Bulletin British Museum (Natural History) Botany Series VOLUME 20 NUMBER 1 28 JUNE 1990 The Bulletin of the British Museum ( Natural History ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. The Botany Series is edited in the Museum's Department of Botany Keeper of Botany: Mr J. F. M. Cannon Editor of Bulletin: Mr J. R. Laundon Assistant Editor: Dr A. J. Harrington 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. A volume contains about 256 pages, made up of two numbers: published Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Sales Department, Natural History Museum Publications, British Museum (Natural History), Cromwell Road, London SW7 5BD Telephone: 07 1-938-9386 Telex: 929437 NH PUBS G 07 1-938-8709 World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.) © British Museum (Natural History), 1990 ISBN 0 565 08026 1 Botany Series ISSN 0068 - 2292 Vol 20( 1 ), pp 1 - 1 5 1 British Museum (Natural History) Cromwell Road London SW7 5BD issued 28 June 1 990 Typeset by Waveney Typesetters, Norwich, Norfolk. Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. Br. Mus. nat. Hist. (Bot.) 20(1): 1-151 Issued 28 June 1990 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys* 17.1111 199 NORMAN K. B. ROBSOJV / fV do Department of Botany, British Museum (Natural History), Cromwell Road, London SW7 5BD PRESENTEC GENERAL CONTENTS Synopsis ................... 1 Introduction .................. 2 Separation of sects Brathys and Trigynobrathys ............ 2 Sect. 29. Brathys 3 (a) Characters ................. 4 (b) Distribution and evolution 4 Sect. 30. Trigynobrathys and its subdivision ............ 5 Morphology of sect. 30. Trygynobrathys ............ 6 (a) Leaves 6 (b) Inflorescences 6 (c) Flowers and fruits ................ 7 Cytology and hybrids in sect. 30. Trigynobrathys ........... 8 Distribution and evolution in sect. 30. Trigynobrathys .......... 8 (a) Principal clades (i) and (ii): relatives of H. rigidum subsp. sellowianum ................ 8 (b) Principal clade (iii): H. brasiliense, H. campestre, and relatives .................. 9 (c) Principal clade (iv): subsect. Knifa ............. 10 (d) Wide disjunctions: discussion 11 Systematic treatment 12 Sect. 29. Brathys (Mut\se\L.f.)Cho\sy 12 Sect. 30. Trigynobrathys (Y. Kimura) N. Robson 47 References 146 Systematic index 148 SYNOPSIS. New collections from Colombia (J. R. I. Wood) and Ecuador (Flora of Ecuador collectors) have necessitated a reorganization of Hypericum sect. 29. Brathys. The mainly woody species, described in Part 7 (Robson, 1987), have increased in number from 61 to 75 by the addition of 11 new species (one as yet unnamed) and the transfer of three species provisionally allocated to sect. Knifa (Robson, 1987). The remainder of this section has been reduced to subsectional status under sect. 30. Trigynobrathys. The treatment of sect. Brathys is completed by accounts of 13 mainly herbaceous species not included in Part 7, two of which are new. A new enumeration of the 88 species of sect. Brathys and a revised key are provided, and the section is divided into four subsections: 1. subsect. Styphelioides N. Robson, subsect. nov., 2. subsect. Phellotes N. Robson, subsect. nov., 3. subsect. Brathys, 4. subsect. Spachium R. Keller. Sect. 30 Trigynobrathys is divided into two subsections: subsect. Connatum (R. Keller) N. Robson, comb. nov. and subsect. Knifa (Adanson) N. Robson, stat.nov., and a systematic account is given of the 52 species. A discussion of the morphology, chromosome numbers, hybrids, distribution, and evolution of each section is included. The following other new taxa (sp. nov. or subsp. nov.) are described in sect. 29. Brathys: Hypericum phellos subsp. marcescens N. Robson (Colombia: Santander), H. matangense N. Robson (Ecuador: Morona-Santiago), H. callacallanum N. Robson (Peru: Amazonas), H. gladiatum N. Robson (Colombia: Boyaca), H. espinaliiN. Robson (Colombia: Cauca), H. castellanoi N. Robson (Colombia: Boyaca, Venezuela: Merida), H. asplundii N. Robson (Ecuador: Pichincha), H. radicans (Columbia: Boyaca), H. paramitanum N. Robson (Venezuela: Trujillo), H. woodianum N. Robson (Colombia: Cundinamarca), H. harlingii N. Robson (Ecuador: Loja), H. rubritinctum N. Robson (Mexico: Guerrero, Mexico), H. beamanii N. Robson (Guatemala: Solola); in sect. 30. Trigynobrathys: H. cordatum subsp. kleinii N. Robson (Brazil: Parana, Sta Catarina), H. pedersenii N. Robson (Brazil: R. G. do Sul), H. campestre subsp. pauciflorum N. Robson (Brazil: Parana to R. G. do Sul, Argentina: Chaco), H. campestre subsp. tenue N. Robson (Paraguay, Argentina: Misiones), H. salvadorense N. Robson (Brazil: R. G. do Sul), H. silenoides subsp. minus N. Robson (Galapagos Is), H. relictum N. Robson (Colombia: Santander), H. killipii N. Robson (Colombia: Santander, Norte de Santander). 'For part 1 see Robson (1977); for part 2 see Robson (1981); for part 3 see Robson (1985); for part 7 see Robson (1987). N. K. B. ROBSON The following changes of rank (stat. nov.) are also made: sect. 29. Brathys - Hyperlcum chamaemyrtus subsp. pseudocaracasanum (Steyerm.) N. Robson (Venezuela: Lara, Trujillo); sect. 30. Trigynobrathys - H. rigidum subsp. meridionale (Lyman B. Smith) N. Robson (Brazil: Sao Paulo to Sta Catarina), H. rigidum subsp. sellowianum (R. Keller) N. Robson (Brazil: Minas Geraes to R. G. do Sul), H. denticulatum subsp. acutifolium (Elliott) N. Robson (south-eastern U.S.A.), H. myrianthum subsp. tamariscinum (Cham. & Schlechtendal) N. Robson (Brazil: R. G. do Sul, Uruguay), H. mutilum subsp. latisepalum (Fern.) N. Robson (south-eastern U.S.A.), H. pumillum subsp. diffusum (Rose) N. Robson (north-central Mexico). One is also a new name (stat. et nom. nov.): H. rigidum subsp. bracteatum N. Robson. One new combination (comb, nov.) is made: H. cordatum (Veil.) N. Robson. INTRODUCTION Separation of sects Brathys and Trigynobrathys One of the main problems in revising the sectional classifica- tion of Hypericum L. has been the recognition of the boundary between Brathys and 'Spachium'. Keller (1893) ranked these taxa as subsections, differentiating them respec- tively as shrubs or subshrubs (subsect. Eubrathys) and herbs (subsect. Spachium). He distinguished two further groups, subsects Connatum (subshrubs with connate leaves - H. connatum Lam.) and Multistamineum (herbs with spike-like inflorescence and numerous stamens - H. angulosum Michaux = H. denticulatum subsp. acutifolium and H. pilosum Michaux = H. setosum), for species that did not fall easily into the major shrub/herb categories. When revising his treatment for the second edition of Die naturlichen Pflanzen- familien (Keller, 1925), he merged these last two subsections with subsect. Spachium and elaborated his differentia: Subsect. Eubrathys. Subshrubs or small shrubs with often dense appressed foliage; flowers often terminal, usually relatively large, i.e. about 1-5 cm in diameter, rarely less. Subsect. Spachium. Annual or perennial herbs; stamens 5- 50, rarely more. Subsect. Eubrathys in Keller's revised treatment still contained two species that are now allocated to other sections (H. nitidum Lam. in sect. 20. Myriandra and H. epigeium R. Keller in sect. 9. Hypericum), as well as several that should have been in his subsect. Spachium. This, in turn, contains some species that would now be allocated to sects 9. Hypericum and 26. Humifusoideum, although most do belong to 'Spachium' . In the preliminary key to sections in Part 1 (Robson, 1977: 342), I still maintained the 'mainly shrubby/mainly herba- ceous' distinction, supplementing it by characters of inflor- escence, leaf margin, and stem internodes and raising Brathys and Spachium to sectional rank. Further work, however, showed this view to be untenable, because (i) sect. Brathys was found to contain herbaceous species, including the type of sect. Spachium (H. gentianoides (L.) Britton, Sterns & Pogg.) and (ii) the subsidiary characters that were used are not completely diagnostic. By combining leaf-insertion and leaf-venation characters with type of inflorescence branching, it seemed possible to extract, partly from each section, a group with long narrow leaves and pinnate or single veins. This was described in sect. Knifa (Adans.) N. Robson in Part 7 (Robson, 1987) and the correct name for sect. 'Spachium' excluding H. gentianoides and its relatives was shown to be sect. Trigynobrathys (Y. Kimura) N. Robson. Scarcely had Part 7 gone to press, however, when a study of new material from Colombia (J. R. I. Wood) and Ecuador (Swedish and Danish collectors from GB and AAU) showed that sect. Knifa in turn was untenable, as it consisted of a shrubby element belonging to sect. Brathys and a herbaceous element belonging to sect. Trigynobrathys. Even then, the dividing line between these two sections was not completely defined and has remained 'fluid' throughout most of the time during which Part 8 has been written. I am now satisfied, however, that all species have been allocated to the appropri- ate section, and the distinctions between the sections have become clear, though still not easy to describe. Sect. Brathys comprises 88 species of shrubs (rarely small trees), subshrubs, shrublets or wiry annuals (rarely peren- nials) with short stem-internodes, leaves plane to completely revolute or completely involute and often united in pairs but not perfoliate, flowers solitary with inflorescence-branching pseudo-dichotomous or (especially in the less shrubby spe- cies) dichasial/monochasial or mixed, stamen fascicles almost never distinguishable, and styles usually 3 but plesiomorphi- cally 5(4) and sometimes apomorphically (4)5. It has two primary centres: (i) two closely related species in Belize and Cuba and (ii) the Venezuela-Colombia border (see Part 7: 7). From the latter area the mainly shrubby part has radiated eastward to Roraima, westward to Costa Rica, and south- westward along the Andes to Bolivia, with a disjunct species (Hpiriai) in south-east Brazil and Uruguay. The subshrub-to- annual part has its primitive species in northern Colombia and north-west Venezuela and a secondary centre in southern Mexico. Thence it has spread south-eastward to Panama, eastward to the Greater Antilles, north-westward to southern Baja California and the Revilla Gigedo Islands and, with a disjunction, north-eastward to the eastern U.S.A. and south- eastern Canada. The 88 species can now be seen to fall into four distinct groups, which are described as subsections. Sect. Trigynobrathys comprises 52 species of shrubs, subshrubs, shrublets, perennial and annual herbs with stem internodes short to elongate, leaves plane to recurved or very rarely incurved and free or perfoliate, flowers solitary or more usually in dichasial/monochasial cymes or with bran- ching mixed cymose/pseudo-dichotomous (rarely wholly pseudo-dichotomous), and styles 5-2 with two independent reversals from 3 to 5. The primary area of speciation here has been south-eastern Brazil, with a primary radiation (to the south of the Amazon) northward to eastern Brazil (Bahia), and (with major disjunction) to south-eastern U.S.A., south- ward to Uruguay and northern Argentina, westward to Bolivia, Peru, and northern Chile and thence north along the Andes to southern Colombia and the Galapagos Islands, and west (with disjunction) to New Zealand, Australia, New Caledonia, and New Guinea and (with further disjunction(s)) to scattered localities in south-east Asia. From a secondary centre in north-east Colombia it has spread into eastern Costa HYPER/CUM Sect. 29 Brathys Relationships wurdackii castellanoi19 \ 20 asplundii | ' I .' / partly,'not|whol'ly/ I I ,' I 3 ;4| 5 styles -*v*"^ -- O / ~. ipseudopetiolate---^ _ — QQ leaf [base i sessile -86/ Fig. 2 Sect. 29 Brathys. Limits of certain characters within the section and related sections. Note the isolated apomorphic occurrences of 4-5- styled species. Revised and expanded from Part 7. sister-group of subsect. Brathys alone rather than of subsects Phellotes + Brathys. (a) Characters The revised character diagram (Fig. 2) shows the limits of the same characters as were depicted in Part 7: 6, fig. 2. Apart from the extensions to subsect. Spachium, these diagrams are basically similar. No more trend reversals (e.g. in Spp. 54-55, 67, and 74) have been revealed, except possibly the partly deciduous leaves in Spp. 4 and 5. Indeed, what appeared to be a reversal to wholly persistent leaves in Spp. 9-10 is now seen not to be one; instead the partly deciduous leaves of Sp. 8 are shown to be a specialization. One additional character in the diagram (leaves free v. united) nicely delimits subsect. Brathys from subsects Styphelioides and Spachium, but sub- sect. Phellotes contains species with both conditions. Finally, the pseudopetiolate leaf base in subsect. Spachium (Spp. 75- 76) is seen to have arisen independently from that in subsect. Brathys. (b) Distribution and evolution (Fig. 3) The six morphological group-headings (a-f) under which the distribution and evolution of sect. Brathys were considered in Part 7: 7-12 have been reduced to three subsections, and a fourth (subsect. Spachium) has been added. Subsect. 1. Styphelioides is confined to Belize and Cuba. Subsect. 2. Phellotes is based (morphologically and geographically) in the Colombia/Venezuela border area and extends eastward to Guyana (Roraima), northward to Costa Rica and adjacent Panama, and south-westward along the Andes to northern Peru and, with disjunction, south-eastern Brazil and Uruguay. Subsect. 3. Brathys has the same centre as subsect. 2 and a similar distribution; it occurs also in Hispaniola and extends along the Andes as far south as Bolivia, but it does not occur in south Brazil. It is present as well in Roraima (but only the Venezuelan part) and also in southern Venezuela (Cerro de Marahuaca). Subsect. Spachium, too, radiates from north-eastern Col- ombia (75. H. cymobrathys), but mainly northward. Apart from 76. H. chamaemyrtus , with a Colombian and a Vene- zuelan subspecies, all the other species are natives of Central and North America and the West Indies. Disjunctions between northern South America (76. H. chamaemyrtus) on the one hand, and southern Mexico (77. H. arbuscula) and Honduras (primitive form of 82. H. gnidioides) on the other, suggest a northward migration through the early, Greater Antilles land-link between the two Americas (Part 7: 9), with subsequent migration and differentiation from the Belize/ HYPER/CUM Sect. 29 Brathys C?£W) ?21CO(nc) E(s)-PE(n) •V. 27 20 ^ C0(c) 25 26 24 _J c°(nc) CO (c) Distribution CO(nc)-V(w) 59 C0(he) E(nh CR'-P'A cocsw}- E(n>PE(n)43 PE(n)42 "w 53 58co(r'<0-v(w) c o(wc) N / c ° (ny 5 7 b v W-c °(n e) 67HA CO(c)-VQw) C0(n)-V(nw) E (s) C0(ne)-V(w) 5 CO(nne)\Co(-ne)V(.v 4 +- 3c— 3b=3a CR-PA(w) CO(ne) 37 40 co PE(n)x (ne - nc) 36 CO(nc) -V(w) 72 PE(s) BO DO V/ n 7 n CO(ne-nw)XD /U 7 1 E^ +CR , 64 69 I >/ ! »^ PE(n-s) CO(ne-nc) 44 — 62+CR C0(nc-c) C0(ne-nc) "68— 7 o _ -7 A COCc)-E(c) 'O ^ '.O E(n)-V(w) 81 ', HAjPRjcU 2c 76a ^30| C0(nc-c)\ USA(e),CAN(se) Fig. 3 Sect. 29 Brathys. Distribution of the 88 species showing major and minor disjunctions (= across sea, across land (major). across land (minor)). Lower-case letters indicate compass points. Revised and expanded from Part 7. For geographical abbreviations see p. 9. Guatemala region. The more primitive species, with longer styles (77. H. arbuscula), contains an indivisible morphocline from southern Mexico (Chiapas) to Guatemala, whence there is a disjunction in this clade to Hispaniola (79. H. fuertesii, 80. H. dichotomum, and 81. H. diosmoides). These species form an increasingly herbaceous and procumbent series, and the most specialized one, H. diosmoides, has extended its area into Puerto Rica and Cuba. Whether the disjunction between Guatemala and Hispaniola is a result of overland or long- distance migration is unclear. The latter seems more likely in view of the relatively specialized form of these species. Northward, //. arbuscula or its predecessor has given rise to 78. H. rubritinctum, which occurs in central Mexico (Guer- rero, Mexico). The initially more southern branch of the Central Amer- ican radiation starts with 82. H. gnidioides, which is very variable in Honduras and, to a lesser extent, Nicaragua, and shows a south-eastward morphocline to western Panama. Here there occurs an extremely reduced prostrate form ('H. woodsonii'), which is linked to the more typical one by a continuous series of intermediates. North-west from Hon- duras the nearest relatives of H. gnidioides are (a) in west- central Mexico and the Revilla Gigedo Islands (83. H. eastwoodianum) with a derivative species in the extreme south of the Baja California peninsula (84. H. peninsulare) and (b) in east-central Mexico (86. H. galinum) with a sister- species in the mountains of Guatemala (85. H. beamanii). Finally, 87. H. drummondii , from south-eastern U.S.A., is morphologically very close to the least shrubby (annual?) Honduran form of 82. H. gnidioides; and 88. H. gentianoides (eastern U.S.A. and extreme southern Canada) is more reduced than H. drummondii. There is no evidence, in the form of relict species (etc.), that H. drummondii has reached the southern United States overland. Indeed, its primitive form is so close morphologically to H. gnidioides that such a long migration without greater differentiation seems unlikely. I am therefore inclined to attribute the disjunction to ancient long-distance migration, possibly the result of carriage of propagules by extra-strong winds or birds. Sect. 30. Trigynobrathys and its subdivision As here delimited, sect. Trigynobrathys comprises 52 species that vary widely in habit from shrubs 2 m tall to prostrate annual herbs (Fig. 4). It can be divided into two subdistinct groups: 1) Shrubs, subshrubs or woody annuals; styles 5(4-3); capsule usually broadly ovoid to globose; mainly south of the Amazon, also Chile, the Andes from Peru to Colombia, N. K. B. ROBSON Sect. 30 Trigynobrathys Relationships 22b ssp. tamariscinum 20 salvadorense 17 polyanthemum anceps 24 25 b ssp. minus 26 caespitosum 35 aphyllum 25a ssp. silenoides 22a ssp myrianthum gramineum 16, 14 31 brevistylum lorentzianum 21 16 brasiliense 24? 30, thesiifolium ssp. campestre 18b 32 paucif lorurr 18a ssp.pauciflorum ssp.acutifolium 24 10b \ •* \ gymnanthum38 re denticulatum 13 cumulicola 10a ssp. denticulatum 24,48 42 japonicum re ssp.diff usum 43a pumillum 43b ssp. pumillum ssp. bracteatum 1 d 1b SSP meridionale ssp. sellowianum 47 anagalloides re 49 globulif erum microlicioides Brathysk 29 la ssp. rigidum scioanum re Knifa legrandii 9 6b ssp.kleinii cavernicola connatum caprifoliatum Campylosporus I * "* arenarioides 40 37 / canadense ssp./ 16 latisepalum 39b 3ga mutilum ssp.mutilum re ^^ 39c ssp. boreale ref is) pleiostylum 45 paucifolium moranense oligandrum Fig. 4 Sect. 30 Trigynobrathys. Relationships of the 52 species and related sections, showing known diploid chromosome numbers. Limits of (named) subsections indicated by bold line. Subspecies linked by paired lines. (a) Leaves Galapagos Is., eastern N. America, and Australasia north to Taiwan, Vietnam, and Bhutan ('Trigynobrathys proper'). Spp. 1-27. 2) Subshrubs, shrublets or herbaceous annuals; styles (2) 3(4- 8); capsule usually narrowly ovoid to ellipsoid or cylindric; mainly north of the Amazon, Andes from Venezuela south to Argentina, Central and North America and Greater Antilles, also Africa, Madagascar, and east Asia, south Australia, and New Zealand ('Knifa'). Spp. 28-52. The presence of 25. H. silenoides, 26. H. caespitosum, and 27. H. gramineum in group 1) and of various other exceptions to the 'broad 5-styled ovary v. narrow 3-styled ovary' division makes it difficult to define Knifa as a subsection. An undivided section of 52 species, however, is rather unwieldy and so an attempt has been made to do so as the subsections in practice are usually easily distinguishable (see pp. 51 and 95). Morphology of sect. 30. Trigynobrathys (Fig. 5) The species in this section vary from tall shrubs (up to 2 m) to erect or prostrate herbaceous annuals. They all lack dark glands completely and have no resin glands on the lower surface of the leaf; and there are no fringing glands on leaves, sepals or petals. All species have marginal punctiform leaf glands (often very small). The laminar glands are usually also wholly punctiform, but very rarely lines of punctate glands may coalesce to produce streaks (30. H. thesiifolium in part). As in sect. 29. Brathys, the leaves in part of the species with the most primitive characters (1. H. rigidum) have parallel and open venation; but even within this species, forms with partially pinnate venation occur. Pinnation occurs in two ways: i) by union of the laterals with the midrib, the outer ones nearest the base, with the venation remaining apparently open (e.g. in 6. H. cordatum, Fig. 13) or becoming closed (e.g. in 4. H. caprifoliatum). ii) by development of secondary laterals along the midrib as the space between it and the primary laterals increases (e.g. in the most primitive form of 16. H. brasiliense). Method i) is often combined with method ii), as in 18. H. campestre, Fig. 15 A-C. Other primary laterals may produce secondaries on the marginal side, and the system may become close (e.g. in 25. H. silenoides, Fig. 17 A-B). In the more shrubby species with free leaf-bases, the leaves are sooner or later deciduous (e.g. in 14. H. denudatum); but where the leaves are herbaceous or the bases united, they persist. In no species are they partially deciduous, as they are in many species of sect. Brathys. Again, in contrast to much of the sect. Brathys, the lamina is usually plane to revolute, rarely involute or conduplicate (e.g. in 9. H. legrandii or 13. H. cumulicola). (b) Inflorescences Unlike the species of sect. Brathys, where the 1 -flowered HYPERICUM Sect. 30 Trigynobrathys Characters 35 39c perennial or """•""•• 5K3(D shrub\subshrub \ shrublet .--'"'annual Fig. 5 Sect. 30 Trigynobrathys. Limits of certain characters within the section and related sections. Note the isolated apomorphic occurrences of 4-5-styled species. inflorescence with pseudo-dichotomous branching is com- moner than the dichasium/monochasium, those of sect. Trigynobrathys more frequently have dichasial/monochasial branching, which can become elaborate (e.g. to the sixth grade in 22. H. myrianthum, Fig. 16 B-C). Pseudo- dichotomous branching, however, is far from absent (e.g. in 50. H. humbertii, Fig. 22 A), and mixed inflorescences with both types of elaboration are of frequent occurrence in subsect. Knifa. In some advanced species of this subsection, only one branch of the pseudo-dichotomy normally develops, resulting in a sympodium (e.g. in 42. H. japonicum variant 5. 'humifusum'). Lateral inflorescences are usually present as well as the terminal one, but they sometimes are slow to develop (e.g. in 27. H. gramineum, Fig. 17 E). Variants of 1. H. rigidum, the species most similar to those of the African sect. 1. Campy losporus, already display most combinations of these types of inflorescence elaboration: 1- flowered with branching wholly lateral (Fig. 12 C), 1-3- flowered with branching wholly lateral (Fig. 12 B), 1-7- flowered with branching at first pseudo-dichotomous then lateral (Fig. 12 A). (c) Flowers and fruits As is usual in sect. Brathys, the perianth in sect. Trigyno- brathys is pentamerous with the sepals appressed to the petals. These are usually longer than the sepals and are always spreading ('flowers stellate'). The sepals vary with the relative width of the base from 1-9-nerved, the nerves often becoming (more) prominent in fruit. There are trends in the petals from presence to absence of an apiculus and pellucid glands, but these are of no more than specific taxonomic value. As in sect. Brathys, the petals and stamens are persistent, the petals rolling downward after anthesis and sometimes enclosing the developed capsule. The number and size of the stamens are correlated in general with flower size as a whole. In the larger flowers the five fascicles are sometimes distinct at the base (e.g. in 16. //. brasiliense; c.f. Reichardt, 1878: fig. 34), but they usually form a narrow complete whorl. In the smallest flowers (e.g. in 37. H. canadense or 51. H. scioanum) the reduced number of stamens separates again into groups, normally three but sometimes five or an irregular grouping that does not correspond to that of the original fascicles. The styles vary from 5 to 3 or rarely 2 (e.g. in forms of 42. H. japonicum). The intermediate number 4, while never uniquely characteristic of a species, is possibly commoner in sect. Trigynobrathys subsect. Connatum than in most other sections of Hypericum. li recurs in several species, nearly all in subsect. Knifa, which are normally 3-styled. As was stated above, however, there are two species (41. H. pleiostylum, 52. H. oligandrum) in which this trend is further reversed, the number of styles increasing in H. oligandrum to 4-5 and in H. N. K. B. ROBSON pleiostylum up to 8. The form and behaviour of the styles do not appear to have taxonomic value, although there is some variation: most curve outward, but some later curve inward again. The form of the stigmas, however, can sometimes be utilized; it varies from not or scarcely enlarged (rare) to very broadly capitate or even peltate, and the style immediately below is often gradually enlarged. This gradual enlargement may occur below a narrow, even rounded, apex (stigma clavate), a state characteristic of the H. lalandii group (Spp. 48-52). Cytology and hybrids in sect. 30. Trigynobrathys The most primitive chromosome number in the genus, n = 12, occurs in subsect. Connatum, in 10. H. denticulatum , 11. H. harperi, and possibly 16. H. brasiliense, and the only polyploid number so far recorded (n = 24) is found in the H. denticulatum group where H. denticulatum subsp. denticula- tum is sometimes diploid, sometimes (auto-?) tetraploid. The two more advanced members of this group, however, have n = 6. This number cannot (as Webb (1980) averred) here represent the basic number of the whole genus, thus making n = 12 tetraploid. The species with this number, 12. H. setosum and 13. H. cumulicola, are clearly derived from 10. H. denti- culatum (n = 12, 24) and, for whatever reason, must be regarded as haploids unless (which seems more unlikely) the whole complement is still present with the chromosomes united in pairs. The numbers n — 11-9 have not yet been recorded in sect. Trigynobrathys except for n = 9 in 39c. H. mutilum subsp. boreale (Kapoor, 1972), where it represents a secondary increase in a taxon that normally has n = 8. Otherwise all chromosome counts of species in subsect. Knifa and else- where in subsect. Connata have yielded n = 8 or 2n = 16, the sole exception being a New Guinea plant of H. gramineum (n = 7) (Borgmann, 1964, as Hypericum sp.). Whether this gap in numbers is real or, more likely, merely the result of insufficient work, remains to be seen. Hybrids have been found only in the H. canadense group (Spp. 36-42), all the North American (i.e. excluding Mex- ican) species of which are apparently interfertile. Webb (1980) reported that hybrids and putative hybrids in this group showed a lower chromosome stainability than did species. From field observations and herbarium studies, he deduced that the name H. dissimulatum Bickn. has been applied to recurrent hybrids between 37. H. canadense and two subspecies of 39. H. mutilum (subsp. mutilum and subsp. boreale} . The only other suspected hybrid that has been reported is from Bhutan, where the distributions of 27. H. gramineum and 42. H. japonicum variant 4c ('calyculatum') overlap (Robson, 1972: 267). Distribution and evolution in sect. 30. Trigynobrathys (Fig. 6) Whereas sect. Brathys is centred north of the Amazon, in the north-eastern Andes and Greater Antilles, sect. Trigynob- rathys has its centre in southern Brazil. The evolutionary divergence pattern of Hypericum as a whole suggests that the split between them occurred early, probably in Africa before the progenitors of either section reached South America (Robson, 1987: 7). The transference of the shrubby species from (subsect.) Knifa to sect. Brathys makes it seem even less likely that there was a later sectional divergence across the Amazon basin. Indeed, the disjunction of subsect. Knifa from subsect. Connatum, which must of course have been later, involves that very region. (a) Principal clades (i) and (ii): relatives o/H. rigidum subsp. sellowianum The most primitive species in sect. Trigynobrathys (i.e. the one that is morphologically nearest to the east African H. revolutum subsp. keniense} is 1. H. rigidum, of which the most primitive subspecies (subsp. a. rigidum) is confined to Parana in southern Brazil and has a closely related derivative species, 2. H. microlicioides, in the same province and also in Santa Catarina. All the other parts of the section are related to other subspecies of H. rigidum. Subsp. b. meridionale, with a somewhat wider distribution (Sao Paulo to Santa Catarina), has apparently given rise to two further subspecies: c. subsp. sellowianum, which has the widest distribution (southern Minas Geraes to Rio Grande do Sul), and d. subsp. bracteatum, which has a relict distribution in Parana. Of the two major clades related to subsp. sellowianum, one, clade (i), is confined to south-eastern Brazil and adjacent regions of Paraguay, Argentina, and Uruguay. In it the tendency to shorten and broaden the leaves in that subspecies is at first elaborated, so that the leaves are very broad in 3. H. teretiusculum and the pairs connate in 4. H. caprifoliatum and 5. H. connatum. In the other derivatives of H. teretiusculum they become smaller and narrower, and this trend finishes in 9. H. legrandii (Uruguay), where they are linear-incurved. In the related 8. H. ternum- they are sometimes in whorls of three. The other major clade related to subsp. sellowianum (ii) begins morphologically not far from H. teretiusculum, but geographically its members are very distant - in south-eastern U.S.A. The affinities of the H. denticulatum (Spp. 10-13) are clearly with the H. teretiusculum group (e.g. they have similar leaves and inflorescence-branching), but both groups seem to be related independently to H. rigidum subsp. sellowianum. Although the American species are woody herbs rather than subshrubs, the morphological differences are not great, which suggests that their ancestor reached the U.S.A. by long- distance dispersal (not overland), from south of Amazonia. H. rigidum and H. teretiusculum are plants of moist habitats, and it is not impossible to imagine that seeds of these or of a related species were transported externally (?) by birds on a northward migration. The suggestion of a long-distance origin of the H. denticulatum group invites a comparison with relatively few examples - 'a remarkable group of 6 species disjunct between the eastern United States and temperate South America' (Raven, 1963). These include Hypericum gentianoides (sect. Brathys}, which must have travelled in the opposite direction more recently, being one of the most highly evolved species in that section and not morphologically distinct in South America (c.f. Smith, 1962; Thorne, 1972). Smith (1962) regards Hypericum as one 'of the best examples ... of a wide-ranging Andean element in southern Brazil'; but this scenario applies only to the H. carinatumlH. silenoides and (possibly) H. relictum groups (Spp. 23-26, 28- 29), the relationships of the other southern Brazilian species being more varied. Examples of birds that regularly migrate from southern Brazil to the south-eastern United States and back have not HYPER1CUM Sect. 30 Trigynobrathys Distribution BR(s)20 BR(s)17 UR BR(s)19 UR 26 CH(c) 27 NZ.AUS.NC, HW, ASIA(e-se) 11 USA(se). 9 UR 23 BR(s),UR A R (h e - e , n w) PY.AR(ne) 18c 7 ' UR BR(se-s)16 PY,BO,AR(ne,nw; 31 CO(sw)-BO / AR(nw) 30/ CO,V,PA,CR ME(se)-HO 34 33 ME(s)-NI(n) BE 35 USA(rO,CAN(s) [GE,F, J J 36 40 37 USA(e),CAN(e) [IR, NE.F] NW 39b ^ USA(se) USA(se)38 393 [GTPOjj^ USA(e),CAN(ef [AZ,EUR,SU(sw), NZ.HW.ME.HO, D,CO,E,PE,PY, BR(se)V] USA(ne) 41 BR (se) 5 BR(s),UR PY,AR(n) CAN (e) T, / ME(nw) / USA(w) / CAN(sw) 48 AFR(nw,ne-se,3w) AFR(e) MA APR (ne-e) 44 ME(c-ne; USA (s) 52 AFR(sw) Fig. 6 Sect. 30 Trigynobrathys. Distribution of the 52 species showing major and minor disjunctions (= across sea, — across land). Paired relationship lines indicate subspecies. Lower-case letters indicate compass points. Introductions (ancient or recent) within square brackets. Geographical abbreviations used in Figs 3 and 6 APR AR ASIA AUS AZ BE BO BR CAN CH Africa Argentina Asia Australia Azores Belize Bolivia Brazil Canada Chile CO Colombia CR Costa Rica CU Cuba D Dominican Republic E Ecuador EUR Europe F France GA Galapagos Islands GE Germany GT Guatemala GY Guyana HA Haiti HO Honduras HW Hawaii IR Ireland J Japan MA Madagascar ME Mexico NC New Caledonia NE Netherlands NI Nicaragua NZ New Zealand PA Panama PE Peru PR Porto Rico PY Paraguay SU Soviet Union UR Uruguay USA United States V Venezuela yet been found;2 but I have little doubt that such migrations occur. At any rate, the bird(s) that hypothetically transported H. gentianoides in relatively recent times would be unlikely to belong to the same species as the ones that were responsible for the ancient conveyance of the H. denticulatum ancestor. Raven (1963) states that all the temperate N. America/S. America disjuncts date from the late Mesozoic or later, and no more accurate dating seems to be possible yet. (b) Principal clade (Hi): H. brasiliense, H. campestre, and relatives The other two principal clades in sect. Trigynobrathys both 2But see Cruden (1966) for a detailed discussion of bird migrants as vectors between California and Chile. derive from Id. H. rigidum subsp. bracteatum, which is represented as yet only by the type specimen, from southern Brazil (Sao Paulo). Despite its apparent rarity, this sub- species forms a morphological link between the basic species, H. rigidum, and the two remaining principle clades. Of these, (iii) is distributed largely to the south of Amazonia and across the southern Pacific Ocean, whereas clade (iv) (subsect. Knifa) is centred to the north of Amazonia and across the northern Pacific Ocean and the Atlantic Ocean (northern and southern). Clade (iii) divides into the H. brasiliense group (clade (v), Spp. 14-17) and the H. campestre group (clade (vi), Spp. 18- 27). Clade (v) is confined to south Brazil, Uruguay, and adjacent Argentina, except for 16. H. brasiliense itself, which extends from Parana northward along the eastern hills, Serra 10 N. K. B. ROBSON da Mantiqueira and S. do Espinhac,o, to Bahia and westward to Paraguay, southern Bolivia, and northern Argentina and possibly into Peru (see p. 87). Clade (vi), based on the variable 18. H. campestre, is also largely restricted to southern Brazil, Uruguay, and adjacent Argentina, but 18c. H. campestre subsp. tenue occurs in Paraguay and north- eastern Argentina. 23. H. carinatum is more widespread, extending south-westward (in Argentina) to Buenos Aires and Cordoba and westward to Paraguay. In Paraguay it is represented by two forms, one tall and more shrubby ('//. altissimutri , H. megapotamicum') and the other short and more herbaceous, even possibly annual ('//. paraguense'). There are intermediate states elsewhere in the range of the species that preclude taxonomic recognition of these forms, but '//. paraguense' links typical H. carinatum with the Andean and trans-Pacific herbaceous species. The most primitive form of 25. H. silenoides is in northern coastal Chile ('//. paposum'}, whence there is a northward morphocline along the coastal foothills to Lima. There is a gap in distribution, though scarcely in variation, between there and the Ecuador/Peru border, where a secondary, higher-altitude, cline originates. This extends northward to Colombia (Antioquia) and southward to Bolivia and north- western Argentina. 26. H. caespitosum, from central Chile, is most similar to to some central Peruvian (Ayacucho) forms of this cline, whereas the endemic Galapagos Is. 25b. H. silenoides subsp. minus is morphologically closest to plants of the type subspecies from southern Ecuador. Like (all?) the other Galapagos biota, therefore, it seems to have reached the islands across ocean, probably in this case by means of wind-borne or bird-borne propagules. The north Chilean form of H. silenoides ('//. paposum') is the most similar part of this species to 27. H. gramineum, which is separated from it by the width of the Pacific Ocean (Robson, 1972: fig. 3 modified). The primitive form of H. gramineum, in turn, occurs in the south of its area (New Zealand, Tasmania, South Australia), whence there are two reduction trends: (a) within Australia northward and west- ward, to small erect annuals and (b) from New Zealand through New Caledonia to New Guinea, to elongate, narrow- leaved grass-like perennials (H. gramineum sensu stricto). Plants similar to those of Queensland and Northern Territory occur in lowland Vietnam and Taiwan, whereas the narrow- leaved form of Yunnan and the eastern Himalaya appears to belong to trend (b). As with H. silenoides, so here the present distribution of H. gramineum is only explicable in terms of long-distance dispersal, first (ancient) to New Zealand and Australia and subsequently to south-east Asia (see p. 95). Its introduction to Hawaii is almost certainly recent. The record of H. gramineum from Baja California cited by Rodriguez Jimenez (1972, 1973) was based on a misidentification of a specimen of H. peninsular -e Eastw. (see p. 40). (c) Principal clade (iv): subsect. Knifa The major disjunction between the areas of Id. H. rigidum subsp. bracteatum (southern Brazil, Parana) and the mostly closely related species in subsect. Knifa, 28. H. relictum (north-eastern Colombia, Santander), immediately poses a problem. Because H. relictum is clearly at the dispersion centre of this geographically very diverse subsection, the migration from south of Amazonia that established its ancestor in the north-east Andean region must have occurred at a geologically remote time, at any rate not later than the early Tertiary. It could, therefore, not have used the Pleistocene Amazonian refugia (Prance, 1978), even if these had been ecologically appropriate. The possible means of migration from south Brazil to north-east Colombia would seem, therefore, to be only two: long-distance dispersal (on birds) or gradually via the embryonic Andes. The intervening Amazonian cerrado and forest, as well as the drier vegetation of these areas in the Pleistocene, would seem to provide no ecologically appropri- ate vegetation in which Hypericum species could flourish on a direct migration route. In view of the varied history of the early paramo vegetation (protoparamo), in which Hypericum is known to have played a part (v.d. Hammen & Cleef, 1986: 175), the gradual migration along the proto- Andean foothills would seem more likely. Having reached the northern Andes, the H. relictum group (Spp. 28, 29) gave rise to three derivative clades (vii-ix). Along the Cordillera in both directions (clade vii), 30. H. thesiifolium occurs from south-west Colombia to eastern Venezuela, and it also penetrates into Panama and Costa Rica. At higher altitudes, the distribution of the more reduced 31. H. brevistylum extends from south-west Col- ombia south to the Andes of Argentina. Apart from the usual gap in lowland Panama, none of these areas includes a disjunction. Clade (viii), on the other hand, is based in central and north Mexico; and its precursors are likely to have reached there via the original, Great Antillean link between South America and the Yucatan-Belize region (see Robson, 1987: 9). Of the two divisions of clade (viii), the one based on 43. H. pumillum (clade xi) remains almost wholly in north and central Mexico apart from 44. H. paucifolium, which just extends its range into south-east Texas, and 47. H. anagal- loides, which extends northwards in the western mountains as far as southern British Columbia. Its near relative (46. H. parvulum) occurs in Hawaii ('//. degenerf) as well as Mexico, but seems to be a recent arrival there. The other division of clade (viii) (clade x) is based on 32. H. pauciflorum (west & south Mexico), from which three subsidiary clades are derived. Clade (xii) comprises a south- eastward development, with 33. H. pratense (extending as far as Nicaragua) and its derivatives 34. H. philonotis (south- eastern Mexico to Honduras) and 35. H. aphyllum (Belize). Clade (xiii) is an originally north-westward trend in which 36. H. majus evolved in north-west U.S.A. and Canada and then migrated eastward, the extreme eastern part becoming specifically distinct (37. H. canadense) and extending its area south in the eastern coastal region and back westward into the area of//, majus (c.f. Utech & Iltis, 1970). Both species have been introduced into Europe (relatively recently?), possibly by birds or (H. majus) human agency. Clade (xiv) is centred in south-eastern U.S.A. (38. H. gymnanthuni) but has become very widely spread indeed. In 39. H. mutilum there is one (type) subspecies widespread in eastern North America, a north-eastern subspecies (39b. subsp. boreale), and a south- eastern subspecies (subsp. 39c. latisepalum). The Cuban endemic 40. H. arenarioides is very similar to a form of subsp. latisepalum. 39a. H. mutilum subsp. mutilum has undergone long-distance dispersal to Central America, the West Indies, South America (both north-west and south-east), Hawaii, New Zealand, Italy, France, and Transcaucasia (Georgia). How many of these introductions are by natural means (birds) is unclear, but some at least must have occurred in HYPERICUM 11 geological time. Thus the eastern Brazil population, which was described as a separate species, H. euphorbioides A. St. Hil., has already speciated, giving rise to 41. H. pleiostylum with 5-8 styles. 38. H. gymnanthum itself has also undergone long-distance dispersal, to Guatemala, Poland, and east Asia. The trans-Pacific migration must have been relatively ancient, because H. gymnanthum no longer occurs there and the consequent 42. H. japonicum has had time to differentiate widely (though not, in my view, to subspeciate) and to spread throughout east and south-east Asia to Nepal, Sri Lanka, Australia, and New Zealand. Finally, clade (ix) includes an ancient crossing of the Atlantic in a south-easterly direction. 48. H. lalandii is a very variable African species with a distribution that extends from the Cape of Good Hope to the Sudan Republic and Nigeria and into Madagascar. Its most primitive form occurs in Natal and the Transvaal, whence it appears to have radiated in all directions. Various forms of H. lalandii are almost indisting- uishable from forms of 30. H. thesiifolium (Colombia) and 44. H. paucifolium (Mexico), the only apparent difference in the African species being the narrower, clavate stigmas. Its nearest relative would appear to be the north-west Col- ombian 28. H. relictum, a statement which again raises the question of how and when the migration from northern South America to south-eastern Africa occurred. It must, in my opinion, be ancient and therefore must have been achieved by natural means, i.e. by birds, or, less likely wind. There are no birds that migrate at present between Central or South America and southern Africa (Maclean, pers. comm., 1988). Some populations of water bird, however, that occur on both sides of the South Atlantic are still no more than subspeci- ficaly distinct (e.g. the knob-billed duck Sarkidiornis melano- tis or the grey-headed gull Lams irrocephalus , which is largely freshwater) or even show no subspecific variation whatsoever (the fulvous and white-faced ducks Dendrocygnus bicolor and D. viduata (Maclean, comm. cit.)). This variation pattern indicates that there must have been some interchange of water birds between the two landmasses within the last 20000-30000 years; and so there seems to be no reason to doubt that, some millions of years earlier, when Africa and South America were less distant than they are now, some south-eastward avian journey was responsible for introducing Hypericum lalandii into southern Africa. From H. lalandii have evolved one species and one species group. In Madagascar the endemic 49. H. globuliferum is very similar to the form of H. lalandii from that island and probably resulted from local speciation there. The species group is centred in the East African mountains, where 50. H. humbertii, the species most closely related to H. lalandii, is endemic in the Ruwenzori and Virunga Mts area. 51. H. scioanum is both more specialized and more widespread, occurring from Ethiopia south to northern Malawi; and the most reduced species (in which there is again a return to flowers with 5 styles), 52. H. oligandrum, is confined to wet habitats in the Kalahari Sands region (Zambia, southern Zaire, Angola, Namibia - Caprivi). (d) Wide disjunctions: discussion From the above outline of evolutionary trends in sect. Trigynobrathys, it is clear that this section, unlike almost all others in Hypericum, includes wide disjunctions in distribu- tion that cannot be explained either as vicarianism resulting from plate tectonics or as the result of gradual overland migration with intermediate extinction. The initial separation of the ancestral species in South America from their relatives in Africa can be attributed to the separation of continental plates; but, apart from the hypothetical migration of the H. pauciflorum group (Spp. 32-47) into Central America via the Greater Antilles land-bridge, the major disjunctions nearly all seem to involve long-distance dispersal. Within the Americas, there are two examples of dispersal between south-eastern U.S.A. and temperate Brazil, one S- N (1. H. rigidum/10. H. denticulatum) and one N-S (39a. H. mutilum subsp. mutilum). Both migrations have been fol- lowed by speciation, unlike the N-S example of this disjunction in sect. Brathys (H. gentianoides), which is therefore likely to have been more recent. H. mutilum subsp. mutilum has also spread to Central America and the West Indies (Mexico, Honduras, Dominican Republic) and the northern Andes (Colombia, Ecuador, Peru) without sub- sequent speciation, and its close relative 38. H. gymnanthum (south-eastern U.S.A.) occurs also in Guatemala. The gap between Id. H. rigidum subsp. bracteatum and 28. H. relictum could, as we have seen, been the result of either long-distance dispersal or gradual northward migration. Of the Atlantic Ocean disjunctions, only one concerns the South Atlantic (28. H. relictum - 48. H. lalandii) and it is clearly the result of ancient dispersal. All the others involve Spp. 36-39, species of the eastern U.S.A. and Canada that have, along with H. gentianoides (sect. Brathys), 'turned up' in scattered localities in Europe and the Caucasus. Despite arguments to the contrary (see discussion under H. cana- dense, p. 113), I believe that all these species have appeared in Europe within the last c. 150 years, some probably as a result of carriage by birds, others possible being introduced accidentally by humans (cf. Bouchard, 1954). There are also northern and southern trans-Pacific Ocean disjunctions. The southern one (25a. H. silenoides subsp. silenoides - 27. H, gramineum) is clearly due to ancient dispersal, almost certainly long-distance, as is one of the northern ones (38. H. gymnanthum - 42. H. japonicum). In both cases there has been subsequent speciation. Two others appear to be more recent in origin: 36. H. majus (northern U.S.A. or Canada - Japan) and 46. H. parvulum (northern Mexico - Hawaii). These may well be results of human activity; the occurrences of 39c. H. mutilum subsp. mutilum in New Zealand and Hawaii and of 27. H. gramineum in Hawaii almost certainly are. It is always possible to attribute 'freak' distributions to the result of unusual winds 'sometime in the past'; but, while not denying that such causes and effects occur, I think that other explanations should be sought first. It is no doubt significant that the Hypericum section that contains most marsh and wet- margin herbs is just the one in which long-distance dispersal is conspicuously present. Mud containing seeds frequently becomes attached to feet or feathers of wading birds; and although migrating birds tend to travel clean, seeds and other propagules are undoubtedly sometimes transported for long distances (Ridley, 1930; v.d. Fiji, 1982; Carlquist, 1983). It seems reasonable, then, to suggest birds as the likely vehicles for carriage of the small seeds of damp-loving Hypericum species over long distances. Correlation of disjunctions with known avian migration routes (present or past) would go far towards verifying this hypothesis, though it seems too much to expect that a bird will be caught in flagrante delicto. 12 N. K. B. ROBSON 7(5) Leaves persistent (deciduous with cortex) (subsect. 2. SYSTEMATIC TREATMENT Pheiiotesp.p.) 8 Leaves deciduous above base or with whole stem internode (subsect. 3. Brathys) 44 Sect. 29. BRATHYS (Mutis ex L.f.) Choisy, Prodr. 8(6,7) Leaves deciduous (partly or apparently completely) monogr. Hyperic.: 38, 58 (1821). 3 without fading and/or with margin recurved to revolute and/or internodes with corky emergences 9 BASIC CHROMOSOME NUMBER (x): 12; ploidy 2 (correction to Leaves persistent (fading in situ, deciduous with Part 7). cortex), margin plane to involute; internodes with- HABITAT. Add: high cold swamps (13. H. callacallanum , 16. H. roraimense); open woods, fields, roadsides, and waste 9(8) Stem internodes with ± prominent lenticels or corky places, 0-1040 m (87. H. drummondii, 88. H. gentianoides); emergences; leaves deciduous at the apparent base open Pinus forest, grassy slopes, roadside, and ditches, often or marcescent in damp places, 300-3000 m (79. H. fuertesii, 80. H. ^ internodes smooth; leaves deciduous above ... 01 u j- -j \ oo • / • 11 u base or persistent, not marcescent 15 dichotomum, 81. H. diosmoides). 88 species (+ 12 subspe- cies). 10(9) Leaves becoming deflexed, marcescent; stem inter- nodes almost smooth 3a. phellos subsp. marcescens DISTRIBUTION. Add: Nicaragua (82. H. gnidioides), Revilla Leaves erect to spreading or, if becoming deflexed, Gigedo Islands (83. H. eastwoodianum), and south-eastern then deciduous; stem internodes with prominent Brazil and Uruguay (10. H. piriai). emergences 11 11(10) Leaves spreading to deflexing ... 3b. phellos subsp. phellos Key tO Sect. 29. Brathys (revised) Leaves remaining or becoming appressed, tetra- stichous 12 1 Leaves deciduous at base leaving rhombic scar, without interfoliar ridge; styles usually 5; flowers 12(n) Leaves Wlth 3 basal vems (branched or not), some- 15-70 mm in diam. (subsect. 1. Styphelioides) .... 2 times spreading at first or deciduous, margin plane to recurved 13 Leaves deciduous at apparent base leaving crescentic Leaves with 1 basal vein (venation wholly pinnate), or round scar with interfoliar ridge, or at least base closely appressed, subpersistent, margin incurved persistent; styles 3(4) or, if 5, then flowers 4-10 mm 3d. phellos subsp. platyphyllum in diam 5 13(12) Leaves deciduous 3c. phellos subsp. oroqueanum 2(1) Leaves narrowly elliptic, acute, 16-30 mm long, Leaves persistent 14 chartaceous to subcoriaceous; sepals 14-22 mm j j terrae-firmae 14(13) Leaves with 3 unbranched basal veins; flowers 25-30 Leaves narrowly obovate or, if oblong to elliptic, then mm in diam- 5 sty'es 9~10 mm long' ^ x as Ion8 as acuminate or shorter (10-16 mm long) or thickly ovary 4- 'razuense coriaceous; sepals 5-15 mm long (2. Styphelioides) 3 Leaves Wlth at least midrib branched; flowers 15-27 mm in diam.; styles 5-9 mm long, 2-4 x as long as 3(2) Leaves narrowly elliptic to narrowly oblong, usually ovary 5. stenopetalum thinly coriaceous, spreading, often outcurving .... 2a Styphelioides subsp. clarense 15(9) Leaves completely revolute (lower surface obscured), Leaves oblanceolate to narrowly obovate, ± thickly linear' deciduous at apparent base or persistent .... 16 coriaceous, ± densely imbricate, straight 4 Leaves Wlth mar§in Plane to recurved or incurved (lower surface at least partly visible), elliptic to 4(3) Leaves with 7-1 1 basal veins, glands not prominent obovate-spathulate or linear, deciduous above base 2b. Styphelioides subsp. Styphelioides or persistent 18 Leaves with 5-7 basal veins, glands prominent 2c. Styphelioides subsp. immense 16(15) Leaves deciduous, 4-10 mm long, elliptic-oblong to linear; sepals ovate to lanceolate; shrub to 3 m tall 5(1) Leaves free, usually with apex of lower interfoliar 8. garciae stem line visible between bases 6 Leaves persistent, (8-) 10-40 mm long, linear; sepals Leaves united, apex of lower interfoliar stem line narrowly lanceolate to linear or oblong; dwarf obscured by interfoliar ridge 7 shrub to 0-8 m tall or subshrub or herb . . 17 6(5) Inflorescence 1-flowered or, if flowers 2-25, then leaves completely revolute or only margin in- curved; trees, shrubs, shrublets or rarely subshrubs or perennial herbs (subsect. 2. Phellotes p.p.) .... Inflorescence (l)2-c. 40-flowered; leaves never com- pletely revolute or only margin incurved; shrubs (rarely), subshrubs or wiry annual herbs (subsect. 4. Spachium) ................................ 88 'Including Sarotha L., Sp. pi.: 272 (1753), Gen. pi., 5th ed.: 13 (1754); Hypericum sect. Perforaria Choisy (1821), pro parte quoad H. dichotomum Lam.; Mania sensu Sprengel, Syst. veg. 3: 333 (1826), pro parte quoad H. dichotomum Lam.; Hypericum sect. Sarothra (L.) Reichardt (1878); Hyper- icum sect. Brathys subsect. Spachium R. Keller (1893); Sarothra sect. Spachium(R. Keller) Y. Kimura (1951); Sarothra sect. Spachium series Eusarothra Y. Kimura (1951); Hypericum sect. Spachium (R. Keller) N. Robson; see Part 1 (Robson, 1977: 338). 17(16) Flowers solitary, inflorescence branching pseudo- dichotomous; flowers stellate; stamens not fascicu- late; dwarf shrub 9. acostanum Flowers 1-25, inflorescence branching monochasial; flowers obconic to pseudotubular; stamens 3(5)- fasciculate; subshrub or perennial herb 10. piriai 18(15) Styles 8-10 mm long; leaves 5-8 mm wide, plane or margin recurved, deciduous above base without twisting 22. paramitanum Styles 3-6(-7) mm long; leaves 0-8-3 mm wide, subrecurved to incurved, persistent or deciduous above base after twisting 19 19(18) Leaves 3-5-5(-7) mm long, plane to subrecurved, midrib prominent beneath; flowers 8-13 mm in diam. . .6. carinosum HYPERICUM 13 20(8) 21(20) 22(21) 23(21) 24(23) 25(23) 26(25) 27(26) 28(27) 29(27) Leaves 10-13 mm long, involute, midrib impressed beneath; flowers c. 25 mm in diam 7. matangense Leaves plane (sometimes except base) or with margin or apex incurved, narrowly oblong or narrowly elliptic or narrowly lanceolate to elongate-linear, bases free or almost so (except in Sp. 12) 21 Leaves incurved or, if plane, then broadly oblong or ovate to subcircular, bases united Leaves elliptic or narrowly oblong to oblanceolate- spathulate (1: b = 2-5-3-5), base cuneate or margin slightly recurved, with 3-7 basal or near-basal veins; styles 4-5-6 mm long Leaves linear-lanceolate or linear-oblong or linear- elliptic to linear (1: b = 4-14), base parallel-sided or, if base narrowed, then margin slightly incurved; styles 1-4-6 mm long 31 22 23 Leaf surface smooth, ± lustrous, bases free or very shortly united; stigmas narrow 11. simonsii Leaf surface papillose, dull, bases united forming cup-shaped sheath; stigmas capitate 12. papillosum Leaves with base narrowly cuneate to angustate, narrowly oblong to narrowly elliptic, discolorous, margin usually slightly incurved 24 Leaves with base parallel-sided, or if slightly nar- rowed then elongate-linear to linear-oblanceolate, concolorous, plane or whole lamina incurved 25 Leaves with 3 basal or near-basal veins, midrib clearly branched; petals 9-11 mm long; inflorescence 1- flowered, branching pseudo-dichotomous 13. callacallanum Leaves 1 -nerved, midrib not or obscurely branched; petals 6-8 mm long; inflorescence 1(3-12)- flowered, branching pseudo-dichotomous to dichasial/monochasial 14. humboldtianum 26 30 30(25) Plant erect, bushy, 1-stemmed Plant spreading or diffuse, stems rooting Leaves broadest below middle, narrowly lanceolate to narrowly elliptic, 1-5-2-5 mm wide, base rounded-amplexicaul to cuneate 15. gladiatum Leaves broadest at or above middle, very narrowly elliptic or very narrowly oblanceolate-spathulate to linear, 0-5-1-8 mm wide, base cuneate or angustate to parallel-sided 27 Leaves chartaceous, spreading to reflexed from above base, apex acute to obtuse, cucullate, base ± sheathing 28 Leaves coriaceous, erect to gradually outcurving, apex sharply acute to acuminate, not cucullate, base plane to incurved but not sheathing 29 Leaves plane, acute, 6-13 mm long, margins straight, becoming reflexed from above base 16. roraimense Leaves incurved, apiculate to obtuse, 3-4-5 mm long, margin undulate, recurving or recurving-ascending from above base 17. sp. Leaves plane to subconduplicate, base parallel-sided to subangustate, glands prominent; stigmas broadly capitate; sepals triangular-lanceolate, 5-ribbed . . . 18. espinalii Leaves incurved at least towards base, base angus- tate, glands impressed; stigmas subcapitate; sepals triangular-linear, 3-ribbed 19. castellanoi Flowers 12-15 mm in diam.; sepals 6-7 mm long; leaves subacuminate, glands not prominent 20. asplundii 31(20) 32(31) 33(32) 34(33) 35(32) 36(35) 37(31) 38(37) 39(37) 40(39) 41(40) 42(39) Flowers c. 6 mm in diam.; sepals 3-5 mm long; leaves acute, glands prominent 21. radicans Leaves sessile, united by broad bases of laminae, often becoming deflexed 32 Leaves pseudopetiolate, united by bases of pseudo- petioles, or sessile but base narrowed (Spp. 34,35), rarely becoming deflexed (Sp. 28 p.p.) 37 Flowers 18-40 mm in diam.; leaf base rounded to cordate-amplexicaul 33 Flowers 6-12 mm in diam.; leaf base parallel to truncate 35 Stem internodes smooth; leaves plane or slightly saccate, (2-)3-7 mm wide, margin not or scarcely hyaline 23. cuatrecasii Stem internodes with corky emergences, at least below upper leaves; leaves saccate or incurved- cucullate, 0-7-3-5 mm wide, margin markedly hyaline 34 Leaves becoming deflexed, saccate; sepals 5-8 mm long 24. goyanesii Leaves spreading only, incurved-cucullate; sepals 4-5 mm long 25. myricariifolium Leaves becoming deflexed, broadly ovate to triangular-ovate or (lower) oblong, basal veins 3(5) 26. quitense Leaves erect or spreading only, triangular-ovate or oblong to linear, basal veins 1(3-5) (27. loxense) .. 36 Leaves broadly triangular-ovate to lanceolate, usually imbricate, base truncate to broadly cuneate 27a. loxense subsp. aequatoriale Leaves narrowly oblong or narrowly elliptic to oblanceolate or linear, usually spreading, base narrowly cuneate to parallel-sided 27b. loxense subsp. loxense Leaves sessile, lamina elliptic-oblong to narrowly oblong, base cuneate to parallel-sided 38 Leaves shortly petiolate or base angustate or, if base narrowly cuneate, then lamina linear to acicular .. 39 Leaves 5-7 mm long, elliptic or oblong to obovate; sepals 1-8-2-3 mm wide, apex plane 34. hartwegii Leaves 2-4 mm long, narrowly elliptic or narrowly oblong to linear; sepals 0-7-1-7 mm wide, apex incurved-cucullate 35. maguirei Leaves with midrib proximally prominent or wholly level with lamina; lamina conduplicate, apex usually acuminate to obtuse 40 Leaves with midrib proximally impressed; lamina incurved, apex usually obtuse to rounded 42 Leaves coriaceous, markedly tetrastichous, apex acu- minate to acute, margin plane 41 Leaves chartaceous, weakly or not tetrastichous, apex acute to rounded, margin usually undulate . . 30. thuyoides Leaves oblong or elliptic to lanceolate, 1-5-3 mm wide, imbricate or ± spreading; petals 13-16 mm long, bright yellow 28. lycopodioides Leaves narrowly lanceolate, appearing curved- acicular, 1-1-5 mm wide, recurved; petals c. 9 mm long, deep yellow 29. woodianum Leaves subpapillose beneath, base cuneate to pseudopetiolate; sepals obtuse to rounded 31. sabiniforme Leaves smooth beneath, base narrowly cuneate to angustate or parallel-sided; sepals acute to obtuse 43 14 N. K. B. ROBSON 43(42) 44(7) 45(44) 46(45) 47(46) 48(44) 49(48) 50(49) 51(50) 52(48) 53(52) 54(53) Plant a bushy or spreading shrub or small tree (0-1)0-3-6 m tall; leaves with glands not or scarcely impressed; sepals usually plane 32. laricifolium Plant a pulviniform shrub 0-08-0-16 m tall; leaves with glands impressed; sepals markedly ribbed . . . 33. martense Leaves and sepals with conspicuous marginal glands that usually secrete clear or milky resin; lamina base ± sheathing 45 Leaves and sepals without conspicuous marginal glands, not sticky; lamina base not sheathing 48 Leaves ± abruptly spreading to deflexed above sheathing base, chartaceous, marcescent, elliptic to oblanceolate 36. magniflorum Leaves erect or gradually outcurving, usually coriaceous, deciduous above base, elliptic to cir- cular 46 Leaves not glaucous, epidermis smooth, elliptic or oblanceolate to obovate; stigmas broadly capitate 47 Leaves densely glaucous, epidermis undulate-papil- lose, broadly elliptic to obovate-spathulate or circular; stigmas narrow 39. baccharoides Flowers solitary, 20-40 mm in diam., sometimes terminating clustered short shoots; leaves 7-10 mm long, always coriaceous; stems erect 37. gleasonii Flowers in 3-13-flowered cymes or, if solitary, less than 20 mm in diam.; leaves up to 17 mm long, sometimes chartaceous and then habit decumbent and rooting 38. mexicanum Leaves sessile, apex cucullate, if 1-nerved then broadest above middle 49 Leaves shortly pseudopetiolate and/or apex not cucullate, if 1-nerved then broadest at or below middle 52 Leaves elliptic-ovate to elliptic-lanceolate, 3-7 mm broad, venation flabellate; flowers 25-40 mm in diam 40. stuebelii Leaves oblanceolate to narrowly oblong, 0-7-3 mm broad, venation pinnate or 1-nerved; flowers 6-25 mm in diam. (petals unknown in 41. prietoi) 50 Sepals acuminate; leaves persistently imbricate- tetrastichous, markedly laterally compressed dis- tally; stigmas capitate 41. prietoi Sepals acute; leaves eventually outcurving, not later- ally compressed distally or, if so (42. cassiopiforme in part) then stigmas narrow 51 Petals c. 14 mm long; styles longer than ovary; stigmas narrow; leaves oblanceolate (1: b = c. 2-5) 42. cassiopiforme Petals 5-9 mm long; styles 0-6-1 x as long as ovary; stigmas capitate; leaves narrowly oblanceolate to linear-elliptic (1: b = 4-11) 43. decandrum Leaves with venation flabellate, some laterals free or almost free from the base; styles stout, stigma usually capitate 44. pimeleoides Leaves with venation pinnate, all laterals originating well above the base, or 1-nerved; styles and stigmas various . 53 55(54) 56(55) 57(56) 58(57) 59(57) 60(59) 61(55) 62(61) 63(53) 64(63) 65(64) Leaves petiolate or abruptly narrowed at the base or stem disarticulating 54 Leaves sessile, gradually narrowed or parallel-sided at the base; stem not disarticulating 63 Styles 6-7 mm long; flowers 25-30 mm in diam.; 66(65) leaves often spreading abruptly above petiole .... 62. jaramilloi Styles 0-8-5 mm long; flowers up to 25 mm in diam.; leaves outcurving or spreading from the base 55 Leaves with midrib prominent or level with lamina beneath, glands usually visible beneath 56 Leaves with midrib ± impressed beneath, glands not visible beneath 61 Stigmas broadly capitate; leaves glaucous, dull 63. cardonae Stigmas clavate or small; leaves not glaucous, lustrous 57 Leaves with margins slightly recurved to incrassate (64. caracasanum) 58 Leaves with margin or whole lamina incurved 59 Leaves 9 x 4-5 mm or larger; petals c. 1-3 x as long as sepals; styles c. 1-3 x as long as ovary 64a. caracasanum subsp. caracasanum Leaves 8 x 3-5 mm or smaller; petals 1-5-2 x as long as sepals; styles 2-2-5 x as long as ovary 64! i. caracasanum subsp. turumiquirense Leaves elliptic to linear, petiolate or angustate, 3-5- 11 mm long; stem not disarticulating 60 Leaves lanceolate-triangular, sessile, amplexicaul, 0-7-2 mm long; stem disarticulating at internodes 67. mi II i- folium Leaf lamina elliptic, 2-4 mm wide, plane with margin incurved; styles 5-6 mm long 65. ekmanii Leaf lamina narrowly elliptic to linear, 0-5-2 mm wide, incurved to canaliculate; styles 3-5-4 mm long 66. pycnophyllum Leaves plane or with margin subincrassate, narrowly ovate to narrowly oblong-elliptic; sepals 1-5-4 mm wide; styles 2-5^ mm long 68. ruscoides Leaves incurved to canaliculate, elliptic or linear- lanceolate to linear; sepals 0-5-1-5 mm wide; styles l-2(-3) mm long 62 Leaf lamina elliptic, 4-6 mm wide; stigmas narrow . . 69. harlingii Leaf lamina linear-lanceolate to linear, 0-5^ mm wide; stigmas broadly capitate 73a. lancioides subsp. lancioides Leaf and sepal apices ± incurved-cucullate; leaf lamina narrowly oblong or narrowly elliptic to linear, narrowed at base 64 Leaf and sepal apices not incurved or cucullate or, if so, then leaves broadened at the base 69 Ovary ellipsoid to globose; styles 0-8-2(-3) mm long, 0-4-1-1 x as long as ovary, stigmas ± broadly capitate; sepals usually oblong to oblanceolate, not ribbed 65 Ovary ovoid; styles either 3-4-5 mm long, 1-3-1-5 x as long as ovary, stigmas scarcely to narrowly capitate, or if styles shorter (1-5-2-5 mm long, 0-6- 0-7 x as long as ovary) and broadly capitate, then sepals lanceolate, ribbed 67 Stems erect or rarely decumbent; leaves 7-15 mm long; sepals 5-8 mm long; styles 3(4), l-2(-3) mm long (73. lancioides) 66 Stems prostrate or ascending; leaves 4-7 mm long; sepals 3-4 mm long; styles (3)4-5, 0-8-1 mm long 74. selaginella Flowers terminal, solitary, inflorescence branching pseudo-dichotomous; habit bushy or caespitose . . . 73a. lancioides subsp. lancioides HYPER1CUM 15 67(64) 68(67) 69(63) 70(69) 71(70) 72(71) 73(72) 74(73) 75(74) 76(75) 77(75) Flowers terminal and lateral in dense racemiform heads, all branching lateral; habit bushy or strag- gling 73b. lancioides subsp. congestiflorum Styles 3^-5 mm long, 1-3-1-5 x as long as ovary; stigmas not or scarcely capitate; leaves 1-5-4 mm wide, margin not or scarcely hyaline 68 Styles 1-5-2-5 mm long, 0-6-0-7 x as long as ovary; stigmas broadly capitate; leaves 0-8-1-5 mm wide, margin narrowly but distinctly hyaline 72. andinum Sepals oblong-oblanceolate, not ribbed; leaves plane or with margin slightly incurved, 6-9 mm long, markedly tetrastichous, glaucous 70. llanganaticum Sepals ovate-lanceolate to oblong, ribbed; leaves with margin markedly incurved, 8-15 mm long, not or obscurely tetrastichous, lustrous 71. struthiolifolium Styles 6-7-5 mm long; flowers 20-30 mm in diam.; leaves with midrib branched, lamina elliptic to oblanceolate, plane or conduplicate, epidermis smooth 45. magdalenicum Styles up to 5 mm long; flowers up to 20 mm in diam. ; leaves with midrib unbranched or, if branched, then lamina narrowly oblong or epidermis papillose above 70 Leaves markedly tetrastichous, narrowly elliptic, papillose above; branches very strict, parallel, with internodes 1-1-5 mm long 56. parallelum Leaves not tetrastichous or if so, then linear, smooth, and branches not as above 71 Sepal margin green or, if hyaline, then inflorescence a racemiform synflorescence and styles shorter than ovary with stigmas capitate 72 Sepal margin hyaline; inflorescence and styles not as above 81 Flowers solitary, not congested; styles 2-5-5 mm long, stigmas narrow 73 Flowers in congested heads or racemiform synflor- escences; styles l-3(-4-5) mm long, stigmas broadly capitate 79 Leaves 2-3 mm wide, narrowly elliptic, slightly incurved 46. valleanum Leaves 0-4-2 mm wide, very narrowly oblanceolate to linear, markedly incurved to conduplicate 74 Styles 2-5-3-5 mm long; leaves lustrous beneath Styles 1-2 mm long; leaves dull beneath Leaves becoming twisted, incurved but not carinate or conduplicate, margin very narrowly hyaline, dorsally usually glandular when mature Leaves remaining erect or becoming recurved, in- curved to conduplicate, often at least partly carin- ate, margin broadly hyaline, dorsally eglandular when mature . 75 78 76 77 Sepals 7-11 mm long; flowers 20-30 mm in diam.; capsule c. 5 mm long; leaves 6-12 x 0-8-1-7 mm; stamens 60-75 47. sprucei Sepals 4-7 mm long; flowers 10-18 mm in diam.; capsule 3-5-4 mm long; leaves 3-7-7 x 0-4-0-6 mm; stamens 30-55 48. aciculare Leaves imbricate at first but not markedly tetra- stichous, recurving, 5-8 mm long; flowers 15-20 mm in diam.; petals 8-12 mm long 49. recurvum Leaves densely and persistently imbricate, 'winged'- tetrastichous, scarcely spreading, 9-14 mm long; flowers 20-25 mm in diam. ; petals 15-18 mm long 50. wurdackii 83(82) 78(74) Flowers 15-20 mm in diam.; leaves 5-15 mm long, acute; stems woody, not rooting 51. costaricense Flowers 6-8 mm in diam.; leaves 3-5 mm long, subacute to obtuse; stems wiry, basally rooting . . . 52. bryoides 79(72) Leaves very narrowly elliptic, 15 x 3 mm or larger; sepals 9-11 mm long, coriaceous; styles c.4-5 mm long 53. bolivaricum Leaves linear, up to 14 x 1-5 mm; sepals 2-7 mm long, chartaceous to membranous; styles 0-6-3 mm long 80 80(79) Stems erect to decumbent; leaves 6-14 mm long; styles 1-5-3 mm long, shorter to longer than ovary 54. juniper in u in Stems prostrate; leaves 2-5 mm long; styles 0-6-0-8 mm long, shorter than ovary 55. prostratum 81(71) Leaves lustrous, margin markedly incurved, lamina twisting and sometimes recurving (57. marahuaca- num) 82 Leaves dull or with metallic sheen, margin plane to incurved, lamina incurved only 84 82(81) Stigmas small or slightly enlarged; ovary and capsule acute; inflorescence branching pseudo-dichot- omous or racemiform 83 Stigmas enlarged to capitate; ovary and capsule ± rostrate; inflorescence branching pseudo-dichot- omous to lateral-congested 57c. marahuacanum subsp. chimantaicum Styles 4-5 mm long; sepals 7-9 mm long; infloresc- ence branching pseudo-dichotomous 57a. marahuacanum subsp. marahuacanum Styles 2-3(-4) mm long; sepals 4-5 mm long; in- florescence branching lateral, racemiform 57b. marahuacanum subsp. strictissimum Leaves plane or slightly incurved, oblong-linear or narrowly oblanceolate, the tips not red when young 58. lancifolium Leaves markedly incurved to canaliculate, linear to linear-acicular, the tips usually red when young ... 85 Plant a prostrate shrublet with branches matted, rooting; leaves often secund, not tetrastichous .... 59. horizon talc Plant an erect shrub with branches strict, not rooting; leaves never secund, usually ± tetrastichous 86 Leaves with metallic sheen, marked tetrastichous, erect to suberect 60. tetrastichum Leaves dull, not tetrastichous (except sometimes in young parts or young plants), usually outcurving (61. strictum) 87 Leaf and sepal apices acicular; sepals 1-3-2 mm wide, not ribbed; leaves 8-13 mm long; branches strict, usually all from one stem . . 61a. strictum subsp. strictum Leaf and sepal apices subacute; sepals 1-1-2 mm wide, clearly ribbed; leaves 5-7(8) mm long; branches ascending, usually branched from the base 61b. strictum subsp. compactum Leaf venation parallel, 5-7-nerved; stigmas narrow; shrub 75. cymobrathys Leaf venation pinnate and/or 1-3-nerved; stigmas capitate to peltate; subshrubs to annuals 89 84(81) 85(84) 86(85) 87(86) 88(6) 89(88) Leaves pseudopetiolate or subsessile, with 2-4 pairs of lateral veins, ovate or elliptic to narrowly oblong or oblanceolate (1: b = 2-4) (76. chamaemyrtus) . . 90 16 90(89) N. K. B. ROBSON 91(89) 92(91) 93(92) 94(92) 95(94) 96(91) Leaves sessile, without or with 1 pair of lateral veins, lanceolate or narrowly elliptic-oblong to linear to linear-subulate (1: b = 6-12) 91 Inflorescence pseudo-dichotomous with ultimate branches dichasial/monochasial; leaves pseudo- petiolate, lamina base broadly cuneate to rounded 76a. chamaemyrtus subsp. chamaemyrtus Inflorescence wholly pseudo-dichotomous; leaves subsessile, lamina base narrowly cuneate to angus- tate .... 76b. chamaemyrtus subsp. pseudocaracasanum Styles 4-5 mm long or, if shorter, then inflorescence branching wholly pseudo-dichotomous; plant a subshrub, shrublet or perennial herb 92 Styles 0-6-3 mm long; inflorescence branching wholly dichasial/monochasial or mixed; plant a subshrub or perennial or annual herb 96 Styles 4-5 mm long; plant a subshrub or shrublet, erect or decumbent, but not rooting; inflorescence branching various 93 Styles 0-5-3 mm long; plant a shrublet or perennial herb, erect to prostrate and rooting; inflorescence branching wholly pseudo-dichotomous/sympodial 94 Sepals 4-5-10 mm long, exceeding capsule; bracts foliaceous; inflorescence branching mixed or wholly pseudo-dichotomous; terminal stem- internodes not relatively elongate 77. arbuscula Sepals 3-5^-2 mm long, shorter than capsule; bracts linear-subulate; inflorescence branching dichasial/ monochasial; terminal stem-internodes relatively elongate 78. rubritinctum Leaves linear or very narrowly oblong to oblan- ceolate, plane or incurved, acute; styles 2-5-3 mm long 79. fuertesii Leaves oblong-elliptic or oblong-spathulate to oblan- ceolate or obovate, plane to revolute, subacute to rounded; styles 0-5-2-5 mm long 95 Leaf margin plane; petals c. 1-5 x as long as sepals; styles 1-5-2-5 mm long 80. dichotomum Leaf margin recurved to revolute; petals about equalling sepals; styles 0-5-1(1-5) mm long 81. diosmoides Inflorescence branching at least partly pseudo- dichotomous; capsule nearly always shorter than sepals; leaves acute to subacute 82. gnidioides Inflorescence branching wholly dichasial/mono- chasial; capsule (unknown in Sp. 84) exceeding or equalling sepals or, if shorter, then leaves obtuse or rounded . 97 lax; subshrub or perennial herb 97(96) 98(97) 99(98) Plant a shrublet or ericoid subshrub or perennial herb; leaves 1-6-5 mm broad, if appressed then margin not entire Plant an annual herb; leaves 0-4-1 mm broad, appressed, margin entire 101 Capsule shorter than to slightly exceeding sepals, usually ellipsoid or cylindric-ellipsoid; leaves coriaceous to thickly chartaceous with margin usually entire 99 Capsule 2-3 times as long as sepals, narrowly ovoid- cylindric or leaves thinly chartaceous with margin undulate-denticulate 100 Styles shorter than ovary; stem branched above base only; inflorescence usually rather dense; subshrub or shrublet 83. eastwoodianum Styles longer than ovary; stem usually with geniculate branches from base; inflorescence rather lax to very 84. peninsulare 100(98) Leaves thinly chartaceous, becoming deflexed, mar- gin undulate-denticulate 85. beamanii Leaves coriaceous, appressed, margin retrorse-denti- culate 86. galinum 101(97) Leaves linear-lanceolate to linear-subulate, 5-22 mm long, margin revolute to subincurved; sepals 3-7 mm long; capsule about equalling sepals 87. drummondii Leaves canaliculate to scale-like, 1-4 mm long, margin incurved; sepals 1-5-2-5 mm long; capsule 2-3 x as long as sepals 88. gentianoides Subsect. 1. Styphelioides N. Robson, subsect. nov. Arbores parvae vel frutices. Folia libera, sessilia, plana vel plusminusve incurva, apice interdum concava, 5-11-nervia, venatione parallela vel subparallela vel flabellata, basi de- cidua cicatricem rhombicum relicta. Inflorescentia 1 -flora ramificatione pseudo-dichotoma vel lateral!. Flores 15-70 mm in diameter, stylis 5 vel raro 4-3. Typus: H. Styphelioides A. Rich. (Species 1-2). 1. Hypericum terrae-firmae Sprague & Riley (Part 7: 18; No. 1). 2. Hypericum Styphelioides A. Rich. (Part 7: 20; No. 2). 2a. subsp. clarense Lippold; 2b. subsp. Styphelioides; 2c. subsp. moaense Lippold. Icon: Leon & Alain, Fl. Cuba 3: 316, f. 141 (1953). Subsect 2. Phellotes N. Robson, subsect. nov. Arbores parvae vel frutices vel fruticuli vel raro suffrutices vel herbae perennes. Folia libera vel binatim conjuncta, sessilia vel rariore breviter pseudopetiolata, plana vel margine recurva vel incurva vel omnino revoluta vel incurva vel involuta, apice interdum concava vel cucullata, 1-7-nervia venatione pinnata vel simplici, ut videtur basi decidua cicatricem lunarem vel circularem relicta vel omnino persis- tentia et interdum marcescentia vel rarissime supra basin decidua (Sp.3). Inflorescentia l(2-5)-flora ramificatione pseudo-dichotoma vel laterali vel (-15)-flora ramificatione cymosa (Sp.14). Flores 5-30 mm in diameter, stylis 3 vel raro 4. Typus: H. phellos Gleason (Species 3-35). 3. Hypericum phellos Gleason (Part 7: 23; No. 3) 3a. subsp. marcescens N. Robson, subsp. nov. 98 Map 1. HYPERICUM 17 Map 1 Sect. 29: 3a. H. p Helios subsp. marcescens •; 7. H. matangense O; 15. H. gladiatum •; 16. H. roraimense A; 17. //. sp. A. Hypericum phellos subsp. phellos, variant 3a (i) (Part 7: 25). Caulis internodii eminentia suberosa vix prominentes in- struct!. Folia deflexescentia et marcescentia ante cadentia, margine recurva, basi 3-7-nervia. Type: Colombia, Santander, vicinity of La Baja, 2700 m, 14- 31 January 1927 (fl), Killip & Smith 18756 (GH!-holotype; GH!,NY!,US!-isotypes). Colombia (Santander). COLOMBIA. Santander: vicinity of Vetas, 3100-3250 m, 16-20 January 1927 (fl & fr), Killip & Smith 17255 (GH, NY); above California, gold-mining valley, 2700-2900 m, 14 September 1985 (fl and fr), Wood 5062 (K). 3b. subsp. phellos (Part 7: 23; No. 3a, variants (ii)-(vi)). Caulis internodii eminentia suberosa plusminusve prominen- tes instruct! . Folia patentia vel deflexescentia ante cadentia sed non marcescentia, margine recurva vel plana, basi 3- nervia. 3c. subsp. oroqueanum N. Robson (Part 7: 25; No. 3b). Caulis internodii eminentia suberosa plusminusve prominen- tia instruci. Folia ad caulem continue adpressa ante cadentia margine plana, basi 3-nervia. 3d. subsp. platyphyllum (Gleason) N. Robson (Part 7: 25; No. 3c). Caulis internodii eminentia suberosa satis prominentia in- structi. Folia ad caulem continue adpressa, persistentia, margine recurva vel plana vel incurva, basi 1-nervia. 4. Hypericum irazuense Kuntze ex N. Robson (Part 7: 30; No. 7). Correction to Part 7: 30: Chromosome number of H. irazuense is 2n = 24, not 2n = 12. 5. Hypericum stenopetalum Turcz. (Part 7: 31; No. 8). 6. Hypericum carinosum R. Keller (Part 7: 33; No. 9). Delete Venezuelan record Gehringer 149 (VEN), see p. 22. A collection from Santander ('about 6 km E. of Cerrito, 3000 m, 5 April 1984', Wood 4335 (K)) extends the range of variation of this species sufficiently to warrant modification to the description in Part 7. Some or all of these variations may be the result of the damp habitat: 'On a scrubby bank between potato fields, growing near where an irrigation ditch flows down the slope'. Shrub . . ., erect or with drooping branches, . . . Stems . . . internodes 2-20 mm long. Leaves sessile or pseudopetiolate with petiole up to 1 mm long, spreading to reflexed from the base or above the pseudopetiole, . . . lamina 3-5-7 x 0-8-3 mm, narrowly elliptic or narrowly oblong to obovate- spathulate, plane or subrecurved, apically subconcave when small, markedly carinate or with midrib only slightly promin- ent beneath, concolorous, slightly to markedly glaucous, chartaceous to subcoriaceous; apex acute to apiculate or rounded, . . . Inflorescence (etc.) as before. 7. Hypericum matangense N. Robson, sp. nov. Map 1. H. phelloti subsp. marcescenti N. Robson affinis, sed caulibus gracilioribus, foliis angustioribus involutis discoloribus tor- quescentibus semper uninervis nervi ramis et venatione 18 N. K. B. ROBSON reticulata indistinctis, inflorescentia uniflori, staminibus paucioribus, stylis brevioribus, differt. Type: Ecuador, Morona-Santiago, Paramo de Matanga, Sigsig-Gualaquiza road, 3000 m, 14 December 1980 (fl), Holm-Nielsen, Jara- millo & Coello 29401 (AAU!-holotype). Shrub 0-8 m tall, erect, with branches strict, lateral. Stems orange-brown, 4-lined and ancipitous when young, eventually terete, cortex exfoliating in strips; internodes 2-4 mm long. Leaves sessile, spreading from above base, twisting, not tetrastichous ?, persistent or sometimes deciduous (breaking off) above base (usually near point of flexure) after fading; lamina 10-13 x 1-1-3 mm, oblanceolate (lower) to linear, involute, not cucullate or carinate, dull ferrugineous above, lucent beneath, not glaucous, subcoriaceous to chartaceous; apex acute, base parallel, not or loosely sheathing, free or united with stem-line apex to form very narrow interfoliar ridge; basal vein 1, impressed beneath, with obscure ascend- ing branches and dense tertiary reticulation; laminar glands dense, visible on both sides, slightly impressed. Inflorescence 1-flowered, terminal and on lateral branches; pedicel c. 2-3 mm long, slightly incrassate upwards; upper leaves foliose. Flowers c. 25 mm in diam., stellate. Sepals 9 x 1-5-2 mm, linear-oblong, acute; veins 3, unbranched ?, not prominent. Petals bright? yellow, 18 x c. 5 mm, c. 2 x sepals, oblanceolate; apiculus acute; glands linear, distally punct- iform. Stamens c. 50, longest 9 mm long, c. 0-5 x petals. Ovary 4x2 mm long, ovoid-pyramidal; styles slightly longer than ovary, proximally suberect, distally divergent-incurved; stigmas slightly enlarged. Capsule and seeds not seen. In scrub paramo; 3300 m. Ecuador (Morona-Santiago). ECUADOR. Morona-Santiago: Paramo de Matanga, Km 36 on road Sigsig-Gualaquiza (old muletrack), E. of the pass, 3300 m, 14 December 1980 (fl), Holm-Nielsen, Jaramillo & Coello 29401 (AAU). H. matangense is known as yet from only one collection, but it is clearly related to H. phellos subsp. marcescens from northern Colombia (Santander) although more specialized in all characters. Thus the leaves are much narrower and involute-twisted, not plane to apically subconcave, and distinctly discolorous; and the bases are loosely appressed to the stem. The flowers are solitary (in the single known specimen). 8. Hypericum garciae Pierce (Part 7: 26; No. 4). 9. Hypericum acostanum N. Robson (Part 7: 27; No. 5). 10. Hypericum piriai Arechav. (Part 7: 28; No. 6). Icon: Lyman B. Smith in/. Wash. Acad. Sci. 48: 312, f. In-p (1958); Rodriguez Jimenez in Reitz, Fl. III. Catar., Hiper- icdc.:6, t. IB (1980). 11. Hypericum simonsii N. Robson (Part 7: 34; No. 10). N.B. The type locality of this species is Maldonado, not Mardonado as in Part 7: 28-29. 12. Hypericum papillosum N. Robson (Part 7: 34; No. 11). 13. Hypericum callacallanum N. Robson, sp. nov. Fig. 7A, Map 2. Map 2 Sect. 29: 13. H. callacallanum O; 14. H. humboldtianum • H. simonsii N. Robson affinis, sed habitu humiliore, foliis angustioribus apice acutis, e basi uninervatis costa media fere in baso et supra ramosa, sepalis angustioribus apice acutis, petalis minoribus, stylis brevioribus, capsula minore, differt. Type: Peru, Amazonas, Chachapoyas, Cerros Calla-Calla, 29-30 km above Leimebamba on road to Balsas, Km 401, 3450-3500 m, 16 October 1964 (fl), Hutchinson & Wright 7003 (K!-holotype; F!, MICH!, MO!, NY!, US!-isotypes). HYPERICUM 19 Fig. 7 A. H. callacallanum: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut away) and ovary; (g) capsule. B. H. asplundii: (h) habit; (i) stem base with roots; (j) stem with leaves; (k) leaf; (1) sepal; (m) petal; (n) stamens (partly cut away) and ovary; (o) young capsule (a, h x Vz; b, i, j, x 2; c-g, k-o x 4). A. Hutchinson & Wright 7003; B. Asplund 17089. 20 N. K. B. ROBSON Shrub 0-2-0-5 m tall, erect, bushy, with branches strict to ascending, pseudo-dichotomous and lateral. Stems orange- brown, 4-lined but scarcely ancipitous when young, even- tually subterete, cortex exfoliating in strips; internodes 2-8 mm long. Leaves sessile, spreading from above base to appressed, persistent until brown, sometimes subsequently deciduous above base; lamina 8-13 x 2-3 mm, very narrowly oblong to very narrowly elliptic, plane or slightly condupli- cate, not carinate or glaucous, concolorous, chartaceous; apex acute, base narrowly cuneate to parallel, not sheathing, united with stem-line apex to form narrow interfoliar ridge; basal vein 1 with 2(3) basal or near-basal branches, midrib branching above, tertiary reticulation not visible; laminar glands rather dense, impressed above, sparser beneath. Inflorescence 1 -flowered, terminal and on short lateral or pseudo-dichotomous shoots; pedicel 3-6 mm long; upper leaves not transitional. Flowers c. 20 mm in diam., stellate. Sepals 6-8 x 1-3-2 mm, rather broadly to narrowly oblong or elliptic-oblong to oblanceolate, acute, distally curved; veins 5, unbranched, midrib not prominent, glands striiform and punctiform. Petals bright yellow, 9-5-11 x 3-6-4-5 mm, c. 1-5 x sepals, obovate-oblanceolate; apiculus acute; glands striiform and punctiform. Stamens 50-60, longest 6-7 mm long, c.0-7 x petals. Ovary c.2-5 x 1-8 mm, ovoid-ellipsoid; styles 3, c.3 mm long, 1-2 x ovary, suberect to spreading, distally incurved; stigmas narrowly capitate. Capsule c. 5 x 2 mm, ovoid-cylindric, shorter than sepals. Seeds c. 1 mm long, not or slightly carinate; testa very finely scalariform- reticulate. In open exposed habitats ('cold swamp' - Wurdack); 3000- 3700 m. Peru (Amazonas). Apparently confined to the Cerros Calla- Calla region. PERU. Amazonas: Chachapoyas, summit of Cerros Calla- Calla between Leimebamba-Balsas road pass and Herredura road, 3500-3700 m, 8 July 1962, (fl & fr), Wurdack 1230 (NY, US); Chachapoyas, Balsas-Leimebamba road, KM 400, 3200 m, 4 June 1977 (fl), Boeke 1920 (BM, NY n.v.); Chacha- poyas, Piscohuanuna Pass, 3000 m, August 1938 (fl), Sande- man 19 (K). H. callacallanum, along with 14. H. humboldtianum, forms a sister group to Spp. 15-21 and is more westerly in origin than the latter. It is markedly disjunct from its nearest Colombian relative in that group, which is in Boyaca (15. H. gladiatum). 20. H. asplundii, from Ecuador, is intermediate geographi- cally between these taxa; but it is morphologically more advanced in most characters than H. callacallanum and is more closely related to H. gladiatum. For the differences between H. callacallanum and H. humboldtianum see p. 21. 14. Hypericum humboldtianum Steudel, Nomencl. hot. 2nd ed. 1: 788 (1840). Type: Colombia, Cundinamarca, prope Santa Fe de Bogota?, June-September 1801 (fl & fr), Humboldt & Bonpland (P-HUM!-holotope; P-isotype; US!-photograph). Map 2. H. thymifolium Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 5: 186, t. 455 (1822); Choisy in DC., Prodr. 1: 550 (1824); Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 297 (1862), pro parte excl. syn. Trevir.; R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. w Torrey hot. Club 56: 103 (1929); non Banks & Sol. (1794). Type as for H. humboldtianum. Brathys thymifolia (Kunth) Spach, Hist. nat. veg. 5: 448 (1836), in Annls sci. nat. (Bot.) II, 5: 366 (1836). H. jussiaei Planchon & Linden ex Triana & Planchon in Annls. Sci. nat. (Bot.) IV, 18: 297 (1862); R. Keller in Bull. Herb. Boissier II, 8: 178 (1908); in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 103 (1929). Types: Colombia, Cundi- namarca, Cordillera de Bogota, 2900 m, 1855 (fl), Triana s.n. (PMectotype); Cundinamarca, paramo de Fortunate, 2870 m, 1842? (fl), Linden s.n. (BR, G-syntypes). H. jussiaei var. saturejifolium R. Keller in Bull. Herb. Boissier II, 8: 181 (1908) ['satureiaefolium']. Type: Col- ombia, Cundinamarca, Cordillera de Bogota, 3000 m, [1855?] 1851-57 (fl), Triana & Linden s.n. (W!-holotype; K!,NY!-isotypes). H. dichotomum Kunth ex R. Keller in Bot. Jb. 42: 130 (1908). Type: Ecuador, Loja, Saraguro, July-August (1802), Humboldt (B-holotype; F!, US! -photographs). Icon: Humb., Bonpl. & Kunth, Nov. gen. sp. 5: t.455 (1822). Shrub or shrublet (0-05-) 0-1-0-75 m tall, erect or decumbent at the base to ascending, with branches pseudo-dichotomous and divergent or lateral and strict, sometimes condensed in leaf axils, not from the base. Stems orange-brown, 4-lined and ancipitous when young, eventually terete, cortex exfoliating in strips; internodes 2-5-14 mm long. Leaves sessile or shortly pseudopetiolate with pseudopetiole up to 0-5 mm long, spreading from above pseudopetiole to subappressed, not tetrastichous, deciduous above base or above pseudopetiole before fading; lamina 6-13 x 1-2-3 mm, narrowly oblong or narrowly elliptic to oblanceolate, plane or ± incurved, not or slightly cucullate, midrib slightly prominent to impressed beneath, concolorous, sometimes glaucous, coriaceous or subcoriaceous; apex acute to obtuse, not mucronate, base angustate, the pseudopetiole sheathing, free or united with stem-line apex to form narrow interfoliar ridge; basal vein 1, without or with obscure lateral branches, tertiary reticulation absent; laminar glands ± dense, impressed above. Inflor- escence l(3-12)-flowered, the branching dichasial or pseudo- dichotomous or mixed; peduncle and pedicels 3-4(-7) mm long, not incrassate upwards; upper leaves transitional. Flowers c. 15 mm in diam., stellate. Sepals 14-5-16-5 x 1-3- 2-2 mm, elliptic-oblong to oblanceolate, acute; veins 3-5, not branched, midrib not prominent; glands linear below and punctiform in upper lA-l/2 or wholly punctiform. Petals bright HYPERICUM yellow, 6-8 x 3-5-5 mm, c. 1-3 x sepals, obovate; apiculus obtuse; glands linear, distally punctiform. Stamens 30-50, longest c.5-5 mm long, 0-6-0-7 x petals. Ovary 1-5-2 x 1 mm, ovoid to cylindric; styles 3, 2-3 mm long, 1-1-5 x ovary, narrowly spreading; stigmas broadly capitate. Capsule 4-6-5 x 2-3 mm, cylindric-ellipsoid, equalling sepals. Seeds 0-5-0-6 mm long, ecarinate; testa finely reticulate-scalariform. Thickets and open slopes of the paramo; (1820) 2500-3500 m. Colombia (Valle de Cauca to Norte de Santander), Vene- zuela (Merida). COLOMBIA. Antioquia: San Jose de Cuerquia, 30 July 1958 (fl), de Garganta 2145 (US). Boyaca: Municipio de Tola, peninsula de Suse, alredores de Lago de Tola, 3050 m, January 1976 (fl), Aguirre & Rangel 411 (COL). Caldas: Cordillera Central, old Quindio trail, 'Laguneta' to 'Magana', 3000-3200 m, 1 August 1922 (fl), Killip & Hazen 9463 (GH, K, NY). Cauca: environs of Purace, Tablon, 2700-2800 m, 22 July 1943, Cuatrecasas 14533 (F). Cundinamarca: Paramo de Guasca, 2940 m, 15 December 1938 (fl), Balls 5701 (BM, COL, F, K); Bogota to Choachf, Km 15, Paramo de Cruz Verde, 3000 m, 9 August 1974 (fl), Grabandt & Idrobo 317 (COL). Norte de Santander: Paramo de Fontibon, 2600-2750 m, 15 October 1941 (fl), Cuatrecasas, Schultes & Smith 12283 (COL); environs of Pamplona, 10 October 1963 (fl), de Garanta 625 (F). Valle: Cordillera Central, W. slope, no Bugalagrande, Barragan, cerro de La Laguna, 2920-2950 m, 17 April 1946 (fl), Cuatrecasas 20836 (F). VENEZUELA Merida: between La Cumbre, San Jose and Mucutuy, 1820-2590 m, 3 May 1944 (fl & fr), Steyermark 56258 (F, NY). H. humboldtianum varies continuously north-eastward from typical '//. thymifolium' (Cundinamarca, Cauca, Valle), with spreading leaves, no axillary branchlets, few or no lateral branches, and a 1-flowered pseudo-dichotomous inflor- escence, to '//. jussiaei var. saturejifolium' (Cundinamarca to Merida), with appressed leaves, axillary branchlets, numer- ous lateral branches, and 3-12-flowered dichasial inflor- escences. Its nearest relative would appear to be H. callacallanum (Peru), which is more bushy and has 3(4)-nerved sessile leaves and larger flowers. Kunth's H. dichotomum, was based on a Humboldt specimen that is labelled as having been collected in 'Sara- guru' (i.e. south Ecuador, Loja, Saraguro). This name remained unpublished (probably because Kunth realized that it would be a later homonym of H. dichotomum Lam.) until Keller discovered a specimen so named in Herb. Humboldt (B) and described it. The photographs of this specimen show that it clearly belongs to the '//. jussiaei' form of H. humboldtianum, not (as the cited locality suggests) to near H. callacallanum. Since no further material of this species has been collected in the botanically relatively well known Saraguro area, and since the species is otherwise unknown south of south-central Colombia, it seems possible that Keller was in error in localizing the Berlin specimen and that the plant was actually collected in Colombia by Humboldt & Bonpland. At any rate, the occurrence of H. humboldtianum in Ecuador must, for the present, remain extremely doubtful. 15. Hypericum gladiatum N. Robson, sp. nov. Map 1. H. simonsii N. Robson affinis, sed foliis angustioribus 1- vel 21 raro 3-nervatis apice acutis, sepalis brevioribus angustioribus- que apice acutis, staminibus paucioribus (circa 60), stylis brevioribus (circa 3 mm longis), differt. Type: Colombia, Boyaca, Arcabuco, alredores de la poblacion, 2739-2850 m, 20 October 1965 (fl & fr), Huertas & Camargo 6294 (COL!- holotype). Shrub, erect, with branches strict, lateral with elongate extension shoots and short determinate laterals, both types flowering, the short shoots occasionally branched pseudo- dichotomously. Stems orange-brown?, 4-lined and ancipitous when young, soon 2-lined, eventually terete, cortex exfoliat- ing in strips; internodes 3-6 mm long. Leaves sessile, appressed to ascending, markedly tetrastichous, persistent (deciduous with cortex); lamina 7-10 x 1-5-2-5 mm, linear- lanceolate to linear-oblong, plane to loosely incurved, sub- cucullate, not or slightly carinate, concolorous, not glaucous, subcoriaceous; apex acute, base parallel to rounded, not sheathing, free; basal veins 1(3), unbranched or with 1-2 pairs of ascending near-basal lateral branches, tertiary reticulation not visible or absent; laminar glands rather sparse, not prominent. Inflorescence 1-flowered, occasionally with pseudo-dichotomous branches from node below; pedicel 3-4 mm long, not or scarcely incrassate upwards; upper leaves foliose. Flowers not observed when expanded. Sepals 5-5-7 x 1-5 mm, narrowly lanceolate, acute, subcucullate; veins 5, laterals branching above, with midrib slightly prominent below; glands linear, distally sparsely punctiform or absent. Petals c.8 x 2-5 mm, c. 1-3 x petals, oblanceolate, apiculus ?; glands?. Stamens c. 60, longest 5-5-6 mm long, c. 0-7 x petals. Ovary not seen; styles 3, c. 3 mm long; stigmas narrowly capitate. Capsule c. 4 x 2-5 mm (? immature), ovoid, shorter than sepals. Seeds not seen. Habitat details unknown; 2739-2850 m. Colombia (Boyaca). COLOMBIA. Boyaca: Arcabuco, 2739-2850 m, 20 Octo- ber 1965 (fl & fr), Huertas & Camargo 6294 (COL). H. gladiatum has been collected only once and the label bears no habitat notes. Nevertheless, it is clearly distinct from all other known species, being intermediate in morphology and 22 N. K. B. ROBSON distribution between 1 1 . H. simonsii (Sierra Nevada de Santa Marta, NE. Colombia) on the one hand, and 16. H. roraimense (Roraima massif) and 20. H. asplundii (central Ecuador) on the other. It has smaller leaves and flowers than H. simonsii and also differs from it in the narrow sword- shaped leaves and sepals, the leaves being l(3)-nerved. For differences from H. roraimense and H. asplundii, see pp. 22 (opposite) and 24. 16. Hypericum roraimense Gleason in Bull. Torrey hot. Club 56: 399 (1929). Type: Guyana, Mt. Roraima, summit, 2595 m, 27 November 1927 (fl), Tate 412 (NY!-holotype; K!- isotype). Map 1. OO Shrub 0-14-0-5 m tall, erect, bushy, rounded, with branches strict, pseudo-dichotomous and lateral. Stems orange-brown, 4-lined and ancipitous when young, eventually terete, cortex flaking irregularly; internodes 1-1-5 mm long. Leaves sessile, spreading from above base, suberect, not tetrastichous, becoming brown and deflexed, persistent until cortex is shed; lamina 6-13 x 1-1-8 mm, very narrowly elliptic or very narrowly oblong to linear, plane, midrib slightly prominent beneath, concolorous, not glaucous, chartaceous; apex acute, base parallel-sided or slightly narrowed above insertion, loosely sheathing, free; basal vein 1, unbranched; laminar glands dense, ± large, prominent beneath. Inflorescence 1- flowered, terminal and on lateral branches, with branching pseudo-dichotomous; pedicel c. 1-5 mm long; upper leaves not transitional. Flowers c. 10 mm in diam., stellate ? Sepals 3-7 x 0-5-1-5 mm, narrowly lanceolate, acute; veins 5, unbranched, midrib rather prominent; glands linear. Petals bright (?) yellow, 5-9 x 2-3 mm, 1-3-1-7 x sepals, oblanceolate-oblong; apiculus acute; glands linear, inter- rupted distally. Stamens c. 25, longest 4-5 mm long, c. 0-5-0-8 x petals. Ovary 1-7x1 mm, narrowly ovoid; styles 3, 4-4-6 mm long, 2-5-2-8 x ovary, erect; stigmas broadly capitate. Capsule c. 5 x 3 mm, ellipsoid, shorter than sepals. Seeds not seen. In swamps; 2595-2880 m. Confined to the summit of the Roraima massif (Guyana, Venezuela). VENEZUELA. Bolivar: summit of Mt. Roraima, S. half, between Summit Camp, Great Central Rift, Central Swamp, and pond at S. end, 2700-2740 m, 28 September 1944 (fl), Steyermark 58898 (K, S, VEN). GUYANA. Mt. Roraima, September-December 1931 (st), Abbensetts 14 (K). H. roraimense, although quite isolated both morphologically and geographically, is most closely related to 15. H. gladiatum. It resembles that species in having long narrow leaves, but these are thinner, narrower, and broader at the middle if at all and become deflexed and marcescent. It would appear to be much smaller in stature and more clumped in habit than H. gladiatum, but has longer styles and broader stigmas. 17. Hypericum sp. Map 1. Shrublet 0-24 m tall, erect from creeping (?) and rooting base, bushy, with branches strict, lateral. Stem with cortex flaking irregularly; internodes 1-2 mm long. Leaves sessile, erect to spreading from above base, not tetrastichous, persistent until cortex is shed; lamina 3-4-5 x 0-5-0-8 mm, oblong-linear, plane to subincurved, slightly cucullate, midrib not or scarcely prominent beneath, concolorous, not glaucous, chartaceous; apex apiculate to obtuse, base parallel-sided, sheathing, free, basal vein 1, unbranched; laminar glands dense, large, prominent beneath. Flowers and fruit not seen. On steep stony slopes; 3400-4500 m. Venezuela (Merida). VENEZUELA. Merida: Paramo de Los Leones, 3400- 4500 m, 31 May 1930 (st), Gehringer 149 (VEN). Although clearly related to H. roraimense, this species is not yet represented by complete enough material to name and describe formally. It is smaller than H. roraimense in all its parts. 18. Hypericum espinalii N. Robson, sp. nov. Map 3. H. gladiato N. Robson affinis, sed foliis angustioribus 1- nervis, in media vel super mediam latioribus, ramificatione inflorescentiae mixta pseudodichotoma cymosaque, floribus minoribus, inter alia differt. Type: Colombia, Cauca, Totoro, 2500 m, 21 October 1968 (fl & e. fr), Espinal & Ramos 3055 (BM!-holotype; MO, VALLE). Shrub c. 0-3 m tall, erect, with branches ± strict, lateral. Stems orange-brown?, 4-lined and ancipitous when young, soon 4-angular, eventually terete, cortex exfoliating in strips; internodes 3-5 mm long. Leaves sessile, appressed to ascend- HYPERICUM Map 3 Sect. 29: 18. H. espinalii D; 19. H. castellanoi •; 20. H. asplundll A; 21. H. radicans A; 22. H. paramitanum •; 29. H. woodianum T; 39. H. baccharoides O. ing, not markedly tetrastichous, persistent (deciduous with cortex); lamina 8-13 x 1-2 mm, linear-oblanceolate, plane or distally conduplicate, not or slightly cucullate, subcarinate, concolorous, not glaucous, subcoriaceous; apex narrowly acute to shortly acuminate with terminal hydathode?, base parallel-sided to subcuneate, not sheathing, free or united with stem-line apex to form narrow interfoliar sheath; basal vein 1 , apparently unbranched or with 1-2 pairs of ascending lateral branches, tertiary reticulation not visible or absent; laminar glands dense, prominent. Inflorescence 1-4 (-7)- flowered, mixed unequally pseudo-dichotomous and (dicha- sial) monochasial, from terminal node only; peduncle and pedicels 2-2-5 mm long, not incrassate upwards; upper leaves foliose or (in cymose parts) reduced. Flowers 6-7 mm(?) in diam., stellate. Sepals 3-3-5 x 0-7-0-8 mm, narrowly triangular-lanceolate, acute, not cucullate; veins 5, un- branched, midrib or all prominent; glands linear. Petals c. 5 x 1-5-2 mm, c. 1-35-1-65 x sepals, oblanceolate, apiculus acute; glands punctiform. Stamens c.25-30, longest 3 mm long, c. 0-6 x petals. Ovary 2 x 0-7 mm, narrowly ellipsoid; styles 1-5 mm long, 0-75 x ovary; stigmas broadly capitate. Capsule (immature) 3-5 x 2 mm, ellipsoid. Seeds not seen. Habitat details unknown; 2500 m. Colombia (Cauca). COLOMBIA. Cauca: Totoro, 2500 m, 21 October 1968 (fl & fr), Espinal & Ramos 3055 (BM). 23 H. espinalii has been collected only once; but, like several of its near relatives (Spp. 15, 17, 19-21), it is morphologically and geographically isolated and cannot be accommodated in any other species. Its 1-nerved leaves resemble those of 20. H. asplundii apart from the prominent glands, but the erect habit (branches not rooting), mixed inflorescence, and smaller flowers distinguish it. 19. H. castellanoi is slenderer in habit with longer, narrower leaves and smaller flowers. 19. Hypericum castellanoi N. Robson, sp. nov. Map 3. H. espinalii N. Robson affinis, sed habitu diffuse ramis basi radicantibus, foliis linearibus incurvatis uninervis nervo haud ramoso, floribus laxe aggregatis, sepalis angustioribus stylis brevioribus, differt. Type: Venezuela, Merida, camino Merida-San Jose, 2360 m, 30 September 1970 (fl), Castellano- Monasterio 80 (VEN!-holotype). Shrub or shrublet 0-15-1 m tall, erect to ascending, with branches ascending, lateral, rooting at the base. Stems orange-brown, 4-angled when young, eventually 2-lined to subterete, cortex eventually exfoliating in strips; internodes 3-6 mm long. Leaves sessile, spreading or outcurving from base, twisting, not tetrastichous, persistent; lamina 12-17 x 0-7-1-2 mm, linear, incurved at first, becoming plane, not cucullate or carinate, concolorous, not glaucous, chartaceous; apex acute, base parallel-sided, not sheathing, free; basal vein impressed beneath, unbranched; laminar glands almost 2-ranked, visible on both sides, not or scarcely impressed. Inflorescence 1 -flowered, terminal and on \-b short lateral branches from c. 3-5 nodes below; pedicel 1^ mm long, incrassate upwards; upper leaves foliose. Flowers c. 8 mm in diam.?, stellate. Sepals 3-5-5 x 0-5-0-6 mm, linear, acute; veins 3, unbranched, not or slightly prominent. Petals bright ? yellow, 6-7 x 1-5-2 mm, c. 1-5 x sepals, narrowly oblan- ceolate, apiculus apiculate; glands linear, interrupted distally. Stamens c. 20-24, longest 3-4 mm long, c. 0-5 x petals. Ovary 1-5-2-5 x 1-1-5 mm, narrowly ovoid-ellipsoid; styles 3(4), 2- 3-5 mm long, c. 1-4 x ovary, proximally suberect, distally spreading; stigmas broadly capitate. Capsule and seeds not seen. Open places among shrubs or in forest; 2300-2360 m. 24 N. K. B. ROBSON Colombia (Boyaca), Venezuela (Merida). COLOMBIA. Boyaca: Rio Pomeca valley below Arca- buco, 2300 m, 7 December 1985 (fl), Wood 5202 (K). VENEZUELA. Merida: Camino Merida - San Jose, 2360 m, 30 September 1970 (fl & fr), Castellano-Monasterio 80 (VEN). H. castellanoi is related to 18. H. espinalii but has a more slender spreading habit with much narrower leaves. The two collections, which are from localities some 500 km apart, are not identical. The Colombian one is from an undershrub up to 1 m tall with dull leaves, the styles being c. 2 mm long with broadly capitate stigmas and the ovary c. 1-5 x 1 mm. The Venezuelan specimen is only 0-15 m tall with sublucent leaves, the styles being c. 3-5 mm long with rather narrowly capitate stigmas and the ovary c. 2-5 x 1-5 mm. Only further collecting will reveal if these differences are significant. 20. Hypericum asplundii N. Robson, sp. nov. Fig. 7B, Map 3. A H. gladiato N. Robson affinis, sed habitu humiliore diffu- sioreque, foliis linearibus, floribus nonnihil minoribus, stig- matis latioribus, differt. Type: Ecuador, Pichincha, NW. slope of Mt. Corazon, c. 3400 m, 27 July 1955 (fl), Asplund 1709 (SNholotype). Shrublet c. 0-22 m tall, diffuse, with branches erect or ascending from rooting base, lateral and pseudo-dichot- omous. Stems reddish-brown, 4-lined and ancipitous when young, eventually terete, internodes 4-8 mm long; cortex exfoliating in strips. Leaves sessile, ascending to spreading from above base, persistent until brown, sometimes subse- quently breaking off above base; lamina 7-13 x 1-3-1-8 mm, linear, plane, apically subconcave, not carinate or glaucous, concolorous, subcoriaceous; apex acute, base parallel or slightly broadened, not or scarcely sheathing, free or united with stem-line apex to form very narrow interfoliar ridge; basal veins 1(3), unbranched, tertiary reticulation not visible; laminar glands rather dense to sparse and not impressed above, sparse or absent beneath. Inflorescence 1(2-3)- flowered, terminal and 1-flowered on short lateral or on pseudo-dichotomous branches; peduncle and pedicels 2-4 mm long; upper leaves foliose, bracts smaller. Flowers 12-15 mm in diam., stellate. Sepals 6-7 x 1-2-1-5 mm, narrowly oblong-lanceolate, acute, cucullate; veins 3-5, branched, midrib not prominent; glands linear to striiform. Petals bright yellow, tinged red in bud, 9-10 x 4-5 mm, c. 1-3 x sepals, obovate, apiculus obtuse; glands linear to striiform. Stamens 35-50, longest 5-6-5 mm long, c. 0-6 x petals. Ovary 2-5 x 1-5 mm, ovoid; styles 3, 2-2-5 mm long, 0-8-1 x ovary, outcurving; stigmas broadly capitate. Capsule (mature) and seeds not seen. At the edge of thickets; c. 3400 m. Ecuador (Pichincha). Only one specimen has been seen. ECUADOR. Pichincha: NW. slope of Mt. Corazon, c. 3400 m, 27 July 1955 (fl), Asplund 17089 (S). H. asplundl is related to H. gladiatum but differs in the more diffuse pseudo-dichotomous branching, the decumbent root- ing stems, and the narrower, usually 1-nerved leaves. 21. Hypericum radicans N. Robson, sp. nov. Map 3. H. asplundio N. Robson affinis, sed habitu humiliori diffu- sion graciliori, foliis supra mediam latioribus, floribus minor- ibus, staminibus circa 2 mm longis, differt. Type: Colombia, Boyaca, no precise locality, 2700 m, 15 April 1964 (fl), Saravia 3938 (COL!-holotype). Shrublet with branches ascending from a rooting base, lateral. Stems orange-brown, 4-lined and ancipitous when young, soon terete, cortex exfoliating in strips; internodes 1-5-4 mm long. Leaves sessile, appressed to ascending, eventually outcurving, persistent; lamina 7-9 x 1-3-1-4 mm, linear- oblong or linear-elliptic to oblanceolate, plane, not cucullate, not carinate, concolorous, not glaucous, chartaceous; apex acute, slightly cucullate, margin plane, base narrowly cune- ate, not sheathing, free or almost so; basal vein 1, un- branched, tertiary reticulation absent; laminar glands dense, not prominent. Inflorescence 1-flowered, branching pseudo- dichotomous? Flowers c. 6 mm in diam. Sepals 3-5 x 0-9 mm, lanceolate, acute; veins 3, laterals branched above, all prominent; glands linear, distally punctiform. Petals c.4 x 2 mm, c. 1-2 x sepals, oblanceolate, apiculus obsolete; glands interrupted-striiform. Stamens ?, longest c. 2 mm long, c. 0-5 x petals. Ovary not seen; styles 3?, c. 1 mm long; stigmas broadly capitate. Capsule and seeds unknown. In mud; 2700 m. Colombia (Boyaca). COLOMBIA. Boyaca: no precise locality, 2700 m, 15 April 1964 (fl), Saravia 3938 (COL). Although clearly related to H. asplundii, H. radicans is smaller in all parts and more diffuse. It also is geographically isolated. HYPER/CUM 22. H. paramitanum N. Robson, sp. nov. Fig. 8, Map 3. H. terrae-firmae Sprague & Riley affinis, sed ramis junioribus valde ancipitis, foliis pinnatinerviis retinerviisque supra basin deciduis, stylis 3, inter alia differt. Type: Venezuela, Trujillo, between Jojo and La Morita, El Paramito, hacia Tuname, 3000 m, August 1958 (fl), Aristeguieta & Medina 3485 (VEN!- holotype; NY!-isotype). Shrub or small tree, 1-4 m tall, erect, with branches strict, pseudo-dichotomous or lateral. Stems orange-brown, 4-lined and markedly ancipitous when young, the lower end of the subfoliar lines projecting between the leaf-pair, eventually terete, cortex exfoliating in strips; internodes 1-5-4 mm long. Leaves sessile, spreading from near base, not tetrastichous, deciduous from near base without fading; lamina 12-23 x 5-8 mm, elliptic or oblong-elliptic to lanceolate, plane or sub- recurved, not cucullate, markedly carinate, concolorous, not glaucous, coriaceous; apex acute to obtuse, base cuneate, not sheathing, free; basal veins 1(3) with c. 5 pairs of ascending or spreading lateral branches, tertiary reticulation often marked, dense, ± impressed; laminar glands dense, obscure or invisible superficially, not prominent. Inflorescence 1(2-3)- flowered, with pseudo-dichotomous branches from node below; peduncle and pedicels 7-9 mm long, gradually incrassate upwards; upper leaves foliose. Flowers 25-30 mm in diam., stellate, petals becoming reflexed. Sepals 8-11 x 3- 4 mm, lanceolate, acute; veins 5-7, branching and anastomos- ing above, with midrib prominent; glands linear, distally punctiform. Petals (bright ?) yellow, 14-15 x 8 mm, c. 2 x sepals, oblanceolate; apiculus acute; glands linear, inter- rupted distally. Stamens 100-120, longest 9-10 mm long, c. 0-6 x petals. Ovary 3-4 x 2-5-3 mm, broadly ovoid; styles 3, 8-10 mm long, c. 2-5 x ovary, suberect; stigmas slightly enlarged. Capsule c. 7 x 3 mm, narrowly ovoid-ellipsoid, shorter than sepals. Seeds not seen. In subparamo and paramo; 2800-3150 m. Venezuela (Trujillo), known from only three collections. VENEZUELA. Trujillo: Paramo de Guaramacal, between Bocono and Guaramacal, 2800 m, 24 February 1971 (fl), 25 Steyermark 104824 (P, VEN); Paramo de Guirigay NE. of Guirigay, base of Piedras Blancas, c. 3150 m, 18 October 1972 (fl), Lopez-Flgueras & Rodriguez 8824 (VEN). H. paramitanum is an isolated relict species, differing from 11. H. simonsii, 12. H. papillosum, and 3. H. phellos in its thickly coriaceous leaves with pinnate and densely reticulate venation, deciduous above the base, and the peduncle and pedicels incrassate upwards. 23. Hypericum cuatrecasii Gleason (Part 7: 41; No. 16). 24. Hypericum goyanesii Cuatrec. (Part 7: 38; No. 14). 25. Hypericum myricariifolium Hieron. (Part 7: 40; No. 15). 26. Hypericum quitense R. Keller (Part 7: 41; No. 17). 27. Hypericum loxense Benth. (Part 7: 43; No. 18). 27a. subsp. aequatoriale (R. Keller) N. Robson; 27b. subsp. loxense. 28. Hypericum lycopodioides Triana & Planchon (Part 7: 36; No. 12). 29. Hypericum woodianum N. Robson, sp. nov. Map 3. H. lycopodioidei Triana & Planchon affinis sed foliis angus- tiore lanceolatis conduplicatis recurvis, aspectu acicularibus, floribus minoribus petalis luteis haud flavis, differt. Type: Colombia, Cundinamarca, between Guasca and Suera, 2900 m, 25 August 1985 (fl), Wood 5040 (K!-holotype; COL). Shrub ('undershrub') to 1-5 m tall, erect, with elongate branches strict, lateral, bearing numerous short ascending lateral branching, only the short shoots c. 3/5-4/s from the base flowering, not branching dichotomously. Stems orange- brown, 4-lined and ancipitous when young, soon 2-lined, eventually terete, cortex exfoliating in strips; internodes 5-6 mm long (long shoots) or c. 1 mm long (short shoots). Leaves sessile, squarrose (spreading above sheathing base), markedly tetrastichous, persistent and becoming deflexed; 26 N. K. B. ROBSON Fig. 8 H. paramitanum: (a) habit; (b) stem with leaf bases; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut away) and ovary; (g) young capsule (a x l/2; b-f x 3; g x 2). All Lopez-Figueras & Rodriguez 8824. HYPER1CUM 27 lamina 4-7 x 1-1-5 mm, lanceolate, markedly incurved, not cucullate, slightly carinate, concolorous, not glaucous, sub- coriaceous; apex acute, margin narrowly pellucid, base parallel to subamplexicaul, somewhat sheathing, united to form projecting interfoliar ridge; basal veins 1(3), un- branched or with 1-2 pairs of ascending near-basal lateral branches, tertiary reticulation obscure; laminar glands rather sparse, not prominent. Inflorescence 1 -flowered, on short lateral branches; pedicel almost absent; upper leaves nar- rowly triangular. Flowers c. 15-18 mm in diam., stellate? Sepals 5-6 x 2-2-5 mm, lanceolate to ovate-lanceolate, acuminate; veins 7, branching and anastomosing above, with midrib slightly prominent; glands linear, distally interrupted. Petals deep? yellow, c. 9 x 6 mm, c. 1-8 x sepals, obovate; apiculus acute; glands linear, distally interrupted to striiform. Stamens c. 75, longest c. 4-5 mm long, c. 0-5 x petals. Ovary c. 3 x 1-8 mm, ellipsoid; styles 3, 4-5 mm long, 1-5 x ovary, divergent from above base; stigma narrow. Capsule and seeds not seen. Roadside bank in cloud forest region; 2900 m. Colombia (Cundinamarca). COLOMBIA. Cundinamarca: Guasca to Suera, 2900 m, 25 August 1985 (fl), Wood 5040 (K). H. woodianum is related to the larger-leaved form of H. lycopodioides, differing from it in its narrow conduplicate recurved leaves, the whole shoot being reminiscent of a small Cryptomerla, and in its smaller flowers. The dried petals are darker yellow than those of H. lycopodioides . The locality of the sole collection of//, woodianum (central Cundinamarca), in conjunction with the morphological differences, suggests that it and H. lycopodioides are vicariant species. 30. Hypericum thuyoides Kunth (Part 7: 37; No. 13). 31. Hypericum sabiniforme Trevir. (Part 7: 45; No. 19). 32. Hypericum lark-Hoi HUM Juss. (Part 7: 47; No. 20). 33. Hypericum mar tense N. Robson (Part 7: 51; No. 21). 34. Hypericum hartwegii Benth., (Part 7: 51; No. 22). 35. Hypericum maguirei N. Robson (Part 7: 52; No. 23). Subsect. 3. Brathys Frutices vel fruticuli. Folia binatim conjuncta, sessilia vel plusminusve breviter pseudopetiolata, plana vel incurva vel incurvo-conduplicata, apice plana vel incurva vel interdum cucullata, 1-9-nervia venatione parallela vel flabellata vel pinnata vel simplici, supra basin decidua vel rarissime omnino persistentia et marcescentia (Sp. 36) vel nodis caulis disarticu- latis (Sp. 67). Inflorescentia 1-flora ramificatione pseudo- dichotoma vel laterali vel raro 2-13-flora ramificatione cymosa (Spp. 38, 39). Flores 5^0 mm in diametro, stylis 3 vel raro 4 (in floribus magnis) vel 4-5 (in floribus parvis). Typus: H. brathys Sm. nom. illegit. (= Brathys juniperina L. f. = H. juniperinum Kunth (Species 36-74). 36. Hypericum magniflorum Cuatrec. (Part 7: 53; No. 24). 37. Hypericum gleasonii N. Robson (Part 7: 55; No. 25). 38. Hypericum mexicanum L. (Part 7: 56; No. 26). 39. Hypericum baccharoides Cuatrec. in Brittonia 11: 165 (1959). Type: Colombia, Magdalena, Sierra de Perija, 48 km from Valledupar, 1 km from Venezuelan border, 3000 m, 5 February 1945 (fl), Grant 10842 (US!-holotype). Map 3. Shrub 0-5-1 m tall, erect, with branches strict, lateral or more rarely paired and pseudo-dichotomous. Stem orange-brown, 4-lined and slightly ancipitous when young, eventually terete, cortex exfoliating in strips or irregular flakes; internodes 1- 1-5 mm long. Leaves sessile, outcurving, ± closely imbricate, tetrastichous, deciduous at or just above the base without fading; lamina 4-8 x 3-5-7 mm, broadly elliptic to circular or obovate-spathulate, plane, not carinate, concolorous, undulate-papillose especially beneath, densely glaucous and (always?) viscid, coriaceous; apex broadly obtuse to rounded, base angustate, sheathing, free or united to form very narrow interfoliar ridge; basal veins 1-7, pinnate with ascending branches or flabellate, ultimate branching and tertiary reticu- lum not visible; laminar glands dense, slightly prominent; marginal glands at least sometimes secreting. Inflorescence 1- flowered, terminal and on lateral branches, occasionally with pseudo-dichotomous branches; pedicel absent; upper leaves transitional. Flowers 15-20 mm in diam., stellate. Sepals 5-7 x 3-8^-5 mm, ovate to subacute; veins 5-7, obscurely branched, with midrib scarcely prominent beneath; glands mostly linear, distally punctiform. Petals bright? yellow, 9-12 x 4-5 mm, c. 2 x sepals, oblong-ovate; apiculus acute; glands linear, distally punctiform. Stamens 70-120, 5-8 mm long, longest c. 0-65 x petals. Ovary l-5-2(-2-5) x 1-1-5 mm, broadly ovoid; styles 3, 3-5 mm long, 2-2-5 x ovary, suberect, outcurved distally; stigmas small. Capsule c. 6 x 5 mm, broadly ovoid, shorter than sepals. Seeds c. 1-3 mm long, ecarinate; testa finely scalariform-reticulate. In forest and paramo; 2700-3350 m. Colombia (Magdalena), Venezuela (Zulia); confined to the Sierra de Perija. COLOMBIA. Magdalena: Sierra de Perija, plain between Cerro Venado and Cerro Avion, 3270-3350 m, 8 November 1959 (fl), Cuatrecasas & Romero Castaneda 25119 (US); Sierra de Perija, east of Manaure, Quebrada de Florida- blanca, 2700-2800 m, 10 November 1959 (fr), Cuatrecasas & Romero Castaneda 25161 (US). VENEZUELA. Zulia: Sierra de Perija, 2800-2900 m, 29 December 1950 (fl), Gines 1993 (US); Perija, Cerro Cetari, April 1952 (fl), Urbano 15 (VEN). H. baccharoides at first glance seems very distinct from other species in sect. Brathys. Closer inspection, however, reveals that it is allied to 37. H. gleasonii, which also occurs in the Colombia- Venezuela border area but much further south. From H. gleasonii it differs in its smaller stature with shorter internodes, smaller and relatively broader leaves that are 28 N. K. B. ROBSON densely glaucous, smaller flowers, 5-7-veined sepals, shorter petals, smaller ovary, and narrower stigmas. 40. Hypericum stuebelii Hieron. (Part 7: 58; No. 27). 41. Hypericum prietoi N. Robson (Part 7: 58; No. 28). 42. Hypericum cassiopiforme N. Robson (Part 7: 59; No. 29). 43. Hypericum decandrum Turcz. (Part 7: 59; No. 30). 44. Hypericum pimeleoides Planchon & Linden ex Triana & Planchon (Part 7: 60; No. 31). 45. Hypericum magdalenicum N. Robson (Part 7: 63; No. 32). 46. Hypericum valleanum N. Robson (Part 7: 64; No. 33). 47. Hypericum spruce! N. Robson (Part 7: 65; No. 34). 48. Hypericum aciculare Kunth (Part 7: 66; No. 35). 49. Hypericum recurvum N. Robson (Part 7: 68; No. 36). 50. Hypericum wurdackii N. Robson (Part 7: 69; No. 37). 51. Hypericum costaricense N. Robson (Part 7: 70; no. 38). 52. Hypericum bryoides Gleason (Part 7: 71; No. 39). 53. Hypericum bolivaricum N. Robson (Part 7: 72; No. 40). 54. Hypericum juniperinum Kunth (Part 7: 72; No. 41). 55. Hypericum prostratum Cuatrec. (Part 7: 75; No. 42). 56. Hypericum parallelum N. Robson (Part 7: 76; No. 43). 57. Hypericum marahuacanum N. Robson (Part 7: 77; No. 44). 57a. subsp. marahuacanum; 57b. subsp. strictissimum N. Robson; 57c. subsp. chimantaicum N. Robson. N.B. The type of 57c is Steyermark & Wurdack 758, not 7581 as cited in Part 7: 80. 58. Hypericum lancifolium Gleason (Part 7: 80; No. 45). 59. Hypericum horizontale N. Robson (Part 7: 81; No. 46). 60. Hypericum let rustic hum Cuatrec. (Part 7: 82; No. 47). 61. Hypericum strictum Kunth (Part 7: 83; No. 48). 61a. subsp. strictum; 61b. subsp. compactum (Triana & Planchon) N. Robson. 62. Hypericum jaramilloi N. Robson (Part 7: 86; No. 49). 63. Hypericum cardonae Cuatrec. (Part 7: 87; No. 50). 64. Hypericum caracasanum Willd. (Part 7: 89; No. 51). 64a. subsp. caracasanum; 64b. subsp. turumiquirense (Steyerm.) N. Robson. 65. Hypericum ekmanii A. H. Liogier (Part 7: 90; No. 52). 66. Hypericum pycnophyllum Urban (Part 7: 92; No. 53). 67. Hypericum millefolium Urban & Ekman (Part 7: 93, No. 54). " 68. Hypericum ruscoides Cuatrec. (Part 7: 93; No. 55). 69. Hypericum harlingii N. Robson, sp. nov. Map 4. H. ruscoidei Cuatrec. affinis, sed foliis minoribus incurvatis Map 4 Sect. 29: 69. H. harlingii D; 75. H. cymobrathys (modified from Part 7, p. 54) O; 76a. H. chamaemyrtus subsp. chamaemyrtus •; 76b. H. chamaemyrtus subsp. pseudocaracasanum •. apice acuminatis, floribus minoribus, sepalis lineari-ellipticis acutis vel subacuminatis, 3-nervatis, stigmatibus angustis, inter alia differt. Type: Ecuador, Loja, Cerro Toledo, 10-12 km SE. of Yangana, 3000-3200 m, 6 April 1985 (fl), Marling & Andersson 23776 (GBl-holotype; BM!-isotype). Shrub 0-5 m tall, erect, branches strict, pseudo-dichotomous and lateral. Stems orange-brown, 4-lined and ancipitous when young, eventually terete, cortex exfoliating in strips; interno- des 2-3 mm long. Leaves petiolate with petiole 1 mm long; lamina 4-6 x 2-3 mm, elliptic, margins incurved, not cucullate or carinate, dull green above, sublucent beneath, not glaucous, chartaceous; apex acuminate, base cuneate- angustate, not sheathing, united to form very narrow HYPER1CUM 29 interfoliar ridge; basal vein 1, impressed beneath, with 2-3 pairs of obscure ascending laterals loosely reticulating; laminar glands dense, visible on both sides, not impressed. Inflorescence 1 -flowered, terminal, repeatedly branching pseudo-dichotomously; pedicel 4 mm long; upper leaves foliose. Flowers 12-15 mm in diam., stellate. Sepals 4—5 x 1-2-1-5 mm, linear-elliptic, acute to subacuminate; veins 3, branching distally, becoming prominent. Petals bright ? yellow, 6-7 x 3 mm, c. 1-5 x sepals, narrowly oblong- oblanceolate; apiculus acute; glands linear, distally striiform. Stamens 35^10, longest 4-^-5 mm long, c. 0-65 x petals. Ovary 2-2-5 x 1-5 mm, broadly ovoid-ellipsoid; styles 3, 3 mm long, 1-3-1-5 x ovary, divergent-incurving; stigmas narrow. Capsule (old) c. 5 x 3 mm, ovoid. Seeds not seen. In montane forest; 3000-3200 m. Ecuador (Loja). ECUADOR. Loja: c. 8 km E. of Yangana, on footpath to Numbala, 3100 m, 31 January 1966 (fl), Knight 11 (WIS). Although H. harlingii is known by only two collections, it is quite distinct. Its nearest relative appears to be 68. H. ruscoides, from which it differs by the smaller, dull green, incurved leaves and the smaller flowers with narrower, 3- veined sepals and slender styles with narrow stigma. The most closely related form of H. ruscoides is not the nearest geographically (central Ecuador) but that in southern Col- ombia. 70. Hypericum llanganaticum N. Robson (Part 7: 95; No. 56). 71. Hypericum struthiolifolium Juss. (Part 7: 96; No. 57). Add first record for Ecuador: ECUADOR, Zamora-Chinchipe: road from Loja to Zamora, 2800 m, 18 September 1961 (fl), Dodson & Thien 685 (WIS pp.). The rest of the WIS material, as well as the BM specimen of this collection, belongs to H. sprucei. 12. Hypericum andinum Gleason (Part 7: 97; No. 58). 73. Hypericum lancioides Cuatrec. (Part 7: 98; No. 59). 73a. subsp. lancioides; 73b. subsp. congestiflorum (Triana & Planchon) N. Robson. 74. Hypericum selaginella N. Robson (Part 7: 102; No. 60). Subsect. 4 Spachium R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 3 (6): 214 (1893), pro parte, see Part 1: 338, excluding synonyms of Sect. 30. Trigynobrathys (Y. Kimura) N. Robson (p. 47). Frutices vel suffrutices vel fruticuli vel herbae perennes vel annuae. Folia libera, pseudopetiolata vel sessilia, plana vel margine recurva vel revoluta vel raro omnino recurva (Spp. 79, 85 p.p., 87), apice rarissime cucullata (Sp. 85 p.p.), 1-3- nervia vel raro 5-7-nervia (Sp. 75) venatione parallela vel pinnata vel simplici, supra basin decidua vel omnino persis- tentia sed rarissime marcescentia (Sp. 75 p.p.) Inflorescentia 1-40-flora ramificatione pseudo-dichotoma vel cymosa vel mixta vel rariori laterali. Flores 3-16 mm in diametro, stylis 3. Typus: H. gentianoides (L.) Britton, Sterns & Pogg. (Species 75-87). 75. Hypericum cymobrathys N. Robson (Part 7: 103; No. 61). Map 4. N.B. The leaves are free, not united as stated in Part 7: 103, and can be marcescent rather than deciduous above the base. 76. Hypericum chamaemyrtus Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 298 (1862); R. Keller in Bull. Herb. Boissier II, 8: 178 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925) ['chamaemyrtos']; Rodriguez Jimenez in Mems Soc. Cienc. nat. La Salle. 33: 16 (1973), in adnot.; non sensu Gleason in Bull. Torrey hot. Club 56: 102 (1929) = H. garciae Pierce. Type: Colombia, Cundinamarca, Gachala [Gacheta?] et Ubala, cordillere de Bogota, 1700 m, August 1855 (fl & fr), Triana 5464 (COL ?-holotype; P!-isotype; BM!, COL!, G!, W!; F!, GH!, NY!-photographs). Subshrub 0-5-1 m tall, erect or decumbent at the base; branches from near base or very rarely lateral, strict and sometimes ascending. Stems reddish-brown, 4-lined, mark- edly ancipitous and eglandular? when young, eventually terete, cortex exfoliating in strips; internodes 2-14(-20) mm long. Leaves pseudopetiolate with pseudopetiole up to 1 mm long or subsessile, spreading from base or outcurving to suberect, longer than internodes, deciduous above base without fading; lamina 8-22 x 1-5-11 mm, broadly ovate or elliptic to very narrowly elliptic-oblong or lanceolate or (lower) oblanceolate, margin ± recurved (plane in narrowest, young leaves), not cucullate, midrib ± prominent beneath or not, not glaucous, somewhat paler beneath, coriaceous; apex acute or apiculate to obtuse, base broadly cuneate or rounded to angustate, not sheathing, the pair free; basal veins 1(3), with or without \-^ pairs of main ascending lateral branches, tertiary reticulation obscure or absent; laminar glands dense, not prominent. Inflorescence 1- c. 30-flowered, branching dichasial/monochasial or pseudo-dichotomous/sympodial or mixed, occasionally with 1-2 pairs of subsidiary branches from node immediately below; peduncle and pedicels 3-8 mm long, not incrassate upward; bracts foliar, gradually reduced. Flowers 10-16 mm in diam., stellate. Sepals 4-9-5 x 1-2^4 mm, unequal, lanceolate or ovate-lanceolate or elliptic to oblong or oblanceolate, acute to acuminate; veins 3-5(7), not or distally branched, all becoming slightly prominent; glands all linear or punctiform in upper !/3-%. Petals bright? to pale yellow, 6-5-10 x 3-5-8 mm, 1-3-1-9 x sepals, obovate; apiculus acuminate to obtuse; glands linear, distally inter- rupted or punctiform. Stamens 35-80, longest 4-6 mm long, 0-5-0-75 x petals, fascicles not distinct. Ovary 2-3 x 1-1-5 mm, ellipsoid; styles 3, 2-5-3-5 mm long, 1-2-1-7 x ovary, spreading-incurving; stigmas ± broadly capitate to peltate. Capsule 5-7 x 2-5-4 mm, narrowly ovoid to cylindric- ellipsoid, shorter than sepals. Seeds 0-6-0-8 mm; testa finely scalarif orm-reticulate . In paramo and in dense woods and damp meadows below; 1700-3290 m. Colombia (Boyaca, Cundinamarca); Venezuela (Lara, Tru- jillo). H. chamaemyrtus is related to H. cymobrathys but is less woody, and the leaves usually have pinnate not parallel venation. Occasionally, however, they have 3 basal leaf veins. The pseudopetiolate leaf base also indicates the relationship with H. cymobrathys. The inflorescence, however, is usually laxer than in that species. The Sachica (Boyaca) population, with broad, sharply acute, 3-nerved leaves, is morphologi- cally closest to H. cymobrathys. 30 N. K. B. ROBSON H. chamaemyrtus occurs in two isolated populations. The Colombian one (from southern Boyaca to central Cundina- marca) is taller with inflorescence branches longer and ultimately dichasial/monochasial and leaves ovate to very narrowly elliptic-oblong, acute to obtuse, definitely pseudo- petiolate and eventually spreading; whereas the Venezuelan population is shorter, less richly branched above, with elliptic, usually mucronate, and often subsessile leaves that remain suberect, and the inflorescence branches are wholly pseudo-dichotomous. There is, however, a certain degree of overlap in variation between these populations that prevents their recognition as species. 76a. Hypericum chamaemyrtus subsp. chamaemyrtus Fig. 9A, Map 4. Subshrub 0-7-1 m tall. Leaves definitely pseudopetiolate, lamina broadly to very narrowly elliptic-oblong, base broadly cuneate to rounded, midrib not prominent beneath. Inflor- escence branching at first pseudo-dichotomous, ultimately dichasial/monochasial. Petals 5-8 mm long, apiculus acumin- ate to acute. Colombia (Boyaca, Cundinamarca). COLOMBIA. Boyaca: Arcabuco, alredores de la pobla- cion, 2739-2850 m, 20 October 1965 (fl), Huertas & Camargo 6265 (COL); Sachica, 1990 m, August 1962 (fl), Saravia 4349 (COL); above and immediately N. of Villa de Leiva, 2300 m, 8 December 1985 (fl & fr), Wood 5207 (K). Cundinamarca: Zipaquira, 'Lungazaque patreros', 2609 m, 17 January 1943 (fl), Huertas & Camargo 546 (F); Fuquene Lake, 24 March 1945 (fl), Schiefer 620 (GH, US); Ubate, 2600 m, 18 May 1952 (fl), K0ie 4517 (C); between Sisga and Macheta, 2800 m, 3 September 1983 (fl), Wood 3960 (K). Subsp. chamaemyrtus has two centres of variation: (i) the main Boyaca population (most similar to H. cymobrathys); (ii) the population in the Bogota area, whence there is a north-eastward trend in leaf shape towards the narrower leaves characteristic of the type. 76b. Hypericum chamaemyrtus subsp. pseudocaracasanum (Steyerm.) N. Robson, stat. nov. Fig. 9B, Map 4. H. pseudocaracasanum Steyerm. in Fieldiana Bot. 28 (2): 394 (1952). Type: Venezuela, Trujillo, La Quebrade Cortijo, by Lara-Trujillo boundary, above Humocara Bajo, 2600-2800 m, 6 February 1944 (fl), Steyermark 55346 (F!-holotype; NY!- isotype). Subshrub 0-5-0-7 m tall. Leaves subsessile, lamina ± nar- rowly elliptic, base narrowly cuneate to angustate, midrib ± prominent beneath. Inflorescence branching wholly pseudo- dichotomous. Petals 3-5-6 mm long, apiculus obtuse. Venezuela (Lara, Trujillo). VENEZUELA. Lara: W. of Humocara Bajo, above Los Aposentos, Las Sabanetas, 2530 m, 5 February 1944 (fl), Steyermark 55292 (F); Ditto. Monen, trail from Humocaro to Buenos Aires (Caserio) below Paramo Los Rosas, 2285-3290 m, 20 June 1979 (fl & fr), Liesner et al. 7987 (BM, MO, VEN). Trujillo; Paramo Agua de Obispo, 2500 m, 24 September 1922 (fl), Jahn 1182 (US, VEN). 77. Hypericum arbuscula Standley & Steyerm. in Publs Field Mus. nat. Hist. (Bot.) 23: 63 (1944); Standley & Williams in Fieldiana Bot. 24(7): 48 (1961). Type: Guatemala, Baja Verapaz, Patal, 4 September 1941 (fl & fr), /. R. Johnston 1812 (F!-holotype). Map 5. H. silenoides sensu Rodriguez Jimenez in C. r. Soc. Biogeogr. 432: 91, map 2 (1972), pro parte, in Mems Soc. Cienc. nat. La Salle 33: 51 (1973), pro parte quoad spec. Breedlove & Raven 8416. Subshrub or shrublet 0-25-0-45 m tall, with taproot, bushy to slender, erect or decumbent but not rooting; branches from all along stem or only from below inflorescence, strict. Stems green to reddish-brown, 4-lined, ancipitous and densely gland-dotted at first, eventually terete, cortex exfoliating in strips or irregularly; internodes 2-20 mm long. Leaves sessile, HYPERICUM 31 Fig. 9 A. H. chamaemyrtus subsp. chamaemyrtus: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) flower (petals and stamens partly cut away); (g) young capsule. B. H. chamaemyrtus subsp. pseudocaracasanum: (h) habit (a, h x '/z; b x 1; c, f x 2; d, e, g x 4). A. Wood 3960; B. Leisner et al. 7987. 32 N. K. B. ROBSON Map 5 Sect. 29: 77. H. arbuscula D; 78. H. rubritinctum T ; 79. H. fuertesii O; 82. H. gnidioides •; 83. H. eastwoodianum A ; 84. H. peninsulare V; 85. H. beamanii ^; 86. H. galinum •; 88. //. gentianoides (part) A. densely imbricate to outcurving or spreading, longer than internodes (sometimes except upper ones), persistent (i.e. deciduous with cortex); lamina 5-20 x 0-6-3-7 mm, lan- ceolate to linear, margin recurved to strongly revolute, entire, not or scarcely cucullate, midrib prominent and smooth beneath, paler beneath or subconcolorous, some- times glaucous above, subcoriaceous to coriaceous; apex subacute to acute, base shortly angustate to subamplexicaul, free; basal veins 1-3, unbranched or sometimes with 1 pair of ascending main lateral branches and rarely 1 pair of midrib branches; laminar glands dense, not prominent. Inflorescence 1-c. 40-flowered, branching mixed dichasial/monochasial and pseudo-dichotomous or repeatedly pseudo-dichotomous/ sympodial, without or with up to 6 -pairs of subsidiary branches; peduncle and pedicels 2-12 mm long, not incrassate upwards; bracts foliar or reduced. Flowers 6-22 mm in diam., stellate (? to infundibuliform). Sepals 4-5-10 x 0-8-3 mm, subequal or unequal, elliptic or linear-oblong or linear- lanceolate to oblanceolate, acute to acuminate; veins (3)5-7, unbranched, becoming prominent; glands linear, distally interrupted or punctiform. Petals bright (?) yellow, not tinged red, 5-10 x 1-4 mm, c. 1-1-1-3 x sepals, oblanceolate to narrowly oblong-spathulate; apiculus acute; glands linear, distally interrupted to punctiform. Stamens c. 40-45, longest 4-5-5 mm long, c. 0-5-0-75 x petals, fascicles not distinct. Ovary c. 1-5-1-8 x 0-8-1 mm, narrowly ovoid-ellipsoid to narrowly ellipsoid, trigonous; styles 3, 3-5-5 mm long, c. 2-3 x ovary, slender, spreading; stigmas narrowly capitate. Capsule 3-5 x 1-5-2 mm, narrowly ovoid to narrowly ellipsoid, trigonous, acute, shorter than sepals. Seeds c. 0-5-6 mm long, ecarinate; testa finely scalariform. Habitat not recorded; 1500 m. Mexico (Chiapas), Guatemala (Baja Verapaz, El Quiche). MEXICO: Chiapas, Municipio de Trinitaria, road to Lagos de Montebello 17-6 km NE. of La Trinitaria, 1590 m, 26 January 1965 (fl & fr), Breedlove & Raven 8416 (F, MICH); a 12 km E. de La Trinitaria por carretera a Montebello, 1570 m, 22 July 1984 (fl & fr), Hernandez & Chacon 544 (BM, MO). GUATEMALA. Baja Verapaz: Santa Rosa, 1500 m, July 1887 (fr), v. Turckheim 1314 (F, K, NY, P, US); Fatal to Salama, 4 November 1941 (fl & fr), /. R. Johnston 1811 (F). El Quiche: no precise locality, 1942 (fl), Aquilar773 (F). H. arbuscula, which typically does seem to merit its epithet (= 'little tree') by remaining unbranched until it produces the first flower, resembles a small, dense-leaved form of H. gnidioides; and the densely imbricate leaves also give it an ericoid appearance. But Johnson 1811, from near the locality of the type4 and with the previous number, has a decumbent stem with longer internodes, longer leaves, and mixed branching in the inflorescence. The Mexican (Chiapas) population has larger flowers and broader leaves than the Guatemalan ones, and the inflorescence branching is mixed. It is thus the most closely related species to 76. H. chamaemyrtus of all the Mexican and Central American members of subsect. Spachium and is also linked with 78. H. 4Standley & Williams (1961: 48) suggest that the type may have been collected from further south than Fatal, perhaps about Santa Rosa. If that was indeed so, it might explain the great similarity between the type and von Turckheim's Santa Rosa collection and the differences between them and Johnson 1811 from between Fatal and Salama. HYPER/CUM 33 rubritinctum and 82. H. gnidioides. This Chiapas population at first glance would seem to merit taxonomic recognition, at least as a subspecies; but Johnson 1811 is nicely intermediate between it and typical H. arbuscula, and so it seems best to maintain the latter species undivided, at least until more is know about the Mexican plants. H. arbuscula is related to both H. gnidioides and H. rubritinctum, differing from the former by the longer styles and from the latter by the smaller flowers. The derivative species related to H. arbuscula are all confined to the Greater Antilles (the H. diosmoides group, Spp. 79-81). 78. Hypericum rubritinctum N. Robson, sp. nov. Fig. 10A,Map5. H. arbuscula Standley & Steyerm. affinis, sed ramificatione inflorescentiae dichasiali vel monochasiali haud pseudo- dichotoma, bracteis bracteolisque lineari-subulatis, floribus maioribus, 13-20 mm in diam., sepalis oblongis vel lanceola- tis vel anguste ovatis, petalis 8-14 mm longis dorsaliter rubritincta, stylis brevioribus, inter alia differt. Type: Mex- ico, Guerrero, Minas Gro., Armenia, 2340 m, 23 October 1936 (fl), Hinton 9757 (US!-holotype; GH!, K!, MICH!, NY!, P!-isotypes). H. gnidioides sensu Rodriguez Jimenez in C. r. Soc. Biogeogr. 432: 91, map 5 (1972), pro parte, in Mems Soc. Cienc. nat. La Salle 33: 114 (1973), pro parte quoad spec. Hinton 7457, 9457. Shrublet 0-2-0-35 m tall, with taproot, fastigiate, erect; branches lateral and sometimes basal (or none), often opposite, strict. Stems reddish-brown, 4-lined, ± ancipitous and ± densely gland-dotted when young, eventually terete, cortex exfoliating in strips; internodes 3-20 mm long. Leaves sessile, erect or appressed, ascending or spreading, subimbri- cate, the upper longer than internodes, the lower shorter, persistent (i.e. deciduous with cortex) or breaking off earlier above base; lamina 6-15 x 1-2-3 mm, linear to narrowly elliptic-oblong, margin revolute, smooth, not cucullate, midrib prominent or not and smooth beneath, paler beneath, and often glaucous above or on both sides, coriaceous; apex subacute, base narrowly cuneate to parallel-sided, free; basal veins 1(3), unbranched, tertiary reticulation absent; laminar glands dense, not prominent or slightly so beneath. Inflor- escence (l)3-9-flowered, branching all dichasial/monochasial (or the lowermost pseudo-dichotomous), without subsidiary branches; peduncle and pedicels 2-4 mm long, not incrassate upwards; bracts and bracteoles linear-subulate. Flowers 13- 20 mm in diam., stellate. Sepals 3-5-4-2 x 1-1-5 mm, subequal, oblong to lanceolate or narrowly ovate, subacute; veins 5-7, unbranched, all becoming ± prominent; glands all linear or punctiform in upper !/3. Petals bright yellow, tinged red outside, 8-14 x 2-5^ mm, 2-3-5 x sepals, oblanceolate; apiculus obsolete; glands few, punctiform. Stamens c. 40, longest 4-5 mm long, c. 0-6 x petals, fascicles 3, subdistinct. Ovary c. 1-5 x 1 mm, narrowly ellipsoid; styles 3, 3-5-4 mm long, c. 2-2-5 x ovary, spreading; stigmas narrowly to rather broadly capitate. Capsule 4-6 x 2-3 mm, ovoid-cylindric, acute, shortly exceeding sepals. Seeds not seen. Rocky places in Quercus woodland and in Pinus forest; 1700- 2340 m. Mexico (Guerrero, Mexico). MEXICO. Guerrero: Minas Gro, Armenia, 2340 m, 23 October 1936 (fl), Hinton 9757 (GH, K, MICH, NY, P, US). Mexico: Temascaltepec, Cumbre, 27 March 1935 (fl & e. fr), Hinton 7457 (F, G, GH, K, MICH, MO, NY, US); Cerro de Ahuacatitlan, Almoylola de Alquisiras, 1700 m, 29-30 March 1954 (fr), Matuda 30605 (MEXU). H. rubritinctum was included in 82. H. gnidioides by Rodriguez Jimenez, but it differs from the latter especially in its longer slender styles (c.f. 77. H. arbuscula) and (both absolutely and relative to the sepals) larger petals, which are red-tinged in bud and become red-veined later. Its other close relative, H. arbuscula, has shorter internodes (particularly below the inflorescence), smaller flowers, and pseudo-dichot- omous/sympodial inflorescence branching. 79. Hypericum fuertesii Urban, Symb. antill. 7: 523 (1913); Mosc., Cat. Fl. doming. 1: 381 (1943). Type: Hispaniola, Dominican Republic, La Vega, Loma Rosilla, 2700 m, 6 July 1912 (fl & fr), Fuertes 1747 (Bt-holotype; BM!, C!, F!, G!, H!, K!, NY!, P!, S!, U!, US!, W!, Z!-isotypes). Fig. 10B, Map 5. Shrublet or wiry perennial herb 0-09-0-2 m tall and to c. 0-4 m long, with taproot, erect or decumbent and rooting, some- times mat-forming; branches from below inflorescence and sometimes lateral, strict or very rarely divaricate. Stems reddish-brown, 4-lined, ± ancipitous and densely gland- dotted when young, eventually terete, cortex exfoliating in 34 Fig. 10 A. H. rubritinctum: (a) habit; (b) stem with leaves; (c) leaf; (d) flower (petals and stamens partly cut away); (e) sepal; (f) petal. B. H. fuertesii: (g) habit; (h) stem with leaves; (i) leaf; (j) sepal; (k) petal; (1) stamens (partly cut away) and ovary; (m) young capsule (a, g x Vv, b, d X 2; c, e, f, h, 1 X 4; i-k, m X 8). A. Hinton 9757; B. Liogier 15969. HYPER/CUM 35 strips; internodes 1-5 mm long. Leaves sessile, appressed to spreading, longer to shorter than internodes, persistent; lamina 0-5-5 x 0-3-1-2 mm, linear to very narrowly oblong, margin plane to incurved, entire, sometimes cucullate, midrib prominent but not scabrid beneath, concolorous, not glaucous, subcoriaceous; apex acute, base parallel-sided, free; basal vein 1, unbranched; laminar glands dense, sometimes prominent beneath. Inflorescence 1 -flowered, branching pseudo-dichotomous, also sometimes from lateral branches; peduncle 2-4-5 mm long, not incrassate upwards; bracts foliar. Flowers 5-10 mm in diam., infundibular. Sepals 2-3(-4) x 0-7-0-9 mm, unequal, narrowly oblong or narrowly elliptic-oblong to lanceolate, acute; veins 3-5 unbranched, becoming ± prominent; glands linear, distally striiform to punctiform. Petals bright yellow to orange, tinged red outside in bud, 4-5-5 x 1-5-2 mm, c. 1-5-2 x sepals, oblong- oblanceolate; apiculus acute; glands striiform to punctiform. Stamens 10-15, longest 3-5-4-5 mm long, c. 0-8 x petals, in 5 ± distinct fascicles. Ovary 1-1-5 x 0-5-1 mm, narrowly ovoid-ellipsoid, trigonous; styles 3, 2-5-3 mm long, c. 2 x ovary, very slender, spreading; stigmas broadly capitate to peltate. Capsule 3-5 x 1-8-2-5 mm, narrowly cylindric to narrowly ellipsoid, acute, exceeding sepals. Seeds 0-5-0-6 mm long, ecarinate; testa finely scalariform. Exposed grassy slopes and open Pinus forest, often in wet areas; (600-) 1200-3000 m. Dominican Republic (La Vega, Benefactor [San Juan], Santiago), Haiti (Quest, Sud). DOMINICAN REPUBLIC. La Vega: vicinity of La Lagunita (Laguita), c. 2800-2900 m, 19 July 1967 (fl & fr), Gastony, Jones & Norris 300 (GH, NY, US); La Rucilla, Cienega de Manabao, Jarabacoa, 2200 m, 10 December 1969 (fr), Liogier 17199 (GH, NY, US); Valle Nuevo, c. 2400 m, 17 October 1929 (fr), Ekman H. 13825 (F, GH, S). Benefactor [San Juan]: Sabana Neuva, Cordillera Central, N. of Rio Arriba del Norte, 1950 m, 17-20 April 1946, R. & E. Howard 9021 (BM, GH, MICH, NY, P, S, US); Pico Duarte, 3000 m, 15-19 June 1974 (fl & fr), Liogier 21779 (NY). Santiago: Mancion, slope of Cerro Lucio, c. 2400 m, 12 June 1929 (fl & fr), Ekman H. 12844 (F, G, GH, K, NY, S, US); bassin du Rio Bao, December 1952 (fl & fr), Humbert 27737 (P). HAITI. Quest: Massif de la Selle, Petionville, Mt. La Visite, c. 2150 m, 12 August 1924 (fr), Ekman H. 1441 (C, S, US). Sud: Massif de la Hotte, W. group, Port-a-Piment, ridge W. of Morne Formont, 2275 m, 2 January 1927 (fr), Ekman H. 7548 (A, MO, S, US). H. fuertesii is most closely related (outside Hispaniola) to H. arbuscula from Guatemala, from which it differs by the smaller size of all parts, the decumbent rooting stems, and the occasional presence of lateral branches. Urban described it as an annual or perennial herb, but it is usually somewhat shrubby. 80. Hypericum dichotomum Lam., Encycl. 4: 167 (1797); Choisy, Prodr. monogr. Hyperic.: 49 (1821), in DC., Prodr. 1: 549 (1824); D. Dietr., Syn. pi. 4: 1236 (1847); Urban, Symb. antill. 7: 284 (1912); Mosc., Cat.fl. doming.: 381 (1943). Types: Hispaniola, Haiti, Thiery s.n. in Herb. Thouin (P-LAM-lectotype (mihi); C!, MPU-syntypes). Map 6. H. diosmoides sensu R. Keller in Bull. Herb. Boissier 6: 261 (1898), pro parte excl. typum. H. christii Urban, Symb. antill. 7: 283 (1912); Mosc., Cat. fl. doming.: 381 (1943). Type: Hispaniola, Haiti, in Morne Bouret, 1700 m, July, Christ 1873 (Bt-holotype?). H. polycladum Urban, Symb. antill. 7: 284 (1912); Mosc., Cat.fl. doming.: 381 (1943). Type: Hispaniola, Dominican Republic, prope Constanza in Valle Nuevo, 2200 m, August 1910 (fl), v. Turckheim 3568 (Bf-holotype; NY!). H. subglaucum Urban & Ekman in sched. (Ekman H. 13610). Shrublet or wiry perennial herb, 0-06-0-2 m tall and to c. 0-4 m long, with taproot, erect or decumbent to trailing and rooting, sometimes mat-forming; branches from below in- florescence and lateral, sometimes also basal, divaricate. Stems reddish-brown, 4-lined, ancipitous and densely gland- dotted at first, eventually terete, cortex exfoliating in strips; internodes 1-18 mm long. Leaves sessile, suberect to spread- ing, longer to shorter than internodes, persistent; lamina 1-5- 6 x 0-5-2-5 mm, oblong-elliptic or oblanceolate to obovate, margin plane, entire, not cucullate, midrib prominent be- neath, concolorous, sometimes ± glaucous, subcoriaceous to chartaceous; apex subacute (or rarely acute) to rounded, base parallel-sided to narrowly cuneate, free; basal vein 1, sometimes obscurely branched; laminar glands ± dense, ± prominent on both sides. Inflorescence 1-5-flowered, branch- ing repeatedly pseudo-dichotomous/sympodial and some- times dichasial-monochasial, without subsidiary branches; peduncle and pedicels 2-7 mm long, not incrassate upwards; bracts and bracteoles foliar, not reduced. Flowers 4-5 mm in diam., infundibuliform? Sepals 2-5-4 x 1-1-8 mm, unequal, narrowly oblong or oblanceolate to obovate-spathulate, acute to rounded; veins 3-5, unbranched, becoming prominent; glands linear, distally punctiform. Petals bright (?) yellow to yellow-orange, 3-5-5 x 1-1-5 mm, c. 1-5 x sepals, oblong- oblanceolate; apiculus acute?; glands distal, punctiform. Stamens c. 10-15, longest 3-5-4 mm long, c. 0-8 x petals, in 5 ± distinct fascicles. Ovary 1-1-5 x 0-5-0-8 mm, ellipsoid, acute to obtuse, exceeding sepals; styles 3, 1-5-2-5 mm long, 2-2-5 x ovary, very slender, spreading; stigma peltate. Capsule 3-5-5 x 2-4 mm, ovoid-ellipsoid to subglobose, acute to obtuse, exceeding sepals. Seeds 0-6-0-7 mm long, ecarinate; testa finely scalariform. Open, often wet places at roadsides and in Pinus forest and cloud forest; 1200-3000 m. Dominican Republic (La Vega, Benefactor [San Juan], Santiago), Haiti (Quest). DOMINICAN REPUBLIC. La Vega: Cordillera Central, 36 N. K. B. ROBSON Map 6 Sect. 29. A (upper): 80. H. dichotomum • . B (lower): 81. H. dlosmoides • (confirmed records), O (additional records, fide Lippold, 1970). Valle Nuevo, c. 2400 m, 17 October 1929 (fl & fr), Ekman H. 13824 (K, NY, S); Jarabacoa, Loma de la Salle, 1200 m, 24 May 1968 (fl & fr), Liogier 11386 (GH, NY, US); Constanza to Jarabacoa, near Paso Bajito, 1300 m, 13 November 1922 (fl & fr), Ekman H. 14121 (C, F, GH, S, US). Benefactor [San Juan]: Sabana Nuevo (Cordillera Central), N. of Rio Arriba de Norte, 1950 m, 18 September 1946 (fr), R. & E. Howard 9081 (BM, GH, MICH, NY, S, US); cercania de Pico Duarte, 3000 m, 15-19 June 1974 (fl & fr), Liogier 21789 (NY). Santiago: Cordillera Central, La Pelona, headwaters of Bao R., 1600 m, 1-7 October 1968 (fl & fr), Liogier 12746 (GH, NY, US). HAITI. Quest: Massif de la Selle, Petionville, between Morne La Visite and Morne Brouet, c. 1500 m, 27 April 1926 (fl & fr), Ekman H. 5999 (G, K, NY, S, US); Massif de la Selle, Fourcy, 10 September 1924 (fl & fr), Ekman H. 1832 (S, US). H. dichotomum has relatively broader leaves than H. fuertesii with the apex usually obtuse to rounded; the sepals, petals, styles, and capsules are also shorter, and the angle of the pseudo-dichotomies is usually wider. Thus, even though the distributional areas of these two species overlap, the species themselves apparently remain distinct. The most primitive form of H. fuertesii is in the Cordillera Central, whereas that of H. dichotomum is in southern Haiti (Massif de la Selle). The latter form is morphologically nearest to H. fuertesii, but the broader (acute) leaves and wider branching are distinc- tive. 81. Hypericum diosmoides Griseb., Cat. pi. Cub.: 40 (1866); Sauvalle in An. R. Acad. Cienc. Habana 5: 203 (1868), Fl. cub.: 8 (1869); R. Keller in Bull. Herb. Boissier 6: 261 (1898), pro parte, quoad typum, in op. cit., II, 8: 180 (1908); Urban, Symb. antill. 4: 412 (1910); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925); Mosc., Cat. fl. doming. 1: 381 (1943); Alain in Leon & Alain, FL Cuba 3: 317 (1953); Lippold in Wiss. Z. Friedr.- Schiller Univ., Jena, Math. -Nat. R. 19: 377 (1970); A. H. Liogier & Martorell, Fl. Puerto Rico & adj. hi. : 110 (1982). Type: Cuba, Pinar del Rio?, San Juan de Buenavista, 15 November 1860-64 (fr), Wright 2124 (GOET-holotype; BM!, GH!, K!, MO!, NY!, P!, S!, W!). Map 6. HYPERICUM H. hecatophyllum C. Wright in An. R. Acad. Ci. Habana 5: 203 (1868), in Sauvalle, Fl. cub.: 8 (1869); Alain in Leon & Alain, Fl. Cuba 3: 317 (1953). Type: Cuba, Pinar del Rio, al lado del no San Sebastian en la loma cerca de la Vega Lagunillas, 1860-64 (fr), Wright 3517 (NY!-holotype; GH!, K!,P!,S!,US!). H. portoricense Kuntze, Revis. gen pi.: 60 (1891). Type: Porto Rico, Cayey, 600 m, 13 July 1874 (fr), Kuntze 445 (NY!-holotype; Kl-isotype). H. constanzae Urban, Symb. antill. 7: 285 (1912); Mosc., Cat. fl. doming. 1: 381 (1943). Type: Hispaniola, Dominican Republic, prope Constanza, 1190 m, June, v. Turckheim 3053 b (Bt). H. ophiticola Britton in Mem. Torrey hot. Club 16: 83 (1920); Alain in Leon & Alain, Fl. Cuba 3: 317 (1953). Type: Cuba, Las Villas, City of Santa Clara, 29-31 March 1910 (st), N. & E. Britton & Wilson 6140 (NY!-holotype). Wiry perennial herb, to c. 0-15 m tall and to c. 0-5 m long, with taproot, decumbent (or rarely erect) to prostrate and rooting, not markedly mat-forming; branches from below inflorescence, lateral and basal, ascending or divaricate. Stems green, 4-lined, ancipitous and densely gland-dotted when young, eventually subterete, cortex sometimes exfoliat- ing in strips; internodes 2-5-12 mm long. Leaves sessile to very shortly pseudopetiolate, ± spreading, longer to shorter than internodes, persistent; lamina 3-9 x 0-5-3-5 mm, oblanceolate to oblong-spathulate, margin recurved to revo- lute, entire, not cucullate, midrib slightly prominent beneath, paler and ± glaucous beneath, subcoriaceous (?) to charta- ceous; apex subapiculate-obtuse to rounded, base narrowly cuneate to angustate, free; basal vein 1, sometimes with 2-3 pairs of lateral branches; laminar glands ± dense, ± prominent beneath. Inflorescence 1-5-flowered, pseudo- dichotomous to sympodial and rarely dichasial/monochasial, without subsidiary branches; peduncle and pedicels 2-6 mm long, not incrassate upwards; bracts foliaceous, not reduced, bracteoles subulate. Flowers 3-7 mm in diam., stellate to infundibuliform? Sepals 2-5-4-3 x 0-5-2 mm, unequal, narrowly oblong or oblanceolate to oblanceolate-spathulate, obtuse or apiculate to rounded, veins 3-5, unbranched, becoming prominent; glands linear, punctiform in distal §. Petals bright (?) yellow, 2-5-3(-4) x 0-8-l(-l-5) mm, about equalling sepals, oblanceolate; apiculus obsolete; glands distal, punctiform. Stamens 10-15, longest 1-5-2 mm long, c. 0-6 x petals, irregularly grouped? Ovary c. I x 0-5-0-8 mm, exceeding sepals; styles 3, 0-5-l(-l-5) mm long, c. 0-5-1-5 x ovary, slender, outcurved; stigma peltate. Capsule 2-5-3-5(- 4-5) x l-5-2(-2-5) mm, cylindric-ellipsoid to subglobose, obtuse, exceeding sepals. Seeds 0-5 mm long, ecarinate; testa finely scalariform. Open or shaded grassy slopes, roadsides or ditches, often in damp places; (200-)300-1350 (-2400?) m. Dominican Republic (La Vega, Santiago, Santiago Rod- riguez, Monte Cristi), Haiti (Quest), Porto Rico, Cuba (Oriente, Camaguey, Las Villas, Pinar del Rio). DOMINICAN REPUBLIC. La Vega: Jarabacoa, 500 m, 18 June 1969 (fl & fr), Liogier 15747 (GH, NY, US); Mata Grande, 900 m, 13 October 1968 (fl & fr), Liogier 13004 (GH, NY). Monte Cristi: Las Rosas, Rio Naranjo, c. 500 m, 6 June 1926 (fr), Ekman H. 6217 (S). Santiago: Sierra del Palo Quemado, 800 m, 10 May 1887 (fl), Eggers 1868 (BM, C, G, NY, P, US); c. 8 km NW. of San Jose de las Matas, Pinar de 37 Caimito, 400-450 m, 9 October 1969 (fr & fr), Liogier 16269 (GH, NY, US). Santiago Rodriguez: toward Rio Mao, Mancion, c. 300 m, 25 May 1929, Ekman H. 12602 (F, GH, S, US). HAITI. Quest: vicinity of Furcy, c. 1300 m, 26 May-15 June 1920 (fr), Leonard 4785 (BM, C, NY, US); Massif de la Selle, Marigot, near Grand Bassin Chotard, 9 June 1928 (fr), Ekman H. 10075 (S). PORTO RICO. Near Cayey, Coamo,5 20 September 1885, Sintenis 2488 (A, BM, F, G, K, MO, NY, P, S, US, W, Z); Monte Torrecilla, 900-1100 m, 19-20 March 1915 (fr), Britton 5201 (F, MO, NY, US); between Ciales and Casablanca, 200- 300 m, 30 January 1979 (fl), Liogier [& Martorell] 28216 (NY). CUBA. Camaguey: Sierra Cubitas to Santa Rosa, 21 February 1909 (fl), Shafer 553 (BM, F, GH, NY, US). Las Villas; Trinidad Mts, Tope de Collanthes, 800 m, 17 July 1957 (fr), Bro. Alain [Liogier] 6317 (GH); El Cumbre, 10 June 1923 (fl), Ekman 13947(8). Oriente: Baracoa, Rio Jojo, c. 1 km N. of Cajovavo, 1 August 1951 (fr), Webster 3998 (GH, MICH); Sabanilla to Yamuri Arriba, 31 January-1 February 1911, Shafer 8415 (GH, K, NY, US). Pinar del Rio: Rio San Sebastian, cerca de la Verga Lagunillas, 1860-1864 (fr), Wright 3517 (GH, K, NY, P, S, US) (see above). H. diosmoides appears to be a lower-altitude derivative of H. dichotomum that has spread from Hispaniola westward to Cuba and eastward to Porto Rico. It is usually easily distinguished from H. dichotomum by the longer, relatively narrower leaves with margin recurved to revolute, the shorter petals relative to the sepals, and the shorter styles relative to the ovary. There are, however, a few specimens from Hispaniola intermediate morphologically and from altitudes at the lower range of the H. dichotomum range but above the normal limit of H. diosmoides. As these specimens are few in number, it seems best to maintain these two taxa as species at present; an intensive study of field variation, however, might show that H. diosmoides should be reduced to subspecific rank under H. dichotomum. The intermediate specimens seen are: DOMINICAN REPUBLIC. La Vega: Constanza to Valle Nuevo, El Montazo, 1600-1800 m, 29 May 1969 (fl & fr), Liogier 15455 (GH, NY, US). HAITI. Quest: Massif de la Selle, Petionville, Morne Brouet, c. 1600 m, 26 July 1924 (fl), Ekman H. 1102 (S, US). In addition Fuertes 1810 (La Vega, in Loma Rosilla, 2700 m, 6 July 1912 (fl & fr) (GH, NY, P, W)), which morphologically is typical H. diosmoides, was growing at an unusually high altitude. 82. Hypericum gnidioides Seemann, Bot. voy. Herald: 88, t. 17 (1853); Triana & Planchon in Annls Sci. nat. (Bot.) II, 18: 298 (1862); R. Keller in Bull. Herb. Boissier II, 8: 176 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey hot. Club 56: 102 (1929); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 91, map 5 pro parte (1972), in Mems Assoc. Cienc. nat. La Salle 33: 114 (1973), pro parte excl syn. H. arbuscula et spec, ex Guatemala et Mexico; N. Robson in Ann. Mo. hot. Gdn 65: 12, f.l (1978), pro parte excl. distrib. Mexic. Type: Panama, Veraguas, Volcano of Chiriqui, February 1849 (fr), Seeman 1640 (GOETMectotype; BM!, GH!, K!, F!, GH!, NY!, US!-photographs). 5Some labels have 'prope Cayey', others 'Coamo'. 38 Map 5. H. hondurasense R. Keller in Bot. Jb. 58: 197 (1923); in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925). Type: Honduras, Zeguagalpa, Cerro Picacho, 24 January 1898 (fl), Niederlein 204 (Bt-holotype; F!, US '.-photographs). H. woodsonii Standley in Ann. Mo. hot. Gdn 26: 294 (1939). Type: Panama, Veraguas, Volcan de Chririqui, Loma Larga to summit, 2500-3380 m, 4-6 July 1938 (fl & fr), Woodson, Allen & Siebert 1040 (F!-holotype; GH!, MO!, NY!, US!-isotypes). H. pinetorum Standley in Publs Field Mus. not. Hist. (Bot.) 23: 65 (1944). Type: Honduras, Dept. Comayagua, near Siguatepeque, 1080-1400 m, 14-27 February 1928 (fr), Standley 55880 (F!-holotype; GH!, US'.-isotypes). H. arbuscula sensu Rodrigues Jimenez in A/eras Soc. Cienc. not. La Salle 33: 114 (1973), qua syn., non Standley & Steyerm. Icones: Seemann, Bot. voy. Herald: t. 17 (1853); N. Robson in Ann. Mo. hot. Gdn 65: 11, f.l (1978). Subshrub or shrublet (sometimes annual ?) 0-09-1-5 m tall, with taproot, usually bushy, erect; branches from base and below inflorescence, strict or lower ones ascending, rarely decumbent and mat-forming (0-03-0-08 m, '//. woodsonii'), not rooting. Stems reddish-brown, 4-lined, ancipitous and densely gland-dotted when young, eventually terete, cortex exfoliating in strips; internodes 3-12 mm long. Leaves sessile, spreading from base to closely imbricate, longer than inter- nodes, deciduous above base or persistent (i.e. deciduous with cortex); lamina (3-)5-18(-23) x 0-8-4(-5) mm, nar- N. K. B. ROBSON rowly elliptic-oblong to linear or more rarely narrowly lanceolate or (the lower) narrowly oblong-spathulate, margin recurved to revolute and rarely scabrid beneath, paler beneath, not glaucous, coriaceous to subcoriaceous; apex acute or apiculate to obtuse, base angustate or narrowly cuneate to parallel-sided, the pair scarcely forming interfoliar ridge or free; basal veins 1(3), with 1(2) pairs of main ascending lateral midrib branches, tertiary reticulation ab- sent; laminar glands dense, not prominent or slightly so beneath. Inflorescence 1-25-flowered, branching pseudo- dichotomous/sympodial or very rarely wholly or mixed dichasial/monochasial, sometimes with 1 pair of subsidiary branches from node immediately below, or lower laterals flowering; peduncle and pedicels 4-10 mm long, not incras- sate upwards;- bracts foliar, gradually reduced. Flowers 6-10 mm in diam., stellate. Sepals 4-8 x 1-2-3 mm, unequal, oblanceolate or oblanceolate-spathulate to oblong or elliptic or linear, acute; veins 5, unbranched, all becoming ± prominent; glands all linear or punctiform in upper 1A. Petals bright yellow to orange-yellow (? or orange), (5-)6-7(-8-5) x 2-5^t-5(-6) mm, l-2(-l-5?) x sepals, oblanceolate to oblong; apiculus acute; glands linear, distally interrupted. Stamens 20-40, longest 3-5-6 mm long, c. 0-75 x petals, fascicles not distinct. Ovary 1-5-1-8 x 0-6-1 mm, narrowly ovoid- ellipsoid; styles 3, 2-3 mm long, c. 1-3-1-5 x ovary, spreading; stigmas broadly capitate. Capsule 3-5-5-5 x 2-3 mm, ovoid-cylindric to cylindric-ellipsoid, trigonous, acute to subrostrate, shorter than or equalling sepals. Seeds c. 0-6 mm long, ecarinate; testa finely scalariform. Open places in dry Pinus or Pinus-Quercus forest (Hon- duras), paramo (Costa Rica), and dry grassland, savanna, or lava flows (Panama); 1050-3820 m. Honduras, Costa Rica (San Jose), Panama (Chiriqui, Code). HONDURAS. Comayagua: Siguatepeque to Comayagua, Barranco El Socorro, 1400 m, 28 November 1958 (fl), Hawkes, Hjerting & Lester 2109 (C, F, K, S); near Guaimaea, Frica Sanson to Cerro Sanson, 18 February 1955 (fl), Carlson 3175 (F); near Siguatepeque, 1050 m, 7 January 1936 (fl & fr), Yuncker, Dawson & Youse 5569 (F, GH, K, MICH, MO, S, U). Distr. Central: near Tamara, 19 June 1970 (fl & fr), Barkley & Ertha 40743 (F, GH, MO). El Paraiso: c. 6 km N. of Guinope, 1400 m, 29 December 1962, Williams, Molina & Williams 23260 (BM, C, F, GH, MICH, MO, NY, P, S, US, W); between Manzaragua & San Lucas, 1500 m, 4 July 1962, Molina 10741 (F). Intibuca: 11 km S. of La Esperanza, 22 May 1970 (fr), Hernandez & Barkley 40369 (GH); Morazan: near Hoya Grande, 1500 m, 28 July 1946 (fl & fr), Williams & Molino Will (F, MO); El Hatillo, 2000 m, 3 August 1978 (fl) Trochez 124 (BM, MO). Ocotepeque: Catarata de Belen, Gualcho, 15 km NE. de La Aldea, 1500-2000 m, 26 June-3 July 1976 (fl & fr), Nelson et al. 3772 (MO); Tegucigalpa: 2 km N. of the city, 30 January 1982 (fl & fr), Blackmore & Heath 1603 (BM). NICARAGUA. Nuevo Segovia: 1 km N. of Dipilto, 1100 m, 16 June 1977 (fl & fr), Neill 2244 (BM, MO). COSTA RICA. San Jose: 6 km NE. of Santa Maria de Dota, 1735 m, 21 December 1963 (fl & fr), Jimenez 1481 (F, NY, US); Paramos de 1'Abejonal pres de Ste Maria de Dota, 1500 m, 5 April 1890 (fl), Pittier 2279 (BM, P, US); Valle de los Conejos (lower), trail to Valle de los Leones, 3250-3450 m, 21-23 August 1971, Burger & Gomez P. 8204 (F). PANAMA. Chiriqui: vicinity of Boquete, Llanos area just S. of town, 1050 m, 10 March 1963 (fl & fr), Stern, Eyde & HYPER1CUM 39 Ayensu 1934 (MICH, MO, US); between Volcan de Chiriqui and Cerro Aquacate, 1950-2160 m, 16 January 1971 (fl), Wilbur & Teen 13310 (DUKE, F, MICH, MO); summit of El Barii, 3225-3474 m, 18 May 1976 (fl & fr), Croat 34927 (MO) ('//. woodsonii'). Code: between Las Margaritas y El Valle, 15 July & 8 August 1938 (fl), Woodson, Allen & Siebert 1758 (F, GH, MO). H. gnidioides varies widely with the habitat, having relatively broad, spreading leaves in moist places and narrow imbricate ones in drier areas. The mature upper leaves, however, are always relatively narrower than those of H. chamaemyrtus . The habit appears to be extremely variable, but this variation is based on two trends, viz. i) the multiplication of flowers by monochasial or sympodial or mixed branching in the secon- dary inflorescences (i.e. those formed above the first pseudo- dichotomy) and ii) the formation of lateral branches below this first dichotomy. H. woodsonii, from exposed habitats at higher altitudes (2500-3800 m) in Panama, is a prostrate form which 'has a low depressed habit and forms dense interlaced mats' [Standley]. The most primitive form of H. gnidioides is in Honduras, whence there is a south-eastward reduction trend to Panama (omitting southern Nicaragua) ending in '//. woodsonii'. 83. Hypericum eastwoodianum I. M. Johnston in Proc. Calif. Acad. Sci. IV, 20: 78(1931). Type: Mexico, Colima, Revilla Gigedo Islands, Socorro I., Grayson's Cove, 4 May 1925 (fr), Mason 1614 (CAS!-holotype; DS!, F!, GH!, MO!, NY!, US!-isotypes; P! -photograph). Map 5. H. insulare Eastw. in sched., non Fouc. & Mandon (1900). H. silenoides sensu Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 89, map 2 (1972), pro parte, in Mems Soc. Cienc. nat. La Salle 33: 51 (1973), pro parte quoad syn. H. east- woodianum. Subshrub or shrublet 0-25-0-6 m tall, with taproot, bushy, erect; branches lateral and below inflorescence, strict or lower ascending, not rooting. Stems reddish-brown, 4-lined, ± ancipitous and ± densely gland-dotted when young, eventually subterete, cortex exfoliating in strips; internodes 3-8-5 mm long. Leaves sessile, spreading from base to subimbricate-ascending, longer than internodes, persistent (i.e. deciduous with cortex) or breaking off earlier above base; lamina 7-30 x 1-3-5 mm, linear or narrowly lanceolate to linear-oblanceolate, margin ± strongly revolute, entire, not cucullate, midrib prominent or not and smooth beneath, usually papillose above, glaucous beneath and sometimes slightly so above, coriaceous; apex obtuse or apiculate to rounded, base angustate to parallel-sided, free; basal vein 1, with 2-3 pairs of ascending lateral branches sometimes visible beneath, tertiary reticulation absent; laminar glands dense, not prominent. Inflorescence (l)5-30-flowered, usually ± dense (c.f. 84. peninsulare) , branching dichasial/monochasial (after 1(2) nodes), without or rarely with subsidiary branches from node below; peduncle and pedicels 1-4-5 mm long; bracts and bracteoles linear. Flowers 4-6(-8?) mm in diam., stellate. Sepals 3-5 x 1-1-5 mm, subequal, oblanceolate to oblong or elliptic-oblong or lanceolate, acute; veins 3-5, unbranched, all becoming ± prominent; glands all linear or punctiform in upper 1A. Petals bright yellow to pale orange, c. 5-6 x 2-2-5 mm, 1-2-1-5 x sepals, oblanceolate to oblong; apiculus obtuse; glands linear, distally interrupted. Stamens (15-)20-40(-50, fide Johnston), longest 2-3 mm long, c. 0-5 x petals, fascicles not distinct. Ovary 1-5-2 x 1 mm, ovoid- ellipsoid; styles 3, (0-6)1-2-5 mm long, (0-4-) 0-7-1 x ovary, spreading; stigmas broadly capitate to peltate. Capsule 3-5- 4(-4-5) x 1-5-2-3 mm, narrowly ovoid-cylindric, trigonous, acute to acuminate, shorter than or equalling or rarely exceeding sepals. Seeds c. 0-6 mm long, ecarinate; testa finely scalariform. Sheltered places in dry rocks; 200-700 m (Socorro I.), 1475- 2100 m (mainland). Mexico (Colima: Revilla Gigedo Islands - Socorro I., Jalisco, Zacatecas). MEXICO. Colima: Socorro I., S. of Mt. Evermann, Cerro de los Dientes, c. 700 m, 11 March 1957 (fl & fr), Moran 5782 (CAS, DS, MEXU, MICH, NY, SD, US); Socorro I., 27 May-3 July 1903 (fr), Barkelew 229 (CAS, DS, GH, US) - 'H. insulare'. Jalisco: Municipio de Talpa, entre Cuala y la Cumbre Blanca, 1475 m, 2 September 1971 (fr & fr), Gonzalez Tamayo 336 (MICH). Zacatecas: Sierra de los Huicholes, 12-18 km SW. of San Juan Capistrano and 20-25 km SW. of Rio Atenco crossing (Chalapagana), 2100 m, 12, 14 January 1975 (fr), McVaugh 25786 (MICH). H. eastwoodianum is related to the most primitive form of H. gnidioides, differing essentially from it in having capsules longer than the sepals and styles relatively longer than the ovary. The leaves are usually spreading, rarely closely imbricate, the lateral branches more numerous, and the inflorescence separated from the foliage by an elongated internode. It is also very closely related to 84. H. peninsulare (q.v.). H. eastwoodianum occurs in three isolated populations, two on the central Mexican mainland (Jalisco, Zacatecas), and the third on Socorro Island, which lies 416 km from the tip of Baja California and 512 km from Cape Corrientes (in Jalisco). The populations are all distinct from one another but, apart from the Socorro Island one, are represented by effectively single collections. It seems unwise, therefore, to attempt to recognize any of the mainland ones taxonomically at present, although the shorter, subimbricate, glaucous 40 N. K. B. ROBSON leaves and capsules exceeding the sepals of the Zacatecas population may prove to be constantly distinct. 84. Hypericum peninsulare Eastw. in Leafl. West. Bot. 3: 257 (1943). Type: Mexico, Baja California Sur, Sierra de la Laguna, c. 1500 m, 13 October 1941 (fl & fr), Hammerley 382 (CAS!-holotype; DS!-isotype). Map 5. H. silenoides sensu Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 89, map 2 (1972), pro parte, in Mems Soc. Cienc. nat. La Salle 33: 51 (1973), pro parte quoad syn. H. peninsulare. H. gramineum sensu Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 92, map 8 (1972), pro parte, in Mems Soc. Cienc. nat. La Salle 33: 97 (1973), pro parte quoad spec. Brandegee 12 September 1892. Subshrub or perennial herb 0-15-0-35 m tall, with taproot; branches lateral and below inflorescence, strict, also basal or near-basal, not rooting, erect from spreading geniculate base, often forming ring round stem base. Stems reddish, persis- tently 4-lined, ± ancipitous and densely gland-dotted when young, cortex exfoliating in strips; internodes 6-30(^5) mm long. Leaves sessile, all spreading from base or upper erect, longer to shorter than internodes, persistent (i.e. deciduous with cortex) or lower breaking off earlier above base; lamina 5-23 x l-4-5(-6-5) mm, linear to narrowly elliptic or narrowly elliptic-oblong or lanceolate, margin ± strongly re volute, entire or subdenticulate, not cucullate, midrib prominent and smooth beneath, finely papillose or almost smooth above, glaucous beneath, subcoriaceous to charta- ceous; apex acute to obtuse, base parallel-sided to cuneate or rounded-amplexicaul, free; basal veins 1(3), without or with 2-4 pairs of ascending lateral branches visible beneath, tertiary reticulation absent; laminar glands dense, often slightly prominent beneath. Inflorescence 5-9-flowered, ± lax (c.f. 83. eastwoodianum), branching dichasial/monochasial (after 1(2) nodes), with subsidiary branches from node below; peduncle and pedicels l-5-3(-5) mm long; bracts and brac- teoles linear. Flowers 5-15 mm in diam., stellate. Sepals 3-5(- 6) x 0-8-1-5 mm, subequal, narrowly oblong or narrowly elliptic-oblong to lanceolate, acute to subacute; veins 3, unbranched, all becoming ± prominent; glands all linear or punctiform in upper 1A to Vi. Petals bright yellow, tinged red in bud, fading pinkish-orange, 5-7-5 x 1-5- c. 2-5 mm, 1-2-2 x sepals, oblanceolate to oblong; apiculus obtuse?; glands linear, distally interrupted. Stamens 15-22, longest 2-5-3 mm long, c. 0-5 x petals, fascicles not distinct. Ovary 1-2 x 0-5-1 mm, narrowly ovoid; styles 3, 1-5-2-5 mm long, 1-2-1-5 x ovary, spreading; stigmas broadly capitate. Capsule 4-7 x 2- 3 mm, ovoid-ellipsoid to cylindric-ellipsoid or ellipsoid, terete, acute to acuminate, exceeding or rarely equalling sepals. Seeds 0-7-0-8 mm long; testa finely scalariform. Open but sheltered dry habitats: meadows, among rocks, and in Pinus-Quercus forest; (200-) 1400-1 800 m. Mexico (Baja California Sur, extreme south). MEXICO. Baja California Sur: Sierra de La Laguna E. of Todos Santos, 1800 m, 27 December 1947 (fl & fr), Carter & Kellogg 2419 (BM, UC n.v.); Sierra de la Victoria, La Laguna Meadow, 20 August 1955 (fl & fr), Chambers 915 (DS, MEXU); Sierra El Taste, 1400-1500 m, 10 November 1955 (fr), Carter & Chisaki 3543 (CAS); ibid., 12 September 1892 (fr), Brandegee s.n. (US). The more shrubby form of H. peninsulare is very similar to 83. H. eastwoodianum, and there could be a case for treating these taxa as subspecies. The style length relative to the ovary, however, provides an absolute distinction (longer in peninsulare, equal or shorter in eastwoodianum), and the trends in branching pattern are different. I prefer, therefore, to treat them as separate species. Whereas the branches in H. eastwoodianum are never basal, in H. peninsulare there are usually several basal or near-basal shoots forming a cupuli- form 'rosette' (when young), as each emerges horizontally and then makes a right-angled bend to become erect. Again, H. eastwoodianum is always woody and usually has a more or less condensed inflorescence, whereas H. peninsulare ranges from shrublet to perennial (or biennial?) with a correspond- ing morphocline in the inflorescence from rather lax to very lax. 85. Hypericum beamanii N. Robson, sp. nov. Map 5. H. galino S. F. Blake affinis, sed folis tenuiter chartaceis margin nonnihil recurvis valde undulato-denticulatis, floribus ut videtur minoribus, differt. Type: Guatemala, Solola, Volcan Atitlan, near summit of mountain c. 3535 m, 10 August 1960 (fl), Seaman 4062 (GH!-holotype; US!-isotype; MSC). Shrublet c. 0-28 m tall, with taproot, erect and bushy or somewhat diffuse; branches basal and lateral, strict or erect from decumbent to prostrate rooting base. Stems reddish- brown, 4-lined, ancipitous and ± densely gland-dotted when HYPER/CUM 41 young, eventually 2-lined to subterete, cortex exfoliating in strips; internodes 3-8 mm long. Leaves sessile, spreading from base, longer than internodes, persistent, becoming deflexed and withering; lamina 8-18 x 1-3-3 mm, linear or narrowly elliptic to narrowly oblanceolate, margin slightly recurved, strongly undulate-denticulate, not cucullate, midrib slightly prominent beneath, smooth on both sides, slightly paler beneath, not glaucous, thinly chartaceous; apex acute to apiculate or obtuse, base narrowly cuneate to angustate, free; basal vein 1, with 1-3 pairs of ascending lateral branches, tertiary reticulation absent; laminar glands rather dense, not prominent. Inflorescence c. 5?-flowered, branching dichasial/ monochasial, with subsidiary branches from 1-2 nodes below; peduncle and pedicels 2-4 mm long; bracts and bracteoles linear. Flowers immature, in bud only. Sepals (immature?) 2-5-3-5 x 0-5-0-7 mm, subequal, narrowly oblong, acute; veins 3, unbranched; glands linear. Petals tinged red in bud. Trough-like depression near fumerole vent; c. 3535 m. Guatemala (Solola). GUATEMALA. Solola: Volcan Atitlan, near summit of mountain, c. 3535 m, 10 August 1960 (bud), Beaman 4062 (GH, MSCn.v.,US). The above single collection represents a quite distinct, isolated species whose affinities lie with 83. H. east- woodianum and 85. H. galinum. Its leaves are thinner than those of either species, but the undulate-denticulate margin is reminiscent of H. galinum. Mature flowers and fruits would enable a more complete differential diagnosis to be written. 86. Hypericum galinum S. F. Blake in /. Wash. Acad. Sci. 14: 289 (1924); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 91, map 5 (1972), in Mems Soc. Cienc. nat. La Salle 33: 116, ff. Ih, 14B (1973). Type as for H. denticulatum Kunth, non Walter. Fig. 11B, Map5. /I \J7 H. denticulatum Kunth in Humb., Bonpl. & Kunth, Nov. gen. sp. 5: 191, t. 458 (1822); D. Dietr., Syn. pi. 4: 1237 (1847); S. Watson in Proc. Am. acad. Arts Sci. 17: 330 (1882); Urbina, Cat. pi. mexic. : 19(1897); R. Keller in Bull. Herb. Boissier II, 8: 180(1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925), non H. denticulatum Walter (1788). Type: Mexico, Guanajuato, 'Guanaxuato Mexicorum', 1944 m ('1080 hex') September 1803, Hum- boldt & Bonpland s.n. (P!-holotype). Brathys denticulata (Kunth) Spach in Annls Sci. nat. (Bot.) II, 5: 367 (1836). Icon: Humb., Bonpl. & Kunth, torn, cit.: t. 458 (1822). Shrublet 0-2-0-4 m tall, with taproot, fastigiate, erect; branches lateral and sometimes basal, often opposite, strict, not rooting. Stems 4-lined, reddish-brown, ancipitous, and densely gland-dotted when young, eventually terete, cortex exfoliating in strips; internodes 3-20 mm long. Leaves sessile, erect to appressed, imbricate, shorter to longer than inter- nodes, persistent; lamina (5-)8-15(-20) x 1-2 mm, linear to narrowly elliptic-oblong, margin recurved or plane to subin- curved, retorse-denticulate to (rarely) subentire, not or slightly cucullate, midrib prominent or not and sometimes denticulate beneath, smooth, concolorous, not or slightly glaucous, coriaceous; apex acute to subacute, base parallel- sided or slightly broadened, free; basal vein 1, not or obscurely branched, tertiary reticulation absent; laminar glands dense, not prominent or slightly so beneath. Inflor- escence 1-12-flowered, branching dichasial/monochasial, often with subsidiary inflorescences or inflorescence branches from node(s) below; peduncle and pedicels 1-5-3 mm long, not incrassate upwards; bracts and bracteoles linear-subulate. Flowers 8-10 mm in diam., stellate. Sepals 2-5-4 x 0-8-1-2 mm, unequal, oblong-spathulate or narrowly oblong to lanceolate, obtuse to acute, ± cucullate; veins 3-5(7), unbranched, all becoming prominent; glands linear. Petals bright yellow, tinged reddish, 4-6 x 2-2-5 mm, c. 1-5 x sepals, oblanceolate; apiculus acute; glands few, punctiform. Stamens 10-14, longest 3-4 mm long, c. 0-7 x petals, fascicles 3, subdistinct. Ovary c. 1-5 x 1 mm, narrowly ellipsoid; styles 3, 1-1-5 mm long, c. 0-7-1 x ovary, suberect; stigmas capitate. Capsule 5-6 x 2-2-5 mm, narrowly ovoid-cylindric, acute, 2-3 x as long as sepals, sometimes with 1-2 vesicles. Seeds 0-5 mm long, ecarinate; testa finely scalariform. Dry, rocky, calcareous areas; 1800-2500 m. Mexico (Guanajuato, San Luis Potosi). MEXICO Guanajuato: Xichii road, 20 August 1947 (fr), Kenoyer 2324 (GH); Jaral, 10 August 1885 (fl & fr), Schumann 350 (P). San Luis Potosi: chiefly in region of San Luis Potosi, 1800-2400 m, 1878 (fr), Parry & Palmer 72 (BM, G, GH, K, MO, NY, P, US); Cerro Tepetate, SW. of San Luis Potosi, 2400-2500 m, 27 July 1934 (fr), Pennell 17655 (GH, MEXU, NY, US); San Luis Potosi, August, October 1877 (fl & fr), Schaffner 607 pp. (GH, K, MEXU, NY), ibid., 1879 (fl & fr), Schaffner 511 (BM, G, MICH, P, US). H. galinum is closely related to the mainland populations of 83. H. eastwoodianum, differing from them inter alia by the plane to incurved leaves with denticulate margin and the elongate capsules. 87. Hypericum drummondii (Grev. & Hook.) Torrey & A. Gray, Fl. N. Amer. 1: 165 (1838); Steudel, Nomencl. hot. 2nd ed. 1: 787 (1840); A. Gray, Manual 5th ed.: 86(1856) et eds subseq.; Chapman, Fl. South. U.S: 42 (1872); Coulter in Bot. Gaz. 11: 110 (1886); Britton, N. L. & Brown, A. H. ///. ft. n. U.S. 2: 435, f. 2462 (1897), 2nd ed. 2: 536 (1913), 3rd ed. 2: 544 (1952); C. Mohr in Contr. U.S. natn. Herb. 6: 621 (1901); R. Keller in Bull. Herb. Boissier II, 8: 180 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 88, map 1 (1972), in Mems Soc. Cienc. nat. La Salle 33: 109, ff. Ig, 14c, t. IB (1973); D. H. Webb, Biosyst. study Hypericum sect. Spachium in eastern N. Amer.: 119 (1980). Type: U.S.A., Missouri, St. Louis, 1832 (fl & fr), 42 N. K. B. ROBSON Drummond (K!-holotype; BM!, GH!, P-isotypes). The lectotype of H. drummondii chosen by Rodriguez Jimenez is superfluous, as the holotype is at Kew. Fig. 11A, Map 7. Sarothra drummondii Grev. & Hook., Bot. misc. 3: 236, t. 107 (1833); Small, Fl. s.e. U.S.: 791 (1903); Y. Kimura in Nakai & Honda, Novafl. jap. 10: 48, 232 (1951). Brathys drummondii (Grev. & Hook.) Spach in Annls Sci. nat. (Bot.) II, 5: 367 (1836). Icon: Rodriguez Jimenez in Mems Soc. Cienc. nat. La Salle 33:110, f. 14C(1973). Annual herb 0-1-0-8 m tall, with taproot, erect; branches lateral from upper part of stem, strict, sometimes opposite, not rooting. Stems eventually orange-brown, 4-lined, ancipi- tous above, densely glandular, cortex exfoliating in strips; internodes 7-12 mm long. Leaves sessile, erect to suberect, longer to shorter than internodes, persistent; lamina 5-22 x 0-5-1 mm, linear or linear-subulate to linear-lanceolate, margin recurved to ± revolute, smooth, not cucullate, midrib prominent, smooth, concolorous, pale but not glaucous, subcoriaceous; apex acute to obtuse, base parallel-sided, free; basal vein 1, unbranched, tertiary reticulation absent; laminar glands dense, not prominent. Inflorescence 1-7 (-12)- flowered, branching dichasial/monochasial, with subsidiary inflorescences and inflorescence branches from up to 6 nodes below; peduncle and pedicels 1-4 mm long, not incrassate upwards; bracts foliar, gradually reduced. Flowers 5-8 mm in diam., stellate. Sepals 3-7 x 0-7-1-3 mm, subequal, narrowly oblong to linear-lanceolate, acute; veins 3-5, unbranched, all becoming ± prominent; glands punctiform in upper Vz-V2. Petals orange-yellow, 4-7 x (1-5)2-2-5 mm, about equalling Map 7 Sect. 29: 87. H. drummondii. Data from Webb (1980). HYPERICUM 43 Fig. 11 A. H. drummondii: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) young capsule. B. H. galinum: (f) habit; (g) stem with leaves; (h) leaf; (i) sepal; (j) petal; (k) young capsule with sepals, petals, and stamen filaments (partly cut away); (1) mature capsule (a, f x 1/2; b, g x 2; c-e x 5; h-1 x 4). A. D. & H. Correll 35993; B. Shaffner 607 p.p. 44 N. K. B. ROBSON sepals, oblong; apiculus subacute, small; glands distally punctiform? Stamens 10-22, longest 2-5-3 mm long, c. 0-5 x petals, obscurely fascicled or irregularly spaced. Ovary c. 1 x 0-5 mm, narrowly ovoid; styles 3, (0-5-)0-8-l-5 mm long, c. 1-1-5 x ovary, spreading; stigmas peltate. Capsule (3-5-)4-5- 6(-7) x 2-5-3 mm, narrowly ovoid to ovoid-cylindric, acute, slightly shorter to slightly exceeding sepals. Seeds 0-9-1-1 mm long, ecarinate; testa finely scalariform. 2n = 24 (n = 12, Webb, 1980). Dry, sandy or clay soils in open woods, old fields, and waste or rocky places; 0-1040 m. Eastern and south-eastern U.S.A. (Texas to NW. Florida, north to Virginia and Ohio, west to Missouri, Kansas, and Oklahoma). U.S.A. Alabama: *Jefferson Co., Red Mountain, 5 Octo- ber 1898, Canby 13 (MO, NY, US); *Tuscaloosa Co., E. of University campus, 9 September 1933, Harper 3108 (GH, MICH, MO, NCU, NY). Arkansas: Ashley Co., Hamburg, 75 m, 27 September 1937 (fr), Demaree 16376 (H); Independ- ence Co., N. of Cushman, 1 September 1960 (fl & fr), Adams 671 (K). Florida: Apalachicola R., August (fr), Curtiss 256 (BM, K, P); *Jackson Co., L. Seminole, N. of Sneeds, 9 September 1958, Godfrey 57779 (GH). Georgia: Richmond Co. ?, circa urbem Augusta, n.d. (fr), Olney & Metcalf 15(K); *Taylor Co., Butler, 1040 m, 14 September 1908, Harper 2239 (GH, MO, NY, P, US). Illinois: *White Co., S. of Norris City, 28 July 1951, Ahles 4680 (ILL); no precise locality, August 1836 (fl), Riedl s.n. (BM). Indiana: "Jeffer- son Co., along Rd. 3, c. 2-4 km N. of Rd. 256, 27 October 1945, Friesner 20247 (MICH, NY, SMU); ? Co., Sapulpa, 21 September 1895 (fl & fr), Bush 1385 (K, P). Kansas: *Chantaqua Co., no precise locality, 1896, Hitchcock 635 (MO, NY, P, US); *Cherokee Co., 2-4 km S. of Galena, 12 October 1968, Magrath & Johnson 3570 (NY). Kentucky: *Madison Co., 3-2 km NE. of Big Hill, 6 August 1940, Wharton 5671 (MICH). Louisiana: *Sabine Par., 7-4 km S. of Many, 9 September 1955, Shinners 21621 (GH, ILL, SMU); Calcasieu Par., L. Charles, 11 September 1915 (fl & fr), Palmer 8524 (K); Ouachita Par., Windsor, 13 October 1915 (fr), Palmer 8936 (K). Mississippi: Jackson Co., Biloxi, 7 October 1897 (fl), Tracy 3577 (BM, GH, MO, NY, US, W); *Perry Co., Maxie, De Soto National Forest, P.O. Beaumont, 22 October 1954, Demaree 3257 (GH, SMU). Missouri: Stoddard Co., Bernie, 2 August 1895 (fl), Bush 85 (K); Jasper Co., near Webb City, 20 August 1927 (fl & fr), Palmer 32589 (K, P). North Carolina: Buncombe Co., Biltmore, 17 August 1897 (fl & fr), Herb. Biltmore 526b (BM, GH, MICH, MO, NY, NCU, US, W, Z); Lee Co., 6-4 km N. of Sanford, near US 1, 18 October 1966 (fr), Radford 45337 (BM, H). Ohio: * Adams Co., L. Adams, 10 October 1956, Braun s.n. (US); *Pickaway Co., Pickaway Towndship, 20 August 1937, Hartley & Pontius 622 (NY). Oklahoma: Le Flore Co., Poteau, 14 July 1915 (fl), Palmer 8290 (K); *Pawnee Co., Cleveland, 1 October 1914, Palmer 6382 (MO). South Carolina: *Berkeley Co., along S.C. 45, Jamestown, Santee Wool-combing Mill, 5 July 1957, Ahles & Haesloop 30826 (NCU, SMU); *Union Co., 3-2 km E. of Carlisle, 25 August 1956, Freeman 56678 (NCU). Tennessee: Dyer Co., N. of Roellin, 16 August 1947 (fl), Sharp & E. & A. Clebsch 6299 (BM, TENN); *Putnam Co., N. of Double Springs, 21 August 1947, Norris & Shanks 7312 (NCU, TENN). Texas: Galveston Co., c. 8 km NE. of Port Bolivar, 0-4 km behind beach, 9 August 1968 (fr), D. & H. Correll 35993 (BM); Lee Co., c. 13 km S. of Dime Box, 26 August 1979 (fl & fr), Fryxell 3123 (BM). Virginia: *Greenville Co., near Three Creek, N. of Emporia, 18 September 1938, Fernald & Long 9378 (GH, NY, US); *Norfolk Co., along RR, 0-8 km W. of Bowers Hill, 6 August 1944, Hubricht B2571 (MO). H. drummondii is most closely related to the possibly annual form of H. gnidioides from Honduras, being smaller or narrower in all parts except apparently the seeds. The wide disjunction in area between Honduras and south- eastern U.S.A. is almost paralleled by that of H. gymnan- thum (south-eastern U.S.A. to Guatemala, see p. 115). 88. Hypericum gentianoides (L.) Britton, Sterns & Pogg., Prelim cat. : 9 (1888); N. L. Britton & A. H. Brown, ///. Fl. n. U.S. 3rd ed. 2: 544 (1952); Lyman B. Smith in J. Wash. Acad. Sci. 48: 313 (1958); Heine in Bauhinia 2: 76 (1962); N. Robson in Fl. Europaea 2: 269 (1968); Utech & Iltis in Wisconsin Acad. Arts Sci. Lett. 58: 346 (1970); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 89, map 2 (1972), in Mems Soc. Cienc. nat. La Salle 33: 112, ff. 2h, 3d (1973), in Fl. III. Catar., Hipericdc.: 9 (1980), pro parte, excl. syn. Sanidophyllum cumulicola et Hypericum cumulicola et H. aphyllum et Sarothra aphyllum; D. H. Webb, Biosyst. study Hypericum sect. Spachium in eastern N. Amer.: 137 (1980), pro parte excl. syn. H. aphyllum; J. M. Gillett & N. Robson in Canad. natn. Mus. nat. Sci., Publs Bot. 11: 11, tt. 4, 16, map 3 (1981). Type as for Sarothra gentianoides L. Maps 5, 8, 9, 38. Sarothra gentianoides L., [Nova pi. gen. II: 14 (1751)] Sp. pi.: 272 (1753), in Amoen. Acad. 3: 11(1756); N. L. Britton & A. H. Brown, ///. fl. n. U.S. 2: 436, f. 2463 (1897), 2nd ed. 2: 536 (1913); Tempere in Act. Soc. linn. Bordeaux 83: 136 (1932); Y. Kimura in Nakai & Honda, Novafl. jap. 10: 231 (1951). Type: U.S.A., Kalm in Herb. Linn. 391: 1 (LINN!- lectotype - Rodriguez Jimenez, 1973). There are three elements in the protologue from which the lectotype must be chosen: 1) the Kalm specimen (LINN), on which Linnaeus based the monotypic genus Sarothra (Gen. Nov. : 1075 (1751)); 2) Clayton 110 (BM), cited by Gronovius in HYP ERIC UM 45 Map 8 Sect. 29: 88. H. gentianoides, natural range. Data from Webb (1980). See also Maps 5 (p. 32), 9 (p. 46) and 38 (p. 123). Fl. Virg.: 29 (1739); 3) a reference to Plukenet, Mant.: 43, t. 342, f.4 (1700), based on a Krieg specimen (BM-H.S. 92: 43, also 102: 162). Gillett & Robson (1981) selected Clayton 110 as lectotype without comment; but Rodriguez Jimenez (1973) had already cited Kalm s.n. as Type'. Since elsewhere in this work she indicates lectotypes and neotypes, there is room for doubt that she did, in fact, intend to choose a lectotype. Nevertheless, Webb (1980, ined.) followed her in citing Kalm s.n. but stated that a strong argument could be made for selecting Clayton 110. Reveal et al. (1987: 217) pointed out that, being the basis of the genus Sarothra, the Kalm specimen is the better lectotype; but, not knowing of the earlier 'choice' by Rodriguez Jimenez, they said that the choice of Clayton 110 could not be overturned. In the circumstances, however, it seems best to do so by crediting Rodriguez Jimenez with the intention of choosing a lectotype. In this way the type of the monotypic genus and its species become one and the same. H. nudicaule Walter, Fl. carol.: 190 (1788); Coulter in Bot. Gaz. 11: 111 (1886)6; R. Keller in Bull. Herb. Boissier II, 8: 179 (1908), in clav. (sphalm. [auct.] Rob Keller), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925). Type: U.S.A., Carolina, Walter L 60 no. 649 (BM!). H. sarothra Michaux, Fl. bor.-amer. 2: 79 (1803), nom. illegit. (Art. 63); Pursh, Fl. Amer. sept. 2: 378 (1814); Torrey & A. Gray, Fl. N. Amer. 1: 165 (1838); Walp., Repert. hot. syst. 1: 387 (1842); D. Dietr., Syn. pi. 4: 1235 (1847); A. Gray, Manual 5th ed.: 86 (1856) et eds subseq.; Chapman, Fl. South. U.S.: 42 (1872); Chodat & Hassler in Bull. Herb. Boissier II, 3: 1126 (1903). Type as for Sarothra gentianoides. Sarothra hypericoides Nutt., Gen. N. Amer. pi. 1: 204 (1818), nom. illegit. (Art. 63); W. P. G. Barton, Fl. N. Amer. 3: 59, t. 92 f. 1 (1823). Type as for 5. gentianoides L. 6Coulter cited Linnaeus's reference to Pluk. aim. 189 under H. setosum as a synonym of H. nudicaule. Plukenet's description (Hypericum virginianum parvum fruticosum, ramulis equiseti. Planta est bicapsularis ex flore pentapetala luteo.) certainly could apply to H. gentianoides. His specimen, however, is lacking at BM; and in any case H. setosum is lectotypified by a Gronovius specimen (see Robson in Taxon 29: 271 (1980)). 46 N. K. B. ROBSON Map 9 Sect. 29: 88. H. gentianoid.es , South American range. Brathys gentianoides (L.) Spach, Hist. nat. veg., Phan. 5: 455 (1836), in Annls Sci. nat. (Bot.) II, 5: 367 (1836). Icon: J. M. Gillett & N. Robson in Canad. natn. Mus. nat. Sci., Publs Bot. 11: 12, t. 4 (1981). Annual herb 0-07-0-6 m tall, with taproot, erect; branches lateral, from upper two-thirds or from most nodes, strict, mostly opposite (at least lower ones), wiry, not rooting. Stem eventually orange- to reddish-brown, 4-lined, ancipitous above, densely glandular, cortex exfoliating in strips; inter- nodes 3-20 mm long. Leaves sessile, appressed, canaliculate or sometimes scale-like, shorter than internodes, persistent; lamina 1^ x 0-4-0-6 mm, all very narrowly triangular- subulate to linear-subulate or first two pairs broader, margin incurved, smooth, ± cucullate, midrib slightly prominent, smooth, concolorous, not glaucous, subcoriaceous to charta- ceous, apex obtuse to rounded, base parallel-sided or slightly broadened, free; basal vein 1, unbranched, tertiary reticula- tion absent; laminar glands often in a single row, not prominent. Inflorescence 1-24-flowered, usually pyramidal overall, branching dichasial/monochasial or often wholly monochasial, with subsidiary inflorescences and inflorescence branches from most or nearly all nodes below; pedicels 1-2-5 mm long, not incrassate upwards; bracts foliar, not reduced. Flowers 3-5 mm in diam., stellate. Sepals 1-5-2-5 x 0-4-0-8 mm, unequal, lanceolate to narrowly oblong or linear- lanceolate, acute; veins 3, unbranched, all becoming ± prominent; glands linear. Petals orange-yellow to yellow, tinged red in bud, 2-3-3(-4) x 0-8-1-2 mm, c. 1-3 x sepals, oblong; apiculus almost absent; glands distally punctiform. Stamens 5-10(11), longest 1-5-2 mm long, c. 0-5-0-65 x petals, obscurely 5-fascicled or 5 single. Ovary 1-1-5 x 0-5- 0-8 mm, narrowly ovoid-trigonous; styles 3, 0-8-1-2 mm long, c. 0-8 x ovary, spreading; stigmas broadly capitate. Capsule 4-5 x 1-1-2 mm, very narrowly cylindric-conic, acute, 2-3 x sepals, with valve midrib ± prominent. Seeds 0-4-0-75 mm long, ecarinate; testa markedly ribbed-scalariform. 2n = 24 (Webb, 1980); n = 12 (Hoar & Haertl, 1932). Dry, sandy soils in open woods, fields, roadsides, and waste or rocky places; lowland in N. America, to 1600 m in Hispaniola. Canada (southern Ontario), eastern U.S.A. (Minnesota and Michigan east to Maine, south to Florida and eastern Texas, north to Oklahoma, Missouri, and Iowa); probably or certainly introduced into Hispaniola (Dominican Republic), Paraguay, Brazil (Rio Grande do Sul), and France (Gironde). CANADA. Ontario: Sandwich, 4 July 1892 (fl), Macoun 18317 (BM, K); Windsor, 23 July 1901 (fl), Macoun 34082 (GH, K, MO, NY, US). U.S.A. Alabama: *Covington Co., 12 km S. of Opps, 18 August 1958, Skinners 27455 (GA, NCU, SMU); *Lee Co., Auburn, 10 September 1899, Earle & Earle 41 (GH, NY, US); *Tuscaloosa Co., 9-6 km E. of University, 28 July 1950, Skinners 12655 (GA, SMU). Arkansas: *Drew Co., P.O. Monticello, 21 September 1937, Demaree 16339A (GH, MO, NY, SMU); *Perry Co., Ouachita Nat. Forest, Lake Sylvia, P.O. Thornburg, 25 August 1937, Demaree 16005 (GA, ILL, MO, NY, SMU); *Stone Co., 8 km SW. of Calico Rock, 14 October 1967, Tucker 6837 (NCU). Connecticut: Ellington, 21 August 1875 (fl), Pease s.n. (BM); *Windham Co., Woodstock, 1 August 1965, Seymour et al. 22973 (MO). Delaware: *New Castle Co., Reybold, 14 October 1939, Benner 9164 (GH) ; *Sussex Co. , Reboboth, 16 October 1933, Larsen 440 (GH, MO). District of Columbia: Washington, July 1897 (fl), Holm s.n. (W); *Mount Hamilton tract, 20 July 1923, Leonard & Kellip 1573 (US). Florida: Brevard Co.: Okeechobee region, 8 June 1903 (fl), Fredholm 5850 (GH, K, MO); Calhoun Co., 19-5 km S. of Blountstown, Creek Swamp, 19 July 1954 (fl & fr), Ford & Arnold 3349 (BM); Gulf Co.: 8 km S. of Wewahitchka on Rt. 71, 7 September 1970 (fl), Mengher310 (BM). Georgia: *Clarke Co., Athens, along Oconee R., behind University campus, 20 September 1946, Cronquist 4149 (GA, GH, MO, NY, SMU, US); Sumter Co., no precise locality, August 1897 (fl & fr), Harper s.n. (BM); *McIntosh Co., beach on Sapelo Island, 26 July 1956, Duncan 20364 (GH, MICH, NCU, SMU, US). Illinois: *Cook Co., West Pullman, 17 September 1900, Chase 1485 (GH, MO); *MasonCo., W. of San Jose, 12 July 1951, Ahles 2873 (ILL); Union Co., no precise locality, 27 July 1878 (fl), French 272 (K). Indiana: *Orange Co., 8 km S. of Chambers- burg, N. of Danner's Chapel, 25 September 1937, Friesner 11572 (NY, SMU); *Porter Co., Mineral Springs, Little Lake, 25 July 1925, Lyon s.n. (MICH). Iowa: Clinton Co., no precise locality, 1 October 1932 (fl), Naers s.n. (W). Kentucky: *Harlan Co., Pine Mountain, August 1893, Kearney 256 (GH, MO, NY, US); *Warren Co., 6-5 km SW. of Shanty Hollow Lake, 4 August 1970, Conrad 1536 (MO). Louisiana: *Rapides Par., N. of Pineville, 29 July 1938, Correll & Correll 9895 (GH, NY); *Sabine Co., 7-2 km S. of Many, 9 September 1955, Skinners 21620 (GH, SMU); St. Tammany Par., Covington, June & August 1932, Drum- mond s.n. (BM, K). Maine: * Hancock Co., S. ridge of Cadillac Mt., 23 July 1952, Rossbach 2436 (NCU); *Lincoln Co., SW. end of Monhegan Island, 12 August 1954, Rossbach 3533 (NCU). Maryland: Baltimore Co., W. of Owings Mills near Deer Park Road, Soldiers' Delight Area, 17 August 1970 (fl & fr), Windier, Keenan & Lombards 3223 (H, MICH, NCU); Calezu Co., Chesapeake Beach, 27 August 1904 (fl), House 366 (K); Massachusetts: Hampshire Co., Hatfield, 31 August 1973 (fl & fr), Ahles 78621 (BM, H); ? Co., Bourne, 15 September 1901 (fl & fr), Kennedy, Williams & Fernald 63 (BM, BR, G, K, P, W). Michigan: *St. Clair Co., near Port Huron, above Tunnel Station, 15 September 1906, Dodge s.n. (MICH). Mississippi: Jackson Co., Petie Bois Island, 5 August 1898 (fl), Tracy 4966 (BM, MICH, MO, NCU, NY); HYPER/CUM *Stone Co., DeSoto Nat. Forest, Beatrice, P.O. McHenry, 24 August 1953, Demaree 34057 (GH, SMU). Missouri: Dunklin Co., Campbell, 15 August 1895 (fl), Bush 90 (K, MO, NY, US); *Polk Co., Graydon Springs, 31 July 1937, Steyermark 23957 (MO); St Louis Co., St. Louis, August 1838 (fl), Riehl 3 (BM, BR, G, MO, P). New Hampshire: Hillsboro Co., 1-1 km NE. of Hillsboro-Cheshire Co. line on US Rt. 202, S. of Peterborough, 15 August 1967 (fl & fr), Ahles 69047 (BM). New Jersey: Burlington Co., c. 9-5 km W. of Chatsworth, 20 August 1948 (fl & fr), Lawrence & Drew 531 (BM); Somerset Co., Watchung, 31 July 1934 (fl), Moldenke 8148 (BM). New York: Nassau Co., Long Island, Montauk, 12 August 1934 (fl & fr), Yamamoto s.n. (TAI); *Tompkins Co., Ithaca, South Hill, 11 August 1921, Muen- scher 14019 (GH, MO). North Carolina: Alamance Co., 7-2 km E. of Snow Camp, 17 July 1956 (fl & fr), Bell 4091 (H); Pitt Co., near US 264, 4-3 km SE. of Grimesland, 10 August 1958 (fl & fr), Radford 39573 (BM); *Wake Co., 11 km NW. of Raleigh, 13 July 1938, Godfrey 4963 (NCU, NY). Ohio: *Hocking Co., Rock House State Park, 1 September 1935, Demaree 11604 (NCU); *Ross Co., near Londonderry, Liberty Township, 8 September 1936, Bartley & Pontius 332 (NY). Oklahoma: *Delaware Co., 6-4 km N. of Little Kansas along Okla. 10, 29 July 1957, Wallis 5343 (SMU). Pennsylva- nia: Alleghany Co., Harrison Township, 16 September 1901 (fr), Shafer 1470 (GH, W); Delaware Co., Swarthmore, 14 July 1947 (fl), Palmer s.n. (W); Northampton Co., Bethle- hem, August 1832 (fl), Moser s.n. (BM, G, MO, NY, P). Rhode Island: Washington Co., junction fo R.I. Rt. 138 and Int. Rt. 95, Wyoming, 19 August 1974 (fl & fr), Ahles 79120 (BM). South Carolina: Beaufort Co., W. end of Lemon Island, just N. of S.C. Rt. 170, slightly above sea level, 12 September 1982 (fr), Spongberg, Boufford & Bartholomew 17189 (BM); Chester Co., Chester, October 1958 (fr), Irvine 877 (BM); *Lexington Co., 12-8 km SE. of Columbia, 8 August 1939, Godfrey & Try on 1330 (GH, NY, US). Tennessee: Cumberland Co. ?, Cumberland Tableland, July 1888 (fl), Middleton (BM); *Grainger Co., Clench Mt., NW. of Bean Station, 28 July 1935, Underwood & Sharp 4182 (NCU, NY, TENN); *Polk Co., below Parksville Dam, 20 July 1969, Rogers 44091 (SMU, TENN). Texas: *Bastrop Co., Bastrop State Park, August 1938, Tharp s.n. (GH, MO, SMU); *Nacogdoches Co., along farm Rt. 1087, 4-8 km E. of Rt. 259, 3 October 1965, Correll & Correll 31822 (NA, GH). *Orange Co., between Orange and Sabine R., 14 September 1968, Correll 36519 (GH, MICH). Virginia: ? Co; near Jamestown, September 1957 (fl), Irvine 93 (K); *Dinwiddie Co., 9-6 km WNW. of McKenny, 29 August 1960, Krai 11310 (GH); * Orange Co., Gordonsville, 1 September 1890, Blan- chard s.n. (MO, NY). West Virginia: *Pocahontas Co., Douthat Creek, near Minnehaha Springs, 6 August 1931, Core 3469 (NY). Wisconsin: * Adams Co., Adams Township, 19 August 1958, Hartley s.n. (GH). DOMINICAN REPUBLIC (introduced ?). Azua: Sierra de Ocoa, San Jose de Ocoa, Sabana de San Juan, c. 600 m, 17 March 1929 (fl & fr), Ekman H. 11947(8). BRAZIL (introduced ?). Rio Grande do Sul: *Osorio, 3 January 1950, Rambo s.n. (US). PARAGUAY (introduced ?). La Cordillera: Cordillera de Altos, 7 November 1902 (fr), Fiebrig 399 (BM, F, G, GH, K); Cordillera central, Cerros de Tobaty, September 1900 (fl), Hassler 6135 (BM, G, GH, K, MICH, P, S, W); near Pirebibuy, by Pirareta waterfall, 18 December 1965 (fr), Pedersen 7599 (C). 47 FRANCE (introduced). Gironde: Cazaux-lac [etang de Cazaux], 30 August 1931 (fl & fr), Tempere (see Tempere (1932), Jeanjean (1961), Heine (1962)). H. gentianoldes is clearly related to H. drummondii but is relatively reduced in all its parts. In particular it differs from H drummondii in being more branched, in the shorter leaves, and in the narrower capsules that much exceed the sepals. The stamens are sometimes reduced to 5, i.e. each fascicle is reduced to a single stamen. Whereas the distribution centre of H. drummondii appears to be Texas, that of H. gentianoides is further east, in the Louisiana to Florida area. The occurrences of this species in Brazil and Paraguay are almost certainly the result of long- distance dispersal; the population in the Gironde region of France, however, may be due to introduction by man. Rodriguez Jimenez (1973) treated H. cumulicola as a synonym of H. gentianoides, but it is a relative of H. denticulatum, not of H. drummondii. H. aphyllum Lundell, described from Belize, was likewise treated as synonymous with H. gentianoides by Rodriguez Jimenez, who thus acted on a suggestion by Adams (1962). Webb (1980) discussed this question and showed that the specimens from Florida that Lundell included in H. aphyllum belong to H. gentianoides. He, too, nevertheless included it in H. gentianoides 'until a series of populations of H. aphyllum from Central America can be critically examined and compared to H. gentianoides'. In my view, H. aphyllum is a species related to and derived from the Central American H. pratense Cham. & Schlechten- dal (see p. 107). Sect. 30. TRIGYNOBRATHYS (Y. Kimura) N. Rob- son in Bull. Br. Mus. nat. Hist. (Bot.) 16: 3 (1987). Ascyrum L.,Sp. PL: 787 (1753); Gen. PI. 5th ed.: 342 (1754), pro min. parte. Knifa Adans., Fam. pi. 2: 444, 541 (1763). Hypericum sect. Perforaria Choisy, Prodr. monogr. Hyperic.: 37, 44 (1821), pro. min. parte. Receveura Veil. Cone., Fl.flumin.: 137 (1829); op. cit., Atlas 5: tt. 119, 120(1831). Tridia Korth. in Tijdschr. Natuurl. Gesch. Physiol. 3: 17 (1836). Hypericum sect. Sarothra sensu Reichardt in Martius, Fl. bras. 12 (1): 185 (1878), pro parte excl. typum. Hypericum sect. Brathys sensu Reichardt in Martius, Fl. bras. 12 (1): 185 (1878), pro parte, excl. typum et H. tenuifolium A. St.-Hil. Hypericum sect. Brathys subsect. Spachium R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 3 (6): 214 (1893), pro parte excl. typum. Hypericum sect. Brathys subsect. Connatum R. Keller, torn. cit.: 214 (1893). Hypericum sect. Brathys subsect. Multistamineum R. Keller, torn, cit.: 214 (1893). Sanidophyllum Small in Bull. Torrey hot. Club 51: 391 (1924). Sarothra sect. Trigynobrathys Y. Kimura in Nakai & Honda, Novafl. jap. 10: 233 (1951). Sarothra sect. Spachium series Knifa (Adans.) Y. Kimura, torn, cit.: 233 (1951) ['Kniffa]. Sarothra sect. Spachium series Japonica Y. Kimura, torn, cit.: 233 (1951). 48 Hypericum sect. Knifa (Adans.) N. Robson in Bull. Br. Mus. nat. Hist. (Bot.) 16: 2 (1987), pro parte, incl. typum. Shrubs, subshrubs or perennial to annual herbs up to 2 m tall, the shrubs evergreen (or deciduous?), glabrous or very rarely with simple hairs, without dark glands, with taproot; branch- ing pseudo-dichotomous or dichasial/monochasial or mixed, from up to 10 nodes. Stems 4-(rarely 6-) lined and ± compressed (ancipitous)when young and in distal nodes of herbaceous species, sooner or later becoming terete in shrubby species, densely glandular (when young in shrubby species); cortex exfoliating in strips or persistent; bark smooth. Leaves opposite or rarely 3-whorled, decussate, sessile, free to completely perfoliate, not forming interfoliar ridge or with base sheathing, deciduous at base or persistent (in herbaceous species); lamina entire, with venation parallel to pinnate or 1-nerved or secondarily with up to 7 basal veins, open? or closed, with tertiary venation laxly to densely reticulate or absent; laminar glands punctiform or rarely united in lines or striae; marginal gland dots dense, very small; ventral resin glands absent. Inflorescence 1-flowered with branching pseudo-dichotomous (usually from uppermost node) or 2-°o-flowered with branching dichasial/monochasial from uppermost node, often with subsidiary branches from lower nodes; bracts and bracteoles foliar or reduced. Flowers stellate, homostylous. Sepals 5(4), free, persistent, erect in fruit, with margin entire; veins 1-11; laminar glands wholly linear to mostly punctiform; marginal, submarginal, and inframarginal glands absent. Petals 5(4), persistent, with apiculus terminal or subterminal to obsolete, acute to acuminate or obtuse; margin entire; marginal glands absent; laminar glands wholly linear or distally interrupted to punctiform or absent. Stamen fascicles basically 5(4?), with 1- c. 80 stamens per fascicle, fascicles free or usually all united to form continuous or interrupted narrow ring of stamens or more rarely united 2 + 2 + 1 or stamens irregularly grouped, persistent; filaments very shortly united or apparently free; anthers yellow, gland amber; pollen type VIII. Ovary with (2)3-5(6-8) parietal placentae, oo.Ovulate; styles (2)3-5(6-8), free from bases contiguous; stigmas clavate to broadly capitate. Capsule (2)3-5-valved, coriaceous to chartaceous, with valves obscurely narrowly longitudinally vittate, some- times with 1-3 flattish vesicles. Seeds ± narrowly cylindric, not carinate, without apical expansion; testa linear- scalariform to ribbed-scalariform or more rarely linear- reticulate to irregularly reticulate. BASIC CHROMOSOME NUMBERS (x). 12, 8(9); ploidy 1 ?,2,4. HABITAT. Open, damp (stream, river, and lake margins, marshes, bogs, damp grasslands, meadows, ditches, rice fields, dripping rocks, wet Pinus woods, Taxodium swamps, paramo) or dry (heaths, forest clearings, pastures, cultivated fields, roadside, dry or short grasslands, dry rocky places), on sand or clay; 0-2021 m (south of the Amazon), 0-4200 m (Andes (Chile) to Costa Rica), 0-3535 m (Nicaragua to Mexico), 0-2825 m (NW. America), 0-500 m (NE. America, Cuba), 0-3400 m (E. Asia, Australasia), 10-3500 m (Africa and Madagascar). DISTRIBUTION. Brazil (south and east), Uruguay, Argentina (north), Chile (central), Paraguay, Bolivia, Peru, Ecuador, Galapagos Is., Colombia, Venezuela, Panama, Costa Rica, Nicaragua, Honduras, El Salvador, Guatemala, Belize, Mexico, Cuba, U.S.A. (western to northern and eastern), N. K. B. ROBSON Canada (south-western to south-eastern); Japan, Ryukyu Is., Taiwan, Philippines Is., Korea (south), China (south-eastern and south), Bhutan, Sikkim, Nepal, India (Himachal Pra- desh, Uttar Pradesh, West Bengal, Meghalaya, Assam, Nagaland, Manipur, Orissa, Madras), Sri Lanka, Burma, Thailand, Laos, Vietnam, Kampuchea, Malaya, Sumatera, Java, Flores, Borneo, Sulawesi, Irian Jaya, Papua New Guinea; Australia (including Tasmania), New Zealand (North, South, and Stewart Islands); New Caledonia, the Hawaiian Islands; Nigeria (Bauchi Plateau), Sudan Republic, Ethiopia, Kenya, Uganda, Tanzania, Zaire (eastern and south-eastern), Burundi, Rwanda, Malawi, Zambia, Angola, Namibia (Caprivi, Waterberg Plateau), Botswana (south- eastern), Zimbabwe, Mozambique (central and south), Swa- ziland, South Africa (Transvaal, Natal, Cape Province south to Cape of Good Hope), Lesotho, Madagascar (centre and east). Introduced into the Azores, Ireland, Holland, France (Landes), Germany (Bavaria), Italy (Tuscany), Poland (Sile- sia), the U.S.S.R. (Georgia), and some species into the Dominican Republic, Japan, New Zealand, and the Hawaiian Islands. Key to sect. 30. Trigynobrathys7 1 Bracts and usually bracteoles foliar; plant a shrub or subshrub Bracts and bracteoles reduced, lanceolate to sub- ulate, or plant a perennial or annual herb 5 2(1) Pedicels 6-32 mm long; flowers 13-25 (rarely to 30) mm in diam.; plant erect or decumbent but rarely diffusely branched (1. rigidum pro parte) 3 Pedicels 2-3(-6) mm long; flowers 25-30 mm in diam.; plant decumbent, diffusely branched 2. microlicioides 3(2) Leaves narrowly elliptic or ± narrowly oblong, (10-) 20-60 mm long, acuminate to acute, base parallel- sided to narrowly cuneate; inflorescence up to 9- flowered Leaves narrowly to broadly elliptic or narrowly to broadly oblong to lanceolate, 10-20 mm long, acute to rounded, base narrowly cuneate to shal- lowly cordate; inflorescence up to 22-flowered .... Ic. rigidum subsp. sellowianum 4(3) Pedicels (9)12-32 mm long; capsule 8-9 mm long; petals 9-17 mm long la. rigidum subsp. rigidum Pedicels 8-10 mm long; capsule 5-7 mm long; petals 6-5-10 mm long lb. rigidum subsp. meridionale 5(1) Leaves coriaceous to subcoriaceous, ovate to broadly elliptic or broadly oblong or, if narrower, then whorled or subulate 6 Leaves chartaceous or thinner or, if coriaceous, then narrower than above but not whorled or subulate 18 6(5) Plant a subshrub; lateral inflorescence-branches, when present, with more than 1 node; leaves coriaceous; styles 3-5 Plant a perennial to annual herb; at least uppermost pair of lateral inflorescence-branches 1-noded; leaves subcoriaceous; styles 3(4) 14 7(6) Styles 5(4); leaf pairs ± connate or, if free, then broadly elliptic, spreading 8 Styles 3(4); leaf pairs free and not broadly elliptic, usually ascending to appressed 10 7N.B. All species in this section are completely glabrous except 12. H. setosum. HYPERICUM 49 8(7) Leaves free or shortly connate; sepals ovate-lanceo- late to lanceolate 3. teretiusculum Leaves % to completely connate; sepals ovate to oblanceolate or spathulate 9 9(8) Leaves chartaceous, oblong to ovate-deltoid (7-10 mm wide), with margin recurved to re volute, thin; sepals 3-4 mm long, subequal 4. caprifoliatum Leaves coriaceous, ovate-deltoid to semicircular (12- 40 mm wide), with margin revolute, incrassate; sepals 6-8 mm long, unequal 5. connatum 10(7) Leaves isomorphic, broadly ovate to narrowly elliptic or linear-lanceolate; petals 8-12 mm long or, if shorter (6-6-5 mm), then outer sepals foliaceous . . 11 Leaves heteromorphic or all linear; petals 6-8 mm long; outer sepals larger but not foliaceous 13 11(10) Outer sepals coriaceous, broadly ovate to rhombic, 5-7 mm long; pedicels 2-3 mm long (6. cordatum) 12 Outer sepals chartaceous, foliaceous, ovate to elliptic-oblong, 7-8 mm long; pedicels absent or almost so 7. cavernicola 12(11) Leaves broadly ovate to ovate-oblong (1: b = 1-2), margin incurved; inflorescence branches from 1-2 nodes; flowers (10)15-25 mm in diam da. cordatum subsp. cordatum Leaves triangular-lanceolate to narrowly elliptic, margin plane or almost so; inflorescence branches from 1-5 or more nodes; flowers 10-13 mm in diam 6b. cordatum subsp. kleinii 13(10) Leaves ovate to linear but not acerose, 1-8 mm wide, paired or in whorls of 3; inflorescence 1-11- flowered, branches from 1(2) nodes 8. ternum Leaves acerose, 0-8-1 mm wide, in whorls of 3: inflorescence c. 14-20-flowered, branches from up to 14 nodes 9. legrandii 14(6) Leaves plane, glabrous; styles 2-4 mm long; stamens 50-80 15 Leaves linear-subulate or scabrous-tomentose to pilose; styles 1-5-2 mm long; stamens 15-40 17 15(14) Stem not thickened at the base; upper leaves ovate or elliptic to obovate or lanceolate; inflorescence from up to 7 nodes; plant terrestrial (10. denticulatum) . . Stem thickened (aerenchymatous) at the base; upper leaves lanceolate to linear-lanceolate; inflorescence from up to 17 nodes; plant semi-aquatic or aquatic 11. harperi 16(15) Sepals 4-8 mm long; leaves ovate to lanceolate or elliptic (or lower obovate), usually appressed; lower internodes usually exceeding leaves lOa. denticulatum subsp. denticulatum Sepals 3-3-5 mm long; leaves lanceolate to oblong- elliptic or obovate, ascending or spreading; lower internodes usually shorter than leaves lOb. denticulatum subsp. acutifolium 17(14) Leaves and stem scabrous-tomentose to pilose; leaves ovate to lanceolate or oblong-elliptic 12. setosum Leaves and stem glabrous; leaves linear-subulate . . . 13. cumulicola 18(5) Uppermost bracteoles lanceolate or narrowly elliptic; leaves with 5-7 basal veins; plant a shrub Id. rigidum subsp. bracteatum Uppermost or all bracteoles linear or subulate, or leaves with 1 basal vein, or plant a perennial or annual herb 19(18) Styles 5(4-3); plant a shrub or subshrub or woody to Id wiry perennial or annual; capsule usually ovoid to globose 20 Styles 3(2) or, if 4-5, then plant a herbaceous perennial or annual; capsule usually narrowly cylindric or ellipsoid to ovoid-ellipsoid or ovoid- cylindric 34 20(19) Capsule cylindric or narrowly ovoid-pyramidal, ex- ceeding or equalling sepals (mature fruit of Sp. 15 unknown) 21 Capsule ovoid to globose, equalling or shorter than sepals (sometimes except sp.22) 23 21(20) Inflorescence l(2-5)-flowered, branching (in Sp. 14) pseudo-dichotomous; leaves 3-13(-18 mm) long, basal or near-basal veins 1-5 22 Inflorescence (at least terminal) c. 9-24-flowered, branching dichasial/monochasial; leaves 8-30 mm long, basal or near-basal veins 3-9 16. brasiliense 22(21) Leaves plane, 4-18 mm long, 3-5-nerved; habit erect to decumbent; sepals 3-5-5 mm long, (3)5-nerved 14. denudatum Leaves incurved, 3-4 mm long, 1-nerved; habit prostrate to ascending; sepals 2-2-5 mm long, 3- nerved 15. pedersenii 23(20) Sepals unequal, often foliaceous, outer broadly rhombic-elliptic or ovate-lanceolate to broadly ovate, 3-5-5 mm long 24 Sepals equal to subequal or, if markedly unequal, then narrow and/or shorter 25 24(23) Leaves linear to narrowly oblong, 0-5-1 -5(-2) mm wide, base not decurrent, basal vein 1; styles 1-1-3 x ovary 17. polyanthemum Leaves narrowly oblong to narrowly lanceolate, 2-5-5 mm wide, base decurrent-amplexicaul, basal veins 3-5; styles 1-3-1-7 x ovary 24. anceps 25(23) Leaf-base decurrent; capsule broadly ovoid to glo- bose; styles 5(4) 26 Leaf-base not decurrent or, if so, narrowly ovoid to cylindric; styles 3(4-5) 27 26(25) Flowers (12-)15-25 mm in diam.; inflorescence l-7(- 17)-flowered; styles 2-5-3 mm long; leaves 5-15 mm wide 23. carinatum Flowers 8-10 mm in diam.; inflorescence 15-c. 50- flowered; styles 2-2-5 mm long; leaves 2-3(-5-5) mm wide 24. anceps 27(25) Capsule (4-)5-6 mm long, ± broadly ovoid; flowers (8-)10-24 mm in diam.; styles mostly 5 (18. campestre) 28 Capsule 2-4-5 mm long, subglobose to globose (or rarely ovoid in 21. lorentzianum); flowers 2-10(- 12) mm in diam.; styles mostly 4-3 30 28(27) Inflorescence 1-5-flowered; flowers 18-24 mm in diam 18a. campestre subsp. pauciflorum Inflorescence 6-c. 65-flowered; flowers 8-15(-18) mm 29 29(28) Leaves 3-7 mm wide (1: b = 4-10); plant suffruticose to perennial; styles 5 18b. campestre subsp. campestre Leaves 1-3-5 mm wide (l:b = 10-15); plant annual; styles 5-4(3) 18c. campestre subsp. tenue 30(27) Inflorescence overall ellipsoid to narrowly cylindric; terminal inflorescence usually 3-25-flowered, lat- eral ones l-10(-14)-flowered 31 Inflorescence overall obovoid to densely corym- biform; terminal inflorescence (7-)30-c. 200- flowered, lateral ones (at least upper) not or scarcely fewer-flowered 32 50 N. K. B. ROBSON 31(30) 32(30) 33(32) 34(19) 35(34) 36(35) 37(36) 38(34) 39(38) 40(39) 41(40) 42(41) Sepals 3-6 mm long; petals 4-8 mm long; stamens 14- 32; styles 1-2 mm long; capsule 4-5-8 mm long .... 25a. silenoides subsp. silenoides Sepals 2-3 mm long; petals 2-5-3-5 mm long; stamens 9-10; styles 0-7-0-9 mm long; capsule 3-4 mm long 25b. silenoides subsp. minus Capsule ± broadly ovoid-ellipsoid, 3-4 mm wide, shorter than sepals; inflorescence mixed pseudo- dichotomous/dichasial/monochasial; lax subshrub to c. 1m tall 28. relictum Capsule narrower and/or exceeding sepals; inflor- escence wholly dichasial/monochasial (rarely mixed in spp. 42, 46, and 47), perennial (sometimes woody) or annual herbs to 0-85 m tall 39 Stigma narrowly to broadly capitate or subpeltate ... 40 Stigma scarcely capitate to clavate (African spp.) ... 61 Leaf-base cuneate or parallel-sided, separated from adjacent one by interfoliar boss; subshrubs or perennial herbs 41 Leaf-base ± amplexicaul or decurrent, not separated from adjacent one; perennial or annual herbs 43 Leaves subcoriaceous, flowers 12-16 mm in diam.; styles 2-5-3 mm long; sepals elliptic 29. killipii Leaves chartaceous; flowers 5-10 mm in diam.; styles 0-5-1-6 mm long; sepals lanceolate to oblong or oblanceolate 42 Stems erect or decumbent, not rooting, usually perennial; leaves elliptic-oblong or lanceolate to linear, 8-27 mm long; capsule ovoid-cylindric to cylindric 30. thesiifolium Stems ascending to prostrate, rooting, usually annual; 43(40) 44(43) Leaves 7-20 mm long, narrowly elliptic to oblan- ceolate, chartaceous, margin plane to recurved; sepals 3-5 mm long, equal 19. linoides Leaves 3-5 mm long, oblong to hastate, sub- coriaceous, margin incurved; sepals 1-5-3 mm long, unequal 20. salvadorense Flowers 7-10 mm in diam.; sepals 2-5-4 mm long, acute; styles (5)4(3) 21. lorentzianum Flowers 2-4 mm in diam.; sepals 0-7-2 mm long, subacute to rounded; styles 3 (22. myrianthum) ... 33 Leaves spreading, margin plane or subrecurved, base cuneate to parallel-sided; sepals narrowly oblong or narrowly elliptic, veins scarcely prominent 22. myrianthum subsp. myrianthum Leaves ascending to erect, margin incurved, base parallel-sided to cordate-aplexicaul; sepals broadly oblong or broadly elliptic to elliptic-ovate, veins prominent 22b. myrianthum subsp. tamariscinum Leaf-bases decurrent-amplexicaul, forming V 35 Leaf-bases not decurrent, if amplexicaul then not forming V 38 Leaves lanceolate or elliptic to oblong or linear, not glaucous; petals oblong-oblanceolate to oblong ... 36 Leaves (at least upper) triangular-lanceolate to 46(45) oblong or obovate, glaucous; petals obovate to oblanceolate 27. gramineum Leaves 1-9 mm wide, lanceolate to elliptic or nar- rowly oblong or, if linear, then apex acute; stems erect to decumbent, not rooting; capsule 1-3-2 x as long as sepals (25. silenoides) 37 Leaves 0-4-1 mm wide, linear, apex rounded; stems decumbent to prostrate, rooting; capsule 1-1-3 x as long as sepals 26. caespitosum 45(44) 47(43) 48(47) 49(48) 50(49) 51(47) 52(51) leaves elliptic to narrowly oblong or oblanceolate, 5-10 mm long; capsule ellipsoid to cylindric 31. brevistylum Capsule ovoid-conic to ovoid, apex acute to acumin- ate; styles (l-)2-5 mm long; leaves coriaceous or subcoriaceous 44 Capsule ovoid-ellipsoid or ellipsoid to cylindric or globose, apex obtuse to rounded; styles 0-5-2(-3) mm long; leaves chartaceous to membranous 47 Leaves oblong or elliptic to linear or oblanceolate (broadest at or above middle), margin often scabrid-denticulate; upper stem internodes about equal 45 Leaves normally lanceolate to linear (broadest below middle), margin entire; upper stem nodes progres- sively elongate 44. paucifolium Styles (2-)2-5-4 mm long, longer than ovary; lower leaves usually spreading; flowers (12-)15-20 mm in diam., petals densely red-tinged 32. pauciflorum Styles 1-2 mm long, shorter than ovary; lower leaves usually ascending to appressed; flowers 10-16 mm in diam., petals usually only red-veined (43. pumil- lum) 46 Leaves with margins and midrib (beneath) scabrid- denticulate or, if smooth, then lamina linear or narrowly elliptic; stems erect to decumbent; styles 1-5-2 mm long; capsule ovoid-conic 43a. pumillum subsp. diffusum Leaves with margins and midrib smooth, lamina oblong-elliptic to obovate; stems decumbent to prostrate; styles 1-1-5 mm long; capsule ovoid .... 43b. pumillum subsp. pumillum Leaves linear or lanceolate or narrowly elliptic to linear-subulate; stems erect, sometimes from de- cumbent base 48 Leaves ovate or oblong to oblong-linear or obovate; stems erect to prostrate 51 Stems caespitose, branching at or near base; inflorescence-branching usually regularly dichasial for at least two nodes; flowering branches from up to 5 nodes 45. moranense Stems single, branching (at least at first) at about the middle; inflorescence-branching usually monocha- sial after first node; flowering branches from 1-2 nodes 49 Leaves plane to recurved, narrowly elliptic or nar- rowly lanceolate or narrowly oblong to linear; capsule 1 -5-4 mm wide 50 Leaves incurved, triangular-subulate to linear- subulate; capsule 1-1-3 mm wide 35. aphyllum Leaf base parallel-sided; styles 1-5-3 mm long; stamens 12-20 33. pratense Leaf base cuneate; styles 0-6-0-8 mm long; stamens 5-7 34. philonotis Stigmas narrowly to scarcely capitate; stems (at least at base) rooting and ± diffuse; capsule usually shorter than or equalling sepals 52 Stigmas broadly capitate; stems erect or rarely (Sp. 42 p.p.) rooting and diffuse; capsule usually exceeding sepals 53 Sepals (outer) and leaves (except lower) narrowly oblong or narrowly elliptic-oblong; leaves 1-3- nerved; capsule broadly ellipsoid 46. par vu him Sepals (outer) and leaves broadly oblong or broadly HYPERICUM 53(51) 54(53) elliptic to obovate; leaves 3-7-nerved; capsule ellipsoid or cylindric or subglobose 47. anagalloides Leaves lanceolate or narrowly elliptic to linear or oblanceolate-linear (1: b = c. 2-5-9); capsule broadest below middle 54 Leaves ovate (rarely lanceolate) to broadly oblong- elliptic or obovate (1: b = 1-2-5); capsule usually broadest at middle 55 Leaves lanceolate to narrowly elliptic, usually 6-12 mm wide, (3)5-7-nerved; inflorescence usually compact 36. majus Leaves linear to oblanceolate-linear, 0-5-4(-5-5) mm wide, l-3(-5)-nerved; inflorescence usually rather lax 37. canadense 55(53) Capsule equalling or exceeding sepals; inflorescence- branching never sympodial 56 Capsule shorter than sepals or, if longer, then inflorescence-branching sympodial 42. japonicum 51 caul, basal veins 5-9, almost equally prominent; plant not glaucous 63 63(62) Pedicels 5-18 mm long; leaf margin plane; styles 3(4), 1-1-5 mm long; stamens 15-45 64 Pedicels 2-4 mm long; leaf margin undulate; styles (4)5, 0-3-0-5 mm long; stamens 12-15 52. oligandrum 64(63) Leaf base cordate; sepals 5-6 mm long; stamens 36- 45; styles 1 -5 mm long 50. humbertii Leaf base rounded to broadly cuneate; sepals 3-4 mm long; stamens 15-30; styles 0-7-1-5 mm long 51. scioanum Subsect. 1. Connatum (R. Keller) N. Robson, comb, nov. Receveura Veil. Cone. Fl. flumin: 137 (1829). Type: R. cordata Veil. Cone. (= H. connatum Lam.). 56(55) Capsule narrowly conic-ellipsoid; branches narrowly Hypericum sect. Brathys subsect. Connatum R. Keller in ascending, often confined to inflorescence; middle Eng!- & Prantl, Nat. Pflanzenfam. 3(6): 214 (1893). Type: and upper leaves ovate-deltoid, usually subacute .. H. connatum Lam. 38. gymnanthum Hypericum sect. Brathys subsect. Multistamineum R. Keller, Capsule narrowly ovoid or cylindric to ellipsoid- torn, cit.: 214 (1893). Type: H. pilosum Walter = H. subglobose; branches ± widely spreading, usually setosum L. from below middle of stem; middle and upper Sanidophyllum Small in Bull. Torrey hot. Club 51: 391 leaves lanceolate-deltoid or ovate-oblong or orbi- (1924) T s cumulicola Small = H. cumulicola (Small) cular, obtuse to rounded 57 r» A j P. Adams. 57(56) Styles 5; bracts foliar; stems branched from base Hypericum sect. Knifa (Adans.) N. Robson in Bull. Br. Mus. 41. pleiostylum nat. Hist. (Bot.) 16: 2 (1987), pro parte excl. typum. Styles 3; if stem branched from base, then bracts subulate 60 TyPe: H~ connatum Lam. co/c-?\ e* h t Shrubs, subshrubs or woody annual herbs with leaves free or 58(57) Stems erect from decumbent and rooting base; . J . inflorescence-branching strictly dichasial/monocha- sometimes ± connate in pairs, styles 5(4-3), and capsules sial (39. mutilum) 59 usually broadly ovoid to globose, or herbaceous perennials Stems decumbent to ascending, rooting above base; with styles 3, capsules narrowly ovoid to cylindric, and leaves inflorescence-branching pseudo-dichotomous at ± decurrent, forming V. first then (dichasial/)monochasial . . 40. arenarioides 1. Hypericum ngidum A. St. Hil., Fl. Bras. mend. 1: 336 59(58) Stem with apical internode shorter than adjacent one (1828); Walp., Repert. hot. syst. 1: 390 (1842); D. Dietr. or absent; inflorescence-branching mostly dichasial Syn pi. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 60 12(1); 189 (1878), pro parte quoad typum; Lyman B. Smith Stem with apical internode well developed, usually in j Wash Acad Sd 48: 314 (1958), pro parte quoad longer than adjacent one; inflorescence-branching typum; Angely, FL Anal. Parana: 451 (1965), FL Anal. S. repeatedly monochasial distally ... Pflu/o j. m (1%9) Rodr{ Jim6nez in ReitZ5 FL HL 39b. mutdum subsp. lat.sepa.um ^^ ^^ £ f lA\mo^ pro parte exd. syn. „. 60(59) Bracts subulate; leaves paler beneath, rarely broadly microlicioides Lyman B. Smith. Type: Brazil, Parana, ovate to suborbicular 39a. mutilum subsp. mutilum prope urbem Curityba, in parte australi provinciae Sancti Bracts foliar; leaves concolorous, broadly ovate to Pauli', March [1820] 1816-1821, St.-Hilaire 1631 pro parte suborbicular 39c. mutilum subsp. boreale (P!-holotype; Fl-photograph). 61(39) Capsule narrowly ovoid, acuminate, exceeding se- p- ^ Macs 10 11 pals; bracts and upper leaves acute, linear; flowers in regular dichasial/monochasial cymes; plant erect Icon: Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc.: or decumbent at base only 48. lalandii ft t j A (1980). Capsule ellipsoid to ovoid-cylindric or subglobose, rounded, equalling or shorter than sepals; bracts Shrub or subshrub (0-25)0-5-2 m tall, erect or decumbent and and upper leaves obtuse to rounded or, if acute, branching at the base, with branches ascending-pyramidal or then linear; flowers terminal or pseudo-axillary, ± diffuse, pseudo-dichotomous or lateral. Stems dull red- usually with shoots in axils of apical node, rarely in brown, 4-lined when young, eventually terete, cortex ex- irregular cymes; plant decumbent to prostrate .... 62 foliating in strips; internodes 3-30(^5) mm long, shorter to longer than leaves. Leaves sessile, spreading; lamina 8-60 x 62(61) Leaf with base narrowly cuneate to PnOeMfed; | linear-oblong or narrowly to broadly elliptic or basal vein 1 or, if 3-5, then with laterals much less f prominent; plant usually ± glaucous ovate-oblong to rarely lanceolate, plane or margin recurved, 49. globuliferum paler and ± reddish beneath, not glaucous, coriaceous; apex Leaf with base broadly cuneate to cordate-amplexi- acute or subacuminate to rounded, base parallel-sided to 52 N. K. B. ROBSON Map 10 Sect. 30: la. H. rigidum subsp. rigidum •; Ib. H. rigidum subsp. meridionale O; 4. H. caprifoliatum •. Records for la and Ib include some from Rodriguez Jimenez (1973). cordate, free; basal or near-basal veins 7, midrib pinnately branched, tertiary reticulum lax; laminar glands dense, prominent beneath or obscure. Inflorescence l-9(-22)- flowered, dichasial and sometimes pseudo-dichotomous, sometimes with branches from up to 4 nodes below, the whole cylindric to pyramidal; primary pedicels 6-20(-32) mm long; bracts foliar, gradually decreasing in size or rarely suddenly reduced, the ultimate ones very rarely lanceolate- subulate (see below). Flowers 13-25(-30) mm in diam., stellate; buds ellipsoid, acute. Sepals 5-13 x 0-9-3 mm, equal, not imbricate, narrowly oblong or oblong-linear to narrowly elliptic-oblong or narrowly elliptic, acute; veins 5(7), midrib sometimes prominent; glands proximally linear, punctiform in upper c. 0-3. Petals yellow to orange (when fading?), 6-5-14(-17) x 3-5-6(-8) mm, 1-2-1-4 x sepals, oblanceolate to oblong-obovate; apiculus acute; glands linear to elongate-punctiform. Stamens (30-)40-75, 5-fascicled, longest 3-5-6 mm long, c. 0-4-0-6 x petals. Ovary 1-5-2-5 x 1-1-6 mm, ovoid; styles 5(4), l-5-3(-3-5) mm long, 1-1-5 x ovary, spreading; stigmas clavate. Capsule 4-5-9 x 3-6 mm, broadly ellipsoid, obtuse to subrounded, shorter than sepals. Seeds 0-6-0-8 mm long; testa ribbed-scalariform. Streamsides, damp meadows, heaths, and bogs; 700-1700 m. Brazil (Minas Geraes and Sao Paulo to northern Rio Grande do Sul). H. rigidum includes the morphologically most primitive forms in sect. Trigynobrathys, some plants from Parana being the nearest to the African H. revolutum subsp. keniense (sect. 1. Campy losporus). Such plants are likewise nearest in their section to H. terrae-firmae (from Belize), the most primitive member of sect. 29. Brathys. H. rigidum was misidentified by Reichardt (1878) as a member of the H. cordatum (=cordiforme) - H. ternum group (Spp. 6-9), and his lead was followed by L. B. Smith (1958). The type specimen, however, belongs to the species that Reichardt described as H. rufescens. H. rigidum is very variable, there being reduction trends from the large- and few-flowered, long-leaved, shrubby forms of Parana to the small- and numerous-flowered, short-leaved, suffruticose forms of Minas Geraes and Santa Catarina/Rio Grande do Sul. The variation along these trends is almost continuous, but it is possible to recognize three main subspecies and one (subsp. bracteatum) that forms an important link with other parts of sect. Trigynobrathys. la. Hypericum rigidum subsp. rigidum Map 10. H. rufescens Klotzsch ex Reichardt in Martius, Fl. bras. 12 (1): 194 (1878), pro parte excj. typum. Shrub, erect, branching pyramidally. Leaves 20-60 x 4-13 mm, narrowly elliptic or narrowly oblong to linear-oblong, apex acute to acuminate, base parallel-sided. Inflorescence 1- 9-flowered. Flowers 20-30 mm in diam., pedicels (9)12-32 mm long, bracts foliar. Petals 9-17 mm long. Stamens (50-) 60-75, longest 5-6 mm long. Styles 5. Capsule 8-9 mm long. Brazil (Parana). BRAZIL. Parana: Serrinha, 8 February 1912 (fl), Dusen 13618 (BM, GH, K, S); mun. Tijucas do Sul, Campina, 14 February 1978 (fl), Kummrow 1214 (BM, MBM); mun. S. Jose dos Pinhaes, Rio Piqueno, 4 March 1980 (fl & fr), Hatschbach 42774 (BM, MBM). Ib. Hypericum rigidum subsp. meridionale (Lyman B. Smith) N. Robson, stat. nov. Map 10. H. rufescens Klotzsch ex Reichardt in Martius, Fl. bras. 12 (1):194 (1878); Angely, Fl. Anal. Parana: 451 (1965), pro parte quoad typum, non Lam. (1795). Type: Brazil, [Parana?], without precise locality or date (fr), Sellow s.n. (Bt [F!, GH!, NY!-photographs]-lectotype; G!, K!, NY!, S!, W!). Reichardt cites five numbered specimens, which I have not seen. The photographs of the Berlin unnumbered specimen, however, show that the specimen was named in 53 Fig. 12 A. H. rigidum subsp. rigidum: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) stamens (partly cut away) and ovary; (f) young capsule. B. H. rigidum subsp. meridionale: (g) habit. C. H. rigidum subsp. sellowianum: (h) habit (a, g, h x '/z; b x 1; c, d x : .4; e, f x 2). A. Kummrow 1214; B. Hatschbach 12027; C. Smith 11002. 54 N. K. B. ROBSON Klotzsch's writing. In the absence of other authentic material, therefore, it can be chosen as lectotype. The existence of photographs of this specimen, even though it itself is no longer extant, would seem to preclude the necessity of selecting a neotype. H. meridionale Lyman B. Smith in J. Wash. Acad. Sci. 48: 311 (1958); Angely, Fl. Anal. Parana: 451 (1965). Type as for H. rufescens Klotzsch ex Reichardt non Lam. H. rigidum sensu Rodriguez Jimenez in Reitz, Fl. III. Catar. Hipericdc.: 29(1980), pro parte quoad syn. H. rufescens Klotzsch ex Reichardt. Shrub orsubshrub, erect or spreading, branching ± diffusely. Leaves 10-30 x 2-8 mm, ± narrowly elliptic or oblong to linear-oblong, apex acute to acuminate, base parallel-sided to narrowly cuneate. Inflorescence 1-7-flowered. Flowers 14-24 mm in diam., pedicels 8-10 mm long, bracts foliar. Petals 6-5- 10 mm long. Stamens 40-65, longest 5-6 mm long. Styles 5. Capsule 5-7 mm long. Brazil (Sao Paulo, Parana, Santa Catarina). BRAZIL. Sao Paulo: mun. S. Paulo, Tres Barras, Exta Sta. Catarina, 26 February 1940 (fl & fr), Mattos & Labouriau R.J. 63284 (P, NY). Parana: Quatro Barras, Rancho Velho, 15 December 1964 (fl), Hatschbach 12027 (F, K); mun. Piraquera, entre Rio Taquari e Rio Divisa, 13 March 1949 (fl), Hatschbach 1213 (US). Santa Catarina: mun. Mafra, 2 km S. of Mafra by Estrada da Rodagem Federal, c. 800 m, 11 March 1957 (fl), Smith & Klein 12077 (F, GH, US). Subsp. meridionale differs from subsp. rigidum in the smaller size of leaf and flower and the diffuse habit. The size characters all overlap except, apparently, the capsule length, but in practice it is usually possible to allot a given specimen to a subspecies without difficulty. Ic. Hypericum rigidum subsp. sellowianum (R. Keller) N. Robson, stat. nov. Map 11. H. sellowianum R. Keller in Bot. Jb. 58: 195 (1923); Lyman B. Smith in J. Wash. Acad. Sci. 48: 311 (1958). Type: Brazil, without precise locality or date (fr), Sellow 468 (Bt- holotype; F!, GH!, NY!-photographs). H. rigidum sensu Rodriguez Jimenez in Reitz, Fl. 111. Catar., Hipericdc.: 29 (1980), pro parte. Shrub or subshrub, erect or spreading, branching ± diffusely. Leaves 10-20 x 4-6 mm, broadly to narrowly elliptic or Map 11 Sect. 30: Ic. H. rigidum subsp. sellowianum •; Id. H. rigidum subsp. bracteatum D. Records for Ic include some from Rodriguez Jimenez (1973). broadly to narrowly oblong or lanceolate; apex acute to rounded, base narrowly cuneate to shallowly cordate. Inflor- escence 1-22-flowered. Flowers 13-20 mm in diam., pedicels 6-16(-20) mm long, bracts foliar or rarely ultimately reduced. Petals 9-12 mm long. Stamens 30-65, longest 3-6 mm long. Styles 4-5. Capsule 4-5-8 mm long. Brazil (Minas Geraes, Sao Paulo, Parana, Santa Catarina, Rio Grande do Sul). BRAZIL. Minas Geraes: Passo Quatro, Faz. Sertao dos Martins, margens do Rio S. Lourenc.o Velho, 22 April 1948 (fl & fr), Barbosa & Aranjo 144 (MO, NY, P). Sao Paulo: Barreiros, Serra da Bocaina, 24 November 1950 (fl), Sagadas- Vianna 3084 (GH, NY, P). Parana: lOp. W. of S. Mateus do Sul, 9 February 1966 (fl), Hatschbach [Haas & Lindeman] 13796 (NY, U, US). Santa Catarina: S. Joachim, Fachinal, 1000 m, 29 January 1950 (fl), Reitz 3400 (NY, US); mun. Chapeco, 24 km W. of Campo Ere, Fazenda Campo Sao Vicente, 900-1000 m, 27 December 1956 (fl), Smith, Reitz & Sufridini 9406 (S, US). Rio Grande do Sul: Bom Jesus, Aparados da Serra, 15 February 1947 (fr), Rambo 35426 (S, US). Subsp. sellowianum differs from subsp. meridionale in its more suffruticose (even herbaceous?) habit, shorter and relatively broader leaves with apex less acute and base never parallel-sided, and the often more numerous flowers in the terminal inflorescence (and the lateral ones). The most HYPERICUM primitive forms are in Sao Paulo (Serra da Bocaina), whence there are trends north to Minas Geraes (Serra da Manti- queira) and south to northern Rio Grande do Sul (Aparados do Serra). These trends are morphologically similar, but the extreme southern form has lanceolate, cordate leaves. The subspecies is apparently rarer in Parana than in the other states, the specimens from there showing a tendency towards 3. H. teretiusculum. Keller described the ovary as 3-styled and the capsule as 3- valved; but the type appears to belong to the primitive, Sao Paulo form, which has 5 styles. Forms with 4 styles occur only towards the respective extremes of the northward and southward trends. Id. Hypericum rigidum subsp. brack-alum N. Robson, stat. et nom. nov. Map 11. 55 H. rigidum var. brevifolium A. St. Hil., Fl. Bras, merid. 1: 336 (1828). Type: Brazil, Parana, 'prope urbem Curityba', [1820] 1816-1821 (fl), St.-Hilaire 1631 pro parte (P!- holotype). Shrub or ? subshrub, erect, with branches strict. Leaves 15-20 x 4-6 mm, narrowly elliptic to narrowly oblong-elliptic; apex acute to subacute, base narrowly cuneate to truncate. Inflorescence c. 5-12-flowered, with 1-3-flowered branches from c. 3 nodes below. Flowers c. 15-18 mm in diam., pedicels 3-6 mm long, bracts reduced foliar, 5-8 mm long, narrowly elliptic. Petals 8-10 mm long. Stamens c. 50, longest c. 4 mm long. Styles 5. Capsule not seen. Brazil (Parana). BRAZIL. Parana, near Curitiba, 1816-1821 (fl), St.- Hilaire 1631 (P). Subsp. bracteatum differs from the more erect form of subsp. meridionale only in having leaves in the flowering region reduced in size and appearing as bracts. The only known specimen, however, forms a link between subsp. rigidum, on the one hand, and (respectively) the brasiliense group (Spp. 14-27) and subsect. Knifa (Spp. 28-52), on the other. Although it shares a collection number (St.-Hilaire 1631) with the type of the species, it is quite distinct from it and may even deserve recognition at the species level. At any rate, it is unlikely that it represents merely intra-population variation. 2. Hypericum microlicioides Lyman B. Smith in J. Wash. Acad. Sci. 48: 311, f.l, g-i (1958). Type: Brazil, Santa Catarina, mun. de Campo Alegre, Morro Iquererim, 1500 m, 10 December 1956 (fl), Smith & Klein 8535 (US!- holotype; P!, NY!-isotypes). Map 12. H. rigidum sensu Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc.: 29 (1980), pro parte, quoad syn. H. microli- cioides. Icon: Lyman B. Smith in /. Wash. Acad. Sci. 48: 312, f. Ig-i (1958). Subshrub or woody herb 0-15-0-45 m tall, decumbent and branching at the base, with branches diffuse, lateral or rarely pseudo-dichomotous. Stems dull red-brown, 4-lined when young, soon terete, cortex exfoliating in strips; internodes l-5-10(-30) mm long, shorter to longer than leaves. Leaves sessile, appressed or tardily spreading; lamina 8-20 x 3-6 mm, narrowly elliptic or narrowly oblong to lanceolate, plane or usually margin subrevolute, paler (? and reddish) beneath, not glaucous, subcoriaceous; apex acute to obtuse, base cuneate, free; basal or near-basal veins 5-7, midrib pinnately branched, tertiary reticulum lax; laminar glands dense, obscure. Inflorescence 1-flowered, terminal and on lateral branches; pedicels 2-3(-6) mm long; bracts foliar. Flowers 25-30 mm in diam., stellate; buds ovoid, acute. Sepals 8-12 x 2-3 mm, equal, not imbricate, narrowly elliptic to linear, acute to acuminate; veins 7-9, midrib prominent; glands mostly punctiform. Petals yellow, 13-16 x 6-8 mm, 1-2-1-4 x Map 12 Sect. 30: 2. H. microlicioides O; 3. H. teretiusculum 56 N. K. B. ROBSON sepals, obovate; apiculus subacute; glands linear, distally absent. Stamens c. 70, 5-fascicled, longest 4-5 mm long, c. 0-3 x petals. Ovary 2-5 x 1 mm, ovoid; styles 5, c. 2 mm long, spreading; stigmas clavate. Capsule and seeds not seen. Alpine bogs (above tree-line); (950-) 1300- 1500 m. Brazil (Parana, Santa Catarina). BRAZIL. Parana: mun. Curitiba, Atuba, 950 m, Novem- ber 1960 (fl), Hatschbach (7468) (US); mun. Guaratuba, Serra do Arac,atuba, 1300 m, 19 November 1971 (fl), Hatschbach 28096 (US). Santa Catarina: see type. H. microlicioides is a higher-altitude derivative of H. rigidum subsp. rigidum, more suffruticose or even herbaceous and more diffuse in habit, with leaves usually shorter and usually appressed, shorter pedicels, and fewer, larger flowers. It seems possible that the bitopic distribution is the result of parallel evolution of two populations, but it is more likely due merely to division and isolation of one. 3. Hypericum teretiusculum A. St. Hil., Fl. Bras, merid. 1: 33 (1828); Walp., Repert. hot. syst. 1: 389 (1842); D. Dietr., Syn. pi. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 12 (1): 194 (1878); Lyman B. Smith in/. Wash. Acad. Sci. 48: 313 (1958); Angely, Fl. Anal. Parana: 451 (1965); Rod- riguez Jimenez in C.r. Soc. Biogeogr. 432: 91, map 6 (1972), in Mems Soc. Cienc. nat. La Salle 33: 127, f. 15B (1973), in Reitz, Fl. III. Catar., Hipericdc.: 27, f. 4B (1980). Type: Brazil, Sao Paulo, 'prope flumen Tarere [Itarare] in parte australi provinciae Sancti Pauli', [1820] 1816-1821 (fl), St.-Hilaire 1385 (P!-holotype; US! -photograph). Map 12. Sarothra teretiuscula (A. St. Hil.) Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 71 (1951); Angely, Fl. Anal. S. Paulo 1: 187(1969). H. subliberum Lyman B. Smith in J. Wash. Acad. Sci. 48: 311, fig. la-f (1958). Type: Brazil, Santa Catarina, mun. Curtibanos/Campos Novos, 22 km W. of Curtibanos towards Campos Novos, 850-950 m, 5 December 1956 (fl), Smith & Klein 8318 (US!-holotype; HER, NY!, P!, R- isotypes). Icones: Lyman B. Smith in/. Wash. Acad. Sci. 48: 312, f.la-f (1958); Rodriguez Jimenez in Reitz, FL III. Catar., Hiper- icdc.: 18, t. 4B (1980). Subshrub 0-4-1 m tall, erect, 1-stemmed or branched near base, with branches ascending-pyramidal or spreading, lateral only. Stems dull red-brown, 4-lined when young, soon 2-lined to subterete, cortex exfoliating in strips; internodes 9-70 mm long, usually exceeding leaves. Leaves sessile, spreading; lamina (10-)14-27 x 5-17 mm, broadly elliptic to broadly ovate, plane or margin recurved, subconcolorous, not glaucous, coriaceous; apex acute to rounded, base broadly cuneate to cordate, free or shortly connate; basal veins 3-5, midrib pinnately branched, tertiary reticulum dense; laminar glands rather dense to sparse, prominent beneath or obscure. Inflorescence (3-)10-°°-flowered, dichasial/monochasial from 1-2 nodes, sometimes with lateral branches from nodes immediately below, the whole corymbiform to broadly paniculate; primary pedicels 3-4 mm long; bracts linear- lanceolate, acute. Flowers 10-12 mm in diam., stellate; buds ovoid-triangular, acute. Sepals 5-7 x 2-2-5 mm, subequal, imbricate, ovate-lanceolate to lanceolate, subacuminate; veins 5-7, midrib prominent; glands linear, sometimes dis- tally punctiform. Petals yellow, 7-10 x 3-5^-5 mm, c. 1-4 x sepals, oblong-obovate; apiculus subacute; glands linear to elongate-punctiform. Stamens c. 40-70, obscurely 5-fascicled, longest 3-4 mm long, c. 0-5 x petals. Ovary c. 2 x I mm, ovoid-ellipsoid; styles 5, 2-2-5 mm long, c. 1-1-2 x ovary, spreading; stigmas subclavate. Capsule 4-5 x 3-5^ mm, ovoid-subglobose, obtuse to shortly rostrate, about equalling sepals. Seeds 0-7-0-8 mm long; testa ribbed-scalariform. Damp meadows and bogs; 840-1000 m. Brazil (Sao Paulo?, Parana, Santa Catarina, Rio Grande do Sul), Paraguay (Guaira)? BRAZIL. Parana: mun. Jaguariaiva, outskirts of Jaguar- iaiva on Arapoli road, 840 m, 17 January 1965 (fl & fr), Smith, Klein & Hatschbach 14639 (US); mun. Piraquara, Nova Tirol, 5 April 1979 (fl & fr), Hatschbach 42186 (BM, MEM). Santa Catarina: mun. Chapeco, Fazenda Campo Sao Vicente, 24 km W. of Campo Ere, 900-1000 m, 27 December 1956 (fl & fr), Smith, Reitz & Sufridini 9342-A (A, S). Rio Grande do Sul: ?(see below). PARAGUAY. Guaira: Cordillera de Villa-Rica, January 1905 (fl & fr), Hassler 8758 (K). H. teretiusculum is related to H. rigidum subsp. sellowianum through the somewhat anomalous, multiflowered Parana populations of that subspecies. It differs in being less woody, with larger, broader, cordate leaves and reduced bracts. The Paraguay record is doubtful, because the Geneva (G) specimen of Hassler 8758 is typical H. connatum. If the type specimen came from the north bank of the rio Itarare, then H. teretiusculum also occurs in Sao Paulo. 4. Hypericum caprifoliatum Cham. & Schlechtendal in Linnaea 3: 125 (1828); Walp., Repert. hot. syst. 1: 389 (1842); D. Dietr., Syn. pi. 4: 1237 (1847); Reichardt in HYPER/CUM 57 Martius, Fl. bras. 12(1): 191 (1878); R. Keller in Bull. Herb. Boissier II, 8: 179 (1908); Briquet in Annu. Conserv. Jard. hot. Geneve 20: 388 (1919); Malme in Ark. Bot. 23 (4): 16 (1930); Lyman B. Smith in J. Wash. Acad. Sci. 48: 311 (1958); Angely, Fl. Anal. Parana: 450 (1965); Rod- riguez Jimenez in C.r. Soc. Biogeogr. 432: 91, map 7 (1972), in Mems Soc. Cienc. nat. La Salle 33: 126, ff. Ic, 15a (1973), in Reitz, Fl. III. Catar., Hipericac.: 21, f. 4A (1980). Type: Brazil, 'in Brasiliae meridionalis provinciis', Sellow 1356 (Bt-holotype, (F!, GH!-photographs); BR (Pi- photograph), G!, S!). Map 10. Icon: Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericac.: 18, t. 4A (1980). Subshrub 0-4-1 m tall, erect, 1-stemmed, with branche^ spreading or ascending, lateral only. Stems dull red-brown, 4- lined and ancipitous when young, soon terete, cortex exfoliating irregularly; internodes 7-25 mm long, exceeding or shorter than leaves. Leaves perfoliate, spreading; lamina 10-22 x 7-10 mm, ovate-deltoid to oblong, margin recurved to revolute, paler beneath, not glaucous, chartaceous; apex acute to apiculate or obtuse, bases % to completely connate; basal vein 1, midrib pinnately branched with 4-5 main laterals, tertiary reticulum dense; laminar glands rather sparse, prominent beneath or obscure. Inflorescence 3-12- flowered, dichasial/monochasial from 1-2 nodes, sometimes with 1-3 pairs of lateral branches, the whole ± broadly pyramidal; primary pedicels 2-3 mm long; bracts and bract- eoles triangular-subulate. Flowers 9-15(-20) mm in diam., stellate; buds ovoid, acute to subacute. Sepals 3-4 x 1-1-5 mm, subequal, imbricate, narrowly oblong to elliptic or oblanceolate, acute to obtuse; veins 5-9, midrib or all slightly prominent; glands linear, sometimes distally punctiform. Petals orange-yellow to orange, 9-11 x 3-5^4 mm, c. 3 x sepals, obovate-oblanceolate; apiculus subacute; glands linear. Stamens 30-50(-60), irregularly grouped, longest c. 3 mm long, c. 0-3 x petals. Ovary 2 x 1-3-1-5 mm, ellipsoid; styles 5, 3-3-5 mm long, 1-5-1-75 x ovary, spreading; stigmas capitate-clavate. Capsule 4-5 x 3-5^-5 mm, globose, rounded to shortly rostrate, slightly shortly than sepals. Seeds 0-5 mm long; testa ribbed-scalariform. Forest clearings, pastures, roadsides, and disturbed habitats; 0-700 m. Brazil (Santa Catarina, Rio Grande do Sul), Argentina (Misiones). BRAZIL. Santa Catarina: mun. Campos Novos, 22 km W. of Campos Novos, c. 700 m, 9-10 February 1957 (fr), Smith & Klein 11174 (K, NY); mun. Concordia, Urugai, Vila Rica by Rio do Peixe, 350-400 m, 24 October 1964 (fl), Smith & Reitz 12923 (NY, P). Rio Grande do Sul: near Canoas, 1-5 km N. of Porto Alegre, 29 December 1966 (fl), Lindeman & de Haas 3898 (K, NY, U); mun. Gravatai, Fonte das Ninfas on road to S. Francisco de Paula, 6 December 1979 (fl & fr), Pedersen 12621 (BM). ARGENTINA. Misiones: Dep. San Javier, Matto Quemado, 26 September 1946 (fl), Bertoni 2964 (F, S). The slenderer stems, strongly connate leaves, and smaller sepals distinguish H. caprifoliatum from its closest relative, H.teretiusculum, which has a more northern distribution. For differences between H. caprifoliatum and H. connatum see below (p. 59). 5. Hypericum connatum Lam., Encycl. 4: 168 (1797); Choisy, Prodr. monogr. Hyperic.: 48 (1821), in DC., Prodr. 1: 548 (1824); A. St. Hil., PI. usuel. bras. 1: t. 61 (1828), Fl. Bras, mend. 1: 329 (1828); Walp., Repert. hot. syst. 1: 389 (1842); D. Dietr., Syn. pi. 4: 1236 (1847); Reichardt in Martius, FL bras. 12(1): 190 (1878); Arechav. in An. Mus. Hist. nat. Montevideo 3: 109 (1890); Chodat & Hassler in Bull. Herb. Boissier II, 3: 1126 (1903); R. Keller in Bull. Herb. Boissier. II, 8: 179 (1908); Briquet in Annu. Conserv. Jard. hot. Geneve 20: 389 (1919); Malme in Ark. Bot. 23A (4): 16(1930); Foster in Contr. Gray Herb. Harv. 184: 131 (1958); Lyman B. Smith in/. Wash. Acad. Sci. 48: 311 (1958); Cabrera, Fl. Prov. Buenos Aires: 225, fig. 65E- H (1965); Angely, Fl. Anal. S. Paulo 1: 186 (1969); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 91, map 7 (1972), in Mems Soc. Cienc. nat. La Salle 33: 128, ff. li, 2b,c, 3 h, 15 D (1973), in Reitz, Fl. III. Catar., Hipericac.: 17, f. 4D (1980). Type: Uruguay, le morne de Monte- Video, May 1767 (fl & fr), Commerson s.n. (P!-holotype, Fl-photograph; G!). Map 13. H. chlorifolium A. St. Hil., Fl. Bras, merid. 1: 329 (1828) ['chhraefolium']; D. Dietr. Syn.pl. 4: 1236 (1847); Walp., Repert. hot. syst. 1: 389 (1842); Angely, Fl. Anal. Parana: 450 (1965). Type: Brazil, Sao Paulo, Campos Geraes, Fazenda da Fortaleza, February [1820] 1816-21 (fl & fr), St. Hilaire 1476 bis (P!-holotype). Brathys connata (Lam.) Spach, Hist. nat. veg. 5: 450 (1836), in Annls Sci. nat. (Bot.) II, 5: 366 (1836). 58 N. K. B. ROBSON H. connatum var. chlorifolium (A. St. Hil.) Reichardt in Martius, Fl. bras. 12 (1): 192 (1878) ['chloraefolium']; Briquet in Annu. Conserv. Jard. hot. Geneve 20: 390 (1919). H. connatum var. obscurum Briquet in Annu. Conserv. Jard. hot. Geneve 20: 389 (1919). Type as for H. connatum Lam. Briquet cites H. connatum Lam. as a synonym of his variety; therefore the selection of a Sellow specimen as lectotype by Rodriguez Jimenez cannot stand. H. connatum var. paraguariense Briquet in Annu. Conserv. Jard. hot. Geneve 20: 390 (1919); Foster in Contr. Gray Herb. Harv. 184: 131 (1958). Type: Bolivia, Tarija, Bermejo, 1400 m, 3 December 1903 (fl & fr), Fiebrig 2361 pro parte (GMectotype; NY, S!, U!, US!, W!). The selection of this specimen as lectotype by Rodriguez Jimenez (1973: 130) is not appropriate because: (i) Briquet chose the epithet paraguariense, and the other syntypes are all from Paraguay; (ii) Briquet described another part of Fiebrig 2361 as the type of his var. fiebrigii. I therefore propose the following specimen as substitute lectotype: Paraguay, Caaguazu, in campis in regione flumine Yhu, November 1905 (fl), Hassler 9686 (GMectotype; BM!, K!, P!, W!). H. connatum var. fiebrigii Briquet in Annu. Conserv. Jard. hot. Geneve 20: 176 (1919); Foster in Contr. Gray Herb Harv. 184: 131 (1958). Type: Bolivia: Tarija, Bermejo, 1400 m, 3 December 1903 (fl), Fiebrig 2361 pro parte (G!- holotype, photograph; NY!, S!, U!, US!, W!). These herbarium symbols indicate where I have seen Fiebrig 2361. I have not noted which specimens are referable to var. paraguariense and which to var. fiebrigii. H. cyathifolium Larranaga in Publ. Inst. Geog. Urug. 2: 239 (1923); Lyman B. Smith in J. Wash. Acad. Sci. 48: 314 (1958). Type: Uruguay; probably no specimen ever ex- isted. Sarothra connata (Lam.) Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 71(1951). Icones: Reichardt in Martius, Fl. bras. 12 (1): t. II (1878): Cabrera, Fl. Prov. Buenos Aires: f. 65E-H (1965). Subshrub or perennial herb with woody base, 0-23-1 m tall, erect, 1-stemmed or branched from base with branches erect, lateral and rarely pseudo-dichotomous. Stems dull red- brown, 4(6)-lined and ancipitous when young, soon terete, cortex exfoliating irregularly; internodes 10-50 mm long, usually exceeding leaves. Leaves perfoliate, spreading or cyathiform; lamina 10-24(-30) x 12-34(^0) mm, ovate- deltoid to semicircular, margin revolute, incrassate, paler beneath, densely glaucous beneath or on both sides, coriaceous; apex obtuse-apiculate to rounded, bases com- Map 13 Sect. 30: 5. H. connatum • (confirmed records), O (additional records from Rodriguez Jimenez, 1973). HYPER1CUM 59 pletely connate or almost so; basal veins 1(3), midrib pinnately branched with 3-4 main laterals, tertiary reticulum dense; laminar glands dense, slightly prominent beneath. Inflorescence c. 10-°° -flowered, rarely pseudo-dichotomous after first flower, otherwise dichasial/monochasial from upper node, very rarely with 1-2 lateral branches from the nodes below, the whole corymbiform; primary pedicels 1-5-5 mm long; bracts and bracteoles linear-lanceolate to subulate. Flowers 10-24 mm in diam., stellate; buds ovoid, acute. Sepals 6-8 x 2-5-3-5 mm, unequal, imbricate, ovate or rhombic to elliptic or spathulate, acuminate to acute; veins 5- 7, midrib or all slightly prominent; glands linear, punctiform in distal %. Petals yellow to orange, 5-13 x 3-4 mm, 1-5-3 x sepals, oblong-oblanceolate to obovate; apiculus obsolete; glands linear, distally interrupted. Stamens 50-60 (-c. 100), obscurely 5-fascicled, shortly monadelphous, longest 3-5 mm long, 0-3-0-4 x petals. Ovary 2-2-5 x 1-5-2 mm, ovoid; styles 5(4), 2-5-3 mm long, c. 1-2 x ovary, spreading; stigma clavate to subcapitate. Capsule 5-6 x 4-5 mm, subglobose, shortly rostrate, almost equalling sepals. Seeds 0-7-1 mm long; testa ribbed-scalariform. Pastures and dry, rocky places; 20-2021 m (Bolivia). Brazil (Sao Paulo to Rio Grande do Sul), Uruguay, Argen- tina (Salta and Chaco south to San Luis and Buenos Aires), Paraguay, Bolivia (Chuquisaca, Tarija). BRAZIL. Sao Paulo: Itapetinga, 24 January 1950 (fl), de Lima RJ 69463 (MO, P). Parana: mun. S. Jose dos Pinhaes, Col. Murici, 30 November 1978 (fl & fr), Hatschbach 41803 (BM, MBM); mun. Jaguariaiva, 730 m, 7 June 1914 (fl & fr), Dusen 15122 (GH, S). Santa Catarina: mun. Lajes, Indies, ± 950 m, 16 December 1967 (fl), Lourteig 2255 (P); mun. Campo Ere, Chapeco, Capetinga, 24 January 1952 (fl), Reitz 4620 (NY); mun. Bom Jardin, Fachinal, 1200 m, 29 January 1950 (fl), Reitz 3409 (NY). Rio Grande do Sul: S. of Vacaria, over 200 km N. of Porto Alegre, 28 December 1966 (fl), Lindeman & de Haas 3953 (K, NY, U); Capao do Leao, prope Pelotas, 3 November 1901 (fl), Malme It. Reg. II 185 (S); Cacapara do Sul, Morro Peraf, 21 February 1948 (fl & fr), Palacios-Cuezzo 1451 (MO). URUGUAY. Artigas: vicinity of Artigas, 5 December 1943 (fl), Bartlett 21063 (GH, MICH). Canelones: Puerto Jackson, 20 m, 3 February 1931 (fr), Herter 86619 (Z). Colonia: Salto S. Luis, 120 m, 24 May 1947 (fr & fr), Herter 99655 (F, MO, S, Z). Maldonado: Cerro Pan de Azucar, 100 m, 11 November 1970 (fl), Lourteig 2480 (P). Rivera: Cunapiru, 180-210 m, 1928 (fl), Wright s.n. (BM). Rocha: 25 km N. Castillos, Palmares de Castillos, 22 January 1944 (fr), Bartlett 21371 (GH, MICH). ARGENTINA. Salta: Olemaina, 1500 m, 5 December 1929 (fl), Venturi 9851 (GH, K). Chaco: Dep. Bermejo, Puerto Bermejo, 2 March 1901 (fl), Kermes 6353 (P). Tucuman: Famailla, San Javier, Villar Nongues, 1200 m, 18 November 1946, Sparre 705 (S). Cordoba: Dep. Santa Maria, Alta Gracia, Puesto 'El Cura', 1800 m, 27 January 1944 (fr), Pierotti (BM, GH, U). San Luis: Sierra de Los Comeching- ones, 'El Rincon', 4 km E. of Merlo, 1000-2000 m, 29 January 1974 (fl), Conrad 2502 (MO). Buenos Aires: near Balcarce, Sierra Bachicha, post El Cruce, W. slope, c. 200 m, 2 April 1966 (fr), Hawkes, Hjerting & Rahn 4036 (BM). Entre Rios: Dep. Concordia, Salto Grande, 12 November 1968 (fl), Cabrera & Sagastegui 19296 (C, P). Corrientes: San Roque, Estancia 'Caaguazu', 1 March 1961 (fl & fr), Pedersen 5854 (C, GH). Misiones: Dep. San Ignacio, San Ignacio (Campo Rosa), 220 m, 16 November 1944 (fl & fr), Monies 465 (BM, F, GH, NY, S). PARAGUAY. Guaira: Cordillera de Villa-Rica [Villar- rica], January 1905 (fr), Hassler 8758 (BM, G, GH, P, W), Caaguazu: Caaguazu, 10 December 1874 (fl & fr), Balansa 2264 (P). Itapua: Itapua, ruderalis Ecclesiae imperfectae pagi Jesus, 12 November 1978 (fl & fr), Bernardi 18542 (G). Misiones: Santiago, Estancia 'La Soledad', 16 November 1956 (fl & fr), Pedersen 4317 (C, G, GH). Paraguari: Paraguarf, Tibicuary Mi, Isla Alta, 17 November 1978 (fr), Bernardi 18762 (BM, G). Alto Parana: Hernandarias, 20 km N. de Hernandarias, 10 January 1974 (fl), Schinini 8036 (G). BOLIVIA. Chuquisaca: Rosal, 2021 m, 3 October 1949 (fl), Brooke 5755 (BM). Tarija: Bermejo, 3 December 1903 (fl & fr), Fiebrig 2361 (NY, S, U, US, W). H. connatum resembles H. caprifoliatum in having the leaf pairs connate, but its leaves are coriaceous, not chartaceous, with a markedly incrassate margin (see Briquet (1919)) and nerves prominent beneath. The leaves are also relatively shorter and broader than in H. caprifoliatum, and the habit is less branched, often indeed with stems simple from the base. The plants nearest in form to H. caprifoliatum, with smaller flowers and leaves and branched stems, are from northern Argentina and Paraguay, whence there is a gradual trend towards basal branching with simple (annual?) stems, larger leaves, and larger flowers. The extreme in leaf size is apparently represented by the population from Sao Paulo that St.-Hilaire named H. chlorifolium; but the larger-leaved, larger-flowered form occurs throughout the range of the species and, in north-eastern Argentina and Paraguay, co- exists with the smaller-flowered form. It was no doubt this overlapping variation that prompted Briquet to describe his varieties; but there is no apparent gap in variation such as would allow the recognition of infraspecific taxa. 6. Hypericum cordatum (Veil. Cone.) N. Robson, comb. nov. Receveura cordata Veil. Cone., Fl. flumin.: 237 (1825), 5: t.119 (1835). Type: Brazil, Vandelli (LISC?-holotype). H. cordiforme A. St. Hil., Fl. Bras, merid. 1: 330 (1828); Walp., Repert. hot. syst. 1: 389 (1842); D.Dietr., Syn. pi. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 12 (1): 190, t.33 f.2 (1878); R. Keller in Bull. Herb. Boissier 6: 259 (1898), op. cit. II, 8 179 (1908); Briquet in Annu. Conserv. Jard. hot. Geneve 20: 390 (1919); Lyman B. Smith in /. Wash. Acad. Sci. 48: 314 (1958); Angely, Fl. Anal. Parana: 450 (1965), Fl. Anal. S. Paulo 1: 186 (1969); Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc: 13, f. 3A (1980), pro parte, excl. syn. var. P A. St. Hil. et var. Glazioui. Type: Brazil, Sao Paulo, in pascuis prope urbem Sancti Pauli, [1819] 1816-1821 (fl), St.-Hilaire 1172 (P!- holotype & isotype). H. cordiforme var. genuinum Briquet in Annu. Conserv. Jard. hot. Geneve 20: 391 (1919). Type: Brazil, Sao Paulo, prope Sao Paulo, February 1839 (fl & fr), Guillemin 315 (G!-holotype; P!). Subshrub (? or perennial herb with woody base), 0-2-0-6(-l) m tall, erect or decumbent, branched from the base and sometimes above, with branches erect, pseudo-dichotomous and/or lateral. Stems dull red-brown, persistently 4-lined, ancipitous when young, cortex exfoliating in strips; inter- nodes 5-15 (-20) mm long, exceeding or rarely equalling leaves. Leaves sessile, isomorphic, erect to appressed; lamina 9-18 x 4-11 mm, broadly ovate to oblong-ovate or elliptic, 60 N. K. B. ROBSON not or only slightly narrower towards base of stem, margin recurved and ± incrassate or rarely plane (when narrow), concolorous or rarely paler beneath, densely glaucous, coriaceous; apex acute or apiculate to obtuse, base cordate or rarely subcordate, free or up to !/3 connate; basal or near- basal veins 1-7, midrib obscurely branched, tertiary reticulum not visible; laminar glands dense, slightly prominent beneath. Inflorescence (3-)7-21(-35)-flowered, dichasial, sometimes branching pseudo-dichotomously below the inflorescence or rarely branching mixed dichasial/pseudo-dichotomous, rarely with 1-2 branches from the node below and very rarely from farther down stem, the whole corymbiform; primary pedicels 2-3 mm long; bracts and bracteoles lanceolate. Flowers (10-) 15-25 mm in diam., stellate; buds ovoid, subacute. Sepals 5-7 x 2-3-3 mm, unequal, imbricate, ovate or ovate-lanceolate to rhombic or rarely lanceolate, acute; veins c. 7, 3 central prominent; glands linear, punctiform in distal !/4 or less. Petals yellow to orange, 8-12 x 4-7 mm, 1-6-2 x sepals, obovate-oblong to obovate; apiculus acute; glands linear. Stamens 35-60, obscurely 3(4)-fascicled, longest 4-5 mm long, c. 0-5 x petals. Ovary 1-5-2 x 1-1-5 mm, ovoid- subglobose; styles 3(4), 3-4 mm long, 1-8-2 x ovary, spreading; stigmas subcapitate. Capsule 4-6(-7) x 3-5 mm, ovoid-subglobose, scarcely rostrate, almost equalling sepals. Seeds 0-8-0-9 mm long; testa ribbed-scalariform. Thickets and dry pastures; 800-1200 m. Brazil (Minas Geraes and Sao Paulo to Rio Grande do Sul). H. cordatum has coriaceous leaves like those of H. connatum but smaller, and they also differ in being free or only slightly connate at the base. The flowers are also usually smaller and have 3(4) styles not 5(4). H. cordatum has hitherto been known as H. cordiforme A. St. Hil., but Vellozo's illustration of Receveura cordata is clearly of the same plant. His epithet therefore has priority. H. cordatum is fairly uniform, with broadly ovate, cordate leaves throughout its range; the plants with the largest leaves and flowers occur in Sao Paulo. In Santa Catarina, however, there is a trend towards narrower leaves (oblong-ovate to elliptic-oblong), more numerous lateral inflorescence bran- ches (-5), and smaller flowers (10-15 mm in diam.). Such plants (e.g. Smith & Klein 8220) at first sight look more like H. ternum, but they have the longer internodes and (usually) 1-noded inflorescence of H. cordatum. These plants, in turn, merge with even smaller-leaved, smaller-flowered individuals (e.g. Reitz & Klein 5223), in which the leaves are as small as those of H. ternum but broader, and they also resemble H. ternum in having numerous lateral branches. The leaves, however, are never narrow or overlapping. Despite some intermediates in Santa Catarina and Parana, it seems most appropriate to place these anomalous plants in a distinct subspecies. 6a. Hypericum cordatum subsp. cordatum. Fig. 13A, Map 14. Leaves broadly ovate or triangular-ovate to ovate-oblong (l:b=l-2:l), margin slightly incurved-incrassate. Inflor- escence from 1(2) nodes, very rarely with lateral branches from further down stem. Flowers (10)15-25 mm in diam. Sepals ± broadly ovate. Distribution of species but rather rare in Santa Catarina. BRAZIL. Minas Geraes: Serra do Picii, 11 December 1886 (fl), Schrenck 1560 (C). Sao Paulo: prope Mugy [Moji], November 1833 (fl), Riedl 1610 (K, P, W); Butantan, 27 August 1917 (fl), Hoehne 2 (F, NY); Itapetininga, 1 March 1951 (fl), de Lima in RJ 78066 (P). Parana: Lagoa Dourada, 13 January 1964 (fl & fr), Pereira 8258 (= Pabst 7533) (NY, P); Ponta Grossa, Passo do Pupa, 10 October 1967 (fl), Hatschbach 17386 (MO). Santa Catarina: mun. Lajes, Indies, c. 950 m, 16 December 1967 (fl), Lourteig 2241 (P). Rio Grande do Sul: prope S. Francisco de Paula, Taimbe, 18 December 1950 (fl & fr), Rambo 49408 (S). 6b. Hypericum cordatum subsp. kleinii N. Robson, subsp. nov. Fig. 13B, Map 14. H. denudatum sensu Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc.: 16 (1980), pro parte, excl. t. 3B, non St. -Hil. (1828). A subsp. cordatum foliis sepalisque angustioribus, floribus minoribus, ramis lateralibus, inflorescentiae interdum e nodis 2-5 ortis, differt. Type: Brazil, Santa Catarina, mun. Lajes, between Palmeiras and Lajes, 800-900 m, 2 December 1956 (fl), Smith & Klein 8100 (US!-holotype; NY!-isotype). Leaves triangular-lanceolate or narrowly oblong to narrowly elliptic (l:b = 3^:1), margin plane or almost so. Inflor- escence from 1-5 nodes, sometimes with lateral branches from lower nodes. Flowers 10-13 mm in diam. Sepals narrowly ovate to lanceolate. Brazil (Santa Catarina, Parana). BRAZIL. Parana: mun. Palmas, 10 km NW. de Palmas, 4 HYPER1CUM 61 Fig. 13 A. H. cordatum subsp. cordatum: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) stamens (partly cut away) and ovary; (f) capsule. B. H. cordatum subsp. kleinii: (g) habit; (h) leaf. C. H. cavernicola: (i) habit (a, g, i x '/2; b, h x 1; c-f x 4). A. Glaziou 10396; B. Hatschbach et al. 28161 ;C. Berro4862. 62 N. K. B. ROBSON Map 14 Sect. 30: 6a. H. cor datum subsp. cor datum cor datum subsp. kleinii O; 7. H. cavernicola I 6b. H. December 1971 (fl), Hatschbach [Smith & Klein] 28161 (BM, MBM). Santa Catarina: mun. Lajes, 15 km N. of Lajes, 900 m, 4 December 1956 (fl), Smith & Klein 8220 (US); Agua Dolce, 3 km NW. of Herciopolis, Campos de Palmas, 1100- 1200 m, 5 December 1964 (fl), Smith & Klein 13628 (US); mun. Campo Alegre, Morro Iquererim, 1300-1500 m, 8 November 1956 (fl), Smith & Klein 7407 (US). 7. Hypericum cavernicola Lyman B. Smith in Wrightia 2: 90 f. 19a-c (1960). Type: Uruguay, Tacuarembo, Gruta de los Cuevos, 17 December 1907 (fl), Berro 4862 (US!-holotype; K!,MVFA). Fig. 13C, Map 14. Subshrub, c. 0-2 m long, erect or ascending, branched from the base and above, with branches erect, lateral. Stems dull red- brown, 4-lined, and ancipitous when young, soon 2-lined, cortex exfoliating in strips; internodes 2-5 mm long, shorter than leaves. Leaves sessile, suberect to spreading; lamina 4-10 x 2-\ mm, ovate-triangular to oblong or (on lateral branches) lanceolate to linear-lanceolate, margin subrecurved, concol- orous, coriaceous; apex acute to subacute, base cordate to truncate, free; basal veins 1(3), midrib not or obscurely branched, tertiary reticulum not visible; laminar glands dense, slightly prominent beneath. Inflorescence l(-ll)-flowered, dichasial/monochasial from 1-3 nodes, with lateral branches from nodes almost immediately below, the whole subcylindric; pedicels almost absent; bracts foliar or transitional; bracteoles appressed to calyx. Flowers c. 10 mm in diam., stellate; buds ovoid, subacute. Sepals 7-8 x 2 or 3-5^ mm, very unequal, imbricate, 2 outer ovate to elliptic-oblong, distally recurved, 3 inner narrowly oblong-lanceolate, all acute to subacuminate; veins 9-11, 5 central or all prominent; glands linear, punctiform in distal c. !/2. Petals yellow, 6-6-5 x 3-5^- mm, shorter than sepals, narrowly obovate; apiculus acute; glands mostly punctiform. Stamens 25-30, obscurely 5-fasciculate, longest c. 4 mm long, c. 0-6 x petals. Ovary 2x1-5 mm, ovoid- subglobose; styles 3, 2-5 mm long, 1-2 x ovary, spreading; stigmas subcapitate. Capsule and seeds not seen. Stony ground; lowland. Uruguay (Tacuarembo). URUGUAY. Tacuarembo: Gruta de los Cuevos, 17 December 1907 (fl), Berro 4862 (K, US). H. cavernicola is known to me only from the type. It is clearly closely related to and derived from H. cordatum subsp. kleinii, but the sessile flowers with unequal sepals, the outer pair much larger, are diagnostic. The large outer pair of sepals are reminiscent of those of the 'Ascyruni' species of sect. Myriandra. 8. Hypericum termini A. St. Hil., Fl. Bras, merid. 1: 330 (1828); Walp., Repert. hot. syst. 1: 389 (1842); D. Dietr., Synpl. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 12 (1): 194 (1878); Lyman B. Smith in/. Wash. Acad. Sci. 48: 312 (1958); Angely, Fl. Anal. Parana: 451 (1965); Fl. Anal. S. Paulo 1: 186 (1969). Type: Brazil, Parana, Distr. Curitiba, 'prope pagulum vulgo Fregesia nova', March [1820] 1815- 1821, St.-Hilaire 1585 (P!-holotype; US!-photograph). Map 15. H. cordiforme [var.] (3 sensu A. St. Hil., Fl. Bras, merid.: 330 (1828); Reichardt in Martius, Fl. bras. 12 (1): 190 (1878). Type: Brazil, Minas Geraes, in pascis mentis vulgo Papagayo, [1822] 1815-1821 (fl), St.-Hilaire D.541 (P!- holotype). H. rigidum sensu Reichardt in Martius, Fl. bras. 12(1): 189 (1878), pro parte, excl. typum et syn. H. sellowianum Klotzsch; Lyman B. Smith in /. Wash. Acad. Sci. 48: 314 (1958). H. cordiforme var. hilairei Briquet in Annu. Conserv. Jard. hot. Geneve 20: 391 (1919). Type as for H. cordiforme var. (3 sensu A. St. Hil. H. cordiforme var. glazioui Briquet in Annu. Conserv. Jard. hot. Geneve 20: 391 (1919). Type: Brazil, Minas Geraes, Serra do Caponema, Caraca, 1883-1884 (fl), Glaziou 14534 (G!-holotype; C!, K!, P!-isotypes). H. denudatum sensu Rodriguez Jimenez in Reitz, Fl. 111. Catar., Hipericdc.: 16 (1980), pro parte, quoad t.3B. HYPER/CUM 63 Icon: Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc.: 14,t. 38(1980). Subshrub, 0-2-0-7 m tall, erect, usually unbranched from the base, ± richly branched above, with branches strict, lateral, rarely in whorls of 3. Stems green to red-brown, persistently 4- lined, ancipitous when young, cortex exfoliating in strips; internodes 5-18 mm long, the upper exceeding leaves. Leaves sessile, ± heteromorphic, erect to appressed, eventually sometimes spreading, paired or very rarely 3-whorled on main stem; lamina 5-14 x l-6(-8) mm, ovate or ovate-triangular to lanceolate or linear, margin recurved or rarely to plane, subconcolorous, coriaceous; apex acute to subacute, base cordate to parallel-sided or subcuneate, free; basal veins 1(3), midrib branched with 1-2 laterals or unbranched, tertiary reticulum not visible; laminar glands dense to rather sparse, slightly prominent beneath. Inflorescence (l-)3-ll -flowered, dichasial (2-3 nodes) then monochasial, from 1(2) nodes, with short lateral branches from up to 10 nodes immediately below, the whole narrowly pyramidal to cylindric; pedicels 1-5-2 mm long; bracts and bracteoles ovate to linear. Flowers (6-)8-12 mm in diam., stellate; buds ovoid, subacute to acute. Sepals 3- 6 x 1-2-5 mm, unequal, imbricate, outer ovate to lanceolate, inner lanceolate to narrowly oblong, acute to subacuminate; veins 3-7, 3 or more usually becoming prominent; glands linear, punctiform in distal 1A - l/2. Petals yellow to orange, (5-)6-8 x (2-5-)3~4 mm, 1-6-2 x sepals, oblong-obovate; apiculus acute; glands linear to punctiform. Stamens 30-63, not or obscurely 3(4)-fascicled, longest 3^ mm long, c. 0-5 x petals. Ovary c. 1-5 x 1 mm, ovoid-subglobose; styles 3(4), 3-4 mm long, 2-2-3 x ovary, spreading; stigmas subcapitate to capitate. Capsule 5 x 2-3 mm, ovoid-subglobose, not rostrate, shorter than sepals. Seeds 0-8-0-9 mm long; testa very finely ribbed-scalariform (markings obscure). Dry, often stony pastures and slopes; 800-1600 m. Brazil (Minas Geraes, Sao Paulo, Parana, northern Santa Catarina, also? Rio Grande do Sul. BRAZIL. Minas Geraes: Rodrigo Silva pres d'Ouro Preto, 9 May 1892 (fl & fr), Glaziou 18913 (C, K, P); Hermello Alves, 1100 m, 24 December 1949 (fl), Duarte 2335 (NY, P). Sao Paulo: Serra da Bocaina, sud vom Itatiaya, 1600 m, 7 December 1952 (fl), Markgraf 10357 (Z); Mogy dos Cruces, 19 April 1889 (veg), Glaziou 17476 (C, K, G, P). Parana: mun. Curitiba, Rio Atuba, 23 November 1970 (fl), Hatsch- bach 25605 (C, MO, NY); Turma no. 23, c. 800 m, 19 October 1914 (fl), Dusen 15724 (BM, GH, MO, S); Capivari, 21 October 1908 (fl), Dusen 54/78 (A, F, G, GH, MO, NY, P, S, Z). Santa Catarina: campos do sul do Rio das Morocubas, n.d.(fl), F. Midler s.n. (P); (?) mun. Campo Alegre, Morro de Iquererim, 1400 m, 18 October 1957 (fl), Reitz & Klein 5223 p.p. (US). Rio Grande do Sul: Sao Francisco de Paula, Taimbezinho, 3 December 1971 (fl & fr), J.C., F.M.L., A.M.G. & M.L.P. ICN 9316 (U). H. ternum can be distinguished from H. cordatum by the smaller flowers and the heteromorphic or uniformly narrow or small leaves. The most primitive forms (in Minas Geraes and Sao Paulo) have ovate or ovate-triangular leaves near the inflorescence, but the lowermost leaves on the shoot are linear-lanceolate to linear. Forms in which the leaves are long, narrow, and with recurved margins occur in all the northern states in its range, as well as in northern Santa Catarina. In Parana, however, there is a trend towards smaller isomorphic leaves which become too small to allow the recurved-margin character to be expressed. In such small- leaved, small-flowered forms, the main stem occasionally bears leaves in whorls of 3, and the type specimen is one of these. The extreme form of the type specimen has resulted in repeated misunderstanding of H. ternum, specimens belong- ing to it having been included by various authors in H. rigidum, H denudatum, and H. sellowianum. As will be seen elsewhere in this work, these names apply to quite different taxa. The inappropriateness of the epithet ternum to all but a small part of the species is no reason (according to the International Code of Botanical Nomenclature) for not using it. Map 15 Sect. 30: 8. H. ternum •; 9. H. legrandii O. 64 N. K. B. ROBSON 9. Hypericum legrandii Lyman B. Smith in /. Wash. Acad. Sci. 48: 314 (1958). Type: Uruguay, Rivera, 17 December 1901 (fl), Berro 1720 (MVM-holotype; US!-isotype). Map 15. H. ericoides Arechav. in An. Mus. Hist. nat. Montevideo 4: 18 (1902), non. L. (1753). Type as for H. legrandii. \J Subshrub 0-45-0-8 m tall, erect, branched above only, with branches strict, lateral, (?) rarely in whorls of 3. Stems dull red-brown, persistently 4-6-lined, ancipitous or triquetrous at first, soon 2-3-lined, eventually terete, cortex exfoliating in strips; internodes 5-7(-10) mm long, shorter than leaves. Leaves sessile, imbricate-appressed, paired or usually 3- whorled; lamina 5-11 x 0-8-1 mm, acerose, margin plane, conduplicate or carinate proximally, concolorous, coriaceous; apex acute, base parallel-sided, free; vein 1, unbranched; laminar glands dense, slightly prominent beneath. Inflor- escence c. 14-20-flowered, trichasial on main stem, dichasial on laterals (2 nodes) then monochasial, from 2 nodes, with short lateral branches from up to 12 nodes immediately below, the whole narrowly cylindric; pedicels 1-2 mm long; bracts and bracteoles linear-subulate. Flowers c. 8 mm in diam., stellate; buds narrowly ovoid, acute. Sepals (4-)5-6 x 1 mm, subequal, subimbricate, linear-lanceolate to linear, acute; veins 5, not prominent; glands linear to mostly punctiform. Petals yellow to orange, c. 5-7 x 3 mm, 1-1-2 x sepals, oblong-obovate; apiculus acute; glands linear. Sta- mens c. 30, obscurely or not fascicled, longest c. 4 mm long, c. 0-6 x petals. Ovary 1-5-2 x 1-1-5 m, elliptic-subglobose; styles 3, 3-4 mm long. 2 x ovary, spreading; stigmas scarcely capitate. Capsule 5-6 x 2-5-3 mm, cylindric-ovoid, about equalling sepals. Seeds not seen. Habitat unknown. Uruguay (Rivera). URUGUAY. Rivera: Rivera, n.d. (fl), Felippone 5070 (K); Chacra Santurio, 9 December 1907 (fl), Berro 4865 (US). H. legrandii shares the tendency shown by reduced forms of H. ternum to have leaves in 3's, but its leaves and sepals are narrower. 10. Hypericum denticulatum Walter, Fl. carol.: 190 (1788); Gmelin, Syst. nat. 2: 1159 (1791); Fern, in Rhodora 44: 43 (1942); Fern. & B. G. Schubert in Rhodora 50: 205 (1948); P. Adams in Rhodora 64: 241 (1962); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 87, map 1 (1972); in Mems Soc. Cienc. nat. La Salle 33: 99, ff. If, 2d (1973); D. H. Webb, Biosyst. Study Hypericum sect. Spachium in eastern N. Am.: 272 (1980). Types: U.S.A., Carolina, Walter (holo- type, specimen not traced); U.S.A., Carolina, 1798-1800, Bosc (P-neotype; Fl, G!, US 8-isoneotypes) (Rodriguez Jimenez, 1973). Perennial herb 0-2-0-75 m tall, erect, branched mostly from the base, sometimes also from upper part of stem. Stems green, 4-lined, densely gland-dotted; internodes 8^17 mm long, exceeding leaves. Leaves sessile, spreading from base to appressed; lamina 5-35(^0) x 3-16 mm, lanceolate or ovate to elliptic or oblong-elliptic or obovate, margin plane but sometimes recurving when dry, concolorous or almost so, not glaucous, subcoriaceous; apex acute to subrounded, base cuneate to rounded-subamplexicaul, sometimes subdecur- rent, free; basal veins 1-5, if 1 then with 2-3 pairs of main lateral branches, the midrib otherwise apparently unbran- ched, tertiary reticulation not visible; laminar glands dense, not or scarcely prominent. Inflorescence up to c. 25-flowered (terminal), monochasial after c. 4th grade, with up to c. 15- flowered lateral dichasial/monochasial branches from up to 6 nodes below, the whole broadly pyramidal to corymbiform, occasionally with lateral flowering branches below; pedicels 2-4 mm long; bracts 3-6 mm long, lanceolate to subulate. Flowers 5-13 mm in diam., stellate. Sepals 3-8 x 1-5-4 mm, ovate or lanceolate to elliptic or obovate, acute to acuminate; veins 3-5, rarely branched, all prominent; glands linear, distally punctiform. Petals yellow to orange-yellow, 5-10 x 4- 6 mm, c. 1-6-2 x sepals, obovate; apiculus obtuse; glands linear, interrupted distally. Stamens 50-80, irregularly grouped, longest 3-5 mm long, c. 0-5-0-6 x petals. Ovary 1-5-2 x 1-1-5 mm, ovoid; styles 3, 2-\ mm long, 1-3-2 x ovary, divergent-incurved; stigmas capitate. Capsule 3-5 x 2- 3 mm, ovoid to rostrate-subglobose, shorter than to shortly exceeding sepals. Seeds 0-45-0-9 mm long, testa obscurely linear-reticulate to finely ribbed-scalariform. 2n = 24, 48. Fields, roadsides, ditches, and open woods on sandy or clay soils; 0-500 m. South-eastern U.S.A. (New Jersey south to Georgia and northern Florida, west to southern Illinois, Tennessee, and Louisiana). H. denticulatum is most closely related to Ic. H. rigidum subsp. sellowianum from south-eastern Brazil, the wide disjunction in distribution being probably the result of ancient long-distance dispersal (see p. 11). The epithet 'denticula- tum' was given by Walter because he was obviously impressed by the apiculus on the petal, whereas H. denticulatum Kunth (= H. galinum S. F. Blake) was named on account of the denticulate leaf margin. The H. denticulatum complex has usually been treated as comprising two taxa, which have been regarded as species or, more recently, as varieties. H. denticulatum sensu stricto is almost confined to the eastern Coastal Plain and has shorter, broader, often more obtuse leaves and sometimes broader sepals than '//. virgatum\ which is absent from the Coastal Plain but occurs from the adjacent uplands westward to the Mississippi. Fernald & Schubert (1948) reviewed the earlier treatments of the group. They divided H. denticulatum into 8See footnote p. 66. 65 three varieties, var. typicum (= var. denticulatum), var. acutifolium, and var. recognitum, the last two comprising H. virgatum sensu lato and differing in leaf-shape. In var. acutifolium they included Georgian specimens described by Keller (1923) as H. harperi. Gleason (1952) and Gleason & Cronquist (1963) recog- nized only two varieties, as did Adams (1973), who speci- fically mentioned that var. recognitum was doubtfully distinct from var. acutifolium. Rodriguez Jimenez (1973) agreed that there are two varieties but used an illegitimate epithet, var. virgatum (Lam.) Rodriguez Jimenez, for the narrow-leaved one. Webb (1980), however, while still adopting the rank of variety for the subordinate taxa of H. denticulatum, showed that Keller's H. harperi was distinct from H. denticulatum var. acutifolium morphologically, ecologically, and chemi- cally, and that it has a distinct distribution. Webb used the name H. incertum Steudel (= Brathys lanceolata Spach, non Hypericum lanceolatum Lam.) for this taxon in anticipation of seeing the type. Subsequently, however (in litt., 1980), he found that Spach's type belonged to H. denticulatum var. acutifolium; and so the correct name for this taxon (which I agree with him in treating as a species) is H. harperi R. Keller. Webb also confirmed earlier reports (Rogers, 1965; Rob- son & Adams, 1968) of the chromosome numbers n = 12 and 24 for H. denticulatum (sensu lato), showing that n = 12 is found in both varieties as well as in H. harperi, whereas n = 24 is apparently confined to var. denticulatum. Lastly, the two taxa that Webb and others treat as varieties differ in leaf shape, in leaf length relative to the internodes, in sepal shape, and in seed size and ornamentation, and they have distinct chromatographic profiles and distributions (Webb, 1980). I therefore regard the rank of subspecies as more appropriate for them than variety. lOa. H. denticulatum subsp. denticulatum Map 16. H. laevigatum Aiton, Hortus Kew. 3: 106 (1789). Type: cultivated, England, Kew, introduced 1772, Samuel Martin (no specimen found). H. angulosum Michaux ex Willd., Sp. pi. 3: 1454 (1802); Michaux, Fl. bor.-amer. 2: 78(1803); Torrey & A. Gray, Fl. N. Amer.: 164 (1838). Type: U.S.A., Carolina, 'in humidis a Carolina ad Floridam', Michaux (B-WILLD- holotype, TENN-photograph; P; for photograph see Rho- dora SO: 202, t. 1109, f.l (1948)). Brathys linoides Spach, Hist. nat. veg. 5: 452 (1836), in Annls. Sci. nat. (Bot.) II, 5: 367 (1836), nom. illegit. (Art. 63). Type as for Hypericum angulosum Michaux ex Willd. H. virgatum sensu Coulter in Bot. Gaz. 11: 106(1886), pro parte excl. var. acutifolium. H. virgatum var. ovalifolium Britton in Trans. N.Y. Acad. Sci. 9: 10 (1889). Type: U.S.A., New Jersey, Spring Lake, 31 July 1887, Lighthipe s.n. (NY-lectotype, Webb (1980), ined.). Rugel 378, selected by Rodriguez Jimenez (1973), is not type material. Webb's selection is confirmed here. H. denticulatum var. ovalifolium (Britton) S. F. Blake in Rhodora 17: 135 (1915); Fern. & B. G. Schubert in Rhodora 50: 206 (1948). H. denticulatum var. typicum Fern. & B. G. Schubert in Rhodora 50: 207 (1948). Type as for H. denticulatum Walter. Map 16 Sect. 30: lOa. H. denticulatum subsp. denticulatum •; lOb. H. denticulatum subsp. acutifolium O; 11. H. harperi D. Data from Webb (1988). 66 N. K. B. ROBSON Icon: Gleason, New Br. & Br. III. Fl. 2: 542 (1952), as H. denticulatum var. ovalifolium. Stems 0-2-0-7 m tall; lower internodes usually exceeding leaves. Leaves 5-20(-24) x 5-15 mm, usually appressed, broadly or narrowly ovate to elliptic or narrowly obovate, acute to subrounded. Sepals 4-8 x 2-\ mm, ovate or lanceolate to elliptic or obovate (especially lower flowers), acute. Petals yellow-orange, 5-10 x 4-6 mm. Seeds (0-45) 0-5-0-65 mm. 2n = 24, 48 (n = 12, 24 - Webb, 1980). Moist ditches, pine barrens, and savannahs; 0-400 m. South-eastern U.S.A. (coastal plain from New Jersey south to Georgia, with disjunct populations in western North Carolina, central Tennessee, and southern Alabama). U.S.A. Alabama;9 *Mobile Co., Dauphin Island, near Hous- ton PI., 15 July 1965, Deramus 568 (NCU). Delaware; *Kent Co., Felton, August 1867, Canby s.n. (NY). *Sussex Co., E. of Ellendale, 5 September 1925, Pennell 12870 (MO). Georgia: *McIntosh Co., 2-1 km N. of Fort Berrington road, 8 July 1972, Bozeman 758 (NCU). New Jersey: Atlantic Co., Hammonton, 18 August 1923 (fl), Bassett (TAI); Burlington Co., c. 23 km S. of Chatsworth, 20 August 1948 (fl), Lawrence & Dress 557 (BM); Camden Co., Winslow, 30 September 1908, Pennell 1777 (NY). North Carolina: Bun- combe Co., near Biltmore, 21 July 1897 (fl), Herb. Biltmore 523b (BM, W); Hoke Co., c. 5.3 km SE. of Antirch, 0-5 km W. of Robeson Co. line, 10 October 1957 (fr), Ahles 36459 (H); *Northampton Co., near Margaretsville, 28 July 1893, Heller 1154 (GH, MO, NY). South Carolina: *Berkeley Co., c. 5 km S. of Bonneau, 13 July 1927, Wiegnand & Manning 2018 (GH); *Georgetown Co., 11 km E. of Andrews, 27 June 1939, Godfrey & Tryon 156 (MO, NY, TENN, US). Tennessee: Coffee Co., Tullaloma, 10 August 1899 (fl), Herb. Biltmore 523h (H, W); *Warren Co. , NE. of Morrison, 18 June 1948, Fairchild et al. 7565 (TENN). Virginia: Sussex Co., c. 6-5 km N. of Homeville, 19-20 August 1936 (fl), Fernald & Long 6280 (K), *6837 (GH, NY, US); York Co., NW. of Grafton, 18 September 1937 (fr), Fernald & Long 7531 (BM, GH, K, MO, NY, US). The disjunct populations of subsp. denticulatum in the more interior regions (North Carolina, Tennessee) were thought by Webb (1980) to be possibly relicts, whereas he regarded the Alabama population as probably a recent introduction. lOb. H. denticulatum subsp. acutifolium (Elliott) N. Robson, stat. nov. Map 16. H. virgatum Lam., Encycl. 4: 158 (1796); Coulter in Bot. Gaz. 11: 106 (1886). Type: U.S.A., without precise locality or collector (P-JUSS-holotype; GH*-photograph; see also Rhodora 5Q: 206, t.llll, f. 1 (1948). H. acutifolium Elliott, Sketch hot. S. Carolina 2: 26(1821); Walp., Repert. hot. syst. 1: 388 (1842). Type: U.S.A., Georgia, Milledgeville, Elliott (?) 19 (CHARL*-holotype; GH*-photograph). Brathys erythraeae Spach, Hist. nat. veg. 5: 452 (1836), in Annls. Sci. nat. (Bot.) II, 5: 367 (1836). Type: U.S.A., Carolina, Leconte s.n. (P*-holotype). Brathys lanceolata Spach, Hist. nat. veg. 5: 453 (1836), in yln this part (Part 8), an asterisk (*) before a locality or after a herbarium symbol indicates that the specimen has been recorded by D. H. Webb, C. Rodriguez Jimenez, or P. Bamps, but not by me. Annls. Sci. nat. (Bot.) II, 5: 367 (1836). Type: U.S.A., 'midi des Etats Unis', Leconte (P*-holotype). H. erythraeae (Spach) Steudel, Nomencl. hot. 2nd. ed. 1: 787 (1840). Type: as for Brathys erythraeae Spach. H. incertum Steudel, Nomencl. hot. 2nd ed. 1: 788 (1840). Type as for Brathys lanceolata Spach (1836), non Hyper- icum lanceolatum Lam. (1797). H. virgatum var. acutifolium (Elliott) Coulter in Bot. Gaz. 11:106(1886). H. denticulatum var. acutifolium (Elliott) S. F. Blake in Rhodora 17: 134 (1915); Fern. & B. G. Schubert in Rhodora 50: 208, t.llll, f.l. (1948); Fern., Gray's Man. Bot. 8th ed.: 1013 (1950), pro parte; Radford, H. E. Ahles & Bell, Man. vase. fl. Carolinas: 716 (1968); D. H. Webb, Biosyst. study Hypericum sect. Spachium in eastern N. Am.: 283 (1980). H. denticulatum var. recognitum Fern. & B. G. Schubert in Rhodora 50: 208, t. 1110 (1948). Type: U.S.A., Tennessee, Knoxville, July 1895, A. Ruth (GH!-holotype; ILL*, US- isotypes). H. denticulatum var. virgatum (Lam.) Rodriguez Jimenez [in C.r. Soc. Biogeogr. 432: 87, 88 (1972), sine basion.j in Mems Soc. Cienc. nat. La Sallett: 100 (1973), nom. illegit. superfl. Icon: Gleason, New Br. & Br. III. Fl. 2: 542 (1952), as H. denticulatum. Stems 0-4-0-75 m tall, usually branched above; lower internodes usually shorter than leaves. Leaves 10-35(^0) x 3-8(-12) mm, ascending to divergent, lanceolate to oblong- elliptic or obovate, acute to obtuse. Sepals 3-3-5 x 1-5-2-5 mm, narrowly ovate to lanceolate, acute to acuminate. Petals yellow to yellow-orange, 6-10 x 5-6 mm. Seeds (0-5)0-6- 0-8(0-9) mm long. 2n = 24 (n = 12 - Webb, 1980). Dry roadsides, fields, and open woods; 0-c. 700 m (or higher?). South-eastern U.S.A. (Maryland to southern Illinois and south to Florida and Louisiana; not east coastal). U.S.A. Alabama: without precise locality, 1832 (fl), Drummond s.n. (K, W); * De Kalb Co., near former Lawson Ford on Town Creek, Sand Mt., 14 August 1945, Harper 399 (US). Arkansas: Saline Co., Bauxite, 90 m, 25 September HYPER1CUM 1965 (fr), Demaree 35005 (BM). Florida: Franklin Co., Apalachicola prope Mount Vernon, August 1843 (fr), Rugel 671 (BM, GH, K, NY, SMU, US, W). Georgia: Thomasville, 30 September 1903 (fr), Mrs A. P. Taylor (H); *Catoosa Co., Chickamauga Creek, near Ringgold, 6-12 August 1895, Small s.n. (NY); Sumter Co., Leslie, 15 September 1900, Harper657 (BM, K, NY, US, W). Illinois: Maysville,-July-, Buckley Ib (K); *Pope Co., S. of Rock, 21 July 1948, Winterringer 4026 (ILL). Indiana: *Spencer Co., c. 5 km NW. of Chrisney, near Little Pigeon Creek, 4 September 1939, Deam 59465 (GH, NY). Kentucky: *Calloway Co., between Murray and New Concord, 20 July 1937, Smith & Hodgdon 4084 (GH, NY, US); *Trigg Co., Kentucky Lake, S. of Frenton, 26 July 1954, Ahles 8123 (ILL, NCU). Louisiana: Covington, August 1832 (fl), Drummond 260 (BM, K); *St. Helena par., c. 6-5 km NE. of Greensburg, 4 August 1971, Allen 1319 (GA, SMU). Maryland: *Prince George's Co., c. 1-5 km SE. of Laurel along Bear Branch. 12 August 1922, Wherry s.n. (US). Mississippi; *Chickasaw Co., c. 6 km SE. of Pontotoc- Chickasaw Co. line on Miss. 41, 23 June 1964, Pullen 64527 (GA, NCU); *Clarke Co., c. 1-5 km SW. of Pachuta, 20 July 1934, Harper 3238 (GH, MO, NY, US). North Carolina: Alexander Co., 3-2 km NW. of Vashti, 9 September 1966 (fl & fr), Bozeman, Ramseur & Radford 4510 (BM); Orange Co., Mason Farm road, c. 200 m N. of Golf Course, 24 June 1961 (fl & fr), Bell 17150 (H); Buncombe Co., Biltmore, 21 July 1897 (fl), Herb. Biltmore 523b (BM, GH*, ILL*, MO*, NY*, US*, W). Ohio: *Jackson Co., near Jackson, 20 August 1937, Drushel 10759 (MO, NY, US). South Carolina: *Aiken Co., Vancluse, 6 August 1898, Eggert s.n. (MO, NY, US); Hampton Co., S.C. Rt. 28, 2-7 km SE. of Early Branch, 29 June 1956, Ahles 15723 (K). Tennessee: Marion Co., Kim- ball, 5 July 1892 (fl), Middleton s.n. (BM); *Van Buren Co., Falls Creek Falls State Park, 20 July 1947, Shanks et al. 2991 (SMU, TENN). Virginia: *Dinwiddie Co., E. of McKenny, 13 October 1941, Fernald & Long 13976 (GH, NY, US); * Greenville Co., NE. of Orion, near Readjuster Bridge over Nottoway R., 13 June 1940, Fernald & Long 12132 (GH). West Virginia: New River, Hawk's Nest, July 1880 (fl & fr), Porter s.n. (K); *Fayette Co., edge of New River, Cotton Hill, 26 June 1941, Sharp s.n. (TENN). The Florida collections ('//. virgatum var. revolutum" R. Keller) have the small flowers and narrow leaves of H. harperi but not the spongy stem base, thus indicating that, although morphologically intermediate in some respects, the habit is dry like that of//, denticulatum subsp. acutifolium. 11. Hypericum harperi R. Keller in Bot. Jb. 58: 198 (1923); in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925). Type: U.S.A., Georgia, Sumter Co., 9 July 1900 (fl), Harper 1028 (Bt-lectotype (Fernald & Schubert (1948); BM!, GH!, MO*, NY*, US*-isolectotypes); for photo- graph see Rhodora 50: t.llll, f. 3 (1948). The selection of Harper 1006 as lectotype by Rodriguez Jimenez (1973) is therefore unacceptable, being later; and the Berlin speci- men, being the only one seen by Keller, must be the lectotype, not the BM one as designated by Webb. Map 16. H. denticulatum var. acutifolium sensu Fern. & B. G. Schubert in Rhodora 50: 208 (1948, pro parte); Fern., Gray's Man. Bot. 8th ed.: 1013 (1950), pro parte. H. denticulatum var. virgatum (Lam.) Rodriguez Jimenez in 67 U7 Mems Soc. Cienc. nat. La Salle 33: 100 (1973), pro parte, quoad syn. H. harperi. H. incertum sensu D. H. Webb, Biosyst. Study Hypericum sect. Spachium in eastern N. Am.: 296(1980), non Steudel (1840). Perennial herb 0-3-1 m tall, erect, branched from the base and usually with flowering branches from median and upper stem nodes. Stems green, 4-lined, densely gland-dotted, aerenchymatous at the base; internodes 8-20 mm long, about equalling leaves. Leaves sessile, ascending, deflexed when fading, lamina 10-30 x 3-5(-8) mm, narrowly oblong-elliptic (lower) or lanceolate to linear-lanceolate, margin plane, concolorous, not glaucous, subcoriaceous?; apex acute, base cuneate to rounded-subamplexicaul, sometimes subdecur- rent; basal or near-basal veins l-3(-5), midrib unbranched or with 1(2) pairs of branches, tertiary reticulation not visible; laminar glands dense, not prominent. Inflorescence up to c. 30-flowered, monochasial after c. 4th grade, with branches from up to 16 lower nodes, the whole broadly pyramidal to subcorymbiform; pedicels 1-5-2 mm long; bracts 2-6 mm long, lanceolate to subulate. Flowers c. 4-10 mm in diam., stellate. Sepals 3-5 x 0-8-1 mm, often unequal, lanceolate, acute to acuminate; veins 3-5, unbranched, all or only midrib prominent; glands linear, distally punctiform. Petals orange- yellow, 6-10 x 5-7 mm, c. 2 x sepals, obovate; apiculus obtuse; glands linear, interrupted distally. Stamens 50-80, irregularly (?) grouped, longest 3-6 mm long, c. 0-5 x petals. Ovary c. 1-1-5 x 0-7-1 mm, ovoid; styles 3, 2-4 mm long, 2-3 x ovary, divergent-incurved; stigmas capitate. Capsule 3—4-5 x 2-2-5(-3 ?) mm, ellipsoid to rostrate-subglobose, shorter than or equalling sepals. Seeds 0-5-0-6(0-65) mm long, testa obscurely linear-reticulate to irregularly reticulate. 2n = 24 (n = 12 -Webb, 1980). Open Taxodium swamps and wet pine barrens; lowland. South-eastern U.S.A. (coastal plain of South Carolina, Georgia, and northern Florida). U.S.A. Florida:* Gadsden Co., c. 5 km NW. of Gretna, 10 October 1959, Godfrey 58994 (GH, NCU, SMU); Jackson Co., Sneads, 20 October 1900 (fr). Curtiss 6731 (GH*, ILL*, 68 N. K. B. ROBSON K, MO*, NY*, US*). Georgia: Sumter Co., W. of Leslie, 9 October 1900 (fr), Harper 1731 (BM, GH);* Miller Co., c. 11 km ESE. of Colquitt, 8 July 1947, Thome 5209 (MO, NY, US);* Seminole Co., 2-4 km E. of Donaldsonville, 26 October 1963, Godfrey 63207 (NCU, SMU, US). South Carolina:* Berkeley Co., 9-5 km SW. of Monck's Corners, 7 August 1939, Godfrey & Try on 1454 (NY, TENN, US); *Orangeburg Co., Eutawville, 6-11 September 1900, Eggles- ton 5017 (GH, MO, NY). Despite the rather scathing comments by Fernald & Schubert (1948: 207) about Keller's habitual assumption that there were eastern North American species awaiting recognition, I agree with Webb (1980) that H. harperi is a good candidate for such recognition. Although clearly closely related to H. denticulatum subsp. acutifolium, it differs from the latter in its aquatic habitat and correlated spongy, aerenchymatous stem base, as well as in having rather smaller flowers and more stem branches. The Floridan collections of H. denticulatum subsp. acutifolium are intermediate in the last two characters, though apparently not in habitat. In addition, Webb (1980) has shown that H. harperi contains C-glycosyl flavones, whereas H. denticulatum does not. In addition to these ecological, morphological, and chemi- cal differences, H. harperi remains distinct in localities where it grows alongside H. denticulatum subsp. denticulatum, presumably because populations of the latter taxon are mostly tetraploid (2n = 48) and would therefore form triploid hybrids with H. harperi. 12. Hypericum setosum L., Sp. pi: 787 (1753); R. Keller in Bull. Herb. Boissier II, 8: 179 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Rodriguez Jimenez in C.r. Soc. Biogeogr. 432: 87, map 1 (1972), in Mems Soc. Cienc., nat. La Salle 33: 132, fig. Id (1973); N. Robson in Taxon 29: 273 (1980); D. H. Webb, Biosyst. Study Hypericum sect. Spachium in eastern N. Am.: 303 (1980). Type: U.S.A., Virginia, Clayton 153 (BM!- lectotype - Robson, 1980). Fig. 14A, Map 17. Ascyrum villosum L., Sp. pi.: 788 (1753); Miller, Card. diet. 8th ed.: No. 2 (1768). Type: Hypericum virginianum frutescens pilosissimum Plukenet, Phytographia: t. 245, f.6 (1691) (lectotype - Robson, 1980). Hypericum villosum (L.) Crantz, Inst. rei herb. 2: 520 (1766), non Crantz, Stirp. austr. fasc. 2: 62 (1763), 2nd ed.( 96 (1769). Hypericum pilosum Walter, Fl. carol.: 190 (1788); Gmelin, Syst. nat. 2: 1159 (1791); Choisy, Prodr. monogr. Hyperic.: 49 (1821), in DC., Prodr. 1: 549 (1824); Elliott, Sketch hot. S. Carolina 2: 25 (1821); Sprengel, Syst. veg. 3: 346 (1826); Torrey & A. Gray, Fl. N. Amer. 1: 163 (1838); D. Dietr., Syn. pi. 4: 1235 (1847); Trevir., Hyper, sp. animadv.: 9 (1861); Chapman, Fl. South. U.S.: 41 (1872); Coulter in Bot. Gaz. 11: 106 (1886); C. Mohr in Contr. U.S. natn Herb. 6: 622 (1901). Types: U.S.A., Carolina, Walter (?- holotype lost or destroyed); S. Carolina, Berkeley Co., c. 22 km S. of Moncks Corners, 11 August 1939, Godfrey & Try on 1424 (NY*-neotype; GH*, US*-isoneotypes - Webb, 1980). H. simplex Michaux, Fl. bor.-amer. 2: 80 (1803); Poiret in Lam., Encycl. Suppl. 3: 698 (1813); Pursh, Fl. Amer. sept. 2: 379 (1814); Elliott, Sketch hot. S. Carolina 2: 25 (1821); Choisy, Prodr. monogr. Hyperic.: 49 (1821), in DC., Prodr. 1: 549 (1824); Sprengel, Syst. veg. 3: 346 (1826), nom. illegit. superfl. (based on Ascyrum villosum L.). Map 17 Sect. 30: 12. H. setosum •; 13. H. cumulicola O. Data from Webb (1980). HYPERICUM 69 Fig. 14 A. H. setosum: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut away) and young capsule. B. H. cumulicola: (g) habit; (h) stem with leaves; (i) petal; (j) old stamens (partly cut away) and ripe capsule (a, g x '/2; b x 1; c, h, x 2; d-f, i, j x 6). A (a-c) Gray 10; (d-f) Sargent 10613. B. Lakela 24561. 70 N. K. B. ROBSON H. nuttallii G. Don, Gen. syst. 1: 607 (1831), nom. illegit. superfl. (based on H. pilosum Walter). Brathys tomentosa Spach, Hist. nat. veg. 5: 453 (1836), in Annls. Sci. nat. (Bot.) II, 5: 367 (1836), nom. illegit. superfl. (based on H. simplex Michaux). Icon: Plukenet, Phytographia: t. 245, f. 6 (1691). Perennial or annual herb 0-2-0-7(-0-8) m tall, erect, unbran- ched below inflorescence or occasionally with 1-2 flowering branches from upper stem nodes. Stems green, 4-lined, sparsely (?) gland-dotted, scabrous-tomentose to pilose; internodes 5-20 mm long, lower equalling leaves, upper longer than them. Leaves sessile, appressed to ascending, spreading when fading; lamina 4-15 x (1-5)2-7 mm, narrowly ovate or lanceolate to narrowly oblong-elliptic (or lower sometimes oblanceolate), margin recurved, concolorous, not glaucous, subcoriaceous, scabrous-tomentose to pilose; apex acute to obtuse, base rounded-subamplexicaul to parallel- sided, not decurrent, free; basal vein(s) l(-5), midrib without or with 1 pair of branches, tertiary reticulation not visible; laminar glands dense, slightly prominent beneath. Infloresc- ence up to c. 30-flowered (terminal), monochasial after 1st or 2nd grade, sometimes with up to 15-flowered dichasial/ monochasial or usually wholly monochasial branches from up to 5 lower nodes, occasionally with 1-2 pairs of subsidiary flowering branches below, the whole cylindric to subcorymbi- form; pedicels 1-5-2 mm long; bracts 2 mm long, lanceolate, setulose-ciliate. Flowers c. 5-11 mm in diam., stellate. Sepals (2-5-)3-5 x 1-5-2-5 mm, subequal, ovate to ovate-lanceolate or obovate, acute; veins 5-7, unbranched, only midrib or all veins slightly prominent, but less so than glands; glands linear, distally punctiform; margin setulose-ciliate, lamina sparsely setulose to glabrous. Petals (deep ?) yellow, 4-7 x 4- 7 mm, 1-5-2 x sepals, obovate; apiculus acute; glands linear, interrupted distally. Stamens (15) 20-40, almost free, longest 2-5-3 mm long, 0-65-0-75 x petals. Ovary c. 1-5 x 0-8 mm narrowly ovoid-ellipsoid; styles 3(4), 1-5-2 mm long, 1-1.3 x ovary, outcurving; stigmas ± broadly capitate. Capsule (3-5-)4-5 x 2-3 mm, ovoid to ellipsoid-subglobose, equalling or slightly exceeding sepals. Seeds 0-4-0-6 mm long, ecarinate; testa linear-reticulate. 2n = 12 (Webb, 1980). Wet ditches, bogs, savannahs, and wet pinelands on sandy soil; lowland. South-eastern U.S.A. (coastal plain from Virginia to north- ern Florida and west to Louisiana and E. Texas (fide Correll & Johnston, 1970)). U.S.A. Alabama: no precise locality, August 1841 (fl), Buckley BM [FI, G, MO]); *Escambia Co., 10-2 km E. of East Brewton on US 29, 16 July 1977, Webb & Ross 53283 (TENN). Florida: Duval Co., near Jacksonville, September 188?, Curtiss 268 (BM, F*, FI*, G, GA*, K, MO*, NY*, P*, US*); Union Co., N. of L. Butler, 7 August 1961 (fr), Gray 10 (BM, GH*, US*). Georgia: Sumter Co., August 1897 (fl), Harper s.n. [? 4403] (BM); *Tattnail Co., c. 3-2 km S. of Glenville, 11 September 1972, Godfrey 72101 (TENN, US). Louisiana: Rapides Par., Alexandria, no date (fl), Hale s.n. (K, NY*); Covington, June-August 1832 (fl), Drummond 246 (BM, K). Mississippi: Pearl River Co., c. 1-7 km W. of Picayune, 31 August 1974 (fl), Sargent 10613 (BM); Smith Co., Taylorville, 21 August 1903 (fl), Tracy 8646 (BM, GH*, MO*, NY*, SMU*). North Carolina: *Carteret Co., c. 0-8 km SE. of co. road 1103, off N.C. 58, S. of Kuhns, 10 September 1966 (fr), Bradley 3489 (BM, NCU*, TENN*); *Hertford Co., E. of Como, Camp Company Forest, entrance to Camp P-D, 23 July 1949, Fox & Godfrey 2793 (GH, NY). South Carolina: Berkley Co., 22-4 km S. of Moncks Corners, 11 August 1939 (fl & fr), Godfrey & Try on 1424 (GH, NY*, US*); Georgetown Co., 8 km S. of Andrews, 11 August 1939 (fl & fr), Godfrey & Tryon 1355 (GH, MO*, NY*, TENN*, US*). Virginia: *Brunswick Co., c. 2-3 km SE. of Triplett, 18 August 1942, Lewis s.n. (GH); *Sussex Co., c. 6-5 km NW. of Waverly, 10 September 1937, Fernald & Long 7532 (GH). H. setosum is related to H. denticulatum subsp. acutifolium , being smaller, without basal branches, and setose or pilose on stem, leaves, and sepals. It is the only species in sects 29-30 with an indumentum. Its nearest relative in H. denticulatum appear to be the population of subsp. acutifolum from Alexander Co., N. Carolina (growing on granite outcrops) that Webb (1980) regarded as somewhat aberrant. If H. setosum is indeed related to H. denticulatum subsp. acutifolium, then Webb's chromosome count (2n = 12) shows it to be haploid, or at least to have half as many chromosomes as its relative (2n = 24). How this halving has occurred is not at all clear; but I cannot agree with Webb that H. setosum has the most primitive chromosome number in Hypericum. 2n = 12 is clearly not a basic diploid number, otherwise the arborescent and shrubby species with 2n = 24 would have to be regarded as tetraploids and a sister group in some obscure way to H. setosum (and H. cumulicola q.v.). 13. Hypericum cumulicola (Small) P. Adams in Rhodora 64: 234 (1962); D. H. Webb, Biosyst. Study Hypericum sect. Spachium in eastern N. Am.: 169 (1980); Abrahamson et al. in Fla Sclent. 47: 221 (1984). Type: U.S.A., Florida, between Avon Park and Sebring, 30-31 August 1922, Small et al. s.n. (NY*-holotype). Fig. 14B, Map 17. Sanidophyllum cumulicola Small in Bull. Torrey hot. Club 51: 391 (1924). Hypericum gentianoides sensu Rodriguez Jimenez in Mem. Soc. Cienc. nat. La Salle 33: 112 (1973), pro parte. Icon: D. Ward in P. Pritchard, Rare & endangered biota of Florida: 34, f. 21 (1980). Perennial herb 0-2-€-7(-0-75) m, erect, glabrous, with bran- ches at or just below ground level, each unbranched below inflorescence. Stems green, 4-lined, rather sparsely gland- dotted; internodes (mature) 3-11 mm long, exceeding leaves. Leaves sessile, subulate, appressed (squamiform and spread- ing in juvenile stems); lamina (l-)2-5-4 x 0-2-0-3 mm, HYPERICUM linear-subulate, margin incurved, concolorous, not glaucous, subcoriaceous; apex acute, base parallel-sided, not decurrent, free; basal vein 1, unbranched; lamina glands in c. 2 lines, not prominent. Inflorescence up to c. 13-flowered, monochasial after 1st grade, sometimes with up to c. 9-flowered dichasial/ monochasial or wholly monochasial branches from 1-2 lower nodes, without subsidiary flowering branches below, the whole subcorymbiform; pedicels 0-5-1 mm long: bracts 1-1-5 mm long, subulate. Flowers c. 3-4 mm in diam., stellate. Sepals 1-5-2 x 0-6-1 mm, unequal, ovate to elliptic or narrowly oblong, acute to subacute; veins 5, unbranched, all slightly prominent, midrib often apically indurated-cucullate; glands linear; margin irregularly ciliolate to entire. Petals yellow, 3-5-5 x 1-3-1-7 mm, c. 2-5 x sepals, obovate-oblong; apiculus subacute; glands linear, interrupted distally. Stamens 20-25, longest 1-5-2 mm long, c. 0-4 x petals. Ovary c. 2 x 0-5 mm, narrowly ovoid-pyramidal; styles 3, 1-5-2 mm long, 0-75-1 x ovary, spreading; stigmas capitate. Capsule (3-5-)4- 6 x 1-1-5 mm, narrowly ovoid-conic, apex subrostrate, exceeding sepals. Seeds 0-5-0-6 mm long, not or scarcely carinate; testa scalariform-reticulate. 2n = 12 (Webb, 1980). Scrub or ancient white-sand dunes; lowland. South-eastern U.S.A. (Florida). U.S.A. Florida: Polk Co., Frostproof, E. of Maintenance Station, L. Clinch, 30 September 1961 (fr), Lakela 24710 (BM, NY*, SMU*); Frostproof, path 630 connecting US 27 and US 27A, 23 August 1961 (fr), Lakela 24561 (BM, GH*); * Highlands Co., 48 km S. of Sebring along Rt. 27, 25 August 1951, Webster 4177 (GH, MICH, SMU). Highlands Co., c. 10-5 km S. of Fla. 70 and US 27, 1 August 1960, Adams 586 (GA*, GH, K, NY*, SMU, TENN*). De Sola Co., Avon Park to Sebring, 1 May 1919, Small & De Winkeler 9040 (K, MICH*, NY*). Webb followed Adams in comparing the highly reduced H. cumulicola with the equally reduced H. gentianoides , and Rodriguez Jimenez included it in that species. The habit and floral details of H. cumulicola, however, are not very like those of H. gentianoides (except possibly the conical capsule); they are much more similar to H. setosum, which has the same unusual chromosome number. Indeed the ciliola on the sepal margin of H. cumulicola can be interpreted as the last remains of the indumentum of H. setosum. Both species grow on sandy soils, but H. cumulicola inhabits drier areas than does H. setosum. H. cumulicola occurs only in the white wind-deposited sands of the Central Florida range, from southern Highlands Co. north to the vicinity of Frostproof, Polk Co. Its very restricted habitat is under threat from 'the bulldozer and the citrus grove' (Ward, 1980: 34). 14. Hypericum denudatum A. St. Hil., Fl. Bras. mend. 1: 336 (1828); Walp., Repert. hot. syst. 1: 390 (1842); D. Dietr., Syn. pi. 4: 1235 (1847); Lyman B. Smith in/. Wash. Acad. Sci. 48: 314 (1958), pro parte quoad typum; Angely, Fl. Anal. Parana: 450 (1965). Type: Brasil, Parana, Campos Geraes, banks of torrent Yapo [lapo] near cottage Barro do Yapo, February 1816-1821 (fl), St.-Hilaire 1530 (P!- holotype; F!, US!-photographs). Map 18. H. rigidum var. humile Reichardt in Martius, Fl. bras. 12(1): 189 (1878). Types: as for H. denudatum A. St. -Hil. 71 (obligate lectotype); Brazil, Sao Paulo, in pratis prope Mugi, /Jfede/ (W?). H. linoides sensu Reichardt in Martius, Fl. bras. 12(1): 192 (1878), pro parte excl. typum, non A. St. Hil. (1828). H. brasiliense var. brasiliense sensu Rodriguez Jimenez in Reitz, Fl. III. Catar., Hipericdc.: 23 (1980), pro parte, non Choisy (1824). Shrub or subshrub 0-3-1(1-5) m tall, erect to decumbent, branching at the base or usually 1 -stemmed, with branches erect or divaricate-ascending, pseudo-dichotomous or lateral, numerous and often fastigiate, soon naked below. Stems dull red-brown, 4-lined and ancipitous when young, eventually terete, cortex exfoliating in strips; internodes 1-25 mm long, usually shorter than leaves. Leaves sessile, appressed to ascending; lamina 4-13(-18) x 0-8-3(^1) mm, narrowly elliptic-oblong to linear, plane, concolorous or almost so, not glaucous, subcoriaceous; apex acute to obtuse or sometimes rounded, base cuneate, not decurrent, free; basal or near- basal veins 5(3), midrib with 1-2 obscure pairs of branches, tertiary reticulum obscure or absent; laminar glands dense, visible above. Inflorescence l(2-5)-flowered, dichasial/ monochasial with lateral branches from up to 18 nodes below, the whole cylindric or fusiform; primary pedicels l-5-3(-6) mm long; bracts and bracteoles linear-subulate. Flowers 10- 16 mm in diam., stellate; buds ovoid-ellipsoid to narrowly ovoid, acute. Sepals 3-5-5 x 0-8-1-5 mm, subequal, not or scarcely imbricate, linear-lanceolate to narrowly lanceolate, acute; veins 5(3), only midrib sometimes prominent; glands wholly linear or punctiform near apex and margin. Petals yellow, 5-12 x 2-4-5 mm, 1-7-2-5 x sepals, oblanceolate; apiculus acute; glands mostly linear, distally punctiform. Stamens 30-50, obscurely (3-?) 5-fascicled, longest 3-6 mm long, 0-5-0-6 x petals. Ovary 1-5-2-5 x 1-1-5 mm, narrowly ellipsoid; styles 3-5, 1-5-3 mm long, 1-1-2 x ovary, outcurv- ing; stigmas broadly capitate. Capsule 5-6-5 x 2-5-3(-4) mm, narrowly ovoid-pyramidal to ovoid, exceeding or rarely 72 N. K. B. ROBSON equalling sepals. Seeds 0-6-0-7 mm long; testa ribbed- scalariform. Wet meadows, marshes, and bogs (thickets in Rio Grande do Sul); 720-1200 m (lowland in Rio Grande do Sul?). Brazil (Parana to northern Rio Grande do Sul), Argentina (Misiones). BRAZIL. Parana: mun. Gal. Carneiro, Lageado, 12 February 1966 (fl & fr), Hatschbach 13732 (F, NY, P. U); Tres Barras, Rio Negro, c. 720 m, 25 January 1916 (fl), Dusen 17527 (BM, F, G, K, NY, S, US); Guajuvira, 22 November 1909 (fl & fr), Dusen 8937 (G, GH, MO, NY, S, Z). Santa Catarina: mun. Agua Doce, 22 km S. of Horizonte (Parana), 2 December 1971 (fl), Smith & Klein 15596 (P, NY); mun. Curitibanos, 2-3 km W. of Curitibanos towards Campos Novos, c. 850 m, 9 February 1957 (fl & fr), Smith & Klein 11096 (NY); mun. Porto Uniao, 42 km S. of Porto Uniao, Matos Costa, c. 900-1100 m, 3 December 1964 (fl), Smith & Klein 10851 (NY, US). Rio Grande do Sul: Cai, 27 April 1949 (fl & fr), Rambo 41254 (K, NY, S); mun. Bom Jesus, Fazenda Bernardo Velho, 2 January 1947 (fl & fr), Rambo 34786 (F, S, US); prope San Leopoldo, Esteio, 20 November 1950 (fl), Rambo 49162 (NY, S). H. denudatum resembles 16. H. brasiliense in having a relatively narrow capsule that nearly always exceeds the sepals. Its branching, however, is sometimes pseudo- dichotomous, whereas that of H. brasiliense is always lateral, the usually shorter leaves are ± concolorous and sub- coriaceous, with only 3-5 basal veins that branch obscurely, if at all. The inflorescence is l(2-5)-flowered (not 8-30- flowered), the stamens are fewer, the capsule usually narrowly ovoid-pyramidal (not cylindric), and the seeds somewhat longer. H. denudatum has been misunderstood since Reichardt treated it as a prostrate variety of//, rigidum, which (in turn) he had wrongly associated with H. cordiforme (= H. cor datum). He also confused it with H. linoides. In fact it is quite distinct from these species and is easily recognizable by its short narrow leaves with 3-5 nerves and its small flowers, usually solitary, with narrow sepals and narrow capsules. The most primitive form (erect with relatively few bran- ches, larger leaves and flowers, and 5 styles) occurs in Parana and adjacent Santa Catarina. From this there are two reduction trends: (i) northward, where the branching be- comes more diffuse, the habit decumbent (or prostrate), the leaves and flowers much smaller, and the styles gradually reduced to 3; (ii) southward, where the stem remains erect but the more numerous fastigiate branches give the plant a broom-like appearance. Here, too, the leaves and flowers are reduced in size and the styles are occasionally reduced to 4 or 3. Although the extreme forms of these morphoclines look very different, the occurrence of a series of intermediates linking both to the primitive form precludes the recognition of infraspecific taxa. 15. Hypericum pedersenii N. Robson, sp. nov. Map 18. //. denudato A. St. Hil. affinis, sed habitu prostrato- adscendente, foliis minoribus acerosis incurvis glaucis 1- nervibus, floribus minoribus, differt. Type: Brazil, Rio Grande do Sul, mun. arroio dos Ratos, Faxinal, 10 November 1977 (fl), Pedersen 11972 (BM!-holotype). Wiry shrublet up to c. 0-18 m tall, ascending from prostrate and diffusely branched base, not rooting, with branches erect or divaricate-ascending, lateral or rarely pseudo- dichotomous, numerous, soon naked below. Stems 4-lined and ancipitous when young, eventually terete, cortex exfoliat- ing in strips; internodes 3-9-5 mm long, equalling or exceeding leaves. Leaves sessile, appressed; lamina 3-4 x 0-5-0-8 mm, acerose, incurved, concolorous, glaucous, sub- coriaceous; apex acute, base subamplexicaul, not decurrent, free; basal vein 1, midrib unbranched; laminar glands rather dense, impressed beneath. Inflorescence 1-3-flowered, dichasial/monochasial with lateral branches from 12 or more nodes below, the whole narrowly pyramidal; primary pedicels 0-5-1 mm long; bracts and bracteoles linear-subulate. Flowers 8-9 mm in diam., stellate; buds ovoid, subacute. Sepals 2-2-5 x 0-5-1 mm, subequal, not imbricate, narrowly lanceolate, acute; veins 3, all or only midrib prominent; glands linear, punctiform in distal 1A. Petals apricot? yellow, c. 4-5 x 2 mm, 1-8-2-2 x sepals, oblong-oblanceolate; apiculus acute; glands Map 18 Sect. 30: 14. H. denudatum •; 15. H. pedersenii O. HYPERICUM 73 linear or mostly striiform. Stamens 30-35, obscurely 3- fascicled, longest 2-5 mm long, 0-55 x petals. Ovary 1-5 x c. 0-8 mm, narrowly ovoid; styles 3, 1-5 mm long, equalling ovary, outcurving; stigmas scarcely enlarged. Capsule c. 3 x 2 mm (?immature), narrowly ovoid-pyramidal. Seeds not seen. Rocky ground. Brazil (Rio Grande do Sul). BRAZIL. Rio Grande do Sul: mun. Arrois dos Ratos, Faxinal, 10 November 1977 (fl), Pedersen 11972 (BM). H. pedersenii is clearly related to the diffusely branched, small-leaved, small-flowered form of H. denudatum from southern Parana, but differs in the more extremely prostrate- ascending habit, the smaller incurved glaucous leaves with a single unbranched vein (midrib), and the smaller flowers. It has been collected only once. 16. Hypericum brasiliense Choisy in DC., Prodr. 1: 547 (1824); A. St. Hil., Fl. Bras, merid. 1: 335 (1828); Cham. & Schlechtendal in Linnaea 3: 121 (1828) [amplified descr.]; D. Dietr., Syn. pi. 4: 1235 (1847); Reichardt in Martius, Fl. bras. 12 (1): 193, t. 34 (1878); Arechav. in An. Mus. Hist. nat. Montevideo 3: 110 (1898), pro parte quoad typum; R. Keller in Bull. Herb. Boissier 6: 266 (1898), in op. cit. II, 8: 181 (1908), in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 183 (1925); Chodat & Hassler in Bull. herb. Boissier II, 3: 1126 (1903); Lyman B. Smith in /. Wash. Acad. Sci. 48: 313 (1958); Forster in Contr. Gray Herb. Harv. 184: 131 (1958); Rodriguez Jimenez in C. r. Soc. Biogeogr. 432: 191, map 6 (1972), in Mems Soc. Cienc. nat. La Salle 33: 118, f. 15c (1973), in Reitz, Fl. III. Catar., Hipericdc.: 23 (1980), pro parte. Type: Brazil, ad Rio-Janeiro, 1830 (fl & fr), Gaudi- chaud 1 (G!-neotype; Pi-photograph). This specimen, which was cited as the type by Rodriguez Jimenez and is so labelled, cannot be the one on which Choisy based his description, as it is dated 1830. There is no specimen of H. brasiliense in Herb. De Candolle (G-DC) and none in the main Geneva herbar- ium (G) or in Paris (P) that could be the holotype. I therefore select Gaudichaud 1, which was annotated by Choisy, as the neotype. Fig. 15D, Map 19. Receveura graveolens Veil. Cone. , Fl. flumin. : 237 (1825), 5: t. 120 (1835), in Archos Mus. nac. Rio de J. 5: 223 (1881). Type: Brazil, Rio de Janeiro (or Minas Geraes), Vandelli (LISU-holotype). H. laxiusculum A. St. Hil., PL usuell. bras.: t. 62 (March 1828), Fl. Bras, merid. 1: 332 (September 1828); Walp., Repert. hot. syst. 1: 389 (1842); D. Dietr., Syn. pi. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 12(1): 192 (1878); R. Keller in Bull. Herb. Boissier 6: 267 (1898); Lyman B. Smith in /. Wash. Acad. Sci. 48: 313 (1958); Angely, Fl. Anal. Parana: 451 (1965), Fl. Anal. S. Paulo 1: 186 (1969). Type: Brazil, Sao Paulo, Campos Geraes, Fazenda de Fortaleza, February [1820] 1816-1821 (fl & fr), St.-Hilaire 1457 ter (Pl-lectotype- Rodriguez Jimenez, 1973); loc. cit., [1820] 1816-1821 (fl & fr), St.-Hilaire 1457, 1457a (P!- syntypes). H. punctulatum A. St. Hil., Fl. Bras, merid. 1: 334 (1828); Walp., Repert. hot. syst. 1: 390 (1842); D. Dietr., Syn. pi. 4: 1236 (1847). Type: Brazil, Minas Geraes, 'haud longo ab urbe Mantequeira', [1816?] October 1816-1821 (fr), St.- Hilaire 123 (P!-holotype). H. brasiliense var. B A. St. Hil., Fl. Bras, merid. 1: 335 (1828). H. brasiliense var. angustifolium Reichardt in Martius, Fl. bras. 12(1): 193 (1878); Arechav. in An. Mus. Hist. nat. Montevideo 3: 111 (1898), pro parte quoad typum; Chodat & Hassler in Bull. Herb. Boissier II, 3: 1126 (1903), pro parte quoad typum. Type: Brazil, Minas Geraes, 'prope praediolum vulgo Caso do Bareto', [1817] October 1816- 1821 (fl), St.-Hilaire 2158 (P!-holotype). H. brasiliense var. latifolium Reichardt in Martius, Fl. bras. 12(1): 193 (1878); Arechav. in An. Mus. Hist. nat. Montevideo 3: 111 (1898), pro parte quoad typum; Lyman B. Smith in/. Wash. Acad. Sci. 48: 313 (1958); Angely, Fl. Anal. Parana: 450 (1965). Type as for H. punctulatum. H. brasiliense var. punctulatum (A. St. Hil.) R. Keller in Bull. Herb. Boissier II, 3: 1126 (1903), pro parte quoad typum. H. stylosum Rusby in Bull. N.Y. hot. Gdn. 4: 326 (1907); Foster in Contr. Gray Herb. Harv. 184: 131 (1958). Type: Bolivia, Yungas, Coripati, 30 March 1894 (fl & fr), Bang 2107 (NY! -holotype; BM!, G!, GH!, K!, MICH!, MO!, US!,W!-isotypes). H. bolivianum R. Keller in Bull. Herb. Boissier II, 8: 189 (1908); Forster in Contr. Gray Herb. Harv. 184: 131 (1958). Type as for H. stylosum (Bang 2107) (G?! or W?!- holotype; BM!, GH!, K!, MICH!, MO!, US!). Sarothra brasiliensis (Choisy) Y. Kimura in Nakai & Honda, Novafl. jap. 10: 71, 238 (1951); Angely, Fl. Anal. S. Paulo 1: 187 (1969). Icon: Reichardt in Martius, Fl. bras. 12(1): t. 34 (1878). Subshrub (? or annual herb) 0-3-1 m tall, erect, 1-stemmed with branches divaricate-ascending, lateral, numerous, be- coming naked below. Stems vinous ? red, 4-lined, and ancipitous when young, eventually terete, cortex exfoliating in strips; internodes (5-) 10-90 mm long, shorter to longer than leaves. Leaves sessile, spreading; lamina 8-30 x 2-7(-ll) mm, narrowly elliptic to narrowly oblong or rarely lanceolate, plane, usually paler beneath, not glaucous, chartaceous; apex acute to rounded, base cuneate, not decurrent, free; basal or near-basal veins 3-9, midrid with 2-c.9 pairs of branches, tertiary reticulum dense (usually obscure) or absent (?); laminar glands dense, not prominent. Inflorescence c. 9-17 (-24)-flowered, dischasial/monochasial, sometimes with short accessory branches from terminal node, with lateral branches from up to 8 nodes below, the whole narrowly cylindric to narrowly or broadly pyramidal; primary pedicels 2-6 mm long; bracts and bracteoles narrowly lanceolate to linear. 74 N. K. B. ROBSON Flowers 10-15 mm in diam., stellate; buds narrowly ovoid, acute to rounded. Sepals 3-5-6 x 0-7-2 mm, subequal, basally imbricate, linear-lanceolate to ovate-lanceolate, acute; veins 5(3), all becoming prominent; glands linear. Petals yellow, 5- 8 x 2-4-5 mm, 1-3-1-5 x sepals, narrowly obovate-oblong; apiculus acute; glands linear. Stamens c. 75, 5-fascicled, longest 3-4-5 mm long. c. 0-6 x petals. Ovary 1-5-2 x 0-8-1-1 mm, narrowly ellipsoid; styles 5 (or very rarely 4), 2-2-5 mm long, 1-2-1-35 x ovary, spreading-incurved; stigmas slightly enlarged to narrowly capitate. Capsule (5-)6 x (2-5-)3 mm, cylindric or rarely ovoid-cylindric, almost always exceeding sepals. Seeds 0-5-0-6 mm long; testa finely ribbed- scalariform. 2n = 24? (see p. 87). Damp meadows, ditches, roadsides, and waste places; 300 (Santa Catarina) - 2525 (Espirito Santo) m. Brazil (Bahia, Espirito Santo, and Minas Geraes to Santa Catarina), Paraguay (Amambay?), Bolivia (La Paz, Santa Cruz), Argentina (Corrientes, Chaco, Tucuman). BRAZIL. Bahia: c. 5-5 km SE. of Barra do Choca on road to Itapetinga, 700-800 m, 30 March 1977 (fl & fr), Harley 20173 (BM, K). Espirito Santo: Pico de Bandeira, Serro Caparao, c. 2525 m, 1 March 1959 (fl), Irwin 2751 (F, NY). Distr. Fed.: Brasilia, Zoobotanico, 16 February 1964 (fl & fr), Heringer 9947 (P). Minas Geraes: Ouro Preto, 930 m, 9 December 1921 (fl & fr), E. & M. Holway 1379 (US); c. 12 km SW. of Barao de Cocais, base of Serra de Caraga, c. 1500 m, 28 January 1971 (fl), Irwin, Harley & Onishi 29300 (C, F, H, K, NY); Maria da Fe, 31 August 1946 (fl), Duarte 258 (MO, NY). Rio de Janeiro: Serro Orgao, Guanabara, Edo. Rio de Janeiro Nat. Parque, 1860-1920 m, 16 December 1959 (fl & fr), B. & C. Maguire 44603 (NY); vicinity of Macieiras, Mt. Itatiaia, Estac.ao Biologica, c. 2000 m, 10 December 1928 (fl), L. B. Smith 1462 (GH, US): mun. Petropolis, km 20 from Itaipava to Tereopolis, 24 February 1963 (fl & fr), Pabst 7309 (K). Sao Paulo: San Bernardo, December 1911 (fl), Brade 5258 (S); Campo de Jordao, 5-20 February 1937 (fl), Porto 2970 (NY, P); Villa Prudente, 20 December 1907 (fl & fr), Usteri s.n. (ZT). Parana: San Cristovao, 18 October 1972 (fl), Hatschbach 30675 (BM, MBM, P); mun. Balsa Nova, San Luis de Puruna, 14 December 1979 (fl & fr), Hatschbach 42642 (BM, MBM); mun. Laranjeiras do Sul, 10 December 1968 (fl), Hatschbach 20597 (BM, MBM). Santa Catarina: mun. Chapeco, W. of Chapeco on road to Guatambu, 300- 400 m, 15 October 1964 (fl), Smith & Reitz 12531 (NY). PARAGUAY. Anambay?: Sierra de Maracayu, prope Ipe hu, November 1898 (fl & fr), Hassler 5250 (A, BM, G, K, NY, P, S, W). BOLIVIA. Santa Cruz: Florida, Samaipata, c. 2000 m, March 1911 (fl & fr), Herzog 1718 (S); Santa Cruz, 1500 m, 12 January 1864 (fr), Pearce s.n. (BM). La Paz: Nor Yungas, Polo-Polo bei Coroico, 1100 m, November 1912 (fl), Buchtien 244 (BM, G, GH, K, US); Inquisivi, Inquisivi 35 km hacia NW. via Circuata, 2300 m, 21 February 1981 (fl & fr), Beck 4555 (BM, UMSA n.v.). ARGENTINA. Corrientes: Beron de Astrada, Arroyo Santa Isabel [al] boca del Parana, 1 December 1945 (fr), Ibarrola 3965 (W); Chaco: Fontana, March 1933 (fr), Meyer 2097 (K). Tucuman: Tafi, Siombon, 20 November 1944 (fl & fr), Obeo 10 (NY). H. brasiliense can be distinguished from all members of the H. campestre group (Spp. 18-24), with which it has been frequently confused, by the cylindric capsule that almost always exceeds the sepals. It is the only 5-styled suffruticose species with such a capsule. For differences between H. brasiliense and 14. H. denudatum, see p. 72. Even though the exclusion of the H. campestre group Map 19 Sect. 30: 16. H. brasiliense •; 17. H. polyanthemum O. HYPERICUM 75 reveals it to be much less variable than has hitherto been thought, H. brasiliense still shows considerable variation. The plants that occur in the northern part of its range (Bahia to Minas Geraes and Rio de Janeiro) include those most similar to H. campestre, with 7-9 veined leaves, a narrow infloresc- ence, and narrow sepals that sometimes almost equal the capsule in length. From this form, two morpho-geographical trends are apparent: (i) Southward the narrow inflorescence persists into Parana and western Santa Catarina, but the leaves and flowers become smaller and the sepals broader. This trend leads to 17. H. polyanthemum. (ii) The forms represented in trend (i) are apparently rare in Sao Paulo, where representatives of H. brasiliense tend to be more laxly branched and more numerous-flowered (H. laxiusculum). Such plants also occur in states adjacent to Sao Paulo, but they are characteristic of a western trend (Paraguay, northern Argentina, Bolivia) ('//. stylosum') in which the flowers become smaller and the leaves sometimes broader - but not with more numerous basal veins. Extreme forms of both these trends are linked to typical H. brasiliense by a series of intermediates, so that no infraspecific taxa can be recognized. 17. Hypericum polyanthemum Klotzsch ex Reichardt in Martius, Fl. bras. 12(1): 189 (1878); Lyman B. Smith in/. Wash. Acad. Sci. 48: 314 (1958); Angely, Fl. Anal. Parana: 451 (1965). Type: Brazil, 'in Brasilia meridionali', n.d. (fl), Sellow 2898 (Bt-holotype, F!, NY!-photographs; K!- isotype). Map 19. H. rivulare Arechav. in An. Mus. Hist. nat. Montevideo 3: 109 (1898); Lyman B. Smith in/. Wash. Acad. Sci. 48: 313, f. 1 j-1 (1958). Type: Uruguay, 'Habita en orillas del arroyo Tambores, en las cercanias del valle Eden', n.d. (fl & fr), Arechavaleta 666 (MVM-holotype; US!-isotype (frag- ment); GH!, US!-photographs). H. brasiliense sensu Rodriguez Jimenez in Mems Soc. Cienc. nat. La Salle 33: 118 (1975), pro parte quoad spec. Reitz & Klein 7933, Reitz 1965, Rambo 51825, 51883; in Reitz, Fl. 111. Catar., Hipericdc.: 23 (1980), pro parte, quoad Reitz & Klein 7933. H. brasiliense var. linoides sensu Rodriguez Jimenez in Mems Soc. Cienc. nat. La Salle 33: 126 (1973), pro parte quoad spec. Reitz 1965, Rambo 51825, 51883. Icon: Lyman B. Smith in /. Wash. Acad. Sci. 48: 312 f. Ik-m (1958). Subshrub 0-1-0-4 m tall, erect or decumbent, branches spreading or divaricate-ascending, lateral, near base or all along stem, very numerous, sometimes naked below. Stems yellow to orange-brown, sharply 4-lined, ancipitous when young, cortex exfoliating in strips; internodes 2-14(-35) mm long, the upper sometimes exceeding leaves. Leaves sessile, spreading; lamina 3-15 x 0-5-1 -5(-2) mm, narrowly oblong to linear, margin revolute, paler beneath, not glaucous, chartaceous, apex obtuse to rounded, base parallel-sided or usually cordate-amplexicaul, not decurrent, free; basal vein 1, sometimes with obscure lateral branches, tertiary reticulum absent; laminar glands dense, ± prominent beneath. In- florescence 3-16-flowered, dichasial/monochasial, with lateral branches from up to 3 nodes below, the whole narrowly cylindric; primary pedicels 0-5-3 mm long; bracts and bracteoles triangular-lanceolate to linear-subulate. Flowers 8-12 mm in diam., stellate or slightly obconic; buds ovoid, acute to subacute. Sepals 3-5^-5(-5) x 1-2-2-4 mm, ± unequal, imbricate, the outer broadly rhombic-elliptic or ovate-lanceolate to ovate, the inner ovate-lanceolate to lanceolate or oblong, acute to obtuse, margin sometimes recurved; veins 5, only midrib becoming prominent; glands linear, distally punctiform. Petals yellow, tinged red outside, 5-7 x 2-5-4 mm, 1-5-1-7 x sepals, oblong; apiculus obtuse; glands striiform to punctiform. Stamens c. 40-50 (or more?), obscurely 3-5-fascicled, longest (2-)3-4 mm long, 0-5-0-7 x petals. Ovary 1-5-2 x 1 mm, ovoid; styles (3-)4-5, 1-5-2-5 mm long, 1-1-35 x ovary, spreading; stigmas slightly enlarged to distinctly capitate. Capsule 3-5-5 x 2-4 m, ovoid to subglobose, about equalling sepals. Seeds 0-6-0-7 mm long; testa finely ribbed-scalariform. Grassland and other open, stony, medium-dry to rather wet ground; 10-1300 m. Brazil (Santa Catarina, Rio Grande do Sul), Uruguay. BRAZIL. Santa Catarina: Sombrio, 10 m, 9 October 1945 (fl), Reitz 1965 (S, US); Bom Jardin, Sao Joaquin, 1300 m, 15 December 1958 (fl), Reitz & Klein 7933 (G, US); Paca, Lagos de Cavera, 22 November 1950 (fl & fr), Hans 418 (P). Rio Grande do Sul: mun. Lavros do Sul, Coxilles de Tabuleiro, 9 November 1976 (fl & fr), Pedersen (BM, C); Passo da Guarda, prope Bom Jesus, 15 January 1952 (fl & fr), Rambo 51825 (S, US); Taimbe, prope Francesca de Paula, 21 February 1951 (fl & fr), Rambo 50128 (S). URUGUAY. Tacuarembo: Arroyo, Tambores, [rec. 1925] (fl & fr), Filippone 5068 (K). Cerro Largo: Sierra Acequa, Huila, January 1926 (fl), Herter 18567 (GH-photograph). H. polyanthemum is a southern derivative of H. brasiliense, sharing with the nearest form of that species the short leaves, narrow inflorescence, and broad sepals. It differs from H. brasiliense in the frequently decumbent, much-branched habit, the revolute, usually cordate-amplexicaul leaves, the broader ovary with styles sometimes reduced to 4 or even 3, and the ovoid to subglobose capsule that rarely exceeds the sepals. 18. Hypericum campestre Cham. & Schlechtendal in Linnaea 3: 122 (1828); Walp., Repert. hot. syst. 1: 388 (1842); D. Dietr., Syn. pi. 4: 1236 (1847); Reichardt in Martius, Fl. bras. 12(1): 192 (1878); Arechav. in An. Mus. Hist. nat. Montevideo 3: 110 (1898), pro parte; R. Keller in Bull. Herb. Boissier 6: 266 (1898), pro parte quoad typum, in op. cit. II, 8: 181 (1908), in Engl. & Prantl, Nat. Pflanzenfam. II, 21: 183 (1925); Malme in Ark. Bot. 23A(4): 17 (1930); Herter, Fl. illust. Urug.: f. 2200 (1957); Lyman B. Smith in J. Wash. Acad. Sci. 48: 313 (1958); Angely, Fl. Anal. Parana: 450 (1965), F. Anal. S. Paulo 1: 185 (1969); non 76 N. K. B. ROBSON Moon, Cat. pi. Ceylon: 56 (1824), nomen (= H. japonicum Thunb. ex Murray). Type: Brazil, Parana, 'ad fluvium Rio Negro aliisque pluribus locis', 1815-1817 (fl & fr), Sellow 1655 (Bt-lectotype; F!, GH!-photographs; K!-isotype); Sellow 385 (Bt-syntype); Sellow 4311 (Bt-syntype). Fig. 15A-C. H. brasiliense sensu Chodat & Hassler in Bull. Herb. Boissier II, 3: 1126 (1903), pro parte, quoad spec, cit.; Rodriguez Jimenez in C. r. Soc. Biogeogr. 432: 91, map 6 (1972), pro parte, in Mems Soc. Cienc. not. La Salle 33: 118 (1973), in Reitz, Fl. III. Catar., Hipericdc.: 23 (1980), pro parte et auct. plur. Icon: Herter, Fl. illust. Urug.: f. 2200 (1957). Subshrub to annual herb, 0-5-2 m tall, erect, 1-stemmed, with branches ascending to spreading or rarely strict, lateral, usually numerous, becoming naked below. Stems dull red to yellow-brown, 4-lined, ancipitous when young, eventually terete, cortex exfoliating in strips; internodes 10-60(-80) mm long, eventually longer than leaves. Leaves sessile, spreading; lamina 12-38 x 2-7 (-10) mm, narrowly elliptic or rarely narrowly oblanceolate to linear, plane or margins recurved to revolute, paler beneath, not glaucous, chartaceous; apex acute to subacute or rarely obtuse, base cuneate to rarely rounded, not decurrent but sometimes forming shallow V, free; basal veins 3-5, midrib with 2-4 pairs of branches, tertiary reticulum obscure or absent (?); laminar glands dense, slightly prominent beneath. Inflorescence 1->100- flowered, diachasial/monochasial, sometimes with accessory branches, with lateral branches from up to 6 nodes below, the whole narrowly cylindric to broadly and laxly pyramidal or rarely subcorymbiform (i.e. not flat-topped); primary pedi- cels l-3(-5) mm long; bracts (rarely) foliar or narrowly lanceolate to linear, bracteoles linear. Flowers (8-) 10-24 mm in diam., stellate; buds ovoid, acute. Sepals 4-6 x (0-7-)0-9- l-l(-l-4) mm, equal, not imbricate, linear-lanceolate, acute; veins 5(7), all becoming prominent; glands linear, distally sometimes striiform. Petals yellow, 5-12 x 2-5-5 mm, 1-65- 1-75 x sepals, obovate-oblanceolate to oblanceolate-oblong; apiculus acute or obsolete; glands linear, distally punctiform. Stamens 65-85, 5-fascicled, longest 3-6 mm long, 0-4-0-6 x petals. Ovary 2-2-5 x 1-1-5 mm, ovoid; styles 5 (rarely 4-3), 2-3 mm long, 1-1-25 x ovary, spreading; stigmas narrowly capitate. Capsule (4-)5-6 x (3-)4 mm, ± broadly ovoid, shorter than or equalling sepals. Seeds c. 0-5 mm long; testa ribbed-scalarif orm . Damp or shaded grassland, roadsides; 90-900 m. Brazil (Sao Paulo?, Parana, Santa Catarina, Rio Grande do Sul), Paraguay, Argentina (Misiones, Corrientes, Chaco), Uruguay. H. campestre has been much confused with H. brasiliense and H. carinatum. It can be distinguished from H. brasiliense by the narrow sepals that are never exceeded by the ± broadly ovoid capsule and from H. carinatum by the narrow leaves with bases not decurrent and the absence of red tinges from the petals. The longer leaves, broader capsule, and usually several- to many-flowered inflorescence differentiates H. campestre from H. denudatum. Even with the exclusion of these species, H. campestre remains very variable. The most primitive form, with a few- flowered inflorescence and large flowers, occurs in Rio Grande do Sul (narrow leaves), Santa Catarina and Parana (broad leaves). From the narrow-leaved form a reduction trend with few-flowered inflorescence leads to 19. H. linoides and 20. H. salvadorense, whereas from the broad-leaved form another reduction trend with an increase in flower number leads to 21. H. lorentzianum and its derivatives. The narrow- leaved form also gives rise to a many-flowered form, but with a very lax inflorescence. This variation can be accommodated in three subspecies, which are mostly easily separable. 18a. Hypericum campestre subsp. pauciflorum N. Robson, subsp. nov. Fig. 15 A, Map 20. A subsp. campestre inflorescentiis terminalibus et lateralibus unifloribus vel paucifloribus differt. Type: Brazil, Parana, mun. Quatro Barras, Rio Taquarf, 28 June 1982 (fl & fr), Oliveira 565 (BM!-holotype; MBM). P/