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Bulletin of the Natural History Museum Bulletin of The Natural History Museum Botany Series THE NATURAL HISTORY MUSEUM VOLUME 32 NUMBER 2 28 NOVEMBER 2002 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: Prof. R. Bateman Editor of Bulletin: Mrs M.J. Stafford 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 before acceptance. SUBSCRIPTIONS Bulletin of the Natural History Museum, Botany Series (ISSN 0968-0446) is published twice a year (one volume per annum) in June and November. Volume 32 will appear in 2002. The 2002 subscription price (excluding VAT) of a volume, which includes print and electronic access, is 88.00 (US $155.00 in USA, Canada and Mexico). The electronic-only price available to institutional subscribers is 79.00 (US $140.00 in USA, Canada and Mexico). ORDERS Orders, which must be accompanied by payment, may be sent to any bookseller, subscription agent or direct to the publisher: Cambridge University Press, The Edinburgh Building, Shaftesbury Road, Cambridge CB2 2RU, UK; or in the USA, Canada and Mexico: Cambridge University Press, Journals Department, 40 West 20th Street, New York, NY 101 1-421 1, USA. EU subscribers (outside the UK) who are not registered for VAT should add VAT at their country's rate. VAT registered members should provide their VAT registration number. Japanese prices for institutions (including ASP delivery) are available from Kinokuniya Company Ltd, P.O. Box 55, Chitose, Tokyo 156, Japan. Prices include delivery by air. Postmaster: send address changes in USA, Canada and Mexico to: Bulletin ofthe Natural History Museum, Botany Series, Cambridge University Press, 1 10 Midland Avenue, Port Chester, New York, NY 105783-4930. Claims for missing issues should be made immediately on receipt of the subsequent issues. Orders and enquiries for issues prior to 2002 should be sent to: Intercept Ltd., P.O. Box 716, Andover, Hampshire SP10 1YG, Telephone: (01264) 334748, Fax: (01264) 334058, Email: [email protected], Internet: http://www.intercept.co.uk ADVERTISING Apply to the Publisher. Address enquiries to the Advertising Promoter. COPYRIGHT AND PERMISSIONS This journal is registered with the Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923, USA. Organizations in the USA who are also registered with CCC may therefore photocopy material (beyond the limits permitted by Section 107 and 108 of US Copyright law) subject to payment to CCC of the per-copy fee of $16.00. This consent does not extend to multiple copying for promotional or commercial purposes. Code 0968-0462/2002 $16.00. ISI Tear Sheet Service, 3501 Market Street, Philadelphia, PA 19104, USA, is authorised to supply single photocopies of separate articles for private use only. Organizations authorised by the UK Copyright Licensing Agency may also copy material subject to the usual conditions. For all other use, permission should be sought from Cambridge University Press. No part of this publication may otherwise be reproduced, stored or distributed by any means without permission in writing from Cambridge University Press, acting for the copyright holder. Copyright The Natural History Museum, 2002 ELECTRONIC ACCESS This journal is included in the Cambridge Journals Online service which can be found at: http://journals.cambridge.org For further information on other Press titles access http://uk.cambridge.org or http://us.cambridge.org World list abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) ISSN 0968-0446 The Natural History Museum Botany Series Cromwell Road Vol. 32, No. 2, pp. 61-156 London SW7 5BD Issued 28 November 2002 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nut. Hist. Mas. Land. (Bot.) 32(2): 61-123 Issued 28 November 2002 Studies in the genus Hypericum L. (Guttiferae) 4(2). Section 9. Hypericum sensu lato (part 2): subsection 1. Hypericum series 1. Hypericum NORMAN K.B. ROBSON 62 N.K.B. ROBSON H. ascyron subsp. ascyron [W. Siberia to Japan, China] Spp. 20-42 [Sakhalin to Borneo] 2-4. H. undulatum group [Macaronesia, Europe Mediterranean, W. Asia] ser. 2. Senanensia Spp. 13-19 [Japan, Korea, C. China] 1 . H. maculatum s.l. 12. H. scouleri [Euro-Siberia] [W. North America 5. H. perforation to C. Mexico] [Eurasia, Med.] 6. H. altenualum [C. Siberia,.China, Korea] l.H. elegans [C. Siberia to C. Europe] 8-1 1 . H. yezoense group [Japan, Sakhalin, S. Kuriles] Fig. 1 Section 9. Relationships and distribution. taxa of subsect. Erecta (H. yamamotoi var. riparium Y. Kimura and derivative of a cross between them, especially as H. perforatum H. ovalifolium Koidz.) have a branching woody stem base, like H. behaves cytologically as a hybrid (see p. 64). However, the nearest ascyron, but the stem internodes are slightly 2-lined or terete, the subspecies of H. maculatum to H. perforatum morphologically is lines being eglandular. Also, the leaves in the H. yamamotoi and H. not the one that occurs in Siberia today (subsp. maculatum) but erectum groups of subsect. Erecta are frequently triangular-lanceo- subsp. immaculatum, which is now confined to the Balkan region. late, like those of//, ascyron subsp. ascyron, whereas a comparable The hypothesis must therefore be that subsp. immaculatum at some form of leaf occurs only apomorphically in subsect. Hypericum, i.e. time was present in Siberia, where it hybridized with H. attenuatum. only in some of the more advanced species. The chromosomes of the resultant diploid hybrid then doubled, giving rise to an allotetraploid species that is partially fertile (see Distribution and evolution (Fig. 1) p. 88). Subsequently the area of//, maculatum subsp. immaculatum became greatly reduced; but that of subsp. maculatum, which is The distributions of subsect. 1 series 2 and subsect. 2 are wholly apomorphic to it and thus a likely derivative of it, extended eastward north and east Asian, but that of subsect. 1 series 1 extends from the into Siberia. Azores to Mexico. The latter series comprises a western group, based in south-eastern on 1. H. maculatum and Europe extending Sect. 9. Hypericum subsect. 1. Hypericum series 1. west to the Azores and east to central Siberia, and an eastern group Hypericum with two centres, one in eastern Siberia and the other in western North America. The North American centre consists only of 12. H. Characters and variation (Figs 2, 3) scouleri, which has its most primitive form in the south-eastern United States and shows trends northward to central British Colum- MORPHOLOGY. The two main divisions of series Hypericum could bia and southward to Michoacan in central Mexico. The basic be separated on whether or not the subsidiary stem lines were species of the east Asian centre is 6. H. attenuatum, which has its present, i.e. whether the stem internodes were 4- or 2-lined, were it most primitive form in eastern Siberia and a range that extends not for (a) 4. H. triquetrifolium, which has 'lost' them in the 4-lined westward to the Altai region and southward to south-central China. group and (b) 5. H. perforatum, which, as has been explained, The basic species of the derivative Japanese H. yezoense group (H. appears to be a hybrid with one parent from each group. Figure 3 tosaense) is morphologically very near H. attenuatum and has been shows that the 4-lined state is primitive in this series, as indeed it is included in it (Kitamura & Murati, 1962), but in my opinion it is in the whole genus. Intermediate states occur in hybrids of H. specifically distinct. Hypericum elegans, a western derivative of H. perforatum, but not in the species. attenuatum, extends the distribution of the eastern group as far as The laminar leaf glands are absent or more rarely sparse in l.H. western Germany. maculatum but more or less dense in the more advanced species. The remaining species in series 1,5. H. perforatum, does not fit They are all or mostly pale except in 6. H. attenuatum, in which easily into the above scheme. Morphologically it is intermediate black glands often predominate. between 1 . H. and 6. maculatum H. attenuatum, and the areas of The sepals are broad with a rounded apex in 1 . H. maculatum and these in species almost overlap western Siberia (Maps 1, 16). Each usually in 12. H. scouleri, as they are in the ancestral (?) H. ascyron of them is diploid (2n=16), whereas H. perforatum is tetraploid subsp. ascyron (sect. 7. Roscyna). In all the other species they are (2n=32); so it is conceivable that H. perforatum is an allotetraploid acute and narrow (Spp. 2, 3 and 5) or acute and broad (6. //. STUDIES IN HYPERICUM 63 3 a. H. tetrupterum var. anagallidifolium 2b . //. undulalum 1 1 . //. yezoense 3b. H. tetrapterum var. boeticum 12. //. scouleri var. corsicum 16 ~~~~~~~~~~~, 3. H. tetrapterum Za. //. undulatum var. tetraplerum var. undulatum 1 0. //. momoseanum 16 16,32 I 9. H. iwatelittorale 4. //. triquetrifolium la //. maculatum 16 .ssp. immaculatum 6. //. attenuatum -8. //. tosaense (16) \ 1 b. //. maculatum 1 c. //. maculatum 1. H. elegans ssp. maculatum ssp. obtusiusculum 5. //. perforatum 32 16 32 16,32 48 ~ f 5a.//. perforatum ssp. perforatum 5b. //. perforatum 32, 48 ssp. songaricum 5d.
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