Introduction 61

Sect. 9. Hypericum 61

Subdivision 61

Characters and variation 61

Distribution and evolution 62

Sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum 62

Characters and variation 62

Hybrids 64

Distribution and evolution 65

Systematic treatment 66

Sect. 9. Hypericum 66

Sect. 9. Hypericum subsect. 1. Hypericum series 1. Hypericum 67

References 119

Systematic index 121

The subdivision of the large sect. 9. Hypericum and four of its segregated sections having been treated in Part 4(1), Part 4(2) is concerned with those members of sect. 9, viz. Hypericum sensu stricto, in which the stem internodes have glandiferous raised lines – subsect. Hypericum series Hypericum. The series comprises 11 species and can be divided into (i) a group centred in Europe and the Mediterranean (4 species) and (ii) one centred in north-east Asia (6 species), one species of which is confined to western North America and another has spread westward into Europe. The remaining species (H. perforatum) is morphologically and geographically intermediate between two taxa, one from each group, and behaves as an allotetraploid hybrid. It is divided into four subspecies: subspp. perforatum, songaricum (Ledeb. ex Rchb.) N. Robson stat. nov., veronense (Schrank) H. Lindb. and chinense N. Robson subsp. nov. The hybrids of H. perforatum are treated in detail and include H. × desetangsii nothosubsp. balcanicum N. Robson, nothosubsp. nov. (H. maculatum subsp. immaculatum × perforatum). Other new hybrids proposed are: H. × laschii nothoforma froelichii N. Robson, nothoforma nov. (H. maculatum subsp. obtusiusculum × tetrapterum), H. undulatum × tetrapterum and H. elegans × perforatum.

Introduction 125

Methods 127

Study site 127

Taxa studied 127

Floral phenology 127

Flower and fruit production 129

Flower visitors 129

Pollen load analysis 129

Results 129

Floral phenology 129

Flower production 131

Fruit production 131

Flower visitors 131

Pollen load analysis 133

Discussion 133

Floral phenology 133

Flower production 133

Fruit production 135

Flower visitors 135

References 136

The phenology and pollination of seven understory species of buzz-pollinated Solanaceae (Solanum erythrotrichum, S. lanceifolium, S. rudepannum, S. cordovense, S. nudum, Lycianthes hypoleuca and L. gorgonea) were investigated at the end of the dry season in the subtropical moist forest at the Las Cuevas Research Station, Chiquibul Forest Reserve, Cayo District, western Belize. Three phenological phenomena were tracked: the opening and closing of flowers, flower production and fruit production. The large short-lived white flowers of S. lanceifolium, S. rudepannum, Lycianthes hypoleuca and L. gorgonea opened around sunrise and closed at sunset. The purple flowers of S. erythrotrichum and the small white flowers of S. nudum and S. cordovense opened more or less randomly. All seven study species flowered at least once during the months of May, June and July; there was substantial overlap in the flowering of some species. Four species, S. rudepannum, S. cordovense, S. lanceifolium and S. erythrotrichum, developed mature fruit during the monitoring period while the remaining species possessed immature fruit at the termination of the study. Thus, it appeared that these seven solanaceous species would provide a fairly constant supply of mature fruit during the rainy season. During observations of pollinators, 17 different bees in the families Colletidae, Halictidae and Apidae were found to visit the buzz-pollinated flowers of Solanum and Lycianthes. Analysis of the pollen loads revealed that bees were highly constant to Solanaceae although it was not possible to determine their constancy to particular species. Very few visits were observed to S. cordovense and L. gorgonea.

Introduction 137

Carter's new taxa: types and illustrations 137

Bibliography 150

References 150

An inventory of John Carter's diatom type slides is given along with a complete bibliography. Details of the whereabouts of Carter's type slides are included along with their geography, references to sources of additional illustrations, and nomenclatural notes.

A neotype is proposed for Calymperes couguiense Besch. and the features are outlined by which the species may be distinguished from C. subintegrum Broth. and C. porrectum Mitt. Owing to its confusion with these latter species, most reports of its occurrence in Malesia are erroneous. The distribution of C. couguiense includes NE Australia and the islands of the Pacific as far east as the Tuamotu Archipelago.