ISSN 0968-0446 Bulletin of The Natural History Museum Sana wea - SDEC1993 PREGENTED GENERAL LIBRARY —w THE NATURAL HISTORY MUSEUM VOLUME 23 NUMBER 2 25 NOVEMBER 1993 TNS es 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. 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(Bot.) 23(2): 55-59 Issued 25 November 1993 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region BRITISH MUSEUS RLATURAL WSTORYS TING-NUNG HO AND SHANG-WU LIU Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China ~SBEC1993 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. chateriT. N. Ho, G. hicksii H. Smith, G. masonii T. N. Ho, G. zekuensis T. 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*Among them three species were diagnosed, described and typified and one species labelled by Harry Smith but were left unpublished at his death in 1971.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 decoratae Diels et Gentiana infelici C. B. Clarke affinis, sed a prima tubo corollae lobis multo longiore (nec 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, S—6 mm longus, lobis ellipticis, 3-3.5 mm longis, c. 2 mm latis, apice acuminatis, mucronatis; corolla alba (in sicco lutescens), calyce triplo 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. 7 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 xX 0.4 mm magna, tenuiter reiculata. China: S Tibet: Tsari, Karkyn la, on open grassy hillside, 4420 m, 24 June 1936, Ludlow & Sherriff 2209 (BM); ibid. on mossy open hillside, 4000-4270 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, 28°45’ N 94°E, on open grassy hillside, 4420 m, 5 June 1938, Ludlow, Sherriff & Taylor 3963 (BM, E); Kongbo, Kusha la near Paka, 29°15’ N 94°2’ 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, © The Natural History Museum, 1993 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); Jula, 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, 34 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; illa caulina 2—3-juga, remota, ovato- lanceolata, 7-18 mm longa et 3-7 mm lata, apice acuta, basi breviter petiolata. Flores saepe 3-4 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 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 (nec 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; illa caulina 34-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. 6mm 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 masonii 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, illa 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 34 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. 1.5mm _ longis; ovarium cylindricum, 11-13 mm longum, breviter stipitatum, stylo breve, stigmate breve. Upper Burma: N’Maikha-Salwin divide, 26°28’ N 98°48’ E, September 1924, Forrest 24944 (K); ibid. 26°50’ N 98°48’ 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 (nec truncato), lobis filiformibus, usque ad 5 mm longis (nec dentiformibus, 1-1.5 mm longis), corolla flavida, caeruleo-punctata, versus limbum caerulea, raro corolla omnino caerulea (nec 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; illa 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 ().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. 1mm 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, S. 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 illa 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; illa 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 TAXA OF GENTIANA ST. 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, 29°59’ N 94°17’ E, 4115 m, 30 August 1938, Ludlow, Sherriff & Taylor 6944 (BM), Kongbo, 30°N 94°20’ 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, J. 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. bot. 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; illa 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 Fig. 2. Gentiana zekuensis S. W. Liu: a, habit ; b, flower; ¢, 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. 1mm _ 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 97°45’ 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 micans 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 (nec acicularibus, caule fere adpressis), lobis calycis brevioribus, latioribus, breviter acutis (nec longe acuminatis), plicis bifidis (nec 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; illa 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 GENTIANA acutis mucronatisque, plicae late ovatae, c. 1 mm longae, lobis breviores, acutae, integrae vel bilobatae; stamina medio corol- lae inserta, filamentis c. 1mm _ longis; ovarium obovato- oblongum cum stipite c. 4mm _ 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, pl. 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 affinis, sed foliis ovatis, lobis calycis triangularibus, multo brevioribus bene recedit. Herba annua, 8-10 cm alta. Caulis ascendens, ramosus. Folia basilaria ignoti; illa 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. 1 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. noy. (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; illa basilaria majora, anguste oblonga, 25-42 mm longa et 3.5-8 mm lata, sessilia vel subsessilia; illa 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 Se 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: Ninggiang, 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. 2mm 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; illa caulina 1—2-juga, eis basilaribus similaria, 2-3.5 cm longa et ad 1.2cm 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. bot. Sada 15: 1-507 (1896-1904)]. Naithani, H. B. 1990. Gentiana. In Flowering plants of India, Nepal and Bhutan: 291-295. asi Rae ea ne: a eee ae 6 oy nd 7 " a wa ~ . we a - , A ee B: - : ru Ce een Wan Ct ea ee ae 4 Sor, es ec — 7 * a ratte - Bull. nat. Hist. Mus. Lond. (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 © The Natural History Museum, 1993 (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), nec Rusby (end 1896) (Mem. Torr. Bot. Club , 6: 79), Type: Bolivia, M. Bang 1153 (K! - isotype). Gentianella bellatula (Gilg) T. N. Ho & S. W. Liu, comb. nov. Gentiana bellatula Gilg, Bot. Jahrb., 50(Beibl. 111): 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., Glett. 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. 62 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, Stuebel 41 (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) (J. 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. T.-N. HO AND S.-W. LIU 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 (Giig) 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. 111): 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, 7. 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. f 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 6051 (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 (P!). 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. Liu, 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: Iles 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. boliviana and G. lobelioides) which are better treated as the following two varieties. The difference between var. peruviana and var. boliviana 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.118): 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. 111): 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), nec 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. Tecn. 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, Herzog 2046b. (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. Tecn. 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, Stuebel 24b, 25 (all 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, Stuebel 35f (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, S. 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 III. 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, Ann. Missouri Bot. Gard., 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 Comastoma (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. Sa Doe: ae —_ Sy —_ aru 7 a > han Lileks Bull. nat. Hist. Mus. Lond. (Bot.) 23(2): 67-70 Studies in Hypericum: validation of new names NORMAN K.B. ROBSON clo Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS MIPEGOMCHONY Sosy aeesh sc sco eac adh pats seb nai a ok okie Nido GIRO LWT a HONG RRA SRI EU SO OE aS Ua ats re RL ID, 67 SECM OnsMyrididid (s0ach) Ro meienT coite ire wae tae cca na cue coals Me Nu Bee Docegan Sua ca NS pony Ses CTS 67 MECHOM 255, HITRAENOLUES FAllbrsae SP ACM occ etki etowhy scence tes uiekg Vue wets area cc deans wena anus Sylacreeredtih were 68 Section Zon vnenoimas: (aula space) ih Keller: sain atc auuadomapeontey haoayehe sok aed dob mnana damien anm doeiwes 68 DECUOM 2 1 cACEMOSCO AEE SP ACIR 52 ot, a wcis cwlse spins amis ste meet nace nes oat ce na aen ols teste Diuiaks Sieieunledadtse Medel sas wee ewe oes 69 IREfETen Ce Stes tars seesd. sane amteeae eck eee tee Oe eae Sear tes seit oleeeeiet a catenin er ee ee tere een ck SeeMa une MUmeN to erence: ss 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,H. 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. Issued 25 November 1993 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. Pl. 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 Hypericum hypericoides (L.) Crantz, /nst. rei herb. 2:520 (1776). Ascyrum hypericoides L., Sp. pl. :789 (1753) excl. syn. Hort. Cliff. et Plukenet., 2nd ed.: 1108 (1763) excl. syn. Plukenet. Type: Hispaniola, Hypericoides frutescens erecta, flore luteo Plumier, Nov. pl. 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 (1703). Ascyrum foliis lanceolato-linearibus, biglandulosis, ramidiffu- sis Burman, Pl. amer.: 146, t.152 f.2 (1758). A subsp. hypericoides habitu prostrato, 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. Jnflorescence- 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. Campylosporus), 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. N.K.B. ROBSON 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—-S-florata, floribus maioribus sepalis crassioribus, capsulis coriaceis valvis leviter angustissime vittatis fere laevibus, inter alia differt. Type: Socotra, Hagghiher Mountains(12°35’N, 54°03’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 xX (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. SA (1836), Hist. nat. vég. 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 + (0.40.5 mm) petiolate, always (?) + incurved-cucullate; lamina 3-6 X 1-2 mm, narrowly oblong to broadly elliptic, acute. Sepals 3.5—S mm long. Petals 6.5—9 mm long. Libya (Cyrenaica—Jebel el Akhdar). STUDIES IN HYPERICUM Section 27. ADENOSEPALUM Spach When Sect. Adenosepalum (sensu Robson, 1977) has been ‘purified’ by the removal of the tropical Asian species(H. 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, Ill. 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.1982 (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 69 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) Fiori 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. bot. Etat Brux. 29, Suppl.: 588 (1959); Moggi & Pisacchi in Webbia 22: 272 (1967) pro parte; Collenette, //l. 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 & Verdc., Fl. Trop. E. Afr. Guttif.:30 (1978). 70 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. N.K.B. ROBSON 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. —— 1953b. Hypericaceae. Jn 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). Jn 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. nat. 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, Villavagen 6, S-752 36 Uppsala, Sweden CHRISTOPHER JOHN HUMPHRIES Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS INtROGUCHON Gece site ees roses ara non ta scisoeetcnaat es nese ee ane sie coe cnaa sine cuss @eusile sama onaertilldennctuislsnsncaniscsantassas 73 Daterals/ aiid MmetnOds 0.2.5 cccysk vanarstaccseu ss ndsrosgs tens stsvens wales a elsows UsteniiebidaRiavaaives sadvie davddn steucuadved adel as Acvetedees 73 DDESCRIGEVE TeTMmINOMO Ye. as:.5, sad fia shard diioe cea sdannoictes ara eleaie nash ju’ug sincan eon bse cies eric sles nal ca nnen anneal tmacede seianses 74 SPAXONOMIC {CONCEDES eee ac cera oh oat tea cias Soe Setlists toss eer chclnaclaniolettina ste isgaetaird Tests tiswle sashes aos aoa eats aenie 74 (OLE (eT T esata ar iran isancetn ha csutad denne ongbed aoGhen Son ans AUR EnacC OGen aceE RA AUBUAr ceione cr inontar nena tnreannncea nn drondcdrc coda 74 RribaliandsubtribalsGivisionsaccctecc csc cte scene eee een acta reas on einen as ie eae eat ec RR eerie socal saat ahs eaetete 74 CSTASSTRICH NON hate Bo 9t Ac ae ero ba ast asics Soe Ore SAN NEES GE accede ac bua aS EGINET SUG ANS SASTIE GA Caan Av cay aanets eee cea cuanes SORE 76 Subtribal classifications 2. 24.c5sccss.c tastes ac fee ces Saeco cease aioaciac se nsileece dene s aaasienenen once ascesacimeasasesedentanssesee ca 76 Generic classifications 4z.1- coe sesun teases a cescsee aes se ase eae ea ous Seon eres sa senee ore earn: Sates acme eealstar nae are noes Ti. Characters Of (he AmMtnegndeaes <5 .25 2c coc cc es ceatsusie suee oars nc scnaneiaeie yn Brn ive mrowsamwnn sak ecunavelsnasne casei bate peets 78 ON APACER. SCONE cso coors sa vacwe anes tacessssex sea sosde wns ceva wean asi eauaalvu ee eues Aiataddansteus4 Aakgeas ya aeeess emlowsabiieesnione 78 IDYINhal a) 011 C0) ole eecesne per ete accion enon atcha Gnd Atma LE Se Bene RES Rrne Aree RBNOE EC ERM R Se AAR OR ANa AOCET co OBCOCDR AC canbe CCE coe 83 OV 10 GOMEM A, ois xictats cr icat fat vances ea sseay dey nis maadoumreronan sie aa deo Gn rey necnce Ate cad wen aSas Donsuing wedcomanyapyoreetea egamwai’ 84 PARISI eae GASB aca a ser neta cm ck zy ca Nath cand Gahnicte nnieniaally ses ca nits Guia deiaslslesise Su MRiLCad Adonis Sabha Let sbad S7ae SeRaetehs 90 POSING, SPECT Oc PAIN DIES esd yo nactacdegesasn taint sie sei nalsetena slevlasteisaucoalinrndenmstendwaicgssetacreacoteetdepeacsanhek oi StH CHOTENICE 2 oicac toe needa ct oasataleniens ca vinte vet hat anys Mone eon MIAO EEA ACRcad Jondsay pave gEreIRs Meh Zaamed eho seta 93 DP LASVOSDETINUAE DAD ARCAS aaa axnasliccs anodised coos aerlnd avaneny sree tienen rangi sb isctder Weaephnsaesmns-enlobasa troneos on unite 94 BPP TEV ITS DEVIN NOSE «coe oh Siok waau sorbic ays tae ee i ovadcany em elo haMkd Ina Read asR FaHEe Ades tech ee cand eoed Sawenipe shana gua mE Ce 94 He PETE TD I NO Gare nis oak od, One aay po swertene 1 ve otea uses Sass ein Ease Ne oa sedate cede Alenia Sala ese ngs eons peed cay RARER 94 SGymnopentzia Benth in Benth .& HOOK. fo oo iccs2 5 es seadsvene vo sessd caasenv exes sp esceuys von sce apbvets eae geeseseose 95 GO; A yIMenOlepis CASS. ics xoceca ch» ahosniis sautansaee eae ihabe nde dovsri sa aputess th iubtedwetarmcuewcidesibteUingdubeNasiaodbsadeats 95 GENE TAA (KS 171 Wee RSPR Srna ape nao ORGO Ron Asad oE TOT IC OA RODR AD ACR CER ann oe col acne eh nance po aconuHen inane acu ocdosce 95 8. Asaemia (Harvey) Harvey ex Benth. in Benth. & Hook. f. ..............cceecceeceeeseeeeneeeeneeenseeeeeenseneceees 96 2. CasiChiininae: Breiner oe FLU DINGIC Set cio ri ti eteo cs atone sns endo’ dinates acoann tale aide saietialeus sie wwtandoiedy soeuepans nee 96 9). Trichantivemis. Regel Be Schitial ts vhs ce vee. cesar aes obs eden ease ese eden sudraah oaanes ema beige side sbedeuanutes ona 97 POE paartia: PAVIOV co tie trna Psst Suitaepasebeeaieruncenesennach marae sntete dm tepannrsunaeieaenins ics sneK eran aude cenvenaggnseuas stds 98 Tl PRichterial kare limi Gs Wilisersectcs sean cae ceo cans eereetnee ehes sec sos as Osenl gee tnt cas oaaseute aaaenereee ene ana ts 98 IIL WA Lars L172) DY S0s 11 aoa a Gee RAR anc eco eo cosenae au caGacee na deice todo bon tins ton oconadnsen oat ce cestode pacudoen Coonan Onn ch 98 US MCGNCPINIAACATCMVOG IRIE dane nc eisihadicros hantsncGeuaeds eels aahhas cgaiouaace iss snssiehd caagie dias cimen eat Riweas Vaneahaes woes 99 AAC BNCHIMICIIE. VZVOICM. eva siclacegiowsss enact xd es vatec sete DS D Dd dd DDD 96 A. elsiae Kallersj6 A. filiformis L. f. A. flexuosa Thunb. A. grandiceps Hilliard & Burtt A. hirsuta Thunb. A. humilis Kallersj6 A. imbricata Harvey A. inopinata (Hutch.) Kallersj6 (Stilpnophyton inopinatum Hutch.) A. juncea (DC.) D. Dietr. A. leptocephala KAllersj6 A. linifolia Burm. (Stilpnophyton linifolium (L. f.) Less., Stilpnophyton longifolium (Thunb.) Less.) A. microcephala DC. A. microphylla DC. A. minuta (L. f.) Kallersj6 A. oocephala (DC.) Kallersjé (Stilpnophyton oocephalum DC.) . pachycephala DC. . pectinata L. f. . pinnata L. f. . pubescens (L.) L. quinquedentata Thunb. . rugulosa E. Meyer ex DC. scabra Thunb. . sertulifera DC. . spathulata (DC.) D. Dietr. . tomentosa Thunb. . trifurcata (L.) L. . vestita (Thunb.) Druce . virgata Jacq. . viridis Kallersj6 paSDPdP.dS ppp pd. pddD 8. ASAEMIA (Harvey) Harvey ex Benth. in Benth. & Hook. f., Gen. pl. 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 (Kallersj6, 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 KaAllersj6 (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. (Ge Bh. SW Asia Asia Cancriniinae Trichanthemis Ugamia Richteria Allardia Cancrinia Cancriniella bi i i i i oa) * 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 9 Trichanthemis Sle 10 Ugamia tc 11 Richteria Cal Eee 12 Allardia 8 je 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, 0 = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. ahaa 1 1 1 11 a: 145057 44483375346 15775555566666175 532718 447983504981 30563567905678511 9. Trichanthemis11a11? 1alla0p00000 ????000000000000p 10. Ugamia al?a1? a11001100000 ?27777222227772727270 11. Richteria alaal? a10000010000 ????000000000000p 12. Allardia alaal? a10000011000 ????0000000000000 13. Cancrinia al?a1? a10000110000 2777727722922722727272?p 14. Cancriniella a1?a1? a10000100111 ?72727272227272772727272770 woody basally, generally with long peduncles. Ugamia has short peduncles. 44 Involucral bracts with dark brown margins. This 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 9) 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. bot. Sada 5: 617 (1877). Type species: T. karataviensis Regel & Schmalh. — Glossanthis Pol}. 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 7. aurea and T. radiata are related to the discoid T. paradoxos and T. karataviensis, respectively. He also indicated the possible relationship between the problematic Xylanthemum tianshanicum (Pyre- thrum tianshanicum) 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.|.), 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 dyjilgense 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, FI. 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 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. discoidea) 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, F/. 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. ‘TANACETINAEF’ Bremer & Humphries, subtrib. nov. Type species: Tanacetum vulgare 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 99 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, 1844a), 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. Tal 15 Tanacetum 16 Opisthopappus [ ie 17 Tanacetopsis TaZ 5 ie 18 Xylanthemum Ta3-— 19 Lepidolopha 20 Hippolytia ras 21 Heliocauta Fig. 4 Cladogram of the ‘Tanacetinae’ produced by the /e option in Hennig86. Cladogram length = 16, consistency index = 100, retention index = 100. Provisional group Tal — subtribe “Tanacetinae’ There is no autapomorphy for this group, but as explained above Tanacetum probably contains the sister groups of the other genera. Tanacetum 175 Embryo sac tetrasporic. The 10 Tanacetum species inves- tigated embryologically (Harling 1951; see discussion under Tanacetum) are all tetrasporic. The other genera of “Tanace- tinae’ have not been investigated. Opisthopappus 163 Pappus of separate, mainly abaxial, subulate scales. 100 K. BREMER AND C. J. HUMPHRIES Table 9 General distribution of Tanacetinae and genera. x=indigenous, o=introduced. C.& E. Asia SW Asia N.Am._ Eur- S.Eur. Asia Tanacetinae Tanacetum Opisthopappus Tanacetopsis Xylanthemum Lepidolopha Hippolytia Heliocauta i i oo N.Afr. S.Afr. Austr. S.Am. N.Zeal. x fe) fe) fe) x e) oO re) Table 10 Data matrix for the Tanacetinae. 1 = presence, 0 = absence, ? = missing data or not applicable, p = polymorphic but scored as the plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. | Ue Wes a a 1 1 1s is a ee | 1 eee | 1 Led SUONas Ie O" 8 G.0. Ooo a 4 ey AO”) Dlr Sao Sra OuOul 76 BO. 2k bee 36 S29 BIC 1 Sse 18428. 2 aS 6 03. 3 S60 O73 15 6. 60 4 426 18 8: 19:4 6.2 15. Tanacetum a liana 1 12 “1 0rp:0 p'0 0p.0 0 p 00.000. 2-70-00 '0:0.0.0 0:00:00 0:0" p pp: pip -0 pp p-p-p 16. -Opisthopappus a l.a-a.i? i? 1000-00000 0.00 0:0:0.2 7 2.0:0.0 0:00 0:00 00000 0:0 0:0:0:0 0.0-0'0 L7