stems few-branched , leafy and with terminal corymbs 28. Handelia Stems loosely branched, almost leaf-less with terminal capitula 29. Sclerorhachis 41 Capitula radiate; rays present 42 Capitula discoid or disciform ; rays absent 109 42 Rays yellow, rarely partly white or reddish or abaxially red- dish 43 Rays white, rarely pink to reddish, bluish violet or creamy orange, but not yellow 62 43 Pappus present in ray or disc cypselas or generally in all cypselas, of scales, an auricle or a corona (sometimes almost absent in Tanacetum but cypselas then apically with an acute rim) 44 Pappus absent but cypselas sometimes apically with an obtuse rim (in Argyranthemum sometimes with pappus-like apically projected cypsela wings) 54 44 Cypselas densely pilose; pappus of 4-12 white scales at least half as long as the corolla 9. Trichanthemis Cypselas glabrous, sometimes glandular; pappus of short scales, a corona or an auricle 45 45 Cypselas dorsiventrally flattened and with 2 lateral broad wings; an annual herb with entire, fleshy leaves .... 101. Adenoglossa Cypselas various, sometimes dorsiventrally flattened but then not or only narrowly winged and often rhombic in cross- section 46 46 Disc corolla 4-lobed; leaves entire 81. Inezia Disc corolla 5-lobed ; leaves various 47 47 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with 86 K. BREMER AND C. J. HUMPHRIES or without 2 weaker ribs; pappus coroniform if present 48 Cypselas various, generally with 5 or more ribs and not with 3 major adaxial-lateral ribs; pappus and habit various 49 48 Leaves pinnatifid-pectinate; cypselas with 3 adaxial-lateral thick ribs and 2 abaxial weaker ribs; pappus absent in disc cypselas but present in ray cypselas 70. Prolongoa Leaves pinnatisect; cypselas with 3 adaxial-lateral acute ribs; pappus present in both disc and ray cypselas . 94. Endopappus 49 Cypselas 8-12-ribbed with dark vallecular secretory canals between the pale ribs; annuals 50 Cypselas angled, prismatic or ribbed, but vallecular secretory canals absent; annuals or perennials 53 50 Pappus a scarious, adaxial, flabelliform auricle, as long as the corolla or longer 76. Glossopappus Pappus a corona, a short auricle, or absent 51 51 Leaves lobed, often trifurcate; disc cypselas with a stiff coroni- form pappus 75. Chrysanthoglossum Leaves serrate-dentate; disc cypselas without or with a scarious, coroniform or auriculiform pappus 52 52 Rays golden yellow; cypselas arcuate with the ribs basally and adaxially fused into a more or less distinct callus 77. Coleostephus Rays pale yellow; cypselas ellipsoid without a basal callus 73. Leucoglossum 53 Cypselas with myxogenic cells on the ribs; pappus a scarious flimsy corona; creeping or caespitose suffruticose perenni- als 68. Leucanthemopsis Cypselas without myxogenic cells; habit and pappus vari- ous 15. Tanacetum 54 Ray cypselas triquetrous, winged; disc cypselas laterally flat- tened and adaxially and abaxially winged or sometimes terete to prismatic 55 Cypselas equal, oblong or obovoid, without wings, or dorsiven- trally flattened and laterally winged 58 55 Shrubs or shrublets 62. Argyranthemum Annual herbs 56 56 Plant covered with viscid hairs; cypsela wings projected to apical spines 60. Heteranthemis Plants not viscid; cypsela wings if present not projected to apical spines 57 57 Disc corolla red; disc cypselas laterally flattened and adaxially and abaxially winged 61. Ismelia Disc corolla yellow; disc cypselas prismatic with a narrow adaxial wing or terete and apparently ribbed 59. Chrysanthemum 58 Disc corolla 4-lobed; rays not true ray florets but outer disc florets with one corolla lobe expanded to a limb; cypselas dorsiventrally flattened 103. Cotula Disc corolla 5-lobed; rays true ray florets with apically 3-lobed limbs; cypselas oblong or obovoid 59 59 Cypselas distinctly ribbed with dark vallecular secretory canals between the pale ribs 71. Leucanthemum Cypselas distinctly or faintly ribbed but vallecular secretory canals absent 60 60 Basally much woody shrublets with few-lobed leaves with linear lobes 30. Brachanthemum Herbs or half-shrubs; leaves generally with rather broad lobes 61 61 An annual herb with few-lobed leaves 33. Tridactylina Perennials; leaves various 31. Dendranthema 62 Pappus present at least in ray cypselas, of scales (bristle-like in Allardia), an auricle or a corona (sometimes almost absent in Tanacetum but cypselas then apically with an acute rim) ... 63 Pappus absent but cypselas sometimes apically with an obtuse rim (mainly abaxial and somewhat toothed in Oncosiphon; cypselas in Argyranthemum sometimes with pappus-like apically projected wings) 88 63 Pappus of 4-12 linear to obovate-oblong scales at least half as long as the corolla, or of many bristle-like scales as long as the corolla or longer 64 Pappus of short scales, an auricle or a corona 66 64 Pappus of 4-12 scales half to equalling the corolla in length; rays white or sometimes pink 65 Pappus of many bristle-like scales equalling or exceeding the corolla in length; rays often bluish-violet, sometimes white or pink 12. Allardia 65 Cypselas densely pilose ; pappus scales white 9. Trichanthemis Cypselas glabrous, glandular; pappus scales brownish 11. Richteria 66 Cypselas 8-12-ribbed with dark vallecular secretory canals between the pale ribs 67 Cypselas angled, prismatic or often ribbed, but vallecular secretory canals absent 69 67 Annuals; cypselas c. 1 mm long 73. Leucoglossum Perennials; cypselas more than 1 mm long 68 68 Leaves entire or serrate or pinnatifid, sessile; cypsela ribs rounded 71 . Leucanthemum Leaves trifid or ternate-pinnatifid and seemingly petiolate; cypsela ribs narrow and somewhat wing-like 72. Rhodanthemum 69 Cypselas triquetrous with 1 adaxial and 2 lateral more or less thick ribs and sometimes with 1-2 abaxial weaker ribs, abaxially and apically with 2 distinct (occasionally fused to 1 or 3-5) resin sacs 98. Tripleurospermum Cypselas various, sometimes triquetrous with 3 adaxial-lateral ribs but not with 2 abaxial-apical resin sacs 70 70 Annual herbs 71 More or less suffruticose perennials, shrublets or shrubs ... 81 71 Cypselas laterally pilose; pappus a fimbriate whitish corona (disc cypselas) or auricle (ray cypselas) 93. Microcephala Cypselas glabrous; sometimes glandular; pappus various ... 72 72 Ray cypselas dorsiventrally flattened and laterally winged; disc cypselas 5-ribbed 91. Daveaua Cypselas equal or subequal, sometimes somewhat flattened but not winged, variously ribbed 73 73 Cypselas somewhat dorsiventrally flattened and with 2 lateral weak ribs; corolla lobes with central resin sacs 99. Aaronsohnia Cypselas with 3 or more, sometimes adaxial, more or less distinct ribs, sometimes slightly flattened; corolla lobes without or sometimes with central resin sacs (in Matrlcarld) 74 74 Cypselas with 1 abaxial (not adaxial) and 2 lateral thick ribs and 2-3 adaxial weaker ribs ; pappus an adaxial stiff auricle 89. Otospermum Cypselas with mainly adaxial-lateral ribs or with ribs all around the cypsela; pappus of scales, a scarious auricle or a corona . 75 75 Cypsela wall and pappus white and spongy, abaxially thin 96. Foveolina Cypsela wall not white and spongy, pappus scarious 76 76 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with or without 2 weaker ribs, or cypselas with 5 adaxial-lateral GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE ribs 77 Cypsela ribs all around the cypsela, not mainly adaxial- lateral 79 77 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with or without 2 weaker ribs; pappus of scales or a stiff corona . 78 Cypselas with 5 adaxial-lateral ribs; pappus an auricle or a small corona 92. Matricaria 78 Cypselas with 3 adaxial-lateral rounded ribs and 2 abaxial weaker ribs, often with 3-5 resin sacs apically in the ribs; pappus of 7-10 obovate scales 90. Heteromera Cypselas with 3 adaxial-lateral acute ribs; pappus a stiff corona 94. Endopappus 79 Pappus a scarious, flimsy corona; cypselas with myxogenic cells on the ribs; leaves pinnatifid-pectinate, spathulate in out- line 69. Hymenostemma Pappus coroniform, of short more or less connate scales, or an auricle; cypselas with or without myxogenic cells; leaves vari- ous 80 80 Cypselas obconical-turbinate, generally with myxogenic cells; slender annuals with several stems from a basal rosette (Anthe- mis subgen. Ammanthus) 57. Anthemis Cypselas more or less oblong, without myxogenic cells; habit various 15. Tanacetum 81 More or less suffruticose perennials 82 Woody shrubs or shrublets 84 82 Cypselas with myxogenic cells on the ribs 83 Cypselas without myxogenic cells 15. Tanacetum 83 Pappus a scarious, flimsy corona; leaves serrate to pinnatifid, generally pectinate and spatulate in outline 68. Leucanthemopsis Pappus of separate, mainly abaxial scales; leaves pinnati- sect 16. Opisthopappus 84 Cypselas dorsiventrally flattened and laterally winged 100. Leucoptera Cypselas more or less terete, not winged 85 85 Leaves obovate and apically serrate 64. Nipponanthemum Leaves pinnatisect to variously lobed 86 86 Pappus an adaxially longer cup, or one large adaxial and one smaller abaxial scale ; cypselas with myxogenic cells 84. Cymbopappus Pappus a small corona, of several scales or an adaxial auricle; cypselas with or without myxogenic cells 87 87 Cypselas with myxogenic cells; pappus of several adaxially longer scales 18. Xylanthemum Cypselas without myxogenic cells; pappus a corona of small scales or an adaxial auricle 15. Tanacetum 88 Ray cypselas triquetrous, winged; disc cypselas laterally flat- tened and adaxially and abaxially winged 89 Cypselas generally equal, sometimes triquetrous or dorsiven- trally (not laterally) flattened and laterally (not adaxially and abaxially) winged 90 89 Annual herb 61. Ismelia Shrubs or shrublets 62. Argyranthemum 90 Disc corolla 4-lobed 91 Disc corolla 5-lobed 97 91 Leaves entire, lanceolate to linear 83. Thaminophyllum Leaves variously lobed or pinnatisect, not entire 92 92 Disc corolla tube very much swollen and brittle 88. Oncosiphon Disc corolla tube not or only slightly swollen 93 87 93 Delicate, somewhat succulent, annual herb; leaves with rounded lobes; capitula small, 5 mm or less in diam 58. Nananthea Annuals or perennials; leaves various; capitula generally more than 5 mm in diam 94 94 Leaves with few, oblong to rounded, apically mucronate lobes; receptacle pilose 82. Lidbeckia Leaves variously pinnatisect ; receptacle glabrous 95 95 Cypselas with 5 adaxial-lateral ribs, somewhat flattened; annu- als 92. Matricaria Cypselas flattened and with 2 lateral ribs, or obovoid and with 2-3 faint ribs; perennials or rarely annuals 96 96 Shrub ; cypselas subglobose with 2-3 adaxial faint ribs 102. Hilliardia Herbs; cypselas dorsiventrally flattened and with 2 lateral ribs . 103. Cotula 97 Leaves opposite Leaves alternate 98 99 98 Shrubs with entire or few-lobed leaves 4. Eumorphia Creeping, suffruticose perennials with serrate-pinnatifid leaves 67. Phalacrocarpum 99 Cypselas triquetrous, with 1 adaxial and 2 lateral more or less thick ribs and sometimes with 2 abaxial weaker ribs, abaxially and apically with 2 distinct (occasionally fused to 1 or 3-5) resin sacs 98. Tripleurospermum Cypselas various, generally with 5 or more ribs, or with 2 lateral ribs only 100 100 Cypselas somewhat dorsiventrally flattened, with 5 adaxial ribs or with 2 lateral ribs only 101 Cypselas more or less terete with 5 or more ribs all around the cypsela 102 101 Cypselas smooth, with 2 lateral weak ribs .... 99. Aaronsohnia Cypselas with 5 adaxial-lateral ribs 92. Matricaria 102 Cypselas distinctly ribbed with dark vallecular secretory canals between the pale ribs 71 . Leucanthemum Cypselas distinctly or faintly ribbed but vallecular secretory canals absent 103 103 Disc corolla tube confluent with the cypsela; cypsela ribs with resin canals; leaves serrate 80. Adenanthellum Disc corolla tube not confluent with the cypsela; cypsela ribs without resin canals; leaves various 104 104 Cypselas 10-18-ribbed with ovoid myxogenic trichomes; shrubs or half-shrubs with entire or few-lobed leaves 3. Phymaspermum Cypselas generally with less than 10 ribs, with or without appressed myxogenic cells but not with ovoid myxogenic tri- chomes; habit and leaves various 105 105 Annual herb with lobed, rather lacerate leaves; cypselas without myxogenic cells 66. Nivellea Perennials; leaves various; cypselas with or without myxogenic cells 106 106 Basally much woody shrublets with few-lobed leaves with linear lobes 30. Brachanthemum Herbs or half-shrubs generally with rather broad leaf-lobes or with linear or serrate leaves 107 107 Cypselas oblong, more or less distinctly ribbed, without myxo- genic cells 108 Cypselas obovoid, faintly ribbed, mostly with myxogenic cells 31 . Dendranthema 108 Leaves mostly rosulate; ray florets fertile . 32. Arctanthemum Leaves alternate; ray florets sterile 65. Leucanthemella K. BREMER AND C. J. HUMPHRIES 109 Cypselas triquetrous with 1 adaxial and 2 lateral more or less thick ribs and sometimes with 2 abaxial weaker ribs, abaxially and apically with 2 distinct (occasionally fused to 1 or 3-5) resin sacs 98. Tripleurospermum Cypselas various, sometimes triquetrous with 3 adaxial-lateral ribs but not with 2 abaxial-apical resin sacs 110 110 Pappus present, of scales (bristle-like in Ugamia), an auricle or a corona (rarely almost absent in Tanacetum but cypselas then apically with an acute rim) Ill Pappus absent but cypselas sometimes apically with an obtuse rim (mainly abaxial and somewhat toothed in Oncosiphon; rarely with a pseudopappus of bristle-like stalked glands in Athanasia) 138 111 Annualherbs 112 Perennial herbs or often half-shrubs, shrublets or shrubs . 120 112 Cypselas conspicuously rugose to tuberculate, without ribs,, apically with a thick, spreading to revolute pappus-like rim 87. Rennera Cypselas often ribbed and not rugose to tuberculate, or if rugose then also with 3 distinct adaxial-lateral ribs; pappus of scales, a corona (but not thick and spreading to revolute) or an auricle 113 113 Cypselas somewhat dorsiventrally flattened and with 2 lateral weak ribs ; corolla lobes with central resin sacs 99. Aaronsohnia Cypselas with 3 or more, sometimes adaxial, more or less distinct ribs, sometimes slightly flattened; corolla lobes without or sometimes with central resin sacs (in Matricarid) 114 114 Cypselas laterally pilose; pappus a fimbriate whitish corona 93. Microcephala Cypselas glabrous, sometimes glandular; pappus various . 115 1 15 Cypsela wall and pappus white and spongy, abaxially thin 96. Foveolina Cypsela wall not white and spongy; pappus scarious 116 116 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with or without 2 weaker ribs, or cypselas with 5 adaxial-lateral ribs 117 Cypsela ribs all around the cypsela, not mainly adaxial- lateral 118 117 Cypselas with 1 adaxial and 2 lateral more or less thick ribs; pappus a stiff corona covered with myxogenic cells 95. Myxopappus Cypselas with 5 adaxial-lateral ribs; pappus an auricle or a small corona 92. Matricaria 118 Cypselas with dark vallecular secretory canals between the pale ribs and with myxogenic cells on the ribs; pappus an adaxial but basally coroniform auricle as long as the corolla or longer 74. Chlamydophora Cypselas without vallecular secretory canals, with or without myxogenic cells; pappus of small scales, a small corona or auricle 119 119 Cypselas obconical-turbinate, generally with myxogenic cells; slender annuals with several stems from a basal rosette (Anthe- mis subgen. Ammanthus) 57. Anthemis Cypselas more or less oblong, without myxogenic cells; habit various 15. Tanacetum 120 Pappus of 4-15 linear and bristle-like to obovate-oblong scales at least half as long as the corolla 121 Pappus of short scales, an auricle or a corona 123 121 Cypselas densely pilose; pappus scales white 122 Cypselas glabrous or sparsely pilose; pappus scales brownish .. 13. Cancrinia 122 Capitula on short, nodding peduncles 10. Ugamia Capitula on elongated, straight peduncles or pedunculoid stems 9. Trichanthemis 123 Cypselas with dark vallecular secretory canals between the pale ribs and with myxogenic cells on the ribs; pappus an adaxial auricle 78. Plagius Cypselas without vallecular secretory canals; pappus of scales, a corona or sometimes an auricle 124 124 Capitula small and numerous in an elongated panicle or a large corymb; basally villous and woody perennials with much pinna- tisect leaves 125 Capitula and leaves various, sometimes small and rather many in a panicle or corymb but plant then not basally villous with much pinnatisect leaves; perennial herbs, half-shrubs, shrublets or shrubs 1 26 125 Capitula in an elongated panicle 25 . Lepidolopsis Capitula in a large corymb 26. Polychrysum 126 Capitula paniculate ; leaves apically few-lobed or entire 42. Crossostephium Capitula solitary or laxly to densely corymbose or closely aggregated; leaves various 127 127 A compact, hirsute, basally woody half-shrub with solitary, pecunculate capitula; cypselas distinctly 10-ribbed with a coroni- form pappus of short wide scales 14. Cancriniella Habit and cypselas various, often perennial herbs or shrubs 128 128 Capitula closely aggregated at the stems; leaves small, ericoid, entire or occasionally few-lobed; pappus of 7-10 oblong, adaxi- ally longer scales 86. Marasmodes Capitula solitary or laxly to densely corymbose; leaves and pappus various 129 129 Capitula narrowly oblong-obconical, slender and few-flowered; pubescent shrubs; indumentum of stellate hairs 6. Hymenolepis Capitula generally urceolate to cyathiform or campanulate; habit various; indumentum if present of simple or bifid hairs 130 130 Shrub with corymbose capitula and pinnatisect leaves; cypselas 8-10-ribbed; pappus of small scales projecting from the cypsela ribs 24. Inulanthera Pappus of scales not distinctly projecting from the cypsela ribs, a corona or an auricle ; habit and cypselas various 131 131 Pollen smooth; a small densely pubescent, basally woody peren- nial with solitary or few capitula on short peduncles (Sphaer- omeria compactd) 39. Sphaeromeria Pollen spiny; habit and capitula various 132 132 Shrublets with entire or 3-lobed leaves; capitula rather narrowly urceolate with involucral bracts in 5-7 rows; pappus of many subulate scales 19. Lepidolopha Habit, capitula and pappus various, if shrubby with entire or 3-lobed leaves (some Pentzia species), then not with involucral bracts in 5-7 rows and not with a pappus of subulate scales 133 133 Capitula disciform; outer female florets present; cypselas with- out myxogenic cells 15. Tanacetum Capitula discoid; all florets hermaphrodite; cypselas with or without myxogenic cells 134 134 Corolla tube generally swollen and with thick vascular strands; shrubs with more or less ericoid leaves 85 . Pentzia Corolla tube not or only slightly swollen and with thin vascular strands; habit and leaves various 135 135 Pappus an adaxial auricle or of several adaxially more devel- GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 89 oped scales 136 Pappus a corona or of scales, adaxially not more devel- oped 137 136 Perennial herbs with corymbose capitula 15. Tanacetum Shrublets with solitary capitula 18. Xylanthemum 137 Leaves entire but marginally crenate-serrate; perennial herb (Tanacetum balsamita) 15. Tanacetum Leaves pinnatisect; habit various 17. Tanacetopsis 138 Leaves opposite; shrubs with sessile or corymbose capitula 139 Leaves alternate, rarely opposite but then habit or capitula different 140 139 Leaves entire; capitula sessile 8. Asaemia Leaves pinnatifid; capitula corymbose 5. Gymnopentzia 140 Capitula solitary or in lax to dense corymbs, small to medium- sized and generally erect; pollen usually spiny, rarely smooth (in Ajaniopsis, Stilpnolepis, Filifolium, and Sphaeromeria) ... 141 Capitula in elongated and paniculate, rarely racemiform or spiciform or subglobose inflorescences, often small and numer- ous and sometimes pendent ; pollen smooth 168 141 Central floret corolla 4-lobed (rarely 3-lobed in Cotuld) ... 142 Central floret corolla 5-lobed 150 142 Cypselas dorsiventrally flattened with 2 lateral more or less distinct ribs or wings 143 Cypselas not or only slightly dorsiventrally flattened without or with 3-5 ribs and no wings 146 143 Shrublets or half-shrubs with corymbose or occasionally solitary capitula 106. Schistostephium Annual or perennial herbs with solitary capitula, rarely few together 144 144 Capitula sessile; outer female florets without corolla and with style persistent and spinescent in fruit 105. Soliva Capitula more or less pedunculate; outer female florets with corolla; style not persistent 145 145 Outer female floret corolla inflated with a hollow space between the outer and inner layer; central florets female-sterile 104. Leptinella Outer female floret corolla not inflated; central florets generally fertile 103. Cotula 146 Perennial (rarely facultative annual) prostrate herbs, regularly rooting at the nodes ; central florets female-sterile 104. Leptinella Annual herbs, not regularly rooting at the nodes; central florets fertile 147 147 Disc corolla tube very much swollen and brittle 88. Oncosiphon Disc corolla tube not or only slightly swollen 148 148 Delicate, somewhat succulent herb; leaves with rounded lobes; capitula small, 5 mm or less in diam 58. Nananthea Habit and leaves various; capitula generally more than 5 mm in diam 149 149 Cypsela wall either white and spongy with 1 adaxial and 2 lateral weak ribs, abaxially thin, or cypselas almost terete and thin- walled all around (additional outer conspicuously rugose cypse- las may be present) 96. Foveolina Cypsela wall not white and spongy, with 5 adaxial-lateral ribs 92. Matricaria 150 Annualherbs 151 Perennial herbs or often half-shrubs, shrublets or shrubs . 156 limb inflated, crateriform; cypselas obovoid-lanceolate, densely glandular 36. Stilpnolepis Involucral bracts obovate-oblong, only marginally scarious; corolla limb more or less distinct but not inflated and crateri- form; cypselas various but not obovoid-oblanceolate and densely glandular 152 152 Corolla lobes densely pilose; capitula few together in dense corymbs; cypselas obovoid with rows of myxogenic cells 37. Ajaniopsis Corolla lobes glabrous, sometimes glandular; capitula often solitary or laxly corymbose; cypselas various 153 153 Cypselas distinctly rugose to tuberculate, more or less 5-angled 87.Rennera Cypselas smooth or ribbed and not rugose to tuberculate (outer conspicuously rugose cypselas in addition to the smooth central cypselas sometimes present in Foveolina) 154 154 Cypselas somewhat dorsiventrally flattened and with 2 lateral weak ribs ; corolla lobes with central resin sacs 99. Aaronsohnia Cypselas smooth and obovoid to terete, or with 3-5 adaxial- lateral, more or less distinct ribs; corolla lobes without or sometimes with central resin sacs (in Matricaria) 155 155 Cypsela wall either white and spongy with 1 adaxial and 2 lateral weak ribs, abaxially thin, or almost terete and thin-walled all around (additional outer conspicuously rugose cypselas may be present) 96. Foveolina Cypsela wall not white and spongy, with 5 adaxial-lateral ribs 92. Matricaria 156 Cypselas dorsiventrally flattened, with or without lateral wings 107. Hippia Cypselas not dorsiventrally flattened, unwinged 157 157 Cypselas with dark vallecular secretory canals between the pale ribs and with myxogenic cells on the ribs .. 71. Leucanthemum Cypselas with or without ribs but vallecular secretory canals absent 158 158 Capitula disciform; outer female florets present 159 Capitula discoid; all florets hermaphrodite 162 159 Central floret corollas soon compressed together in a resinous mass; cypselas obliquely obovoid; a perennial herb with basal fibrous leaf sheaths and filiform leaf lobes 38. Filifolium Central floret corollas not compressed in a resinous mass; cypselas straight and obovoid to obovate-oblong; habit and leaves various 160 160 Pollen smooth ; small perennial herbs or half-shrubs 39. Sphaeromeria Pollen spiny (sometimes with short spines only); habit vari- ous 161 161 Style-branches brownish; corolla lobes erect . 35. Phaeostigma Style-branches yellowish; corolla lobes spreading .. 34. Ajania 162 Cypselas slender and somewhat arcuate, tuberculate with numerous obtuse excrescences; a basally villous and woody perennial with much pinnatisect leaves and densely corymbose capitula 27. Pseudohandelia Cypselas obovoid-oblong, straight or somewhat oblique, smooth or ribbed but not tuberculate; habit, leaves and capitula various 163 163 Cypsela ribs acute, with secretory cavities (best seen in cross- section) ; corolla gradually expanded and funnel-shaped 7 . Athanasia Cypsela ribs faint or rounded, without secretory cavities; corolla more or less distinctly divided into tube and limb 1 64 151 Involucral bracts widely obovate and largely scarious; corolla 164 Cypselas 10-18-ribbed 3. Phymaspermum 90 K. BREMER AND C. J. HUMPHRIES Cypselas with fewer than 10 ribs 165 more rows, tubular, rarely without corollas. Central disc florets 5- or 4-lobed, rarely 3-lobed, yellow or rarely whitish 165 Anthers tailed; perennial herbs with much pinnatisect, basally - ,. J . ?n Hi nnn iv*i or red, rare y somewhat zygomorphic, perfect or functionally more or less rosulate leaves 20. Hippolytia J Anthers not tailed; habit various; leaves entire or rather few- male. Anthers generally obtuse at base, rarely shortly tailed; j obed 166 filaments basifixed. Style-branches (in central florets) almost always truncate and penicillate, with stigmatic areas in two 166 Perennial herbs with laxly corymbose capitula . 36. Stilpnolepis marg j na i stripes, but sometimes undivided in functionally Shrubs or shrublets with solitary capitula 1 staminate florets. Cypselas variable, homo- or heteromor- 167 Leaves with linear lobes 30. Brachanthemum phic, generally terete to weakly angled or ribbed or flattened, Leaves ericoid with short lobes 85. Pentzia sometimes winged, thin- or thick-walled, without a carbon- ized layer, often with secretory canals and myxogenic cells. 168 Capitula disciform; outer female florets present; mvolucral * b t ' 2-5 ws 169 Pappus generally of rather few scanous scales, a corona or an Capitula discoid; all florets hermaphrodite; involucral bracts in auricle, rarely of many flat bristle-like scales, often absent. 4-7 rows, unequal, the outer short and rounded, the inner Embryo sac monosporic or sometimes tetrasporic, rarely gradually longer and linear 41. Seriphidium bisporic. Chromosome number generally x=9, sometimes x=10, rarely x=6, 7, 8, 11 or 18. Irregular monoterpenes 169 Corolla lobes pilose present in high concentrations. Corolla lobes glabrous, sometimes glandular 173 170 Cypselas densely pilose .. 171 DISTRIBUTION (Table 3). Worldwide but with main concen- Cypselas glabrous 172 trations in central Asia, the Mediterranean region and South Africa. - 12 subtribes, 108 genera, 1741 spp. 171 Outer female florets without corolla and with dilated, lan- ceolate, flat style-branches; a virgate shrub 46. Mausolea The interrelationships of the 12 proposed subtribes are uncer- Outer female florets with a tubular corolla and linear style- ta j n but one possible hypothesis is presented in the cla- branches; a woody shrublet with older branches transformed dogram (Fig. 1). It should be noted that a number of equally into s P ines 47 - Picrothamnus parsimonious solutions are possible , and that the strict con- 172 Leaves pinnatisect; capitula densely congested in glomerules sensus tree of these solutions is totally collapsed. The chosen arranged in spikes, or solitary in interrupted partly congested cladogram is offered simply as a suggestion for further spikes ; indumentum of simple of bifid hairs analysis and test . The characters used to identify the various 45. Turaniphytum clades, indicated in the cladogram, are listed with comments Leaves entire or few-lobed; capitula few in an elongated below panicle, at the summit fasciculate; indumentum of stellate hairs 40. Kaschgaria 173 Suffruticose perennial with entire or 3-5-lobed leaves; capitula i AO _Jl JP^ ^Ursiniinae 30-50-flowered in a pyramidal to elongated panicle (Sphaer- 1C 2 Cancriniinae omeria ruthiae) 39. Sphaeromeria 3 Tanacetinae Habit, leaves, and capitula various, usually with fewer than 30 IU:A2= = 4 Gonosperminae florets 174 5 Handeliinae = 6 Artemisiinae 174 Central florets of two kinds; outer perfect, inner completely ._ 7 Achilleinae sterile with reduced ovaries; panicle spiciform; leaves pectinate- r= 8 Anthemidinae pinnatisect with filiform, apically somewhat swollen and it=A3= =A4 9 Chrysantheminae mucronulate lobes 44. Neopallasia F= 10 Leucantheminae Central florets all perfect or all female-sterile with reduced J=A5^= 11 Thaminophyllinae ovaries ; inflorescence and leaves various 43 . Artemisia ^ Matricarnnae Fig. 1 Cladogram (of five possible) of the 12 subtribes; produced by the mhennig* option in Hennig86. Cladogram length = 39, consistency index = 84, retention index = 53. ANTHEMIDEAE Cass. /. Phys. Chim. Hist nat. 88: 192 (1819). Type species: Anthemis maritima L. Aromatic annual or perennial herbs, subshrubs or shrubs, rarely spinescent. Leaves alternate or rarely opposite or fasciculate or rosulate, generally variously dissected, pinnati- sect, pinnatifid, lobed or serrate-dentate or rarely entire. Capitula solitary or corymbose or paniculate, rarely aggre- gated, often pedunculate, sometimes sessile, variable in size, radiate or disciform and heterogamous or homogamous and discoid. Involucral bracts in three or more rows, rarely in two rows, imbricate, almost always with scarious margins and apex. Receptacle paleate or epaleate, rarely pilose or hirsute. Ray florets (in radiate capitula) female, fertile or sterile, or neuter; limb white, yellow or rarely blue-violet, pink or reddish. Outer female florets (in disciform capitula) in one or Clades and characters - Fig. 1, Tables 2 & 4. Clade AO 15 Leaves variously deeply lobed or divided. Dissected leaves are characteristic of most Anthemideae as well as of its postulated relatives in the Heliantheae. Some of the Anthemideae genera and species have entire leaves, but these taxa are clearly related to other taxa with dissected leaves. 43 Involucral bracts with scarious margins. This is characteris- tic of almost all Anthemideae as well as of its postulated relatives in Heliantheae, but also in some austral and grangeoid Astereae. At the generic level. Myxopappus with subulate involucral bracts represents a reversal. Clade Al - tribe Anthemideae GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 91 Table 4 Data matrix for the Anthemideae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. Ill 1 1 1 1 11 1 111 1450574597 4616 1319758857 422746 532718565944192865132124251101706 1. Ursiniinae alaal?al 1 lOOpOOOOpOOppOOpOppOOOOO 2. Cancriniinae alaal?OOOOaapOOOOpOOOpOOO?0000000 3. Tanacetinae alaalapOOOpplppppppOppOOOppOOpOOp 4. Gonosperminae alaal?aOp0001alOOpOOOOOOO?0000000 5. Handeliinae al?al?p000001001alOOppOOO?0000000 6. Artemisiinae alaalapOOOpplOOOppalapOOOOpOOOOOO 7. Achilleinae alaalaaOOOOplOOOpppOaaalOOppOpOOO 8. Anthemidinae alaalapOOOOplOOOOppOpapOalpOOOOOp 9. Chrysantheminae alaal?0000001000p0001a001pallpOOO lO.Leucantheminae alall?pOOOOplOOOOpaOpapOOOppOaOOp H.Thaminophyllinaealall?pp000010000000ppOOO?OOOalOO 12.Matricariinae aaallapOpOOOlOOOppOOpapOOpppOOpaa 52 Ray floret limb white. Rays in the Anthemideae are mostly white and sometimes yellow (rarely pink, reddish or blue- violet). In the Heliantheae relatives rays are yellow. 107 Cypselas terete to weakly angled, or flattened. The fruits are very variable in the Anthemideae, but the sharply angled type, present in all of its postulated immediate relatives in the Heliantheae, is hardly ever present. 151 Cypsela wall without carbonized layer. A carbonized layer in the fruit wall is characteristic of many Heliantheae and of all Eupatorieae, including the immediate relatives of the Anthemideae, but it is totally absent in this tribe. The absence of this feature is thus interpreted as a derived condition for the tribe. 178 'Irregular' monoterpenes in high concentrations present (Greger, 1977). Clade A2 161 Pappus of short (not large and obovate or bristle-like) scales, an auricle, a corona, or absent. In subtribe Cancrinii- nae, which is not a member of this clade, the pappus consists of many subulate, bristle-like scales and it looks very anoma- lous within the tribe. The same pappus type is found in the immediate relatives within Heliantheae. Ursinia in subtribe Ursiniinae, also not a member of this clade, has a pappus of large and obovate scales, similar to some of the wider pappus scales in Cancriniinae and Heliantheae. Judging from other characters, Ursinia is related to a number of South African genera, all united in subtribe Ursiniinae. In these genera the pappus is absent and we consider it secondarily lost. How- ever, in one genus, Hymenolepis , a pappus of small scales has evolved. In all other Anthemideae, comprising clade A2 and excluding Cancriniinae and Ursiniinae, the pappus, if not secondarily lost, consists of scales, an auricle or a corona. Clade A3 51 Floral parts with resin canals. Resin canals or sacs are frequently present in style-branches, corolla lobes and some- times also anther tips in a majority of the Anthemideae. They seem to occur mainly in the subtribes of this clade, though present also in some genera of other subtribes. Clade A4 152 Cypsela wall several cell layers thick, partially or com- pletely sclerified. Clade A5 21 Leaves serrate-dentate. In the two subtribes of this clade non-dissected leaves dominate, though there are exceptions in several genera. 1. URSINIINAE Bremer & Humphries, subtrib. nov. Type species: Ursinia paradoxa Gaertner (U. chrysan- themoides (Less.) Harvey). Herbae annuae vel perennes vel frutices. Corollae floscu- lorum radii cellulis epidermalibus tabularibus. Antherae cel- lulis endothecii polaratis. Pappus squamiformis magnus aut parvus vel nullus. Furanosesquiterpena adsunt. Annual or perennial herbs or shrubs. Leaves alternate or sometimes opposite, variable in shape. Capitula solitary or laxly to densely corymbose, pedunculate or rarely sessile along the stems, radiate or discoid. Receptacle paleate or epaleate. Ray floret limb yellow, white or rarely red; epider- mis cells tabular. Disc corolla 5-lobed. Anthers with endoth- ecial tissue polarized. Cypselas 5- to many-ribbed, glabrous or pubescent, papillose to copiously villous, sometimes with myxogenic cells. Pappus of large or small scales, or often absent. Furanosesquiterpenes present. DISTRIBUTION (Table 5). Southern Africa, one Ursinia spe- cies in Ethiopia and one Lasiospermum species in Egypt (Sinai), some Ursinia species introduced in Australia and New Zealand. - 8 genera, 114 spp. Table 5 General distribution of Ursiniinae and genera. x=indigenous, o=introduced. S. Eur. N. Afr. S. Afr. Austr. N. Zeal. Ursiniinae X X o Ursinia X X Lasiospermum X X Phymaspermum X Eumorphia X Gymnopentzia X Hymenolepis X Athanasia X Asaemia X This subtribe was first identified as a group by Bohlmann and collaborators (Greger, 1977), who discovered that a number of South African Anthemideae genera contained furanoses- quiterpenes rather than the common polyacetylenes. Subse- quently, the classification of the group has been revised by Kallersjo (1986, 1991), who added a number of micromor- phological characters (see cladogram in Fig. 2). Kallersjo circumscribed it more precisely, by moving a number of Athanasia species (to Inulanthera in the Gonosperminae) and by including some chemically unknown genera, now placed in Phymaspermum and Hymenolepis, which have now been investigated chemically. The tribal position of Ursinia within Anthemideae, discussed below, has gained further support. 92 K. BREMER AND C. J. HUMPHRIES There is one alternative equally parsimonious cladogram to that shown here. In the alternative cladogram Phymasper- mum and Gymnopentzia are sister groups based on a shared loss of receptacular paleae (character 45), whereas in the presented cladogram Gymnopentzia and Eumorphia are sis- ter groups based on their opposite leaves (character 14). Clades and characters - Fig. 2, Tables 2, 6. =Ur3 |=Url=ij= 1 Ursinia rp= 2 Lasiospermum H=Ur2=! rF=Ur4^= 3 Phymaspermum U=Ur5^= 4 Eumorphia l!= 5 Gymnopentzia rp= 6 Hymenolepis i=Ur6^ 7 Athanasia '^= 8 Asaemia Fig. 2 Cladogram (of two possible) of the Ursiniinae produced by the ie option in Hennig86. Cladogram length = 39, consistency index = 82, retention index = 79. Table 6 Data matrix for the Ursiniinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. Ill 1111111 111 1 1 11 1 1 145057 4597566735 222441341567395682231 532718 56599412302894544562194242096344 1. Ursinia llaal? 2 . Lasiospermum 11111? 3 . Phymaspermum a 1 1 1 1 ? 4. Eumorphia all 11? 5 . Gymnopentzia 1 1 ? 1 1 ? 6. Hymenolepis 7. Athanasia 8. Asaemia 1111 1 lOpOOOOOOOOOpOOOOOOOOOOOOOO 111 1001 HlOOOOOOOpOOOOOOOOOOOOOO 01 1 1001 1001 1 a 1 1 aOpOOOOOOOOOOOOpO all 10 OllOOllallOaOOOOOOOOOOOOOO 0?1 1001 1001 1 1 1 101 1 10000000000000 a?11001000111000010111 110000000 a?l 1001 1001 laOOOOlOll 11 lallalOOO 0?1 1001 1001 10000110?! Ill 11 1001 11 1111 11 1 177675555566666171 240 337 356600357905678511121997853 I. Ursinia ???OOOOOOOOOOOOOOOppppOOOOO 2. Lasiospermum ?00??OOOOOOOOOOOOOpOOOOOOOO 3.P/i;7mwperwiMjn??????000000000000000000000 4. Eumorphia ?????0000000000000000000000 5. Gymnopentzia ?????????????????0000000000 6. Hymenolepis ???000???????????OOOOOpOOOO 7. Athanasia ?????0???????????OOpOOppppO 8. Asaemia ??????????????????00000000p Clade Url - subtribe Ursiniinae 45 Receptacle paleate. The distribution of this character is difficult to interpret. Receptacular paleae are absent in Phymaspermum, Gymnopentzia, Asaemia, a few species of Athanasia, and one species of Hymenolepis. This is most parsimoniously interpreted as secondary losses within the subtribe. 56 Ray floret limb epidermis cells tabular (senecioid or muti- sioid type) (Baag0e, 1977). 95 Anthers with endothecial tissue partly or wholly polarized. 179 Furanosesquiterpenes present. Ursinia 159 Cypsela wall with a continuous ring of fibre-like cells. 164 Pappus of 5 convolute-contorted scales. Clade Ur2 161 Pappus of short (not large and obovate or bristle-like) scales, an auricle, a corona, or absent. 172 Pappus absent in ray and disc cypselas. Hymenolepis has a pappus of small scales, most parsimoniously interpreted as secondarily evolved within this clade. Lasiospermum 133 Cypselas with costal resin canals or sacs. These structures are similar to those occurring in the subtribe Matricariinae, for example, but different from those in Hymenolepis, Atha- nasia, and Asaemia (character 132). 150 Cypselas copiously villous, seemingly covered in 'cotton wool'. Clade Ur3 2 Plants shrubby. 28 Leaves with secretory cavities. 29 Capitula densely corymbose. Asaemia and some species of Athanasia and Phymaspermum have solitary capitula. Clade Ur4 124 Cypselas with 10 (8-12) multicellular epicarpic ribs. Occasionally there are up to 18 ribs. 145 Cypselas papillose. Phymaspermum 45 reversed. See clade Url. 144 Cypselas with scattered, ovoid, myxogenic trichomes. In a few species the trichomes are not myxogenic, though similar in structure. Clade Ur5 14 Leaves opposite. Eumorphia There is no autapomorphy for Eumorphia. The matter is further discussed under the genus. Gymnopentzia 35 Capitula discoid. 45 reversed. See clade Url. 146 Cypselas long-papillose, seemingly pubescent. Clade Ur6 12 Plants with stellate hairs. 35 Capitula discoid. 51 Floral parts with resin canals. 69 Corolla gradually expanded, rather thin and funnel-shaped. 74 Corollas with continuous veins extending into the lobes. 132 Cypsela ribs with ellipsoid secretory cavities forming longitudinal ducts. Hymenolepis 172 reversed. See clade Ur2. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE Clade Ur7 94 Anthers with an apical resin sac. This character occurs in most but not all species of Athanasia. 152 Cypsela wall several cell layers thick, partially or com- pletely sclerified. 160 Cypsela wall with a continuous ring of sclerified isodiamet- ric cells. Athanasia 89 Corolla tube with long-stalked glands; stalk cells elongated. Some species of Athanasia have glabrous corolla tubes, apparently secondarily. 126 Cypsela ribs protruding, narrow and somewhat wing-like. Asaemia 14 Leaves opposite. 15 reversed. Leaves not variously deeply lobed or divided, but entire. 23 Leaves entire, ericoid. 29 reversed. See clade Ur3. 34 Capitula sessile along the stems. 45 reversed. See clade Url. 114 Cypselas dorsiventrally flattened. 1. URSINIA Gaertner, Fruct. sem. pi. 2: 462 (1791). Type species: U. paradoxa Gaertner (U. chrysanthemoides (Less.) Harvey). - Sphenogyne R. Br. - Ursiniopsis E. Phillips. Annual or perennial herbs or shrublets. Leaves alternate, entire or generally variously lobed. Capitula solitary or laxly corymbose, pedunculate, radiate or occasionally dis- coid. Receptacle paleate; paleae scarious, often enveloping florets, sometimes narrow with an apical limb. Ray florets generally neuter or occasionally female, sterile or fertile; limb yellow, white, or rarely red, dorsally often reddish; epidermis cells tabular. Disc corolla shallowly 5-lobed. Anthers with a widely ovate apical appendage; endothecial tissue polarized. Cypselas slender or obovoid, straight or curved, 5-ribbed, with a basal tuft of long hairs or glabrous. Pappus of 5 (rarely 8-10) large, convolute-contorted, whit- ish scales, or of 5 outer such scales and 5 inner subulate, whitish scales, or occasionally absent. Furanosesquiterpe- nes present. DISTRIBUTION. S. Africa mainly in the SW Cape, also in Namibia, Botswana, and Ethiopia. - 38 spp. Ursinia was revised by Prassler (1967). Species without a basal tuft of cypsela hairs earlier constituted Ursinia s. s. (excluding Sphenogyne; the two genera differed also in other characters). Species with glabrous cypselas are considered derived by Prassler so we assume that presence of cypsela hairs is a diagnostic character for the whole genus, though secondarily lost in some species. Another former genus, Ursiniopsis, was distinguished simply by female rather than neutral rays. It was reduced to synonomy by Prassler. Earlier authors, e.g. Bentham (1873a), placed Ursinia in the tribe 93 Arctoteae sensu Norlindh (1977) mainly because of its well developed pappus scales. Cassini (1816) followed by Beau- verd (1915) and Prassler considered it a member of Anthemideae. Robinson & Brettell (1973) argue that inclu- sion in the Anthemideae of the anomalous genus Ursinia with its conspicuous pappus scales, widely ovate apical anther appendages, and different pollen (exine without columnar structure) would destroy a workable tribal concept. Hence they proposed a monotypic new tribe, Ursinieae. The large pappus scales and the shape of the apical anther appendage may be plesiomorphies within Anthemideae, since similar structures occur in the outgroup. The pollen was investigated by Stix (1960) and she concluded that Ursinia belongs in Anthemideae. The presence of unique furanosesquiterpenes in Ursinia and a number of other South African Anthemideae corroborates its tribal position. Although mainly South African, there is one species (U. nana) also known from Ethiopia and U. anthemoides is introduced into western Australia. The following list of species is taken mainly from Prassler's revision. U. abrotanifolia (R. Br.) Sprengel U. anethoides (DC.) N. E. Br. U. anthemoides (L.) Poiret U. brachyloba (Kunze) Bremer & Humphries, comb. nov. Basionym: Sphenogyne brachyloba Kunze in Linnaea 20: 21 (1847). U. cakilefolia DC. U. caledonica (E. Phillips) Prassler U. calenduliflora (DC.) N. E. Br. U. chrysanthemoides (Less.) Harvey (U. paradoxa Gaertner) U. coronopifolia (Less.) N. E. Br. U. dentata (L.) Poiret U. discolor (DC.) N. E. Br. U. dregeana (DC.) N. E. Br. U. eckloniana (Sender) N. E. Br. U. filipes (E. Meyer ex DC.) N. E. Br. U. frutescens Dinter U. heterodonta (DC.) N. E. Br. U. hispida (DC.) N. E. Br. U. macropoda (DC.) N. E. Br. U. merxmuelleri Prassler U. montana DC. U. nana DC. U. nudicaulis (Thunb.) N. E. Br. U. oreogena Schltr ex Prassler U. paleacea (L.) Moench (U. crithmoides (P. Bergius) Poiret) U. pilifera (P. Bergius) Poiret U. pinnata (Thunb.) Prassler U. punctata (Thunb.) N. E. Br. U. pygmaea DC. U. quinquepartita (DC.) N. E. Br. U. rigidula (DC.) N. E. Br. U. saxatilis N. E. Br. U. scariosa (Aiton) Poiret U. sericea (Thunb.) N. E. Br. U. serrata (L. f.) Poiret U. speciosa DC. U. subflosculosa (DC.) Prassler U. tenuifolia (L.) Poiret U. trifida (Thunb.) N. E. Br. 94 K. BREMER AND C. J. HUMPHRIES 2. LASIOSPERMUM Lagasca, Gen. sp. pi.: 31 (1816). Type species: L. pedunculare Lagasca (L. erectum (Lam.) Druce). Annual or perennial herbs. Leaves alternate, variously pinna- tisect. Capitula solitary, pedunculate, radiate or discoid. Receptacle flat or convex, paleate; paleae thin and scarious with a conspicuous resin canal. Ray florets female, fertile; limb very short to long, white or reddish; epidermis cells tabular. Disc corolla 5-lobed; lobes sometimes reddish. Anthers with endothecial tissue polarized. Cypselas copiously villous, with resin canals. Pappus absent. Furanosesquiterpe- nes present. DISTRIBUTION. S. Africa in the Cape, Namibia, and Egypt in Sinai. -4 spp. Lasiospermum is a well defined genus with copiously villous cypselas and the development of the cypsela wool deserves detailed investigation. One annual species (L. brachyglos- sum) occurs also in Sinai. Similar disjunctions are known also from other groups. The first three species in the list are taken from Flora capensis (Harvey, 1865). L. bipinnatum (Thunb.) Druce (L. radiatum Trevir.) L. brachyglossum DC. L. erectum (Lam.) Druce (L. pedunculare Lagasca) L. poterioides Hutch. - Note: Description in Hutchinson, 1946. 3. PHYMASPERMUM Less., Syn. gen. compos.: 253 (1832). Type species: P. junceum Less. - Adenachaena DC. - Brachymerls DC. - locaste E. Meyer ex Harvey Shrubs or half-shrubs. Leaves alternate, entire or lobed, often ericoid with secretory cavities. Capitula solitary or corymbose, generally pedunculate, rarely sessile along the stems, radiate or discoid. Receptacle flat to conical, epaleate. Ray florets female, fertile; limb white; epidermis cells tabu- lar. Disc corolla 5-lobed, rarely pubescent with long hairs, with a narrow tube and a distinct limb. Cypselas 10-18- ribbed, generally minutely papillose especially on the ribs and generally with scattered ovoid myxogenic trichomes, with a more or less distinct apical rim. Pappus absent. Furanoses- quiterpenes present. DISTRIBUTION. S. Africa in the Cape, Orange Free State, and Transvaal, Swaziland, Zimbabwe and Namibia. - 18 spp. Phymaspermum was formerly diagnosed as South African shrubs with epaleate and radiate capitula and papillose and glandular cypselas. Only the cypsela character represents a useful synapomorphy for the genus. Phymaspermum cypselas have a peculiar and unique type of myxogenic trichome, which are ovoid or almost subglobose and scattered over the cypsela surface. Brachymeris was first described as a mono- typic genus and the type species, B. scoparia, is a shrub with much reduced leaves, small capitula sessile along the stems, and a pubescent corolla. Hutchinson (1917) described four new species and transferred one Pentzia species to Brachym- eris, mainly because they all, like B. scoparia, possessed cypselas without a pappus. Hutchinson's species differ from B. scoparia in several characters; they have rather closely set long leaves, capitula in corymbs, and glandular but not pubescent corollas. Kallersjo (1986), who investigated these genera, concluded that B. scoparia as well as Hutchinson's species could be transferred to Phymaspermum because of their similar cypsela morphology. All Brachymeris species have the Phymaspermum type of ovoid cypsela trichomes, although they are not always myxogenic. Kallersjo also transferred a group of Athanasia species with the same cypsela morphology to Phymaspermum. These Athanasia species are also in habit similar to P. aciculare for example. The species of Phymaspermum s. s. (excluding the former Athanasia and Brachymeris species but including Aden- achaena and locaste) are taken from Flora capensis (Harvey, 1865) with three species described later added. The former Athanasia and Brachymeris species are best identified using the treatments by Hilliard (1977) and Hutchinson (1917), respectively. P. acerosum (DC.) Kallersjo (Athanasia acerosa (DC.) D. Dietr. P. aciculare (E. Meyer ex Harvey) Benth. ex B. D. Jackson (locaste acicularis E. Meyer ex Harvey) P. appressum Bolus - Note: Description in Bolus, 1905. P. argenteum Brusse - Note: Description in Brusse, 19896. P. athanasioides (S. Moore) Kallersjo (Brachymeris athana- sioides (S. Moore) Hutch.) P. bolusii (Hutch.) Kallersjo (Brachymeris bolusii Hutch.) *P. equisetoides Thell. - Note: Description in Thellung, 1923. P. erubescens (Hutch.) Kallersjo (Brachymeris erubescens Hutch.) P. junceum Less. P. leptophyllum (DC.) Benth. ex B. D. Jackson (Aden- achaena leptophylla DC.) P. montanum (Hutch.) Kallersjo (Brachymeris montana Hutch.) P. parvifolium (DC.) Benth. ex B. D. Jackson (Adenachaena parvifolia DC.) P. peglerae (Hutch.) Kallersjo (Brachymeris peglerae Hutch.) P. pinnatifidum (Oliver) Kallersjo (Athanasia pinnatifida (Oliver) Hilliard) P. schroteri Compton - Note: Description in Compton, 1931. P. scoparium (DC.) Kallersjo (Brachymeris scoparia DC.) P. villosum (Hilliard) Kallersjo (Athanasia villosa Hilliard) P. woodii (Thell.) Kallersjo - Athanasia woodii (Thell.) Hilliard) 4. EUMORPHIA DC., Prodr. 6: 2 (1838). Type species: E. dregeana DC. Shrubs. Leaves opposite, rarely alternate, entire or lobed, more or less ericoid, with secretory cavities. Capitula gener- ally solitary or corymbose, generally pedunculate, radiate. Receptacle flat or slightly convex, rarely conical, generally paleate, sometimes epaleate. Ray florets female, fertile; limb white; epidermis cells tabular. Disc corolla 5-lobed, with a narrow tube and a distinct limb. Anthers with endothecial tissue polarized. Cypselas 10-12-ribbed, rarely up to 18-ribbed, minutely papillose especially on the ribs, with an apical rim. Pappus absent. Furanosesquiterpenes present. DISTRIBUTION. S. Africa in the central Cape, Natal and Transvaal, and in Lesotho and Swaziland. - 6 spp. Traditionally Eumorphia comprises South African Anthemideae with radiate capitula (a plesiomorphy), a pale- ate receptacle and cypselas without a pappus, characters common to several other genera. E. prostrata has a partly GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE epaleate receptacle. The apical cypsela rim present in Eumor- phia occurs also in other genera, e.g. Phymaspermum. Eumorphia has opposite leaves, which also characterize Gymnopentzia. One species, E. davyi, is aberrant in having alternate, closely set leaves and a conical receptacle. It is provisionally retained in Eumorphia by Kallersjo (1985). The circumscription of this genus obviously needs further consid- eration. E. corymbosa E. Phillips - Note: Description in Phillips, 1950. E. davyi Bolus - Note: Description in Bolus, 1906. E. dregeana DC. - Note: Description in Harvey, 1865. E. prostrata Bolus - Note: Description in Milliard, 1977. E. sericea J. M. Wood & M. Evans - Note: Description in Milliard, 1977. E. swaziensis Compton - Note: Description in Compton, 1976. 5. GYMNOPENTZIA Benth. in Benth. & Hook, f., Gen. pi. 2(1): 537 (1873). Type species: G. bifurcata Benth. A shrub. Leaves opposite, pinnatifid, somewhat ericoid, with secretory cavities. Capitula corymbose, discoid. Receptacle flat or slightly convex, epaleate. Corolla 5-lobed, with a narrow tube and a distinct limb. Anthers with endothecial tissue polarized. Cypselas 10-ribbed, densely long-papillose and thus seemingly pubescent. Pappus absent. Furanoses- quiterpenes present. DISTRIBUTION. S. Africa in the E. Cape, Natal and Trans- vaal, and in Lesotho. - Monotypic. In related genera, i. e. Eumorphia and Phymaspermum, the cypselas are minutely papillose, the epidermis cells being more or less projected. This is especially pronounced in Gymnopentzia, the papillae often being much longer than wide and similar to unicellular hairs. The cypselas are thus often described as pubescent. The opposite leaves is another distinguishing feature from Phymaspermum and possibly synapomorphic with Eumorphia. 6. HYMENOLEPIS Cass. in Bull. Sci. Soc. philom. Paris 1817: 138 (1817). Type species: H. parviflora (L.) DC. - Phaeocephalus S. Moore. Pubescent shrubs; hairs stellate. Leaves alternate, lobed, serrate-dentate or entire, with secretory cavities. Capitula slender, few-flowered, corymbose, sometimes rather densely aggregated, discoid. Receptacle flat, generally paleate, rarely epaleate. Corolla gradually expanded and funnel-shaped, 5-lobed, with short-stalked glands and with continuous veins also in the lobes. Anthers with endothecial tissue polarized. Cypselas 5-10-ribbed, with ellipsoid secretory cavities form- ing longitudinal ducts. Pappus a corona of fimbriate scales. Furanosesquiterpenes present. DISTRIBUTION. S. Africa in the Cape. - 7 spp. Harvey's (1865) treatment of Hymenolepis as a separate genus rather than as a section of Athanasia was recently revived by Kallersjo (1986), since Athanasia including Hyme- nolepis is paraphyletic. Athanasia s. s. is more closely related to Asaemia than to Hymenolepis, as shown in the cladogram (Fig. 2). Hymenolepis differs by its slender, few-flowered 95 capitula and scaly pappus. The monotypic Phaeocephalus was reduced to a synonym of this genus by Kallersjo. A key to the species was provided by Bremer & Kallersjo (1986). H. cynopus Bremer & Kallersjo H. dentata (DC.) Kallersjo (Athanasia schizolepis Harvey) H. gnidioides (S. Moore) Kallersjo (Phaeocephalus gnidio- ides S. Moore) H. incisa DC. H. indivisa (Harvey) Kallersjo H. parviflora (L.) DC. H. speciosa (Hutch.) Kallersjo 7. ATHANASIA L., Sp. pi. 2nd ed.: 1180 (1763). Type species: A. crithmifolia (L.) L. (Bremer & Wijnands, 1982). - Stilpnophyton Less. Glabrous or pubescent shrubs; hairs stellate. Leaves alter- nate, entire, dentate, or lobed, generally ericoid, with secre- tory cavities. Capitula generally corymbose, more rarely solitary, discoid. Receptacle flat, generally paleate, rarely epaleate. Corolla gradually expanded and funnel-shaped, 5-lobed, glabrous or with long-stalked glands, with continu- ous veins also in the lobes. Anthers often with an apical resin sac; endothecial tissue polarized. Cypselas 5-12-ribbed gen- erally with protruding and narrow ribs, glabrous or occasion- ally glandular, with ellipsoid secretory cavities forming longitudinal ducts. Pappus absent or often a pseudopappus of stalked glands. Furanosesquiterpenes present. DISTRIBUTION. S. Africa, mainly in the SW Cape, one species in Natal (A. grandiceps). - 39 spp. Athanasia traditionally embraced all South African, discoid Anthemideae with a paleate receptacle. Epaleate species were placed in other genera, e. g. Pentzia and Stilpnophyton. Athanasia and Stilpnophyton (as well as Asaemia) have similar cypsela wall anatomy, with a continuous ring of sclerified isodiametric cells. Stilpnophyton was reduced to a synonym by Kallersjo (1986). Several former species of Athanasia have now been trans- ferred by Kallersjo to the genera Hymenolepis, Phymasper- mum, and Inulanthera. The remaining part, Athanasia s. s. is a homogeneous and monophyletic group. Characteristically, the cypsela ribs are narrow and somewhat wing-like, and the cypselas are often furnished with a peculiar 'pseudopappus' of long-stalked glands. In some species such glands are also present on the corolla tube, and in others they are totally absent, probably secondarily. In her most recent treatment of Athanasia (Kallersjo, 1991) the distinctive characters of Asaemia are considered to be autapomorphic and A. minuta is considered to be very similar to Athanasia humilis Kallersjo. The list of species is based on Kallersjo (1986, 1991). A. adenantha (Harvey) Kallersjo A. alba Kallersjo A. bremeri Kallersjo A. calophylla Kallersjo A. capitata (L.) L. A. cochlearifolia Kallersjo A. crenata (L.) L. A. crithmifolia (L.) L. A. cuneifolia Lam. A. dentata (L.) L. 96 A. elsiae Kallersjo A. filiformis L. f. A. flexuosa Thunb. A. grandiceps Hilliard & Burtt A. hirsute Thunb. A. humilis Kallersjo A. imbricata Harvey A. inopinata (Hutch.) Kallersjo (Stilpnophyton inopinatum Hutch.) A. juncea (DC.) D. Dietr. A. leptocephala Kallersjo A. linifolia Burm. (Stilpnophyton linifolium (L. f.) Less., Stilpnophyton longifolium (Thunb.) Less.) A. microcephala DC. A. microphylla DC. A. minuta (L. f.) Kallersjo A. oocephala (DC.) Kallersjo (Stilpnophyton oocephalum DC.) A. pachycephala DC. A. pectinata L. f. A. pinnata L. f. A. pubescens (L.) L. A. quinquedentata Thunb. A. rugulosa E. Meyer ex DC. A. scabra Thunb. A. sertulifera DC. A. spathulata (DC.) D. Dietr. A. tomentosa Thunb. A. trifurcata (L.) L. A. vestita (Thunb.) Druce A. virgata Jacq. A. viridis Kallersjo 8. ASAEMIA (Harvey) Harvey ex Benth. in Benth. & Hook, f., Gen. pi. 2(1): 433 (1873). Type species: A. axillaris (Thunb.) Harvey ex Hoffmann (A minuta (L. f.) Bremer). A glabrous, sometimes spiny shrub. Leaves opposite, sheath- ing, entire, ericoid, with secretory cavites. Capitula sessile, solitary along the branches and on lateral branchlets, discoid. Receptacle flat, epaleate. Corolla gradually expanded and funnel-shaped, 5-lobed, glandular, with continuous veins also in the lobes. Anthers with an apical resin sac; endothecial tissue polarized. Cypselas 2-5-angled but generally dorsiven- trally flattened with 1 adaxial and 2 lateral ribs, glabrous or basally with a few hairs, with few ellipsoid secretory cavities forming longitudinal ducts, apically with a smooth or denticu- late thickened rim. Pappus absent. Furanosesquiterpenes present. DISTRIBUTION. S. Africa in the Cape and in Namibia. - Monotypic. Asaemia minuta is habitually very distinct, a shrub with ericoid leaves and small sessile capitula. It is sometimes spinescent (ssp. minuta) and sometimes unarmed (ssp. iner- mis (E. Phillips) Bremer). Asaemia is related to Athanasia. They have similar cypsela wall anatomy with a continuous ring of sclerified isodiametric cells (Kallersjo, 1986). Both Asaemia and most species of Athanasia also have anthers with apical resin sacs. The genus was revised by Bremer (1983) but sunk into Athanasia by Kallersjo (1991). K. BREMER AND C. J. HUMPHRIES 2. CANCRINIINAE Bremer & Humphries, subtrib. nov. Type species: Cancrinia chrysocephala Karelin & Kir. Plantae perennes, herbaceae vel suffruticosae, compactae, plusminusve scaphoideae. Bracteae involucri plerumque mar- gine atrofuscae. Pappus e squamis vel setis planis pluribus, obovatis vel linearibus, albidis vel brunneis, longitudine quam corolla 2-plo brevioribus vel longioribus formatus. Compact, more or less scaphoid perennial herbs or half- shrubs. Leaves alternate to rosulate, pinnatifid to pinnatisect. Capitula solitary, pedunculate, radiate or discoid. Involucral bracts generally with dark brown margins. Receptacle epale- ate, glabrous or sometimes pilose or hirsute. Ray floret limb white, yellow, pinkish or bluish violet. Disc corolla 5-lobed, sometimes pilose. Cypselas 5-15-ribbed, glabrous or pilose. Pappus of several, obovate to linear, whitish or brownish scales or flat bristles, at least half as long as the corolla. DISTRIBUTION (Table 7). genera, 26 spp. Asia, mainly central part. - 6 Table 7 General distribution of Cancriniinae and genera. x=indigenous, o=introduced. C.&E. SW Asia Asia Cancriniinae X X Trichanthemis X Ugamia Richteria X X X Attar dia X Cancrinia X Cancriniella X This subtribe comprises some of the most plesiomorphic representatives of the tribe Anthemideae. In habit species of Trichanthemis and Richteria, and in pappus structure Ugamia and Allardia, are similar to members of the outgroup in tribe Heliantheae. Most species are restricted to mountain habitats in central Asia. Ugamia, Cancriniella and some Trichanthe- mis species, are small, compact, much woody, cushion- formed half-shrubs. Cancrinia and Allardia are compact perennial herbs. This habit character, together with the dark brown-margined involucral bracts, are both considered syna- pomorphies for the genera of the subtribe. It is possible that groups of species in Tanacetum, subtribe Tanacetinae', also sharing these features, are related to members of Cancrinii- nae rather than Tanacetinae', and should be transferred to Cancriniinae. The cladogram for the genera is only one of several equally parsimonious ones, and shown only to display one possible hypothesis of generic interrelationships. The strict consensus tree for the alternative cladograms is totally collapsed. Clades and characters - Fig. 3, Tables 2, 8. Clade Cal - subtribe Cancriniinae 4 Plants compact and more or less scaphoid. Allardia and Cancrinia are compact, more herbaceous perennials with rather short peduncles, whereas the other genera are much GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 97 f t 9 Trichanthemis 10 Ugamia 11 Richteria 12 Allardia 13 Cancrinia 14 Cancriniella Fig. 3. Cladogram (of 21 possible) of the Cancriniinae produced by the ie option in Hennig86. Cladogram length = 14, consistency index = 85, retention index = 60. Table 8 Data matrix for the Cancriniinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. Ill 145057 532718 1 1 1 44483375346 447983504981 11 11 15775555566666175 30563567905678511 9. Trichanthemis I lalll 10. Ugamia al?al? 11. Richteria alaal? 12. Allardia alaal? 1 3 . Cancrinia a 1 ? a 1 ? 14. Cancriniella al?al? lallaOpOOOOO allOOHOOOOO alOOOOOlOOOO alOOOOOHOOO alOOOOHOOOO alOOOOlOOlll ????000000000000p ????000000000000p ????0000000000000 9999999999999?99 Q woody basally, generally with long peduncles. Ugamia has short peduncles. 44 Involucral bracts with dark brown margins. T-his character is absent in some Trichanthemis species. Clade Ca2 147 Cypselas densely pilose; hairs subulate, with a few basal cells and one long apical cell. Cancrinia and some species of Allardia sometimes have sparsely pilose cypselas. Trichanthemis 49 Receptacle densely hirsute. 88 Disc corolla tube pilose. Some species of Trichanthemis have glabrous corollas. Ugamia 33 Capitula on short and nodding peduncles. 35 Capitula discoid. Some species of Trichanthemis, Can- crinia and Cancriniella are also discoid. Clade Ca3 170 Pappus scales brownish. Richteria There is no obvious autapomorphy for this genus, and it appears undefined compared to Allardia and Cancrinia. Allardia 54 Ray floret limb bluish violet. It is not clear if this character occurs in all Allardia species. Some may have whitish rays. Cancrinia 35 Capitula discoid. Cancriniella 35 Capitula discoid. 39 Involucral bracts in 1-2 rows, rather wide. According to Tzvelev (in Komarov, 1961) this character distinguishes Can- criniella from Cancrinia. 48 Receptacle pilose. 161 Pappus of short (not large and obovate or bristle-like) scales, an auricle, a corona, or absent. 9. TRICHANTHEMIS Regel & Schmalh. in Trudy imp. S.-Peterb. hot. Sada 5: 617 (1877). Type species: T. karataviensis Regel & Schmalh. - Glossanthis Polj . Basally much woody half-shrubs with erect annual stems basally covered in sheathing leaf bases. Leaves alternate and basally rosulate, pinnatisect. Capitula solitary, pedunculate, radiate or discoid. Involucral bracts with or without dark brown margins. Receptacle convex, generally densely hir- sute, epaleate. Ray florets female, fertile; limb white, pale pink, or yellow, many-veined; tube generally pilose. Disc corolla 5-lobed, generally pilose; hairs subulate, with a few basal cells and one long apical cell. Cypselas 7-10-ribbed, densely pilose with the same type of hairs as on the florets, sometimes with myxogenic cells. Pappus of 4-12 large, white, linear to oblong scales at least half as long as the corolla. DISTRIBUTION. C. Asia. - 9 spp. Trichanthemis has several unusual features such as the hirsute receptacle and the pilose corollas and cypselas. The relation- ship to Cancrinia and its relatives was noted by Poljakov (1959), who placed several of the present discoid Trichanthe- mis species in Cancrinia. He also defined Trichanthemis as discoid and removed the radiate species to a new genus Glossanthis. His treatment has not been followed by later authors, e.g. Tzvelev in Flora URSS (Komarov, 1961). Tzvelev noted that the radiate T. aurea and T. radiata are related to the discoid T. paradoxes and T. karataviensis, respectively. He also indicated the possible relationship between the problematic Xylanthemum tianshanicum (Pyre- thrum tianshanicuni) and Trichanthemis, notably T. butkovii. Another intergeneric relationship in need of investigation is that of Ugamia and Trichanthemis. Some discoid, small- leaved species of Trichanthemis may be more closely related to Ugamia than to other Trichanthemis species. It appears that Trichanthemis may be paraphyletic. Small segregates, possibly Ugamia for example, or even larger ones in Tanace- tum may have their sister groups within Trichanthemis. The list of species is taken from Flora URSS and Flora iranica but with one former synonym re-established as a species (T. simulans; Pavlov, 1966). T. afghanica Podl. T. aulieatensis (B. Fedtsch.) H. Kraschen. *T. aurea H. Kraschen. *T. butkovii Kovalevsk. T. karataviensis Regel & Schmalh. *T. litwinowii (H. Kraschen) Tzvelev T. paradoxos (Winkler) Tzvelev T. radiata H. Kraschen. & Vved. *T. simulans (Pavlov) Pavlov 98 10. UGAMIA Pavlov in Vest. Akad. Nauk Kazakh. SSR 8:25 (1950). Type species: U. trichanthemoides Pavlov (U. angrenica (H. Kraschen.) Tzvelev). A compact, tomentose, basally much woody half-shrub. Leaves alternate to rosulate, densely set, small, pectinate. Capitula solitary on short, nodding peduncles, discoid. Involucral bracts with dark brown margins. Receptacle almost flat, epaleate. Corolla 5-lobed. Cypselas 10-15- ribbed, densely pilose; hairs subulate, with a few basal cells and one long apical cell. Pappus of 10-15 white linear scales roughly as long as the corolla. DISTRIBUTION. C. Asia. - Monotypic. Ugamia angrenica is very characteristic with its compact and woody habit, small, pectinate leaves and campanulate capitula on short, nodding peduncles. Though discoid as in Cancrinia and Cancriniella, it is possibly most closely related to part of Trichanthemis , which also has discoid representa- tives, with densely pilose cypselas. 11. RICHTERIA Karelin & Kir. in Bull. Soc. Nat. Moscou 15:126 (1842). Type species: R. pyrethroides Karelin & Kir. Tomentose half-shrubs, basally woody with stems covered in sheathing leaf bases. Leaves alternate and basally rosulate, pinnatisect. Capitula solitary, pedunculate, radiate. Involu- cral bracts with dark brown margins. Receptacle convex, epaleate. Ray florets female; limb white. Disc corolla 5-lobed. Cypselas faintly 6-10-ribbed, with sessile glands. Pappus of 6-10 obovate, apically brownish scales at least half as long as the corolla. DISTRIBUTION. Iran, Afghanistan, C. Asia, Mongolia, China in Xinjiang and Tibet, Himalaya. - 3 spp. Richteria is commonly treated as part of Pyrethrum (i.e. Tanacetum s.l.), but since it was originally described as a genus we find it suitable to retain it as such and improve the circumscription of Tanacetum (incl. Pyrethrum) by restricting it to those species with a short coroniform or auriculate pappus. Richteria approaches Trichanthemis in habit; the somewhat pedunculoid stems are basally covered in more or less sheath- ing leaf bases and the leaves are mainly basally arranged. This appears to be a plesiomorphic condition within the tribe, since a similar habit is shown by representatives of the outgroup. The pappus scales are apically brownish as in Cancrinia and Allardia, though in the latter genus they are more narrow and numerous. Being white-rayed rather than discoid or blue-rayed, Richteria is plesiomorphic and appears undefined compared to the latter two genera. Two species of Richteria have been described and one more is added here but there may be more species hidden within Tanacetum. R. djilgense (Franchet) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum djilgense Franchet in Bull. Mus. Hist. nat. Paris 2:345 (1896) (Pyrethrum djilgense (Franchet) Tzvelev, Tanacetum djilgense (Franchet) Podl.). R. leontopodium Winkler R. pyrethroides Karelin & Kir. (Pyrethrum arassanicum (Winkler) O. & B. Fedtsch., Pyrethrum neglectum Tzvelev, K. BREMER AND C. J. HUMPHRIES Pyrethrum pyrethroides (Karelin & Kir.) B. Fedtsch. & H. Kraschen., Pyrethrum transiliense (Herder) Regel & Schmalh., Tanacetum pyrethroides (Karelin & Kir.) Schultz-Bip.) - Note: Synonymy after Kovalevskaja (in Vvedensky, 1962: 133) and Podlech (in Podlech et al., 1986). 12. ALLARDIA Decne in Jacquemont, Voy. Inde 4: 87 (1842-7). Type species: A. tomentosa Decne. - Waldheimia Karelin & Kir. Glabrous to densely tomentose perennial herbs. Leaves alter- nate to rosulate, densely set, pinnatifid. Capitula solitary, pedunculate, radiate. Involucral bracts with dark brown margins. Receptacle convex, epaleate. Ray florets female, fertile or sterile, or neuter; limb white, pink, or bluish-violet. Disc corolla 5-lobed, with a yellow or bluish-violet limb. Cypselas faintly 5-10-ribbed, generally with sessile glands, sometimes pilose. Pappus of many bristle-like, subulate, apically brownish scales as long as or longer than the corolla. DISTRIBUTION. Afghanistan, C. Asia, Mongolia, China in Sinkiang and Tibet, and Himalaya. - 8 spp. With its white to pink and blue-violet florets and cypselas with pappus bristles Allardia may seem out of place in the Anthemideae but it is clearly a member of this tribe. The pappus 'bristles' are subulate, much elongated scales, similar to those of Ugamia (in shape and number) and Richteria and Cancrinia (in being apically brownish). Several narrow bristles are probably plesiomorphic, and apically brownish bristles are probable synapomorphies for Allardia and the latter two genera. Tzvelev in Flora URSS (Komarov, 1961) also stated that the genus is related to Richteria. The publication dates of the various parts of Jacquemont's Voyage dans I' Inde are still in dispute, but it appears that pp. 1-88 of volume 4 were published before 1842 (Stafleu & Cowan, 1979). Allardia is then prior to Waldheimia, which was published in 1842. Allardia was typified by Tzvelev in Flora URSS (Komarov, 1961). He used Waldheimia for the section name and Waldheimia tomentosa (Allardia tomen- tosa) as the type species. *A. huegelii Schultz-Bip. (Waldheimia huegelii (Schultz-Bip.) Tzvelev) A. lasiocarpa (G. X. Fu) Bremer & Humphries, comb. nov. Basionym: Waldheimia lasiocarpa G. X. Fu in Shih & Fu, Acta phytotax. sin. 17: 113 (1979). A. nivea Hook. f. & Thomson ex C. B. Clarke (Waldheimia nivea (C. B. Clarke) Regel) *A. stoliczkae C. B. Clarke (Waldheimia stoliczkae (C. B. Clarke) Ostenf.) A. tomentosa Decne (Waldheimia tomentosa (Decne) Regel, Tanacetum tomentosum (Decne) Muradyan) A. transalaica (Tzvelev) Bremer & Humphries, comb. nov. Basionym: Waldheimia transalaica Tzvelev in Komarov, Ft. URSS 26: 875 (1961). A. tridactylites (Karelin & Kir.) Schultz-Bip. (A. glabra Decne, Tanacetum glabrum (Decne) Muradyan, Waldhe- imia glabra (Decne) Regel, Waldheimia tridactylites Kare- lin & Kir.), Note: A. glabra was reduced by Kovalevskaja (in Vvedensky, 1962:188). A. vestita Hook. f. & Thomson ex C. B. Clarke (Waldheimia vestita (C. B. Clarke) Pampan.) GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 99 13. CANCRINIA Karelin & Kir. in Bull. Soc. Nat. Moscou 15: 124 (1842). Type species: C. chrysocephala Karelin & Kir. Compact tomentose perennial herbs. Leaves alternate to rosulate, densely set, pinnatifid. Capitula solitary, peduncu- late, discoid. Involucral bracts with dark brown margins. Receptacle convex, epaleate. Corolla 5-lobed with a narrow tube and a distinct limb. Cypselas faintly 7-9-ribbed, glabrous or sparsely pilose. Pappus of 5-12 lanceolate, apically brown- ish scales as long as or slightly longer than the corolla. DISTRIBUTION. C. Asia, Mongolia and China in Xinjiang. - 4 spp. Cancrinia, originally described as monotypic, was expanded to include some 20 species, mainly from Tanacetum, by Poljakov (1959) and Tzvelev in Flora URSS (Komarov, 1961). Tzvelev divided the genus into four sections. Most of the species do not belong together with the type species, C. chrysocephala and its sister species, C. tianshanica. Tzvelev's sect. Polychrysum and sect. Tanacetopsis were elevated to genera by Kovalevskaja (in Vvedensky, 1962). The mono- typic section Matricarioides (C. discoided) is transferred to Matricaria. Cancrinia and Allardia are possibly more closely related than indicated by the cladogram. They are similar in habit and cypsela morphology, though Allardia has more and narrower pappus scales. C. chrysocephala Karelin & Kir. *C. krasnoborovii V. Khan. C. pamiralaica (Kovalevsk.) Kovalevsk. *C. tianshanica (H. Kraschen.) Tzvelev 14. CANCRINIELLA Tzvelev in Komarov, Fl. URSS 26:876 (1961). Type species: C. krascheninnikovii (Rubtzov) Tzvelev. A compact, hirsute, basally woody half-shrub. Leaves alter- nate to rosulate, densely set, pectinate. Capitula solitary, long-pedunculate, discoid. Involucral bracts in 1-2 rows, rather wide, subequal. Receptacle convex, sparsely pilose, epaleate. Corolla 5-lobed. Cypselas 10-ribbed. Pappus a corona of scales. DISTRIBUTION. C. Asia. - Monotypic. This monotypic genus is supposed to be related to Cancrinia. Poljakov (1959) stated that it is related to Trichanthemis karataviensis , and he placed both species in Cancrinia. 3. 'TANACETINAE' Bremer & Humphries, subtrib. nov. Type species: Tanacetum vulgar e L. Herbae perennes vel frutices vel raro herbae annuae. Cypse- lae plerumque oblongae et plus quam quinquecostatae. Pap- pus coroniformis e squamis distinctis vel auricula adaxiali formatus, vel nullus. Perennial herbs or shrubs, rarely annuals. Leaves alternate or sometimes rosulate, generally pinnatisect, rarely entire or few-lobed. Capitula solitary or corymbose, generally pedun- culate, radiate, disciform or discoid. Involucral bracts some- times with dark brown margins. Receptacle flat to conical, epaleate or rarely paleate (Heliocauta) . Ray floret limb white, yellow or pink. Disc corolla 5-lobed. Cypselas often oblong and more than 5-ribbed, rarely dorsiventrally flat- tened (Heliocauta), sometimes with sessile glands and myxo- genic cells. Pappus a corona, or of separate scales, or an adaxial auricle, or absent. DISTRIBUTION (Table 9). C. Asia but also in N. America and N. Africa, some Tanacetum species introduced in the S. hemisphere. - 7 genera, 213 spp. The genera of this subtribe, 'Tanacetinae', have no synapo- morphies in common. The subtribe is probably paraphyletic (see for example, Schultz Bipontinus, 18440), and hence put within inverted commas following the convention suggested by Patterson & Rosen (1977) and Wiley (1981). There is still a lot of work to be undertaken on the classification of Anthemideae, and Tanacetum particularly is a key genus of the tribe. At the present state of knowledge we have felt it necessary to adopt a provisional subtribe comprising Tanace- tum and a number of odd genera apparently related to this genus. There are several segregate genera and groups of genera, possibly even whole subtribes, which are related to parts of Tanacetum, which is thus paraphyletic as presently circum- scribed. Apparent examples of such genera are those classi- fied in this subtribe. Subtribes which probably have their sister groups within Tanacetum include Gonosperminae, Handeliinae, and Artemisiinae. The remaining subtribes, excluding Ursiniinae and Cancriniinae, may form one group (as indicated by the cladogram) or a number of groups also with their sister groups within Tanacetum. In fact, Tanacetum is a paraphyletic group basal to large parts of the tribe. The cladogram shown here is the single most parsimonious one derivable from the present data matrix, but the picture may be quite different when Tanacetum is resolved into smaller monophyletic units. The whole complex is in need of detailed investigation, both on the generic and the specific level. Clades and characters - Fig. 4, Tables 2, 10. f=Tal^j= 15 Tanacetum ||= 16 Opisthopappus [P= 17 Tanacetopsis rj= 18 Xylanthemum jJL= 19 Lepidolopha E20 Hippolytia i 1 ?010000pOOpOOOOO ????????????OOOOpOOOOO o o o 18. Xylanthemum a 1 a a 1 ? 1 ? a 1 100000000000 ???0000000000000000000 19 Lepidolopha a 1 9 a 1 1 1 ?011011000pOOOOO ???????????????OOpOOOO (1 o o 20 Hippolytia a 1 9 a 1 "> 1 ?010?OOaalalOOOO ????????????????OOpOOO o o fl 21 Heliocauta a 1 9 a 1 "> 1 ?010?00aal00alll ?0??????????????000000 o o (1 Clade Ta2 35 Capitula discoid. For reasons of parsimony the five genera of this clade are grouped together, though they probably have different sister groups within Tanacetum. The radiate species Xylanthemum tianshanicum is probably misplaced in this genus. Tanacetopsis There is no autapomorphy for this genus. Clade Ta3 2 Plants shrubby. Xylanthemum 166 Pappus adaxially long. Lepidolopha 37 Involucre rather narrowly urceolate. 38 Involucral bracts in 4-7 rows. Clade Ta4 44 Involucral bracts with dark brown margins. This character is present also in several Tanacetum and some Tanacetopsis species. 51 Floral parts with resin canals. Floral resin canals also occur in scattered species of the other genera of the subtribe. 172 Pappus absent in ray and disc cypselas. Hippolytia 29 Capitula densely corymbose. Corymbose capitula are com- mon in Tanacetum and present also in some species of Tanacetopsis and Lepidolopha. Some specialized alpine spe- cies of Hippolytia have solitary capitula. 92 Anthers caudate. Heliocauta 45 Receptacle paleate. 100 Style immersed in a lobed nectary. 114 Cypselas dorsi ventrally flattened. 138 Cypselas with scattered elongated resin sacs. 15. TANACETUM L., Sp. pi.: 1028 (1753). Type species: T. vulgar e L. - Balsamita Miller - Gymnocline Cass. - Hemipappus K. Koch - Pyrethrum Zinn - Spathipappus Tzvelev. Perennial, rarely annual herbs or seldom small, basally woody half-shrubs. Leaves alternate or rarely rosulate, pin- natifid to pinnatisect or rarely entire, serrate. Capitula soli- tary or corymbose, radiate, disciform, or discoid. Involucral bracts sometimes with a dark brown margin. Receptacle flat or convex, epaleate. Ray florets female or neuter; limb yellow, white, or pink. Disc corolla 5-lobed. Cypselas gener- ally oblong, 5-12-ribbed, often with sessile glands, never with myxogenic cells. Pappus a short or well developed corona, sometimes of short free scales, rarely an adaxial auricle (very rarely absent). Embryo sac tetrasporic. DISTRIBUTION. Europe and temperate Asia, also in N. Africa (T. annuum, T. corymbosum) and several species in N. America, some species introduced in the S. hemisphere. - 152 spp. Tanacetum s. 1. is a large and poorly understood boreal genus with many little-known Asian representatives. Variation in habit, foliage, inflorescences, and capitula structure is consid- erable. As presently circumscribed, the genus may be recog- nized by the epaleate receptacle and the non-myxogenic, generally straight and ribbed cypselas with a coroniform or rarely auriculate pappus. Occasional species with epappose cypselas currently placed in Tanacetum are probably mis- GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 101 placed and are likely to be transferred to other genera. One example of a probable misplaced species is T. micro- phyllum, with triangular apical anther appendages and thin- walled, obovoid cypselas with minute pappus scales and no ribs. These characters are reminiscent of subtribe Artemisii- nae, where this species probably should be accommodated. Pyrethrum differs from Tanacetum s. s. by its white or pink rays. Tanacetum s. s. has discoid or disciform capitula or, if radiate, with yellow rays. Tzvelev (in Komarov, 1961) diag- nosed Pyrethrum as white-rayed and Tanacetum as yellow- rayed or disciform (heterogamous). He classified discoid (homogamous) species in Cancrinia. Tzvelev also noted the heterogeneous nature of Pyrethrum and speculated that Can- crinia (s. 1.) evolved from Pyrethrum by loss of rays, where- after Tanacetum (s. s.) evolved from Cancrinia by transformation of the outer, yellow disc florets into short, yellow rays. Except for the often short lamina the yellow rays of Tanacetum (s. s.) are similar to the white or pink ones of Pyrethrum. In other Anthemideae, e.g. Cotula, where pseu- dorays have evolved from the disc florets, their discoid nature is clearly recognized. Also ray colour is very homoplasious within the tribe, many well-defined genera having species with both white and yellow rays. The relationship between Pyrethrum and Tanacetum s. s. is close and involves several sister group relationships between species and groups of species currently classified in the two genera. We follow Heywood in Flora europaea (Tutin et al., 1976; also Hey- wood, 1954) and Grierson in Flora of Turkey (Davis, 1975) in uniting the two genera. Tzvelev (in Komarov, 1961) recognized 14 sections of Pyrethrum and four sections of Tanacetum. Sections Leucan- themopsis and Pyrethrum section Richteria are considered by us as separate genera in different subtribes. Section Balsamita (T. balsamita) is sometimes considered as a separate genus distinguished by its entire, serrate leaves. Hading (1951) in describing the unusual tetrasporic embryo sac development and Favarger (1966) referring to chemistry both recom- mended that Balsamita should be kept separate from Tanace- tum. The actual presence of a tetrasporic embryo sac in Balsamita seems to strengthen its position within Tanacetum (together with the type species T. vulgare and others), though there is a difference in embryo sac development. Chemical evidence is far too scattered to support a removal of Bal- samita from Tanacetum. In morphology Balsamita is similar to several representatives of Pyrethrum sensu Tzvelev and we follow him and Grierson in Flora of Turkey (Davis, 1975) in reducing Balsamita to synonymy. Tzvelev's other sections of Pyrethrum and Tanacetum are a mixture of isolated and, in most cases, vaguely defined entities. Pyrethrum sect. Trichanthemopsis with the single species P. tianshanicum has been transferred to Xylanthe- mum. Other more or less isolated sections are the woody Pyrethrum sect. Xylopyrethrum and Tanacetum sect. Asterot- richa. The woody sections, subtribe Cancriniinae, and the shrubby genera Xylanthemum and possibly also Lepidolopha may be more closely related than indicated by the present classification. Harling (1951) reported tetrasporic embryo sacs in all 10 species of Tanacetum so far investigated (T. balsamita, T. camphoratum, T. cinerariifolium, T. coccineum (Chrysanthe- mum marschallii), Tanacetum corymbosum, T. macrophyl- lum, T. millefolium, T. parthenium, T. roseum and T. vulgare. Interestingly, these species represent nine of Tzvelev's different sections, but not the more isolated ones mentioned above. Tzvelev accepted only heterogamous (disciform or yellow- rayed) species in Tanacetum s. s. Thus the discoid T. argen- teum was accommodated in the genus Hemipappus, further distinguished by its auriculate pappus. On other characters Hemipappus fits well into Tanacetum and it was not accepted as a genus by Grierson in Flora of Turkey, who included not only T. argenteum but also some related Turkish endemic species (T. depauperatum, T. haradjanii, T. tomentellum) in Tanacetum. This solution is followed here, and we include other species with an auriculate pappus in Tanacetum (for- merly Spathipappus , discussed below). Except for the pappus there are no indications that Hemipappus and Spathipappus are closely related sister groups. Spathipappus described by Tzvelev in Flora URSS, is supposed to differ from Pyrethrum by its auriculate pappus and sterile rays. Muradyan (1970) found that the fruits are very similar to those of other Tanacetum species and trans- ferred S. griffithii to Tanacetum. The other two species, S. chitralensis and 5. porphyrostephanus , are recombined here. The list of species is compiled mainly from Flora europaea (Tutin et al., 1976), Flora of Turkey (Davis, 1975), and Flora URSS (Komarov, 1961, including Pyrethrum) with species described later or from other areas added. T. abrotanifolium (L.) Druce. Turkey, Caucasus, Iran. T. abrotanoides Bremer & Humphries, notn. nov. Basionym: Pyrethrum abrotanifolium Bunge ex Ledeb., Fl. ross. 2: 549 (1845) - Note: In Tanacetum the epithet abrotanifolium is already occupied by the preceding species in the list. C. Asia, China in Xinjiang. T. achilleifolium (M. Bieb.) Schultz-Bip. E. Europe, C. Asia. T. akinfievii (Alexej.) Tzvelev. Caucasus. T. alatavicum Herder (Pyrethrum alatavicum (Herder) O. & B. Fedtsch.). C. Asia, Mongolia, China in Xinjiang. T. albipannosum Huber-Mor. & Grierson. Turkey. T. alyssifolium (Bornm.) Grierson. Turkey. T. annuum L. SW Europe in France, Spain, and Portugal, N. Africa in Morocco. *T. arctodzhungaricum (Golosk.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum arctodzhungaricum Golosk. in Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Kazakh. SSR 7: 35 (1971). C. Asia. T. archibaldii Podl. Iran. T. argenteum (Lam.) Willd. (Hemipappus argenteus (Lam.) Tzvelev, Hemipappus canus K. Koch). Turkey, Middle East, Caucasus. *T. argyranthemoides (Boiss. & Kotschy) Schultz-Bip. Iran. T. armenum (DC.) Schultz-Bip. (Pyrethrum heldreichianum Fenzl ex Tchich.). Turkey. T. aromaticum (Tzvelev) Bremer & Humphries, comb. nov. - Basionym: Pyrethrum aromaticum Tzvelev in Komarov, Fl. URSS 26: 222 (1961). - Note: the name Pyrethrum aromati- cum must be considered a new species described by Tzvelev, and not a new combination based on Tripleuro- spermum aromaticum Rupr. ex Boiss. as stated by Tzvelev. That name was only published pro syn. by Boissier (1875: 334). Caucasus. *T. artemisioides Schultz-Bip. in Hook. f. Himalaya. *T. atkinsonii (C. B. Clarke) Kitam. (Pyrethrum atkinsonii (C. B. Clarke) Ling & Shih). China, Himalaya. T. aucheranum (DC.) Schultz-Bip. (Pyrethrum aucheranum DC.). Turkey, Caucasus. 102 K. BREMER AND C. J. HUMPHRIES T. aucheri DC. Turkey, Syria, Lebanon, Israel. T. audibertii (Req.) DC. Sardinia. T. balsamita L. (Balsamita major Desf., Pyrethrum balsamita (L.) Willd., Pyrethrum balsamitoides (Nab.) Tzvelev, Pyre- thrum majus (Desf.) Tzvelev). SW Asia from Turkey, Caucasus, Iran, and Afghanistan, also widely cultivated and naturalized. T. bamianicum Podl. Afghanistan. *T. bardayanum DC. (T. turlanicum (Pavlov) Tzvelev). C. Asia, China in Xinjiang. T. bipinnatum (L.) Schultz-Bip. NE Europe, Siberia, Alaska. T. boreale Fischer ex DC. Siberia, C. Asia and Far East, China, Korea, Alaska, Canada. T. budjurnense (Rech. f.) Tzvelev. Iran. *T. cadmeum (Boiss.) Heyw. Turkey. T. camphoratum Less. Western N. America from British Columbia S. to California. T. canescens DC. (T. gilliatii (Turrill) Parsa, T. modestum (Heimerl ex Stapf) Parsa). Turkey, Caucasus, Iran. T. cappadodcum (DC.) Schultz-Bip. Turkey. T. changaicum (H. Kraschen. ex Grubov) Bremer & Humphries, comb. nov. Basionym: Pyrethrum changaicum H. Kraschen. ex Grubov in Bot. Mater. Gerb. hot. Inst. V. A. Komorova 27: 23 (1955). Mongolia. T. chiliophyllum (Fischer & C. Meyer) Schultz-Bip. (T. heimerli (Nab.) Parsa, T. oligocephalum (DC.) Schultz- Bip.). Turkey, Caucasus, Iraq, Iran. T. chitralense (Podl.) Bremer & Humphries, comb. nov. Basionym: Spathipappus chitralensis Podl., Fl. iranica 158: 151 (1986). Pakistan. T. dlidum (Boiss.) Grierson. Turkey, Middle East, Iraq. T. dnerariifolium (Trevir.) Schultz-Bip. (Pyrethrum dnerarii- folium Trevir). SE and E. Europe, Caucasus, C. Asia, China, often cultivated as the source of the insecticide pyrethrin. T. cocdneum (Willd.) Grierson (Pyrethrum cocdneum (Willd.) Vorosch., Pyrethrum chamaemellfolium (Sommier & Levier) Sosn.). Turkey, Caucasus, Iran, often cultivated as the garden 'Pyrethrum'. T. corymbiforme (Tzvelev) Bremer & Humphries, comb, nov. Basionym: Pyrethrum corymbiforme Tzvelev in Komarov, FL URSS 26: 873 (1961). C. Asia, China in Xinjiang. T. corymbosum (L.) Schultz-Bip. (Pyrethrum corymbosum (L.) Willd., Pyrethrum clusii Fischer ex Reichb.). N. Africa in Morocco and Algeria, most of Europe except N. parts, Turkey, Caucasus, C. Asia. *T. crassipes (Stchegl.) Tzvelev. C. Asia, China in Xinjiang. T. daghestanicum (Rupr. ex Boiss.) Bremer & Humphries, comb. nov. Basionym: Chamaemelum daghestanicum Rupr. ex Boiss., Fl. orient. 3: 334 (1875) (Pyrethrum daghestanicum (Rupr. ex Boiss.) Rupr. ex Flerov). Cauca- sus. *T. demetrii (Manden.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum demetrii Manden. in Zametki Sist. Geogr. Rast. 22: 60 (1961). Caucasus. T. densum (Labill.) Schultz-Bip. Turkey. T. depauperatum (Post) Grierson. Turkey. T. dolomiticum (Galushko) Bremer & Humphries, comb, nov. Basionym: Pyrethrum dolomiticum Galushko in Novit. Syst. PI. Vase. Acad. Sci. URSS 6: 218 (1970). Caucasus. *T. dumosum Boiss. Iran. T. eginense (Hausskn. ex Bornm.) Grierson. Turkey. *T. elongatum (Bornm. & Gauba) Parsa. Iran. *T. falcatolobatum H. Kraschen. (Cancrinia maximoviczii Winkler). China. T. falconeri Hook. f. Himalaya. T. ferulaceum (Webb ex Berth.) Schultz-Bip. Canary Islands. *T. funkii Schultz-Bip. ex Willk. SW Europe in Spain. *T. galae (Popov) Nevski (Pyrethrum galae Popov). C. Asia. T. galushkoi (Prima) Bremer & Humphries, comb. nov. Basionym: Pyrethrum galushkoi Prima in Nov. Sist. Vysshikh Rast. 11: 277 (1974). Caucasus. T. germanicopolitanum (Bornm. & Heimerl) Grierson. Tur- key. T. ghoratense Podl. Afghanistan. T. glanduliferum (Sommier & Levier) Bremer & Humphries, comb. nov. Basionym: Pyrethrum glanduliferum Sommier & Levier, Decas Comp. Nov. Cauc. 87 (1895). Caucasus. *T. griff ithii (C. B. Clarke) Muradyan (Spathipappus griff ithii (C. B. Clarke) Tzvelev). Afghanistan, C. Asia, Himalaya. T. grossheimii (Sosn.) Muradyan (Pyrethrum grossheimii Sosn.). Caucasus, Iran. T. haradjanii (Rech. f.) Grierson. Turkey. *T. haussknechtii (Bornm.) Grierson. Turkey. *T. herderi Regel & Schmal. SW Asia. *T. heterotomum (Bornm.) Grierson. Turkey. T. hissaricum (H. Kraschen.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum hissaricum H. Kraschen. in Feddes Reprium 26: 26 (1929). C. Asia. T. hololeucum (Bornm.) Podl. Iran. T. huronense Nutt. N. America in Alaska, Canada and Michigan. T. karelinianum Bremer & Humphries, nom. nov. Basionym: Pyrethrum karelinii H. Kraschen. in Nov. Sist. Vysshikh Rast. 9: 157 (1946). C. Asia. *T. karelinii Tzvelev. C. Asia. T. kaschgarianum Bremer & Humphries, nom. nov. Basionym: Pyrethrum kaschgaricum H. Kraschen. in Nov. Sist. Vysshikh Rast. 9: 158 (1946) (non T. kaschgaricum H. Kraschen.). China. T. kelleri (Krylov & Plotn.) Takht. (Chrysanthemum kelleri Krylov & Plotn., Pyrethrum kelleri (Krylov & Plotn.) H. Kraschen.). C. Asia. T. khorassanicum (H. Kraschen.) Parsa (T. czerniakowskae (H. Kraschen.) Parsa). Iran. T. kotschyi (Boiss.) Grierson (Pyrethrum kotschyi Boiss.). Turkey, Caucasus, Iraq, Iran. T. krylovianum (H. Kraschen.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum krylovianum H. Kraschen. in Bot. Mater. Gerb. hot. Inst. V. A. Komorova9: 155 (1946). C. Asia, China in Xinjiang. *T. kubense (Grossh.) Muradyan (Pyrethrum kubense Grossh.). Caucasus. *T. lanuginosum Schultz-Bip. & Herder (Pyrethrum lanugi- nosum (Schultz-Bip. & Herder) Tzvelev). E. Siberia, Mon- golia. T. leptophyllum (Steven) Schultz-Bip. (Pyrethrum leptophyl- lum Steven). Caucasus. *T. longipedunculatum (Sosn.) Tzvelev. Caucasus. *T. macrocephalum Pampan. Himalaya. T. macrophyllum (Waldst. & Kit.) Schultz-Bip. (Pyrethrum macrophyllum (Waldst. & Kit.) Willd.). C. and SE Europe, Turkey, Caucasus. T. marionii (Albov) Bremer & Humphries, comb. nov. Basionym: Pyrethrum marionii Albov in Bull. Herb. Bois- sier3: 92 (1895). Caucasus. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE T. maymanense Podl. Afganistan. T. microphyllum DC. SW Europe in Spain and Portugal. *T. mikeschinii (Tzvelev) Takht. (Pyrethrum mikeschinii Tzvelev). C. Asia. T. millefolium (L.) Tzvelev (T. kittaryanum (C. Meyer) Tzvelev). E. Europe, S. Siberia, C. Asia. *T. mucroniferum Huber-Mor. & Grierson. Turkey. T. mucronulatum (Hoffsgg & Link) Heyw. SW Europe in Portugal. T. nitens (Boiss. & Noe) Grierson. Turkey. T. nivale Schultz-Bip. Iraq. T. niveum (Lagasca) Schultz-Bip. (Pyrethrum fruticulosum Biehler). Caucasus. T. nuristanicum Podl. Afghanistan. *T. odessanum (Klokov) Tzvelev. E. Europe. T. oltense (Sosn.) Grierson. Turkey. T. ordubadense (Manden.) Bremer & Humphries, comb, nov. Basionym: Pyrethrum ordubadense Manden. in Nov. Sist. Vysshikh Rast. 19: 358 (1959). Caucasus. T. oxylepis (Bordz.) Grierson. Caucasus, Turkey. T. oxystegium (Sosn.) Grierson. Turkey. *T. paczoskii (Zef.) Tzvelev. Krym. T. pakistanicum Podl. Pakistan. T. paleaceum Podl. Afghanistan. *T. paradoxum Bornm. Iran. T. parthenifolium (Willd.) Schultz-Bip. (Pyrethrum partheni- folium Willd.). SW Asia from Turkey to Caucasus, Iran, Aghanistan, and C Asia. T. parthenium (L.) Schultz-Bip. (Pyrethrum par thenium (L.) Smith). N. Africa, SE and E. Europe, SW Asia from Turkey to Caucasus, Iran, Aghanistan, and C. Asia, also widely naturalized. *T. petiolosum Pampan. Himalaya. T. petrareum (Shih) Bremer & Humphries, comb. nov. Basionym: Pyrethrum petrareum Shih in Bull. hot. Lab. n.-eastFor. Inst. 6: 10 (1980). China. *T. peucedanifolium (Sosn.) Bremer & Humphries, comb, nov. Basionym: Pyrethrum parthenifolium Willd. var. peu- cedanifolium Sosn. in Trudy tiflis. hot. Sada 17: 35 (1915) (Pyrethrum peucedanifolium (Sosn.) Manden.). Caucasus. *T. pinnatum Boiss. (T. flavovirens (Boiss.) Tzvelev, T. tamrutense (Sosn.) Sosn.). Turkey, Caucasus, Iraq, Iran. T. podlechii Bremer & Humphries, nom. nov. Basionym: Pyrethrum komarovii Sosn. in Dokl. Akad. Nauk armyan. SSR 2 (4): 119 (1945) (non Tanacetum komarovii (Winkler) Muradyan). Caucasus. T. polycephalum Schultz-Bip. (T. argyrophyllum (K. Koch) Tzvelev, T. duderanum (Boiss.) Tzvelev, T. heterophyllum Boiss., T. junesarense (Bornm.) Parsa, T. myriophyllum Willd.). Turkey, Caucasus, Iraq, Iran. T. porphyrostephanum (Rech. f.) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum porphyrostepha- num Rech. f., Symb. Afghan. 2: 47 (1955) (Spathipappus porphyrostephanus (Rech. f.) Tzvelev). Iran. T. poteriifolium (Ledeb.) Grierson (Pyrethrum poteriifolium Ledeb.). Turkey, Caucasus. T. praeteritum (Horw.) Heyw. Turkey. T. pseudoachillea Winkler. C. Asia. T. ptarmidflorum (Webb & Berth.) Schultz-Bip. Canary Islands. *T. pulchellum (Turcz.) Schultz-Bip. (Pyrethrum pulchellum Turcz.). E. Siberia. T. pulchrum (Ledeb.) Schultz-Bip. (Pyrethrum pulchrum Ledeb.). C. Asia, Mongolia, China in Xinjiang. 103 T. punctatum (Desr.) Grierson (Pyrethrum punctatum (Desr.) Bordz. ex Sosn.). Turkey, Caucasus. *T. richterioides (Winkler) Bremer & Humphries, comb, nov. Basionym: Chrysanthemum richterioides Winkler in Trudy imp. S.-Peterb. hot. Sada 10: 86 (1887) (Pyrethrum richterioides (Winkler) H. Kraschen.). China. *T. robustum Hook. f. & Thomson ex C. B. Clarke. Hima- laya. T. roseum (Adams) Schultz-Bip. (Pyrethrum roseum (Adams) M. Bieb.). Caucasus. T. roylei (DC.) Podl. W. Himalayas. T. salsugineum Podl. Iran. *T. sanguineum (Parsa) Parsa. Iran. T. santolina Winkler. S. European USSR, S. Siberia. C. Asia, China in Xinjiang. *T. saxicolum (H. Kraschen.) Tzvelev. C. Asia. T. sderophyllum (H. Kraschen.) Tzvelev. S. European USSR. *T. scopulorum (H. Kraschen.) Tzvelev. C. Asia, China in Xinjiang. T. semenovii Herder (Pyrethrum semenovii (Herder) Winkler ex O. & B. Fedtsch.). C. Asia. T. sericeum (Adams) Schultz-Bip. (Pyrethrum sericeum (Adams) M. Bieb.). Turkey, Caucasus. T. sevanense (Sosn.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum sevanense Sosn. ex Gross., Fl. Kauk. 4: 137 (1934). Caucasus, Iran. T. silaifolium (Steven) Schultz-Bip. (Pyrethrum silaifolium Steven). Caucasus. T. silvicola Podl. Afghanistan. T. sinaicum (Fresen.) Del. ex Bremer & Humphries, comb, nov. Basionym: Santolina sinaica Fresen., Mus. senckenb. 1: 83 (1833) (Pyrethrum santolinoides DC.) - Note: The combination Tanacetum sinaicum has hitherto not been validly published; it was only published pro syn. by de Candolle (1837: 59). Middle East in Sinai. *T. sipikorense (Bornm.) Grierson (Pyrethrum oxylepis (Bordz.) Tzvelev). Turkey, Caucasus. T. songaricum (Tzvelev) Bremer & Humphries, comb. nov. Basionym: Pyrethrum songaricum Tzvelev in Komarov, Fl. URSS 26: 874 (1961). C. Asia. T. sorbifolium (Boiss.) Grierson (Pyrethrum sorbifolium Boiss.). Turkey, Caucasus. *T. stapfianum (Rech. f.) Podl. Iran. T. tabrisianum (Boiss.) Sosn. & Takht. Turkey, Caucasus, Iran. T. tanacetoides (DC.) Tzvelev. C. Asia, Mongolia, China in Xinjiang. T. tatsienense (Bureau & Franchet) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum tatsienense Bureau & Franchet in /. Bot. 5: 72 (1891) (Pyrethrum tatsienense (Bureau & Franchet) Ling ex Shih). China in Tibet. T. tenuisectum (Boiss.) Podl. Iran. T. tenuissimum (Trautv.) Gross. Caucasus, Iran, Afghani- stan. T. tirinense Podl. Afghanistan. T. tomentellum (Boiss.) Grierson. Turkey. T. trichophyllum (Sosn.) Bremer & Humphries, comb. nov. - Basionym: Pyrethrum trichophyllum Sosn. in Zametki Sist. Geogr. Rast. 15: 2 (1949) (non Griseb.) (Pyrethrum tri- cholobum Sosn. ex Manden.). Caucasus. T. trifoliolatum Podl. Iran. *T. turcomanicum (H. Kraschen.) Tzvelev. Iran, C. Asia. *T. ulutavicum Tzvelev. C. Asia. 104 K. BREMER AND C. J. HUMPHRIES T. uniflorum (Fischer & C. Meyer ex DC.) Schultz-Bip. Turkey, Caucasus, Iran. *T. uralense (H. Kraschen.) Tzvelev. E. Europe, S. Siberia, C. Asia. *T. vahlii DC. SW Europe in Spain. T. vulgare L. Throughout most of Europe and temperate Asia, introduced in America, Australia and New Zealand. T. walteri (Winkler) Tzvelev. Iran, C. Asia. *T. willkommii Schultz-Bip. SW Europe in Spain. *T. yabrudae (Mout.) Charpin & Dittrich (Pyrethrum yabrudae Mout.). Middle East in Syria. T. zahlbruckneri (Nab.) Grierson. Turkey. 16. OPISTHOPAPPUS Shih in Acta phytotax. sin. 17: 110 (1979). Type species: O. taihangensis (Ling) Shih. Perennial herbs with basally somewhat woody stems. Leaves alternate, pinnatisect. Capitula solitary, radiate. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb white or pinkish, many-veined. Disc corolla 5-lobed, with sessile glands. Cypselas obovoid, 3-5-ribbed, thin-walled, with myxogenic cells. Pappus of separate, mainly abaxial, subulate scales. DISTRIBUTION. NE central China. - 2 spp. The position of this genus is unclear. It appears to be a derivative of Tanacetum, including Pyrethrum, where the type species was formerly accommodated. According to Shih it differs from Pyrethrum and Spathipappus (here included in Tanacetum) by its thickly ribbed cypselas and by its mainly abaxial pappus of separate scales, rather than a corona or an auricle. The cypsela ribs as such are not thick but since they are invested with large myxogenic cells, dry cypselas appear thickly ribbed. The presence of myxogenic cells is indeed a character distinguishing Opisthopappus from Tanacetum, which has non-myxogenic cypselas. O. longilobus Shih O. taihangensis (Ling) Shih 17. TANACETOPSIS (Tzvelev) Kovalevsk. in Vvedensky, Fl. uzbekistana 6: 138 (1962). Type species: T. mucronata (Regel & Schmalh.) Kovalevsk. - Cancrinia sect. Tanacetopsis Tzvelev. Perennial herbs or basally woody half-shrubs. Leaves alter- nate, pinnatisect. Capitula solitary or generally laxly to densely corymbose, discoid. Involucral bracts sometimes with dark brown margins. Receptacle convex, epaleate, rarely sparsely pilose. Corolla 5-lobed. Cypselas 5-ribbed, some- times with sessile glands, generally with myxogenic cells. Pappus a corona of free or united scales. DISTRIBUTION. Iran, Afghanistan and C. Asia. - 21 spp. Tanacetopsis is based on a section of Cancrinia described by Tzvelev (1971) and in Komarov (1961). Tzvelev transferred a number of Tanacetum and Lepidolopsis species to Cancrinia, although he noted that his Cancrinia sections could be treated as separate genera. He argued that their relationship was shown by similar floret, cypsela, and pollen morphology. The type species of Cancrinia (see below) is very different from Tanacetopsis species, in habit as well as floret and cypsela morphology. The removal of Tanacetopsis from Cancrinia is well justified (Kovalevskaja, 1972), but we doubt its status as a genus separate from Tanacetum, at least as presently circumscribed. Tzvelev noted that some species of Cancrinia, i.e. Tanac- etopsis, have a sparsely pilose receptacle, a character which does not occur in Tanacetum. Since only some species are involved, however, it cannot be used as a synapomorphy for the genus. The alleged difference in shape of involucre, also noted by Tzvelev, campanulate in Cancrinia and Tanacetopsis and cyathiform in Tanacetum, breaks down after examination of the numerous species involved. Tzvelev also noted that his monotypic section Leptanthe- mum of Pyrethrum (Tanacetum leptophyllum) is almost iden- tical to some species of Cancrinia sect. Tanacetopsis (e.g. the type species) except in being radiate rather than discoid. In fact, Tanacetopsis differs from Tanacetum simply by homoga- mous (discoid) as opposed to heterogamous capitula. With the inclusion of Hemipappus in Tanacetum there are also discoid representatives of Tanacetum. Podlech (in Podlech et al., 1986) included Tanacetopsis as a section of Tanacetum. Since we have seen only limited material of Tanacetopsis and since this whole Tanacetum group is in need of generic revision, we retain Tanacetopsis provisionally as a genus. T. afghanica (Gilli) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum afghanicum Gilli in Feddes Reprium Spec. nov. veg. 68: 93 (1963) (Tanacetum afghani- cum (Gilli) Podl.) *T. botschantzevii (Kovalevsk.) Kovalevsk. (Cancrinia botschantzevii (Kovalevsk.) Tzvelev) T. doabensis (Podl.) Bremer & Humphries, comb. nov. Basionym: Tanacetum doabense Podl., Fl. iranica 158: 131 (1986). *T. eriobasis (Rech. f.) Kovalevsk. (Tanacetum eriobasis (Rech. f.) Kovalevsk.) *T. ferganensis (Kovalevsk.) Kovalevsk. (Cancrinia ferganen- sis (Kovalevsk.) Tzvelev) T. freitagii (Podl.) Bremer & Humphries, comb. nov. Basionym: Tanacetum freitagii Podl., Fl. iranica 158: 132 (1986). T. golovskovii (Polj.) Karmysch. (Cancrinia golovskovii (Polj.) Tzvelev) T. hedgei (Podl.) Bremer & Humphries, comb. nov. Basionym: Tanacetum hedgei Podl., Fl. iranica 158: 130 (1986). *T. kamelinii Kovalevsk. T. karataviensis (Kovalevsk.) Kovalevsk. (Cancrinia karatav- ica Tzvelev) * T. kjurendaghii Kurbanov *T. krascheninnikovii (Nevski) Kovalevsk. (Cancrinia nevskii Tzvelev) T. mucronata (Regel & Schmalh.) Kovalevsk. *T. pjataeviae (Kovalevsk.) Karmysch. (Cancrinia pjataeviae (Kovalevsk.) Tzvelev) *T. platyrachis (Boiss.) Kovalevsk. T. santoana (H. Kraschen., Popov & Vved.) Kovalevsk. (Cancrinia santoana (H. Kraschen., Popov & Vved.) Tzvelev) *T. setacea (Regel & Schmalh.) Kovalevsk. (Cancrinia seta- cea (Regel & Schmalh.) Tzvelev) T. submarginata (Kovalevsk.) Kovalevsk. (Cancrinia sub- marginata (Kovalevsk.) Tzvelev) *T. subsimilis (Rech. f.) Kovalevsk. (Cancrinia subsimilis GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 105 (Rech. f.) Tzvelev, Tanacetum subsimile (Rech. f.) Kova- levsk.) *T. tripinnatifida (Oliver) Kovalevsk. T. urgutensis (Popov) Kovalevsk. (Cancrinia urgutensis (Popov) Tzvelev) 18. XYLANTHEMUM Tzvelev in Komarov, Fl. URSS 26: 877 (1961). Type species: X. fisherae (Aitch. & Hemsley) Tzvelev. Shrublets generally with virgate eventually leaf-less stems, sometimes much woody basally. Leaves alternate, pinnati- sect. Capitula solitary, pedunculate, discoid or possibly also radiate with white rays. Involucral bracts in several rows, the outer much smaller than the inner. Receptacle flat, epaleate. Corolla 5-lobed. Cypselas oblong, 5-6-ribbed, generally with sessile glands and myxogenic cells. Pappus an adaxial auricle or a corona of several adaxially more developed scales. DISTRIBUTION. Iran, Afghanistan and C. Asia. - 9 spp. According to Tzvelev (in Komorov, 1961) Xylanthemum differs from Tanacetum by its homogamous (discoid) capitula, flat receptacle, and auriculate pappus. However, these characters occur also in Tanacetum with Hemipappus and Spathipappus included (see discussion under Tanace- tum). Furthermore, X. pamiricum has a pappus of free scales developed more strongly on the adaxial side, probably a plesiomorphic condition when compared to the auriculate pappus in X. fisherae. Thus, the genus as a whole has not a pappus of a single auricle but at most, as expressed by Tzvelev, an auricle sometimes cleft to the base. Nevertheless, it is possible that Xylanthemum as circum- scribed by Tzvelev is a monophyletic group defined not only by these characters as synapomorphies, but also by its shrubby habit and involucral bracts in several rows. Tzvelev also stated that the species are closely related geographical- ecological races of one natural unit. Muradyan (1970) investi- gated fruits of Tanacetum and Xylanthemum and concluded that they are different and that Spathipappus belongs in Tanacetum rather than together with Xylanthemum, despite the similarity in pappus structure. The relationship of Xylan- themum to Tanacetum or probably rather to a part of Tanacetum, which then becomes paraphyletic by the exclu- sion of Xylanthemum, deserves further investigation. Lepi- dolopha with a similar virgate shrubby habit is clearly a related genus. The list of species is compiled from Flora URSS (Komarov, 1961) with one radiate species transferred from Pyrethrum by Muradyan (1970) added. The position of this species, X. tianshanicum, within Xylanthemum is provisional. According to Tzvelev it is more closely related to Trichanthemis , and differs from that genus simply by its glabrous cypselas. Tzvelev placed it in the monotypic Pyrethrum section Trichanthemopsis. Muradyan, when investigating cypsela morphology, transferred it to Xylanthemum mainly because of the presence of myxogenic cells on the cypselas. In other characters, e.g. of pappus and habit, the species appears more close to Trichanthemis or Richteria. Podlech in Podlech et al. (1986) considers this genus as a section of Tanacetum. X. fisherae (Aitch. & Hemsley) Tzvelev X. gilletii (Podl.) Bremer & Humphries, comb. nov. Basionym: Tanacetum gilletii Podl., Fl. iranica 158: 140 (1986). X. lingulatum (Boiss.) Bremer & Humphries, comb. nov. Basionym: Pyrethrum lingulatum Boiss., Fl. orient. 3: 357 (1875). X. macropodum (Hemsley & Lace) Bremer & Humphries, comb. nov. Basionym: Tanacetum macropodum Hemsley & Lace in J. Linn. Soc. 28: 324 (1891). X. paghmanense (Podl.) Bremer & Humphries, comb. nov. Basionym: Tanacetum paghmanense Podl., Fl. iranica 158: 141 (1986). X. pamiricum (Hoffm.) Tzvelev X. polycladum (Rech. f.) Tzvelev X. rupestre (Popov ex Nevski) Tzvelev X. tianshanicum (H. Kraschen.) Muradyan (Pyrethrum tians- hanicum H. Kraschen.) 19. LEPIDOLOPHA Winkler in Trudy imp. S. Peterb. hot. Sada 13: 236 (1894). Type species: L. komarovii Winkler. Shrublets with virgate, basally sometimes much woody stems. Leaves alternate or basally on the woody stems also fascicu- late on brachyblasts, entire or 3-lobed. Capitula solitary or corymbose, discoid. Involucre rather narrowly urceolate; involucral bracts in 4-7 rows, the outer much smaller than the inner. Receptacle flat to convex, epaleate. Corolla 5-lobed; tube basally swollen in fruit. Cypselas oblong, 6-10-ribbed, with sessile glands. Pappus of many subulate scales. DISTRIBUTION. C. Asia. - 9 spp. Lepidolopha is well characterized, for example by the entire to 3-lobed leaves and the narrowly urceolate capitula with involucral bracts in several rows. The species are closely related with vicarious distributions and they differ mainly in foliage and inflorescence characters. A possibly related genus is Xylanthemum, also shrubby and with involucral bracts in several rows, though with a wider involucre and different pappus. Apart from the treatment by Knorring in Flora URSS (Komarov, 1961), there is also a more detailed discussion by the same author (Knorring, 1959). L. fedtschenkoana Knorr. *L. filifolia Pavlov *L. gomolitzkii Kovalevsk. & Safralieva L. karatavica Pavlov *L. komarovii Winkler (Tanacetum komarovii (Winkler) Muradyan) L. kraschenennikovii Kovalevsk. & Safralieva L. mogoltavica (H. Kraschen.) H. Kraschen. L. nuratavica H. Kraschen. (Tanacetum nuratavicum (H. Kraschen.) Muradyan) *L. talasica Kovalevsk. & Safralieva 20. HIPPOLYTIA Polj. in Bot. Mater. Gerb. hot. Inst. V. A. Komarova 18: 288 (1957). Type species: H. darvasica (Winkler) Polj. Pubescent, sometimes densely tomentose perennial herbs. Leaves alternate and basally more or less rosulate, much pinnatisect. Capitula densely corymbose to glomerulate or rarely solitary, discoid. Involucral bracts with dark brown margins. Receptacle flat to convex, epaleate. Corolla 5-lobed. Anthers tailed. Cypselas 5-8-ribbed, sometimes with 106 K. BREMER AND C. J. HUMPHRIES resin canals, with sessile glands, with a more or less distinct apical rim. Pappus absent. DISTRIBUTION. C. Asia, Mongolia, China in Xinjiang and Tibet, and Himalaya (map by Shih, 1979: 68). - 19 spp. Poljakov (1957) discussed the wide circumscription and loose definition of Tanacetum (then including also Ajanid) and he found it difficult to accept both homogamous (discoid) and heterogamous (disciform) species in the same genus. Conse- quently Poljakov removed a number of central Asian discoid Tanacetum species to a new genus, Hippolytia. The cypselas have no pappus as opposed to the Tanacetum species. The species have a characteristic habit with much pinnatisect leaves, pedunculoid stems with crowded (rarely solitary) capitula, and involucral bracts with dark brown margins. Some high-altitude species have densely tomentose leaves crowded on vegetative shoots and short flowering stems. This alpine habit is best explained as a secondary development within the genus. Tzvelev in Flora URSS (Komarov, 1961) stated that Hip- polytia is related to Dendranthema and Ajania and that Hippolytia is intermediate between these genera, a view analogous to the position of 'Cancrinia' (s. 1. incl. Tanacetop- sis) between Pyrethrum and Tanacetum s. s. According to Tzvelev radiate species of Dendranthema (cf. Pyrethrum) evolved into discoid species of Hippolytia (cf. Tanacetopsis) , which further evolved into disciform species of Ajania (cf. Tanacetum s. s.). Podlech et al. (1986) included Hippolytia as a section of Tanacetum. In our opinion Hippolytia, probably together with Heliocauta, is related to part of Tanacetum as discussed under Heliocauta. Hippolytia is retained here pending a reinvestigation of the whole Tanacetum complex. It is pos- sible that there are species of Ajania that should be trans- ferred to Hippolytia. Tzvelev removed some of Poljakov's species of Hippolytia to Ajania. Shih (1979) revised the genus and added a number of Chinese species. The list of species is taken from Shih's paper but with H. alashanensis listed as a synonym. H. crassicollum (Rech. f.) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum crassicollum Rech. f. in Biol. Skr. 8 (2): 43 (1955) (Tanacetum crassicollum (Rech. f.) Podl.) H. darvasica (Winkler) Polj . H. delavayi (W. Smith) Shih H. desmantha Shih H. dolichophylla (Kitam.) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum dolichophyllum Kitam. in Acta. phytotax. geobot. Kyoto 23: 73 (1968) (Tanacetum dolichophyllum (Kitam.) Kitam.) *H. glomerata Shih H. gossypina (C. B. Clarke) Shih H. herderi (Regel & Schmalh.) Polj. H. kaschgarica (H. Kraschen.) Polj. (H. alashanensis (Ling) Shih) - Note: In recent treatments of Hippolytia (Shih, 1979) H. kaschgarica has disappeared. It was based on Tanacetum kaschgaricum H. Kraschen. (1933) from Tien- Shan range, Mount Ischma (China in Xinjiang). Material of this species in St. Petersburg, annotated by Kraschenin- nikov, appears to be the species currently known as Hip- polytia alashanensis (Ling) Shih originally described as Tanacetum alashanense Ling from Ala-Shan, Inner Mongo- lia (China). At least, they must be closely related with the shrubby, subspinose habit (see Ling, 1983: plate 13,1). Furthermore, H. kaschgarica which is equal to H. alashan- ensis is aberrant in Hippolytia and thus deserves further study. *H. kennedyi (Dunn) Ling H. longifolia (Wallich) Shih (Tanacetum himachalense Aswal & Mehrotra) H. megacephala (Rupr.) Polj. *H. nana (C. B. Clarke) Shih H. schugnanica (Winkler) Polj. *//. senecionis (Besser) Polj. *H. syncalathiformis Shih H. tomentosa (DC.) Tzvelev *//. trifida(Turcz.)Po\]. H. yunnanensis (Jeffrey) Shih 21. HELIOCAUTA Humphries in Bot. Notiser 130: 155 (1977). Type species: H. atlantica (Litard. & Maire) Humphries. A perennial rosulate or creeping herb. Leaves in a basal rosette, pinnatisect. Capitula solitary, pedunculate, discoid. Involucral bracts with dark brown margins. Receptacle coni- cal, paleate. Corolla 5-lobed; lobes soon brownish. Style immersed in a lobed nectary. Cypselas somewhat dorsiven- trally flattened, 4-5-ribbed with 2 major lateral ribs, with scattered elongated resin sacs and with an apical erose rim. Pappus absent. DISTRIBUTION. N. Africa in Morocco. - Monotypic. This recently described monotypic genus is difficult to place and as stated by Humphries (1977) it is not related to Anacyclus, wherein it was formerly classified. Humphries compared Heliocauta to a number of genera, notably Achil- lea, but refrained from indicating a possible sister group. The Achillea species discussed by Humphries, A. barrelieri and A. oxyloba, are possibly plesiomorphic within the genus and within the subtribe Achilleinae. The similarities to Heliocauta may be interpreted as symplesiomorphies, shared also by members of Tanacetum. Another genus mentioned by Humphries is Sclerorhachis , supposed to have the same type of scattered epicarpic resin sacs. The elongated resin sacs in Heliocauta have transverse walls and are morphologically similar to the elongated rows of epicarpic myxogenic cells present in many genera. In Sclerorhachis, however, they are not resiniferous but myxo- genic as in other genera, and a close relationship between Heliocauta and Sclerorhachis cannot be assumed. Hippolytia is another possible relative of Heliocauta. They share a number of albeit homoplasious characters and can be placed together within Tanacetinae'. The reason for this placement is the presence of rather similar species within Tanacetum. It appears that both genera, singly or together, have their sister group within Tanacetum. One interesting species is T. tatsienense, a radiate species with short rays but otherwise very similar to Heliocauta and the single-headed species of Hippolytia. 4. GONOSPERMINAE Bremer & Humphries, subtrib. nov. Type species: Gonospermum fruticosum (C. Smith ex Link) Less. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 107 Frutices vel herbae perennes, basi fruticosae. Folia multilo- bata lobis multis rotundatis vel interdum paucilobata vel Integra et margine dentata. Capitula laxe vel dense corym- bosa. Receptaculum paleaceum vel raro epaleaceum. Cypse- lae 5-10(-12)-costatae. Pappus e squamis parvis formatus, squama quoque costam terminanti. Shrubs or perennial, basally woody herbs. Leaves alternate, with many rounded lobes, sometimes few-lobed or entire and dentate. Capitula laxly to densely corymbose, radiate or discoid. Receptacle flat or conical to elongated, paleate or rarely epaleate. Ray floret limb white. Disc corolla 5-lobed. Cypselas 5-10(-12)-ribbed. Pappus of small scales, each terminating a rib. DISTRIBUTION (Table 11). Canary Islands and southern Africa, mainly in Natal, one species also in Madagascar. - 3 genera, 15 spp. Table 11 General distribution of Gonosperminae and Handeliinae and genera. x=indigenous, o=introduced. Table 12 Data matrix for the Gonosperminae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. C.&E. Asia SW Asia N.Afr. S.Afr. Gonosperminae X X Lugoa X Gonospermum X Inulanthera X Handeliinae X X Lepidolopsis X X Polychrysum X X Pseudohandella X X Handelia X X Sclerorhachis X The interesting close relationship between the two Macaron- esian genera Lugoa and Gonospermum and the southern African Inulanthera has only recently been revealed (Kallersjo, 1986). Furthermore, the southern African group of species were formerly erroneously classified in Athanasia, obscuring their true relationships. The Gonosperminae is probably related to a part of Tanacetum, in the subtribe 'Tanacetinae'. Of the two Macaronesian Tanacetum species, T. ferulaceum and T. ptarmiciflorum , the latter at least is a possible close relative. Clades and characters - Fig. 5, Tables 2, 12. = 22 Lugoa tj= 23 Gonospermum =Go 2Ji= 24 Inulanthera Fig. 5 Cladogram of the Gonosperminae produced by the ie option in Hennig86. Cladogram length = 9, consistency index = 100, retention index = 100. Clade Gol - subtribe Gonosperminae 19 Leaves large with many rounded lobes. Inulanthera is very variable in leaf shape, but there are also species of Inulan- thera with these kinds of leaves, very similar to those of Gonospermum and Lugoa. 45 Receptacle paleate. One species of Inulanthera is epaleate. 162 Pappus of scales or teeth projected from the ribs. 1111 1 1 11 11 1450576 146 23992 177555556666617 5327181 952295254 356356790567851 22. ^3 Lugoa Gonospermum alaal?! lalOOOOOO lallllOOO ???000000000000 *><\ aPal 9 ! aalllllll 999999999999999 Lugoa There is no obvious autapomorphy for this genus. Clade Go2 2 Plants shrubby. 29 Capitula densely corymbose. 35 Capitula discoid. Gonospermum There is no obvious autapomorphy for this genus, which appears undefined by comparison to Inulanthera. Inulanthera 92 Anthers caudate. 95 Anthers with endothecial tissue partly or wholly polarized. 124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 22. LUGOA DC., Prodr. 6: 14 (1838). Type species: L. revoluta (C. Smith ex Link) DC. A suffruticose, basally woody perennial. Leaves alternate, pinnatisect, large and flat with rounded lobes. Capitula laxly corymbose, radiate. Receptacle conical, paleate. Ray florets female, fertile; limb white. Disc corolla 5-lobed, slightly thickened at base. Cypselas 5-ribbed. Pappus of teeth pro- jected from the ribs. DISTRIBUTION. Canary Islands. - Monotypic. Lugoa with radiate capitula is the most plesiomorphic mem- ber of subtribe Gonosperminae. There is no obvious autapo- morphy for this monotypic genus, though the one species is clearly distinct from the species of Gonospermum. It is sometimes included in Gonospermum, but retained here in accordance with recent practice (e.g. Bramwell & Bramwell, 1974). Furthermore, inclusion of Lugoa in Gonospermum would result in a clearly paraphyletic taxon since the discoid Gonospermum species are more closely related to Inulan- thera than to Lugoa. 23. GONOSPERMUM Less., Syn. gen. Compos.: 263 (1832). Selected type species: G. fruticosum (C. Smith ex Link) Less. Rather elaborate shrubs. Leaves alternate, pinnatisect, large and flat with many rounded lobes. Capitula corymbose, discoid. Receptacle elongated, paleate. Corolla 5-lobed, 108 K. BREMER AND C. J. HUMPHRIES slightly thickened at base. Cypselas 5-ribbed. Pappus of scales or teeth projected from the ribs. DISTRIBUTION. Canary Islands. - 4 spp. Gonospermum is related to Inulanthera and Lugoa. Inulan- thera differs by its caudate anthers and many-ribbed cypselas. By comparison to Inulanthera, Gonospermum is undefined and paraphyletic. Bramwell & Bramwell's (1974) Wild flow- ers of the Canary Islands is useful for species identification. G. canariense (DC.) Less. G. elegans (Cass.) DC. G. fruticosum (C. Smith ex Link) Less. G. gomerae Bolle 24. INULANTHERA Kallersjo in Nord. J. Bot. 5: 539 (1986). Type species: /. calva (Hutch.) Kallersjo. Shrubs or rarely basally woody half-shrubs. Leaves alternate, variously lobed, dentate or entire. Capitula corymbose, dis- coid. Receptacle flat, paleate, rarely epaleate. Corolla 5-lobed, with a narrow tube and a distinct limb. Anthers caudate; endothecial tissue polarized. Cypselas 8-10-ribbed, glabrous or sometimes glandular. Pappus of small scales or teeth, each terminating a rib. DISTRIBUTION. S. Africa in the E. Cape, Natal, and Trans- vaal, and in Lesotho, Angola (/. schistostephioides), Zimba- bwe (/. nudd) and Madagascar (/. brownii). - 10 spp. Inulanthera is recently described and is comprised of a number of species formerly included in Athanasia. The species of Inulanthera were found to differ considerably from Athanasia in cypsela anatomy, having sclerenchyma of longi- tudinal bundles or elongated cells in the ribs rather than sclerified parenchyma cells as in Athanasia and lacking resin canals which are present in Athanasia. The species of Inulan- thera investigated chemically also have the common poly- acetylenes rather than furanosesquiterpenes, which characterize Athanasia and the group of related genera within the Ursiniinae. In these features Inulanthera is similar to most Anthemideae but the genus is further characterized by its basally caudate anthers, a very unusual character in Anthemideae (occurring elsewhere only in Hippolytia and Osmitopsis). Cypselas with ribs projected into small pappus scales or teeth, and the characteristic leaves with many rounded leaf lobes of some species (/. brownii, I. nuda, I. schistostephio- ides) clearly indicate a relationship to Gonospermum and Lugoa. Other species have few-lobed or dentate leaves, presumably apomorphic conditions within the genus. The South African species, all occurring in Natal, are at the species level adequately described under Athanasia by Hill- iard (1977). /. brownii (Hochr.) Kallersjo (Athanasia brownii Hochr.) - Note: Description in Hochreutiner, 1908. /. calva (Hutch.) Kallersjo (Athanasia calva Hutch.) /. coronopifolia (Harvey) Kallersjo (Athanasia coronopifolia Harvey) I. dregeana (DC.) Kallersjo (Athanasia dregeana (DC.) Har- vey, Athanasia punctata (DC.) Harvey) /. leucoclada (DC.) Kallersjo (Athanasia leucoclada (DC.) Harvey) /. montana (J. M. Wood & M. Evans) Kallersjo (Athanasia montana J. M. Wood & M. Evans) /. nuda Kallersjo (Pentzia schistostephioides M. Taylor) - Note: Description in Taylor, 1940. /. schistostephioides (Hiern) Kallersjo (Athanasia schis- tostephioides Hiern) - Note: Description in Hiern, 1898. /. thodei (Bolus) Kallersjo (Athanasia thodei Bolus) /. tridens (Oliver) Kallersjo (Athanasia tridens Oliver) 5. HANDELIINAE Bremer & Humphries, subtrib. nov. Type species: Handelia trichophylla (Schrenk) Heimerl. Herbae perennes, basaliter suffruticosae et villosae- tomentosae, caule haud vel pauceramoso crassiusculo medulla molli. Folia alterna vel rosulata, pinnatisecta lobis filiformibus. Capitula dense vel laxe corymbosa, raro anguste paniculata, discoidea. Cypselae plerumque parvae, quinque- costatae. Pappus coroniformis e squamis parvis formatus vel nullus. Basally more or less woody and villous-tomentose perennials with generally few- or unbranched rather thick stems with a soft pith. Leaves alternate or rosulate, pinnatisect with fili- form lobes. Capitula generally small in dense to lax corymbs or rarely in a long narrow panicle, discoid. Receptacle flat to conical, paleate or epaleate. Corolla 5-lobed. Cypselas gener- ally small and 5-ribbed, rarely slender with obtuse excres- cences, sometimes with myxogenic cells. Pappus a corona of small scales or absent. DISTRIBUTION (Table 11). Asia, mainly central part. - 5 genera, 8 spp. This subtribe forms a homogeneous and monophyletic group, characterized by a number of synapomorphies. The relation- ship of Lepidolopsis , Poly chry sum, Handelia and Pseudohan- delia was first noted by Tzvelev (in Komarov, 1961). Sclerorhachis was originally part oiAnthemis, mainly because of its paleate receptacle but after it was placed as a section of Anthemis (Rechinger, 1944) Rechinger elevated it to generic rank of uncertain position within the tribe (Rechinger 1968). We consider that Sclerorhachis is related to Handelia and Pseudohandelia and a true member of this subtribe. This relationship has probably been obscured by the distinct habit of Sclerorhachis, with basal leaf- rosettes and laxly branched, leafless aerial stems. We interpret these stems as branches of a secondarily lax inflorescence, homologous to the dense corymbs of the other genera. From the data matrix there is only one parsimonious solution to the generic interrelationships within Handeliinae, shown in the cladogram. The sister group of Handeliinae is probably to be found within Tanacetum, where there are several thick-stemmed species with corymbose inflores- cences. Clades and characters - Fig. 6, Tables 2, 13. |=Hal=jj= 25 Lepidolopsis rj= 26 Polychrysum |l TT^O-II 27 Pseudohandelia 28 Handelia 29 Sclerorhachis Fig. 6 Cladogram of the Handeliinae produced by the ie option in Hennig86. Cladogram length = 13, consistency index = 92, retention index = 80. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 109 Table 13 Data matrix for the Handeliinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 111 1111 11 1450576 13323701457 1555556666617775674 5327181 865190293511 3356790567851560606 111100000000 111011000000 111010111000 HlOlOlOOaOO lalOOOlOOaal 25.Lepidolopsis 26. Polychrysum al?al?l 27 . Pseudohandelia a 1 ? a 1 ? 1 2%. Handelia al?al?l 29.Sclerorhachis Clade Hal - subtribe Handeliinae 8 Plants basally villous-tomentose with rather thick stems and a soft pith. 16 Leaves much pinnatisect with filiform lobes. 35 Capitula discoid. Lepidolopsis 31 Capitula in a long narrow panicle or raceme. Clade Ha2 29 Capitula densely corymbose. In Sclerorhachis the capitula are terminal on the loosely branched, leafless aerial stems, which are considered homologous to the branches of a corymbose inflorescence. Polychrysum 30 Capitula very small and numerous in a large, dense, semiglobose corymb. Clade Ha3 172 Pappus absent in ray and disc cypselas. Pseudohandelia 109 Cypselas arcuate. 113 Cypselas slender and tuber culate with numerous obtuse excrescences. Clade Ha4 45 Receptacle paleate. Handelia There is no obvious autapomorphy for this genus. Sclerorhachis 29 reversed. See clade Hal. 51 Floral parts with resin canals. 71 Corolla apically contracted. 25. LEPIDOLOPSIS Polj. in Bot. Mater. Gerb. hot. Inst. V. A. Komarova 19: 374 (1956). Type species: L. turkestanica (Regel & Schmalh.) Polj. A basally woody perennial with rather thick unbranched stems with a soft pith, villous-tomentose at base and the leaf bases. Leaves alternate, much pinnatisect with filiform lobes. Capitula small, numerous in a long panicle, discoid. Recep- tacle conical, epaleate. Corolla 5-lobed, with sessile glands. Cypselas small, 5-ribbed. Pappus of several mainly abaxial acute scales. DISTRIBUTION. Iran, Afghanistan and C. Asia. - Monotypic. Vegetatively Lepidolopsis is very similar to Polychrysum, Handelia and Pseudohandelia. All four genera have single, unbranched or apically few-branched, basally pubescent stems with much pinnatisect leaves with filiform lobes. Lepi- dolopsis differs by its much elongated inflorescence and its sessile floret glands. Originally Poljakov included a number of discoid species in Lepidolopsis. These are now in other genera, mainly Tanacetum and Tanacetopsis . Both Poly- chrysum tadshikorum and Pseudohandelia umbellifera were also classified in Lepidolopsis by Poljakov. 26. POLYCHRYSUM (Tzvelev) Kovalevsk. in Vvedensky, Fl. uzbekistana 6: 148 (1962). Type species: P. tadshikorum (Kudr.) Kovalevsk. - Cancrinia sect. Polychrysum Tzvelev. A basally woody perennial, one to few-branched with rather thick light stems and a soft pith, villous-tomentose at the leaf bases. Leaves alternate, much pinnatisect with filiform lobes. Capitula small, numerous in a large, dense, semiglobose corymb, discoid. Receptacle convex, epaleate. Corolla 5-lobed, with stalked glands. Cypselas small, 5-ribbed, with stalked glands. Pappus a corona of many small scales. DISTRIBUTION. Afghanistan, C. Asia. - Monotypic. Polychrysum is based on a monotypic section of Cancrinia, described by Tzvelev in Flora URSS (Komarov, 1961). It is clearly different from the type species of Cancrinia, and more closely related to Lepidolopsis, Handelia and Pseudohande- lia. One character stressed by Tzvelev is the capitulum with only one mature fruit. This is obviously correlated to the small and numerous capitula. 27. PSEUDOHANDELIA Tzvelev in Komarov, Flora URSS 26: 878 (1961). Type species: P. umbellifera (Boiss.) Tzvelev. A basally woody perennial with unbranched, rather thick stems with a soft pith, villous-tomentose at the stem and leaf bases. Leaves alternate, basally approaching rosulate, much pinnatisect with filiform lobes. Capitula corymbose, discoid. Receptacle conical, epaleate. Corolla 5-lobed, with many stalked glands. Cypselas slender and somewhat arcuate, with 5 vascular strands, tuberculate with numerous obtuse excres- cences. Pappus absent. DISTRIBUTION. Iran, Afghanistan, C. Asia and China in Xinjiang. - Monotypic. Pseudohandelia and Handelia are very similar and closely related. Their close relationship has been more or less concealed by the traditional practice of keeping taxa with and without receptacular paleae far apart in classification. The two genera are often confused. Apart from the receptacle character Pseudohandelia is also distinguished by its slender and tuberculate cypselas and the more dense, umbelliform corymb. The close relationship of Pseudohandelia to Hande- lia as well as to Polychrysum (treated by Tzvelev as a section of Cancrinia), and Lepidolopsis was first noted by Tzvelev (in Komarov, 1961). 110 The cypsela excrescences are homologous with gland stalks; glands with such stalks are present on the corolla. 28. HANDELIA Heimerl in Ost. hot. Z. 71: 215 (1922). Type species: H. trichophylla (Schrenk) Heimerl. A basally woody perennial with few- or unbranched, rather thick stems with a soft pith, basally and at the leaf bases villous-tomentose. Leaves alternate, basally becoming almost rosulate, leaves pinnatisect, with deep sinuses and filiform lobes. Capitula small, numerous, corymbose, discoid. Recep- tacle conical, paleate. Corolla 5-lobed, with stalked glands. Cypselas small, 5-ribbed, with an abaxial rim. Pappus absent. DISTRIBUTION. Afghanistan, Pakistan, C. Asia and China in Xinjiang. - Monotypic. Handelia is very similar to and closely related to Pseudohan- delia, though the two genera have been placed far apart. Handelia has a paleate receptacle, whereas Pseudohandelia is epaleate. They are often confused but they are easily distin- guished by the receptacle character and by their different cypselas. Handelia also has a more laxly branched inflores- cence than Pseudohandelia. The relationship to Sclerorhachis is discussed below. 29. SCLERORHACHIS (Rech. f.) Rech. f. inAnz. ost. Akad. Wiss. Mathematische Naturwissenschaftliche Klasse 105: 242 (1968). Type species: 5. caulescens (Aitch. & Hemsley) Rech. f. - Anthemis sect. Sclerorhachis Rech. f. Rosulate basally pubescent perennials with a basal leaf- rosette and almost naked, loosely branched stems, basally rather thick and with a soft pith. Leaves pinnatisect, with a persistent, sclerified rachis. Capitula laxly corymbose, pedun- culate, discoid. Receptacle flat to convex, paleate; paleae filiform. Corolla 5-lobed, with a distinct, apically contracted limb. Cypselas 5-ribbed, with scattered rows of myxogenic cells. Pappus absent. DISTRIBUTION. Iran and Afghanistan. - 4 spp. The type species of Sclerorhachis was originally placed in Anthemis, though considered highly isolated (Rechinger, 1944). At a first glance Sclerorhachis seems isolated but it is a member of subtribe Handeliinae together with Handelia and Pseudohandelia (Rechinger, 1955, 1968). In Sclerorhachis the leaves are concentrated into a basal leaf-rosette and we consider the loosely branched, leafless aerial stems with terminal capitula homologous to the corymbose inflorescence of the other genera. The receptacle is furnished with filiform bristles in the same position as receptacular paleae, hence they appear homologous to paleae. S. caulescens (Aitch. & Hemsley) Rech. f. (Anthemis caule- scens Aitch. & Hemsley) *S. leptoclada Rech. f. S. platyrhachis (Boiss.) Podl. ex Rech. f. 5. polysphaera Rech. f. K. BREMER AND C. J. HUMPHRIES 6. ARTEMISIINAE Less, emend. Bremer & Humphries, emend, nov. Lessing in Linnaea 5: 163 (1830) ('Artemisieae'). Type spe- cies: Artemisia vulgaris L. Herbae annuae vel perennes, suffrutices vel frutices. Capitula interdum radiata, saepe disciformia vel discoidea, corymbosa vel paniculata. Receptaculum plerumque epaleaceum. Appendix apicalis antherarum subtriangularis vel lanceolato- linearis, parietibus cellularum aliquantum incrassatis. Cypse- lae oblongo-obovoideae, quinque-costatae vel saepe obovoideae ecostatae, parietibus tenuibus, raro pilosae. Pap- pus nullus vel raro coroniformis e squamis parvis formatus. Perennial, rarely annual herbs, half-shrubs or shrubs; indu- mentum frequently of dolabriform hairs. Leaves variously dissected, rarely entire. Capitula radiate or generally disci- form or discoid, often rather small; inflorescence various but often corymbose or paniculate. Receptacle epaleate or very rarely paleate. Ray floret limb white, yellowish or pink. Outer florets (in disciform capitula) in one row (rarely two) , female, fertile. Disc or central florets perfect and hermaphro- dite or female-sterile, 5-lobed. Apical anther appendages (sub)triangular-lanceolate-linear, of rather thick-walled cells. Pollen often without or with short spines. Cypselas oblong- obovoid, 5-ribbed or generally rather small, obovoid and faintly ribbed or without ribs, thin-walled, with or without myxogenic cells in rows, rarely pilose. Pappus absent or very rarely coroniform, of small scales. DISTRIBUTION (Table 14). Worldwide but mainly northern hemisphere and especially central and E. Asia, some Artemi- sia species widespread as weeds. - 18 genera, 634 spp. The main genera of this subtribe are familiarly known as the Artemisia group, including the large genera Artemisia and Seriphidium (commonly considered a section of Artemisia}, eight small Asian genera, and two small North American genera. The group is characterized by disciform or discoid, commonly paniculate capitula, smooth or short spined pol- len, and obovoid, thin-walled, and ribless cypselas without a pappus. However, within this subtribe we have included the prob- able, more plesiomorphic relatives of the Artemisia group (Fig. 7, Clades Arl-Ar5), which in previous classifications were hidden within a broad concept of Chrysanthemum. These relatives include the radiate genera Brachanthemum, Dendranthema, Arctanthemum and Tridactylina with solitary or laxly corymbose capitula, and the disciform Ajania and Phaeostigma with densely corymbose capitula. These genera have rather thin-walled and more or less faintly ribbed cypselas always without a pappus. The apical anther append- ages are synapomorphic for the whole subtribe. They are (sub)triangular to lanceolate-linear, often acute to acumi- nate, and composed of rather thick- walled cells, best expressed in the apomorphic representatives of the Artemisia group. The sister group of Artemisiinae is to be found within Tanacetum, where there are species very similar to represen- tatives of Dendranthema and Brachanthemum. The delimita- tion between Tanacetum and these two genera, notably Dendranthema, traditionally rests on two characters, pres- ence of pappus and myxogenic cells on the cypselas. Pappose species without myxogenic cells are classified in Tanacetum GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE Table 14 General distribution of Artemisiinae and genera. x=indigenous, o=introduced. Ill N. Am. Eur- Asia C. & E. SW S. Eur. N. Afr. S. Afr. Austr. S. Am. Asia Asia N. Zeal. Artemisiinae x x X X X X X X Brachanthemum X Dendranthema x X Arctanthemum x X X Tridactylina X Ajania X Phaeostigma X Stilpnolepis X Ajaniopsis X Filifolium X Sphaeromerla x Kaschgaria X Serlphldium x x XXX Crossostephium X Artemisia x x X X X X X X Neopallasia X Turaniphytum X Mausolea X Picrothamnus x and epappose species with or without myxogenic cells are classified in Dendranthema. (Pappose species with myxogenic cells represent another problem, and are more or less provi- sionally relegated to other genera, as exemplified by Xylan- themum tianshanicum) . The relationship of Dendranthema and Artemisiinae in general to Tanacetum cannot be assessed in detail before the latter genus has been properly revised and dismantled. The cladogram (Fig. 7) is one of many (71) equally parsimonious trees. In this case the analysis was performed with characters 29 and 31 combined into one multistate character with two apomorphic states, densely corymbose and paniculate-racemose capitula. A preliminary analysis with the binary characters as presented in the matrix (Table 15) yielded an artificial grouping of Stilpnolepis and Seriphidium supported by a reversal in character 29, i.e. loss of the dense capitula corymbs, but the two genera are totally different in inflorescence structure. The strict consensus tree of all the equally parsimonious cladograms is almost totally collapsed, indicating the instability in our current hypothesis of Artemisiinae generic interrelationships. However, clade Ar2 with Dendranthema, Arctanthemum and Tridactylina, as well as clade ArlO, Mausolea and Picrothamnus, were sup- ported by all cladograms and hence retained in the consensus tree. Clades and characters - Fig. 7, Tables 2, 15. Clade Arl - subtribe Artemisiinae 11 Plants with dolabriform hairs. Dolabriform, i.e. T-shaped, or Y-shaped hairs occur in most genera of this subtribe. They are also frequent in Leucantheminae, Anthemis (Anthemidi- nae) and some genera of Achilleinae. The character is possibly a synapomorphy at a lower level within the tribe. 93 Anthers with triangular-linear-lanceolate apical append- ages, of rather thick-walled cells. This character is present in all genera, though variously expressed. 172 Pappus absent in ray and disc cypselas. Crossostephium and one species of Sphaeromeria both have a pappus of short scales. Brachanthemum 2 Plants shrubby. In Artemisiinae there are many transitions from woody perennials to half-shrubs or shrubs. Hence the character is difficult to apply and it is present within several genera not scored in the matrix. Clade Ar2 44 Involucral bracts with dark brown margins. This character is present also within Ajania, Phaeostigma, and Tanacetum, where the sister group of the subtribe is found. Hence the character may be plesiomorphic within the subtribe. Dendranthema This genus is plesiomorphic compared to Arctanthemum, Tridactylina and possibly also compared to the rest of the subtribe. Arctanthemum 6 Plants rhizomatous with rosulate, spathulate-obovate-linear leaves. 15 reversed. Leaves not variously deeply lobed or divided, but entire. Tridactylina 1 Plants annual. 52 reversed. Ray floret limb not white, but yellow. There are several species of Dendranthema with yellow rays. Clade Ar3 29 Capitula densely corymbose. Inflorescences are variable and complicated in subtribe Artemisiinae. Most of the Arte- misia group of genera have paniculate inflorescences. In Stilpnolepis the capitula are, probably secondarily, solitary or laxly corymbose. 112 K. BREMER AND C. J. HUMPHRIES [=Arl= 1= 30 Brachanthemum 31 Dendranthema =Ar2:j!= 32 Arctanthemum l!= 33 Tridactylina = 34 Ajania [f= 35 Phaeostigma l rj= 36 Stilpnolepis U=Ar5| n= 37 Ajaniopsis ^ 38 Filifolium li= 39 Sphaeromeria L= =Ar7 40 Kaschgaria 41 Seriphidium 42 Crossostephium j= 43 Artemisia = 44 Neopallasia = 45 Turaniphytum rr= 46 Mausolea ^= 47 Picrothamnus Fig. 7 Cladogram (of 71 possible) of the Artemisiinae produced by the bb option in Hennig86. Cladogram length = 48, consistency index = 79, retention index = 82. Table 15 Data matrix for the Artemisiinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 11111 11 1 1 111 1450576 197 4 23905369741318312916936700466 8 5327181 132246 1966245578331208629761249468783 1 1111 11 11 5677755556175531 147 46788 06056357905161744657983223 30. Brachanthemum a 1 a a 1 ? 1 31. Dendranthema a 1 a a 1 1 1 32. Arctanthemum 01111?! 33. Tridactylina a 1 ? a 1 ? 1 34 Ajania a 1 9 a 1 9 1 all 100000000pOOOO?OpOOOOOOOOOOOOO 00000 a!10aOOpOOOOOOOOO?00000000000000000000 al!01100000000000?00000000000000000000 ?1101010000000000?00000000000000000000 a!10pOOalOp000000 9 00000000000000000000 ?????000000000000000000000 ?????000000000000000000000 ???00000000000000000000000 ?????000000000000000000000 9999999999999ppOOOOOOOOOOO 35. Phaeostigma a 1 ? a 1 ? 1 36. Stilpnolepis a 1 ? a 1 ? 1 37. Ajaniopsis a 1 ? a 1 ? 1 38 Filifolium a 1 ? a 1 ? 1 allOp001111000000?00000000000000000000 ?11000p001011?pOO?00000000?0000000??00 ?1 10001 11 10101 110?00000000000000000000 al 10000 11 10101 101 ?00000000000000000000 ?????????????0000000000000 ?????????????pOpOOOOOOOOOO ????????????00000000000000 99999999999990000000000000 39 Sphaeromeria a 1 ? a 1 ? 1 allOOOOal 10101 lOOlOpOOOOOOOOOOOOOOOOOp 9999999999999000pOOOOOOOOO 40 Kaschgaria a 1 9 a 1 9 1 all 00000 110 10 11 00? 11 100000000000000000 99999999999990000000000000 41 . Seriphidium a 1 ? a 1 ? 1 42. Crossostephium a 1 ? a 1 ? 1 43 Artemisia a 1 ? a 1 1 1 allpOOp001011?1001aOOHOOO?0000000??00 ?10100001 10001 1 00 ?10000100000000000000 allppOpOl 10101 IGOaapOOOOlOOOOOOppOOOpO ?????????????OpOOpppOOOOOO ?00??????????0000000000000 ?????????????0p0pp00pppppp 44. Neopallasia a 1 ? a 1 ? 1 allOOOaOl 10101 100?1000001 11 10000000000 ?????????????OOOOpOOOOOOOO 45. Turaniphytum a 1 ? a 1 ? 1 ?1 100000 110101 100? 10000010001 1 10000000 ?????????????0000000000000 46. Mausolea a 1 ? a 1 ? 1 all 10000110101 100? 10000010000001 111100 ?????????????0000000000000 47. Picrothamnus a 1 ? a 1 ? 1 all 100001 10101 1001 10000010000001 1 1001 1 ?????????????0000000000000 36 Capitula disdform. Stilpnolepis and Seriphidium have discoid capitula. Within this subtribe discoid capitula are most parsimoniously interpreted as derived from disciform, rather than from radiate capitula. Ajania There is no autapomorphy for this genus, which appears to be a paraphyletic grade when compared to Phaeostigma and the Artemisia group. Clade Ar4 96 Pollen grains with short spines or without spines. Phaeostigma 102 Style-branches brownish. Clade Ar5 - the Artemisia group 154 Cypselas thin-walled, obovoid to oblanceolate, devoid of ribs. Faint ribs are sometimes present, most clearly seen in Crossostephium and sometimes within Sphaeromeria. Stilpnolepis 29 reversed. See clade Ar3. 35 Capitula discoid. See character 36 under clade Ar3. Clade Ar6 65 Outer female floret corollas 'flask-shaped', tapering above or narrowly cylindrical. 97 Pollen grains without spines. Ajaniopsis 1 Plants annual. 78 Corolla apically with erect, straight hairs. Filifolium 143 Cypselas with myxogenic glands in 2 distinct adaxial- lateral rows. Clade Ar7 13 Plants with interxylary cork. Interxylary cork has been studied in Sphaeromeria, Seriphidium, Artemisia and Picroth- amnus only. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE Sphaeromeria There is no obvious autapomorphy for this genus. Clade Ar8 29 reversed. See clade Ar3. 31 Capitula in a long narrow panicle or raceme. See character 29 under clade Ar3. Kaschgaria 12 Plants with stellate hairs. 80 Corolla apically with stellate hairs. Seriphidium 35 Capitula discoid. See character 36 under clade Ar3. 38 Involucral bracts in 4-7 rows. 116 Disc cypselas laterally flattened. Crossostephium 2 Plants shrubby. See note under Brachanthemum. 22 Leaves entire or apically tridentate. 154 reversed. See clade Ar5. 172 reversed. See clade Arl. Clade Ar9 99 Style slender, parallel-sided at base. Artemisia There is no autapomorphy for Artemisia when compared to the remaining four genera. Neopallasia 1 Plants annual. 17 Leaves pectinate-pinnatisect with filiform, apically some- what swollen and mucronulate lobes. 66 Outer female floret corollas without teeth. 91 Central florets of two kinds; outer perfect, inner completely sterile with reduced ovaries. Turaniphytum 32 Capitula in glomerules arranged in long spikes. 64 Outer female florets subtended by scaphoid bracts. 79 Corolla apically with long, reddish hairs. Clade ArlO 2 Plants shrubby. See note under Brachanthemum. 104 Disc floret style-branches fused. This is characteristic also of Artemisia sect. Dracunculus. 106 Central floret ovaries reduced; florets functionally male. 148 Cypselas cobwebby pilose. Mausolea 67 Outer female florets without corollas. 68 Outer female floret style-branches lanceolate, flat, acute. Picrothamnus 113 3 Plants spiny. 81 Corolla cobwebby pilose. 30. BRACHANTHEMUM DC., Prodr. 6: 44 (1838). Type species: B. fruticulosum (Ledeb.) DC. Small, more or less procumbent shrublets, woody at the base. Leaves alternate, few-lobed. Capitula solitary or laxly corym- bose, pedunculate, radiate or rarely discoid. Receptacle flat or convex to conical, epaleate. Ray florets female, fertile; limb yellow or yellowish. Disc corolla 5-lobed; tube with sessile glands. Apical anther appendage subtriangular. Cypselas obovoid to oblong, faintly 5-7-ribbed, thin-walled, with myxogenic cells. Pappus absent. DISTRIBUTION. C. Asia, Mongolia and China (map by Kra- scheninnikov, 1949: 199). - 10 spp. According to Krascheninnikov (1949) and Tzvelev (in Koma- rov, 1961) most species of Brachanthemum are closely related, although B. baranovii is different and placed in the monotypic sect. Dendranthemopsis . It differs from sect. Brachanthemum by its oblong rays and flat, shortly pilose receptacle, as opposed to the otherwise shorter rays and convex to conical, glabrous receptacle. On the other hand the characteristic shrubby habit, the few-lobed leaves, and the rather small, cyathiform to urceolate capitula may distinguish this genus as monophyletic. Tzvelev noted the close relation- ship to Dendranthema, emphasized also by the similar thin- walled, myxogenic fruits without a pappus (described in detail by Savczenko, 1949). The list of species is taken from Krascheninnikov's (1949) revision and Flora URSS (Koma- rov, 1961) with species from China and Mongolia added. *B. baranovii (H. Kraschen. & Polj.) H. Kraschen. B. fruticulosum (Ledeb.) DC. B. gobicum H. Kraschen. B. kasakhorum H. Kraschen. B. kirghisorum H. Kraschen. *B. krylovii Serg. B. mongolicum H. Kraschen. *B. mongolorum Grubov B. pulvinatum (Hand.-Mazz.) Shih (B. nanshanicum H. Kraschen.) B. titovii H. Kraschen. 31. DENDRANTHEMA (DC.) Des Moul. in Act. Soc. linn. Bordeaux 20: 561 (1860). Type species: D. indicum (L.) Des Moul. - Pyrethrum sect. Dendranthema DC. Perennial herbs or half-shrubs. Leaves alternate, pinnatisect, lobed, serrate or rarely entire. Capitula laxly corymbose or solitary, radiate. Involucral bracts generally with dark brown margins. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb white, pink, or yellowish. Disc corolla 5-lobed; tube generally with sessile glands. Apical anther appendage subtriangular. Cypselas obovoid, faintly 5-8- ribbed, thin-walled, generally with myxogenic cells in rows. Pappus absent. DISTRIBUTION. Asia, mainly in China and Japan, one species extending to E. Europe (D. zawadskii). - 37 spp. Dendranthema differs from Tanacetum (i.e. Chrysanthemum 114 K. BREMER AND C. J. HUMPHRIES s. 1.) by its obovoid, thin-walled, generally myxogenic cypse- las without a pappus. It was revised by Tzvelev (in Komarov, 1961). Later Tzvelev (1985) transferred three species of Dendranthema to a new genus Arctanthemum and later erected the new genus Hulteniella based on Arctanthemum integrifolium (Richardson) Tzvelev (Tzvelev, 1987). Other related genera are Brachanthemum and Tridactylina. There is still much work to be undertaken on this species- rich and horticulturally interesting genus. There are many, poorly understood species, known only from the original description, and probably several undescribed ones. Possibly there are other species to be transferred from Tanacetum. Future revision should be undertaken with Ajania, since some species of Dendranthema may be more closely related to Ajania than to other species within Dendranthema. In other words, Dendranthema may be paraphyletic when Aja- nia and its relatives (i.e. Artemisia etc.) are excluded. As presently understood, Dendranthema is always radiate and Ajania always disciform. Most species of Dendranthema also have involucral bracts with dark brown margins, a character that may have been secondarily lost in the small-headed species of Ajania. The well-known autumn-flowering chrysanthemums of horticulture are derived from D. grandiflorum and D. indi- cum. The list of species is compiled mainly from Flora URSS (Komarov, 1961) and the Flora of the People's Republic of China (Ling & Shih, 1980, 1983; Shih & Fu, 1983), as well as from accounts from Japan by Kitamura (1940, 1978, 1979). These authors transferred a number of species from Chrysan- themum to Dendranthema. Kitamura also included four disci- form species, D. pallasianum, D. rupestre, D. pacificum, and D. shiwogiku, in section Ajania. The latter we consider as a separate genus in agreement with various other authors and so the disciform species are here listed under Ajania. The generic distinction, as discussed under that genus, is in doubt but pending a detailed study of both genera we consider it best to transfer a few disciform Japanese Dendranthema species to Ajania rather than to recombine the numerous Chinese Ajania species under Dendranthema. *D. aphrodite (Kitam.) Kitam. Japan. *D. argyrophyllum (Ling) Ling & Shih. China. D. arisanense (Hayata) Ling & Shih. Taiwan. D. boreale (Makino) Ling ex Kitam. China, Korea, Japan. D. chalchingolicum (Grubov) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum chalchingolicum Grubov in Bot. Zhurn. 15: 1592 (1972). Mongolia. D. chanetii (A. Leveille) Shih (D. erubescens (Stapf) Tzvelev). China, Korea. D. coreanum (A. Leveille & Vaniot) Vorosch. Korea. *D. crassum (Kitam.) Kitam. Japan. D. cuneifolium (Kitam.) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum cuneifolium Kitam. in Acta phytotax. geobot. Kyoto 7: 68 (1938). Japan. *D. dichrum Shih. China. *D. glabriusculum (W. Smith) Shih. China. D. grandiflorum (Ramat.) Kitam. (D. morifolium (Ramat.) Tzvelev). China, much cultivated. D. hypargyrum (Diels) Ling & Shih. China. D. indicum (L.) Des Moul. China, Korea, Japan, much cultivated. D. japonense (Nakai) Kitam. Japan. D. japonicum (Makino) Kitam. Japan. D. lavandulifolium (Fischer ex Trautv.) Kitam. China. *D. littorale (Mackawa) Tzvelev. Far East, Japan. D. maximowiczii (V. Komarov) Tzvelev. Far East, China, Korea. D. miyatojimense (Kitam.) Hind. Japan. D. mongolicum (Ling) Tzvelev. E. Siberia, China. *D. morii (Hayata) Kitam. Taiwan. D. nankingense (Hand.-Mazz.) Y. R. Ling. China. *D. okiense (Kitam.) Kitam. Japan. D. oreastrum (Hance) Ling (D. sicho tense Tzvelev). Far East, China, Korea. D. ornatum (Hemsley) Kitam. Japan. *D. parvifolium (Chang) Shih. China. D. potentilloides (Hand.-Mazz.) Shih. China. *D. rhombifolium Ling & Shih. China. *D. sinchangense (Ueki) Kitam. Korea. D. sinuatum (Ledeb.) Tzvelev. C. Asia, Mongolia. *D. vestitum (Hemsley) Ling. China. *D. weyrichii (Maxim.) Tzvelev. Far East, Japan. *D. xeromorphum Khokr. Russia. D. yezoense (T. Maek.) Hind. Japan. D. yoshinaganthum (Makino ex Kitam.) Kitam. Japan. D. zawadskii (Herbich) Tzvelev (D. naktongense (Nakai) Tzvelev). From E. Europe through Russia to China, Mon- golia, and Japan. 32. ARCTANTHEMUM (Tzvelev) Tzvelev in Nov. Sist. Vysshikh Rast. 22: 274 (1985). Type species: A. arcticum (L.) Tzvelev (including Hulteniella Tzvelev). Perennial rhizomatous herbs. Leaves rosulate to alternate, obovate-spathulate to linear, apically lobed-serrate or entire. Capitula solitary, radiate. Involucral bracts with dark brown margins. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb white. Disc corolla 5-lobed; tube gener- ally with sessile glands. Apical anther appendage subtriangu- lar. Cypselas oblong, somewhat 5-8-ribbed, without myxogenic cells. Pappus absent. DISTRIBUTION. Arctic Eurasia, Siberia, Far East, Japan, Arctic N. America. - 4 spp. Arctanthemum was formerly a section of Dendranthema. It consists of more or less rosulate herbs with an arctic distribu- tion, whereas most Dendranthema species are leafy herbs or herbaceous perennials from China and Japan. It is worth noting that Tzvelev (1987) transferred A. integrifolium to a new genus, Hulteniella Tzvelev, which we do not recognize here. A. arcticum (L.) Tzvelev (Dendranthema arcticum (L.) Tzvelev). Far East and Arctic America. A. hultenii (A. & D. Love) Tzvelev (Dendranthema hultenii (A. & D. Love) Tzvelev). Arctic Eurasia and Arctic America. A. integrifolium (Richardson) Tzvelev (Dendranthema inte- grifolium (Richardson) Tzvelev, Hulteniella integrifolium (Richardson) Tzvelev). NE Siberia and Arctic N. America. A. kurilense (Tzvelev) Tzvelev (Dendranthema kurilense (Tzvelev) Tzvelev). Far East, Japan. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 33. TRIDACTYLINA (DC.) Schultz-Bip. in Webb & Berthelot, Hist. nat. lies Canaries 3 (2,2): 245 (1844). Type species: T. kirilowii (Turcz. ex DC.) Schultz-Bip. An annual herb. Leaves alternate, few-lobed. Capitula laxly corymbose, radiate. Involucral bracts with dark brown mar- gins. Receptacle flat to convex, epaleate. Ray florets neuter; limb yellow. Disc corolla 5-lobed. Apical anther appendage subtriangular. Cypselas 5-ribbed, with myxogenic cells and with an apical rim. Pappus absent. DISTRIBUTION. E. Siberia. - Monotypic. According to Tzvelev in Flora URSS (Komarov, 1961) this species is clearly related to Dendranthema but differs by its annual habit and neutral rays. It is similar for example in foliage to some species of Dendranthema and Arctanthemum. 34. AJANIA Polj. in Bot. Mater. Gerb. hot. Inst. V. A. Komarova 17: 419 (1955). Type species: A. pallasiana (Fischer ex Besser) Polj . Perennial herbs or half-shrubs. Leaves alternate, pinnatisect, lobed, serrate, or rarely entire. Capitula small, corymbose or rarely solitary, disciform. Receptacle convex to conical, epaleate. Outer female florets in one row. Corollas of central florets 5-lobed; tube generally with sessile glands. Apical anther appendage subtriangular. Cypselas obovoid, faintly 4 6-ribbed, thin-walled, generally with myxogenic cells in rows. Pappus absent. DISTRIBUTION. C. Asia, mainly in China, also in Japan. - 34 spp. Poljakov removed a number of species from Artemisia to a separate genus Ajania. He noted that Ajania has fertile florets and the common form of distinct 5-lobed corollas with spreading lobes, whereas in Artemisia only some of the florets in each capitulum are fertile, producing mature fruits, and the corollas have small erect lobes. Ajania also has a corymbose inflorescence as opposed to the elongated inflorescences of Artemisia. Poljakov speculated that Ajania is a very old group derived from the same ancestors as Artemisia. Tzvelev (in Komarov, 1961) accepted Ajania although he retained some of Poljakov's Ajania species in Artemisia. He also noted the close affinity between Dendranthema and Ajania, and speculated that Ajania and Artemisia are two convergent and habitually similar lines independently evolved from the same 'dendranthemoid' ancestors. More parsimoniously, Ajania may be considered the plesiomorphic sister group to Artemisia and all its relatives. Possibly Ajania or part of the genus is the sister group of Artemisia and the other genera with smooth or short-spined pollen, as indicated in the cladogram. Dendranthema is even more plesiomorphic and possibly paraphyletic with Ajania, and Artemisia etc. excluded. Ajania differs from Dendranthema by its smaller, disciform, densely corymbose capitula. No doubt this species- rich and little-known genus deserves a detailed study together with Dendranthema. A few species of Ajania have recently been placed in a separate genus, Phaeostigma. There may be several species of Ajania (Muldashev, 1982, 1983) that should be transferred to Phaeostigma, should the latter be kept distinct. The type species of Ajania, A. pallasiana, and A. latifolia, A. rapes tris, and A. shiwogiku are all examples of species similar to 115 Phaeostigma. The matter is discussed further under Phaeo- stigma. Tzvelev suspected that the distinction between Ajania and Dendranthema would become difficult after examination of the extensive Chinese material. Nevertheless Ling & Shih (1980, 1983) kept them separate in their accounts of the Chinese species. Our list of species is compiled mainly from their treatments and from Flora URSS (Komarov, 1961). The species from Tibet were listed by Shih & Fu (1979) and the Japanese species are from Kitamuras's (1978) Dendranthema sect. Ajania (see discussion under Dendranthema). A. achilleoides (Turcz.) Polj. ex Grubov. Mongolia. A. adenantha (Diels) Ling & Shih. China. *A. brachyantha Shih. China. A. breviloba (Franchet ex Hand.-Mazz.) Ling & Shih. China. *A. elegantula (W. Smith) Shih. China. A. fastigiata (Winkler) Polj. C. Asia, China. A. fruticulosa (Ledeb.) Polj. (A. aureoglobosa (W. Smith & Farrer) Muld.). C. Asia, E. Siberia, Mongolia, China. A. gracilis (Hook. f. & Thomson) Polj. ex Tzvelev. C. Asia, China, Himalaya. *A. gruboviiMuld. Mongolia. *A. junnanica Polj. China. A. khartensis (Dunn) Shih (A. mutellina (Hand.-Mazz.) Muld.). China. M. kokanica (H. Kraschen.) Tzvelev. C. Asia. A. latifolia Shih. China. A. myriantha (Franchet) Ling ex Shih (A. oresbia (W. Smith) Muld.). China. *A. nana (H. Kraschen.) Muld. China. M. nematoloba (Hand.-Mazz.) Ling & Shih. China. M. nitida Shih. China. A. nubigena (Wallich) Shih. China. A. pacit]c;i (Nakai) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum pacificum Nakai in Bot. Mag., Tokyo 42: 462 (1928) (Dendranthema pacificum (Nakai) Kitam.). Japan. A. pallasiana (Fischer ex Besser) Polj. (Dendranthema palla- sianum (Fischer ex Besser) Vorosch.). Far East, China, Korea, Japan. A. parviflora (Griin.) Ling. China. A. potaninii (H. Kraschen.) Polj. China. A. przewalskii Polj. China. M. purpurea Shih. China. *A. remotipinna (Hand.-Mazz.) Ling & Shih. China. *A. roborowskii Muld. China. A. rupestris (Matsum. ex Koidz.) Muld. Japan. A. scharnhorstii (Regel & Schmalh.) Tzvelev. C. Asia. *A. sericea Shih. China. A. shiwogiku (Kitam.) Bremer & Humphries, comb. nov. Basionym: Chrysanthemum shiwogiku Kitam. in Acta phy- totax. geobot. Kyoto 4: 71 (1935). Japan. A. tenuifolia (Jacquem.) Tzvelev. Himalaya. A. tibetica (Hook. f. & Thomson) Tzvelev. C. Asia, China, Himalaya. *A. trilobata Polj. C. Asia. *A. tripinnatisecta Ling & Shih. China. 35. PHAEOSTIGMA Muld. in Bot. Zhurn. 66: 586 (1981). Type species: P. salicifolium (Mattf.) Muld. Perennial herbs or half-shrubs. Leaves alternate, pinnatifid to shallowly lobed or entire. Capitula small, densely corymbose, 116 K. BREMER AND C. J. HUMPHRIES disciform. Receptacle convex, epaleate. Outer female florets in one row. Central floret corolla 5-lobed; lobes erect. Apical anther appendage subtriangular. Style-branches brownish. Cypselas 4-6-ribbed; ribs projected into minute teeth. Pap- pus absent. DISTRIBUTION. China. - 3 spp. Muldashev made a detailed comparison of Ajania fruticulosa and the three species of Ajania transferred by him to Phaeo- stigma. According to Muldashev Phaeostigma is distinguished by its brownish style-branches and erect corolla lobes. The latter character, as well as the suffruticose habit and the less spiny pollen, are characters similar to those of Artemisia. Muldashev stated that phylogenetically Phaeostigma is related to the ancestors of Ajania and distantly related to Artemisia, mainly on account of its pollen morphology. The brownish style-branches is a difficult character to evaluate. Similar style-branches may be present in related genera. Although Shin & Fu (1983) reduced Phaeostigma to synonomy under Ajania we have kept them distinct. In that case Phaeostigma may be another grade group between Ajania and Artemisia and its relatives. The actual circum- scription of Phaeostigma will have to be assessed in a wider study of Ajania and especially those Ajania species that are similar in habit to Phaeostigma. P. quercifolium (W. Smith) Muld. (Ajania quercifolia (W. Smith) Ling & Shih) P. salicifolium (Mattf.) Muld. (Ajania salicifolia (Mattf.) Polj.) P. variifolium (Chang) Muld. (Ajania manshurica Polj., Ajania ramosa (Chang) Shih, Ajania variifolia (Chang) Tzvelev) 36. STILPNOLEPIS H. Kraschen. in Nov. Sist. Vysshikh Rast. 2: 207 (1946). Type species: 5. centiflora (Maxim.) H. Kraschen. - Elachanthemum Ling & Y. R. Ling. Annual or perennial herbs. Leaves opposite or alternate, pinnatisect, few-lobed or entire. Capitula laxly corymbose, discoid. Involucral bracts widely obovate and largely scari- ous. Receptacle convex-subconical, epaleate. Corolla 5-lobed, with a distinct tube and more or less crateriform limb, glandular. Apical anther appendage acuminate- triangular. Cypselas obliquely obovoid to narrowly obovoid- oblanceolate, thin-walled, without ribs, densely glandular. Pappus absent. DISTRIBUTION. Mongolia and China. - 2 spp. According to Krascheninnikov (1946) S. centiflora resembles Artemisia, but does not share the obovoid cypselas, the slender corollas, the acuminate anther appendages and the paniculate inflorescence of Artemisia and closely related genera. Though elongated, the cypselas are basically the same as in other genera of subtribe Artemisiinae; obovoid (oblanceolate), thin-walled and devoid of ribs. There is no doubt about the subtribal position of Stilpnolepis . According to Ling & Y. R. Ling (1978) the second species, 5. intricata, formerly classified in Artemisia or Seriphidium, should be separated from those genera because of the long peduncles and the laxly corymbose inflorescence, the cen- trally abortive disc florets, and the multicostate appearance of the rows of myxogenic cells on the cypselas. Hence they created a new genus Elachanthemum for this species. How- ever, Ling and Y. R. Ling did not consider a possible relationship with Stilpnolepis. Recently Shih (1985) argued that Elachanthemum intricatum is closely related to Stilpnol- epis centiflora and his treatment is followed here. Both species have discoid and laxly corymbose capitula, most parsimoniously interpreted as apomorphic within the Artemi- sia group, and hence synapomorphies uniting the two species. 5. centiflora (Maxim.) H. Kraschen. 5. intricata (Franchet) Shih (Elachanthemum intricatum (Franchet) Ling & Y. R. Ling 37. AJANIOPSIS Shih in Acta phytotax. sin. 16: 86 (1978). Type species: A. penicilliformis Shih. A pubescent annual herb. Leaves alternate, pinnatisect and few-lobed. Capitula rather small, few together in dense corymbs, disciform. Receptacle convex, epaleate. Outer female florets in one row, tapering above, apically densely pilose with erect, straight hairs. Central floret corolla 5-lobed, apically densely pilose with erect, straight hairs. Apical anther appendage subtriangular. Cypselas obovoid, thin-walled with 3-6 rows of myxogenic cells. Pappus absent. DISTRIBUTION. China, Tibet. - Monotypic. This is a distinctive species and undoubtedly a member of Artemisiinae, though its immediate relatives are difficult to indicate. Shih (1978) compared it to both Ajania and Artemi- sia. It is distinguished by its annual habit, the densely corymbose capitula and apically pilose corollas. The genus differs from Ajania only by the corollas being pilose at the apex. The cypsela ribs, described and illustrated by Shih, appear to be rows of myxogenic cells. We have only exam- ined the holotype and avoided consuming type material for a detailed cypsela investigation. 38. FILIFOLIUM Kitam. in Acta phytotax. geobot. Kyoto 9: 157 (1940). Type species: F. sibiricum (L.) Kitam. A perennial herb, basally somewhat woody and covered with fibrous leaf sheaths. Leaves alternate, pinnatisect with long filiform lobes. Capitula rather small, corymbose, disciform. Receptacle conical, epaleate. Outer female florets tapering above, minutely 4-dentate. Central florets apparently her- maphrodite but functionally male and female-sterile; corollas compressed together in a resinous mass, 5-lobed. Apical anther appendage triangular. Cypselas obliquely obovoid, thin-walled, with 2 adaxial-lateral rows of myxogenic cells. Pappus absent. DISTRIBUTION. Far East, China and Korea. - Monotypic. Filifolium was earlier included in Artemisia sect. Dracunculus on account of the female-sterile disc florets. However, the inflorescence is corymbose, the style is distinctly divided and the apical appendages of the anthers are triangular, not linear-lanceolate. In habit Filifolium is more similar to some species of Ajania than to Artemisia. In pollen and cypsela characters Filifolium is apomorphic compared to Ajania. Hence, it occupies an intermediate position within the sub- tribe, as expressed in the cladogram. The obliquely obovoid cypselas with two distinct rows of myxogenic cells appear to be autapomorphic for this genus. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 117 39. SPHAEROMERIA Nutt. in Trans. Am. phil. Soc. 2 (7): 401 (1841). Type species: S. capitata Nutt. - Chamartemisia Rydb. - Vesicarpa Rydb. Perennial herbs or half-shrubs, sometimes rather compact and basally woody. Leaves alternate to rosulate, pinnatisect and few-lobed or entire. Capitula apparently corymbose but sometimes somewhat paniculate or capitate or solitary, disci- form. Receptacle flat to conical, epaleate, rarely pubescent. Outer female florets tapering above, sometimes glandular, rarely with dolabriform hairs at the apex. Central florets with corolla 5-lobed, sometimes glandular, rarely with dolabri- form hairs at the apex. Apical anther appendage subtriangu- lar. Cypselas obovoid-oblong, sometimes faintly ribbed, sometimes with myxogenic cells, often with an apical rim. Pappus usually absent or rarely minutely coroniform, with small scales. DISTRIBUTION. Western N. America in California, Colo- rado, Montana, Nevada, Oregon, Utah and Wyoming, and in Mexico. - 9 spp. Sphaeromeria was originally proposed by Nuttall to circum- scribe S. capitata and 5. argentea. Torrey & Gray (1843) reduced Sphaeromeria to a section of Tanacetum simply on combination of three plesiomorphic characters, the yellow 5-lobed corollas, the woody base and the alternate leaves. Rydberg (1916) returned Sphaeromeria to its generic status and erected two new genera, Vesicarpa to include Artemisia potentilloides A. Gray with a hairy receptacle and Chamarte- misia for Tanacetum compactum Hall with pappose cypselas. A. Holmgren et al. (1976) presented a detailed study of the three genera Sphaeromeria, Vesicarpa and Chamartemisia, and concluded that they are more closely related to one another than to any other group. Sphaeromeria appears to be more closely related to Artemisia than Tanacetum. However, of the 12 characters discussed by A. Holmgren et al. not one is autapomorphic for Sphaeromeria. They are present in all other genera of Artemisiinae. The presence of a small coroniform pappus in one species is an unusual feature within Artemisiinae but a parallel case is Crossostephium. Sphaeromeria is similar in habit to Kaschgaria. The inflo- rescences of Sphaeromeria are variable and the transforma- tions within the genus are difficult to assess. In the cladogram the corymbose condition is considered plesiomorphic but there are species with slightly elongated inflorescences resem- bling those of Kaschgaria. The two genera are possibly closely related. The list of species is taken from A. Holmgren et al., who also provided a key. 5. argentea Nutt. 5. cana (D. C. Eaton) A. A. Heller S. capitata Nutt. S. compacta (H. M. Hall) A. Holmgren, Shultz & Lowrey (Chamartemisia compacta (H. M. Hall) Rydb.) S. diversifolia (D. C. Eaton) Rydb. *S. martirensis (Wiggins) A. Holmgren, Shultz & Lowrey S. potentilloides (A. Gray) A. A. Heller (Vesicarpa potentil- loides (A. Gray) Rydb.) S. ruthiae A. Holmgren, Shultz & Lowrey 5. simplex (Nelson) A. A. Heller 40. KASCHGARIA Polj. in Bot. Mater. Gerb. hot. Inst. V. A. Komarova 18: 282 (1957). Type species: K. brachanthemoid.es (Winkler) Polj. Rather woody half-shrubs; indumentum of stellate hairs. Leaves alternate, entire or few-lobed. Capitula rather small and few in an elongated panicle, at the summit fasciculate, disciform. Receptacle conical, epaleate. Outer female florets few, tapering above. Central floret corolla apically with stellate hairs, 5-lobed. Apical anther appendage linear- lanceolate. Cypselas obovoid, thin-walled. Pappus absent. DISTRIBUTION. C. Asia and China. - 2 spp. Poljakov removed the two species of Kaschgaria from Arte- misia mainly because of their stellate corolla hairs. He also stated that Kaschgaria is closer to Seriphidium than to Artemisia s. s., but Kaschgaria differs from Seriphidium, for example, by its disciform (heterogamous) versus a discoid (homogamous) capitulum. The possible relationship to Sphaeromeria (then part of Tanacetum) was not mentioned by Poljakov, who probably did not consider the North American species of Tanacetum. He closed his discussion noting that the systematic position of Kaschgaria could not be settled before the whole Asian part of the Artemisia group had been properly considered. We agree that Kaschgaria is a distinct genus probably related to Seriphidium, or possibly more closely to Sphaeromeria, although there is no obvious synapomorphy uniting the two genera. K. brachanthemoides (Winkler) Polj . K. komarovii (H. Kraschen & N. Rubtzov) Polj. 41. SERIPHIDIUM (Besser ex Hook.) Fourr. in Annls. Soc. linn. Lyon. II, 17: 89 (1869) Type species: 5. maritimum (L.) Polj. -Artemisia sect. Seriphidium Besser - Artemisiastrum Rydb. Perennial herbs, half-shrubs or annual herbs. Leaves alter- nate, pinnatisect. Capitula small and few-flowered, oblong, numerous in a long panicle, discoid. Involucral bracts in 4-7 rows, unequal; the outer short and rounded, the inner gradually longer and linear. Receptacle small, conical, epale- ate or occasionally paleate. Corolla 5-lobed, tubular, yellow to purple. Apical anther appendage linear-lanceolate. Cypse- las small, obovoid to ellipsoid, somewhat flattened or trique- trous, thin-walled. Pappus absent. DISTRIBUTION. From Europe throughout temperate Asia to N. America (5 spp.), though mainly in central Asia. - 134 spp. Seriphidium, established as a genus by Poljakov (1961), is almost always considered a section or subgenus of Artemisia (e.g. Ward, 1940). We agree with Poljakov (1961), Leonova (1970) and Filatova (1981, 1982a, b 1984) that it should be kept distinct. Seriphidium differs in a number of characters, representing synapomorphies. Furthermore, its sister group appears to be found outside Artemisia s. s., and Seriphidium cannot be considered a specialized segregate of Artemisia. Seriphidium is most closely related either to a larger group of genera including Artemisia or some other genus (or genera), e.g. Kaschgaria, within the Artemisiinae. The genus is char- acterized by its discoid, homogamous capitula, not disciform and heterogamous as in Artemisia. The most parsimonious interpretation for this character is to assume that the outer 118 K. BREMER AND C. J. HUMPHRIES female florets present in related genera have been lost. This was originally suggested by Krascheninnikov (1946) 'although hotly contested by Poljakov (1961). It must be noted that this character breaks down in 5. bigelowii. This species is usually included in Artemisia section Abrotanum because there are at least some capitula within inflorescences that have one or two female outer florets. However, on the basis of involucral (many-rowed bracts) and anther (slender apical appendages) characters S. bigelowii is certainly a member of Seriphidium. It is also virtually identical to the American 5. tridentatum, which is homogamous and always included in Seriphidium (see Weber, 1984). Heterogamy in 5. bigelowii is plesiomor- phic or a secondary reversal from the homogamous condi- tion, which then can be interpreted as a synapomorphy for the genus. The corollas of Seriphidium are tubular and the teeth usually infolded for much of anthesis. During anthesis the short flat lobes of the style are often still together and enclosed in the anther tube, a feature mostly associated with cleistogamy. Mature cypselas are hard to find and there are many occasions when only one will mature in a head of 3-10 florets. The apical anther appendages of Seriphidium are very slender, narrowly lanceolate to linear and notably different from those of Artemisia and related genera. The involucre is specialized with 4-7 rows of overlapping bracts, as compared to the 2^-rowed involucre of related genera. The monotypic Artemisiastrum (A. palmeri=S. palmeri) was separated by Rydberg (1916) on the presence of recep- tacular scales. Parallel cases are legion in the Anthemideae and in Asteraceae as a whole. Here the presence of paleae is best considered a gain character, a feature also noted by Hall & Clements (1923). In all characters S. palmeri is virtually identical to other taxa within Seriphidium. The list of species has been compiled from the standard floras, cf. under Artemisia, and by original research with Y. R. Ling (1991a, b) during his sabbatical year at The Natural History Museum, London. Except for cases of basionyms, synonymous Artemisia names are not included when the same epithet is involved. Also, combinations from Artemisia by Bremer & Humphries have been validly published in Ling (19910). S. algeriense (Filat.) Y. R. Ling. Algeria. *S. amoenum (Polj.) Polj. C. Asia. S. aralense (H. Kraschen.) Polj. SW C. Asia. 5. arbusculum (Nutt.) W. A. Weber. W. United States. 5. arenicolum (H. Kraschen. ex Polj.) Y. R. Ling. Afghani- stan, Iran, C. Asia. *S. argilosum (Beetle) Bremer & Humphries, comb. nov. Basionym: Artemisia argilosa Beetle, Rhodora 61: 84 (1959). United States in Colorado. *S. assurgens (Filat.) Bremer & Humphries in Y. R. Ling. Russia. *S. aucheri (Boiss.) Ling & Y. R. Ling. Iran, Afghanistan, China. S. badhysi (Krasch. & Lincz. ex Polj.) Polj. C. Asia. S. balchanorum (H. Kraschen.) Polj. SW C. Asia. *S. baldshuanicum (H. Kraschen. & Zaprj.) Polj. Afghani- stan, C. Asia. 5. barrelieri (Besser) Sojak. Spain, N. Africa. S. bicolor (Rech. f. & Wagenitz) Bremer & Humphries, comb. nov. Basionym: Artemisia bicolor Rech. f. & Wagenitz in Anz. ost. Akad. Wiss. Mathematische Natur- wissenschaftliche Klasse., 98: 78 (1961). Afghanistan. S. bigelowii (A. Gray) Bremer & Humphries, comb. nov. Basionym: Artemisia bigelowii A. Gray in Torrey Pacific R. Rep. 4: 110 (1857). S. and W. United States. *S. borotalense (Polj.) Ling & Y. R. Ling. C. Asia, China. 5. botschantzevii (Filat.) Y. R. Ling. Russia. *S. brevifolium (Wallich ex DC.) Ling & Y. R. Ling. Afghanistan, Pakistan, India, China. 5. caerulescens (L.) Sojak. S. Europe. *S. camelorum (H. Kraschen.) Polj. C. Asia. 5. canum (Pursh) W. A. Weber. W. North America. S. chitralense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia chitralensis Podl., Fl. iranica 158: 198 (1986). Afghanistan, Pakistan. *5. ciniforme (H. Kraschen. & Popov ex Polj.) Polj. Iran, C. Asia. S. cinum (P. Bergius ex Polj.) Polj. C. Asia, China. S. compactum (Fischer ex DC.) Polj. C. Asia, S. Siberia, China in Sinkiang, Mongolia. *S. cretaceum (Fiori) Bremer & Humphries, comb. nov. Basionym: Artemisia caerulescens var. cretacea Fiori in Fiori & Paoletti, Fl. Italia 3: 251 (1904). S. Europe in Italy. S. densifolium (Filat.) Y. R. Ling. Algeria 5. deserti (H. Kraschen.) Polj. Afghanistan, Iran, SW C. Asia. S. diffusum (H. Kraschen. ex Polj.) Y. R. Ling. Iran, Afghanistan, Pakistan. *5. dubjanskyanum (H. Kraschen. ex Polj.) Polj. C. Asia. S. dumosum (Polj.) Polj. C. Asia. *S. dzevanovskyi (Leonova) Sojak. Krym. *S. elongatum (Filat. & Ladyg.) Bremer & Humphries in Y. R. Ling. C. Asia. *5. eremophilum (H. Kraschen. & Butkov ex Polj.) Bremer & Humphries in Y. R. Ling. C. Asia. 5. federovii (Rzazade) Y. R. Ling. C. Asia. *5. fedtschenkoanum (H. Kraschen.) Polj. C. Asia, China in Xinjiang. *5. ferganense (H. Kraschen. ex Polj.) Polj. C. Asia. *S. finitum (Kitagawa) Ling & Y. R. Ling. China. 5. fragrans (Willd.) Polj. Afghanistan, Caucasus, Iran, C. Asia. S. freitagii (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia freitagii Podl., Fl. iranica 158: 193 (1986). Afghanistan. *S. fulvellum (Filat. & Ladyg.) Bremer & Humphries in Y. R. Ling. C. Asia. S. ghazniense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia ghazniensis Podl., Fl. iranica 158: 213 (1986). Afghanistan. S. ghoratense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia ghoratensis Podl., Fl. iranica 158: 197. Afghanistan. *5. glanduligerum (H. Kraschen. ex Polj.) Polj. Afghanistan, Pakistan, C. Asia. *S. glaucinum (H. Kraschen. ex Polj.) Bremer & Humphries in Y. R. Ling. C. Asia. 5. gorjaevii (Polj.) Y. R. Ling. C. Asia. S. gracilescens (H. Kraschen. & Iljin) Polj. C. Asia, S. Siberia, Mongolia, China. 5. grenardii (Franchet) Y. R. Ling & Humphries. China. *5. gurganicum (H. Kraschen.) Bremer & Humphries in Y. R. Ling. C. Asia. *S. gypsaceum (H. Kraschen., Popov & Lincz. ex Polj.) Polj. Iran, SW & C. Asia. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 119 *S. halophilum (H. Kraschen.) Polj. C. Asia. *S. heptapotamicum (Polj.) Ling & Y. R. Ling. C. Asia, China. S. herba-album (Asso) Sojak. SW Europe, Turkey, Middle East, Iran, Himalayas. 5. incultum (Del.) Y. R. Ling. Egypt. S. issykkulense (Polj.) Polj. C. Asia, China. S. junceum (Karelin & Kir.) Polj. C. Asia, China in Xinjiang. S. kandaharense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia kandaharensis Podl., Fl. iranica 158: 217. Afghanistan. *S. karatavicum (H. Kraschen. & Abolin ex Polj.) Ling & Y. R. Ling. C. Asia, China. *S. kasakorum (H. Kraschen.) Bremer & Humphries, comb. nov. Basionym: Artemisia maritima ssp. kasakorum H. Kraschen., Otch. Rab. Pochv.-Bot. Otr. Kazakhst. Exped. Akad. Nauk SSSR 4(2): 272 (1930). C. Asia. *S. kaschgaricum (H. Kraschen.) Polj. C. Asia, China in Xinjiang. *S. kemrudicum (H. Kraschen.) Polj. SW & C. Asia. S. kermanense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia kermanensis Podl., Fl. iranica 158: 206 (1986). Iran. *S. khorassanicum (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia khorassanica Podl., Fl. iranica 158: 210 (1986). Iran, Afghanistan. *S. knorringianum (H. Kraschen.) Polj. C. Asia. *5. kochiiforme (H. Kraschen. & Lincz. ex Polj.) Polj. Afghanistan, C. Asia. *S. kopetdaghense (H. Kraschen. ex Polj.) Polj. Afghanistan, Iran, Afghanistan, SW C. Asia. S. korovinii (Polj.) Polj. Afghanistan, C. Asia. 5. korshinskyi (H. Kraschen. ex Polj.) Y. R. Ling. Afghani- stan. S. kurramense (Qaz.) Y. R. Ling. Afghanistan, Pakistan. *S. lehmannianum (Bunge) Polj. Afghanistan, C. Asia. S. lerchianum (G. Weber in Stechm.) Polj. From SE Europe in Bulgaria through Russia and C. Asia to S. Siberia. S. lessingianum (Besser) Polj. SE Russia, C. Asia, S. Siberia. 5. leucodes (Schrenk) Polj. C. Asia. S. leucotrichum (H. Kraschen. ex Polj.) Bremer & Humphries in Y. R. Ling. Afghanistan, Pakistan, C. Asia. *S. longilobum (Osterh.) Bremer & Humphries, comb. nov. Basionym: Artemisia spiciformis var. longiloba Osterh. in Muhlenbergia 4: 69 (1908). N. America. S. maritimum (L.) Polj. W., N. and E. Europe, Iran, C. Asia and the Himalayas. S. mendozanum (DC.) Bremer & Humphries, comb. nov. Basionym: Artemisia mendozana DC., Prodr. 6: 105 (1837). W. North America. *5. minchunense Ling & Y. R. Ling. China. S. mogoltavicum (Polj.) Y. R. Ling. C. Asia. *S. mongolomm (H. Kraschen.) Ling & Y. R. Ling. China in Mongolia. S. mucronulatum (Polj.) Y. R. Ling. C. Asia. *S. namanganicum (Polj.) Polj. C. Asia. *S. nigricans (Filat. & Ladyg.) Bremer & Humphries in Y. R. Ling. C. Asia. *S. nitrosum (G. Weber ex Stechm.) Polj. SE Russia, C. Asia, S. Siberia, China in Xinjiang. *5. novum (Nelson) W. A. Weber. W. United States. S. nutans (Willd.) Sojak. SE Russia. *S. oliverianum (Gay ex Besser) Bremer & Humphries in Y. R. Ling. Iran, Afghanistan, Pakistan, C. Asia. S. oranense (Deb. ex Filat.) Y. R. Ling. NW Africa (Alge- ria). 5. oratense (Deb. & Filat.) Y. R. Ling. Algeria. S. palmeri (A. Gray) Bremer & Humphries, comb. nov. Basionym: Artemisia palmeri A. Gray in Proc. Am. Acad. 11: 79 (1876) (Artemisiastrum palmeri (A. Gray) Rydb.). N. America in California and Baja California (Mexico). S. pauciflorum (G. Weber in Stechm.) Polj. SE Russia, C. Asia, S. Siberia. S. poljakovii (Filat.) Y. R. Ling. Russia. 5. polystichum (Polj.) Y. R. Ling. C. Asia. S. porrectum (H. Kraschen. ex Polj.) Polj. C. Asia. *S. prasinum (H. Kraschen. ex Polj.) Polj. Afghanistan, C. Asia. *S. prolixum (H. Kraschen. ex Polj.) Polj. C. Asia. *S. pygmaeum (A. Gray) W. A. Weber. W. United States. *S. quettense (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia quettensis Podl., Fl. iranica 158: 212 (1986). Iran, Pakistan. S. rhodanthum (Rupr.) Polj. C. Asia. *S. rigidum (Nutt.) W. A. Weber. W. United States. S. rothrockii (A. Gray) W. A. Weber. W. United States. 5. saharum (Pomel) Y. R. Ling. Algeria, Tunisia. *S. saissanicum (H. Kraschen.) Bremer & Humphries in Y. R. Ling. C. Asia. S. santolinum (Schrenk) Polj. (5. lobulifolium (Boiss.) Polj.). Iran, C. Asia. S. santonicum (L.) Sojak (S. monogynum (Waldst. & Kit.) Polj.). SE and E. Europe to C. Asia, Turkey. S. sawanense Y. R. Ling & Humphries. China. S. schrenkianum (Ledeb.) Polj. C. Asia, S. Siberia, China in Xinjiang, Mongolia. *S. scopiforme (Ledeb.) Polj. C. Asia, S. Siberia. *5. scotinum (Nevski) Polj. Afghanistan, C. Asia. *S. semiaridum (H. Kraschen. & Lavrenko) Ling & Y. R. Ling. C. Asia. S. serotinum (Bunge) Polj. C. Asia. *5. sieberi (Besser) Bremer & Humphries in Y. R. Ling. Middle East, Iraq, Iran, Afghanistan, SW C. Asia. S. skorniakowii (Winkler) Bremer & Humphries in Y. R. Ling. C. Asia. *5. spicigerum (Koch) Polj. Turkey, Caucasus, Iran. *S. stenocephalum (H. Kraschen. ex Polj.) Polj. Afghanistan, Pakistan, C. Asia. *S. subchrysolepis (Filat.) Bremer & Humphries, comb. nov. Basionym: Artemisia subchrysolepis Filat. in Nov. Sist. Vysshikh Rast. 18: 224 (1981). C. Asia. 5. sublessingianum (Kell.) Polj. C. Asia, S. Siberia, Mongo- lia. *S. subsalsum (Filat.) Bremer & Humphries in Y. R. Ling. C. Asia. 5. szowitzianum (Besser) Polj. Caucasus. *S. tauricum (Willd.) Polj. Krym, Caucasus, Turkey. *S. tecti-mundii (Podl.) Bremer & Humphries, comb. nov. Basionym: Artemisia tecti-mundii Podl., Fl. iranica 158: 206 (1986). Afghanistan, Pakistan. 5. tenuisectum (Nevski) Polj. C. Asia. S. terrae-albae (H. Kraschen.) Polj. C. Asia, Mongolia, China. *S. thomsonianum (C. B. Clarke) Ling & Y. R. Ling. China. S. tianshanicum (H. Kraschen.) Y. R. Ling. C. Asia, China in Xinjiang, Mongolia. S. transiliense (Polj.) Polj. C. Asia. S. tridentatum (Nutt.) W. A. Weber. W. North America. 120 K. BREMER AND C. J. HUMPHRIES S. tripartitum (Rydb.) W. A. Weber. W. United States. *S. turanicum (H. Kraschen.) Polj. Afghanistan, Iran, Paki- stan, C. Asia. *S. turcomanicum (Gand.) Polj. Iran, SW & C. Asia. *S. vachanicum (H. Kraschen. ex Polj.) Polj. Afghanistan, Pakistan, C. Asia. *S. validum (H. Kraschen. ex Polj.) Polj. C. Asia. S. valesianum (Lam.) Y. R. Ling. C. Europe. S. vallesiacum (All.) Sojak. S. Europe. *S. vaseyanum (Rydb.) W. A. Weber. W. United States. 42. CROSSOSTEPHIUM Less, in Linnaea 6: 220 (1831). Type species: C. artemisioides Less. (C. chinense (L.) Makino). A tomentose shrub. Leaves alternate, narrowly spathulate, apically few-lobed or entire. Capitula rather small and rounded, paniculate, disciform. Outer involucral bracts tomentose, inner scarious. Receptacle hemispherical, epale- ate. Outer female florets tubular, 2-3-lobed, glandular. Cen- tral florets with corolla tubular, 5-lobed, glandular. Apical anther appendage subtriangular. Cypselas obovoid, weakly 5-ribbed, glandular. Pappus coroniform, of small scales. DISTRIBUTION. The Philippines, Taiwan, S. Japan, and China, where it is also widely cultivated. - Monotypic. Crossostephium chinense is a former species of Artemisia of uncertain systematic position. The rounded capitula with their pubescent outer involucral bracts recall many species of Artemisia. It appears from inflorescence, floret and pollen structure that it is related to the Artemisia group of genera. It differs by its weakly ribbed cypselas, furnished with a distinct coroniform pappus. On the basis of these characters a pos- sible relationship to the North American Artemisia califor- nica has been suggested by Gray (1884) and Rydberg (1916). Rydberg even transferred some species of Artemisia and relatives of Crossostephium on the basis of ribbed cypselas being present. However, no pappus is present and the presumed relationship was questioned by Hall & Clements (1923). 43. ARTEMISIA L., Sp. pi.: 845 (1753). Type species: A. vulgaris L. - Oligospoms Cass. (including Artemisiella A. Ghafoor) Annual and perennial herbs, half-shrubs or shrubs. Leaves alternate, variously lobed or dissected, rarely entire. Capitula disciform; inflorescence usually a long panicle but sometimes much reduced and racemose, spiciform or subglobose. Receptacle flat to conical, epaleate, sometimes pilose. Outer female florets usually tapering above, with 2-4 teeth, or truncate, commonly oblique at orifice. Central florets her- maphrodite and fertile or female-sterile and functionally male; corolla 5-lobed, yellow or sometimes purplish. Apical anther appendage lanceolate-linear to subulate. Cypselas obovoid, thin-walled, with or without rows of myxogenic cells, usually glabrous but occasionally hairy. Pappus absent. DISTRIBUTION. Predominantly N. hemisphere but with a few species also from S. America, Africa S. of Sahara and the Hawaiian Islands. Most species in temperate Eurasia and W. N. America. - 388 spp. Artemisia is the largest genus of the Anthemideae (see Krasheninnikov, 1946). Because there are so many species different authors have made numerous attempts to divide it up in some way. Problems have arisen because two of the four commonly recognized sections appear not to be mono- phyletic and attempts to separate individual genera have been carried out for a variety of different reasons, in regional isolation, without an appraisal of either all of the characters or all of the taxa. The division of Artemisia goes back to Tournefort (1700). He recognized three genera, Abrotanum, Absinthium, and Artemisia. These were based on gross morphological charac- ters and general habit. They are not the same groups as those recognized today, although the names have been retained at sectional or subgeneric level. Linnaeus (1753) united Tourne- fort's three genera into one, establishing more or less the concept of Artemisia which has been recognized ever since. Cassini (1817) established a new genus, Oligosporus, to accommodate those species with functionally separate sexes, outer female florets and central, functionally male florets with fused style-branches and reduced, abortive ovaries. This genus corresponds with the present day section (or subgenus) Dracunculus. All of the remaining taxa were included in Artemisia, Absinthium not being recognized. The next impor- tant development was by Besser (1829). Although he never completed his monograph, his results were published by De Candolle (1837). Besser established three subdivisions which were expanded to four by de Candolle as follows: Sect. Abrotanum (^Artemisia). Capitula heterogamous (dis- ciform); outer florets female, fertile; central florets perfect, fertile; receptacle glabrous. Sect. Absinthium. As Abrotanum but receptacle hairy. Sect. Dracunculus. As Abrotanum but central florets female- sterile. Sect. Seriphidium. Capitula homogamous (discoid); florets all perfect, fertile; receptacle glabrous. This arrangement has more or less persisted ever since and most treatments have fused or separated the different groups. Grenier & Godron (1850) amalgamated all four sections into one genus, Euartemisia, but Rouy (1903) by contrast raised three sections, Seriphidium, Abrotanum and Absinthium to the status of subgenera. Later Rydberg (1916) promoted Dracunculus to the same rank. Gray (1884) kept sections Seriphidium and Dracunculus but united Abrotanum and Absinthium into one new section Euartemisia. Hall & Clem- ents (1923) attempted the first phylogenetic interpretation of the four sections, on the basis of three transformations. These were receptacle becoming hairy, loss of female fertility in disc florets and a complete reduction of the female florets. The three sections Absinthium, Dracunculus and Seriphidium were considered to be three coherent groups derived as three separate lines from an ancestral Abrotanum. The most recent treatments have been by Russian and Chinese botanists (Drokhina, 1978; Poljakov, 1961, 1967; Wang, 1979; Leonova, 1971, 1980; Korobkov, 1979). Kra- scheninnikov (1946) re-sorted Gray's section Euartemisia, raising it to subgeneric rank but kept Dracunculus and Seriphidium as two separate subgenera. The most radical treatment is that of Poljakov (1961). He used the section Artemisia s. s. to accommodate distinctive species of the 'Artemisia vulgaris' -group as distinct from section Abro- tanum. This, together with sections Abrotanum and Absin- thium, comprised a smaller genus Artemisia. Seriphidium and Dracunculus (=Oligosporus) were raised to generic rank GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 121 together with some other small genera (Kaschgaria, Neopal- lasia, Turaniphytum and Mausolea). The most recent classifications of Artemisia and its allies are those of Y. R. Ling (19806, 1982, 1984 19880, b, 19916). In his 1982 treatment he recognizes nine sections of Artemisia and considers Abrotanum and Absinthium the primitive ones, from where the others, as well as a number of related genera have evolved. It is worth noting that Ling considers Seriphidium as a separate genus. Ling (1984) divides the genus into two subgenera, Artemisia and Dracunculus, and maintains the three traditional sections, Abrotanum, Absin- thium and Dracunculus. Seriphidium is kept as a separate genus and a number of new combinations are made concern- ing Chinese species. Absinthium has been variably maintained as a section, sunk into subgenus Artemisia (i. e. Abrotanum), or raised to subgeneric rank. The main distinguishing feature, and in fact the only criterion used for recognition, is the presence of a ring of receptacular hairs around the base of each flower. However, even Gray (1884) noted that in certain species there is only a partial presence, or indeed, a complete absence of this character. Poljakov (1961) maintains that it is incorrect to sink Absinthium because most species do in fact have the character. He notes also that many species of Absinthium have a dense, woolly pubescence of white silky hairs. As far as we can judge, certain species of different sections are artificially separated by this character. Further- more, there are those taxa with a dense indumentum but without receptacular hairs, and there are those without a silky indumentum but with receptacular hairs. The problem is left to an internal generic study beyond the scope of this work. The section or subgenus Dracunculus is distinguished by the fact that the pistil of the central florets is abortive and they are therefore functionally male. The central ovaries are uniformly sterile and very reduced. This condition is found also in Mausolea and Picrothamnus . We agree with Hall & Clements (1923) that Dracunculus is a monophyletic group, but only so if Mausolea and Picrothamnus are included. Future work on generic delimitation of Artemisia s. 1. will probably result in Dracunculus being removed from Artemi- sia. As a separate genus it will be named Oligosporus, following Cassini (1817) and Poljakov (1961). (Dracunculus Miller applies to plants of Araceae.) The problem of Mauso- lea and Picrothamnus and their sister group, Oligosporus as a whole or only a part of it, also has to be considered, Many species are involved and several new combinations necessary. For these reasons we have for the time being provisionally retained Oligosporus as an infrageneric taxon Dracunculus within Artemisia. Seriphidium, commonly treated as a section or subgenus of Artemisia, is recognized here as a separate genus, following Poljakov (1961) and Y. R. Ling (1982, 1984). Neopallasia and Turaniphytum are two small generic segre- gates established by Poljakov (1955, 1961). Apparently they have their sister group(s) within Artemisia, paraphyletic as presently circumscribed. A proper generic circumscription of Artemisia, considering also these segregate genera as well as Oligosporus (Artemisia sect. Dracunculus), Picrothamnus and Mausolea, is a major task considering the numerous species involved. Work on this problem has been undertaken by Yeou-Ruenn Ling at The Natural History Museum, Lon- don and Institute Sinica, Guangzhou. The list of species is compiled from the major floras, with recently described species added. From those areas, notably China, without recent floristic accounts the list must naturally be taken as rather preliminary. In those cases where major floras disagree on synonymy, the more recent treatments have generally been followed. *A. abaensis Y. R. Ling & S. Y. Zhao. China. A. abrotanum L. Eurasia, widely cultivated and introduced, also in N. America. A. absinthium L. Eurasia and N. Africa, widely introduced, also in N. America. A. abyssinica Schultz-Bip. Saudi Arabia. A. adamsii Besser. S. Siberia, Mongolia, China. * A. afghanica Rech. f. Afghanistan. A. afra Jacq. Africa S. of Sahara. *A. aksaiensis Y. R. Ling. China. A. alaskana Rydb. N. America in Alaska. A. albicerata H. Kraschen. C. Asia. A. aleutica Hulten. Aleutian Islands. *A. altaiensis H. Kraschen. S. Siberia, Mongolia. *A. amygdalina Decne. Himalayas. A. andersiana Podl. Afghanistan. A. anethifolia G. Weber in Stechm. E. Siberia, Mongolia, China. A. anethoides Mattf. China. *A. angustissima Nakai. China, Japan. A. annua L. Eurasia, widespread and introduced, also in N. America. A. anomala S. Moore. China. *A. aquatica Lour. China. A. arborescens L. S. Europe, Turkey, Middle East, N. Africa from Libya to Morocco. A. arctica Less. Siberia, Japan, W. North America. *A. arctisibirica Korobkov. Siberia. A. argyi A. Leveille & Vaniot. Far East, Mongolia, China, Korea. *A. argyrophylla Ledeb. Mongolia, China. A. armeniaca Lam. SE European Russia, Turkey, Iran, S. Siberia. A. aschurbajewii C. Winkler. C. Asia. A. atlantica Cosson & Durieu. N. Africa in Tunisia, Algeria and Morocco. A. atrata Lam. C. Europe. A. atrovirens Hand.-Mazz. China. A. aucheri Boiss. Iran, Pakistan. *A. aurata V. Komarov. Far East, China, Korea. A. australis Less. Hawaii Islands. A. austriaca Jacq. E. and E.G. Europe, Turkey, Iran, Afghanistan, C. Asia, W. Siberia, Far East, China. A. austro-himalayensis (Y. R. Ling & H. S. Puri.) Y. R. Ling & H. S. Puri. N. India. *A. austro-yunnanensis Ling & Y. R. Ling. China. *A. avarica Minat. Caucasus. *A. baimaensis Y. R. Ling & Z. C. Z. Y. Zhuo. China. *A. banihalensis Kaul & Bakshi. India. *A. bargusinensis Sprengel. E. Russia, Siberia. *A. bejdemaniae Leonova. Siberia. A. biennis Willd. Eurasia, widespread and widely introduced, also in N. America. *A. blepharolepis Bunge. Mongolia, China. A. borealis Pallas - Note: The delimitation towards A. campestris is unclear. In Flora europaea (Tutin et al., 1976) this species is treated as a subspecies of A. campestris. N. Europe, Siberia, Mongolia, China, N. America. A. borealo-siamensis Y. R. Ling. N. Thailand 122 K. BREMER AND C. J. HUMPHRIES *A. brachyloba Franchet. China. *A. brachyphylla Kitam. China, Korea. *A. brevis Pampan. China. *A. burmanica Pampan. China. A. caespitosa Ledeb. S. Siberia, Mongolia, China. A. californica Less. United States in California and Mexico in Baja California. *A. calophylla Pampan. China. A. campbellii Hook. f. & Thomson. Himalayas, China in Tibet. A. campestris L. - Note: See note under A. borealis. Wide- spread in Eurasia, N. America, and N. Africa. A. camphorata Villars. Europe. A. canariensis (Besser) Less. Canary Islands. *A. cannabifolia A. Leveille. China. A. cantabrica (Lainz) Lainz. SW Europe in Spain. A. capillaris Thunb. Far East, China, Japan, SE Asia in Malaya. A. carruthii Wood. W. United States. A. caruifolia Buch.-Ham. in Roxb. Himalayas. *A. cashimirica Kaul & Bakshi. India. A. caucasica Willd. S. European Russia, Caucasus, Turkey. A. chamaemelifolia Villars. C. and SW Europe, Caucasus, Turkey, Iran. M. chiajeana Kunze. Iran. M. chiarugii Pampan. China. *A. chienshanica Ling & W. W. Wang. China. M. chingii Pampan. China. A. chitachensis Cosson ex Battand. & Trabut. N. Africa. *A. chrysolepis Kitagawa. China. M. conaensis Ling & Y. R. Ling. China. M. congesta Kitam. Japan. A. copa Philippi. S. America (Argentina & Chile). M. coracina W. W. Wang. China. A. crithmifolia L. Europe. M. cuspidata H. Kraschen. E. Siberia. *A. daghestanica H. Kraschen. & Pors. Caucasus and adja- cent parts of Russia (Daghestan). M. dahurica (Turcz.) Polj. China. M. dalai-lamae H. Kraschen. China in Tibet. *A. demissa H. Kraschen. C. Asia, China. *A. densifolia Filat. Algeria. *A. depauperata H. Kraschen. Mongolia. A. desertorum Sprengel. Far East, E. Siberia, Mongolia, China. *A. dlmoana Popov. SW & C. Asia. A. disjuncta H. Kraschen. Mongolia, China. *A. divaricata (Pampan.) Pampan. Mongolia, China. *A. diversa Diels. China. M. dolichocephala Pampan. Himalayas. A. douglasiana Besser in Hook. f. W. United States. *A. dracunculiformis H. Kraschen. Arctic Siberia. A. dracunculus L. (A. glauca Pallas ex Willd.). Eurasia, widely cultivated and introduced, also in N. America. A. dubia Wallich ex Besser. Himalayas, China. *A. dudinensis V. P. Amel'chenko. Siberia. *A. duthreuil-de-rhinsi H. Kraschen. China in Tibet. * A. edgeworthii Balakr. China. A. eldarica Rzazade. Russia. *A. elegantissima Pampan. W. Himalayas. A. emeiensis (Chang) Y. R. Ling. China. A. eriantha Ten. C. and SW Europe. M. eriocephala Pampan. W. Himalayas. A. eriopoda Bunge. China. *A. erlangshanensis Ling & Y. R. Ling. China. *A. faurieri Nakai. Korea, Japan. A. filifolia Torrey. W. United States. A. filiformilobulata Y. R. Ling & H. S. Puri. India. A. flaccida Hand.-Mazz. China. *A. flahaultii Emb. & Maire. N. Africa in Morocco. A. flava Jurtzev. Siberia. *A. flavifolia Gilli. Afghanistan. *A. forrestil W. Smith. China. A. franserioides Greene. W. N. America. A. freyniana (Pampan.) H. Kraschen. Far East, Mongolia, China, Korea. A. frigida Willd. SE Russia, C. Asia, Siberia, Mongolia, China, N. America. M. frigidioides H. C. Fu & Z. Y. Zhu. China. M. fukudo Makino. Korea, Japan. *A. fulgens Pampan. China. A. furcata M. Bieb. E. Siberia, N. America in Alaska and Canada. M. gabriellae Braun-Blanquet. SW Europe. A. gangsuensis Ling & Y. R. Ling. China. A. genipi G. Weber in Stechm. C. Europe. *A. gilvescens Miq. China, Japan. A. giraldii Pampan. China. *A. glabella Karelin & Kir. China, C. Asia, W. Siberia, Mongolia. A. glacialis L. C. Europe. M. globosa H. Kraschen. Mongolia, China. *A. globosoides Ling & Y. R. Ling. China. A. globularia Cham, ex Besser. E. Siberia, N. America in Alaska. A. glomerata Ledeb. Arctic and E. Siberia, Japan, N. America in Alaska and Canada (Yukon). A. gmelinii G. Weber in Stechm. Himalayas, C. Asia, Siberia, Far East, Mongolia, China, Korea, Japan. A. gongshanensis Y. R. Ling & Humphries. China. *A. gorgonum Webb in Hook. Cape Verde Islands. A. granatensis Boiss. SW Europe in Spain. *A. graveolens Minat. Caucasus. *A. gyangzeensis Ling & Y. R. Ling. China. M. gyitangensis Ling & Y. R. Ling. China. M. haichowensis Chang. China. A. hallaisanensis Nakai. Korea. *A. halodendron Turcz. ex Besser. E. Siberia, Mongolia, China. *A. hancei (Pampan.) Ling & Y. R. Ling. China, Indo- China. M. haussknechtii Boiss. Turkey, Iraq, Iran. A. hedinii Ostenf. & Pauls in Hedin. China in Tibet. *A. henriettae H. Kraschen. Arctic Siberia. *A. hillebrandii Skottsb. Hawaii Islands. *A. hippolytii Butkov. Russia. A. hispanica Lam. Spain. A. hololeuca M. Bieb. ex Besser. S. European Russia. A. hulteniana Vorosch. Aleutian Islands. A. hultenii Maksimova. Far East. A. idilongensis Y. R. Ling. China. M. ifranensis J. Didier. N. Africa in Morocco. M. igniaria Maxim. China. *A. implicata Leonova. Mongolia. *A. imponens Pampan. China. A. incana (L.) Druce. Turkey, Caucasus and adjacent parts of Russia (Daghestan), Iraq, Iran. A. incisa Pampan. (A. nuristanica Kitam.). Himalayas. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 123 A. indica Willd. Himalayas, China, Taiwan, Japan. A. insipida Villars. C. Europe in France. A. insulana H. Kraschen. E. Siberia in Bering Island. A. integrifolia L. Siberia, Far East, Mongolia, China, Korea. *A. intramongolica H. C. Fu & Z. Y. Zhu. China. A. jacutica Drob. E. Siberia. A. japonica Thunb. Afghanistan, Pakistan, Far East, China, Korea, Japan, Taiwan. A. javanica Pampan. Indonesia. *A. jaxatica Polj. C. Asia. A. jilongensis Y. R. Ling & Humphries. China. A. judaica L. Middle East and N. Africa in Egypt, Libya and Algeria. *A. kabylica Chabert. N. Africa in Algeria. *A. kanashiroi Kitam. N. China. *A. kangmasensis Ling & Y. R. Ling. China. *A. karavajevii Leonova. Siberia. A. kauaiensis (Skottsb.) Skottsb. Hawaii Islands. A. kawakamii Hayata. Taiwan. A. keiskeana Miq. Far East, China, Korea, Japan. A. kelleri H. Kraschen. C. Asia. A. kitadakensis Hara & Kitam. Japan. *A. klementzae H. Kraschen. ex Leonova. Mongolia. A. klotzschiana Besser. N. America in Mexico. A. koidzumii Nakai. E. Siberia, Far East, Japan. *A. komarovii Polj. Far East. *A. kulbadica Boiss. & Buhse. Iran, C. Asia. *A. kumykorum Minat. Caucasus. A. kuschakewiczii Winkler. C. Asia. A, laciniatiformis V. Komarov. E. Siberia, Far East, N. America in Alaska. A. lactiflora Wallich ex DC. China, Taiwan. A. lagocephala (Fischer ex Besser) DC. Siberia, Far East, China. A. lagopus Fischer ex Besser. E. Siberia, Far East. A. lamprocaulos Rech. f. Iran. A. lancea Vaniot (A. feddei Levl. & Vaniot). China, Korea, Japan. A. latifolia Ledeb. Russia, C. Asia, Siberia, Mongolia, China. A. lav andulif olia DC. Far East, China, Korea. *A. lavei Kostel. China. A. ledebouriana Besser. E. Siberia. *A. leontopodioides Fischer ex Besser. E. Siberia, Aleutian Islands. A. leptophylla D. Don. Himalayas (Nepal). A. leucophylla (Turcz. ex Besser) C. B. Clarke. S. Siberia, Mongolia, China. *A. limosa Koidz. Far East. *A. limprichtii (Pampan.) Ling & Y. R. Ling. China. *A. lipskyi Polj. C. Asia. *A. littoricola Kitam. Far East, Japan. A. longifolia Nutt. Canada and W. United States. A. ludovidana Nutt. Canada and W. United States. *A. macilenta (Maxim.) H. Kraschen. Far East, China. *A. maciravae Hutch. & Dalziel. Africa in Sahara. A. macrantha Ledeb. European Russia, Siberia, Mongolia, China. A. macrocephala Jacq. Iran, Afghanistan, Himalayas, S. Siberia, Far East, Mongolia, China. *A. macrorhiza Turcz. E. Siberia. A. magellanica Schultz-Bip. S. America in Chile. A. mairei A. Leveille. China. *A. manshurica (V. Komarov) V. Komarov. Siberia, China. A. maroccana Cosson. Morocco. *A. marschalliana Sprengel. China. *A. martjanovii H. Kraschen. ex Polj. E. Siberia. A. mattfeldii Pampan. China. A. mauiensis (A. Gray) Skottsb. Hawaii Islands. A. maximovicziana (F. Schum.) H. Kraschen. ex Polj. Far East. *A. medioxima H. Kraschen. ex Polj. Far East, N. China. A. melanolepis Boiss. & Kotschy. Iran. A. mesatlantica Maire. N. Africa in Morocco. A. michauxiana Besser. W. North America. A. minor Jacq. in Besser. Himalayas, China in Tibet. *A. molinieri Quezel, Barbero & R. Loisel. SW Europe in France. *A. molluccana Roxb. SE Asia in the Moluccas. A. momiyamae Kitam. Japan. A. mongolica (Fischer ex Besser) Nakai. China. A. monophylla Kitam. Japan. A. monosperma Del. Turkey, Middle East, N. Africa in Egypt and Libya. A. montana Pampan. Far East, China, Japan. *A. montevidensis Sprengel. S. America in Argentina. *A. moorcroftiana Wallich ex DC. China. A. morrisonensis Hayata. Taiwan. *A. multisecta Leonova. C. Asia. A. mutellina Villars. C. & S. Europe. M. myriantha Wallich ex DC. China, Himalyas, Burma. *A. nakaii Pampan. China, Korea. *A. nanshanica H. Kraschen. China in Tibet. M. neglecta Leonova. C. Asia. *A. negrei Ouyahya. Morocco. *A. nesiotica Raven. W. United States. A. niitakayamensis Hayata. Taiwan. A. nilagirica (C. B. Clarke) Pampan. India, Burma. A. nitida Bertol. C. Europe. *A. nivalis Braun-Blanquet. C. Europe in Switzerland. *A. nortonii Pampan. China in Tibet. A. norvegica Fries. N. Europe, Arctic America. *A. nujianensis (Ling & Y. R. Ling) Y. R. Ling. China. A. obscura Pampan. Mongolia, China. A. obtusiloba Ledeb. C. Asia, S. Siberia, Mongolia. *A. occidentali-sichuansensis Y. R. Ling. China. A. occidentali-sinensis Y. R. Ling. China. A. occidentali-yunnanensis Ling & Y. R. Ling. China. A. oelandica (Besser) V. Komarov. N. Europe in Sweden. *A. olchonensis Leonova. Siberia. *A. olgensis (Vorobiev) Vorosch. Russia. A. oligocarpa Hayata. Taiwan. A. opulenta Pampan. N. Japan, E. Russia. *A. oranensis Filat. Algeria. *A. ordosica H. Kraschen. China. *A. orientalis (Pampan.) Ling & Y. R. Ling. China. A, orientali-hengduangensis Ling & Y. R. Ling. China. A. orientali-xizangensis Y. R. Ling & Humphries. China. A. orientali-yunnanensis Y. R. Ling. China. *A. orthobotrys Kitagawa. China. *A. oxycephala Kitagawa. China. *A. packardiae Grimes & Ertter. W. United States. *A. pollens Wallich ex Besser. India. A. palustris L. S. Siberia, Far East, Mongolia, China. Mon- golia, China, Korea. A. pancicii (Janka) Ronniger. E.G. Europe. *A. pannosa H. Kraschen. Far East. A. papposa Blake & Cronq. W. United States in Idaho. 124 K. BREMER AND C. J. HUMPHRIES A. parry i A. Gray. W. United States. A. parviflora Buch.-Ham. ex Roxb. China. A. pattersonii A. Gray. W. United States. M. pedatifida Nutt. W. and C. United States. A. pedunculosa Miq. Japan. M. pengchuoensis Y. R. Ling & S. Y. Zhao. China. A. persica Boiss. Iran, Afghanistan, Himalayas, C. Asia. M. pewzowii Winkler. China in Tibet. M. phaeolepis H. Kraschen. S. Siberia, Mongolia, China. M. phyllobotrys (Hand.-Mazz.) Ling & Y. R. Ling. China. M. polybotryoidea Y. R. Ling. China. A. pontica L. C. and E. Europe, W. Siberia, introduced in N. America. M. porteri Cronq. W. United States in Wyoming. M. praticola Klokov. Ukraine. M. prattii (Pampan.) Ling & Y. R. Ling. China. A. princeps Pampan. China, Korea, Japan, Taiwan. M. przewalskii H. Kraschen. China. A. pseudopontica Schur. E. Europe. M. pubescens Ledeb. China. M. punctigera H. Kraschen. Far East, European Russia, E. and W. Siberia, Mongolia, N. America. M. pycnorhiza Ledeb. C. Asia, S. Siberia, Mongolia. *A. quinlingensis Ling & Y. R. Ling. China. *A. quinqueloba Trautv. C. Asia. M. ramosa C. Smith. Canary Islands. M. rehan Chiov. Africa in Ethiopia. M. remotiloba H. Kraschen. ex Polj. E. Siberia. A. reptans C. Smith ex Link. SW Europe in Spain, N. Africa in Morocco, Canary Islands. M. robusta (Pampan.) Ling & Y. R. Ling. China. M. rosthornil Pampan. China. A. roxburghiana Besser. Afghansitan, Pakistan, Himalayas, China. M. rubripes Nakai. Far East, Mongolia, China, Korea, Japan. A. rupestris L. N. Europe, C. Asia, W. Siberia, Mongolia, China, N. America in Canada. A. rutifolia Stephen ex Sprengel. Iran, Afghanistan, Himala- yas, C. Asia, Siberia, Mongolia, China. A. sacrorum Ledeb. China, Korea, Japan, Himalayas, C. Asia, Afghanistan. M. saitoana Kitam. Far East, Korea. A. salsoloides Willd. Russia, W. Himalayas, W. Siberia, China in Tibet. *A. samoiedorum Pampan. Arctic Siberia. A. santolinifolia Turcz. ex H. Kraschen. Afghanistan, China in Tibet, European Russia, C. Asia, S. Siberia, Mongolia, Pakistan. M. saposhnikovii H. Kraschen. ex Polj. C. Asia. A. schimperi Schultz-Bip. ex Schweinf. Africa in Ethiopia. *A. schischkinii H. Kraschen. Mongolia. A. schmidtiana Maxim. Far East, Japan. A. scoparia Waldst. & Kit. C. and E. Europe, Turkey, Middle East, Iran, Himalayas, C. Asia, Siberia, Mongolia, China, Japan, N. Africa in Egypt. A. scopulorum A. Gray. W. United States. A. selengensis Turcz. ex Besser. E. Siberia, Far East, Mongo- lia, China. A. senjavinensis Besser. E. Siberia, N. America in Alaska. A. sericea G. Weber in Stechm. Russia, Siberia, Mongolia, China. A. serrata Nutt. W. United States. *A. serreana Pampan. China. *A. shangnanensis Ling & Y. R. Ling. China. *A. shennongjaensis Ling & Y. R. Ling. China. *A. sichuanensis Ling & Y. R. Ling. China. A. sieversiana Ehrh. in Willd. European Russia, Himalayas, C. Asia, Siberia, Far East, China. *A. simulans Pampan. China. A. sinanensis Yabe. Japan. A. sinensis (Pamp.) Ling & Y. R. Ling. China. A. smithii Mattf. China. M. somai Hayata. Taiwan. A. songarica Schrenk. C. Asia, China. M. speciosa (Pampan.) Ling & Y. R. Ling. China. *A. sphaerocephala H. Kraschen. Mongolia, China. A. splendens Willd. Turkey, Caucasus, Iraq, Iran. A. stelleriana Besser. Far East, China, introduced in N. Europe and N. America, Japan. A. stenophylla Kitam. China, Korea, E. Russia. A. stipularis Urb. & Ekman. Haiti. A. stolonifera (Maxim.) V. Komarov. Far East, China, Japan. A. stracheyi Hook. f. & Thomson ex C. B. Clarke (Note added in proof. This taxon was recently removed by Ghafoor (1992) and described as a monotypic genus under the name Artemisiella stracheyii (Hook. f. & Thomson ex C. B. Clarke) Ghafoor). Himalayas, China in Tibet. A. stricta Edgew. Iran, Himalayas, China in Tibet. *A. subulata Nakai. Soviet Far East, China, Korea. A. subviscosa Turcz. E. Siberia. M. succulenta Ledeb. C. Asia, W. Siberia. *A. succulentoides Ling & Y. R. Ling. China. A. suksdorfii Piper. W. Canada and United States. *A. super ba Pampan. Mongolia. A. swatensis Podl. Pakistan. A. sylvatica Maxim. Far East, Mongolia, N. China. *A. tafelii Mattf. China in Tibet. A. taibaishanensis Y. R. Ling & Humphries. China. *A. tainingensis Hand.-Mazz. China. A. tanacetifolia L. China, N. Korea, C. and W. Russia, Europe, N. America. A. tangutica Pampan. China. A. tenuifolia Y. R. Ling & H. S. Puri. China. A. thellungiana Pampan. SW China, N. India, Sikkim. A. tilesii Ledeb. Arctic Eurasia and Arctic N. America. A. tomentella Trautv. C. Asia. A. tournefortiana Reichenb. Turkey, Caucasus, Iran, Afghanistan, Himalayas, C. Asia, Mongolia, China. M. transbaicalensis Leonova. Siberia. M. trautvetteriana Besser. S. European Russia. A. tridactyla Hand.-Mazz. China. M. triniana Besser. Arctic Siberia. A. tschernieviana Besser. E. Europe, C. Asia, China. *A. tsugitakaensis (Kitam.) Ling & Y. R. Ling. China. *A. tsuneoi Tatewaki & Kitam. Japan. *A. tukuchaensis Kitam. Himalayas (Nepal). A. tyitangensis Ling & Y. R. Ling. China. A. unalaskensis Rydb. Far East, N. America in Alaska. *A. ussuriensis Polj. Far East. *A. velutina Pampan. China. M. verbenacea (V. Komarov) Kitagawa. China. A. verlotorum Lamotte. China, Himalayas, Malaya, natural- ized in W. and C. Europe and in N. Africa, S. America. A. vestita Wallich ex Besser. W. Himalayas. A. vexans Pampan. China, Himalayas. *A. viridisquama Kitam. China. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 125 *A. viridissima (V. Komarov) Pampan. China. *A. viscida (Mattf.) Pampan. Himalayas, China. *A. viscidissima Ling & Y. R. Ling. China. A. vulgaris L. Widespread in Eurasia and N. America, also in N. Africa, widely introduced e. g. in Australia. *A. waltonii J. R. Drumm. ex Pampan. China in Tibet. M. wellbyi Hemsley & Pears. China, Himalayas. *A. wudanica Liou & W. Wang. China. *A. xanthochloa H. Kraschen. Mongolia, China. *A. xerophytica H. Kraschen. Mongolia, China. *A. xigazeensis Ling & Y. R. Ling. China. *A. yadongensis Ling & Y. R. Ling. China. A. yongil Y. R. Ling. China. *A. younghusbandii J. R. Drumm. China in Tibet. A. yunnanensis Jeffrey ex Diels. China. M. zayuensis Ling & Y. R. Ling. China. A. zhongdianensis Y. R. Ling. China. 44. NEOPALLASIA Polj. in Bot. Mater. Gerb. hot. Inst. V. A. Komarova 17: 429 (1955). Type species: N. pectinata (Pallas) Polj . Annual or biennial herbs. Leaves alternate, pectinate- pinnatisect with filiform, apically somewhat swollen and mucronulate lobes. Capitula rather small and rounded in a narrow spiciform panicle, disciform. Receptacle narrowly conical, epaleate. Outer female florets narrowly tubular, without teeth. Central florets of two kinds; outer perfect, inner completely sterile with reduced ovaries. Apical anther appendage ovoid-lanceolate and acuminate. Cypselas arranged around the base of the receptacle, oblong-obovoid, somewhat compressed or triquetrous, thin-walled, with many rows of myxogenic cells. Pappus absent. DISTRIBUTION. C. Asia, S. Siberia, Mongolia and China. - 3 spp. Poljakov (1955) distinguished Neopallasia from Artemisia by the characteristic pectinate leaves, the apically truncate (with- out teeth) outer female florets, the presence of completely sterile central florets (in addition to perfect ones) situated at the apex of a narrowly conical receptacle, the ovoid- lanceolate and attenuate anther appendages and the rosette- shaped arrangement of the cypselas around the receptacle. These characters are autapomorphies, though the shape of the anther appendages is hardly spectacular considering the variation present within the subtribe, and the cypsela arrangement follows from the sex distribution within the head, fertile florets being restricted to the outer part or the base of the conical receptacle. The immediate relatives or the sister group of Neopallasia is not easy to identify. Poljakov suggested that the genus is related to Artemisia sect. Dracunculus because some of the central hermaphrodite florets are sterile. It does seem clear that Neopallasia has its sister group within a presently para- phyletic Artemisia, possibly within sect. Dracunculus as sug- gested by Poljakov. Y. R. Ling (1980a) has recently added two Chinese species. The material of those were formerly considered part of N. pectinata s. 1. N. pectinata (Pallas) Polj . N. tibetica Y. R. Ling *N. yunnanensis (Pampan.) Y. R. Ling 45. TURANIPHYTUM Polj. in Komarov, Fl. URSS 26: 880 (1961). Type species: T. eranthemum (Bunge)Polj. Perennial herbs, somewhat woody at the base. Leaves alter- nate to rosulate, pinnatisect. Capitula disciform; inflores- cence a spike of glomerules with densely congested capitula, or rarely capitula solitary in interrupted, partly congested spikes. Receptacle convex to hemispherical, epaleate. Outer female florets subtended by scaphoid inner involucral bracts, unequally crenate at the apex. Central florets 5-lobed, api- cally with long rigid somewhat reddish hairs at the apex. Apical anther appendage lanceolate, acuminate. Cypselas obliquely oblong-obovoid, thin-walled, with rows of myxo- genic cells. Pappus absent. DISTRIBUTION. C. Asia. - 2 spp. Turaniphytum was distinguished from Artemisia mainly because of the peculiar inflorescence, with the capitula aggre- gated into glomerules and arranged in long spikes. These are presumably transformed paniculate inflorescences of the common Artemisia type. Turaniphytum also has scaphoid inner involucral bracts, subtending the outer female florets. The immediate relatives of Turaniphytum are unknown. It may have its sister group within Artemisia. T. codringtonii (Rech. f.) Polj. (T. kopetdaghense Polj.). Afghanistan. T. eranthemum (Bunge) Polj. 46. MAUSOLEA Polj. in Trudy Inst. Bot. Alma-Ata 11: 170 (1961). Type species: M. eriocarpa (Bunge) Polj. A virgate shrub. Leaves alternate, few-lobed or entire. Capitula small and subglobose, rather few and more or less sessile in a reduced panicle, disciform. Receptacle epaleate. Outer female florets without corolla; style-branches dilated, lanceolate, flat, acute. Central florets 5-lobed, apically with bifurcate hairs, hermaphrodite and female-sterile; ovaries reduced and style-branches fused. Apical anther appendage narrowly lanceolate-linear. Cypselas obovoid, densely pilose. Pappus absent. DISTRIBUTION. Iran, Afghanistan, and C. Asia. - Mono- typic. Mausolea was separated from Artemisia mainly because of the corollaless marginal flowers. The styles of the marginal flowers are also further modified compared to those of Artemisia, being wider and lanceolate. It is probably related to Picrothamnus as discussed under that genus. The cypsela hairs are straighter and less cobwebby in Mausolea compared to Picrothamnus. 47. PICROTHAMNUS Nutt. in Trans. Amer. Philippi Soc. II, 7: 417 (1841). -Type species: P. desertorum Nutt. A basally much woody shrublet with older branches trans- formed into long spines. Leaves alternate, few-lobed. Capitula small and subglobose, solitary or few together along the branches, almost sessile, disciform. Receptacle epaleate. Outer female florets tubular. Central florets 5-lobed, with long cobwebby bifurcate hairs, hermaphrodite and female- sterile; ovary reduced and style-branches fused. Apical 126 K. BREMER AND C. J. HUMPHRIES anther appendage lanceolate-linear. Cypselas obovoid, thin- walled, densely cobwebby-pilose with bifurcate hairs. Pappus absent. DISTRIBUTION. N. America in western United States. - Monotypic. Picrothamnus was established by Nuttall but reduced by Eaton (in Watson, 1871), a classification accepted by most later authors. Hence it is generally known as Artemisia spinescens D. C. Eaton. Hall & Clements (1923) consider it 'in all essentials an Artemisia of the section Dracunculus\ The spiny habit and the cobwebby-pilose corollas are autapo- morphies of Picrothamnus. The cobwebby-pilose cypselas are shared with Mausolea, the probable sister group. Together they are related to Artemisia sect. Dracunculus because of their functionally male central florets with reduced ovaries and fused style-branches. Pending a revised generic delimita- tion of Artemisia we think these genera should be retained rather than sunk in Artemisia. 1. ACHILLEINAE Bremer & Humphries, subtrib. nov. Type species: Achillea millefolium L. Herbae annuae vel perennes vel suffrutices. Receptaculum paleaceum. Corolla flosculorum disci tubo varie saccato et incrassato, saltern adaxialiter basi saccato. Cypselae parieti- bus tenuibus, plerumquefasciculis vascularibus duobus later- alibus interdum etiam fascicule uno adaxiali vel nonnunquam fasciculis 4-5 instructae, interdum compressae, saepe cellulis mucilaginis instructae. Pappus nullus. Annual or perennial herbs or shrublets. Leaves variously dissected, sometimes vermiform, rarely few-lobed or entire. Capitula solitary or corymbose, radiate or discoid. Recep- tacle variously shaped, often conical, paleate. Ray floret limb white or yellow. Disc corolla 5-lobed; tube variously saccate and thickened in fruit, basally saccate at least adaxially. Cypselas thin-walled, generally with 2 lateral and with or without 1 adaxial strand, sometimes 4-5-stranded, sometimes flattened, often with myxogenic cells. Pappus absent. DISTRIBUTION (Table 16): Eurasia and N. Africa, mainly in S. Europe, the Mediterranean and SW Asia, also in N. America, some species of Achillea widely introduced also in the S. hemisphere and one species of Santolina introduced in N. America. - 9 genera, 147 spp. Some groupings within this subtribe have been recognized earlier. The relationship between Achillea, Anacydus, and Leucocyclus has been pointed out by Humphries (1979) and phytochemical investigations have indicated a close relation- ship between Chamaemelum and Cladanthus. Both genera accumulate similar thiophene derivatives (Greger, 1977). Phytochemistry has also contributed to the recognition of subtribe Achilleinae. Achillea and Anacydus, Chamaemelum and Cladanthus, as well as Otanthus, all synthesize amides (Greger, 1977; Bohlmann et al., 1973). Santolina has a different chemistry, plesiomorphic in being similar to Tanace- tum, but it is here provisionally accepted in Achilleinae because of its adaxially saccate corolla and the paleate receptacle. Leucocyclus, Mecomischus, and Rhetinolepis , three little-known North African genera, are also included in Achilleinae and considered related to Chamaemelum and Cladanthus on a number of morphological characters. Meco- mischus and Rhetinolepis have not been investigated chemi- cally. The cladogram is one of nine equally parsimonious cla- dograms. They involve rearrangements at the base and among the last four genera. In all cladograms and hence also in the strict consensus tree, clades Ac4 (Achillea, Anacydus, Leucocyclus), Ac5 (Anacydus and Leucocyclus), Ac6 (Meco- mischus, Chamaemelum, Rhetinolepis, Cladanthus), and Ac8 (Rhetinolepis and Cladanthus) were present. Clades and characters - Fig. 8, Tables 2, 17. ^F= 48 Santolina 49 Otanthus Ac 2 lt= 50 Achillea 51 Anacyclus 52 Leucocyclus 53 Mecomischus 54 Chamaemelum fi= 55 Rhetinolepis = 56 Cladanthus [7= =Ac3 Fig. 8 Cladogram (of nine possible) of the Achilleinae produced by the ie option. Cladogram length = 34, consistency index = 88, retention index = 85. Clade Acl - subtribe Achilleinae 45 Receptacle paleate. 82 Disc corolla tube thickened in fruit. This character is variously strongly expressed in the different genera, most clearly in Otanthus. It is hardly evident in Achillea, Anacy- dus, and Leucocyclus. Table 16 General distribution of Achilleinae and genera. x=indigenous, o=introduced. Achilleinae Santolina Otanthus Achillea Anacydus Leucocyclus Mecomischus Chamaemelum Rhetinolepis Cladanthus N.Am. Eur- Asia C.&E. SW Asia Asia S.Eur. N.Afr. S.Afr. Austr. S.Am. N.Zeal. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 127 Table 17 Data matrix for the Achilleinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 11 1 111 111 1 14505765 4887 38 28351116284358 351471 53271811 524225199608479175194017401836 4&.Santolina al?allla 49. Otanthus 01?al?la 50. Achillea alaallla 51. Anacyclus alaal?la 52.Leucocyclus alaal?10 53. Mecomischus alaal?la 54.Chamaemelumalaallla 55.Rhetinolepis al?al?la 56.Cladanthus a!0al?la allll lOOOOOOOOOOpOOOOOOOOOpOOO al 1 101 1 1 1 100000000000000000000 aOHOplOpOlllOOpOOOOOOOOOOpOOO aOl lOplOOOl 1 1 1 1 aOOOOOOpOOOOOOO a01100?0001 1 1 1 101 1000000000000 al 1 10070001 OOOOOOOal 1 ?pOOOOOOO alllOplOOOlOOOOOOOalllpOOOOOOO all 10170001000000001 171111 1000 all 1001000100000001 111 111101 11 1111 11 1 167772555556666617 141383 3605613567905678513246726 4&.Santolina 49. Otanthus ??????????????????0000000 50.Achillea ???OOOOOOOOOOOOOOOOOppppO Sl.Anacyclus ???OOOOOOOOOOOOOOOOOpOOOO 52.Leucocyclus ?????00000000000000000000 53.Mecomischus ?????00000000000000000000 54.C/w/nfle/ne/Hm???OOOOOOOOOOOOOOOOOOOOOp 55.Rhetinolepis ???????????????????000000 56.Cladanthus ???0000000000000000000000 84 Disc corolla tube basally saccate at least adaxially. 172 Pappus absent in ray and disc cypselas. Santolina 2 Plants shrubby. 35 Capitula discoid. Some Anacyclus and Chamaemelum species are also discoid, as well as Otanthus and Rhetinolepis. Clade Ac2 181 Amides present. Leucocyclus, Mecomischus , and Rhetin- olepis have not been investigated chemically. Otanthus 9 Plants covered with a dense greyish-white indumentum. 15 reversed. Leaves not variously deeply lobed or divided, but entire or crenulate only. 29 Capitula densely corymbose. Most species of Achillea also have densely corymbose capitula. 35 Capitula discoid. 86 Corolla basally copiously swollen and spongy, almost enclosing the cypsela especially laterally. Clade Ac3 130 Cypselas with 2 lateral vascular strands, sometimes also with 7 adaxial strand. This cypsela vascularization is also characteristic of a large part of subtribe Matricariinae, but the two groups do not seem closely related. Clade Ac4 58 Ray and disc floret tube dorsiventrally flattened. 82 reversed. See clade Acl. 114 Cypselas dorsiventrally flattened. Flattened cypselas occur in various subtribes. Achillea There is no obvious autapomorphy for this large genus. Clade Ac5 117 Cypselas laterally winged. Some genera of the Cotula group (Matricariinae) also have winged cypselas. As noted above (clade Ac3) the groups are not closely related. 119 Cypselas with sclerenchymatic lateral wings. Anacyclus 61 Ray floret tube persistent on the cypsela. This character occurs also in some genera of subtribe Thaminophyllinae. Leucocyclus 27 Leaves vermiform. Some species of Santolina also have vermiform leaves. 51 reversed. Floral parts without resin canals. 85 Disc corolla deeply and equally saccate both abaxially and adaxially. Clade Ac6 41 Involucral bracts wide, flabelliform. This character reverses in the small-headed genus Rhetinolepis. 139 Cypselas completely covered with rows ofmyxogenic cells. 154 Cypselas thin-walled, obovoid to oblanceolate, devoid of ribs. The same type of cypselas is characteristic of the unrelated Artemisia and allies in subtribe Artemisiinae. Mecomischus There is no obvious autapomorphy for this little-known genus. Clade Ac7 180 Particular thiophene derivatives present. Rhetinolepis has not been investigated chemically. Chamaemelum There is no autapomorphy for Chamaemelum. The genus is variable in several characters. Clade Ac8 I Plants annual. Some species of Chamaemelum and other genera of the Achilleinae are also annuals. 7 Plants with branches in whorls below the first capitula. The capitula of Rhetinolepis and Cladanthus are very different in size and shape, but sessile (following character) and arranged similarly. 34 Capitula sessile along the stems. 50 Receptacular paleae pilose. This character also occurs in some species of Chamaemelum, as well as in the unrelated Eriocephalus in subtribe Matricariinae. Rhetinolepis II Plants with dolabriform hairs. 35 Capitula discoid. 128 K. BREMER AND C. J. HUMPHRIES 41 reversed. See clade Ac6. Cladanthus 48 Receptacle pilose. Some genera of subtribe Thaminophylli- nae also have pilose receptacles. 52 reversed. Ray floret limb not white, but yellow. 73 Disc corolla lobes with dorsal appendages. Corolla lobe appendages occur also in various genera of subtribes Leucan- theminae and Matricariinae. 116 Disc cypselas laterally flattened. Laterally flattened but otherwise quite different cypselas occur in subtribe Chrysan- theminae. 48. SANTOLINA L., Sp. pi: 842 (1753). Type species: S. chamaecyparissus L. Shrublets. Leaves alternate, dentate to pinnatifid or pinnati- sect, sometimes vermiform. Capitula solitary, pedunculate, discoid. Receptacle convex, paleate; paleae with a central resin canal. Corolla 5-lobed, basally saccate around the cypsela especially adaxially, with a rather long bent tube and a distinct limb. Cypselas 3-5-angled, sometimes with myxo- genic cells. Pappus absent. DISTRIBUTION. S. Europe, mainly in Spain but extending to Jugoslavia, and N. Africa in Morocco and Algeria, one species (S. rosmarinifolid) introduced in N. America. - 8 spp. Santolina is provisionally placed in Achilleinae. It differs in chemistry from the other genera of this subtribe, containing polyacetylenes and other substances similar to those in Tan- acetum (Greger, 1977). It may be a derivative of that genus rather than the sister group of the genera of Achilleinae. Although some efforts have already been made to identify taxa based on chromosome numbers (e.g. Arrigoni, 1977) Santolina is in need of revision. There are two species, S. chamaecyparissus and S. rosmarinifolia, with numerous syn- onyms and many variants. A few 'species' have been described from North Africa. Apparently they belong under the complex European species. S. chamaecyparissus L. S. elegans Boiss. ex DC. *S. insularis (Gennari ex Fieri) Arrig. *S. ligustica Arrig. *S. marchi Arrig. 5. oblongifolia Boiss. 5. rosmarinifolia L. 5. viscosa Lagasca 49. OTANTHUS Hoffsgg & Link, Fl. portug. 2: 364 (1889). Type species: O. maritimus (L.) Hoffsgg & Link - Diotis Desf. A suffruticose perennial covered with a dense greyish-white indumentum. Leaves alternate, entire or crenulate. Capitula corymbose, discoid. Receptacle convex, paleate; paleae with a central resin canal. Corolla 5-lobed, basally copiously swollen and spongy, almost enclosing the cypsela especially laterally. Cypselas with 4-5 weak ribs, thin-walled, glandular. Pappus absent. Amides present. DISTRIBUTION. Europe, N. Africa, and SW Asia extending from Ireland to Caucasus, along sea shores, mainly in the Mediterranean. - Monotypic. This characteristic species is known also as Diotis candidis- sima Desf. The interrelationships of Otanthus have been obscure but it seems that it is an autapomorphic member of subtribe Achilleinae, sharing the same chemistry as most members of the subtribe. The copiously swollen corolla base also indicates the same relationship, though the character is not so extremely developed in the other genera. 50. ACHILLEA L., Sp. pi.: 896 (1753). Type species: A. millefolium L. Perennial herbs generally with rhizomes. Leaves alternate, pinnatisect, lobed or rarely entire. Capitula comparatively small, generally corymbose or rarely few together or solitary, radiate or rarely discoid. Receptacle flat to convex or conical or rarely much elongated, paleate; paleae sometimes with a central resin canal. Ray florets female, fertile; limb rather short and wide, white or yellow; tube more or less flattened. Disc corolla 5-lobed, more or less flattened, basally slightly saccate around the cypsela especially adaxially, white, yel- low, or pink. Cypselas dorsiventrally flattened, with two lateral ribs with vascular strands, a third adaxial vascular strand rarely present. Pappus absent. Amides present. DISTRIBUTION. Europe and temperate Asia, some species also in N. Africa, a few species, in general A. millefolium (s. 1.) naturalized in N. America and also in the S. hemisphere; most species in SE Europe and SW Asia. - 115 spp. Achillea is a large genus, but with respect to floral characters, homogeneous and well defined. The cypselas are flattened and thin-walled with 2 lateral ribs and the corolla is basally slightly saccate around the cypsela (Khandzhyan, 1983). The list of species is compiled from standard floras. A. abrotanoides (Vis.) Vis. SE Europe. A. absinthoides Hal. SE Europe (Greece). A. acuminata (Ledeb.) Schultz-Bip. E. Siberia, Far East, Mongolia, China, Japan. A. aegyptiaca L. SE Europe (Greece). A. ageratifolia (Smith in Sibth. & Smith) Boiss. SE Europe. A. ageratum L. S. Europe and N. Africa in Morocco. A. aleppica DC. Turkey, Middle East, Iraq, Iran. *A. alpina L. E. Siberia, Far East, Mongolia, China, Himala- yas. *A. ambrosiaca (Boiss. & Heldr.) Boiss. SE Europe (Greece). A. arabica Kotschy. Middle East. *A. asiatica Serg. C. Asia, Siberia, Far East, Mongolia, China. A. asplenifolia Vent. E. Europe. A. atrata L. C. Europe. A. aucheri Boiss. Iran. A. barbeyana Heldr. & Heimerl in Heimerl. SE Europe (Greece). A. barrelieri (Ten.) Schultz-Bip. S. Europe (Italy). A. biebersteinii Afan. (A. micrantha Willd., A. micranthoides Klokov). E. Europe, S. European Russia, Turkey, Cauca- sus, Middle East, Iran, Afghanistan, C. Asia. A. biserrata M. Bieb. Caucasus. *A. boissieri (Hausskn.) Boiss. Turkey. *A. brachyphylla Boiss. & Hausskn. in Boiss. Turkey. *A. bucharica Winkler. C. Asia. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 129 A. callichroa Boiss. Iran. A. camtschatica Rupr. ex Heimerl. E. Siberia. A. cappadocica Hausskn. & Bornm. Turkey. A. cartilaginea Ledeb. ex Reichenb. (A. septentrionalis (Serg.) Botsch.). E. Europe, Siberia, C. Asia. A. chamaemelifolia Pourret. SW Europe. A. chrysocoma Friv. SE Europe. A. clavennae L. C. and SE Europe. A. clypeolata Sibth. & Smith SE Europe. A. coarctata Poiret in Lam. SE Europe, Turkey. A. collina J. Becker ex Reichenb. C. and SE Europe. A. compacta Willd. SW Europe, S. Russia. A. conferta DC. Syria, Iraq, Iran. A. cretica L. SE Europe, Cyprus, Turkey. A. crithmifolia Waldst. & Kit. C. and SE Europe. *A. cucullata (Hausskn.) Bornm. Turkey. *A. cuneatiloba Boiss. & Buhse. Caucasus, Iran. *A. decolorans Schrader. Turkey. A. depressa Janka. E. and SE Europe. A. distant Waldst. & Kit. ex Willd. C. Europe. A. erba-rotta All. C. and SE Europe. A. falcata L. Turkey, Middle East, Iraq. A. filipendulina Lam. Caucasus, Iran, Afghanistan, C. Asia. A. fraasii Schultz-Bip. Turkey, SE Europe. A. fragrantissima (Forssk.) Schultz-Bip. N. Africa in Egypt, Middle East, Iraq. A. gerberi Willd. W. Asia, S. Europe, Russia. A. glaberrima Klokov. S. European Russia. A. goniocephala Boiss. & Bal. in Boiss. Turkey. A. grandifolia Friv. SE Europe, Turkey. A. griseo-virens Albov. Caucasus. A. gypsicola Huber-Mor. Turkey. A. holosericea Sibth. & Smith. SE Europe. A. impatiens L. E. Europe (Romania), Siberia, China. A. inundata Kondr. in Wissjul. S. European Russia. *A. japonica Heimerl. Far East, China, Japan. M. kellalensis Boiss. & Hausskn. in Boiss. (A. haussknechtii Boiss.). Iran. A. kotschyi Boiss. Turkey. M. latiloba Ledeb. ex Nordm. Caucasus. *A. ledebourii Heimerl. S. Siberia, China. A. leptophylla M. Bieb. E. Europe, S. European Russia, N. Africa in Morocco and Algeria. A. ligustica All. S. Europe, N. Africa in Morocco, Algeria and Tunisia. A. lingulata Waldst. & Kit. SE Europe. A. lucana Pign. Italy. A. lycaonica Boiss. & Heldr. in Boiss. Turkey. *A. macrocephala Rupr. Far East, Japan. A. macrophylla L. C. Europe. A. magnified Huber-Mor. Turkey. A. maura Humbert. N. Africa in Morocco. A. membranacea (Labill.) DC. Turkey, Middle East, Iraq. A. millefolium L. (A. lanulosa Nutt., A. sudetica Opiz). Widespread in Eurasia and N. America, introduced in Australia and New Zealand. *A. monocephala Boiss. & Bal. in Boiss. Turkey. A. multifida (DC.) Boiss. Turkey. A. nana L. C. Europe. A. nobilis L. (A. neilrichii A. Kerner). S. and C. Europe, European Russia, W. Siberia, Turkey, Caucasus, Iran, C. Asia. A. ochroleuca Ehrh. E. Europe. A. odorata L. C. and SW Europe, N. Africa in Morocco and Algeria. A. oligocephala DC. Turkey, Middle East, Iraq, Iran. A. oxyloba (DC.) Schultz-Bip. C. and E. Europe. A. oxyodonta Boiss. Iran. *A. pachycephala Rech. f. Iran. A. pannonica Scheele. C., E. and SE Europe. A. phrygia Boiss. & Bal. in Boiss. Turkey. A. pindicola Hausskn. SE Europe (Greece). *A. pseudoaleppica Huber-Mor. Turkey. A. ptarmica L. Widespread in Eurasia, introduced in N. America. A. ptarmicifolia (Willd.) Rupr. ex Heimerl. Caucasus. A. ptarmicoides Maxim. E. Siberia, Far East, China, Japan. A. pyrenaica Sibth. ex Godron in Gren. & Godron. SW Europe. A. roseo-alba Ehrend. C. Europe. M. sachokiana Sosn. Caucasus. A. salicifolia Besser. European Russia, Siberia, C. Asia, China. A. santolina L. Throughout N. Africa, Middle East, Iraq, Pakistan. A. santolinoides Lagasca. SW Europe, N. Africa in Morocco and Algeria. *A. schischkinii Sosn. Turkey, Caucasus. M. sedelmeyeriana Sosn. Caucasus. *A. serbica Nyman (A. schurii Schultz-Bip.). SE Europe. A. setacea Waldst. & Kit. S., C. and SE Europe, European Russia, S. Siberia, Turkey, Iran, Afghanistan, C. Asia, China. A. sibirica Ledeb. Siberia, Japan, N. America in Alaska and Canada. A. sieheana Stapf. Turkey. A. sintenisii Huber-Mor. Turkey. A. sipikorensis Hausskn. & Bornm. Turkey. *A. spinulifolia Fenzl ex Boiss. Turkey. A. stricta (Koch) Schleicher ex Gremli. C. Europe. A. talagonica Boiss. (A. oxylepis Boiss. & Hausskn. in Boiss.). Iran. A. tanacetifolia All. Europe. A. taygetea Boiss. & Heldr. in Boiss. SE Europe (Greece). A. tenuifolia Lam. Turkey, Caucasus, Iran. *A. teretifolia Willd. Turkey. A. thracica Velen. E. Europe. A. tomentosa L. SW Europe. A. umbellata Sibth. & Smith. SE Europe (Greece). A. vermicularis Trin. Turkey, Caucasus, Iraq, Iran. M. virescens (Fenzl) Heimerl in A. Kerner. SC Europe. A. wilhelmsii Koch (A kermanica Gand.). Turkey, Cauca- sus, Syria, Iraq, Iran, Afghanistan, Pakistan, C. Asia. *A. wilsoniana Heimerl ex Hand.-Mazz. China. 51. ANACYCLUS L., Sp. pi: 892 (1753). Type species: A. valentinus L. Annual or perennial herbs. Leaves alternate, rarely rosulate, pinnatisect. Capitula solitary or laxly corymbose, peduncu- late, rarely closely aggregated, radiate or discoid. Receptacle flat to conical, paleate. Ray florets female, fertile; tube flattened, persistent on the cypselas; limb white or yellow, abaxially sometimes reddish. Disc corolla 5-lobed, sometimes slightly zygomorphic with 2 larger lobes; tube somewhat flattened and adaxially slightly saccate. Cypselas dorsiven- trally flattened and laterally winged, rather thick-walled, 130 apically sometimes coroniform, sometimes with myxogenic cells; wings thick and sclerenchymatic. True pappus absent. Amides present. DISTRIBUTION. Mainly W. Mediterranean; N. Africa, S. Europe and the Middle East. - 12 spp. Anacyclus was revised and discussed in detail by Humphries (1979), who also indicated Leucocydus as the sister group. A. davatus (Desf.) Pers. A. homogamos (Maire) Humphries A. incomtans Pomel A. latealatus Huber-Mor. A. linearilobus Boiss. & Reuter A. maroccanus (Ball) Ball A. monanthos (L.) Thell. (A cyrtolepidiodes Pomel) A. nigellifolius Boiss. A. offidnarum Hayne A. pyrethrum (L.) Lagasca A. radiatus Lois. A. valentinus L. 52. LEUCOCYCLUS Boiss. in Diagn. pi. orient. I (II): 14 (1849). Type species: L. formosus Boiss. A perennial herb. Leaves alternate, vermiform, pinnatisect. Capitula solitary, pedunculate, radiate. Receptacle flat to convex, paleate. Ray florets female, fertile; tube flattened, both adaxially and abaxially vaginate around top of cypsela; limb white. Disc corolla 5-lobed; tube flattened, both adaxi- ally and abaxially but not laterally vaginate around top of cypsela. Cypselas dorsiventrally flattened and laterally winged, rather thick-walled; wings thick and sclerenchymatic. Pappus absent. DISTRIBUTION. SW Asia in Turkey. - Monotypic. This monotypic genus is the sister group of Anacydus, as shown by Humphries (1979). Recent work by Valent- Vetschera (1982) has indicated that Leucocydus is similar in flavonoid chemistry to certain members of Achillea sect. Santolinoidea. This is opposed to cypsela morphology, group- ing Leucocydus with Anacydus, and requires further investi- gation. 53. MECOMISCHUS Cosson ex Benth. in Benth. & Hook, f., Gen. pi. 2: 418 (1873). Type species: M. geslini (Cosson) Cosson (M. pedunculatus (Cosson & Durieu) Maire). Annual or perennial herbs. Leaves alternate or sometimes partly opposite, few-lobed or entire. Capitula solitary, pedunculate, radiate. Receptacle paleate; paleae with a cen- tral resin canal. Ray florets neuter; limb white or yellow. Disc corolla 5-lobed, adaxially slightly saccate, with a rather long tube and a partly enervate limb. Cypselas with 1 adaxial and 2 lateral vascular strands, thin-walled, with myxogenic cells. Pappus absent. DISTRIBUTION. N. Africa in Morocco and Algeria. - 2 spp. Mecomischus with two rather different species appears to be related to Chamaemelum, Cladanthus, and Rhetinolepis. They all have thin-walled, obovoid, myxogenic cypselas. M. halimifolius (Munby) Hochr. K. BREMER AND C. J. HUMPHRIES M. pedunculatus (Cosson & Durieu) Maire 54. CHAMAEMELUM Miller in Card. Diet. abr. 4th edn. (1754). Lectotype: C. nobile (L.) All. - Ormenis (Cass.) Cass. Annual or perennial herbs or half-shrubs. Leaves alternate, pinnatifid or variously pinnatisect. Capitula solitary or laxly corymbose, pedunculate, radiate, disciform, or discoid. Receptacle conical or elongated, paleate; paleae flat or often canaliculate, sometimes enclosing florets, often with a central resin canal, glabrous or abaxially pilose. Ray florets female, fertile or sterile; limb white or yellow. Disc corolla 5-lobed, basally saccate around the cypsela especially adaxially, with a rather long tube and a more or less distinct, generally enervate limb. Cypselas obovoid, with 1 adaxial and 2 lateral very thin ribs with vascular strands, thin-walled and covered with myxogenic cells in longitudinal rows. Pappus absent. Amides and particular thiophene derivatives present. DISTRIBUTION. Mediterranean, from the Canaries in N. Africa and S. Europe to the Middle East. - 6 spp. The species of Chamaemelum were formerly placed in Orme- nis or as species oiAnthemis s. 1. The generic name Chamae- melum of Miller is prior to Cassini's Ormenis, however. Cassini distinguished Ormenis from Anthemis by the basally saccate corolla. Later, it has also been shown that Ormenis has a specialized cypsela morphology, different from that of Anthemis (Briquet, 1916). C. fuscatum, C. mixtum, and C. nobile (including Ormenis santolinoides (Munby) Harling) also differ in embryology; in contrast to Anthemis they have the normal monosporic type of embryo sac development (Harling, 1960). When Miller described the genus he included several species now in other genera. Only two of his species are presently classified in Chamaemelum, C. nobile and C. mixtum. The former is more well-known and the latter is the type of Ormenis, hence our choice of C. nobile as type species of Chamaemelum. In Flora europaea (Tutin et al., 1976) the three widespread species are recognized under Chamaemelum (C. fuscatum, C. mixtum, C. nobile). There are also a number of North African species described from Morocco. Those that seem distinct are transferred from Ormenis to Chamaemelum following recent treatments of Benedf Gonzalez (1986, 19880, b; see list of species). The North African species are still in need of revision. C. eriolepis (Cosson ex Maire) Benedf. *C. flahaulti (Emb.) Benedf. C. fuscatum (Brot.) Vase. (Anthemis fuscata Brot., Anthemis praecox Link, Ormenis praecox (Link) Briq. & Cavill.) C. mixtum (L.) All. (Anthemis mixta L., Ormenis mixta (L.) Dumort.) C. nobile (L.) All. (Anthemis nobilis L., Ormenis nobilis (L.) Gay) C. scariosum (Ball) Benedf. 55. RHETINOLEPIS Cosson in Bull. Soc. hot. Fr. 3: 707 (1856). Type species: R. lonadioides Cosson. An annual herb, branched from the base; hairs dolabriform. Leaves alternate, entire or few-lobed. Capitula solitary or few closely together, almost sessile, comparatively small, discoid. Receptacle paleate; paleae scarious with a central resin canal, abaxially pilose. Corolla 5-lobed, adaxially shal- GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 131 lowly saccate, with a narrow tube and a distinct, enervate limb. Cypselas narrowly obovoid, without ribs, with 1 adaxial and 2 lateral vascular strands, thin- walled, covered with myxogenic cells in longitudinal rows. Pappus absent. DISTRIBUTION. N. Africa in Algeria, Tunisia, and Libya. - Monotypic. Rhetinolepis is a curious small annual very different in habit from its relatives Chamaemelum, Cladanthus, and Mecomis- chus. However, they have a similar specialized cypsela mor- phology, and many floral characters in common. In North African floras Rhetinolepis lonadioides is also often called Ormenis lonadioides (Cosson) Maire. 56. CLADANTHUS Cass. in Bull. Sci. Soc. philom. Paris 1816: 199 (1816). Type species: C. arabicus (L.) Cass. An annual herb with branches in whorls below the first capitulum. Leaves alternate, below the capitula whorled, pinnatisect. Capitula solitary, sessile, radiate. Receptacle conical, paleate and pilose; paleae canaliculate and half- enclosing cypsela and basal part of corolla, with a central resin canal, pilose on both sides. Ray florets female, sterile; limb yellow, with comparatively large apical lobes. Disc corolla 5-lobed, basally saccate around the cypsela especially adaxially; lobes with acute appendices. Cypselas obovate, laterally flattened with 1 lateral and 2 marginal very thin ribs with vascular strands, thin-walled and covered with myxo- genic cells in longitudinal rows. Pappus absent. Amides and particular thiophene derivatives present. DISTRIBUTION. Mediterranean, S. Spain and N. Africa in Morocco, Algeria, Tunisia, and Libya. - Monotypic. This species is known as a relative of Chamaemelum, which has relatively plesiomorphic characters when compared to Cladanthus. The sister group, however, appears to be Rhetin- olepis, with small discoid capitula. Thus although the two genera seem rather different they both have a similar branch- ing habit and sessile capitula. 8. ANTHEMIDINAE Dumort. emend. Bremer & Humphries, emend, nov. (Dumortier, FL belg.: 69 (1827) ('Anthemideae')). Type species: Anthemis maritima L. Herbae annuae vel perennes vel suffrutices. Capitula solitaria vel laxe corymbosa. Receptaculum paleaceum vel interdum epaleaceum. Corolla flosculi disci 5- vel raro 4-lobata tubo plerumque basaliter incrassato. Cypselae plerumque turbina- tae parietibus crassis. Pappus coroniformis vel auriculiformis vel nullus. Sacculus embryonis tetrasporus. Annual or perennial herbs or suffrutices. Leaves pinnatisect to variously lobed, rarely entire. Capitula solitary or laxly corymbose, radiate or rarely disciform or discoid. Receptacle convex to narrowly conical, paleate or sometimes epaleate. Ray floret limb white or yellow. Disc corolla 5- or rarely 4-lobed; tube mostly basally swollen in fruit. Cypselas gener- ally turbinate and thick- walled, rarely with myxogenic cells. Pappus a corona, an auricle, or absent. Embryo sac tet- rasporic. DISTRIBUTION (Table 18). Eurasia, N. and E. Africa, some Table 18 General distribution of Anthemidinae, Chrysantheminae, and genera. x=indigenous, o=introduced. N. Eur- SW S. N. S. Austr. S. Am. Asia Asia Eur. Afr. Afr.N.Zeal.Am. Anthemidinae o XXX X O Anthemis o XXX X O O Nananthea X Chrysantheminae XXX X O O o Chrysanthemum XXX X 00 Heteranthemis X X Ismelia X Argyranthemum X Anthemis species widespread as weeds also in the S. hemi- sphere. - 2 genera, 213 spp. Traditionally subtribe Anthemidinae represents all Anthemideae with a paleate receptacle. In our classification it is essentially restricted to the large genus Anthemis, the immediate relatives of which are unknown. Chrysantheminae (s. s.), also with thick-walled cypselas, is a possible sister group candidate. We have also provisionally included the isolated monotypic Nananthea in Anthemidinae. The reasons are given below in the discussion of Nananthea. The cladogram also includes the Chrysantheminae. If the matrices of both subtribes are analysed together, Nananthea appears as the sister group to Chrysantheminae rather than to Anthemis. Nananthea and the Chrysantheminae share the annual habit and absence of pappus. We would consider the tetrasporic embryo sac uniting Nananthea and Anthemis a stronger character than these two characters together, how- ever. The possible relationship of Nananthea to Anthemis is further discussed under the former genus. Clades and characters - Fig. 9, Tables 2, 19. == 57 Anthemis 58 Nananthea jp 59 Chrysanthemum U=Ch2^= 60 Heteranthemis U 61 Ismelia l!= 62 Argyranthemum Fig. 9 Cladogram of the Anthemidinae and the Chrysantheminae produced by the ie option in Hennig86. Cladogram length = 23, consistency index = 86, retention index = 76. Table 19 Data matrix for the Anthemidinae and Chrysantheminae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 11 1 1 1 11 11 1 11 1 145057655 71480 774271212857 7 15763461321 532718112 51528122105610225726 30065414616 57. Anthemis alaalllal 58. Nananthea alaal?100 SV.ChrysanthemumalQalllal 60. Heteranthemis a!001?lal 6l.Ismelia Ila01?lll 62.ArgyranthemumnaO\7W\ 1 aa 1 apppOOOOOOOOOOOO l?0001alOOOOOOOOOOOO 00000101111000000000 10000101110111110000 00000101110110001100 000000011 101 lOOOOpll ?OOpppppOOO ????OOOOpOO ????OOOOOpO ????0000000 ????0000000 ????00000pp Clade Anl - subtribe Anthemidinae 175 Embryo sac tetrasporic. 132 K. BREMER AND C. J. HUMPHRIES Anthemis 11 Plants with dolabriform hairs. Many species of Anthemis have T- or Y-shaped hairs (Napp-Zinn & Eble, 1980), which occur also in Artemisiinae, Leucantheminae and some genera of Achilleinae. 45 Receptacle paleate. Some species of Anthemis, subgenus Ammanthus, have epaleate receptacles. 82 Disc corolla tube thickened in fruit. A thickened corolla tube is characteristic also of many genera in Achilleinae, Leucantheminae and Matricariinae. 108 Cypselas turbinate. Nananthea 1 Plants annual. 51 reversed. Floral parts without resin canals. 72 Disc corolla 4-lobed. This character occurs also within Thaminophyllinae and Matricariinae. 152 reversed. Cypsela wall not several cell layers thick, not partially or completely sclerified. 172 Pappus absent in ray and disc cypselas. 51. ANTHEMIS L., Sp. pi.: 893 (1753). Type species: A. maritima L. Annual or perennial herbs or half-shrubs; indumentum fre- quently of dolabriform hairs. Leaves alternate, pinnatisect to variously lobed, occasionally entire. Capitula solitary or laxly corymbose, pedunculate, radiate or discoid. Receptacle con- vex to narrowly conical, paleate, rarely basally or totally epaleate; paleae scarious and oblong or subulate, truncate to acute to acuminate, occasionally with a central resin canal. Ray florets female, fertile, or neuter; limb white or rarely yellow or reddish. Disc corolla 5-lobed, yellow or rarely reddish; tube basally much swollen in fruit, rarely pilose; lobes rarely with acute appendices. Cypselas generally turbi- nate, smooth to prismatic to c. 10-ribbed, sometimes dor- siventrally compressed, sometimes tuberculate, thick-walled, rarely with myxogenic cells. Pappus a shallow, often adaxially more developed corona, or an adaxial auricle, or absent. Embryo sac tetrasporic. DISTRIBUTION: Europe, Asia, and N. Africa, mainly in S. Europe and SW Asia, one species also in tropical E. Africa (A. tigrensis) and 2-3 species widespread as weeds also in N. America and the S. hemisphere (A. arvensis, A. cotula, A. tinctoria). -211 spp. The large genus Anthemis is, despite its size, morphologically homogeneous and the numerous species stick together in a number of groups. As a whole the genus also appears to be monophyletic based on its turbinate (obconical, in some derived species more obovoid), thick-walled fruits and the invariably basally swollen corolla tube. The south-east Euro- pean species with an epaleate receptacle, hence earlier classi- fied in a separate genus Ammanthus, have now been transferred to Anthemis (Greuter, 1968). Ammanthus is treated by Fernandes in Flora europaea (Tutin et al., 1976) as a subgenus and it represents a monophyletic group of derived species within Anthemis. The infrageneric classification of Anthemis follows the common pattern with recognition of a number of apomorphic sections and subgenera in addition to a plesiomorphic and probably paraphyletic subgenus Anthe- mis. Yavin (1970, 1972) has proposed an elaborate infrage- neric classification of Anthemis. She also revised section Maruta. Anthemis was typified with A. maritima by Britton & Brown (1913). Later Green (in Hitchcock & Green, 1929) proposed A. arvensis as type species, since it is a well-known species also occurring in Sweden, the home-country of Lin- naeus. Both species fit Linnaeus' generic description in Genera plantarum (Linnaeus, 1757). Linnaeus did not coin the generic name; it was adopted from Micheli (1729) who included A. maritima but not A. arvensis. Hence, we consider the original typification with A. maritima by Britton & Brown to be the correct choice. A. aaronsohnii Eig. Middle East. *A. abagensis Fed. Caucasus. A. abrotanifolia (Willd.) Guss. Greece. A. aciphylla Boiss. Turkey. *A. adonidifolia Boiss. Turkey. *A. aeolica Lojac. Italy. A. aetnensis Schouw in Sprengel. Italy. A. alpestris (Hoffsgg & Link) R. Fernandes. SW Europe. A. altissima L. S. Europe, Krym, Turkey, Caucasus, Iraq, Iran, Afghanistan, C. Asia. A. amblyolepis Eig. Turkey, Cyprus, Middle East. A. ammanthus Greuter. Greece. A. ammophila Boiss. & Heldr. in Boiss. Turkey. A. anatolica Boiss. Greece, Turkey. A. anthemiformis (Freyn & Sint.) Grierson. Turkey. *A. antilibanotica Eig. Middle East. A. antitaurica Grierson. Turkey. A. arenicola Boiss. Turkey. *A. argyrophylla (Hal. & Georgiev) Velen. Bulgaria. A. armeniaca Freyn & Sint. Turkey. A. arvensis L. Europe, N. Africa, W. Asia, also widespread as a weed in N. America, S. Africa, Australia and New Zealand. *A. atropatana Iranshar. Iran. A. auriculata Boiss. SE Europe, Turkey. A. austriaca Jacq. C. and E. Europe, Krym, Turkey, Cauca- sus, Iran. A. austro-iranica Rech. f., Aellen & Esfand. Iran. A. bornmuelleri Stoy. & Acht. (A. galilaea Eig). Egypt, Middle East. *A. bourgaei Boiss. & Reuter. Spain, Morocco. A. boveana Gay. N. Africa in Morocco, Algeria and Libya. M. brachmannii Boiss. & Heldr. in Boiss. Greece. A. brachycarpa Eig. Middle East. A. br achy Stephana Bornm. & Gauba. Iran. A. brevicuspis Bornm. (A. feinbruniae Eig., A. rayatensis Eig). Middle East, Iraq, Iran. *A. breviradiata Eig. Middle East. *A. bulgarica N. N. Thin. Bulgaria. *A. bushehrica Iranshahr. Iran. A. calcarea Sosn. Turkey, Caucasus. A. candidissima Willd. ex Sprengel. Caucasus, Iran, C. Asia. A. carpatica Waldst. & Kit. ex Willd. C. and S. Europe. A. chia L. SE Europe, Middle East, Egypt. A. chrysantha Gay. Spain, Algeria. A. coelopoda Boiss. SE Europe, Turkey, Middle East, Iran, Afghanistan. A. cornucopiae Boiss. Middle East. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 133 A. corymbulosa Boiss. & Hausskn. in Boiss. Middle East. A. cotula L. Europe, N. Africa, W. Asia, also widespread as a weed in America, S. Africa, Australia and New Zealand. *A. cretacea Zefirov. Krym, Caucasus. A. cretica L. (A. anahytae Woronow ex Sosn., A. iberica M. Bieb., A. montana L., A. panachaica Hal., A. pindicola Heldr. ex Hal., A. ptarmiciformis K. Koch, A. tempskyana Freyn & Sint.). S. Europe, N. Africa, Turkey, Middle East, Iran. A. cuneata Huber-Mor. & Reese. Turkey. *A. cypria Boiss. Cyprus. A. cyrenaica Cosson. Libya. A. damascena Boiss. & Gaill. Middle East. A. davisii Yavin. Turkey. A. debilifolia Eig. Middle East. A. deserticola H. Kraschen. & Popov. C. Asia. A. deserti-syriaci Eig. Middle East. *A. didymaea Mout. Middle East. A. dipsacea Bornm. Turkey. *A. dubia Steven. Krym. A. edumea Eig. Middle East. A. ellezrae Eig. Middle East, Egypt. A. emasensis Eig. Middle East. A. emiliae Sosn. Caucasus. A. filicaulis (Boiss. & Heldr.) Greuter. Greece. A. fimbriata Boiss. Turkey. A. flexicaulis Rech. f. Greece. A. freitagii Iranshahr. Afghanistan. A. fruticulosa M. Bieb. Caucasus. A. fulvida Grierson. Turkey. A. fumariifolia Boiss. Turkey. M. fumarioides Hochst. Middle East. *A. fungosa Boiss. & Hausskn. Iran. A. gaudium-solis Velen. Bulgaria. A. gay ana Boiss. Iran. A. gerardiana Jordan. France. A. gilanica Boiss. Iran. *A. gilletti Iranshahr. Iraq, Iran. A. glaberrima (Rech. f.). Greuter. Greece. A. glareosa Durieu & Barratte. Libya. *A. gracilis Iranshahr. Iran. A. gross heimii Sosn. Caucasus. A. halophila Boiss. & Bal. in Boiss. Turkey. *A. hamrinensis Iranshahr. Iraq. A. handel-mazzettii Eig. Middle East. A. haussknechtii Boiss. & Reuter in Boiss. Syria, Iraq, Iran. A. hebronica Boiss. & Kotschy. Middle East, Egypt. A. hemistephana Boiss. Middle East, Iran. *A. hermonis Eig. Middle East. *A. hinkovae N. N. Thin. Bulgaria. A. hirtella Winkler. C. Asia. A. homalolepis Eig. Middle East. A. hyalina DC. Turkey, Middle East, Iraq, Iran. A. hydruntina Groves. Italy. A. indurata Del. Middle East, N. Africa in Egypt and Libya. A. ismelia Lojac. Italy. *A. jailensis Zefirov. Krym. *A. jordanovii Stoy. & Acht. Bulgaria. A. kandaharica Iranshahr. Afghanistan, Pakistan. *A. karabaghensis Mikheev. Caucasus. A. kitanovii N. N. Thin. Bulgaria. A. kitenensis N. N. Thin. Bulgaria. A. kotschyana Boiss. Turkey, Middle East, Iraq, Iran. *A. krugeriana Pampan. Libya. *A. kurdica Iranshahr. Iraq. *A. kuzmanovii N. N. Thin. Bulgaria. A. laconica R. Franzen. Greece. *A. leptophylla Eig. Iraq, Iran. A. leucanthemifolia Boiss. & Blanchet. Middle East, Egypt. A. leucolepis Eig. Middle East. *A. linczevskyi Fed. C. Asia. *A. lithuanica (DC.) Besser ex Trautv. Europe in Lithuania and Russia. A. lorestanica Iranshahr. Iran. A. lyonnetioides (Boiss. & Kotschy) Boiss. Middle East. *A. macedonica Boiss. & Orph. in Boiss. SE Europe. A. macrantha Heuffel. SE Europe. *A. macroglossa Sommier & Levier. Caucasus. A. maris-mortui Eig. Middle East. *A. maris-nigri Fed. Caucasus. A. maritima L. SW Europe, N. Africa. *A. markhotensis Fed. Caucasus. A. marschalliana Willd. Caucasus. A. mauritiana Maire & Sennen. Morocco. A. mazandaranica Iranshahr. Iran. A. melampodina Del. (A. deserti Boiss.) Middle East, Egypt. A. melanacme Boiss. & Hausskn. in Boiss. Turkey, Middle East. A. melanoloma Trautv. Turkey. *A. meteorica Hausskn. Greece. A. micrantha Boiss. & Hausskn. Iraq. A. microcephala (Schrenk) B. Fedtsch. (A. straussii Bornm., A. tenuiflora Gilli). C. Asia, Iraq, Iran, Afghanistan. A. microlepis Eig. Middle East. A. microsperma Boiss. & Kotschy. Middle East, Egypt. *A. mirheydari Iranshahr. Iran. A. moghanica Iranshahr. Iran. *A. monantha Willd. Krym. M. monilicosta Pomel. N. Africa in Morocco, Algeria and Libya. A. muricata (DC.) Guss. Italy. *A. nabataea Eig. Middle East. A. odontostephana Boiss. (A tubicina Boiss. & Hausskn. in Boiss.). Iran, C. Asia. *A. orbelica Pancic. Bulgaria. A. orientalis (L.) Degen (A. pectinata (Bory & Chaub.) Boiss. & Reuter). Greece, Turkey. A. oxylepis (Boiss.) Boiss. Turkey. A. palestina Reuter in Boiss. (A. melanolepis Boiss., A. syriaca Bornm.). Turkey, Cyprus, Middle East, SE Europe. A. parnassica (Boiss. & Heldr.) R. Fernandes. SE Europe. A. parnesia Boiss. & Heldr. in Boiss. Greece. M. parviceps Dobrocz. & Fed. Krym. A. parvifolia Eig. Middle East. *A. patentissima Eig. Middle East. A. pauciloba Boiss. Turkey, Iraq. A. pedunculata Desf. N. Africa in Morocco, Algeria and Libya. A. persepolitana Boiss. Iraq, Lebanon, Syria. A. persica Boiss. Iran. A. pestalozzae Boiss. Turkey. A. plebeia Boiss. & Noe. Iraq. *A. plutonia Meikle. Cyprus. A. pseudocotula Boiss. (A. behboudiana Rech. f. & Esfand.). Turkey, Cyprus, Middle East, Iraq, Iran, N. Africa in Egypt and Libya. 134 K. BREMER AND C. J. HUMPHRIES A. punctata Vahl. Italy, N. Africa in Morocco, Algeria and Tunisia. *A. pungens Yavin. Turkey. A. rascheyana Boiss. Middle East. *A. regis-borisii Stoy. & Acht. Bulgaria. A. retusa Del. Egypt. *A. rhodensis Boiss. Turkey. *A. rhodocentra Iranshahr. Iran, Afghanistan, Pakistan. A. rigida (Sibth. & Smith) Boiss. & Heldr. SE Europe, Turkey, Cyprus. A. rosea Smith in Sibth. & Smith. Turkey. A. rumelica (Velen.) Stoy. & Acht. Bulgaria. A. ruthenica M. Bieb. C. and SE Europe, Caucasus. A. sabullfolia Pomel. N. Africa. A. saguramica Sosn. Caucasus. A. samuelssonii Rech. f. Middle East. A. sancti-johannis Turrill. Bulgaria. *A. saportana Albov. Caucasus. A. scaettae Pampan. Libya. A. scariosa Banks & Sol. in Russell. Turkey, Middle East, Iraq, Iran. *A. schischkiniana Fed. Caucasus. A. schizostephana Boiss. & Hausskn. Iraq, Iran. A. scopulorum Rech. f. Greece. A. scrobicularis Yavin. Middle East. A. secundiramea Biv. SW Europe, N. Africa in Algeria and Tunisia. A. segetalis Ten. (A. brachycentros Gay ex W. Koch). S. Europe. A. semiensls Pichi-Serm. Ethiopia. A. sibthorpii Griseb. Greece. A. sintenisii Freyn. Turkey. A. sosnovskyana Fed. Caucasus. A. spruneri Boiss. & Heldr. in Boiss. Greece. A. sterilis Steven. Krym. A. stiparum Pomel. N. Africa. *A. stribrnyi Velen. Bulgaria. *A. susiana Nab. Iraq, Iran. A. talyschensis Fed. Caucasus, Iran. M. taubertii Durieu & Barratte. Libya. A. tenuicarpa Eig. Middle East. A. tenuiloba (DC.) R. Fernandes. SE Europe. A. tigrensis Gay ex A. Richards. E. Africa. A. tinctoria L. (A. debilis Fed., A. euxina Boiss., A. subtinc- toria Dobrocz.). Europe, W. and C. Asia, also naturalized in N. America. A. tomentella Greuter. Greece. A. tomentosa L. (A. peregrina L.). SE Europe, Turkey. *A. tranzcheliana Fed. Krym. A. tricolor Boiss. Cyprus. A. tricornis Eig. Turkey. A. tripolitana Boiss. & Blanchet in Boiss. Middle East. A. triumfettii (L.) DC. in Lam. & DC. (A. dumetorum Sosn., A. khorassanica Rech. f., A. rigescens Willd.). S. Europe, Turkey, Caucasus, Iran. A. trotzkiana Claus ex Bunge. S. Russia, C. Asia. A. tuberculata Boiss. Spain. *A. virescens Velen. Bulgaria. A. wallii Huber-Mor. & Reese. Turkey. *A. werner i Stoy. & Acht. Greece. A. wettsteiniana Hand.-Mazz. Iraq, Iran. A. wiedemanniana Fischer & C. Meyer. Caucasus, Turkey. A. woronowii Sosn. Caucasus. A. xylopoda O. Schwarz. Turkey. A. yemenensis Podl. Yemen. *A. zephyrovii Dobrocz. Krym. A. zoharyana Eig. Middle East, Egypt. A. zyghia Woronow. Caucasus. 58. NANANTHEA DC., Prodr. 6: 45 (1838). Type species: N. perpusilla (Lois.) DC. A delicate, somewhat succulent, annual herb. Leaves alter- nate, pinnatifid, with obovate lobes. Capitula solitary, long- pedunculate, very small (2-5 mm diam.), radiate or disciform. Involucre of 5-10 wide bracts in 1 to 2 rows. Receptacle conical, epaleate. Ray florets female, fertile, with or without a white lamina. Disc corolla 4-lobed, with a short tube and large lobes. Cypselas obovoid, with myxogenic cells; ray cypselas with 2 lateral vascular strands; disc cypselas with 2 lateral, 1 adaxial and 1 pseudolateral vascular strand. Pappus absent. Embryo sac tetrasporic. DISTRIBUTION. S. Europe, Corsica and Sardinia. - Mono- typic. Nananthea is a genus of uncertain position. Traditionally it has been included in a widely circumscribed and heteroge- neous Cotula group but it shares no synapomorphies with the Cotula group, which is part of subtribe Matricariinae, as understood here. Nananthea is here provisionally placed together with Anthemis, following a suggestion by Reitbrecht (1974). It could be a highly specialized Anthemis derivative, related to part of Anthemis, possibly subgenus Ammanthus. Both genera have tetrasporic embryo sacs. This also occurs in Tanacetum, Tripleurospermum, and Heteranthemis, but Nan- anthea is hardly reminiscent of these genera. 9. CHRYSANTHEMINAE Less, emend. Bremer & Humphries, emend, nov. (Lessing in Linnaea 6: 167 (1831) ('Chrysanthemeae')). Type species: Chrysanthemum coronarium L. Herbae annuae vel perennes vel suffrutices vel frutices. Capitula solitaria vel laxe corymbosa. Bracteae involucri latae, plurinerves. Receptaculum epaleaceum. Cypselae pari- etibus crassis, heteromorphae; cypselae flosculorum radii triquetrae, alatae; eae flosculorum disci plerumque lateraliter compressae et abaxialiter adaxialiterque alatae vel raro ter- etes ad prismaticae. Pappus nullus. Annual or perennial herbs, half-shrubs or shrubs. Leaves serrate-dentate-pinnatifid to variously dissected. Capitula solitary or laxly corymbose, pedunculate, radiate. Involucral bracts wide, many- veined. Receptacle convex to conical, epaleate. Ray floret limb white and/or yellow. Disc corolla 5-lobed, yellow or rarely red. Cypselas thick-walled, without myxogenic cells, heteromorphic; ray cypselas triquetrous, winged; disc cypselas generally laterally flattened and abaxi- ally and adaxially winged, or rarely terete to prismatic. Pappus absent. DISTRIBUTION (Table 18). Eurasia, N. Africa and Macaron- esia, Chrysanthemum coronarium widespread as a weed also in the S. hemisphere. - 4 genera, 28 spp. Traditionally subtribe Chrysantheminae comprises all of the Anthemideae species with an epaleate receptacle. As circum- scribed here it is a small and homogeneous taxon. The close relationship between Chrysanthemum s. s. and the three GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE other genera of this subtribe has also been recognized by earlier authors and is discussed by Humphries (1976). Clades and characters - Fig. 9, Tables 2 and 19. Clade Chi - subtribe Chrysantheminae 1 Plants annual. Argyranthemum consists of shrublets or half-shrubs, here considered a secondary development within the subtribe. A shrubby habit is commonly evolved in island groups. 41 Involucral bracts wide, flabelliform. This character also occurs in several genera of other subtribes. 120 Cypselas heteromorphic; ray cypselas triquetrous, winged; disc cypselas terete to prismatic to laterally flattened. 172 Pappus absent in ray and disc cypselas. Chrysanthemum 52 reversed. Ray floret limb not white, but yellow. 75 Disc corolla lobes with central resin sacs. Clade Ch2 107 reversed. Cypselas not terete to weakly angled, but acutely angled. 116 Disc cypselas laterally flattened. 121 Disc cypselas abaxially and adaxially winged. Heteranthemis 10 Plants covered with viscid hairs. 52 reversed. Ray floret limb not white, but yellow. 122 Cypsela wings as apical spines. 175 Embryo sac tetrasporic. The embryo sac of Ismelia is monosporic (Hading, 1951). 182 Flavonol 5-glycosides present. Most of the related genera have not been investigated chemically. Ismelia 55 Ray floret limb deeply emarginate. 77 Disc corolla red. Argyranthemum 1 reversed. See clade Chi. 2 Plants shrubby. See clade Chi under character 1. 51 reversed. Floral parts without resin canals. 176 Embryo sac disporic. 59. CHRYSANTHEMUM L., Sp. pi.: 887 (1753). Type species: C. coronarium L. Annual herbs. Leaves alternate, deeply serrate-dentate and pinnatifid to pectinate, somewhat amplexicaul. Capitula soli- tary or laxly corymbose, pedunculate, radiate. Involucral bracts wide, many- veined, with resin canals. Receptacle convex, epaleate. Ray florets female, fertile; limb yellow or white distally, many-veined. Ray cypselas triquetrous, later- ally winged, adaxially with a narrow wing or ribbed; pappus absent. Disc corolla 5-lobed; lobes with central resin sacs. Disc cypselas prismatic with a narrow adaxial wing or terete, 135 with a thick undulating wall, thus apparently ribbed; pappus absent. DISTRIBUTION. Europe, Asia and N. Africa, C. coronarium widespread as a weed. - 2 spp. The adaxial cypsela wings of C. coronarium are similar to those in Heteranthemis, Ismelia, and Argyranthemum, whereas C. segetum has terete disc cypselas and only laterally winged ray cypselas. The adaxial cypsela wing is conceived as a parallelism, since the two species of Chrysanthemum are united for example by their corolla lobe resin sacs. C. carinatum Schousboe is more closely related to Heteran- themis and Argyranthemum than to the two Chrysanthemum species adopted here, and thus it is transferred to Ismelia. C. coronarium L. C. segetum L. 60. HETERANTHEMIS Schott in Isis, Oken 1818 (5): 822 (1816). Type species: H. viscidehirta Schott. An annual herb covered with viscid glandular hairs. Leaves alternate, serrate-dentate to pinnatifid. Capitula solitary or laxly corymbose, pedunculate, radiate. Involucral bracts wide, many-veined, with resin canals. Receptacle convex, epaleate. Ray florets female, fertile; limb yellow, many- veined. Ray cypselas triquetrous, laterally and adaxially winged; wings projected to apical spines; pappus absent. Disc corolla 5-lobed. Disc cypselas laterally flattened, winged; adaxial wing projected to an apical spine; pappus absent. Embryo sac tetrasporic. DISTRIBUTION. SW Europe in Spain and Portugal, N. Africa in Morocco and Algeria. - Monotypic. This species, also known as Chrysanthemum viscidehirtum (Schott) Thell., is distinguished from Ismelia and Argyranthe- mum by its pubescence of viscid glandular hairs and the apical spines on the cypsela wings. 61. ISMELIA Cass. in Diet. Sci. Nat. 41: 40 (1826). Type species: /. versicolor Cass. (/. carinata (Schousboe) Schultz-Bip.). An annual herb. Leaves alternate, pinnatisect. Capitula solitary or laxly corymbose, pedunculate, radiate. Involucral bracts wide, many-veined, with resin canals. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb yellow, basally reddish or white, many-veined, deeply emar- ginate. Ray cypselas triquetrous, laterally and adaxially winged; pappus absent. Disc corolla 5-lobed, red to purple. Disc cypselas laterally flattened, winged; pappus absent. DISTRIBUTION. N. Africa in Morocco but frequently escaped from cultivation. - Monotypic. This handsome species, frequently cultivated as an ornamen- tal, is commonly known as Chrysanthemum carinatum Schousboe. It is, however, more closely related to Heteran- themis and Argyranthemum than to Chrysanthemum coro- narium and Chrysanthemum segetum, the two species here retained in that genus. Ismelia, Heteranthemis, and Argyran- themum form a monophyletic group based on their laterally compressed disc cypselas and their especially strongly devel- oped cypsela wings on the adaxial side. Interestingly, Schultz- Bipontinus (1844ft) expanded the concept of Ismelia to 136 K. BREMER AND C. J. HUMPHRIES include various Canary Island endemics of Argyranthemum (see Humphries, 1976, for details). Argyranthemum, how- ever, forms a monophyletic group, so Ismelia is related to Argyranthemum as a whole rather than to part of it. Uniting Ismelia and then by consequence also Heteranthemis with Argyranthemum because of their similar fruits is hardly desirable, since it necessitates recombination of all Argyran- themum specific names, Argyranthemum being the youngest name. 62. ARGYRANTHEMUM Webb ex Schultz-Bip. in Webb & Berthelot, Hist. nut. lies Canaries 3 (2,2): 245, 258 (18446). Type species: A. frutescens (L.) Schultz-Bip. Shrublets or half-shrubs. Leaves alternate, variously dis- sected. Capitula solitary or laxly corymbose, pedunculate, radiate. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb white, rarely yellow or pink, many- veined. Ray cypselas triquetrous, generally laterally and adaxially strongly winged, sometimes coalesced into groups; wings sometimes reduced, often apically projected to a pappus-like corona; true pappus absent. Disc corolla 5-lobed, yellow; lobes rarely reddish purple. Disc cypselas generally laterally flattened and adaxially and abaxially winged, some- times prismatic to terete and wingless, sometimes coalesced with ray cypselas, apically often coroniform; true pappus absent. Embryo sac bisporic. DISTRIBUTION. Macaronesia in the Canary Islands, Madeira (A. dissectum, A. haematomma, A. pinnatifidurri), and the Salvage Islands (A. thalassophiluni) . - 24 spp. Argyranthemum was revised by Humphries (1976). As already pointed out by him, it is related to Ismelia and Heteranthemis. Argyranthemum has, unlike these genera, evolved into many species with even more specialized some- times coalesced fruits often with folded wings or sometimes secondarily wingless (Borgen, 1972). A new species, A. sundingii, was described by Borgen (1980) and a study of variation within A. pinnatifidum was undertaken by Rustan (1981). A. adauctum (Link) Humphries A. broussonetii (Pers.) Humphries A. callichrysum (Svent.) Humphries A. coronopifolium (Willd.) Humphries A. dissectum (Lowe) Lowe A. x escarrei (Svent.) Humphries A. filifolium (Schultz-Bip.) Humphries A. foeniculaceum (Willd.) Webb ex Schultz-Bip. A. frutescens (L.) Schultz-Bip. A. gracile Schultz-Bip. A. haematomma (Lowe) Lowe A. haouarytheum Humphries & Bramwell A. hierrense Humphries A. jacobiifolium Kunkel A. lemsii Humphries A. lidii Humphries A. maderense (D. Don) Humphries A. pinnatifidum (L. f.) Lowe A. sundingii Borgen A. sventenii Humphries & Aldridge A. tenerifae Humphries A. thalassophilum (Svent.) Humphries A. webbii Schultz-Bip. A. winteri (Svent.) Humphries 10. LEUCANTHEMINAE Bremer & Humphries, subtrib. nov. Type species: Leucanthemum vulgare Lam. Herbae annuae vel perennes vel interdum suffruticosae. Folia saepe serrata-dentata vel interdum pectinata-lobata, trifur- cata vel integra. Capitula solitaria, pedunculata. Flosculi disci tubo plerumque incrassato. Cypselae plerumque 10(8-12) vel interdum pauciores costas praebentes, inter costas saepe lacunis vallecularibus et canalibus secretoriis et fasciculis vascularibus (Leucanthemum et genera sequentia), in costis plerumque cellulis mucilaginis, instructae. Pappus coronifor- mis vel adaxialiter auriculiformis vel nullus. Annual or perennial herbs, sometimes suffruticose; indumen- tum frequently of dolabriform hairs. Leaves often serrate- dentate, sometimes pectinate-lobed, trifurcate or entire. Capitula solitary, pedunculate, generally radiate or some- times discoid. Involucral bracts sometimes with dark brown margins, sometimes wide and flabelliform. Receptacle flat to conical, epaleate. Ray floret limb white, yellow or rarely reddish. Disc corolla 5-lobed or rarely 4-lobed; tube generally swollen in fruit. Cypselas mostly 10(8-12)-ribbed, sometimes with fewer ribs, often with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs (Leucanthemum group of genera), generally with myxogenic cells along the ribs. Pappus a corona, an adaxial auricle, or absent. DISTRIBUTION (Table 20): Eurasia and N. Africa, mainly in the Mediterranean region, some Leucanthemum species widely introduced, also in N. America and the S. hemisphere. - 16 genera, 75spp. Leucantheminae consists of one well defined monophyletic subclade, the Leucanthemum group of genera, a smaller subclade comprised of Leucanthemopsis , Hymenostemma and Prolongoa, as well as a number of isolated genera of uncertain position, here provisionally included in this sub- tribe. The Leucanthemum group of genera (71-78; see Fig. 10), are all characterized by their specialized cypsela wall with vallecular lacunae, secretory canals, and vascular strands. Their status as a group has been recognized by several earlier authors (for example, see Briquet, 1916). Leucanthemopsis is apparently allied to the two Iberian monotypic genera Hymenostemma and Prolongoa, a relation- ship that hitherto has not been clearly recognized. The subtribal description above does not cover all diver- gent characters of the genera, but applies mainly to the Leucanthemum and Leucanthemopsis groups. A relationship between Leucanthemopsis and Leucanthe- mum has been suggested several times, e. g. by Heywood (1954, 1976). It is supported by presence of flavonol 5-glycosides in both genera. They are also found in Coleoste- phus and Plagius and constitute a possible synapomorphy for the Leucanthemum and Leucanthemopsis groups. Other gen- era of these groups have not been investigated chemically, as far as we know. Included in Leucantheminae are also a number of odd, mainly monotypic genera of uncertain affinity. Lepidopho- GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE Table 20 General distribution of Leucantheminae and genera. x=indigenous, o=introduced. 137 N.Am. Eur- Asia C.& Asia E.SW. Asia S.Eur. N.Afr. S.Afr. Austr. S.Am. N.Zeal. Leucantheminae o x X X X O Lepidophorum x Nipponanthemum X Leucanthemella x X Nivellea X Phalacrocarpum X Leucanthemopsis x X X Hymenostemma X X Prolongoa X Leucanthemum o x X X O O Rhodanthemum X X Leucoglossum X X Chlamydophora X X Chrysanthoglossum X Glossopappus X X Coleostephus X X Plagius X X rum, Nipponanthemum, Leucanthemella, Nivellea and Phala- crocarpum are classified in this subtribe mainly because of their serrate-dentate leaves (not in Nivellea) but also on account of some floral similarities. These genera have 10(8-12)-ribbed cypselas as in the Leucanthemum group. An exception is Lepidophorum with 5-ribbed cypselas. The posi- tions of these genera are further discussed under each genus. There is one most parsimonious cladogram produced from the data matrix. It differs from the cladogram pre- sented by having Hymenostemma and Prolongoa as the sister group to Lepidophorum rather than to Leucanthe- mopsis. However, we have chosen the latter arrangement, which is one step longer. We consider the peculiar pappus (character 165) shared by Hymenostemma, Prolongoa and Leucanthemopsis a stronger character than the annual habit and the loss of brown involucral bracts together (characters 1 and 44), which the former two genera share with Lepidophorum. Clades and characters - Fig. 10, Tables 2, 21. Clade Lei - subtribe Leucantheminae theminae, notably those provisionally included in the early part of the account, have dolabriform hairs. The character occurs also in Artemisiinae, Anthemis (Anthemidinae), and some genera of Achilleinae. Possibly it is a synapomorphy at a lower level within the tribe. Lepidophorum 1 Plants annual. 45 Receptacle paleate. 52 reversed. Ray floret limb not white, but yellow. 53 Ray floret limb golden yellow. Similar rays occur in subclade LelO of the Leucanthemum group of genera. 133 Cypselas with costal resin canals or sacs. Clade Le2 124 Cypselas with 10 (8-12) multicellular epicarpic ribs. Fewer ribs occur in most species of Leucanthemopsis and in Hymenostemma and Prolongoa. 11 Plants with dolabriform hairs. Most genera of Leucan- Nipponanthemum k=Lel=n= 63 Lepidophorum lL =Le 2== 64 Nipponanthemum f = L= 65 Leucathemella lL Le3 _ 66 Nivellea LzLe4=n= 67 Phalacrocarpum [L Le 5_ L e6:jr= 68 Leucanthemopsis lL= Le7 _ 69 Hymenostemma IL 70 Prolongoa r= 71 Leucanthemum 72 Rhodanthemum E73 Leucoglossum :1JCJLU 74 Chlamydophora it=Le8: LL e 9 IT* " 17= 75 Chrysanthoglossum L=LeiaJ=Lel2Tf= 76 Glossopappus L= 77 Coleostephus IL= 78 Plagius Fig. 10 Cladogram of the Leucantheminae produced by the bb option in Hennig86. Cladogram length = 29, consistency index = 58, retention index = 79. 138 K. BREMER AND C. J. HUMPHRIES Table 21 Data matrix for the Leucantheminae. 1 = presence, = absence, 7 = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 111 1111111111111 1 1111 1450576521 4532 784107860573368221236474026 532718111 1153342224444253714564060258131959 111 1 1577755556666617 3056056790567857 63. 64. 65. Lepidophorum Nipponanthemum Leucanthemella a a a 1 1 1 1 1 1 1 1 1 7 1 1 7 1 1 7 1 1 1 1 1 1 ? 1 1 1 1 1 1000000000000000000000 00001 10000000000000000000 0000101000000000000700000 00000000 00000000 00000000 7000000000000000 ????000000000000 ?????00000000000 66. Nivellea 1 1 1 1 1 7 1 1 9 1000101 1 00000000000700000 00000000 7700700000000000 67. Phalacrocarpum a 1 1 1 1 7 1 1 1 1 0000101 1 1 1 1 10000000700000 00000000 7777700000000000 68. Leucanthemopsis 1 1 a 1 1 7 1 1 1 00000001 10001 100000000000 00000000 7777000000000000 69. Hymenostemma 1 1 1 1 1 ? 1 1 1 10000001 0000 ? 100000000000 00000000 7777000000000000 70. Prolongoa 1 1 1 1 ? 1 1 1 10000000 0000 7 111100000000 00000000 7777000000000000 71. Leucanthemum a 1 a 1 1 7 1 1 1 00000 1 Op 1 10001 OOOOllalOOOO pOOOOOOO 7700000000000000 72. Rhodanthemum 1 1 1 1 1 7 1 1 9 00001001 1000? 000011101100 00000000 ????00000000000p 73. Leucoglossum a 1 1 1 1 7 1 1 1 9 10001001 0000 7 000111100010 00000000 7777000000000000 74. Chlamydophora a 1 7 1 1 7 1 1 9 10701001 0000? 000711100011 1 1000000 9?990 999 '>'> 9 '> 99 ' ? 75. Chrysanthoglossum 1 1 1 1 7 1 1 9 aO 1 1 00 1 0000 7 00001 1000000 00111000 7777000000000000 76. Glossopappus a 1 1 1 7 1 1 1 1 10101001 0000 ? 00001110000000110111 7700000000000000 77. Coleostephus a 1 1 1 7 1 1 1 1 10101001 00001 000011100000 1 1 1 aO 7700000000000000 78. Plagius a 1 7 1 1 7 1 1 1 9 00701001 00001 000711100000 10010110 9999099999999990 2 Plants shrubby. 51 reversed. Floral parts without resin canals. Leucanthemella 172 Pappus absent in ray and disc cypselas. Clade Le3 82 Disc corolla tube thickened in fruit. This character is sometimes vaguely expressed and absent at least in Prolon- goa. Nivellea 1 Plants annual. 21 reversed. Leaves not serrate-dentate. 172 Pappus absent in ray and disc cypselas. Clade Le4 44 Involucral bracts with dark brown margins. This character is absent in several genera (see cladogram) and present in many genera of other subtribes (Cancriniinae, Tanacetinae, Artemisiinae). It may be a synapomorphy at a lower level with reversals in several instances. Phalacrocarpum 14 Leaves opposite. 104 Disc floret style-branches fused. 172 Pappus absent in ray and disc cypselas. 174 Testa epidermis cells thick-walled and dark reddish. Clade Le5 182 Flavonol 5-glycosides present. In this subtribe only Leu- canthemopsis, Leucanthemum, Coleostephus and Plagius have been investigated for these compounds. Clade Le6 21 reversed. Leaves not serrate-dentate. 124 reversed. See clade Le2. 165 Pappus a scarious, flimsy corona. Leucanthemopsis There is no autapomorphy for this genus. Clade Le7 I Plants annual. 44 reversed. See clade Le4. Hymenostemma There is no autapomorphy for this genus. Prolongoa 52 reversed. Ray floret limb not white, but yellow. 82 reversed See clade Le3. 103 Disc floret style-branches long-penicillate. 157 Cypsela wall with rod-shaped crystals in small packets. Ill Pappus absent in disc cypselas, but present in ray cypselas. Clade Le8 - The Leucanthemum group of genera 134 Cypselas with vallecular secretory canals. 135 Cypselas with vallecular vascular strands. 166 Pappus adaxially long. In Chrysanthoglossum the pappus is coroniform and hardly longer adaxially. Leucanthemum II reversed. Plants without dolabriform hairs. 184 Anthocyanin present in root tips. Rhodanthemum 20 Leaves spathulate in outline, ternate to ternately pinnate. 21 reversed. Leaves not serrate-dentate. 126 Cypsela ribs protruding, narrow and somewhat wing-like. Clade Le9 1 Plants annual. The character reverses in Plagius, which is perennial. 44 reversed. See clade Le4. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE Clade LelO 110 Cypselas ellipsoid, small, c.l mm long. Leucoglossum 111 Pappus absent in disc cypselas, but present in ray cypselas. Chlamydophora 21 reversed. Leaves not serrate-dentate. 22 Leaves entire or apically tridentate. 35 Capitula discoid. 168 Pappus a large, scarious, adaxial but basally coroniform auricle, as long as the corolla or longer. Clade Lell 41 Involucral bracts wide, flabelliform. The character reverses in Plagius, where the involucral bracts are not flabelliform. 52 reversed. Ray floret limb not white, but yellow. 53 Ray floret limb golden yellow. 73 Disc corolla lobes with dorsal appendages. Chrysanthoglossum 21 reversed. Leaves not serrate-dentate. 141 Cypselas with dense rows of myxogenic cells also on the corona. 166 reversed. See clade Le8. Clade Lel2 109 Cypselas arcuate. 125 Cypsela ribs basally fused into a more or less well developed foot callus. This character is variously strongly expressed in the different species of the three genera involved. Glossopappus 169 Pappus a large, scarious, adaxial, flabelliform auricle, as long as the corolla or longer. Coleostephus There is no autapomorphy for this genus. Plagius 1 reversed. See clade Le9. 35 Capitula discoid. 41 reversed. See clade Lell. 63. LEPIDOPHORUM Necker ex Cass. in Diet. Sc. Nat. 26: 36 (1823). Type species: L. repandum (L.) DC. An annual or biennial herb. Leaves alternate, oblong to obovate-spathulate, serrate. Capitula solitary, pedunculate, radiate. Involucral bracts wide, many-veined. Receptacle paleate; paleae scarious with a conspicuous resin canal. Ray florets neuter or female, sterile; limb yellow. Disc corolla 5-lobed. Ray cypselas (sterile) flat, with 2 lateral and 1 adaxial resin canal; pappus of c. 4 free or basally connate 139 scales. Disc cypselas 5-angled with 5 thin ribs covered with myxogenic cells; pappus absent. DISTRIBUTION. SW Europe in Spain and Portugal. - Mono- typic. Lepidophorum, formerly a member of Anthemis because of its paleate receptacle, is difficult to place. Several authors have suggested a relationship to Leucanthemum mainly because of the habitual (foliage) similarities, though Lepi- dophorum does not have the specialized fruits of the Leucan- themum group of genera. Harling (1960) investigated the embryology of many Anthemis species, including A. repanda (= Lepidophorum repandum). Lepidophorum has the common (Polygonum) type of monosporic embryo sacs, while Anthemis proper has tetrasporic embryo sacs. Harling also described the cypselas of Lepidophorum in detail and discussed the relationships of the genus. He concluded that Lepidophorum should be recognized as a separate genus, not related to Anthemis but possibly distantly so to Coleostephus, i.e. the Leucanthemum group of genera. Lepidophorum is here provisionally placed as a basal member of subtribe Leucantheminae, where the Leucanthemum group is a subclade (Fig. 10, Le8). 64. NIPPONANTHEMUM Kitam. in Acta phytotax. geobot. Kyoto 29: 169 (1978). Type species: N. nipponicum (Franchet ex Maxim.) Kitam. A shrub. Leaves alternate, obovate, apically serrate leaves rather densely set at the ends of the branches. Capitula rather large, solitary, pedunculate, radiate. Involucral bracts wide, many-veined. Receptacle convex, epaleate. Ray florets female, fertile; limb white, many-veined. Disc corolla 5-lobed. Cypselas oblong, 8-10-ribbed, rather thin-walled, without myxogenic cells. Pappus a corona of small scales. DISTRIBUTION. Japan. - Monotypic. The monotypic Nipponanthemum is based on a handsome species formerly known as Chrysanthemum nipponicum Franchet ex Maxim. The position of Nipponanthemum is very unclear. It is provisionally placed as an aberrant member of subtribe Leucantheminae. It may on the other hand be more closely related to the relatively plesiomorphic members of Artemisiinae, i. e. Dendranthema or certain species in that genus. When he described the genus Kitamura considered it related to Argyranthemum but we find no support for his conclusion. 65. LEUCANTHEMELLA Tzvelev in Komarov, Fl. URSS 26: 137 (1961). Type species: L. serotina (L.) Tzvelev. Perennial herbs. Leaves alternate, entire or serrate. Capitula solitary or laxly corymbose, pedunculate, radiate. Receptacle convex, epaleate. Ray florets female, sterile; limb white or reddish, many-veined. Disc corolla 5-lobed, with a distinct enervate limb with sessile glands. Cypselas distinctly 7-12- ribbed, without myxogenic cells, with an apical rim but pappus absent. DISTRIBUTION. E. Europe (L. serotina) and Far East, China in Manchuria, Korea and Japan (L. linearis). - 2 spp. This genus is difficult to place (see Dienst, 1983). Tzvelev 140 also noted its isolated position. In habit it is similar to other members of subtribe Leucantheminae, where it is provision- ally placed. It may also be related to Dendranthema or some species of that genus. Leucanthemella has distinctly many- ribbed, non-myxogenic fruits. This may be related to habitat; the two species grow in marshy places. L. linearis (Matsum.) Tzvelev L. serotina (L.) Tzvelev 66. NIVELLEA Wilcox, Bremer & Humphries, gen. nov. Type species: N. nivellei (Braun-Blanquet & Maire) Wilcox, Bremer & Humphries. Herba annua. Folia lobata sublacerata. Capitula solitaria pedunculata radiata. Receptaculum epaleaceum. Flosculi radiati limbo albo. Flosculi disci tubo basaliter incrassato. Cypselae oblongae, 5-8-costata, sine canalibus secretoriis, sine cellulis mucilaginis. Pappus nullus. An annual herb. Leaves lobed and rather lacerate. Capitula solitary, pedunculate, radiate. Receptacle flat to convex, epaleate. Ray florets female, fertile; limb white. Disc corolla 5-lobed; tube basally swollen in fruit. Cypselas oblong, 5-8-ribbed, without resin canals and myxogenic cells. Pappus absent. DISTRIBUTION. N. Africa in Morocco. - Monotypic. The position ofNivellea is difficult to assess. It is superficially similar to some members of the Leucanthemum group, e. g. Leucoglossum, but does not have the specialized cypselas of that group. It is here provisionally placed as a basal member of subtribe Leucantheminae. N. nivellei (Braun-Blanquet & Maire) Wilcox, Bremer & Humphries, comb. nov. Basionym: Chrysanthemum nivellei Braun-Blanquet & Maire in Bull. Soc. Hist. nat. Afr. N. 13: 187 (1922). 67. PHALACROCARPUM (DC.) Willk. in Bot. Ztg 22: 252 (1864). Type species: P. oppositifolium (Brot.) Willk. Creeping, suffruticose perennials. Leaves opposite, sheath- ing, serrate to pinnatifid. Capitula solitary, pedunculate, radiate. Involucral bracts with dark brown margins. Recep- tacle convex, epaleate. Ray florets female, fertile; limb white or purplish. Central disc florets male (style-branches fused) or neuter. Disc corolla 5-lobed. Cypselas 7-9-ribbed; testa epidermis thick-walled and dark reddish, of elongate-sinuate cells. Pappus absent. DISTRIBUTION. SW Europe in Spain and Portugal. - 2 spp. Phalacrocarpum with opposite leaves is in habit and leaf shape similar to other members of Leucantheminae. The subtribal position of this genus must nevertheless be consid- ered provisional. P. hoffmannseggii (Samp.) Lainz P. oppositifolium (Brot.) Willk. (P. anomalum Cout.) K. BREMER AND C. J. HUMPHRIES 68. LEUCANTHEMOPSIS (Giroux) Heyw. in An. Inst. hot. A. J. Cavanilles 32: 181 (1975). Type species: L. alpina (L.) Heyw. Creeping or caespitose suffruticose perennials. Leaves alter- nate, serrate to pinnatifid, generally pectinate and spathulate in outline. Capitula solitary, rather long-pedunculate, radi- ate. Involucral bracts sometimes with dark brown margins. Receptacle convex, epaleate. Ray florets female, fertile; limb white to pinkish or yellow. Disc corolla 5-lobed; tube basally somewhat swollen in fruit. Cypselas 3-10- ribbed, with myxo- genic cells especially on the ribs, often with large 2-celled glands. Pappus a scarious flimsy corona. Flavonol 5-glycosides present. DISTRIBUTION. S. Europe mainly in Spain but extending to SW Russia (L. alpina), one species in N. Africa in Morocco (L. longipectinata). - 9 spp. Leucanthemopsis was originally a subsection of Tanacetum (sect. Pyrethrum), Tanacetum then taken in a very wide sense. Heywood (1954, 1976) noted its intermediate position between Tanacetum and Leucanthemum. Obviously, the Leu- canthemopsis species could not be accommodated in any of these two large genera, and the subsection was elevated to the rank of genus. New taxa have recently been described by Marchi (1980). In floral morphology and foliage Leucanthemopsis agrees with Hymenostemma, a possible sister group. Species of Leucanthemopsis are creeping or caespitose suffruticose perennials, whereas Hymenostemma is an annual herb. It seems to be the only reliable difference. On the other hand Hymenostemma groups with the specialized Prolongoa, both being annuals. The three genera clearly form a monophyletic group, but their interrelationships are uncertain. It is also possible that Leucanthemopsis is paraphyletic with Hymenos- temma and Prolongoa excluded. The matter requires further study. L. alpina (L.) Heyw. L. flaveola (Hoffsgg & Link) Heyw. L. longipectinata (Font Quer) Heyw. L. minima (Villars) Marchi L. pallida (Miller) Heyw. L. pectinata (L.) Lopez Gonzalez & Jarvis (L. radicans (Cav.) Heyw.) L. pulverulenta (Lagasca) Heyw. L. tatrae (Vierh.) Holub L. tomentosa (Lois.) Marchi 69. HYMENOSTEMMA (Kunze) Willk. in Bot. Ztg 22: 253 (1864). Type species: H. pseudanthemis (Kunze) Willk. - Prolongoa sect. Hymenostemma Kunze, pro parte. An annual herb. Leaves alternate, pinnatif id-pectinate, spathulate in outline. Capitula solitary, rather long- pedunculate, radiate. Receptacle convex to conical, epaleate. Ray florets female, sterile; limb white. Disc corolla 5-lobed; tube basally swollen in fruit. Cypselas 5-6-ribbed, with myxo- genic cells along the ribs. Pappus a scarious, flimsy corona. DISTRIBUTION. SW Europe in Spain and N. Africa in Morocco. - Monotypic. Hymenostemma is very similar in habit to Prolongoa, though GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 141 the latter is distinguished by a number of floral autapomor- phies. Hymenostemma differs from Leucanthemopsis by its annual habit, a feature shared by Prolongoa. The sister group relationships of these three genera are somewhat uncertain and it is even possible that Hymenostemma and Prolongoa, together or separately, have their sister groups within Leu- canthemopsis. 70. PROLONGOA Boiss., Voyage hot. Espagne 2: 320 (1840). Type species: P. hispanica Lopez Gonzalez & Jarvis. An annual herb. Leaves alternate, pinnatif id-pectinate, gen- erally spathulate in outline. Capitula solitary, rather long- pedunculate, radiate. Receptacle convex, epaleate. Ray florets neuter; limb yellow; pappus a large flimsy corona. Disc corolla 5-lobed. Style-branches long-penicillate. Cypse- las with 1 adaxial and 2 lateral rather thick ribs and 2 abaxial ribs, with myxogenic cells along the ribs; cypsela wall with rod-shaped crystals in small packets; pappus absent. DISTRIBUTION. SW Europe in Spain. - Monotypic. Prolongoa has been proposed and now accepted for conser- vation (Bremer et al., 1987). The species has been known erroneously as Prolongoa pectinata, a name that cannot be used since the basionym is a species of Leucanthemopsis. Prolongoa is related to Hymenostemma and Leucanthemop- sis. The matter is further discussed under these genera. Prolongoa has a number of autapomorphies, e. g. a reduced disc floret pappus and more strongly developed cypsela ribs. 71. LEUCANTHEMUM Miller, Card. Diet. abr. 4th ed. (1754). Type species: L. vulgare Lam. Perennial herbs with red-tipped roots. Leaves alternate, entire, serrate, or pinnatifid. Capitula solitary or laxly corym- bose, pedunculate, radiate or discoid. Receptacle convex or sometimes conical, epaleate. Ray florets female, fertile; limb white, pink, or rarely yellow. Disc corolla 5-lobed; tube basally swollen and spongy in fruit, especially abaxially. Cypselas c. 10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells along the ribs. Pappus a corona or an adaxial auricle, sometimes absent. Flavonol 5-glycosides present. DISTRIBUTION. Throughout Europe but mainly C. and S. parts, one species (L. discoideum) also in N. Africa in Morocco, Algeria, and Tunisia, some species (e.g. L. vul- gare) widespread as weeds. - 33 spp. With the removal of L. paludosum (=Leucoglossum palu- dosum) and L. arundanum (=Rhodanthemum arundanum) as well as the North African Leucanthemum species to Leucoglossum and Rhodanthemum, Leucanthemum becomes morphologically homogeneous and defined by its anthocya- nin red root tips. The latter are present in all herbaceous perennials (Favarger, 1966) which now constitute the genus Leucanthemum s. s. The genus is a polyploid complex (Vil- lard, 1970). L. adustum (Koch) Gremli L. aligulatum Vogt L. atratum (Jacq.) DC. L. burnatii Briq. & Cav. L. catalaunicum Vogt L. chloroticum A. Kerner & Murb. L. corsicum (Less.) DC. L. crassifolium (Lange) Willk. in Willk. & Lange L. cuneifolium Le Grand ex Coste L. delarbrei Timb.-Lagr. L. discoideum (All.) Coste (L. fontanesii Boiss. & Reuter, Chrysanthemum fontanesii (Boiss. & Reuter) Quezel & Santa) L. favargeri Vogt L. gaudinii Dalla Torre L. gracilicaule (Duf.) Alavi & Heyw. L. graminifolium (L.) Lam. L. heterophyllum (Willd.) DC. L. ircutianum DC. L. laciniatum Huter, Porta & Rigo L. lacustre (Brot.) Samp. L. leucolepis (Briq. & Cav.) Horvatic L. maestracense Vogt & Hellwig L. maximum (Ram.) DC. L. meridionale Le Grand L. merinoi Vogt & Castroviejo L. monspeliense (L.) Coste L. montserratianum Vogt L. pollens (Gay in Perreymond) DC. L. praecox (Horvatic) Horvatic L. pluriflorum Paul. L. subglaucum De Laramb. L. sylvaticum (Brot.) Nyman L. vulgare Lam. L. waldensteinii (Schultz-Bip.) Pouzar 72. RHODANTHEMUM Wilcox, Bremer & Humphries, comb, et stat. nov. Type species: R. arundanum (Giroux) Wilcox, Bremer & Humphries (Leucanthemum sect. Rhodanthemum Vogt, Leucanthemum subg. Chrysanthemopsis Maire, nom. nud.). Plantae perennes stolonibus saepe tegetes formantes. Folia alterna aggregata rosulata trifida vel ternato-pinnata ut vide- tur longi-petiolata. Capitula solitaria longepedunculata radiata. Bracteae involucri margine atro-fuscae. Receptacu- lum convexum epaleaceum. Flosculi radii feminei, vel fertiles vel steriles, limbo albido vel roseo vel rubro vel cremeo- aurantici multi-venoso. Flosculi disci corolla quinquiloba, lutea vel rubra tubo basi inflato et spongioso in fructus maturitate. Cypselae 5-12-costae valleculis lacunis et canali- culis secretoriis cum filis vascularibus et cellulis myxogenis intercostalibus instructis; costae protusae angustae et aliquan- tum aliformes. Pappus e corona scariosa vel auricula adaxiali basi coroniformi sistens. Stoloniferous and often mat-forming perennials. Leaves alternate, closely set, becoming rosulate, trifid or ternate- pinnatifid and seemingly long-petiolate. Capitula solitary, with long peduncles, radiate. Involucral bracts with dark brown margins. Receptacle convex, epaleate. Ray florets female, fertile or sterile; limb white, pink, reddish, or creamy orange, many-veined. Disc corolla 5-lobed, yellow or red; tube basally swollen and spongy at maturity of the fruit. Cypselas 5-12-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, and with myxogenic cells along the ribs; ribs protruding, 142 K. BREMER AND C. J. HUMPHRIES narrow and somewhat wing-shaped. Pappus with a scarious corona or an adaxial basally coroniform auricle. DISTRIBUTION. N. Africa in Morocco and Algeria, one spe- cies (R . arundanum) also in SW Europe in Spain. - 12 spp. Rhodanthemum (see Vogt, 1991) is a well characterized group of North African, Atlas montane perennials which form mats at relatively high altitudes (700-1200 m). They were formerly classified in Leucanthemum or Chrysanthe- mum but are distinguished by a number of synapomorphies. R. arundanum (Boiss.) Wilcox, Bremer & Humphries, comb, nov. Basionym: Pyrethrum arundanum Boiss., Voy. hot. Espagne 2: 317 (1840) (Leucanthemum arundanum (Boiss.) Cuatrec., Leucanthemum mairei Humbert). R. atlanticum (Ball) Wilcox, Bremer & Humphries, comb, nov. Basionym: Chrysanthemum atlanticum Ball in /. Bot. , Lond. 11: 366 (1873) (Leucanthemum atlanticum (Ball) Maire). R. briquetii (Maire) Wilcox, Bremer & Humphries, comb, nov. Basionym: Leucanthemum briquetii Maire in Bull. Soc. Hist. not. Afr. N. 15: 88 (1924). R. catananche (Ball) Wilcox, Bremer & Humphries, comb, nov. Basionym: Chrysanthemum catananche Ball in/. Bot. Lond. 11: 366 (1873) (Leucanthemum catananche (Ball) Maire). R. depression (Ball) Wilcox, Bremer & Humphries, comb, nov. Basionym: Chrysanthemum gayanum var. depressum Ball in J. Linn. Soc. 16: 509 (1878) (Leucanthemum depressum (Ball) Maire). R. gayanum (Cosson & Durieu) Wilcox, Bremer & Humphries, comb. nov. Basionym: Pyrethrum gayanum Cosson & Durieu in Bull. Soc. hot. Fr. 4: 15 (1857) (Chrysanthemum gayanum Ball, Leucanthemum gayanum (Cosson & Durieu) Maire). R. hosmariense (Ball) Wilcox, Bremer & Humphries, comb, nov. Basionym: Chrysanthemum maresii var. hosmariense Ball in J. Bot. Lond. 11: 366 (1873) (Leucanthemum hosmariense (Ball) Font Quer). R. maresii (Cosson) Wilcox, Bremer & Humphries, comb, nov. Basionym: Pyrethrum maresii Cosson in Bull. Soc. hot. Fr. 4: 16 (1857) (Leucanthemum maresii (Cosson) Maire). R. maroccanum (Battand.) Wilcox, Bremer & Humphries, comb. nov. Basionym: (Chrysanthemum maroccanum Battand. in Bull. Soc. Hist. not. Afr. N. 12: 189 (1921) (Leucanthemum maroccanum (Battand.) Maire). R. mesatlanticum (Emb. & Maire) Wilcox, Bremer & Humphries, comb. nov. Basionym: Leucanthemum mesat- lanticum Emb. & Maire, PI. Maroc Nov. (Arch. Sci. Maroc.) Fasc. II: 5 (1929). R. pseudo-catananche (Maire) Wilcox, Bremer & Humphries, comb. nov. Basionym: Leucanthemum pseudo- catananche Maire in Mem. Soc. Sci. nat. Phys. Maroc 15:37 (1926). R. redieri (Maire) Wilcox, Bremer & Humphries, comb. nov. Basionym: Leucanthemum redieri Maire in Mem. Soc. Sci. nat. Phys. Maroc 15:38 (1926). 73. LEUCOGLOSSUM Wilcox, Bremer & Humphries, gen. nov. Type species: L. paludosum (Poiret) Wilcox, Bremer & Humphries - Prolonged sect. Hymenostemma Kunze pro parte. Herbae annuae. Folia dentata-serrata vel pinnatifida. Capitula solitaria radiata. Receptaculum epaleaceum. Flos- culi radiati limbo albo vel flavido, basaliter luteo; pappus coroniformis scariosus vel adaxialiter auriculiformis. Flosculi disci corolla parum zygomorpha tubo incrassato; pappus plerumque nullus. Cypselae ellipsoideae parvae 7-10- costatae, inter costas lacunis vallecularibus canalibus secre- toriis et fasciculis vascularibus, in costis cellulis mucilaginis instructae. Annual herbs. Leaves alternate, dentate-serrate to pinnatifid. Capitula solitary, pedunculate, radiate. Receptacle convex or conical, epaleate. Ray florets female, fertile; limb white or pale yellow with a yellowish base; pappus a scarious corona or an adaxial auricle. Disc corolla 5-lobed, slightly zygomor- phic; tube basally swollen in fruit. Cypselas ellipsoid, com- paratively small, c. 1 mm long, 7-10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells along the ribs; generally the pappus is absent. DISTRIBUTION. SW Europe (L. paludosum) in Spain and N. Africa (all species) in Morocco, Algeria, Tunisia, and Libya. - 3 spp. The type species of this new genus has been recognized as anomalous within Leucanthemum e. g. in Flora europaea (Tutin et al., 1976). It differs by its annual habit, the small cypselas and the pappus, present in ray florets but absent in disc florets. Together with two North African species it is more closely related to Chlamydophora, another annual with small cypselas. Leucoglossum paludosum is often cultivated as an orna- mental. L. decipiens (Pomel) Wilcox, Bremer & Humphries, comb. nov. Basionym: Leucanthemum decipiens Pomel, Nouv. mat.fl. flf/.:59(1860). L. paludosum (Poiret) Wilcox, Bremer & Humphries, comb. nov. Basionym: Chrysanthemum paludosum Poiret, Voy. Barbarie 2: 241 (1789). (Leucanthemum paludosum (Poiret) Bonnet & Barratte, Leucanthemum setabense DC., Hymenostemma paludosum (Poiret) Pomel). L. reboudianum (Pomel) Wilcox, Bremer & Humphries, comb. nov. Basionym: Leucanthemum reboudianum Pomel, Nouv. mat. fl. all.: 291 (1860). 74. CHLAMYDOPHORA Ehrenb. ex Less., Syn. gen. Compos.: 265 (1832). Type species: C. tridentata (Del.) Ehrenb. ex Less. An annual herb. Leaves alternate or basally opposite, entire or tridentate, somewhat fleshy. Capitula solitary, peduncu- late, discoid. Receptacle convex, epaleate. Corolla 4- or 5-lobed, yellow or sometimes reddish; tube basally swollen and spongy in fruit. Cypselas ellipsoid, comparatively small, c. 1 mm long, 6-10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells on the ribs. Pappus a large, scarious, adaxial but basally coroniform auricle, as long as the corolla or longer. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE DISTRIBUTION. N. Africa in Tunisia, Libya, and Egypt, SE Europe in Greece and in Cyprus. - Monotypic. Chlamydophora is the monotypic and discoid sister group of Leucoglossum. The former C. pubescens is transferred to Aaronsohnia. C. tridentata is known also as Tripleurosper- mum indentation Hoffm. 75. CHRYSANTHOGLOSSUM Wilcox, Bremer & Humphries, gen. nov. Type species: C. trifurcatum (Desf.) Wilcox, Bremer & Humphries. Herbae annuae vel raro biennes. Folia pinnatisecta vel saepe trifurcata. Capitula solitaria, pedunculata, radiata. Bracteae involucri latae, plurinerves, flabelliformes. Receptaculum epaleaceum. Flosculi radiati limbo aureo; pappus adaxialiter auriculiformis scariosus vel nullus. Flosculi disci tubo incras- sato spongioso in fructu; pappus coroniformis rigidus extus cellulis mucilaginis instructus. Cypselae c. 10-costata, inter costis lacunis vallecularibus canalibus secretoriis et fasciculis vascularibus, in costis et corona cellulis mucilaginis instruc- tae. Annual or rarely biennial herbs. Leaves alternate, pinnati- sect, often trifurcate. Capitula solitary, pedunculate, radiate. Involucral bracts wide, many-veined, flabelliform. Recep- tacle convex, epaleate. Ray florets female, sterile or some- times fertile; limb golden yellow, many-veined; pappus a scarious adaxial auricle or absent. Disc corolla 5-lobed; tube basally swollen and spongy in fruit; lobes with appendages; pappus a rather stiff corona with myxogenic cells on the outside. Cypselas dorsiventrally somewhat compressed, c. 10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells along the ribs and also on the pappus. DISTRIBUTION. N. Africa in Morocco, Algeria, Tunisia and Libya. - 2 spp. Both species are readily recognizable as sister species by the distinctive leaves and cypselas. Chrysanthoglossum is the sister group of the three genera Glossopappus , Coleostephus , and Plagius. C. deserticola (Murb.) Wilcox, Bremer & Humphries, comb. nov. Basionym: Pyrethrum deserticola Murb., \r\Acta Univ. lund. 33(12): 98 (1897) (Chrysanthemum deserticola (Murb.) F. Buxbaum). C. trifurcatum (Desf.) Wilcox, Bremer & Humphries, comb. nov. Basionym: Chrysanthemum trifurcatum Desf., Fl. atlant. 2: 281 (1799) (Leucanthemopsis trifurcata (Desf.) Alavi). 76. GLOSSOPAPPUS Kunze in Flora, Jena 29: 748 (1846). Type species: G. chrysanthemoides Kunze (G. macrotus (Durieu) Briq. in Burnat). An annual herb. Leaves alternate, serrate-dentate, obovate- spathulate. Capitula solitary, pedunculate, radiate. Involu- cral bracts wide, flabelliform. Receptacle conical, epaleate. Ray florets female, fertile; limb golden yellow, many-veined. Disc corolla 5-lobed; tube basally much swollen and spongy in fruit, especially abaxially; lobes with appendages. Cypselas somewhat arcuate, 8-10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells along the ribs; basally with a 143 bulbous callus; pappus a large, scarious, adaxial, flabelliform auricle, as long as the corolla or longer. DISTRIBUTION. SW Europe in Spain and Portugal, N. Africa in Morocco, Algeria and Tunisia. - Monotypic. Glossopappus macrotus is related to Coleostephus and Plagius. Possibly Glossopappus has its sister group within a paraphyletic Coleostephus. The three genera form a mono- phyletic group. They are further discussed under Coleoste- phus. 77. COLEOSTEPHUS Cass. in Diet. Sci. Nat. 41: 43 (1826). Type species: C. myconis (L.) Reichenb. f. Annual herbs. Leaves alternate, serrate-dentate, spathulate. Capitula solitary or laxly corymbose, pedunculate, radiate. Involucral bracts wide, more or less flabelliform. Receptacle convex to conical, epaleate. Ray florets female, fertile; limb golden yellow, many-veined. Disc corolla 5-lobed; tube basally much swollen and spongy in fruit, especially abaxially; lobes with more or less developed appendages. Cypselas arcuate, 8-10-ribbed with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs, with myxogenic cells on the ribs; ribs basally and adaxially fused into a bulbous callus; pappus an oblique, adaxially more or less strongly developed scarious corona. Flavonol 5-glycosides present. DISTRIBUTION. S. Europe and N. Africa from Morocco to Libya. - 3 spp. Coleostephus is related to Plagius and Glossopappus (Alavi, 1968, 1988; Alavi & Heywood, 1976). The six species of these three genera form a monophyletic group but their interrela- tionships are unclear. Plagius with two species is diagnosed by being discoid and perennial (probably secondarily) but these characters are homoplasious and not supported by stronger synapomorphies. The monotypic Glossopappus has a specialized large pappus auricle. There is no synapomorphy to diagnose Coleostephus from the other two genera and it is possibly paraphyletic. Coleostephus and Plagius appear to be related by their more arcuate cypselas but the character is variable within Coleostephus. C. multicaulis (Desf.) Durieu (Chrysanthemum multicaule Desf.) C. myconis (L.) Reichenb. f. (Chrysanthemum myconis L., Kremeria myconis (L.) Maire) C. paludosus (Durieu) Alavi (C. clausonis Pomel, Chrysan- themum clausonis (Pomel) Battand., Kremeria paludosa Durieu) 78. PLAGIUS L'Herit. ex DC., Prodr. 6: 135 (1838). Type species: P. ageratifolius (Desf.) L'Herit. ex DC. (P. flosculosus (L.) Alavi & Heyw.). Herbaceous or suffruticose perennials. Leaves alternate, serrate-dentate, spathulate. Capitula solitary or laxly corym- bose, pedunculate, discoid. Receptacle flat or convex, epale- ate. Disc corolla 5-lobed; tube basally swollen and spongy in fruit; lobes with appendages. Cypselas arcuate, c. 10-ribbed, with vallecular lacunae and vallecular secretory canals as well as vascular strands between the ribs; basally with a bulbous callus, with myxogenic cells on the ribs. Pappus an adaxial auricle. Flavonol 5-glycosides present. 144 K. BREMER AND C. J. HUMPHRIES DISTRIBUTION. S. Europe in Corsica and Sardinia (P. floscu- losus) and N. Africa in Algeria and Tunisia (P. grandis). - 2 spp. The two species are very different from each other. Alavi & Heywood (1976) united them into one genus because both are discoid. P. grandis is instantly recognizable by the larger solitary capitula on unbranched, basally hairy stems. P. flosculosus , by contrast, has smaller capitula on branched stems and it is similar to species of the related genera Coleostephus and Glossopappus , except for the discoid capitula. Plagius, Coleostephus, and Glossopappus form a monophyletic group but their interrelationships deserve fur- ther study (see under Coleostephus). The species known as P. virgatus (Desf.) DC. is synonymous with Leucanthemum discoideum, following Alavi & Heywood (1976). P. flosculosus (L.) Alavi & Heyw. P. grandis (L.) Alavi & Heyw. (Chrysanthemum grandiflo- rum (Desf.) Battand.) 11. THAMINOPHYLLINAE Bremer & Humphries, subtrib. nov. Type species: Thaminophyllum multiflorum Harvey Herbae perennes vel suffrutices vel frutices. Folia serrata vel dentata vel integra vel solum pauciloba. Capitula solitaria, radiata. Cypselae oblongae vel ellipsoideae. Pappus coroni- formis vel nullus. Perennial herbs, half-shrubs or shrubs. Leaves serrate, den- tate, entire, or few-lobed only. Capitula solitary, radiate. Receptacle flat to conical, paleate or epaleate, sometimes pilose. Ray floret limb white or yellow. Disc corolla 4- or 5-lobed. Cypselas oblong to ellipsoid, 3-8-ribbed (or angled) or rarely 10-ribbed with costal veins and resin canals (Ade- nanthellum), without or rarely with large myxogenic cells in scattered groups (Thaminophyllum). Pappus a corona or absent. DISTRIBUTION (Table 22). S. Africa. - 5 genera, 17 spp. Table 22 General distribution of Thaminophyllinae and genera. x=indigenous, o=introduced. S. Afr. Thaminophyllinae Osmitopsis Adenanthellum Inezia Lidbeckia Thaminophyllum This subtribe comprises some small South African genera, which have received little attention in general treatments of the Anthemideae. Thaminophyllum, Lidbeckia, Inezia, and Adenanthellum are related as noted by earlier authors. Bond (1980) and Goldblatt (1980) stated that Thaminophyllum is related to Lidbeckia and Inezia. The unusual chromosome number x=10, known in Thaminophyllum and Inezia, is a feature mentioned by Bond and Goldblatt. Inezia was origi- nally described as a Lidbeckia. Adenanthellum was assumed to be most closely related to Inezia by Nordenstam (1976) when he originally described the genus. Osmitopsis was earlier classified in Inuleae because of the tailed anthers and its proper relationship within Anthemideae has hitherto not been considered. A preliminary chromosome count of 2n=10 as well as remarkable similarities between some species of Osmitopsis and the other genera indicate that these genera are related, and it is probable that together they form a monophyletic group. The cladogram is the single most parsimonious one derivable from the data matrix. Clades and characters - Fig. 11, Tables 2, 23. =jj= 79 Osmitopsis n= 80 Adenanthellum JJ= 81 Inezia h2^| n= 82 Lidbeckia 1!= 83 Thaminophyllum Fig. 11 Cladogram of the Thaminophyllinae produced by the ie option in Hennig86. Cladogram length = 25, consistency index = 92, retention index = 80. Table 23 Data matrix for the Thaminophyllinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 1 1111 11 1 1111 145057652 745966 5892337574081 24 177675 532718111 75620157784132928538262 365600 79.Osmitopsis allll?10a 1111000000000pOOOp?OpOO ???00p 80. Adenanthellum allll?lll 700011 11 111 11 1000070000 ?????? 81. Inezia al011?101 10001111000000110070000 ???00? 82.Lidbeckia 111117100 70001100000001011111000 ?????? B3.ThaminophyllumOinmOG 10001100000001011110111 ?????? Clade Thl - subtribe Thaminophyllinae 51 reversed. Floral parts without resin canals. Floral resin canals are present in a number of subtribes and also in Thaminophyllinae, although only in Adenanthellum. This is most parsimoniously explained as a reversal for the subtribe with a reappearance in Adenanthellum. Clearly, the character is difficult to interpret. 177 Chromosome number x=10. The chromosome number is unknown in Adenanthellum and Lidbeckia, but is neverthe- less best interpreted as a synapomorphy for the whole sub- tribe. Osmitopsis 45 Receptacle paleate. 56 Ray floret limb epidermis cells tabular (senecioid or muti- sioid type). This ligule epidermis type is uncommon in Anthemideae and reported for example in Osmitopsis (Baag0e, 1977). 92 Anthers caudate. Clade Th2 60 Ray floret tube confluent with the cypsela. 61 Ray floret tube persistent on the cypsela. In Adenanthellum and Inezia the tube sinus extends to the base and the tube is then virtually absent. Nevertheless the ray corolla, in these cases the limb, is both confluent with and persistent on the cypsela. Clade Th3 GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 145 5 Plants with one or few sparsely branched stems arising from a woody villous caudex. Some species of Osmitopsis are similar to Adenanthellum and Inezia in this aspect, as men- tioned by Nordenstam (1976) when grouping the latter two genera together. 57 Ray floret tube sinus extending to the base. Adenanthellum 51 Floral parts with resin canals. See note under clade Thl. 87 Disc corolla tube confluent with the cypsela. 98 Pollen grains hexa-panto-colporate. 124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 131 Cypselas 10-ribbed with costal veins and resin canals. 133 Cypselas with costal resin canals or sacs. This is character- istic, for example, of Lepidophorum in Leucantheminae and large part of Matricariinae but in Adenanthellum the cypselas are also 10-ribbed. Whether the resin canals are homologous in the two groups or not is difficult to assess. It is interpreted here as a parallelism. 172 Pappus absent in ray and disc cypselas. Inezia 52 reversed. Ray floret limb yellow not white. 59 Ray floret tube and cypsela pilose laterally; ray floret limb pilose abaxially. This character also occurs in one of the two Lidbeckia species (L. pectinata). 72 Disc corolla 4-lobed. Clade Th4 21 reversed. Leaves not serrate-dentate. 48 Receptacle pilose. 72 Disc corolla 4-lobed. 105 Stylopodium large and persistent in fruit. This character occurs also in some species of Osmitopsis. 172 Pappus absent in ray and disc cypselas. 183 Dehydrofalcarinone and dehydrofalcarinol present. The other genera of this subtribe are unknown chemically. Lidbeckia 18 Leaves with few, oblong to rounded, apically mucronate lobes. Thaminophyllum 2 Plants shrubby. 15 reversed. Leaves entire, not variously deeply lobed or divided. 26 Leaves closely set, lanceolate to linear. 142 Cypselas with large myxogenic cells in rounded, scattered groups. 79. OSMITOPSIS Cass. in Bull. Sci. Soc. philom. Paris 1817: 154 (1817). Type species: O. asteriscoides (P. Bergius) Less. Shrubs or half-shrubs. Leaves alternate, lobed, dentate, serrate, or entire, often glandular-punctate. Capitula solitary or laxly corymbose, radiate. Receptacle flat or convex to conical, paleate. Ray florets female, fertile or sterile, or neuter; tube occasionally pilose; limb white, many-veined, occasionally abaxially pilose; epidermis cells tabular. Disc corolla 5-lobed, glandular. Anthers caudate. Stylopodium sometimes large and persistent in fruit. Cypselas somewhat 3-4-angled or -ribbed. Pappus a corona of subulate to trian- gular scales, or absent. DISTRIBUTION. S. Africa in the Cape. - 9 spp. Until recently Osmitopsis has been placed in the Inuleae because of its tailed anthers. The traditional concept of the Anthemideae comprised genera without anther tails, a condi- tion which must be interpreted as a symplesiomorphy, how- ever. Tailed anthers have evolved independently within Anthemideae in the relatively unrelated genera Osmitopsis, Inulanthera (Gonosperminae), and Hippolytia (Tanacetii- nae). Stix (1960) snowed that Osmitopsis has anthemoid pollen and the tribal position in the Anthemideae was con- firmed by Bremer (1972) and Reitbrecht (1974). The position of Osmitopsis within Anthemideae has hith- erto remained unclear (Heywood & Humphries, 1977). It is placed here in Thaminophyllinae with other small South African genera with similar habit, foliage, many-veined rays, and the less usual chromosome number of x=10. A prelimi- nary count of 2n=20 was made by Bremer (unpubl., on O. pinnatifida) but it needs confirmation. Various but not all species of Osmitopsis are similar to other genera of subtribe Thaminophyllinae in a number of characters, particularly in habit, pilose and sterile rays, the loss of pappus and a large Stylopodium. The genus was revised by Bremer (1972, 1976). O. afra (L.) Bremer O. asteriscoides (P. Bergius) Less. O. dentata (Thunb.) Bremer O. glabra Bremer O. nana Schltr O. osmitoides (Less.) Bremer O. parvifolia (DC.) Hofmeyr O. pinnatifida (DC.) Bremer O. tenuis Bremer 80. ADENANTHELLUM B. Nord. in Bot. Notiser 132: 160 (1979). Type species: A. osmitoides (Harvey) B. Nord. - Adenanthemum B. Nord. A perennial herb with erect stems from a subterranean caudex. Leaves alternate, serrate, glandular-punctate. Capitula solitary, radiate. Receptacle convex, epaleate. Ray florets female, fertile; limb white, many-veined, confluent with the cypsela; tube absent (sinus extended to the base). Disc corolla 5-lobed, confluent with the cypsela. Pollen hexa-panto-colporate. Cypselas oblong, 10-ribbed, with 10 veins and 10 resin canals. Pappus absent. DISTRIBUTION. S. Africa in Natal and Transvaal and in Swaziland. - Monotypic. This recently described monotypic genus has a number of 146 K. BREMER AND C. J. HUMPHRIES autapomorphies, notably the unique pollen and the 10-ribbed cypselas with 10 resin canals. In habit and life-form, with a subterranean caudex generating herbaceous flowering stems, it is similar to Inezia, as indicated also by Nordenstam (1976) in his original description. A chromosome number of 2n=30 was reported by Goldb- latt (1980) who cited a preliminary, unpublished count by Nordenstam. From that one count (Nordenstam, pers. comm.) it is impossible to tell whether it represents an occasional triploid cell or a triploid specimen or taxon. The name Adenanthellum (Nordenstam, 1979) replaces Adenanthemum (Nordenstam, 1976), which turned out to be illegitimate. 81. INEZIA E. Phillips in Bull. misc. Inf. R. hot. Gdns, Kew: 297 (1932). Type species: /. integrifolia (Klatt) E. Phillips. Perennial herbs with erect stems from a subterranean caudex. Leaves alternate, entire, glandular-punctate. Capitula soli- tary, radiate. Receptacle convex, epaleate. Ray florets female, fertile; limb yellow, many-veined, abaxially and basally laterally pilose, confluent with the laterally pilose cypsela; tube absent. Disc corolla 4-lobed, glandular. Cypse- las oblong, 4-angled. Pappus of minute scales. DISTRIBUTION. S. Africa in Transvaal and in Swaziland. - 2 spp. Inezia is based on the transfer of /. integrifolia from the related genus Lidbeckla. Brusse (1989a) has shown that the two genera differ by several floral characters. Inezia has a stylopodium of thick-walled cells as compared with thin- walled cells in Lidbeckia and the nectaries are conspicuously larger in Lidbeckia. Also, there are 9-10 rows of cells in the filament collars of Lidbeckia as compared with 5-8 rows of cells in Inezia. Both genera have a 4-lobed corolla but nevertheless the sister group of Inezia is taken here to be Adenanthellum (with a 5-lobed corolla). These two genera have subterranean caudices from which herbaceous stems emerge (Nordenstam, 1976). They also have rays without a tube. It is more parsimonious to place Adenanthellum rather than Lidbeckia (as well as Thaminophylluni) as the sister group of this genus. /. integrifolia (Klatt) E. Phillips /. speciosa Brusse 82. LIDBECKIA P. Bergius, Descr. pi. Cap.: 307 (1767). Type species: L. pectinata P. Bergius Half-shrubs. Leaves alternate, glandular-punctate with few, oblong to rounded, apically mucronate lobes. Capitula soli- tary, rather long-pedunculate, radiate. Receptacle convex, epaleate, pilose. Ray florets female, sterile, or neuter; limb white, many-veined; sometimes dorsally pilose; tube some- times laterally pilose, confluent with the sometimes laterally pilose cypsela. Disc corolla 4-lobed, glandular, sometimes pilose; lobes sometimes with short acute appendages. Sty- lopodium large and persistent in fruit. Cypselas ellipsoid, 3-8-ribbed. Pappus absent. Dehydrofalcarinone and dehy- drofalcarinol present. DISTRIBUTION. S. Africa in the Cape. - 2 spp. The two species of Lidbeckia are related to Inezia and Thaminophyllum. They are held together as a pair by their similar foliage and habit. L. quinqueloba is more pubescent with pilose rays as in Inezia. On the other hand both species share a number of synapomorphies, for example the pilose receptacle, with Thaminophyllum. L. pectinata P. Bergius L. quinqueloba (L. f.) Cass. (L. lobata Thunb.) 83. THAMINOPHYLLUM Harvey, Fl. cap. 3: 155 (1865). Type species: T. multiflorum Harvey Shrublets. Leaves alternate, closely set, entire, lanceolate to linear, ericoid, glandular. Capitula solitary or laxly corym- bose, radiate. Receptacle convex or conical, epaleate, pilose. Ray florets female, sterile; limb white to somewhat purple- pink, many-veined; tube confluent with the cypsela. Disc corolla 4-lobed, glandular; lobes with acute or rounded appendages. Stylopodium large and persistent in fruit. Cypse- las ellipsoid, 3-4-angled, with large myxogenic cells in rounded, scattered groups. Pappus absent. Dehydrofalcar- inone and dehydrofalcarinol present. DISTRIBUTION. S. Africa in the Cape. - 3 spp. Thaminophyllum is a small and distinct genus of ericoid South African shrublets. The sister group is Lidbeckia. The two genera differ much in foliage but have a similar floral morphology and they both have a pilose receptacle. The myxogenic cells on the cypselas are of a type different from the common Anthemideae pattern (of rows of large myxo- genic cells). In Thaminophyllum the myxogenic cells are arranged in rounded groups, scattered on and hardly elevated above the epicarp. Thaminophyllum was revised by Bond (1980). T. latifolium Bond T. multiflorum Harvey T. mundtii Harvey 12. MATRICARIINAE Bremer & Humphries, subtrib. nov. Type species: Matricaria recutita L. Herbae annuae vel perennes vel suffrutices vel frutices. Capitula saepe solitaria vel raro corymbosa. Receptaculum epaleaceum vel raro paleaceum. Cypselae plerumque 5(4-6)- costatae costis varie dispositis vel interdum dorsiventraliter compressae et lateraliter alatae, plerumque abaxialiter et in costis sed non adaxialiter cellulis mucilaginis instructae. Pap- pus plerumque adaxialiter longior, coroniformis vel auriculi- formis vel e squamis discretis compositus vel nullus. Annual or perennial herbs, shrublets or shrubs. Leaves often much dissected, sometimes few-lobed or entire or ericoid. Capitula solitary or rarely corymbose, pedunculate or rarely sessile, radiate, disciform or discoid. Receptacle variously shaped, epaleate or rarely paleate. Ray floret limb white or yellow. Outer female florets (in disciform capitula) in one or more rows, sometimes stalked. Disc or central florets 5- or 4-, rarely 3-lobed. Cypselas generally 5(4-6)-ribbed with differ- ent rib arrangements or sometimes dorsiventrally flattened and laterally winged, generally with myxogenic cells abaxially and on the ribs but not adaxially. Pappus generally adaxially long, a corona, an auricle, of separate scales, or absent. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE DISTRIBUTION (Table 24). Worldwide, most genera in the Mediterranean region and in S. Africa, some Matricaria, Tripleurospermum, Cotula and Soliva species widespread as weeds. - 25 genera, 252 spp. 147 the absence of synapomorphies with genera of other subtribes they are provisionally retained in Matricariinae. The species of those two genera were formerly classified in Pentzia and Tripleurospermum. Rennera and Oncosiphon appear as sister Table 24 General distribution of Matricariinae and genera. x=indigenous, o=introduced. N.Am. Eur- Asia C.&E. SW S.Eur. N.Afr. Asia Asia S.Afr. Austr. S.Am. N.Zeal. Matricariinae x x X X X X XXX Cymbopappus X Pentzia X X Marasmodes X Rennera X Oncosiphon o X Otospermum X X Heteromera X Daveaua X X Matricaria x x X X X X O O Microcephala x X Endopappus X Myxopappus X Foveolina X Lonas X X Tripleurospermum o x X X X X o o o Aaronsohnia x X Leucoptera X Adenoglossa X Hilliardia X Cotula x o X XXX Leptinella X X Soliva x o X X Schistostephium X Hippia X Eriocephalus X The Matricariinae is our largest subtribe with respect to the number of genera (although the Artemisiinae, with Artemi- sia, contains more species). It consists mainly of a large group of predominantly annual and Mediterranean genera such as Matricaria and Tripleurospermum, and the predominantly South African Cotula group. Various South African genera are related to the Eurasian/North African taxa, for example, the annual genera Adenoglossa, Myxopappus, Foveolina, and possibly Rennera and Oncosiphon, as well as the South African shrubby genera Leucoptera, Cymbopappus, Pentzia and Marasmodes. The Cotula group is more narrowly and distinctly circum- scribed here than traditionally. It comprises the last seven genera in this account, characterized by their dorsiventrally flattened fruits. Hilliardia and Eriocephalus are both isolated and must be regarded as provisionally placed here, however. The whole subtribe is characterized by the arrangement of myxogenic cells on the cypselas, abaxially and on the ribs, but not on the adaxial surface. Furthermore, they have a pappus that is adaxially long, whether coroniform, auriculiform or of separate scales. Some genera do not have cypselas with myxogenic cells or with a pappus, for example, parts of the Cotula group. In the cladogram (Fig. 12) these two characters unite all Matricariinae, except the Cotula group, for reasons of parsimony. However, it is possible that they are synapo- morphies for the whole subtribe, since they occur also within the Cotula group. The positions of Oncosiphon and Rennera, both with epappose and non-myxogenic cypselas, are more doubtful. In groups in the cladogram. This is because they are similar in their mutual absence of some features, viz. myxogenic cells and pappus. In future studies they may turn out not to be closely related. The matter is further discussed by Kallersjo (1988). The generic interrelationships within the subtribe as shown in the cladogram (Fig. 12) must be regarded as very prelimi- nary. Many of the groupings are supported only by one or a few homoplasious characters, and numerous (226) equally parsimonious solutions are possible. We consider some sub- clades well supported, however. These include Ma3 with Cymbopappus, Pentzia and Marasmodes, Ma8 Heteromera and Daveaua, Ma9 Matricaria and Microcephala, Mal2 Myxopappus and Foveolina, Ma 16 Leucoptera and Adeno- glossa, Mal7 the Cotula group of genera (with the possible exclusion of Hilliardia and Eriocephalus}, Mal9 with Cotula, Leptinella, and Soliva, and Ma21 with Schistostephium, Hip- pia, and Eriocephalus (the last genus possibly to be excluded). All those clades are retained in the strict consen- sus tree of all the equally parsimonious cladograms found. Clades and characters - Fig. 12, Tables 2 and 25. Clade Mai - subtribe Matricariinae In this cladogram there are no synapomorphies for the subtribe but as explained above, characters 140 and 166 of clade Ma2 may be synapomorphies for the subtribe, with reversals in the Cotula group (Mal7). They appear as such in the alternative equally parsimonious cladograms. 148 K. BREMER AND C. J. HUMPHRIES |=Mal= tj= 84 Cymbopappus T=Ma3| |j= 85 Pentzia i!= 86 Marasmodes li= 87 Rennera r:Ma7=!i= 88 Oncosiphon F=Ma6:j!= 89 Otospermum 90 Heteromera 91 Daveaua 92 Matricaria Ma 9 93 Microcephala rp= 94 Endopappus f=Mall| ij= 95 Myxopappus ltMa!2^ 96 Foveolina n= 97 Lonas Mal3=] rj= 98 Tripleurospermum I M=1 A -I >&= 89OtC iLMasE =Mal4: 102 Hilliardia 99 Aaronsohnia fj= 100 Leucoptera Mai 6^= 101 Adenoglossa =Mals 103 Cotula rj= 104 Leptinella a20^ 105 Soliva 106 Schistostephium rf= 107 Hippia a22=!i= 108 Eriocephalus Fig. 12 Cladogram (of 226 possible) of the Matricariinae produced by the bb option in Hennig86. Cladogram length = 97, consistency index = 60, retention index = 68. Table 25 Data matrix for the Matricariinae. 1 = presence, = absence, 7 = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 1111 11 1 11 11111111 111 11111 111 1 1 11 111 14505765 46 89333 77578226712174424372433784415541291279760736601545 53271811 06220543132323387359816779059576531126824457564724063711500 1 111 1755555666671 82 3635679056806922 84. Cymbopappus 11111710 85. Pentzia 86. Marasmodes 87. Rennera 88. Oncosiphon 11111711 89. Otospermum 11111711 90. Heteromera 11111711 91. Daveaua 11111711 92. Matricaria 11111111 93. Microcephala 11111711 94. Endopappus llall?ll 95. Myxopappus 10711711 96. Foveolina 11111711 97. Lonas 11711711 98. Tripleurospermum 11111111 99. Aaronsohnia 11111711 100. Leucoptera 11111711 101. Adenoglossa 01011711 102. Hilliardia 11111710 103. Cotula 11111711 104. Leptinella 11111711 105. Soliva ll?ll?la 106. Schistostephium al 71 1710 107. Hippia 11711710 108. Eriocephalus 11111710 Hal 100000000000000000000000000070000000000000000000000000? 1 11 Illp00pp0000000?00?000000p000?00000000000000000000 70000? 1111111000000000007007000000000070000000000000000000070000? 070001001 1 1 10000007007000000000070000000000000000000070000? 0?010p0011101110000000pOOOOOpOOOOOOOOOOOOOOOOOOOOOOOOOOOOO? 1100000011000001110000000000000070000000000000000000000000? 11000001 11 000000001 100000000000070000000000000000000000000? 1101000111000000001011000000000070000000000000000000000000? HOOOOOalOpOpOOOOOOOOplllOOpOOOOOOOOOOOOOOOOOOOOOOOpOOOOOO? lOOOpOOlOOOOOOOOlOOOOal 1 100000070000000000000000000000000? 110000001 0000000000 10000001 00000701 HOOOOOOOOOOOOOOpOOOOOO? 110001 001 OOOpOOOOO? 10700001 00000701001 110000000000000 700000 HOOOpOOlOOOpOOOOOOaOOOOOOl 00000700001 lOOOOOOOOOOOOpOOOOOO? 11000 101 10000000007107000010 11 1000000000000000000000000000? a lOOOpOlaOpOOOOOOOOlOOOOOOOapOOl 11000000000000000000070000? 11010p0110p000000000000000110000?OOOOOOOOOOOOOOOOOOpOOOOOO? 11 100001 000000000000000000 laOOOO?00100001 11 100000000000000? 1100000110000000000000000010000070010000111111100000000000? 0710000000 101 00000000000001 00000??OOOOOOOOOa0001apOappOOOO? P?000p0ap010100000000000001p000001000000100a0001011app0000? P?000p0a0010100000?00?00001p0000??000000a001000?0101111000? 070000 101 010 100000700? 0000 100000770000001000000100000001 10? 0?100pOOOO 10 100000700? 000010aOOO??00000010000000aOOaOOOOOO? 0710000000 10000000700? 0000 10aOOO??000000 100100000 101 000000? 07100000001 000000000000000 lOa 100? 1000000100000000 lOpOOOOOl a 7700000000000000 7000000000070000 7700000000000000 7700000000000000 7700000000000000 ?OOOOOOOOOOOpOOO 7700000000000000 7700000000000000 ?0?????????00000 ?OOOOOOOOOOOpOOO 7700000000000000 7700000000000000 7700000000000000 7700000000000000 ?0000000000?OOpO 7777777700070000 ????????000?000p 7700000000070000 7777777700070000 7777777700070000 7700000000070000 Clade Ma2 140 Cypselas with myxogenic cells on ab axial surface and on the ribs of the adaxial surface. This character is assumed to be secondarily lost in Rennera and Oncosiphon. Several species of Tripleurospermum are also devoid of myxogenic cells, presumably secondarily. The character occurs also within Cotula and Leptinella. 166 Pappus adaxially long. This character comprises a variety of different but homologous pappus types. The pappus is secondarily lost in Rennera and Oncosiphon. Some species of Matricaria, Tripleurospermum and Pentzia also lack pappus, presumably secondarily. Clade Ma3 2 Plants shrubby. 51 reversed. Floral parts without resin canals. GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 149 82 Disc corolla tube thickened in fruit. This character is widespread in Achilleinae and Leucantheminae and occurs also in other genera of Matricariinae, notably Oncosiphon and Aaronsohnia. 90 Disc corolla tube and and cypsela ribs with thick vascular strands. Cymbopappus There is no clear autapomorphy for this genus. Clade Ma4 35 Capitula discoid. Pentzia There is no autapomorphy for this genus. Marasmodes 34 Capitula sessile along the stems. Clade Ma5 1 Plants annual. Clade Ma6 73 Disc corolla lobes with dorsal appendages. Clade Ma7 140 reversed. See clade Ma2. 172 Pappus absent in ray and disc cypselas. Rennera 35 Capitula discoid. 153 Cypsela thick-walled and conspicuously rugose. Oncosiphon 72 Disc corolla 4-lobed. Cotula and related genera also have 4-lobed corollas. 82 Disc corolla tube thickened in fruit. 83 Disc corolla tube very thick and brittle. 123 Cypselas with a mainly abaxial entire or toothed rim. Otospermum 128 Cypselas with 1 abaxial and 2 lateral thick ribs and 2-3 adaxial ribs. The arrangement of the three major ribs is abaxial-lateral and thus opposite to that of Tripleurospermum and many other genera. 167 Pappus a stiff adaxial auricle. 173 Testa epidermis cells spirally arranged around the embryo. A similar testa occurs in Microcephala. Clade Ma8 115 Ray cypselas dorsiventrally flattened with 3 adaxial ribs. 133 Cypselas with costal resin canals or sacs. Heteromera 129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. The same rib arrangement occurs in Tripleurospermum and related genera of clade MalO. Daveaua 82 Disc corolla tube thickened in fruit. 118 Ray cypselas laterally winged; wings projected to apical teeth. 111 Pappus absent in disc cypselas, but present in ray cypselas. Clade Ma9 46 Receptacle narrowly conical to subulate. 47 Receptacle hollow. 127 Cypselas with 5 mainly adaxially arranged ribs. Matricaria 133 Cypselas with costal resin canals or sacs. Microcephala 149 Cypselas with rather stiff unbranched hairs of 3-8 cells with spiral wall thickenings. 173 Testa epidermis cells spirally arranged around the embryo. A similar testa occurs in Otospermum. Clade MalO 129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. The same rib arrangement occurs also in Heteromera. Foveolina has rather weak, but similarly arranged ribs. 130 Cypselas with 2 lateral vascular strands, sometimes also with 1 adaxial strand. Tripleurospermum has a different autapomorphic type with 5 strands, with 2 abaxial strands associated with the two apical-abaxial resin sacs. Clade Mall 141 Cypselas with dense rows of myxogenic cells also on the corona. This character is not expressed in Foveolina. Endopappus 41 Involucral bracts wide, flabelliform. 107 reversed. Cypselas not terete to weakly angled or flattened, but sharply angled. 112 Cypselas large, with 3 thick protruding sclerenchymatous ribs, somewhat winged in ray cypselas. Clade Mal2 156 Cypsela wall white and spongy; pericarpic cells isodiamet- ric with thin spiral wall thickenings. (Kallersjo, 1988). 158 Cypsela wall with numerous druses in the pericarp. (Kallersjo, 1988). Myxopappus 35 Capitula discoid. 42 Involucral bracts subulate. 43 reversed. Involucral bracts without scarious margins. Foveolina 141 reversed. See clade Mall. Clade Mal3 133 Cypselas with costal resin canals or sacs. Lonas 150 K. BREMER AND C. J. HUMPHRIES 29 Capitula densely corymbose. 35 Capitula discoid. 45 Receptacle paleate. 137 Cypselas with a single resin sac apically in the adaxial rib. Clade Mal4 75 Disc corolla lobes with central resin sacs. The character is expressed in most but not all species of Tripleurospermum, in Aaronsohnia and in Leucoptera. It reverses in Adenoglossa. Tripleurospermum 130 reversed. See clade MalO. 136 Cypselas abaxially and apically with 2 distinct, occasion- ally 1 or 3-5, resin sacs. 175 Embryo sac tetrasporic. 183 Dehydrofalcarinone and dehydrofalcarinol present. These products in polyacetylene synthesis are common in the Aster- aceae but uncommon in Anthemideae, occurring also in Cotula and Eriocephalus. They may represent a secondarily simplified metabolism (Greger, 1977). The position of the character on the cladogram is uncertain because most genera of the subtribe have not been investigated chemically. Clade Mal5 129 reversed. See clade MalO. Aaronsohnia 82 Disc corolla tube thickened in fruit. Clade Mal6 24 Leaves rather fleshy , few-lobed or entire. 41 Involucral bracts wide, flabelliform. 95 Anthers with endothecial tissue partly or wholly polarized. 114 Cypselas dorsiventrally flattened. This is a traditional character of the Cotula group. 117 Cypselas laterally winged. Winged fruits occur also within the Cotula group, clade Mal7. Leucoptera 1 reversed. See clade Ma5. 2 Plants shrubby. Several other genera in the Cotula group, as well as Pentzia and related genera, are also more or less shrubby. Adenoglossa 15 reversed. Leaves not variously deeply lobed or divided, but entire. 25 Leaves rather fleshy , entire, linear. 52 reversed. Ray floret limb not white, but yellow. 75 reversed. See clade Mal4. 76 Disc corolla zygomorphic with 2 smaller adaxial lobes with marginal resin canals extending from the base of the corolla and with 3 larger abaxial lobes. 94 Anthers with an apical resin sac. Clade Mal7 - The Cotula group of genera 72 Disc corolla 4-lobed. Some Cotula and Soliva species have 3-lobed corollas, another step in the reduction of the basically 5-lobed corolla. Hippia and Eriocephalus have 5-lobed corol- las and the character is thus assumed to be reversed in these two genera. Oncosiphon also has 4-lobed corollas. 130 Cypselas with 2 lateral vascular strands, sometimes also with 1 adaxial strand. 172 Pappus absent in ray and disc cypselas. 177 Chromosome number x=10. A variety of chromosome numbers are known from Cotula but 10 appears to be the base number. Most of the related genera are not investigated karyologically but Hilliardia also has x=10. Eriocephalus, however, has x=9, the plesiomorphic base number of the tribe. 183 Dehydrofalcarinone and dehydrofalcarinol present. Most of the related genera have not been investigated chemically. However, these substances have been found also in Tripleu- rospermum. See further comments under that genus. Hilliardia 2 Plants shrubby. 51 reversed. Floral parts without resin canals. Ill Cypselas subglobose, with 2-3 very thin lateral-adaxial ribs. 155 Cypsela wall very thin, translucent and showing brownish black, rounded, very thick-walled testa epidermis cells. Clade Mal8 114 Cypselas dorsiventrally flattened. Clade Mal9 117 Cypselas laterally winged. 133 Cypselas with costal resin canals or sacs. Note: Leptinella and Soliva probably have their sister group(s) within Cotula as discussed under these genera. This is based on information other than that used to construct the cladogram. Thus Cotula could be paraphyletic and the char- acters appearing for Cotula in the cladogram should rather be interpreted as defining this clade, including all three genera, with reversals in Leptinella and Soliva. Cotula 62 Outer florets stalked. This peculiar receptacular structure is a classical character for Cotula, though not particularly well developed in some species. Quite clearly it occurs also in Schistostephium and it is tempting to assume that it is a character for clade Mal8 although secondarily lost in the other genera of that clade. With this cladogram it is more parsimoniously interpreted as a parallelism between the two genera, however. Clade Ma20 104 Disc floret style-branches fused. The same character occurs in Hippia and Eriocephalus. Leptinella GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 151 70 Corolla inflated with a hollow space between outer surface and inner layer. Soliva 1 Plants annual. Many species of Cotula are also annuals. 34 Capitula sessile along the stems. 36 Capitula disciform. Most species of Cotula as well as Hippia and Schistostephium are also disciform and it may be that the character is a synapomorphy for these genera. It is in conflict with other characters, however, and the presence of plesiomorphic, radiate species of Cotula, as well as the inclusion of Eriocephalus in clade Mal8, makes interpreta- tion difficult. 63 Outer female florets in several rows. This character also occurs within Cotula, possibly indicating the sister group of Soliva within Cotula. See note under clade Mal9. 67 Outer female florets without corollas. 101 Style persistent and spinescent in fruit. 133 reversed. See clade Mal9. Clade Ma21 2 Plants shrubby. 29 Capitula densely corymbose. 51 reversed. Floral parts without resin canals. Ill reversed. See clade Mal7. Schistostephium 36 Capitula disciform. See comment under Soliva. Some Schistostephium species are discoid, here considered a further development from the disciform condition. 62 Outer florets stalked. See comment under Cotula. Clade Ma22 72 reversed. See clade Mal7. 104 Disc floret style-branches fused. Hippia 36 Capitula disciform. See comment under Soliva. 117 Cypselas laterally winged. Eriocephalus 40 Involucral bracts in 2 unequal series. 45 Receptacle paleate. 50 Receptacular paleae pilose. 84. CYMBOPAPPUS B. Nord. in Bot. Notiser 129: 150 (1976). Type species: C. lasiopodus (Hutch.) B. Nord. (C. piliferus (Thell.) B. Nord.). Shrubs or half-shrubs. Leaves alternate, variously lobed or dentate, more or less ericoid. Capitula solitary, pedunculate, radiate. Receptacle convex to hemispherical or subconical, epaleate. Ray florets female, fertile; limb white or pinkish. Disc corolla 5-lobed; tube swollen and with thick vascular strands. Cypselas 5-ribbed, with large 2-celled glands and with myxogenic cells mainly abaxially; ribs with thick vascular strands. Pappus an oblique, adaxially longer whitish corona or cup, or of one large adaxial and one smaller abaxial scale. DISTRIBUTION. S. Africa in the Cape, Transkei, and Trans- vaal. -4 spp. Cymbopappus based on species formerly classified as Chry- santhemum and Marasmodes (Nordenstam, 1976) is close to Pentzia. They are distinguished by Cymbopappus being radi- ate and Pentzia discoid. Hence the species of Cymbopappus have not been usually associated with Pentzia although Nordenstam (1976) noted the relationship. Cymbopappus also has an almost tubular pappus but a similar pappus occurs in species of Pentzia. The distinction between these two genera deserves further study. C. adenosolen (Harvey) B. Nord. C. hilliardiaeB. Nord. C. piliferus (Thell.) B. Nord. C. lasiopodus (Hutch.) B. Nord. 85. PENTZIA Thunb. in Prodr. pi. cap.: 145 (1800). Type species: P. crenata Thunb. (P. dentata (L.) OK.). Shrubs. Leaves alternate, variously pinnatisect, occasionally entire or apically dentate, more or less ericoid. Capitula solitary or corymbose, pedunculate, discoid. Receptacle con- vex or hemispherical to conical, epaleate. Corolla 5-lobed; tube generally swollen and with thick vascular strands. Cypselas 5-ribbed, often with large 2-celled glands and with myxogenic cells abaxially and on the ribs; ribs with thick vascular strands. Pappus an adaxial auricle, or an oblique adaxially longer cup, or of free adaxially longer whitish scales, sometimes absent. DISTRIBUTION. S. Africa mainly in the Cape, also in Namibia, two species (P. hesperidum and P. monodiana) also in N. Africa in Morocco and Algeria. - 23 spp. Traditionally (sensu Hutchinson, 19l6a,b) Pentzia was all South African discoid Anthemideae with pinnatisect leaves and an epaleate receptacle. Two North African, typical Pentzia species, have also been described. Kallersjo (1988) has recently re-classified Pentzia and most South African species included in Tripleurospermum have been assigned to a number of genera, including Pentzia, Oncosiphon, Myxo- pappus and Foveolina. Pentzia now comprises discoid shrubs (no annuals) with more or less ericoid leaves and 5-ribbed, myxogenic cypselas. The separation of Pentzia from the small genera Cymbopa- ppus and Marasmodes is problematical. In future it might turn out that the latter genera have their sister group within a presently paraphyletic Pentzia. Clearly the generic delimitation of Pentzia is in need of detailed investigation, even after the removal of the annuals, as was undertaken by Kallersjo. At the species level Pentzia is also in need of revision. Some of the 'species' in the list below are probably conspe- cific but the list is taken from Kallersjo (1986). References are also given to species described after Hutchinson's ) revision since they were not included in his key. P. argentea Hutch. P. bolusii Hutch. P. calcarea Kies - Note: Description in Kies, 1945. 152 K. BREMER AND C. J. HUMPHRIES P. calva S. Moore P. cooperi Harvey P. dentata (L.) OK. P. elegans DC. P. globosa Less. P. hesperidum Maire & Wilczek - Note: Description in Maire, 1936. P. incana (Thunb.) OK. P. lanata Hutch. P. monocephala S. Moore P. monodiana Maire & Wilczek - Note: Description in Maire, 1929. P. nana Burch. P. peduncularis B. Nord. - Note: Description in Nordenstam, 1987. P. pinnatisecta Hutch