African tig trees and tig wasps

Koninklijke Nederlandse Akademie van Wetenschappen Verhandelingen Afdeling Natuurkunde, Tweede Reeks, Deel 89

African fig trees and fig wasps

C.C. BERG AND J.T. WIEBES

North-Holland, AmsterdamlOxford/New YorklTokyo, 1992 C.C. BERG The Norwegian Arboretum, N-5067 Store Milde, Norway l .T . WIEBES Florijn 13, 2353 TC Leiderdorp, The Netherlands

ISBN 0-444-85741-9 Contents

I. Introduction 7 Referenee 8 11. Biology of figs and fig wasps Introduetion 9 The development of the figs 9 The habits of fig trees II The leaves l3 Stipules and buds 14 The position and size of the figs 14 The strueture of the fig 15 The flowers 17 The staminate flowers 18 The pistillate flowers 19 The pollinators 22 Colonization of the syeonia 24 Oviposition and pollination 25 Eclosion from the galls and mating of the was ps 26 Eclosion from the syeonium 28 The number of otTspring and the sex-ratio of the wasps 28 Phenology 28 The infrueteseenee and the fruitlets 30 Dispersal 31 Germination 31 Hybridization 32 The classifieation of the figs 33 The classifieation of the pollinators 36 Comparison of the classifieations of figs and wasps 37 The origin and development of the symbiosis 39 Parasitoids and predators 41 References 43 In. The genus Ficus in Afriea Introduction 47 Taxonomie history 47 Distribution 48 Diagnosis of the genus Ficus L. 51 Keys to the sections 51 Keys to the species 52 African groups and species 62 Introduced species 166 Reference 167

IV. The African fig wasps Introduction 169 Key to the genera 170 African groups and species 173 References 192

V. The poUinators (Agaoninae) Diagnosis 195 Introduced species 195 Classitication 196 Key to the genera 196 African genera and species 200 References 273

VI. Bibliography 275

VII. Index Scientitic names of animals (and Bacteria) 287 Scientitic tig names 291 I. Introduction

The present publication is the result of a long-standing co-operation between the authors, Berg studying the taxonomy of Ficus and Wiebes describing the insects that could be reared from the figs. The explicit understanding was from the beginning, that the botanica I and entomological taxonomic studies would be independent. These activities led to a number of papers on the figs and fig wasps of the African region (including Madagascar), resulting in the check-lists of Ficus (Berg, 1990c) and the Agaonidae (Wiebes & Compton, 1990). The co-operation was considerably intensified and broadened by the par­ ticipation in a french CNRS-Écotrop-programme (see e.g., Lachaise, 1984), which also involved other biologists studying other groups of fig-symbionts and their biology. High-lights of our concerted efforts (not mentioning many indi­ vidual trips, mainly by Berg) certainly were the collecting trip that the authors made into Ivory Coast (1982), together with Dr. G. Michaloud, and the sym­ posiums in Paris (1983) and Montpellier (1984). Two grants from the dutch ZWO-BION, the first one allowing Dr. S.A. UIen berg to study a genus of fig­ wasp-parasitoids, i.e., Apocrypta (UIenberg, 1982- 1985), and a second one to Dr. W. Verkerke, for a study of the fig-morphology (Verkerke, 1986-1990), substantially supported our study. Gradually, the knowledge of African figs and fig was ps attained a level at which the publication of a synopsis has become desirabIe, if only to show the state of our present knowledge to other students of the fig-symbiosis, and to all ow them to identify the figs and wasps that they find. The plan of tbis book is as follows. In four consecutive chapters the various aspects of figs and fig-wasps are treated, folio wed by a bibliography of African figs and fig wasps. The references that are not included in the bibliography, because they do not pertain to Africa, are given in a paragraph of references, after each cha pter. Our activities, and especially the participation in the Écotrop-program, pro­ moted contacts with many other students of figs and fig-insects. To all, we are greatly indebted for their collaboration and friendsbip, which we value very much: Getachew Aweke (Addis Ababa), Himansu Baijnath (Durban), Mike Bingham (Lusaka), Steve Compton (Grahamstown), Ib Friis (Copenhagen), Jacob Galil (Tel-Aviv), Veldie van Greuning (Pretoria), Finn Kjellberg (Mont­ pellier), Daniel Lachaise (Gif-sur-Yvette), Géorges (and the late Sylvie) Micha­ loud (Montpellier), Tom Müller (Harare), Anders Nilsson and Börge Pettersson

7 (Uppsala), Simon van Noort (Grahamstown), Shamilla Ramcharun (Durban), Sandrine UIenberg (Amsterdam), Géorges Valdeyron (Montpellier), Wouter Verkerke (fonnerly in Amsterdam), Anthony Watsham (Harare). The provenance of the figures, most of which were published before, is ac­ knowledged in the captions. Some of the entomological drawings were borrowed from publications by Guido Grandi and one or two by Zdenek Boucek, Jacob Galil, and Anthony Watsham, but most were made by Wiebes. For the botanical drawings, and for the kind pennission to use them in the present publication, thanks are due to E Croizier and D. Molez, Laboratoire de Phanérogamie, Muséum National d'Histoire Naturelle, Paris; EM. Bata-Gillot, E. Jens, J. Wil­ liamson and I. Zewald, Department of Plant Taxonomy and Geography, Agri­ cultural University of Wageningen; E.M. Hupkens van der EIst and T. Schipper, Institute for Systematic Botany, University of Utrecht; W. Verkerke, Hugo de Vries Laboratory, University of Amsterdam; S. Dawson, Royal Botanic Gar­ dens, Kew. S. Herland, of the Botanical Institute, University of Bergen, adapted some of the drawings to our present requisite. Dr. S. Edwards, of the Manchester Museum (U.K.), is thanked for making available the coloured plate of the cover, showing Ficus sansibarica subspecies macrosperma in full fruit.

REFERENCE

Lachaise, D. (ed.) - Programme Écotrop. Interface plantes-insectes en milieu tropical: relations entre les stratégies adaptives des figuiers et celles des insectes des figues. Courrier de I'Écotrop, Numéro spécial, 1...{j9 (avril 1984).

8 II. Biology of figs and fig wasps

Introduction Figs (Ficus L.) and the pollinating tig-insects (Agaoninae) form an unbreakable unity. It is true th at individual tig trees can grow, but for their propagation pollinators are indispensable. Outside the tig inflorescence, the pollinators live only for at most a few days. Ficus is the only genus of the tribe Ficeae, one of the tive recognized in the family Moraceae (Berg, 1989b). Because of its urceolate inflorescence, the genus Sparattosyce has been regarded as related to Ficus, but such inflorescences ap­ pear to occur also in other genera of the Moraceae and even in the Urticaceae. Both the tri bes Ficeae and Dorstenieae are characterized by a general occurrence of (basically) bisexual inflorescences. The Ficeae cannot readily be related with any other tribe of the Moraceae, which may imply that Ficus represents an early branch-off in the history of the Moraceae. The pollinators form the subfamily Agaoninae of the Hymenoptera Chalcidoidea. They have some resemblance to the Sycoecinae (see Chapter IV) which, however, is based on the very instru­ ments used for entering the tig, and thus may result from a parallel development. The wing-venation is distinctiy different. The males of the Agaoninae are wing­ less, they are winged in the Sycoecinae. Also the tig-pollinators appear to form an old group. In a recent review, Berg (1990a) treated the botanical aspects of the symbiosis of tigs and wasps; some years earlier, Wiebes (1986a) had discussed general features from the entomological point of view. Among older reviews are Janzen (1979) and Wiebes (1977a, 1979c). In 1988, a research colloquium under the auspices of the american NSF and the french CNRS, brought together in Miarni a number of botanists and entomologists interested in the tig-wasp-mutualism (multi-author review, see references under Miarni). In a nurnber of paragraphs, some morphological and biological features interesting to our purpose, will be discussed below, to begin with the two peculiar features of tigs, viz., the develop­ ment of the tigs and the pollination of their flowers by tig wasps, and the hemi­ epiphytic life-habit of many of the tig-species.

The development of the figs The fruit-like inflorescence of Ficus - tig or syconium - consists of a more or less globular (or urn-shaped) hollow receptacle, fully enclosing the flowers borne

9 --'II,..,JJc:---- 0 S T lOL E

b c

Fig. I. Anatomy of a fig-sycone, and of staminate (a) and pistillate flowers (b, short-, and c, long­ styled flower). After Wiebes (1982a, fig. I). on the inner side (fig. I). There are three main types of flowers, viz., staminate flowers (fig. Ia) and pistillate flowers, divided into long-styled flowers ('seed­ flowers', fig. Ic) and short-styled flowers ('gall-flowers', fig. I b). If the two types of pistillate flowers, together with the staminate flowers, occur in one and the same syconium (as in the monoecious species), intermedia te style-Iengths are found and all pistillate flowers can produce seed (Neeman & Galil, 1978). If the two types of pistillate flowers occur in different figs (as in the gynodioecious species), intermediate style-Iengths are not found and usuallly only the long­ styled flowers produce seed. In gynodioecious species the short-styled flowers occur together with staminate flowers in 'gall figs' , while figs containing the long-styled flowers ('seed figs') lack staminate flowers. The two types of fig are found on different trees. A survey of features connected with monoecy and gyno-dioecy in Ficus is presented in table I. Most of the African species of Ficus are monoecious, only Ficus pa/mala and the species of section Sycidium (together 10 out of 105 African species) are gynodioecious. In the development of the syconium Galil & Eisikowitch (l968c) distinguished between a number of phases, as follows (fig. 2). A. Pre-female phase, during which the inflorescence and the flowers develop until the receptiveness of the stigmas of the pistillate flowers. B. Female phase, in which the ovules become fully developed, the stigmas are receptive, and the ostiole becomes pene tra bie for the pollinators: they enter, pollinate and oviposit. C. Interfloral phase, in which the larvae of the pollinators develop to imagines and the fig-seeds develop to maturity. This ph ase separates the anthesis of the pistillate flowers from that of the staminate flowers. D. Male phase, in which the insects emerge from their galls. The pollen was already mature in an earlier (i .e. , the female) phase, but now it becomes exposed by the elongation of the filament andlor the opening of the thecae. The male

10 WASP ~ 11 F>oviposition-Iarva-d' x ~ phase A- r-phase B-phase C-phase D-phase E FIG sycone (pre-female) (female) (inter-floral) (male) (post-floral> !:;.pollination seed t I

WASP ~ 11 ~oviposition-Iarva-d' x;

GALL fopollination -t SYCONE phase A- f-phase B - phase C-phase 0 (pre-female) (female) (inter-floral) (male) FFIG SEED phase A- r-phase B phase E SYCONE (pre-female) (female) ( post-floral) .pollination seed J

Fig. 2a (above). Generalized cycle of a monoecious tig and its pollinator wasp. The developmental phases of the tig-sycone (A - E) are indicated in the lower quadrangIe, those of the wasp in the upper. Fig. 2b (below). Generalized cycle of a gynodioecious tig and its pollinator wasp. The developmental phases of the tig-sycone containing staminate and gall-flowers ('capriticus') are indicated in the middle, those of the syconium with only pistillate seed-f1owers ('domestica') in the lower part, the development of the wasp in the upper. After Wiebes (1979c, figs . 1- 2).

pollinators mate with the females while these are still in their galls, then also the females emerge, collect pollen and leave the syconium through the ostiole or via tunnels made by the males. E. Post-floral phase, in which the fruitlets become mature and the syconium becomes soft and attractive to organisms that disperse the fruits.

The habit of fig trees

With regard to habit, growth- and life-forrn, Ficus is one of the most diverse genera, especially so in Asia and Australasia, where it comprises hemi-epiphytes (many of them being potential 'strangiers' and some exhibiting the 'banyan'­ habit), holo-epiphytes, climbers (among which a group of root-climbers), creep­ ing shrubs, rheophytes, etc. (Corner, 1952, 1967, 1976; van Steenis, 1981). The diversity is less in Africa and it is much less in the Neotropics. A variety of architectural models are represented (Caraglio, 1985). About 50% of the tig-species are (potentially) hemi-epiphytic and about 50% have a terrestrial life-fofffi. In Mrica, hemi-epiphytism is restricted to the sub­ genus Urostigma, where it is exhibited in most species. Hemi-epiphytes start their life as epiphytes at some site suitable for settlement on a host tree, such as

II crevices in the bark, a rotten stump of a broken branch, or in forks of branches (see Michaloud & Michaloud-Pelletier, 1987). Saplings of hemi-epiphytic species have a swollen base of the stem and main root ('Iignotuber'), which may be of importance for anchoring the plant. It usually sends down a ('tap')root along the trunk of the host. Until this reaches the soil, giving the plant access to more nutrients, allowing a more rapid growth, the hemi-epiphyte grows slowly and normally does not reproduce. The root-system expands and by anastomosing may form a root-basket around the trunk ofthe host, and a dense root-mat undemeath the host-tree. By reducing the nutrient-supply, shading its crown and probably also by cutting ofT the sap-streams, a hemi-epiphytic tig may overpower the host-tree, 'strangling' it by the root-basket around the trunk. In more or less open vegetation, like savanna wood land and riverine forest,

Table I. A survey of features connected with monoecy and (gyno)dioecy in Ficus.

Monoecious Dioecious

Inllorescences with: staminate Il. >40-48% a. gall ligs: short-styled Il. 40-48% staminate Il. 3-35% long-styled 11 3-> 10% short-styled Il. 95-65% b. seed ligs: long-styled Il. 1000/0 or with neuter Il.

Heterostyly: imperfect perfect

Ovaries at anthesis: not lined up lined up

Ovules: all can form seeds a. short-styled Il.: do not (usually) form seeds b. long-styled Il.: all can form seeds

Flowering: mostly (excl. Sycomorus) a. gall lig trees: individuals synchronous individuals often populations asynchronous asynchronous, tig crops not separate, or ± synchronous b. seed tig trees: (probably) often synchronous with separate (seasonal?) lig crops

Represented in: ca. 50010 of the species ca. 500/0 of the species In Old and New World: In Old World: 'Pharmacosycea group' 'Ficus group' 'Urosligma group' 'Sycidium group' 'Sycomorus group' p.p. 'Sycomorus group' p.p.

12 hemi-epiphytes, settling not far from the soil, soon form trees of considerable size and 'kiB' their host. The 'secondary' trunk formed by the cylinder of anasto­ mosed roots, often is strong enough to bear the (true) trunk and branches after the host has died and its trunk become rotten, thus replacing the host tree. In high forest, however, where the hemi-epiphytes may settle far from the soil, it takes much more time for overpowering the host-tree and the cylinder of roots around the host-trunk often is too weak to prevent it from falling down with the rotten trunk of the host. Only some very vigorous species, like Ficus elas­ ticoides and F. cyathistipuloides, can form a broad root-infrastructure (often achieved by sending down roots along neighbouring trees), sufficiently strong to replace the host tree. Hemi-epiphytic species are often found on rock surfaces (or on waUs of build­ ings), where they settle in crevices and send down tap-roots over the rock. Some species, like F. abutilifolia and F. tettensis, are mostly or even only found on rocks: they are called hemi-epilithic, and may split the rocks. On plants hosts, hemi-epiphytic tigs may occasionally split trunks, like the Asian F. religiosa L. often does: it can be indicated as a 'tree-splitter' (Galil, 1983). In some hemi­ epiphytic tigs one can see aerial roots hanging from branches. They may reach the soil (or water surface), but rarely form 'pillar-roots'. The typical 'banyan'­ habit, as found e.g., in the Asian F. benghalensis L. , is not c1early exhibited in Africa. Some representatives of the subgenus Urostigma are usually or always ter­ restrial, like F. platyphylla, with (fire-resistant) trees in savanna woodland, F. verruculosa, with shrubs or treelets along streamlets, and F. trichopoda, with large shrubs or small trees with stilt- and/or pillar-roots, in marshy places. A few species are more or less lianescent, with a slender trunk and a narrow crown, as in (some forms of) F. ottoniifolia and in F. lingua, if occurring terrestrially in forest (or scrub). Most Urostigma-species, as also those of the subdivisions Sycomorus (not F. mucuso), Sycidium, Oreosycea (not F. variifolia) and Ficus, which are terrestrial, have short (true) trunks and broad crowns. Ficus mucuso and F. variifolia are forest trees and they have taU trunks. Most African tig species are evergreen, but some are deciduous (e.g., F. ca­ lyptrata and F. sycomorus).

The leaves The leaves vary in size from a few centimeters (e.g., in F. humbertii and F. lingua) to half a meter long (e.g., in F. wildemaniana). They are mostly coriaceous, with an entire margin, in Urostigma, but mostly chartaceous to subcoriaceous, often with a dentate to crenate margin, in the other groups. In some species, of (sec­ tion) Pharmacosycea and Sycidium, the leaves are pinnatitid, especially when juvenile. The lamina is glabrous to more or less densely hairy, but this (as weil as the hairiness of leafy twigs and tigs) can vary considerably within species. In Sycidium the upper Ieaf-surface is often scabrous.

13 In most species the lamina is distinctly longer than wide, but in some it is cordi- or reniform, and about as long as wide. Leaves with a cordiform, (broad­ ly) ovate or reniform lamina usually have relatively long petioles. This combi­ nation of characters, as weil as hairy surfaces, a chartaceous texture and a den­ tate margin, are usually associated with more or less open and dry habitats. On the lower surface of the lamina one or more waxy (glandular) spots occur: in sub genus Urostigma one at the middle of the base of the midrib, in subgenus Pharmacosycea often two (± lateral) at the base of the midrib. In the other groups the glands occur in the axils of the (main) basal lateral veins, or some­ times also in the axils of other lateral veins, or in the main furcations of lateral veins. In section Sycocarpus (Asia-Australasia) waxy glandular spots also occur at the nodes of the leafy twigs. The function of these glandular spots is not known.

Stipules and buds The stipules are often fully amplexicaul, and then leave an annular scar. They are mostly free, but in some species of subsection Cyathistipulae they are partly fused. In this subsection the stipules are often coriaceous and subpersistent. In some species pairs of young figs are enclosed by a calyptrate bud cover, in texture and indument similar to stipules, and probably representing enlarged and fused prophylls of the (usuaUy abortive) axillary shoot bearing the two syconia. These calyptrate buds can be some centimeters long, like in F amadien­ sis, F calyptrata, and F ovata. The bud covers, e.g. in F craterostoma, may open with lateral slits; the adaxial part can be subpersistent.

The position and size of the figs The figs are often borne in the leaf-axils, mostly in pairs, less commonly solitary. In some species, e.g., Ficus adolfi-friderici, small short-shoots ('spurs') develop in the leaf-axils, and they can be ar more than two figs at the same time and continue to produce syconia on older wood. In some species of subsection Cau­ locarpae, spurs al ready formed in the leaf-axils may function for a long time, producing the successive crops, or the spurs initiated in the leaf-axils start to develop only on the older wood, on the main branches or the trunk. These spurs are leaf-less and become mostly peg-shaped (up to ten cm long in F sansibarica subsp. macrosperma), or less commonly become more or Ie ss cushion-shaped (as the result of branching of the short-shoot). In some species (e.g., Ficus calyp­ trata), the (paired or solitary) figs are normally (or always?) borne on the pre­ vious season's growth, which is apparently related to the seasonal growth of the branches, in combination with deciduousness. In several representatives of Sycomorus and Sycidium, the figs are also borne on the older wood (in some species down to the base of the trunk) on leaf-less branchlets, which can be branched; they have long internodes, and can become much longer than ten cm. In the geocarpic Ficus vogeliana these branchlets are

14 all, or for the greater part, formed at the base of the trunk, and they become stolon-like, many meters long, bearing the syconia at the soil surface or in the litter. The production of figs helow the leaf-axils (ramiflory, cauliflory and geo­ carpy) give Ficus the possibility to increase the number of syconia per crop, and to make their production independent of the seasonal growth of the terminal branches (see Berg, 1990a). It also gives species with slender leafy twigs the possibility to produce large syconia. Some species (e.g., F lingua) have figs less than 0.5 cm in diameter (containing some tens of flowers), but forms of F sansibarica and F sycomorus have figs more than 5 cm (up to 10 cm) in diameter (with some thousands offlowers). The majority of the Mrican fig species have figs with sizes in between, with a distinct peak in the I cm diameter c1ass (Berg, 1990a). Two sizes of sycones are rather well represented, viz. , those of about I cm and those of 2.5-3 cm in diameter. There is a positive correlation between the size of the sycones borne in leafaxils, with the diameter of the leaf twigs. Species with slender leafy twigs may have large sycones, but these are borne on the older wood. In a species, figs may considerably vary in size and (consequently) in the number of flowers per fig. If the figs are borne in the leafaxils, there appears to exist a positive correlation between the size and the diameter of the leaf twig. Species with slender leafy twigs may have large figs, but these are borne on the older wood. The size of the figs is of importance for various aspects of the pollination system, as it is related to the number of pistillate flowers, the number of (poten­ tiai) breeding sites for the wasps, and the number of fig wasps needed to occupy these sites: one pollinator may be sufficient, or more than one are needed to oviposit in all or most short-styled flowers of one syconium. The position and size of the syconia can also be regarded as adaptations to dispersal, as they may increase the range of animals that can he involved.

The structure of the fig In most groups the receptacle is subtended by a whorl of three or two 'basal bracts' , which may he small or large, persistent or sometimes caducous. In Sy­ cidium these basal bracts are usually lacking: a number of bracts occur scattered on the peduncle ('peduncular bracts'), in addition to a caducous bract at its base. Sometimes these bracts occur on the outer surface of the receptacle ('lateral bracts'). The margin of the receptacle bears numerous, more or less tightly packed bracts ('ostiolar bracts'), as a barrier in the entrance of the fig cavity. In the subgenera Pharmacosycea and Urostigma, bracts occur among the flowers ('in­ terftoral bracts'), but they are lacking in the other groups - there often are (bristle-Iike) hairs instead. In Sycomorus, however, a pair of 'bracteoles' are subtending and often enclosing the staminate flowers. The diameter of the ostiole varies, as weil as the number of (visible) uppermost ostiolar bracts: two, three, or more. In most groups all, or at least the upper ostiolar bracts are interlocking, from more or less tightly to more or less loosely.

15 In several the lower ostiolar bracts are descending, in the subgenus Sycomorus (fig. 3e, f) eventually forrning a plug pressing on the developing (pistillate) flo­ wers (Verkerke, I 988a). In the section Galoglychia (fig. 3g, h) all ostiolar bracts are descending. The two outermost are large, and they leave between them a narrow, bilobate, slit-shaped opening towards the interior, but three bracts oc­ casionally occur, forming a triradiate slit. In most cases the braets are so tight that pollinators can only enter with diffieulty, and that escape is not possible. In the section Sycidium (fig. 3e, d), however, the ostiolar braets are more looseiy interlocked, at least in the female and male phases, and fig wasps may escape without tunneiing: this appears to represent a eomparatively primitive state. The figs are peduneulate or sessile. In some species (e.g., F. cyathistipuloides

Fig. 3. Ostioles of various subdivisions of Ficus. a-b, sections Ficus; c--d, section Sycidium; e- f, subgenus Sycomorus; g- h, section Galoglyclzia; i- k, subgenus and section Pharmacosycea; I, subgenus Urostigma, section Americana. a-j in female phase, k-I, in post-floral phase, I with tunnels. The bar indicates ca. 0.5 mmoAfter Verkerke (1989, fig. 2).

16 and F densistipulata) the receptacle is narrowed above the basal bracts into a 'stipe'. The figs, therefore, may be both pedunculate and stipitate. In Sycomorus the cavity of the figs often (or usually?) contains a (slightly mucilaginous) fluid during (part of) the interftoral phase. The receptacle - the wall of the syconium - varies considerably in thickness, from less than one to more than one cm, often in relation to its size. The texture varies from solid and firm, to sometimes spongy (F cyathistipula). At full ma­ turity the wall can become soft and free of latex, and the inner layers may become mucilaginous, and the wall is often coloured: yellow, orange, pink, red, or purplish. In many species the wall is spotted. In gynodioecious species 'gall figs' do not show the features of the mature 'seed figs', in which the wall becomes fleshy and coloured. Ripe figs do not usually have pronounced smells, but those of F carica L. do and also those of the subgenus Sycomorus. Ripe figs of F calyptrata smell after rotten fish. The functional anatomy of the syconium was treated by Verkerke (1989).

Theflowers Fig-flowers are unisexual, at least functionally, but there is an Asian subgroup of Sycidium in which the flowers are morphologically bisexual, with a short­ styled pistil in which the larva of the pollinator may deveIop, but it does not produce seed (see below). In some groups pi still odes occur in staminate flowers, but even in groups where they are usually lacking, bisexual flowers occasionally occur. In 'seed figs' of gynodioecious species one can find 'neuter flowers', as substitutes of staminate flowers, occurring fully developed in 'gall figs' . The flowers are small, one to some millimeters long - their size generally is positively correlated with the size of the syconium. They are either sessile or pedicellate. There are 2- 5(- 7) free or connate tepals, in the last-mentioned situa­ tion forming a tubular, lobate or dentate perianth. The flowers and floral parts are mostly glabrous, with the notabie exception of the section Sycidium, usually bearing short bristles on the pedicel, the (margin of the) tepals and on the style. The flowers are packed together, more tightly in the monoecious than in the gynodioecious species. The development of the syconia in the pre-floral phase is shown in fig. 4. The number of flowers per syconium increases with its size, from some tens of flowers in a sycone of up to 0.5 cm in diameter, 300-400 in a sycone of about 1 cm, about 2500 in a sycone of about 2.5-3 cm, to several thousands in larger sycones (different species of tig are being compared). Also the size of the flowers is often increasing with the size of the sycone, less so in gynodioecious than in monoecious taxa, which may be explained by a difference in the arrangement of the ovaries in the female phase: in one or two layers, respectiveIy. Quantitative data per tig are very important. Compton & Nefdt (1990) gave them for Ficus burtt-davyi and its pollinator Elisabethieila baijnathi, as follows. An average of about 275 female flowers and 14 male flowers (with an average

17 Fig. 4. Syconia of various subdivisions of Ficus. a--<:, section Galoglychia; d-f, subgenus Sycomorus; g- i, section Sycidiwn; j-I, section Ficus. The bar indicates ca. 0.2 mm. After Verkerke (1989, fig. 1).

of 5.403 pollen grains per anther) were present per sycone (which measure about 0.4-1.2 cm in diameter). Infertile flowers were the most abundant category, 50.2%; bladders 21.4%; galls containing insects 18.6% (7.3%, pollinators); fio­ wers containing healthy seeds 10%. In the figs of the section Oreosycea and subgenus Urostigma an average of about 10% of the flowers are staminate (cf. Hili, 1967b). In the subgenus Sy­ comorus the percentage is lower, i.e. up to 5%, but the flowers may bear two stamens and the anthers are rather large. In the sections Ficus and Sycidium the percentage of staminate flowers in the 'gall figs' often is quite variabie.

The staminate fiowers The staminate flowers are either dispersed among the pistillate flowers (fig. 5), or they are arranged in one or several rows near the ostiole (fig. 6a). They have

18 Fig. 5. Staminate and pistillate ftowers of Ficus ottoniifolia (Miq.) Miq. a, in the fernale fase of syconiurn developrnent; b, in the late interftoral phase (s = fruit with seed, exocarp rernoved; g = 'gall fruit'. After Verkerke (1986, figs. 6a and toa, resectively). one, two, or sometimes three stamens. The anthers are in most species ca. 0.5-1 mm long, in species of subgenus Sycomorus mostly 1- 1.5 mm long. The staminate ftowers of species of subgenus Sycomorus are distinct from those of the other subdivisions. The two (or three) stamens are packed in the tubular perianth with imbricate lobes. The stamens become exposed by elonga­ tion of the filaments, by which the upper part of the perianth is tom open or off (fig. 7). The flowers are initially entirely or partly enveloped by two bracte­ oles. Staminate ftowers occurring dispersed among the pistillate flowers (as in the subgenus Urostigma) are more or less hidden by these flowers and undemeath the synstigma. The anthers usually become exposed by e1ongation of the fila­ ments and/or the pedicels (figs. 7 and 8). In Urostigma the thecae usually open with narrow slits; fig wasps collecting pollen may even need to widen the slit (Baijnath & Ramcharun, 1988).

The pistillate flowers The pistillate flowers show differences in the leng th of the style (heterostyly) and often also in shape and dimensions of the ovaries (Bronstein, 1988; Compton & Nefdt, 1990). The stigmas are variously shaped. There are two filiform to subulate stigmas in the subgenus Pharmacosycea, in F palmata, and sometimes in the section Galoglychia, but the majority of the African species have a single stigma, being more or less filiform in the subgenus Urostigma, (sub)infun­ dibuliform in the section Sycidium (fig. 9) and (sub)infundibiliform to subclavate to tongue-shaped in the subgenus Sycomorus (fig. 6b). The stigmas of the flowers of a syconium are always lined up at anthesis of the pistillate flowers: they form a more or less continuous layer of stigmas touch­ ing each other, adnate, or cohering and then forrning a distinct 'synstigma', at

19 Fig. 6. Flowers of Ficus sur Forssk. a, staminate flowers (enveloped by bracteoles); b, pistillate flowers at anthesis; c, pistillate flowers at fruit (s = fruit with seed; g = 'gall frui!'). After Verkerke (1988c, figs. 6f, 7b, and 9, respectively).

20 Fig. 7. StamÎnate flowers of Ficus sur Forssk. a, flower enveloped by bracteoles; b, stamen; c, after elongation of the filaments. After Verkerke (l988c, fig. 6). a certain di stance from the wall of the syconium. This is realized by the presence of sessile flowers andJor flowers with different pedicel-Iengths, often in combina­ tion with difTerences in shape and length of the ovaries: relativeJy short and ovoid or oblongoid to obovoid, with or without a stipitate base. At anthesis then, the ovaries are arranged in at least two layers (figs. 5 and 6). In monoecious species, heterostyly can be said to he imperfect. The difTerentiation of the flowers probably is realized ontogenetically, during their deveJopment (Verkerke, I 988c). In monoecious species all pistillate flowers can produce seed, which sometimes occur in nature if the pollinators fail to oviposit, as was demonstrated in experiments by Neeman & Galil (1978). On the other hand, not all ovaries can he occupied by larvae of the pollinator, as it cannot reach all ovules: the ovipositor may be too short for the long-styled flowers andJor the long styles may he too slender, too flexible or too much curved. There may he also other reasons, such as the numher of eggs carried by the pollinator, the number or species of pollinator, etc. (Bronstein, 1988; Compton & Nefdt, 1990; Michaloud 1988b). Normally, up to 10% of the pistillate flowers do not deveJop fruits, and in general, about 50% of the deveJoping fruits contain seed, the remainder insect larvae. In gynodioecious species the ovaries are situated at about the same distance from the syconium wall: they are uniseriate at anthesis (fig. 9a), with all stigmas forming a more or less continuous (not always clearly synstigmatic) layer. Mter anthesis, difTerential eJongation of the pedicels does occur and the developing fruits hecome arranged in some layers (fig. 9c). Alllong-styled flowers can pro­ duce seed. Verkerke (l987b) found in F. asperifolia, that after pollination (and subsequent fertilization) the development of the embryo as weil as endosperm was initiated, although the embryo soon aborted and the endosperm tissue con­ tinued to proliferate only if the ovule was occupied by an wasp-egg (or a larva). Therefore, one can say that F. asperifolia is functionally dioecious, although

21 Fig. 8. Development of staminate fiowers in Ficus ottoniifolia (Miq.) Miq. After Verkerke (1986, fig . Ig). morphologically gynodioecious, and th is likely is similar in ot her gynodioecious taxa (cf. Condit, 1932). In gynodioecious species, heterostyly can be said to be perfect; it is determined genetically (Valdeyron & Lloyd, 1979). Heterostyly may lead to loss of the fe­ male function in short-styled morphs (Beach & Bawa, 1980; Ganders, 1979). In Ficus the process of reduction to fully non-functional ovules and ovaries appar­ ently is arrested at a stage, in which at least endosperm-production can still be initiated by fertilization, which allows occasional seed-production by short-sty­ led flowers, as in F carica. Further reduction beyond this stage (cf. Beck & Lord, 1988a, b) would terminate the pollination system. Stimulative parthenogenetic development of some ovular tissue (nucellus or endosperm?), which allows insects to breed without preceding pollination, has been found in some species (F carica, F microcarpa, F sycomorus; see Con dit, 1932; Galil, 1990; Storey. 1975). This prevents abortion of the syconia, which normally drop if there is no development of the fruitlets.

The pollinators The two sexes of the pollinators are different in body-form. The female fore and hind legs, especially the tibiae, are short, strong and spiny, and help push the head forward when the female enters the ostiole, whilst the mandibles pull it.

22 Fig. 9. Long-styled flowers of Ficus asperifolia Miq. a, at anthesis; b, styles and stigmas; c, after anthesis. After Verkerke (l987b, figs . Sa, b, and 6a, respectively).

The mandibles have mostly strong apical teeth and ventral lamellae, and they be ar an appendage provided with ventral rows of small teeth, crenulations or lamellae, probably preventing the head from slipping back.

23 Also the male has strong fore and hind legs; the mid legs usually are slender, or they may be rudimentary. The mandibles mostly are rather strong, and they are used (not necessarily in all groups!) for biting an opening in the wall of the gall fruit containing a female, and for tunneling the wall of the receptacle. The segments of the thorax, especially the dorsal parts, often are for the greater part fused, which may make them rather robust. The males are wing-less and often blind; the gaster is long, shaped like a tube, enabling the mating with the female still enclosed in its gall.

Colonization of the syconia The female fig wasp approaches a fig with sycones in the female phase, probably attracted by volatile compounds ernitted by the fig-leaves andJor -syconia, e.g., F. ingens attracts its pollinator Platyscapa soraria by using odours (Barker, 1985; van Noort, Ware & Compton, 1989). Possibly, one should distinguish between long-distance pollinator-attraction and their short-di stance guidance towards the syconium and its ostiole, but about this little is known. For the Australian fig species Ficus macrophy/la Desf. ex Vent., Pemberton (1921 : 305-306) described how the female of Pleistodontes froggatti Mayr en-

Fig. 10. Ventral view of the thorax of a female fig-wasp (Platyscapa etiennei Wiebes), showing the right fore leg (bent at the knee) and the left poUen pocket. The left leg is taken off at the coxa. After Wiebes (1981, pI. 11 fig. 5).

24 counters considerable difficulties in gaining entrance in the syconium. Several never succeed and may die in the attempt. The head is pushed in the ostiole, the mandibles are fixed into the tissue in front, and the whole body is drawn for­ ward. The action of the mandibles and their appendages draws the body further in. The antennal hooks, i.e., the elongated appendages of the third segment (not obvious in all groups) may be used as a sort of crowbar for prying apart the ostiolar scales, and the spines (if present) on the axial surface of the pedicel, directed backwards, may help prevent the head from slipping back. After the head has passed into the open interior, the legs are brought into use and are the sole means of pushing the body completely through the ostiole. The difficulties encountered usually result in the loss of the wings and the outer antennal seg­ ments; also, a quantity of fluid is discharged from the abdomen. In other groups (e.g. , Sycidium) the passage through the ostiole may he less difficult. In Ficus burtt-davyi, a mean of 1.62 pollinators entered each sycone, the ma­ jority receiving only a single pollinator, carrying a mean of about 314 pollen grains, which approxirnated to just over one pollen grain for every female flower in the sycone (Compton & Nefdt, 1990). A minimum of two pollen grains per stigma seem to he normally required to achieve areasonabie seed-set (Cruden, 1977). An important advantage of the synstigma may he that a pollen grain posited on a certain stigma, may form a tube entering the style of another flower. Also in F ottoniifolia, the sycones were usually colonized by a single female of Courtella gabonensis (andlor several of C. camerunensis; Michaloud et al. , 1985: lOl). When a female finally has reached the interior, it immediately begins egg­ laying and pollination.

Oviposition and pollination For a long time it remained a enigma how the pollen was introduced into the fig sycone by the wasp that evidently pollinated it: seed-set required its presence, but it was found to carry at most only a few pollen grains on its almost glabrous body. Then Galil & Eisikowitch (1969a) discovered the thoracal pollen pockets (fig. 10) and Ramirez (1969) added the description of the coxal corbiculae, pol­ len-holding cavities located one on each coxa of the front legs. Later, many more, refined observations were made on various groups of figs and wasps, inc\uding those by Galil & Neeman (1977) and Okamoto & Tashiro (1981), who described two different modes of passive transport, ensuing from the shrinking after leaving the gall, and partial swelling of the wasp's body during oviposition (attempts) in the young, colonized sycone. The act of pollination is so integrated in the oviposition process that it is not possible to understand the one without the other (Galil & Eisikowitch, 1969a: 6, from wh om also the following description of the behaviour of Ceratosolen arabicus in Ficus sycomorus is borrowed). The wasp stands on the synstigma and pro hes with her ovipositor. When an appropriate site is located, the ovipositor

25 fable 2. Figs, the pollination of which was described.

Ficus Pollinator Authors carica B. psenes Galil & Neeman (1977) erecta B. nipponica Okamoto & Tashiroi (1981) asperifolia K. ges/ra i Verkerke (1987b) fistulosa C. hewilli Galil (1973b) racernasa C. fusciceps Joseph & Abdurahiman (1981) sycomorus C. arabicus* Galil & Eisikowitch (1969a) sur C. capensis * Baijnath & Ramcharun (1983) costaricana Pe. estlzerae Galil et al. (1974) Iz emsleyana Pe. tonduzi Galil et al. (1974) religiosa P quadraticeps Galil & Snitzer-Pasternak macrophy /la PI. froggatti Pemberton (1921) (1970) lwea A. hel.-morplzum Newton & Lomo (1979) burtt-davyi E. baijna/hi Baijnath & Ramcharun (1988) ottoniifolia Cou. gabonensis* Michaloud et al. (1985)

* other Agaonids were recorded from the sycones as weil, viz. , sycomorus C. galili c\eptoparasite sur C. flabella/us pollinator C. silvestrianus pollinator ottoniifolia COU. camerunensis pollinator

moves downwards along the style. Then the wasp energetically bites the stigmata within reach. A few seconds before the withdrawal of the ovipositor, the fore legs of the wasp fold simultaneously or alternately until the tarsi and pulvilli touch the lower margins of the pollen pockets (fig. 11). Now the wasp lowers its legs and moves them delicately to and fro, caressing the stigmatic surface below. In other species (see table 2) such pollination movements may be differ­ ent, e.g., striking the tarsi on each other as e.g. , in Pegoscapus estherae (Grandi) in the New World Ficus costaricana (Liebm.) Miq. (Galil et al. , 1974). The mean duration of oviposition and pollination was 31.31 seconds for Cour­ tella gabonensis in Ficus ottoniifolia (Michaloud et al., 1985: 103), but twice as long for C. camerunensis (69.22 seconds). The pollen tube grows through the stylar canal and through the micropyle into the embryo-sac, and the embryo develops and endosperm is formed. The egg is deposited in the fig-ovule, outside the nucellus and beneath the insertion point of the inner integument at the raphal si de (Verkerke, 1988c). The development of the wasp-Iarva and the fig-embryo takes place in what was called the inter­ floral phase (C) of the sycone.

EcIosion from the gaUs, and mating of the wasps The male wasps come out of their galls first, and immediately go in search for ga lis containing females in which they gnaw a hole, enter their tubular abdomen

26 Fig. 11. Ovipositing female of Ceralosolen arabicus Mayr at the pollination act. After GaW & Eisi­ kowitch (1969a, fig . 16).

and copulate with (most of) the females inside (Godfray, 1988 estimated that, for the Papuan Ceratosolen dentifer Wiebes, 2% of the females leaving the figs of Ficus hispidioides S. Moore were unmated). Michaloud et al. (1985: 104) gave the duration of this behaviour of the wasps in Ficus ottoniifolia, viz., 14.5 minu­ tes (s.d. 3.8) spent on opening the gall and copulation for CourtelIa camerunensis, 9.6 minutes (s.d. 2.79) for C. gabonensis; the copulation took 37 and 47 seconds, respectively. A number of males copulated again with mated females; the num­ ber of copulations per male could vary from 10 to 15. After each copulation, the male (and/or the female) widens the hole in the gall through which the female will emerge. In species of Alfonsiella, the males look rather fierce. Instead of fighting, however (although they may push each other), they use their large, falcate mandibles to gently remove the female antennal envelope, before she leaves the gall. Then, the females load pollen. Galil (1973) distinguished between the active ('ethodynamic') and passive ('topocentric') transfer of pollen. In the active mode the female approaches the anthers and, if necessary, with her mandibles widens

27 the opening slit, and starts to lift the pollen onto the ventral surface of the thorax, shovels it backwards with the fore coxal combs of setae, and pushes it into the pockets (Ceratosolen arabicus in Ficus sycomorus, Galil & Eisikowitch, 1974: 524).

Eclosion from the syconiom The ostiolar scales may loosen slightly towards the end of the male phase, and the female wasps crawl out of the syconium through the slit thus formed (e.g., Platyscapa quadraticeps in Ficus religiosa, Galil & Eisikowitch, 1968c: 359), or the males may tunnel through or near the ostiole, and the females fly out (e.g., Ceratosolen capensis in Ficus sur, Baijnath & Ramcharun, 1983: 885), or else­ where (e.g., Pleistodontes froggatti in Ficus macrophylla, Pemberton, 1921: 309 and tig. 7; Elisabethiella stuckenbergi in F thonningii, Boueek et al., 1981: 154, tig. 6E).

The nomber of ofIspring and the sex-ratio of the fig wasps In Ficus ottoniifolia, the average offspring in sycones colonized by one female was found to be ca. 250 for CourtelIa gabonensis and ca. 200 for C. camerunensis; for both, the sex-ratio was ca. 0.08 (Michaloud et al., 1985: 104-105). In the New World examples mentioned by Herre (1989, tigs. 2, 3), the mean number of offspring per (single) foundress was about 150 for a large tig with many flowers (± 1100) pollinated by a relatively sm all wasp, as also for a small sycone with relatively few flowers (± 350) pollinated by a relatively small wasp, while it was about 350 for a large sycone with many flowers (± 975) pollinated by a large wasp. As the number of foundress fema1es per sycone increased, the number of their offspring dropped, that of the seeds tended to rise, and also the sex-ratio 1 increased (approximately by a factor 2 for two foundresses, 2 / 2 for three, 2% for four, etc.). Herre (1989: 646) conc1uded that, although this works against the reproductive interest of the tig, generally, when comparing natural numbers and trying to assess the reproductive output of tigs and wasps, the tig seems to be the 'controling factor' in the mutualism. Hamilton (1967) had been the tirst to show that the number of founding females is positively correlated with the frequency of males produced in a sy­ conium, after he had found the sex-ratio biases to depend on pattems of male competition for access to mates (see Frank, 1989, for a recent review). The explanation lies in the fact th at riyal Agaonine males in one sycone are likely to be brothers and th us they are not expected to tight. Other relevant papers are e.g., those by Frank (1985) and Herre (1985).

Phenology Fig phenology is described as synchronous within each crown of an individual and asynchronous among individuals (Frank, 1989: 676). Wharton et al. (1980)

28 described asynchrony in a population of Ficus sycomorus in South West Mrica, in which any stage of the reproductive cycle could be found throughout the year. Very important in this respect is the longevity of the polJinator, which, according to Kjellberg et al. (1988), for Blastophaga psenes in Southem France is 48 hours

o4

7 8 _-, , r" 1 , , \ / 0\~ ,.. , ,/ 12 9 : 1

14 15

Fig. 12. Schematic drawings of fruit types. 1--4, dehiscent drupe1et as occurs in many Moraeeae: I, longitudinal section; 2, cross section; 3, empty exocarp; 4, endocarp body (pyrene). 5-8, section Galoglychia-fruit type I: 5, longitidinal section; 6, empty exocarp; 7, endocarp body surrounded by mucilaginous layer; 8, endocarp body with 'pseudohilum'. 9-12, section Galoglychia-fruit type 2 : 9, longitudinal section; 10-11, empty exocarps; 12, endocarp body with a mucilaginous layer. 13, section Galoglychia- fruit type 3: 13, indehiscent fruit with the upper part surrounded by a muci­ laginous layer. 14-15, section Galoglychia- fruit type 4: indehiscent dry fruit with mucilage (of eeUs of the syconium wall) at the base; 15, indehiscent dry fruit. After Berg (1984a: 33).

29 (in the wild). Grandi (1929: 107) had kept several individuals of B. psenes alive in the laboratory for 4-5 days, Joseph (1958: 209) for 2-3 days. The various levels at which one can observe synchrony of life-cycles were discussed by Addicott et al. (1990: 151-155). They distinguished between syn­ chrony within individual syconia (rather strict, because of the obligate associa­ tion), synchrony among syconia within a tree (to some extent, density-depend­ ent), and that among trees within a population (see below). Most of the monoecious Ficus-species in Africa, i.e., the representatives of Urostigma and Oreosycea (for Sycomorus, see above) show synchronous and intermittent flowering of individuals, but asynchronous and continuous flower­ ing in populations. Unfavorable (seasonal) growth-conditions may influence flo­ wering, by reducing the number and size of fig crops and by slowing down the development of the sycones. In the gynodioecious species, gall-sycones are produced throughout the year, continuously or in overlapping crops; seed-sycones are usually produced only once a year. Outside Africa, seed-fig-production can occur less often (cf. Med­ way, 1972) or in a pattern similar to th at of gall-figs (cf. Corlett, 1987). In Panama, Windsor et al. (1989) followed fruit- and leaf-iniation by 26 trees representing five Urostigma species, for 5--8 years. There proved to be a high variation, indicating th at the species initiate fruit-crops the year around, but for four species the mean crop-initiation feil within the four-month dry season and for all species, the mean leaf-flushes feil within the first three months of the dry season. Phenology may vary with latitude, but little is known in this respect. Kjellberg & Maurice (1989) tried to determine, through simulations, the consequences of the general property of Ficus - so it would seem - to have a non-uniform year­ round distribution of crops and independence among trees, in connection with the maintenance of a population of their pollinators. In order to ensure the survival of the wasps over five years in 95% of the runs, 120 crops were necessary for the uniform distribution. The chosen probability (95% over 5 years) would mean 50% over 65 years: rare wasp transfers between adjacent populations may be sufficient to counteract local extinction (e.g., predation by fruit-bats, or by typhoons) and keep the system going. Another model investigated how the sea­ sonal pattern may evolve within a fig population as a result of individu al selec­ tion on the trees, and it showed how the breeding system of Ficus limits the expansion of the pollinators into highly seasonal habitats.Seasonal habitats may also lead to seasonal adjustment of male versus female investments and to the evolution of dioecy. Seasonality data on fig trees were generally discussed by Wolda (1987), who referred to several other papers on th is subject.

The infructence and the fruitlets In the postfloral phase the sycone is fully mature and it is in th is stage that the whole 'fruit', or only the fruitlets will be dispersed.

30 The fruitlets are quite diverse (Berg, 1984a; fig. 12). In some groups (e.g., ti subgenera Sycomorus and Pharmacosycea) the fruitlet is achene-like, with a dl crustaceous pericarp, or with only the outermost layer(s) of eells soft. Dr achene-like fruitlets may have mucilage from the wall of the syconium adherin to their bases. In other groups (e.g., the section Sycidium, F. palmata, and th subsection Caulocarpae of section Galoglychia) the fruitlets are similar to th basic type of fruit in Moraceae: the 'dehiscent drupe', with white, unevenly thick ened fleshy exocarp and a woody to crustaceous endocarp, which become relea sed - pushed out or ejected - by pressure of the exocarp. The released endocar bodies (pyrenes) look like seeds (with a 'pseudohilum') and are surrounded b: a mucilaginous layer. In subsection Cyathistipulae of the section Galoglychia the upper part of tht fleshy exocarp becomes inflated and mucilaginous. The ditTerenees in the fruitlets may have a bearing on predation, and the dispersal may be exozoochoric, because of the sticky layers, in addition to being (normally) endozoochoric. The waU of ovaria occupied by insect larvae develop ditTerently from th at of normal fruitlets: the pericarp becomes dry.

Dispersal Syconia are eaten by numero us species of birds, monkeys, bats, squirrels, as well as by terrestrial species eating fallen figs or cauliflorous and geocarpic figs. The diaspores (fruitlets and endocarp bodies) are usuaUy endozoochorously disper­ sed, but mucilage adhering to fruitlets or endocarps may promo te exozoocho­ rous dispersal. Breitwisch (1983) listed the frugivores at a fruiting Ficus in Came­ roon, while Gauthier-Hion & Michaloud (1989) focussed on the main food con­ sumed by Cercopithecine monkeys in the lean season, but also included other taxa. They concluded that figs are rather infrequently and opportunistically ea­ ten by frugivorous birds and mammaIs, although the large bats, such as Hypsi­ gnathus monstruosus, weU ab Ie to visit quite distant trees in a single night, are more relying on figs for food, and the same may apply to the bulbul (Ixonotus gustatus). Several insects are known as predators of 'seeds' and accidentaUy disperse them, e.g. , ants and Heteropterous insects (Slater, 1972; Couturier, 1984). In some taxa dispersal by water may be important. The spongy wall of the fig in F. cyathistipula may be an adaptation to hydrochory.

Germination

Most Ficus-species appear (or are supposed) to demand light for germination, but apparently in different degrees. For hemi-epiphytic species this implies dif­ ferent preferences or sites for establishment on host trees. In F. lutea and F. saussureana, both found at lower strata in hight forest, germinated seed have been found inside syconia.

31 ~ Fig. 13. Schematic diagrams of key features in Ficus. I, receptacle with peduncular, lateral, and ostiolar bracts (section Sycidium); 2, receptacle with basal and ostiolar bracts (other sections); 3, stipitate receptacle; 4, basal bracts two (section Galog/ychia); 5, basal bracts three (F. palmata, subgenus Sycomorus, sections Oreosycea, Urostigma and Conosycea); 6, most ostiolar bracts in­ terlocking (F. pa/mata, section Sycidium, subgenus Sycomorus, sections Urostigma and Conosycea); 7, the middle ostiolar bracts not interlocking (section Oreosycea); 8, all ostiolar bracts descending (section Ga/og/ychia); 9, ostiole circular, several ostiolar bracts visible (F. palmata, section Syci­ dium, subgenus Sycomorus); 10, ostiole circular, three ostiolar bracts visible (section Oreosycea, section Urostigma and Conosycea); 11 , ostiole slit-shaped (section Ga/oglychia); 12, stigma ± infun­ dibuliform (section Sycidium, subgenus Sycomorus); 13, stigma ± infundibuliform to tongue­ shaped (subgenus Sycomorus); 14, stigmas two (F. palmata, section Oreosycea, and sometirnes in section Galog/ychia); 15, stigma one, elongate (sections Urostigma, Conosycea and Galoglychia); 16, long-styled ('seed') ftower, often sessile or short-pedicellate; 17, sbort-styled ('gal!') ftower, (± long- pedicellate); 18, staminate ftower with distinet pistillode and hairy tepals (section Sycidium); 19, staminate ftower with saccate perianth and two stamens, being enveloped by two large bracts or bracteoles (subgenus Sycomorus); 20, staminate ftower with one stamen (sections Oreosycea, Urostigma, Conosycea and Ga/oglychia); 21, calyptrate bud-<:over (as found in, e.g., F. craterostoma and F. ovata); 21a, diagram of same; 22, glandular spots in the axils of tbe basal (or main) lateral veins beneath (F. palmata, section Sycidium, subgenus Sycomorus); 23, glandular spots two, on the base of the midrib beneath (section Oreosycea); 24, glandular spot one, on the base of the midrib beneath (sections Urostigma, Conosycea and Galoglychia); 25, basal lateral veins branched; 26, basal lateral veins unbranched; 27, tertiary venation scalariform; 28, tertiary venation predomi­ nantly parallel to the lateralveins; 29, tertiary venation reticulate. After Berg & Hijman, 1989 (fig. 16).

Seeds can keep viability during many years, if stored under dry and cool conditions.

Hybridization Among the specimens examined there were none that could he supposed to he species-hybrids. This may partly he the result of the selection of the pollinators, which do not normally confuse tig species, but also in species pairs sharing one and the same species of poJ1inator, such as F. mucuso and F. sycomorus, or F. dicranostyla and F. variifolia, hybrids appear to be absent. Pollen exchange be­ tween these species may be reduced by their occurrence in different habitats, viz. savanna woodland and rain forest. There is, however, some evidence that the arrival of pollen from foreign spe­ cies (even helonging to another section or subgenus) can result in the production of viabie seed (Compton, 1990; Condit, 1947). Compton reported on F. lutea pollinated by wasps of F. thonningii and F. sur: the seeds germinated, but they did not develop heyond the cotyledon-state (personal communication by Comp­ ton, 1991) - which inay be another reason for the absence of hybrids. Condit reported on two-years-old hybrids of F. carica and F. pumila, but it is not known wh at happened to them afterwards.

32 The c1assificatioD of the figs The most recent cIassification of Ficus was proposed by Corner (1965); it is adopted for the taxonomie part of the present publication, aithough not without some comments on details related to the African representatives of the genus. In later publications, Corner hirnself implicitly commented on bis cIassification, but without proposing changes. An (informal) modification of Corner's cIas­ sification was proposed by Ramirez (1974), not based on strong arguments with

33 Table 3. The subgenera and sections of the genus Ficus, after Corner (1965). The section Galoglychia was sub-divided by Berg (1986b). The nurnbers of the African species are given according to Berg (199Ob), and the genera of the pollinators are indicated. subgenus Number of species Genera of section (and subspecies) in pollinating subsection Africa and Madagascar wasps

UROSTIGMA Urostigrna 5 (+ 2) Platyscapa Leucogyne Conosycea 2 Platyscapa Stilpnophyllurn Americana Malvanthera Galogiychia Galoglychia 3 (+ 2) Allotriozoon Platyphyllae 18 Chlamydodorae 13 (+ 5) } table 6 Crassicostae 8 Cyathistipulae 19 (+ 3) Agaon Caulocarpae II (+ 8) CourtelIa PHARMACOSYCEA Pharmacosycea Oreosycea 4 Dolichoris FICUS Ficus Blastophaga Rhizocladus Kalosyce Sinosycidium Sycidiurn 9 (+ I) Kradibia Adenosperma Neomorphe Sycocarpus SY COMOR US 12 Ceratosolen

Total 105 (+ 21)

reference to morphological characters of Ficus, but in a way connected with his wish to match the classifications of the Agaonidae and Ficus (see Corner, 1985a). The existing discordances between the two indeed are stimuli to reconsider the two classifications. Some suggestions for remodelling the subdivision of Ficus were made by Berg (1989a, 1990a), but, awaiting thorough morphological stu­ dies on some groups of species, a formal modified classification cannot yet be proposed. Still, there are already several (rather) solid elements for such a new classification, as follows (cf. fig. 13; for the African species see table 3; a tenta­ tive, pseudo-cl ad is tic diagram is presented in the right part of fig . 14). There is: (1) a dichotomy between one group (A) comprising (the 'subgenera') Pharma-

34 cosycea and Urostigma, with a world-wide distribution, and another group (B) comprising the rest of the genus (Corner's subgenera Ficus and Sycomorus) con­ fined to the Old World. The differences between these groups are mainly found in monoecy (A) and gyno-dioecy (the majority of B) - see table l. (2) a dichotomy between (the 'subgenera') Pharmacosycea and Urostigma, but without very profound differences. (3) a dichotomy in Urostigma separating the species with slit-shaped ostioles (Galoglychia and Malvanthera) and the rest, with interlocked ostiolar bracts (forming a helicoidal passage to the interior ofthe syconium, cJ. Ramirez, 1974). One possible exception, i.e., E elastica Roxb., is the only species of the Asian section Stilpnophy/lum, having several features in common with Malvanthera, but not the structure of the ostiole and the single theca (Corner, 1985b). (4) a dichotomy in Pharmacosycea separating the New World species (in 'sec­ tion' Pharmacosycea) and the Old World species (in 'section' Oreosycea). (5) a dichotomy separating the African ('section') Galoglychia from the Aus­ tralasian ('section') Malvanthera (including E elastica?). (6) a dichotomy separating the New World ('section') Americana from the Old World species ('sections' Urostigma, Conosycea and Leucogyne). (7) in group B a subgroup Sycomorus (comprising Corner's subgenus Sy­ comorus and the sections Neomorphe and Sycocarpus) and a subgroup Sycidium, are quite distinct morphologically, far more distinct than the two subgroups in group A.

Elisabethiella sec!. Galoglychia also see table 6 Nigeriella

'-----Agaon Cyathistipulae

L..------Courtella Caulocarpae

'------Allotriozoon Galoglychia

L------Paragaon

'------Alfonsiella

r-----Platyscapa Urostigma I-- ,...----Dolichoris Oreos ycea A] '------Blastophaga Ficus ------,

r---- Kradibia Sycidium ----+-----B

'----Ceratosolen Sycomorus ----' Fig. 14. A cladogram for the Mrican Agaoninae (left) compared with that for the subgroups of Ficus (right).

35 (8) a dichotomy, in the subgroup Sycomorus separating a entity comprising Corner's subgenus Sycomorus and section Neomorphe, from another entity, i.e., Corner's section Sycocarpus. (9) a major problem is Corner's subgenus Ficus, which is morphologically quite heterogeneous and not likely to be monophyletic. The 'sections' Rhizocla­ dus and Kalosyce form a distinct entity of root-climbers, but when these are taken out, the rest still lacks suflicient uniformity: some subgroups showing relations with the Sycomorus-group and others with Sycidium.

The classification of the pollinators The main division (see Wiebes, 1982b) is in two tribes (for a discussion of the rank of the groups see the introduction to Chapter IV), viz., the Agaonini and the Blastophagini, although the connection of the new World genus of the Aga­ onini (i.e., Tetrapus Mayr, from 'section' Pharmacosycea) and the Old World genera (viz., Pleistodontes Saunders in Australia, from Malvanthera, and Agaon etc. in Africa, from Galoglychia) may be paraphyletic rather than monophyletic. For Africa, the division between the two tribes is obvious (tabie 4 and fig. 14). In the Agaonini, the genera Elisabethiel/a and Nigeriel/a are close in the female sex, but the morphology of the males separates Elisabethiel/a from the pair Nigeriel/a and Alfonsiel/a. In several groups of fig-wasps, it is difficult to find a parallel classification of the two sexes. Overall, Alfonsiel/a seems aberrant, and

Table 4. The genera of the African Agaoninae and the number of species, according to Wiebes & Compton (1990). The groups of host figs are indicated.

Tribe Number of species and Group of genus subspecies in Africa host fig and Madagascar

BLASTOPHAGINI Blastophaga Ficus Kradibia 5 Sycidium Ceratosolen 12 Sycomorus Dolichoris Oreosycea [parapristina I, introduced Conosycea] Platyscapa 6 Urostigma + Conosycea AGAONINI Allotriozoon 3 Galoglychia Elisabethiella 14 (+ I) Nigeriella 4 A1fonsiella 7 } ~b", Paragaon 2 Agaon II (+ I) Cyathistipulae CourtelIa 13 (+ 2) Caulocarpae

Total 80 (+ 4)

36 also Allotriozoon obviously differs from the complexes of Elisabethiella and Ni­ geriella, Agaon, and from CourtelIa. The situaton of Paragaon is not very dear. In the Blastophagini, the group of Ceratosolen and relatives (in Africa, Kradi­ bia) is dearly different from Blastophaga, as weil as also from Platyscapa. Tentatively, the relations of the African genera are depicted in the dadogram of fig. 14, and compared with the host-groups of Ficus.

Comparison of tbe classificatioDS of figs and wasps There are various levels at which one can compare the dassifications of figs and fig-wasps. At the lowest level: in general each species of Ficus has its own species of pollinator wasp, but some exceptions do occur. Some can be explained by special biological circumstances, such as the co-occurrence of Ceratosolen arabi­ cus and C. galili in the sycones of Ficus sycomorus, in which C. galili behaves like a clepto-parasite (Galil & Eisikowitch, 1969a). The question theri remains: where did it come from? For a time, Ficus mucuso seemed a good candidate for an original host of C. galili, but also there, it seems to act as a cuckoo (Wie bes, 1989b: 261). The species of wasp co-occurring with Ficus ottoniifolia (Michaloud et al., 1985) show other possible explanations. Subspeciation might, for the different subspecies of the fig, lead to different species of pollinators, such as F. ottoniifolia multinervia in Ivory Co ast with C. gabonensis [never mind the name!]; but F. o. ottoniifolia in Cameroon and Gabon, F. o. macrosyce in Zambia, and F. o. ulugurensis in Kenya with C. camerunensis; and F. o. lucanda in Uganda with C. scobinifera. The co-occurence of C. gabonensis and C. camerunensis in Gabon, then, may be explained by a secondary sympatrism of the wasp species (with some niche-se para ti on). AH exceptions known for the African species of fig are mentioned in tab Ie 5. The list is rather long, but it contains instances in which the pollinators of more than one species of fig are not distinct morphologically, although they may be different biologically. More interesting are those mentioned above, the excep­ tions that are found in the sycones of Ficus sur, viz., Ceratosolen capensis and C. jlabellatus, and the co-occurence of species of Alfonsiella with other Agaonini. At a taxonomic level, higher than that of the specific relations, a phylogenetic specificity is apparent. Related species of fig have related pollinators: the species of section Ficus have pollinators of the genus Blastophaga, those of Sycidium: Kradibia, Sycomorus: Ceratosolen, Oreosycea: Dolichoris, Urostigma: Platy­ scapa; the Galoglychia have Agaonini, as against the ot her groups (botanically, however, not an unity!) Blastophagini, the subsection Galoglychia has Allotrio­ zoon, those of the Cyathistipulae: Agaon, and Caulocarpae: CourtelIa. Also here, there are interesting exceptions, next to the misfit of the Agaonine tri bes already mentioned. From tables 4 and 6 it is apparent th at there is no exact fit between the classification of the figs in the subsections Platyphyllae, Chlamydodorae and

37 Crassicostae, and that of the pollinators in the genera Alfonsiella, Nigeriella, Paragaon and Elisabethiella (Wiebes, 1990). Wiebes (1989a: 118) expressed the opinion that the classifieation of the fig species should be reeonsidered, with th at of the pollinators as a suggestion. Berg (1989a: 610-611), although expeeting that part of the diserepaney between the classifieations will disappear through further taxonomie studies and re-evalua-

Table 5. Exceptions to the specificity of the symbiosis.

Ficus Fig-insect(s) (Agaoninae)

1. palmata B. psenes, also from F. carica 2. exasperata K gestroi, also from F. asperifolia; and from F. capreifolia 5. asperifolia K gestroi, sec 2 and 6; from the urceolaris-form: K hilli 6. capreifolia K gestroi, sec 2 and 5 11. sycomorus C. arabicus and C. galili; both also from F. mucuso; 'var.' sakalavarum: C. namorakensis 12. mucuso C. arabicus and C. gali/i; both also from F. sycomorus 13. sur C. capensis or C. silvestrianus, and C. flabellatus, sec II - 13; and sec IS 15. vallis-choudae C. megacephalus and silvestrianus? see 13 23. variifolia D. flabellata, also from F. dicranostyla 24. dicranosty/a D. flabellata, also from F. variifolia 40. vasta E. socotrensis, also from F. wakefieldii; and from F. n. nata/ensis; Alfonsiella, see remarks below: 60, 66, 67 41. wakefieldii E. socotrensis, see 40 47. abutilifolia E. comptoni, also N. fusciceps 58. craterostoma A. micha/oudi, also from F. I. lingua 59. lingua A. micha/oudi, also from F. craterostoma 60. natalensis E. socotrensis, also from F vasta; also Alfonsiella natalensis?; sec 66; A. fimbriata from F. n. /eprieurii; sec 67; A. longiscapa from F. n. natalensis; sec 66; 66. thonningii E. stuckenbergi, also A. longiscapa; and also A. natalensis; see 60 67 . kamerunensis A. fimbriata, also from F. n. leprieurii 76. conraui A. kiellandi, also [rom F. c. pringsheimiana; and from F. densistipu/ata 77 . tesse/ata A. taiense, ? also A. balio/urn 82. pringsheimiana A. kiellandi, also from F. conraui; and from F. densistipu/ata 86. densistipulata A. kiellandi, also from F. conraui; and from F. pringsheimiana 95. ottoniifolia C. camerunensis, gabonensis & scobinifera 97. artocarpoides C. penicula, also C. h/adikae

38 Table 6. Agaoninae from Platyphy/lae, Chlamydodorae and Crassicostae: the 'hosts of Alfonsie/la, Nigeriella, Paragaon and Elisabethie/la.

Agaonidae

Ficus Alfonsiella Nigeriella Paragaon Elisabethiella

Platyphyllae 37 platyphy/la near pectinata 40 vasta socotrensis 41 wakefieldii socotrensis 42 glumosa glumosae 43 stuhlmannii binghami 44 nigropunctata bergi 45 tettensis excavata 47 abutilifolia fusciceps comptoni 50 trichopoda bergi 52 rubra avico/a

Chlamydodorae 55 ca/yptrata pectinata 56 fischeri platyscapa 58 craterostoma michaloudi 59 lingua micha/oudi 60 nata/ensis fimbriata socotrensis /ongiscapa 62 burtt-davyi baijnathi 63 i{jcina enriquesi 65 reflexa reflexa 66 thonningii brongersmai stuckenbergi longiscapa 67 kamerunensis fimbriata

Crassicostae 68 elasticoides artieu/ata 69 burretiana /ongiscapa 71 pseudomangifera letouzeyi 74 louisii josephi

tion of the characters used, suggested the subdivision of the Agaoninae to he related to the type of ostiole, which shows similarities in the sections Galoglychia, Malvanthera and Pharmacosycea (all three with long-faced 'Agaonini' as pol­ linators). Indeed, the pattems in the functional significance of features (taken as taxonomic characters) of the syconium, flowers, fruits, and of the wasp's mandibulae, antennae, armature of the tibiae, etc., are not yet fully traced.

The origin and development of the symbiosis Describing the development of the symbiosis leads to little more (nor less!) than

39 explaining all adaptations of figs and wasps to their symbiosis, our knowledge of which still is very deficient. Still, several suggestions can be made about the ancestral way of life of figs and fig-wasps. One may regard an more or less open, discoid to cup-shaped receptacle with extemal, or at least marginal, bracts and with numerous small unisexual flowers tightly packed together, as the (Compositae-like) type of inflorescence, from which the syconium developed and the relation between (pre-)Agaonids and (? the ancestors of) Ficus originated. An inflorescence like this would have attrac­ ted insects, beetles and fEes in particular, for breeding (Berg, 1990a, I 990d). Several characteristic features of the syconium, among them those of importance for the pollination system, can be explained as being developed to protect the inftorescence (ftowers, and developing ovaries and seeds in particular) from damage by insect larvae, e.g.: (I) barring out unwelcome visitors from the interior of the inflorescence (urceolate receptacle, ostiolar bracts); (2) crowding of the ftowers, by various interftoral bracts or bristles, and for­ ming of a synstigmatic (matted) layer; (3) filling the cavity with ftuid; (4) hiding the staminate among the pistillate ftowers, under the stigmatic layer, or in large bracteoles; or by separating starninate and pistillate flowers spatially (dioecy) or temporary (by postponement of the exposure of the an­ thers). The degree of perfection, in relation to morphology, behaviour, timing, etc., appears to .be different for different groups of figs. In the Sycidium-group, the inftorescences have many 'prirnitive' morphological features, and the pollina­ tion-system appears to be less 'weil organized' than in other groups. The subdivi­ sion of the genus Ficus and its morphological differentiation may suggest that the relation with the fig wasps was established in an already more or less di­ versified group. This implies the possibility that a pollination-system fully de­ pendent on Agaoninae, did not become established in the same period through­ out the genus. Pattems in the distribution of Ficus appear to comply with this possibility. Crucial in all scenarios for the pollinators, is the attainment of entrance to the (c1osing) receptacle and carry pollen. The sister-group of the Agaoninae most probably is formed by the Sycoecinae (Wiebes, 1982b: 405, 407), but a compa­ rison is difficult, because also they have many adaptations to entering the sy­ conium. In most characters, the Sycoecinae are Ie ss reduced than are the Agao­ nmae. Wiebes (1963) suggested that the Agaoninae could have been descended from gall-forming Chalcidoidea or from para si tic Chalcidoidea living on other insects, in the ftowers or seeds of the pre-Ficus, but Ramirez (1976) suggested that they evolved from pollen-feeding gall-makers. Wiebes (1979c) argued that it is easy to be imagined that the active carrying of pollen in the digestive tract (such as may occur in New World Tetrapus) was from the beginning one of the

40 strategies followed, an alternative being to carry it in pockets. The structure of the pockets is insufficiently known, but they may not all be the same: for instance, the relatively open pockets of the Agaonini and e.g., Ceratosolen, differ from the closed pockets, as e.g., described by Galil & Snitzer-Pastemak (1970) for Platyscapa quadraticeps.

Parasitoids and predators The data and references of this paragraph are rather anecdotal, and not at all complete; also not all groups are mentioned, e.g., Curculionidae (Perrin, 1991), and Staphylinidae, about we know nothing but for the fact that they may be associated with figs! Best known of the several groups of organisms known to be predatory on the fig-sycones, fig seeds, andJor the (larvae of the) pollinat­ ing fig wasps, mainly by the efTorts of D. Lachaise (e.g., Lachaise, 1977; Lachaise et al., 1982), are the ca. twenty-odd species of the Drosophilid genus Lissocep­ hala, all strictly specialized on Ficus. Pendants are formed by the sycophagous species of the genus Zaprionus and the group of Drosophila fima, like Lissocep­ hala endemic to Africa. The eggs are laid in or near the ostiole, through which the larvae enter, and they breed inside the immature sycone. The third-instar larvae leave the syconium again, also through the ostiole, and drop to the soil, where they pupate. Several species may exploit the same syconium, neatly sepa­ rating the niches, and one species of Lissocephala may infect more than one species of fig. Compton & Disney (1991) described how the larvae of five species of Me­ gaselia (scuttle tlies; Diptera: Phoridae) feed in mature sycones of several species of Ficus. Adult phorids were regularly observed congregating on the surface of the sycones as the female wasps were emerging, which they pursued and cap­ tured. The (female) flies pierce the cuticle of the fig wasps and suck out the body contents; the male flies are non-predatory, but - like also the females - feed on the broken edges of the sycones. Barnes (in Williams, 1928: 21 - 24) described two gall-midges (Diptera, Ceci­ domyidae), the larvae of which feed in the rind of fig-sycones in the Philippines; Roskam & Nadel (1990) redescribed an American gall-midge of the genus Ficio­ myia, inhabiting syconia of an American fig . Several Lepidoptera, representing various families (see e.g. , New, 1982) are usually reared from galls (see also Compton, 1989, for catterpillars on fig leaves); earlier, Williams (1928: 24-26) gave quite a list of (Philippine) Lepidoptera at­ tacking figs: green sycones, leaves, and twigs included. There also were many other insect-groups mentioned, not all of which are repeated in this survey. DiakonofT (1983) described a species of Stathmopoda from African Ficus lu tea, coUected in the Seychelles Islands. Already Gasparrini (1864) recorded nematodes carried by Blastophaga psenes. Martin et al. (1973) gave a short review of the pertinent literature and listed six Rhodesian species of fig, in the sycones of which they found nematodes to a

41 number of as many as 50.000 in one sycone! Poinar (1979) described Parasitodi­ plogaster sycophilon from Ficus thonningii. It occurs in the body cavity of the fig wasp Elisabethiella stuckenbergi. The biology is not weil known, but they must be transported by the female wasp to the newly colonized sycones, where they deve\op. See also Poinar & Herre (1991). Another phoretic relationship exists between fig-wasps and mites. That rnÎtes inhabit fig-trees is weil known (e.g., Rasmy & Abou-Awad, 1972), but the many rnÎtes seen inside mature sycones and on the body of fig-insects, remained un­ noted until Wiebes (1968c: 117) mentioned phoretic Tarsonemids on the gaster of Pleistodontes rennellensis Wiebes, and Wiebes & Compton (1990: 208) recor­ ded Tarsonemella africana (Hirst, 1923) to be phoretic on Kradibia gestroi in Sénégal. Mites like these may be found on several parts of the (female) Agaonid body, such as sheltered places on the thorax and gaster, or in a row on the ovipositor-valves. Ants may show a rather complex relationship to figs and fig-insects (Compton & Robertson, 1988), protecting seeds and pollinators by tending homopterans and fighting off all other inftuences. Thomas (1988) examined the effect of ag­ gressive arboreal weaver-ants (Oecophylla longinoda) on fruit removal from the African Ficus sur by fruit-bats. Grimont et al. (1979, 1981) isolated Serratia-bacteria from figs (Ficus carica) and fig-wasps (Blastophaga psenes): this is just mentioned here to give an entry - there may be much more known! Lachaise et al. (1979) mentioned the yeast-ftora of figs. The many Cha\cidoid fig-symbionts are specially treated in Chapter IV. Most are quite specific, also for the groups of Ficus. Figs of the subgenus Sycomorus, for in stance, have a characteristic fauna, a.o. including a species of Sycophaga (entering the syconium for oviposition) and one of Apocrypta (ovipositing from the outside), and lacking e.g., Otitesellinae. In Africa, Sycoecinae are exclusive to Galoglychia-figs, such as e.g., Sycoecus with Agaon in the subsection Cyathi­ stipulae and Seres with CourtelIa in the Caulocarpae. Wiebes (in Boucek et al., 1981 , table 1) gave a survey of groups and selected genera of fig insects, with the Ficus-species from which they were reared. Most of the forms with long ovipositor-valves are clepto-parasites and most, if not all, are phytophagous, but too little is known of their biology to generalize. In contrast with the males of the pollinators, those of many of the parasitoids do fight, to the extent of infticting severe injuries. Murray (1989: 186) mentioned a mortality of 50% or higher in New World ldarnes. In a species of Philotrypesis, i.e., P pilosa Mayr from Malayan Ficus hispida Linn.f., the cost of fighting, estimated from the length and frequency of fights and by the extent of male injury, was highest at an intermediate density of males (Murray, 1987). Murray's data and model (Murray & Gerrard, 1985) led Frank (1987: 260) to a (mild) critique on the accurateness of the assumptions about male movement and sperm competition. Many more observations are urgently required.

42 REFERENCES (only those not mentioned in the bibliography)

Addicott, J.F., J. Bronstein & F. Kjellberg - Evolution of mutualistic life-cycles: Yucca moths and fig wasps. In: F. Gilbert (ed.), Genetics, evolution, and coordination of insect life cycles. Springer Verlag, London, pp. 143--161 (1990). Beach, J.H. & K.S. Bawa - Role of pollinators in the evolution of dioecy from distyly. Evolution 34, 1138- 1142 (1980). Beek, N.G. & E.M. Lord - Breeding system in Ficus carica, the common fig. i. Floral diversity. Amer. J. Bot. 75, 1904--1912 (l988a). Beek, N.G. & E.M. Lord - Breeding system in Ficus carica, the common fig. ii . Pollination events. Amer. J. Bot. 75, 1913-1922 (I 988b). Berg, C.C. - Floral difTerentiation and dioecism in Ficus (Moraceae). Acta Bot. Neerl. 32, 344-345 (1983). Berg, e.e. - Differentiation of fruitlets of Ficus, especially of subg. Urostigma sect. Galoglychia. Minisymposium: Figs and fig insects (1983), CNRS-Centre L. Emberger, Montpellier, 29-33 (I 984a). Berg, e.C. - Classification and distribution of Ficus. Experientia 45, 605-611 (1989a). Berg, e.e. - Systematics and phylogeny ofthe Urticales. In: P.R. Crane & S. Blackmore (eds.), Evolu­ tion, systematics, and fossil history of the Hammamelidae, vol. 2 ('Higher Hammamelidae'). Syst. Assoc., Oxford, 193--220 (l989b). Berg, C.C. - Reproduction and evolution in Ficus (Moraceae): traits conneeted with the adequate rearing of pollinators. In: G. Gottsberger & G.T. Prance (eds.), Reproductive biology and evolu­ tion of tropical woody angiosperms. Mem. New York Bot. Gdn. 55, 169-185 (l990a). Berg, e.e. - DifTerentiation of fiowers and infiorescences of Urticales in relation to their proteetion against breeding insects and to pollination. Sommerfeltia 11, 13- 34 (l990d). Bronstein, J.L. - Mutualism, antagonism, and the fig-pollinator interaction. Ecology 69, 1298- 1302 (1988). Caragiio, Y. - Architecture de quelques espèces du genre Ficus L. Thesis, Montpellier, 1- 73 (1985). Compton, S.G. - Sabotage of latex defences by caterpillars breeding on fig trees. S. Mr. J. Sci. 85, 605-606 (1989). Compton, S.G. & R.H.L. Disney - New species of Megaselia (Diptera: Phoridae) whose larvae live in fig syconia (Urticales: Moraceae), and adults prey on fig wasps (Hymenoptera: Agaonidae). J. Nat. Hist. 25, 203--219 (1991). Compton, S.G. & H.G. Robertson - Complex interactions between mutualisms: ants tending ho­ mopterans proteet fig seeds and pollinators. Ecology 69: 1302- 1305 (1988). Condit, 1.1 . - The structure and development of fiowers in Ficus carica L. Hilgardia 6, 443-481 (1932). Condit, 1.1. - The fig. Chronica Botanica Co., Waltham, xv + 222 pp. (1947). Corlett, R.T. - The phenology of Ficus jistulosa in Singapore. Biotropica 19, 122-124 (1987). Corner, EJ.H. - Wayside trees of Malaya (ed. 2), 1. Govt. Printing OfT. , Singapore, 772 pp. (1952). Corner, EJ.H. - Cheek-list of Ficus in Asia and Australasia, with keys to identification. Gdns' Bull. Singapore 21 , 1-186 (1965). Corner, EJ.H. - Ficus in the Solomon islands and its bearing on the post-Jurassic history of Melanesia. Philos. Trans. Roy. Soc. London (B) 253, 23--159 (1967). Corner, EJ.H. - The climbing species of Ficus: derivation and evolution. Philos. Trans. Roy. Soc. London (B) 273, 359- 386 (1976). Corner, EJ.H. - Ficus (Moraceae) and Hymenoptera (Chalcidoidea): figs and their pollinators. Biol. J. Linn. Soc. 25, 187- 195 (1985a). Corner, EJ.H. - Essays on Ficus. Allertonia 4, 125-168 (1985b). Couturier, G. - Les héteroptères vivant au pied des Ficus en Cöte d'Ivoire. Courrier de I'Ecotrop, Num. spèc., 42-53 (1984). Cruden, R.W. - Polien-ovule ratios: a conservative indicator of breeding systems in fiowering plants. Evolution 31 , 32-46 (1977).

43 DiakonofT, A. - A new species of Stathmopoda. Ent. Ber. Amst. 43, 117-118 (1983). Frank, S.A. - Hierarchical selection theory and sex ratios 11. On applying the theory, and a test with fig wasps. Evolution 39, 949-964 (1985). Frank, S.A. - Weapons and fighting in fig wasps. Treee 2, 259- 260 (1987). Frank, S.A. - Ecological and evolutionary dynarnics of fig cornrnunities. Experientia 45, 674-680 (1989). Gali1, J. - Topocentric and ethodynamic pollination. In N.B.M. Brantjes (ed.), Pollination and dispersal, 85-100. Nijmegen, The Netherlands (1973). Galil, 1. - Ficus religiosa L. - the tree splitter. Bot. J. Linn., Soc. 88, 185-203 (1983). GaliI, J. & D. Eisikowitch - On the pollination ecology of Ficus religiosa in Israel. Phytomorph. 18, 356-363 (1968c). Galil, J. & G. Neeman - Pollen transfer and pollination in the cornrnon fig (Ficus eariea L.). New Phytol. 79, 163- 171 (1977). Galil, J. , W. Rarnirez B. & D. Eisikowitch - Pollination of Ficus eostarieana and F hemsleyana by Blastophaga estherae and B. tonduzi in Costa Rica (Hymenoptera: Chalcidoidea, Agaonidae). Tijdschr. Ent. 116, 175- 183 (1974). Galil, J. & Y. Snitzer-Pasternak - Pollination in Ficus religiosa L. as connected with the structure and mode of action of the pollen pockets of Blastophaga quadratieeps. New Phytol. 69, 775-784 (1970). Ganders, F.R. - Tbe biology of heterostyly. New Zealand J. Bot. 17, 607-635 (1979). Gasparrini, G . - Sulla maturazione e la qualita dei fichi dei contorni di Napoli. Atti Accad. Pon­ taniana 9, 99-118 (1864). Godfray, H.CJ. - Virginity in haplodiploid populations: a study on fig wasps. Econ. Entom. 13, 183- 291 (1988). Grandi, G. - Studio morfologico e biologico della Blastophaga psenes (L.). 2a edizione riveduta. Boli. Lab. Ent. Bologna 2, 1-147 (1929). Grirnont, P.A.D., F. Grimont & M.P. Starr - Serratia flearia spec. nov., a bacterial species as­ sociated with Smyrna figs and the fig wasp Blastophaga psenes. Current Microbiology 2, 277-282 (1979). Grimont, P.A.D., F. Grimont & M .P. Starr - Serratia species isolated from plants. Current Micro- biology 5, 317- 322 (1981). Hamilton, W.D. - Extraordinary sex ratios. Science 156, 477-488 (1967). Herre, E.A. - Sex ratio adjustment in fig wasps. Science 228, 896-898 (1985). Herre, E.A. - Coevolution of reproductive characteristics in 12 species of New World figs and their pollinator wasps. Experientia 45, 637-647 (1989). HilI, D.S. - Figs (Ficus spp.) of Hong Kong. Hong Kong Univ. Press, i-viii, 1-\30 (1967b). Hirst, S. - On some new or little known species of Acari. Proc. Zool. Soc. London (1923), 971 - 1000 (1923). Janzen, D.H. - How to be a fig. Ann. Rev. Ecol. Syst. 10, 13-51 (1979). Joseph, K.J. - Recherches sur les Chalcidiens Blastophaga psenes (L.) et Philotrypesis earieae (L.) du Figuier Ficus eariea (L.). Ann. Sci. Nat., Zool. (ser. 11) 20, 187- 160 (1958). Kjellberg, F., B. Doumesche & J.L. Bronstein - Longevity of a fig wasp (Blastophaga psenes). Proc. Kon. Ned. Akad. Wet. (C) 91 , 117- 122 (1988). Kjellberg, F. & S. Maurice - Seasonality in the reproductive phenology of Ficus: its evolution and consequences. Experientia 45, 653-660 (1989). Lachaise, D. - Niche separation of African Lissoeephala within the Ficus Drosophilid cornrnunity. Oecologia (Berl.) 31, 201 - 214 (1977). Lachaise, D ., M.-C. Pignal & J. Rouault - Yeast flora partitioning by Drosophilid species inhabiting a tropical African savanna of the Ivory Coast (Diptera). Ann. Soc ent. France (N.S.) 15, 659-{j80 (1979). Lachaise, D ., L. Tsacas & G. Couturier - The Drosophilidae associated with tropica1 African figs . Evolution 36, 141 - 151 (1982).

44 Martin, G .e., A.M. Owen & J.l. Way - Nematodes, figs and wasps. J. Nemat. 5, 77- 78 (1973). Medway, L. - Phenology of a tropical rain forest in Malaya. Biol. J. Linn. Soc. 4, 117-146 (1972). Miami (multi-author review) - The comparative biology of figs. Experientia 45: 599...{j80 (1989). Murray, M.G. - The closed environment of the fig receptacle and its inftuence on male conflict in the Old World fig wasp, Phi/otrypesis pi/osa. Anim. Behav. 35, 438-506 (1987). Murray, M.G. - Environmental constraints on fighting in ftightless male fig wasps. Anim. Behav. 38, 186-193 (1989). Murray, M.G. & RJ. Gerrard - Putting the challenge into resource exploitation: a model of contest competition. J. theor. Biol. 115, 367- 389 (1985). Neeman, G . & J. Galil - Seed set in the 'male syconium' of the common fig, Ficus carica L. (caprificus). New Phytol. 81 , 375-380 (1978). New, T.R. - Lepidoptera from Uromyc/adium galls on Acacia. Austr. J. Zool. 30, 357- 364 (1982). Okamoto, M. & M. Tashiro - Mechanism of pollen transfer in Ficus erecta by Blastophaga nip­ ponica. Bull. Osaka Mus. Nat. Hist. 34, 7- 16 (1981). Pemberton, e.E. - The fig wasp in relation to the development of fertile seed in the Moreton Bay fig. Hawaii. Plant. Rec. 24, 297- 319 (1921). Perrin, H . - Biosystématique et écologie évolutive des Curculio (Coleoptera: Curculionidae). Double radiation sur Ficus (Moraceae) et sur Fagales. Thesis, Univ. Paris-VI, 266 pp. (1991). Poinar, G .O. - Parasitodiplogaster sycophilon gen. n. , sp. n. (Diplogasteridae: Nematoda), a pa­ rasite of Elisabethiella stuckenbergi Grandi (Agaonidae: Hymenoptera) in Rhodesia. Proc. Kon. Ned. Akad. Wet. (C) 82, 375-381 (1979). Poinar, G .O. & E.A. Herre - Speciation and adaptive radiation in the fig wasp nematode, Parasitodip/ogaster (Diplogasteridae: Rhabdita) in Panama. Revue Nématol. 14, 361-374 (1991). Rarnirez B., W - Fig wasps: mechanism of pollen transfer. Science 163, 580--581 (1969). Rarnirez B., W - Coevolution of figs and Agaonidae. Ann. Missouri Bot. Gard. 61 , 770--780 (1974). Ramirez B., W - Evolution of blastophagy. Brenesia 9, 1-4 (1976). Rasmy, A.H . & B.A. Abou-Awad - Mites inhabiting fig trees in Egypt. Zeitschr. angew. Ent. 70, 314-316 (1972). Roskam, J.e. & H. Nadel - Redescription and immature stages of Ficiomyia perarticulata (Diptera: Cecidomyidae), a gall midge inhabiting syconia of Ficus citrifolia. Proc. ent. Soc. Wash. 92, 778- 792 (1990). Slater, JA - Lygaeid bugs (Hemiptera: Lygaeidae) as seed predators of figs. Biotropica 4, 145-151 (1972). Steenis, e.G.G.J. van - Rheophytes of the world. SijthotT & NoordhotT, Alphen aan den Rijn - Rockvilie, xv + 407 pp. (1981). Storey, WB. - Figs. In: J. Janicke & J.N. Moore (eds.), Advances in fruit breeding. Westlafayette, 568- 5589 (1975). Thomas, O.W. - Tbe inftuence of aggressive ants on fruit removal in the tropicaJ tree, Ficus capensis (Moraceae). Biotropica 20, 49--53 (1988). Valdeyron, G. & O.G. Lloyd - Sex ditTerences and ftowering phenology in the common fig, Ficus carica L. Evolution 33, 673...Q85 (1979). Verkerke, W - Structure and function of the fig. Experientia 45, 612--622 (1989). Wiebes, J.T. - Taxonomy and host preferences of Indo-Australian fig wasps of the genus Cerato­ solen (Agaonidae). Tijdschr. Ent. 106, 1-112 (1963). Wiebes, J.T. - A new Pleistodontes (Hymenoptera Chalcidoidea, Agaonidae) from Rennell Island. Nat. Rist. Rennell isl., Brit. Solomon !sIs. 5,115-117 (l968c). Wiebes, J.T. - A short history of fig wasp research. Gdns' Bull. Singapore 29, 207-232 (I 977a). Wiebes, J.T. - Co-evolution of figs and their insect pollinators. Ann. Rev. Ecol. Syst. 10: 1-12 (1979c). Wiebes, J.T. - Fig wasps (Hymenoptera). In: J.L. Gressitt (ed.), Biogeography and ecology of New Guinea. Monogr. Biol. (W. Junk) 42, 735-755 (l982a).

45 Wiebes, l.T. - The phylogeny of the Agaonidae (Hymenoptera, Chalcidoidea). Neth. l. Zool. 32, 395--411 (I 982b ). Wtlliams, F.X. - Studies in tropical wasps - their hosts and associates (with descriptions of new species). Bull. Hawaü. Sug. Plant. Exp. Sta., Ent. Ser. 19, 1-179 (1928). Windsor, O.M., D.W. Morrison, M.A. Estribi & B. de Leon - Phenology of fruit and leaf produc­ tion by 'strangier' figs on Barro Colorado Island, Panama. Experientia 45, 647-653 (1989). Wolda, H. - Seasonality and the community. In: l.H.F. Gee & P.S. Giller (eds.), Organization of communities. Past and present. Blackwell, Oxford, etc., pp. 69-95 (1987).

46 111. The genus Ficus in Africa

Introduetion The present study of the African representatives of Ficus is a compilation of data accumulated through preparation of treatments for various regional ftoras, supplemented by descriptions and data not published in these treatments. The greater part of the collections of African Ficus species have been examined, essen ti al type collections and literature studied, and some field work carried out. The present study misses the profoundness of a revision, but the taxonomy of African Ficus appears to be sorted out satisfactorily. Some problems, however, still exist, as there still are uncertainties about the delimitation of some species (e.g., those of the section Sycidium in Madagascar) and the correct application of the ranks of species or subspecies. In Ficus thonningii the pattems of variation are so complex and confusing, that possibly too broad a species-concept had to be adopted. The classification by Corner (1965) was adopted for the present treatment. Several changes in this classification were suggested (see p. 33- 36), but these have no consequences for the grouping of the African species, except for the question whether or not the distinction between the sections Urostigma and Conosycea sho­ uld be maintained. Corner did not propose a subdivision of the sections outside the region of Asia and Australasia. A revised subdivision of the exclusively African section Ga­ loglychia was proposed by Berg (1986b), replacing subdisions proposed previ­ ously by e.g. , Mildbraed & Burret (1911), Hutchinson (1916) and Lebrun (1934). There remain some doubts about the correct position of some species, as dis­ cussed below under the subsections.

Taxonomie history The taxonomic history of African Ficus is similar to that of most larger genera of Angiosperms: same authors described only one species, several a few, and others (like Baker, 1877- 1894; De Wildeman, 1901-1922; Hiem, 1900; and Warburg, 1894-1906) many. Some authors also attempted to bring some order in the genus, in that way setting mi1estones such as those by Vahl (1805), Miquel (1848-1867), Mildbraed & Burret (1911), and Hutchinson (1916-1917). For re­ gional ftoras this was done by Perrier de la Bäthie (1928) for Madagascar, Lebrun & Boutique (1948) for Zaïre, Eggeling & Da1e (1951) for Uganda,

47 Andrews (1952) for Sudan, Keay (1958) for West Africa, Aubréville (1959) for Ivory Coast, Dale & Greenway (1961) for Kenya, Palmer & Pitman (1972) and Palgrave (1977) for South(ern) Africa, and Berhaut (1979) for Sénégal. In recent years, several more critical revisons were made for regional floras, viz., Aweke (1979) for Ethiopia, van Greuning (1982, 1990) for South Africa, Troupin (1982) for Rwanda, and Friis (1982, 1989) for Somalia and Ethiopia, Geerling (1982) for the Sahel, while Berg, often with colJaborators such as van Heusden en Weerdenburg, prepared treatments of Ficus for several regional floras (1984- 1990).

Distribution The distribution of the African taxa was discussed by Berg (1990b), from which paper the following data are taken. For the African flora region 105 species of Ficus are recognized, a number somewhat smaller than the estimated number (125-150) for the Neotropics, and about one-fifth of that for the Asian-Aus­ tralasian region. African Ficus are more diverse than Neotropical Ficus, but less diverse than Ficus in Asia and Australasia. Seven out of the 17 sections recognized by Corner (1965) occur in the African flora region. The section Galoglychia is the largest of the sections re­ presented, and it is almost confined to the region (a few species extend to the southern part of the Arabian peninsula and Socotra). This section has nine species in the Madagascar (sub)region, three of them also occurring in continen­ tal Africa. Second to Galoglychia is subgenus (or section) Sycomorus with 12 species, only one of which (i.e., F. racemosa L.) occurs outside the region. This group has eight species in the Madagascar (sub)region, one ofwhich (i.e., F. sycomorus) also occurs on the continent. The other African species are representatives of sections centered in the Asian­ Australasian region, viz., Sycidium with nine species, five of which in the Mada­ gascar (sub)region and four in continental Africa; Urostigma and Conosycea together, with seven species, four of which in the Madagascar (sub)region and three in continental Africa; Oreosycea with four species, two of which in the Madagascar (sub)region and the other two in continental Africa; and Ficus with F. palmata in north-eastern Africa, as an extension of its Asian range of distribu­ tion. The only other African species with a range so far outside Africa, is F. exasperata (section Sycidium). Twenty-eight Ficus species are known from the Madagascar (sub)region (Ma­ dagascar, the Mascarene Islands, the SeychelJes, the Aldabra Islands and the Comoro Islands), four of which are shared with the African continent, which in addition has 77 species. Most of these 81 species are confined to the continent, some extend to the islands in the Gulf of Guinea and the Cape Verde Islands, some to the southern part of the Arabian Peninsuia and Socotra, and two species (F. exasperata and F. palmata) extend to India.

48 The groups (sections) centered outside Mrica are relatively better represented in the Madagascar (sub)region than in continental Africa. The same applies to the Sycomorus (sub)group, regarded as part of a predominantly Asian-Aus­ tralasian subgenus (?), also comprising the sections Neomorphe and Sycocarpus, rather than as a distinct subgenus in itself. Forty-nine (out ofthe 81) species found on the African continent are elements of (lowland) forests of the Guineo-Congolian region, the Lake Vicoria region, and the transitional zone to the Zambezian region (cf. White, 1983). Only two of them (F. pachyneura and F. scott-elliotiz) are confined to the western part of this region (West of the 'Dahomey gap'). Moreover, F. ottoniifolia subsp. multi­ nervia (and a yet undescribed subspecies of F. cyathistipu/a) are confined to this subregion. F. lyrata and F. sagittifolia, both also occurring in western Ca­ meroun, can be regarded as centered in the western subregion. In addition to the two species last-mentioned, 33 species occur both East and West of the 'Dahomey gap'. Eleven species are confined to the central and eastern part of the Guineo­ Congolian region. In this part also two montane species occur, as weIl as two species (F. amadiensis and F. scassellatiz) occurring disjunctly in eastern Mrica (see below). Several species of the Guineo-Congolian region extend to the Zambian region, sometimes with more or Ie ss distinct forms, as in F. craterostoma, F. nata/ensis subsp. /eprieurii, and F. sansibarica subsp. macrosperma, or even a distinct subspecies (F. ottoniifolia subsp. macrosyce). Twelve species, widely distributed in the Guineo-Congolian region, occur dis­ junctly in forested areas in eastern (south-eastern) Mrica (Zanzibar-Inhambane and Tongaland-Pondoland mosaics), including forested (submontane) areas in the Zambezian region (cf. White, 1983), five of them with distinct subspecies (F. lingua, F. nata/ensis, F. ottoniifolia, F. sansibarica, and F. tremu/a; cf. Berg, 1988). This region has, moreover, three species with small ranges of distribution (F. bizanae and F. burtt-davyi in its southern-most part and F. usambarensis in Tanzania). FinaIly, this region is inhabited by three more wide-spread species, viz. , F. bussei (very closely related to the Guinea-Congelian F. recurvata), F. chirindensis, F. scassellatii (centered in eastern Africa and known from one col­ lection in E. Zaire), and F. amadiensis (centered in the eastern part of the Guineo-Congolian region and from a few collections known in the area North of Lake Malawi). Twenty-three species (excluding F. sur and F. thonningii, see below) found in continental Mrica are components of drier types of vegetation, varying from savanna woodland to semi-desert. Fourteen of them occur in the Soudanian & Somalia-Masai regions (cf. White, 1983). Four species (F. pa/mata, F. platyphylla, F. populifolia, and F. vasta) are confined to these regions. F. cordata subsp. lecardii (very closely related to F. cordata subsp. cordata of SW. Mrica) and the 'gnaphalocarpa' form of F. sycomorus (the only form occurring in Sw. Mrica) are typical for the western part of the Soudanian region. The

49 distribution of these two entities might indicate the occurrence of a (historical) link between the two regions. The other species extend to southem Africa, only one of them (F. abutilifolia) disjunctly, with a slightly different forrn. Six (out of the 23) species are elements of the Zambezian region; one (F. pygmaea) is confined to the Kalahari (transitional) region; two species (F. muelleriana and F. tettensis) are confined to eastem Southem Africa and one (F. ilicina) is an element of the Karoo-Namib region. Two taxa occur in the Sahara region: F. ingens and F. corclata subsp. sali­ cifolia. Two of the continental African species are more or less distinctly transitional with regard to the vegetations in which they occur: F. sur and F. thonningii. For the last-mentioned species a very broad (possibly too broad) species concept has been adopted. Two species (F. oreodryadum and F. oresbia) are montane. In addition to these two species three montane subspecies can be recognized (F. chlamydocarpa subsp. chlamydocarpa, F. polita subsp. brevipedunculata, and F. tremuia subsp. acuta). Some species occurring in relatively dry regions are distinctly associated with habitats like banks of streams and lakes and/or marshes (F. verruculosa, F. pygmaea, F. capreifolia, and F. trichopoda). Such an association is, among species occurring in humid climatical conditions, only clear for the terrestrial species F. asperifolia and F. vogeliana; it might less clearly occur in some hemi­ epiphytic species (e.g., F. cyathistipula) or 'ecotypes' (e.g., of F. natalensis subsp. leprieurii in Gabon) as weU. Somewhat less than 1/3 of the species of the African continent are associated with more or less dry climatical conditions (woodland to semi-desert). All sec­ tions are represented in this group of species. Only one of the five subsections of section Galogly chia (Berg, 1986b), i.e., subsection Platyphyllae, is with most of its 18 species weU represented in this group, the other subsections hardly or not. This strong representation of subsec­ tion Platyphyllae can be related to some morphological features common in the subsection: long-petiolate leaves, cordiforrn to reniform blades, the dense indu­ ment often occurring on leaves and other parts, and the chartaceous texture of the leaf blade. The region comprising Madagascar, the Comoro Islands, the Aldabra Islands, the Seychelles and the Mascarene Islands comprises 28 species, of which four are shared with the continent: F. lutea (Madagascar, Comoro Islands, Aldabra Island, and the Seychelles), F. polita (Madagascar), F. sycomorus (Mad­ agascar and Comoro Islands), and F. trichopoda (Madagascar). About %of the species are associated with (relatively) humid vegetation, and the rest with more of Ie ss dry vegetation (the proportions being roughly similar to those on the African continent). Madagascar has 24 species of which 14 are endemic; the Comoro Islands bas eight species, one of them (F. karthalensis) being endemic, the Mascarene Islands has five species, three of them (F. densifolia, F. lateriflora, and F. mauritiana) being endemic, the Aldabra Islands has three wide-spread

50 species, and the Seychelles have four species with a wide distribution; only F reflexa subsp. sechellensis is endemic.

Diagnosis of the genus Ficus Trees, shrubs or sometimes lianas, terrestrial or hemi-epiphytic (and then with aerial roots, often strangling and secondarily terrestriai), monoecious or (gyno)dioecious. Leaves in spirals, distichous, subopposite or subverticillate; lamina pinnately veined with glandular (sometimes waxy) spots in the axils of at least the basal lateral veins beneath or (mostly a single spot) at the base of the midrib beneath; stipules fully amplexicaul, semi-amplexicaul or lateral, free or partly connate. Inflorescences ('figs' or 'syconia') solitary or in pairs in the leaf-axils, mostly several together on small spurs in the leaf-axils down to the lesser branches or on distinct spurs on the lesser branches down to the trunk, or on leafless branchlets on the older wood down to the base of the trunk; figs consist of an urceolate receptacle with a ± narrow apical opening ('ostiole'): it encloses the fiowers; figs are functionally bisexual, containing staminate flowers, long-styled fiowers (destined to produce seeds) and short-styled flowers (destined to hatch the larvae of the pollinator) or occur on one plant as 'gall figs' containing starninate flowers and short-styled flowers (which do not produce seeds but are used as breeding sites for the pollinator) and on another plant as 'seeds figs' containing only long-styled (and fully functional) pistillate flowers; bracts 2- 3(-4) on the peduncle (peduncular bracts) or 2 or 3 subtending the receptacle (basal bracts), sometimes on the outer surface of the receptacle (lateral bracts), numerous bracts in the ostiole (ostiolar bracts), and often bracts among the fiowers (interfloral bracts). Staminate fiowers near the ostiole or dipersed, mostly pedicellate; tepals 2-6, free or connate; stamens 1 or 1-3; pistillode absent or sometimes present. Pistillate flowers sessile or pedicellate; tepals 2-6(-7), free or connate; ovary free; stigmas 1, filiform or (sub)infundibuliform, (sub)clavate or tongue-shaped, or 2 and ± filiform, or subulate, stigmas cohering or not. Pis til late flowers in (gyno )dioecious taxa distincly differentiated in long-styled and short-styled flowers and in monoecious taxa (in which all pistillate flowers can set seed) intermediate style lengths often occur. Fruits (see fig. 12) achene­ like or often ± drupaceous, releasing the endocarp-body (pyrene) or not; at fruit the wall of the fig mostly ± fleshy, coloured or green; gall fruits achene-like.

KEY TO THE SECTIONS

I. Ostiole slit-shaped (or occasionally triradiate); all ostiolar bracts descending; basal bracts in a whorl of 2 (occasionally 3); staminate flowers always disperse ...... section Galoglychia - Ostiole circular with at least 3 ostiolar bracts visible; only the lower ostiolar bracts descending; basal bracts in a whorl of 3 or bracts scattered on the pedunde; staminate flowers mostly near the ostiole ...... 2

51 2. Waxy gJanduJar spots on the lower surface of the larnina in (at least) the axils of the (main) basal lateral veins (only in F pachyclada partlyon the midrib); mostly several, more than 3 ostiolar bracts visible; leaf margin often dentate to crenate . . . . 3 - Waxy gJandular spot(s) on tbe lower surface of the larnina on the base of the midrib; mostly only 3 ostiolar bracts visible; leaf margin (of adult plants) entire ...... 5

3. Plants monoecious (all figs with starninate flowers and both long- and short-styled pistillate flowers); starninate flowers subtended by 2 bract(eole)s; lamina always equal­ sided and never lobed to fid ...... section Sycomorus - Plants (gyno)dioecious (there are 'seed figs' with only long-styled pistillate flowers and 'gall figs' with both staminate and short-styled pistillate flowers); starninate flowers not subtended by bract(eole )s; larnina often unequal-sided and/or lobed to fid...... 4

4. StipuJes fully amplexicaul; usually with 3 basal bracts in a whorl; starninate flowers without a pistillode; lamina equal-sided ...... section Ficus - StipuJes not fully amplexicaul; usually without distinct basal bracts but some bracts scattered on the peduncle; starninate flowers with a pistillode; larnina often unequal­ sided ...... section Sycidium

5. Lenticels concentrated in the uppermost part of the internodes; 2 stigmas ...... section Oreosycea - Lenticels usually scattered on the internodes; (usually) I stigma . . . section Urostigma

KEYS TO THE SPECIES In order to make the keys not too complicated, the species of the African con­ tinent (and the islands in the Gulf of Guinea) and those of Madagascar and neighbouring Indian Ocean islands are keyed out separately. The introduced species (see p. 166-167) are not included in the keys.

I. Key to the species of sec!. Sycidium

A. Continent

I. Larnina unequal-sided (asymmetricaJ) ...... 5. F asperifolia - Larnina equal-sided (symmetrical) ...... 2

2. La teral veins 3- 5 pairs; trees ...... 2. F exasperata - Lateral veins up to 8 or 12 pairs; shrubs . 3

3. Leaves usually partly subopposite or subverticillate; larnina often with a subentire to faintly crenate margin and a 3-dentate (to 3-lobed) apex; stipules 0.5-1 cm long ...... 6. F capreifolia - Leaves always alternate; lamina irregularly serrate-dentate to pinnately fid; stipules 0.2-0.6 cm long ...... 7. F pygmaea

B. Madagascar

I. Larnina hirtellous (to puberulous) beneath . 2 - Larnina hispidulous beneath ...... 6

2. Hairs on the leafy twigs (partly) brown with ± swollen bases; gJanduJar spots (for the greater part) on the midrib beneath ...... 9. F brachyclada Hairs on the leafy twigs white or, if brownish, then without swollen bases; gJandular spots in the axils of the (main) basal lateral veins beneath ...... 3

52 3. Base of the lamina cordate or subcordate andlor the petiole 1.5-7(- 13) cm long . 4 - Base of the lamina acute to obtuse or the petiole up to I cm long...... 5

4. Stipules (0.3- )0.5-1.5 cm long; Réunion ...... 4. F lateriflora - Stipules 0.1-0.5 cm long; Madagascar (and Comoro Islands?) . ... . 3. F pachyclada

5. Lamina usually up to 10 cm long or, if longer, then mostly sublinear or lobed; figs when dry 0.5-0.8(- 1) cm in diameter, usually in the leafaxils or just below the leaves . .. . · ...... 10. F politoria - Lamina usually subobovate to obovate; glandular spots (for the greater part) on the midrib beneath; figs when dry 0.8--1.2 cm in diameter, often on spur-Iike branchlets (already forrned in the leafaxils) on the older wood...... 9. F brachyclada

6. Stipules (0.3--)0.5-1.5 cm long; Réunion...... 4. F lateriflora - Stipules 0.1-0.5 cm long; other islands ...... 7

7. Larnina usually up to 10 cm long, if longer, then sublinear or lobed; gall fruits stipitate · ...... 10. F politoria - Larnina usually longer than 10 cm, neither sublinear nor lobed; gall fruits non-stipitate · ...... 8. F bojeri

11. Key to the species of section Sycomorus

A. Continent

I. Figs solitary in the leafaxils (or just below the leaves) ...... 2 - Figs on leafless branchlets on the older wood, often down to the trunk . 3

2. Larnina scabrous above; hairs on the petiole distinctly different in length . · ...... 11. F sycomorus - Lamina smooth above; hairs on the petiole not distinctly different in length ...... · ...... 15. F. vallis-choudae

3. Stipules subpersistent; basal bracts 3--5 mrn longs; trees geocarpic . . 14. F vogeliana - Stipules caducous; basal bracts 2-3 cm long; trees not geocarpic . . 4

4. Hairs of leafy twig and petiole conspicuously different in length; periderm of leafy twigs and epidermis of petiole flaking off ...... 5 - Hairs of leafy twig and petiole not conspicuously different in length (or absent); peri­ derm of leafy twigs and epidermis of petiole usually not flaking off...... 13. F sur

5. Apex of lamina rounded to obtuse; petiole up to 4(-6) cm long; trunk short ...... · ...... 11 . F sycomorus - Apex of lamina acuminate to acute; petiole up to 9 cm long; trunk tall ...... · ...... 12. F mucuso

B. Madagascar

I. Indument of the leafy twigs (partly) brown ...... 18. F torrentium - Indument of the leafy twigs whitish or yellowish ...... 2

2. Basal lateral veins branched . . 3 - Basal lateral veins unbranched 6

3. Stipules glabrous, sparsely puberulous or ciliolate ...... 4 - Stipules subsericeous to subhirsute ...... 5

4. Figs 0.5- 1 cm in diameter when dry; petiole up to 2(- 2.5) cm long . . . . · ...... 22. F karthalensis - Figs 1- 1.5 cm in diameter when dry; petiole up to 13 cm long .. . . 16. F mauriliana

53 5. Apex of lamina short-acwninate to subacute; petiole up to 9(- 13) cm long; figs in the leafaxils or just below the leaves, sometimes also on branchlets on the older wood, 1.5-2.5 cm in diameter when dry ...... 17. F. tiliifolia - Apex of lamina rounded; petiole up to 3 cm long; figs on branchlets on the older wood, if in the leafaxils or just below the leaves, then when dry 4-5 cm in diameter . . . . . · ...... 11. F. sycomorus

6. Figs when dry 0.8- 1 cm in diameter, only 3 or 4 ostiolar bracts visible ...... · ...... 22. F. karthalenis - Figs usually 1.5-3.5 cm in diameter when dry, more than 4 ostiolar bracts visible . .. 7

7. Figs subsessile or up to 0.4 cm long pedunculate; a few large lenticelIs in the upper part of the intemodes ...... 19. F. polyphlebia - Figs on peduncles at least 0.4 cm long; such large lenticelIs absent ...... 8

8. Lateral veins 4-6 pairs; petiole up to 13 cm long; Mauritius and Réunion . · ...... 16. F. mauritiana - Lateral veins (5-)7-13(-15) pairs; petiole up to 3 cm long; Madagascar...... 9

9. Basal bracts ca. 4 mm long; peduncle (0.7- )1-8 cm long ...... 20. F. botryoides - Basal bracts 1-2 mrn long; peduncle 0.4-0.8 cm long ...... 21. F. trichoc/ada

111. Key to the species of section Oreosycea

A. Continent

I. Lateral veins (5-)8- 13 pairs; hunina usually smooth above ...... 23 . F. variifolia - Lateral veins 5- 8(- 9); lamina scabrous above ...... 24. F. dicranostyla

B. Madagascar

I. Apex of lamina acute to faintly acuminate; lateral veins 7- 14 pairs .. . 25. F. assimilis - Apex of lamina rounded to obtuse or very shortly and obtusely acwninate; lateral veins &-8 pairs ...... 26. F. ampana

IV. Key to the species of sections Urostigma and Conosycea

A. Continent

I. Figs sessile (or subsessiie) . . 28. F. cordata - Figs pedunculate ...... 2

2. Shrubs (growing along streams and in marshy pi aces) . . .29. F. verruculosa - Trees (of dry habitats) ...... 3

3. Basal lateral veins usually branched, almost straight (not running parallel to the leaf margin), the other lateral veins often dividing far from the margin; figs when fresh 1- 2 cm in diameter, when dry wall mostly wrinkled ...... 27. F. ingens - Basal lateral veins unbranched, ± curved, running parallel to the leaf margin, the other lateral veins dividing not far from the margin; figs when fresh 0.5-1 cm in diameter, when dry wall usually smooth ...... 28. F. cordata

B. Madagascar

I. Figs pedunculate . . 33. F. humbertii - Figs sessile ...... 2

2. Lateral veins (3--)12- 22 pairs; petiole up to 2 cm long ...... 32. F. menabeensis - Lateral veins 7-12 pairs; petiole up to 5 cm long ...... 3

54 3. Stipules 0.5--1.5(- 8) cm long; figs 0.5--1 cm in diameter when dry; Mauritius and Réunion ...... 31. F densifolia - Stipules 0.3--0.5 cm long; figs ca. 0.5 cm in diameter when dry; Madagascar ...... 30. F madagascariensis V. Key to the species of section Galoglychia

A. Continent

I. Lamina constricted below the rniddle (± fiddle-shaped, pandurate) . 2 - Larnina not (distinctly) constricted in the lower half ...... 6

2. Stipules caducous 3 - Stipules persistent 4

3. Lamina 10-17 x 2-5 cm ...... 94. F crassicosta - Larnina (15--)20--60 x 6-25 cm ...... 92. F wildemaniana

4. Lateral veins 6-7 pairs ...... 89. F lyrata - Lateral veins at least 9 pairs ...... 5

5. Larnina at the base (usually) with a coarsely dentate margin; figs ca. 1.5 cm in diameter when dry ...... 90. F sagittifolia - Larnina at the base with an entire margin; figs 2.5--3.5 cm in diameter when dry ...... 91. F subsagittifolia

6. Stipules andJof parts ofaxillary calyptrate bud covers (enclosing pairs of young figs) persistent (or subpersistent) ...... 7 - Stipules andJor parts ofaxillary calyptrate bud covers caducous (- on flush stipules may be subpersistent) ...... 32

7. Tertiary leaf venation partly scalariform; leafy twigs, stipules andJor lower leaf surface partly hirsute ...... 34. F saussureana - Tertiary leaf venation reticulate or almost parallel to the lateral veins; leafy twigs, stipules andJor lower leaf surface glabrous, puberulous, or at most short-pubescent . 8

8. Stipules up to 8 cm long ...... 9 Stipules up to 3 cm long (- on flush up to 6 cm long) ...... 10

9. Figs ca. 1- 1.5 cm in diameter when dry, largely concealed by the stipules; apex of the receptacle crateriform andJor swollen ...... 81. F abscondita - Figs ca. 2.5--3.5 cm in diameter, not concealed by the stipules; apex of the receptacle rounded ...... 91. F subsagiltifolia

10. Stipules and axillary calyptrate bud covers densely hairy inside .. . . 57. F amadiensis - Stipules and axillary calyptrate bud covers glabrous inside . II

11. Figs sessile ...... 12 - Figs pedunculate ...... 25

12. Figs with a distinctly stipitate base ...... 86. F densistipulata - Figs with a broad or at most slightly narrowed base . 13 13. Figs when dry up to ca. I cm in diameter . . . 14 - Figs when dry at least ca. 1.5 cm in diameter . 21

14. Stipules 1- 2 cm long; petiole when dry 2-2.5 mm thick . 15 - Stipules up to 1 cm long (often only subpersistent); petiole when 1.5--2 mm thick . 17 15. Lateral veins 8-11 pairs; petiole when dry 1- 2 mm thick ...... 76. F conraui - Lateral veins 6-8 pairs; petiole when dry 2- 2.5 mm thick ...... 16

55 16. Lateral veins in the lower part of the larnina distinctly weaker than those in the middle part of the lamina ...... 78. F. pachyneura - Lateral veins in the lower part of the larnina about equally thick as those in the midde part of the lamina ...... 77. F. tesse/ata

17. Midrib not reaching the apex ofthe lamina...... 18 - Midrib reaching the apex of the larnina ...... 19

18. Petiole up to 2 cm long; ligs when dry ca. 0.5 cm in diameter; young ligs in up to I cm long axillary calyptrate bud covers ...... 58. F. eraterostoma - Petiole up to 0.8 cm long; ligs when dry 0.3-D.4 cm in diameter; young ligs not in well-developed axillary bud covers ...... 59. F. lingua

19. Well-developed lateral veins arising from the midrib 0.5-2 cm from the base of the lamina ...... 79. F. ardisioides - Well-developed lateral veins arising from the midrib at or near the base of the larnina 20

20. Lamina when dry brown, often folded, at least the margin involute; apex of larnina mostly distinctly acuminate ...... 67. F. kamerunensis - Larnina when dry mostly greyish-green to brown, including the margin plane; apex of larnina, if acuminate, then faintly andlor shortly so ...... 66. F. thonningii

21. Petiole ca. 4 mm thick when dry; basal bracts ca. 5 mm long; lamina mostly 20-30 cm long ...... 80. F. preussii - Petiole up to 3 mm thick when dry; basal bracts at most 4 mm long; larnina mostly 10-20 cm long ...... 22

22. Larnina usually (sub)ovate and with a (sub)cordate base ...... 87. F. subcostata - Larnina usually broadest in or above the middle and with an acute to obtuse, sometimes to truncate base ...... 23

23. Lateral veins 8-11 pairs. .76. F. eonraui - Lateral veins 5-7(- 8) pairs...... 24

24. Wall of the lig spongy; basal bracts ca. 4 mm long ...... 82. F. eyathistipu/a - Wall of the lig firm; basal bracts ca. 2 mm long...... 77. F. tesse/ata

25. Stipules 1.5- 3.5 cm long, connate ...... 26 - Stipules up to 1(-1.5) cm long, free ...... 27

26. Peduncle up to 0.4 cm long; ligs distinctly stipitate; wall of lig when dry 0.5-1 mm thick, not spongy ...... 86. F. densistipu/ata - Peduncle (usually) 0.5-2.5 cm long; figs with a broad base or slightly stipitate; wall of fig when dry 2- 5 mm thick, spongy ...... 82. F. eyathistipu/a

27. Midrib not reaching the apex of the lamina ...... 28 - Midrib reaching the apex of the lamina ...... 29

28. Peduncle (0.3-)0.5-1.5 cm long; figs when dry 0.4-û.8(- 1.2) cm in diameter ...... 61. F. fau/kneriana - Peduncle up to 0.5 cm long; figs when dry 0.3-D.4 cm in diameter .. . . . 59. F. lingua

29. Figs when dry 2- 3 cm in diameter ...... 82. F. cyathistipu/a - Figs when dry up to 1.5 cm in diameter ...... 30

30. Well-developed lateral veins arising from the midrib 0.5-2 cm from the base of the lamina ...... 79. F. ardisioides - Well-developed lateral veins arising from the midrib at or near the base of the larnina 31

56 31. Lamina when dry brown, folded or at least the margin involute; apex of lamina mostly distinctly acuminate ...... 67. F. kamerunensis - Lamina when dry greyish-green to brown, inc1uding the margin plane; apex of lamina, if acuminate, then faintly andJor shortly so . . 66. F. thonningii

32. Lamina distinctly hairy beneath ...... 33 - Lamina glabrous or at most sparsely puberulous beneath ...... 51

33. 8asal lateral veins reaching the margin above (or sometimes at) middle of the lamina 34 - 8asal lateral veins reaching the margin (far) below the middle of the larnina . 38

34. Epidennis of the petiole ftaking off . . . 35 - Epidennis of the petiole not fiaking off . 36

35. Stipules 2- 5 cm long (on ftush up to 8 cm long); basal bracts caducous . . 40. F. vasta - Stipules 0.5-1.5 cm long (on ftush up to 4 cm long); basal bracts persistent ...... · ...... 41. F. wakefie/dii

36. Lateral veins 7-9 pairs; petiole up to 10(- 18) cm long ...... 47. F. abutilifo/ia - Lateral veins 3--7 pairs; petiole up to 4(--8) cm long ...... 37

37. Hairs on leafy twigs prominently different in length, the longer ones usually yellowish; stipules 0.5- 1.5 cm (on ftush up to 4 cm) long ...... 42. F. g/umosa - Hairs on leafy twigs not c1early different in length, whitish; stipules 0.2-D.6 cm long · ...... 45. F. tettensis

38. Epidennis of the petiole ftaking off . 39 - Epidennis of the petiole not fiaking off . 41

39. Lateral veins 6--8(- 10) pairs; figs when dry usually less than 1.5 cm in diameter . · ...... 36. F. /utea - Lateral veins 8--20 pairs; figs when dry more than 1.5 cm in diameter ...... 40

40. Stipules up to 8(- 12) cm long; apex of the lamina distinctly acuminate, base mostly acute to obtuse ...... 34. F. saussureana - Stipules up to 2.5 cm long; apex of the lamina usually faintly and shortly acuminate, base mostly subcordate to truncate ...... 45. F. ch/amydocarpa

41. Figs (usuaUy) distinctly longer than wide and initially enc10sed by up to 2.5 cm long calyptrate axillary bud covers, leaving conspicuous triangular scars . . . . 105. F. ovata - Figs (usually) globose and axillary buds minute or absent...... 42

42. Lateral veins 3--7 (or 8) pairs ...... 43 - Lateral veins at least 8 (or 7) pairs ...... 47

43. Figs when dry (0.7- )1 - 1.8 cm in diameter; vein-reticulum of lamina prominent beneath; periderm of leafy twigs ftaking off; indument whitish to pale yellow ...... · ...... 43. F. s tuh/mannii - Figs when dry 0.5- 1 cm in diameter; vein-reticulurn of lamina not prominent beneath; periderm of leafy twigs not ftaking off or indurnent partly bright yellow...... 44

44. Hairs on leafy twigs prominently different in length, the longer hairs usually yellow; periderm of leafy twig ftaking off ...... 42. F. g/umosa - Hairs of leafy twigs not c1early different in length; indurnent whitish; periderm of leafy twigs not fiaking off ...... 45

45. Petioles of leaves on the same twig quite varia bie in length independent of size of the lamina and position on the twig ...... 66. F. thonningii - Petioles of leaves on the same twig ± equal in length, or if different, then the length proportional to the size of the larnina...... 46

57 46. Petiole (1.2-)3.5-5.5 cm long, 1- 1.5 mrn thick; leaf margin entire; leafy twigs when dry red-brown ...... 46. F. muelleriana - Petiole 0.3-2 cm long, 0.5-1 mrn thick; leaf margin crenate; leafy twigs when dry usually blackish with conspicuous whitish lenticels ...... 44. F. nigropunctata

47. Peduncle at least I cm long ...... 48 - Peduncle at most I cm long ...... 50

48. Stipules densely hairy ...... 37. F. platyphylla - Stipules glabrous or subglabrous ...... 49

49. Stipules 0.3-1.2 cm (on ftush up to 3 cm) long . . 38. F. bussei - Stipules 1-2.5 cm (on ftush up to 4.5 cm) long ...... 39. F. recurvata

50. Stipules 1.5-4.5 cm (on ftush up to 8 cm) long ...... 50. F. trichopoda - Stipules up to I (- I. 5) cm long...... 66. F. thonningii

51. Stipules and/or axillary bud covers (enclosing young figs) hairy inside ...... 52 - Stipules and, if present, axillary bud covers glabrous inside ...... 54

52. Lamina usually slightly longer than wide ...... 55. F. calyptrata - Lamina usaIly 2 x as long as wide ...... 53

53. Stipules 0.5-1.5 cm long; lamina (4-)7-17 x (2-)3-8.5 cm ...... 57. F. amadiensis - Stipules ca. 0.5 cm long; lamina 3-8 x 2-4.5 cm ...... 58. F. craterostoma 54. Epidermis of petiole ftaking oIT...... 55 - Epidermis of petiole not ftaking oIT...... 58

55. Figs when dry less (usually) less than 1.5 cm in diameter; lateral veins up to 10 pairs 56 - Figs when dry usuaIly more than 1.5 cm in diameter; lateraI veins mostly more than 10 pairs ...... 57

56. Base of lamina usally cordate to truncate; basaI bracts ca. 1.5 mrn long...... 75. F. leonensis - Base of lamina usually obtuse to acute; basal bracts 3-6 mrn long . . . . . 36. F. lutea

57. Stipules up to 8(- 12) cm long; apex of lamina usally distinctly acuminate, base acute to obtuse ...... 34. F. saussureana - Stipules up to 2.5 cm long; apex of lamina usually faintly and/or shortly acuminate, base mostly subcordate to rounded ...... 35. F. chlamydocarpa

58. Basal lateral veins usually reaching the margin above the middle of the lamina ...... 47. F. abutilifolia - Basal lateral veins usuaIly reaching the margin (far) below the middle of the lamina 59

59. Figs in pairs in the leafaxils or somtimes below the leaves (on previous season's growth), if more than 2 together, then (aIrnost) confined to the leafaxils; figs sessile or pedunculate ...... 60 - Figs usually more than 2 together on ± distinct spurs below the leaves, often on older wood (down to the trunk) but sometimes also in the leafaxils; figs pedunculate (but sessile in F. adolfi-friderici and sometimes in F. louisii) ...... 94 60. Figs sessile or subsessile (peduncle up to I mrn long for figs up to I cm in diameter or up to 2 mrn long for larger figs) ...... 61 - Figs pedunculate ...... 72

61. Figs when dry ca. 1.5 cm or more, rarely down to 1.2 cm in diameter. . . 62 - Figs when dry up to ca. I cm in diameter ...... 67

58 62. Stipules up to 1(- 1.5) cm long; lamina usuaIly up to 10 cm long .... 66. F. thonningii - Stipules at least ca. 1.5 cm long andJor lamina usually longer than 10 cm . 63 63 . Figs when dry 2.5-4.5 cm in diameter; lamina usually 20-{j() cm long ...... 92. F. wildemaniana - Figs up to ca 2. cm in diameter, or if more than 2 cm, then the lamina usuaIly 10--20 cm long ...... 64 64. Base of lamina usuaIly cordate to truncate; figs with ± dense rigid hairs ...... 87. F. subcostata - Base of lamina acute to obtuse, if subcordate then figs subglabrous ...... 65

65. LateraI veins 7-8 pairs ...... 93. F. oresbia - LateraI veins 8-18 pairs ...... 66

66. Petiole 1.5-2 mrn thick; figs warty and with a plane apex ...... 70. F. oreodryadum - Petiole 2- 3 mrn thick; figs smooth and with a ± protracted apex . . . 83. F. scassellatii 67. Midrib not reaching the apex of the lamina . 68 - Midrib reaching the apex of the lamina . . . 69 68. Petiole up to 2 cm long; figs when dry ca. 0.5 cm in diameter; young figs in up to I cm long axillary caIyptrate bud covers ...... 58. F. craterostoma - Petiole up to 0.8 cm long; figs when dry 0.3-û.4 cm in diameter; young figs not in distinct axillary bud covers ...... 59. F. lingua 69. Lateral veins 3-7(-8) pairs . . 70 - Lateral veins at least 7 pairs . 71

70. Stipules glabrous ...... 79. F. ardisioides - Stipules hairy ...... 42. F. glumosa 71. Lamina when dry brown, folded, or at least the margin involute; apex of lamina mostly distinctly acuminate ...... 67. F. kamerunensis - Lamina when dry greyish-green to brown, including the margin plane; apex of lamina, if acuminate, then faintly andJor shortly sa . . 66. F. thonningii 72. BasaI bracts (their free parts!) caducous . 73 - Basal bracts persisten t ...... 79 73. Midrib reaching the apex of the lamina . 74 - Midrib usually not reaching the apex of the lamina ...... 75

74. Figs when dry 0.5-1.5 cm in diameter; lamina 2.5-10 cm long . . 60. F. natalensis - Figs when dry 0.3-û.4 cm in diameter; lamina 0.5-5 cm long . . 59. F. lingua 75. Base of lamina usually cordate to truncate . 76 - Base of lamina usually acute to obtuse . . . 77

76. Stipules 1-4 cm long; lateral veins 6-11 pairs ...... 48. F. populifolia - Stipules 0.3-D.8 cm long; lateral veins 9-15 pairs ...... 56. F. fischeri

77. Stipules 0.5-1 cm long; lateraI veins 10--15 pairs ...... 69. F. burretiana - Stipules 1- 2.5 cm long; lateral veins 10--20 pairs ...... 78

78. Figs when dry 0.5-1 cm in diameter; peduncle (0.5-)1-2.5 cm long ..... 88. F. barteri - Figs when dry ca. 1.5 cm in diameter; peduncle 0.5-1 cm long ... . 68. F. elasticoides 79. Peduncle at least I cm long ...... 80 - Peduncle at most I cm long ...... 82

59 80. Stipules densely hairy ...... 37. F. platyphylla - Stipules gJabrous or only at the base hairy ...... 81

81. Stipules 0.3-1.2 cm (on ftush up to 3 cm) long ...... 38. F. bussei - Stipules 1- 2.5 cm (on ftush up to 4.5 cm) long ...... 39. F. recurvata

82. Figs when dry ca. 1.5 or more (but sometimes down to I cm) in diameter . 83 - Figs when dry up to I cm in diameter...... 88

83. Stipules densely hairy, up to 4.5 cm (on ftush up to 8 cm) long .. .. 50. F. trichopoda - Stipules gJabrous or sparsely bairy, at most 2 cm long . 84

84. Lateral veins 6-10 pairs ...... 85 - Lateral veins (normally) 10-20 pairs...... 86

85. Stipules up to 2 cm long; figs ± stipitate and with protruding apex . . . . . · ...... 84. F. cyathistipuloides - Stipules up to 1 cm long; figs with broader bases and plane to slightly protruding apex · ...... 85. F. scott-e/liotii

86. Base of lamina cordate ...... 94. F. crassicosta - Base of lamina acute to obtuse ...... 87

87. Tertiary venation reticulate; apex of fig protruding ...... 83. F. scassellatii - Tertiary venation largely parallel to the lateral veins; apex of fig plane ...... · ...... 68. F. elasticoides

88. Stipules 1.5--4.5 cm (on ftush up to 8 cm) long...... 50. F. trichopoda - Stipules up to I cm (sometimes up to 1.5 cm) long . 89

89. Midrib not reaching the apex of the lamina . 90 - Midrib reaching the apex of the larnina . . . 91

90. Peduncle (0.3- )0.5- 1.5 cm long; figs when dry 0.4-0.8(- 1.2) cm in diameter. · ...... 61. F. faulkneriana - Peduncle up to 0.5 cm long; figs when dry 0.3-0.4 cm in diameter . .. . 59. F. lingua 91. Lamina (at least when dry) ± revolute, especially towards the base . 62. F. burtt-davyi - Lamina with aplane margin ...... 92

92. First well-developed lateral veins 0.5-2 cm from the base of the lamina . · ...... 79. F. ardisioides - First well-developement at or near the base of the la mina ...... 93

93. Petioles of leaves on the same twig about equally long or, if unequally long, then length proportional to size of the lamina (SW. Africa) ...... 63. F. ilicina - Petioles of leaves on the same twigs usually quite different and independent of size of the lamina and position on the twig ...... 66. F. tllOnningii

94. Fig-bearing short-shoots minute; figs often also in the leafaxils; figs when Jry up to I cm in diameter ...... 95 - Fig-bearing short-shoots well-developed, mostly becoming 0.5 cm or more in diameter; figs occasionally also in the leafaxils; figs when dry at least I cm in diameter . 99

95. Figs sessile ...... 96 - Figs pedunculate ...... 97

96. Lamina usually less than 10 cm long; petiole ca. 1.5 mm thick . . 74. F. louisii - Lamina usally more than 10 cm long; petiole 1.5-2.5 cm long ... 73. F. adolfi-friderici

60 97. Lateral veins 14-27 pairs; stipules greyish to yellowish to brown strigose to sub­ sericeous ...... 71. F. pseudomangifera - Lateral veins up to 14 pairs; stipules g1abrous or puherulous ...... 98

98. Basal bracts persistent; stipules up to 0.5 cm long ...... 72. F. usambarensis - Basal bracts (their free parts) caducous; stipules up to I cm long ...... 49. F. jansii

99. Basal bracts persistent (or subpersistent) ...... 100 - Basal bracts caducous ...... 104

100. Figs usually ellipsoid, when dry 2-3 cm in diameter and the wall 3-7 cm thick; base of lamina acute to subattenuate ...... 99. F. bizanae - Figs g1obose, if ellipsoid, then when dry at most 2 cm in diameter, the wall up to 2 mm thick and/or base of lamina cordate to rounded ...... 101

101. Petiole 1-2 mm thick; bud sca1es of (fig-hearing) spurs (sub)g1abrous ...... 102 - Petiole 2-4(- 5) mm thick; bud scales of (fig-hearing) spurs puherulous or with a ru- minate surface ...... 103

102. Basal bracts 3-5 mm thck, coriaceous; petiole up to 12 cm long; figs when dry 1.4-4 cm in diameter ...... 98. F. polita - Basal bracts 2- 3 mm long, chartaceous; petiole up to 4(- 5.5) cm long; figs when dry (1-)1.2-1.8 cm in diameter ...... 95e. F. ottoniifolia subsp. ulugurensis

103. Lamina usually broadest at (or above) the middle, usually drying greyish-green; base of lamina mostly (sub)cordate ...... 104. F. bubu - Lamina usually broadest helow the middle, usually drying brown; base of lamina truncate to rounded (or subcordate with a wide sinus) ...... 103. F. umbelIata

104. Figs ellipsoid or if (sub)g1obose, then at most 1.8 cm in diameter when dry; wallof fig usually not wrinkled when dry; bud scales of (fig-hearing) spurs (sub)g1abrous . . . . . 105 - Figs (sub)g1obose, usually more than 2 cm in diameter when dry; wallof fig when dry usually wrinkled; bud sca1es of (fig-hearing) spurs ± densely hairy (except in F. chi- rindensis) ...... 106

105. Petiole up to 9 cm long and/or lamina 6.5-15(-22) cm long ...... 95. F. ottoniifolia - Petiole up to 3(-4) cm long and lamina 2.5-8 cm long ...... 96. F. tremuIa 106. Bud scales of (fig-hearing) spurs subglabrous; stipules with white appressed hairs; base of lamina usually cordate to truncate ...... 100. F. chirindensis - Bud scales of (fig-hearing) spurs densely hairy; stipules g1abrous or only ciliolate, or if minutely puherulous, then base of lamina usually acute to obtuse ...... 107

107. Lateral veins 10-16 pairs; (fig-hearing) spurs curved ...... 97. F. artocarpoides - Lateral veins usually up to 10 pairs (in juvenile leaves sometimes up to 14 pairs); (fig-hearing) spurs usually straight or ± cushion-shaped ...... 108

108. Lateral veins (in leaves of adult specimens!) loop-connected 1-2 mm from the margin; petioles usually drying (red-)brown; figs whitish puherulous ..... 101. F. sansibarica - Lateral veins loop-connected 3-8(- 10) mm from the margin; petioles usually drying blackish; figs yellow to brownish puherulous to hirtellous (densely so near the ostiole) ...... 102. F. dryepondtiana B. Madagascar

I. Figs ovoid to ellipsoid ...... 54. F. bivalvata - Figs (sub)g1obose to obovoid ...... 2

2. Figs sessile (or subsessiIe, peduncle at most 1.5 mm long) . 3 - Figs pedunculate (peduncle at least 2 mm long) ...... 5

61 3. Petiole (1.5--)4-13(- 17) cm long, its epidemis flaking 01T ...... 36. F lutea - Petiole 0.3-3(-4) cm long, its epidennis persistent ...... 4

4. Stipules 0.2-D.5 cm long, caducous; lamina usually elliptic to obovate; figs when dry 0.8-1.3 cm in diameter ...... 64. F antandronarum - Stipules 0.5--2 cm long, usually subpersistent; lamina usually oblong to subobovate; figs when dry 0.5-D.8 cm in diameter ...... 65 F reflexa

5. Figs on spurs on the older wood, when dry (1 - )2-4 cm in diameter . . .. 98. F polita - Figs in the leafaxils or just below the leaves, when dry up to 1.5 cm in diameter . .. 6

6. Basal bracts caducous ...... 51. F grevei - Basal bracts persistent ...... 7

7. Stipules 0.2-D.5 cm long; petiole up to 2 cm long. . . 64. F antandronarum - Stipules 0.5--1.5 cm long; petiole up to 7 cm long . 8

8. Stipules densely hairy ...... 50. F trichopoda - Stipules glabrous ...... 9

9. Leafy twigs 3- 5 mm thick, g1abrous (or minutely puberulous); apex of lamina obtuse, subacute or subacuminate; figs when dry 0.6-1 cm in diameter, g1abrous (or minutely puberulous) ...... 52. F rubra - Leafy twigs 1.5- 3 mm thick, densely puberulous; apex of lamina ± distinctly and often long-acuminate; figs when dry 0,2-.0.8 cm in diameter, puberulous or hirtellous ...... 53. F marmorata

AFRICAN GROUPS AND SPECIES

Ficus L. subgenus Ficus section Ficus

Shrubs or trees, terrestrial (or sometimes epiphytic), (gyno)dioecious; sap rnilky or watery. Leaves in spirals to al most distichous, entire or lobate, margin entire to crenate to dentate, waxy glandular spots in the axils of the basal lateral veins beneath; stipules fully amplexicaul. Figs solitary (or in pairs) in the leafaxils or just below the leaves; basal bracts 3, usually in a whorl, bracts at the outer surface of the receptacle lacking; ostiole circular and several ostiolar bracts visible, the lower ostiolar bracts descending, interfloral bracts lacking, receptacle inside hairy; staminate flowers near the ostiole or also among the pistillate flowers, with 4-6 tepals and 2- 5 stamens, pistillode usually lacking; pistillate flowers with 4-6 tepals, stigma ± bifid; pedicels, tepals and styles hairy or glabrous; in 'seed figs' often 'neuter flowers' as substitutes of the starninate flowers; fruits drupaceous, endocarp body (often) released.

The section Ficus comprises about 60 species in the Asian mainland and Malesia; only F pa/mala extends to north-eastern tropical Africa. The section shows quite a diversity. F pa/mala has been ranked under subsection Ficus series Cariceae together with two other species, including F carica, by Corner (1965). The pollinators belong to the genus Blaslophaga Gravenhorst.

62 1. Ficus palmata Forssk. (fig. 15) Forsskäl, Ft. Aegypt.-Arab.: 179 (1775); Aweke, Meded. Landbouwhogeschool Wageningen: 48, fig. 12 (1979).

F. caricoides Roxb. (1832), F. forsskalii Vahl (1805), F. morifolia Forssk. (1775), non Lam. (1788), F. pa/mala var. morifo/ia (Forssk.) Warb. (l904b), F. pa/mala var. petitiana (A. Rich.) Fiori (19\Ob), F. pa/mala var. pseudocarica (Miq.) Warb. (l904b), F. pa/mala var. soma/ensis Warb. (l904b), F. pa/mala var. slipitala Warb. (l904b), F. petitiana A. Rich. (1851), F. pa/mala var. pseudocarica (Miq.) Fiori (19\Ob), F. pseudocarica Miq. (1848), F. pseudocarica var. lomenlosa A. Rich. (1851), F. pseudosy­ comorus Decne (1834).

Shrub up to 5 m tall. Leafy twigs 2--4 mm thick, minutely puberulous. Larnina ovate to cordiforrn, entire or 3-5-lobed to -fid, (2.5-)4-10 x (1.5- )28 cm, chartaceous, apex acute to subacurninate, base cordate to truncate (to subacute), margin entire to serrate-dentate; upper surface scabrous, hispidulous, lower surface scabrous to scabridulous to smooth, hispidulous to hirtellous to sub­ tomentose; lateral veins 5-8 pairs, tertiary venation partly scalariforrn; petiole 1--4.5(-12) cm long, hispidulous to puberulous to hirtellous to subtomentose; stipules 0.5-1.5 cm long, puberulous to pubsecent, caducous. Figs solitary (or in pairs) in the leafaxils or just bel ow the leaves; peduncle 0.3-1.7 cm long, often with a bract at its base; basal bracts 3, mostly in a whorl, semicircular to ovate, 1-2 mm long, puberuIous or only ciliolate; receptacle subglobose and up to 0.5 cm long, stipitate or subpyriforrn, when fresh 2- 3 cm, when dry 1-1.5 cm in diameter, puberulous to hispidulous or to tomentose, at maturity yellowish to greenish, often pinkish to purplish tinged, the ostiolar bracts (scarlet-)red to dark purple.

Distribution. Ethiopia to India and Nepal. Mostly in thickets along water courses, mostly at altitudes between 1700 and 2400 m. The species is very closely related to F. carica L.; the two are probably conspe­ cific. The pollinator is Blastophaga psenes (L.), also known as the pollinator of F. carica.

Ficus subgenus Ficus section Sycidium Miq. Trees or shrubs, terrestrial, (gyno)dioecious; sap watery. Leaves (almost) dis­ tichous (and altemate) or subopposite, margin often dentate, when juvenile often ± deeply pinnate1y or palmately incised; glandular spots in the axils of the basal lateral veins beneath (or in F. pachyclada on the base of the midrib), sometirnes also in the axils of other lateral veins; stipules lateral to semi-amplexicaul. Figs in the 1eafaxils or on the older wood, pedunculate, usually with a bract at the base of the peduncle and with 2--4, usually disperse (peduncular) bracts, several bracts on the outer surface of the receptacle, ostiole circular with several ostiolar bracts visible, only the lower ostiolar bracts descending; interfioral bracts lack­ ing, receptacle inside setose; staminate fiowers usually near the ostiole, with

63 Fig. IS. Ficus pa/mala Forssk. I, leafy twig with figs; 2, 1eaf; 3-4, staminate fiowers; 5, stamen; 6-9, pistillate fiowers; 10, pistil; 17, ostiole. After Aweke (1979, fig. 12).

(2- )3-6 free tepals, stamen(s) 1 (or 2), pistillode present; pistillate flowers with 1-6 free tepals, stigma 1, truncate and infundibuliform; pedicels, (margin of the)

64 tepals and (usually also the) style hairy; fruit drupaceous, endocarp body released; wall of the fig in fruit soft, red, orange, or yell ow.

Tbe section Sycidium comprises about 105 species, most of them in Asia­ Australasia. Four species accur in continental Africa. Tbe frutescent species occur along streams. It is not always easy to teil these species apart. The fourth species, F. exasperata, is arborescent and accurs in non-riparian habitats. Five species are recognized for Madagascar and neighbouring islands. The label data of the collections studied do not supply information from which differences in the ecology of these species are dear. The pollinators belong to the genus Kradibia Saunders.

2. Ficus exasperata Vahl Vahl, Enum. 2: 197 (1805); Berg et aI ., FI. Cameroun 28: 121 , fig. 39 (1985); Berg et aI, Fl. Trop. East Africa, Moraceae: 52 (1989).

F. punctifera Warb. (1904), F. scabra WiUd.(1801), F. silicea Sim (1909), F. serrata Forssk. (1775).

Shrub or tree up to 20 m taU. Leafy twigs 1-5 mm thick, hispidulous. Leaves almost distichous and altemate, sometimes subopposite; lamina ovate to elliptic to obovate, sometimes to oblong or to suborbicular, 2.5-20 x 1- 12 cm, usually equal-sided, sometimes ± unequal-sided, subcoriaceous to coriaceous, apex shortly acuminate, sometimes acute, obtuse or rounded, base acute to obtuse (to subcordate), margin dent(icul)ate to subentire; upper surface scabrous, his­ pidulous, lower surface scabrous, hispidulous or partly hirtellous to sub­ tomentose; lateral veins 3- 5(-6) pairs, basal pair ± distinctly branched and reaching the margin above or at the rniddle of the lamina, tertiary venation for the greater part scalariform; petiole 0.5-2.5(-6) cm long; stipules 0.2--0.5 cm long, strigillose to strigose, caducous. Figs in pairs or solitary in the leafaxils, just below the leaves or sometimes on the older wood; pedunde 0.5-1 cm long; bracts 1- 5, ca. 1 mm long, scattered on the pedunde and 1-4 sirnilar ones on the outer surface of the receptade; receptade (sub )globose, when fresh 1-2.5 cm, when dry 0.8-1 .5 cm in diameter, hispidulous, at maturity yellow, orange or reddish.

Distribution. From NW. Angola, Zambia, Zimbabwe and Mozambique to Sé­ négal and Ethiopia; also in Vemen, Sri Lanka and India. In areas with evergreen (e.g., rain) forest, often in secondary growth; at altitudes up to 1800 m. Tbe pollinator is Kradibia gestroi afrum (Wiebes).

3. Ficus pachycIada Baker Baker, J. Linn. Soc., Bot. 22: 522 (1887); Perrier, FI. Madagascar, Moracées: 68, fig. 13 (1952); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 19 (1986a).

Tree up to 25(-35) m tall. Leafy twigs 1.5-4 mm thick, usually scabrous, whitish

65 hispidulous or sometimes to brownish hirtellous, the bases of the hairs not swol­ len, periderm (of older parts) ftaking ofT. Leaves in spirals; lamina cordiform to ovate, sometimes subobovate or elliptic, (2- )4.5- 14(- 22) x (1.8-)3-8(- 14) cm, equal-sided (or unequal-sided), chartaceous, apex short-acuminate, base cordate to subcordate (to rounded to obtuse), margin crenate-dentate; both surfaces scabrous, ± densely hispidulous, the lower surface hirtellous or subhirtellous to puberulous as weil; lateral veins (3- )4-6 pairs, (main) basal pair ± distinctly branched, reaching the margin at about the middle of the lamina, below the ma in basal pair of lateral veins 1-3 pairs of minor basal veins, these usually on both sides of the midrib not equal in number and position, tertiary vena ti on ± scala­ riform, in small leaves reticulate; petiole (1 - )2.5-5(-13) cm long, hispidulous; stipules 0.1-0.5 cm long, hispidulous, caducous. Figs usually solitary in the leaf axils or below the leaves on previous season's growth; peduncle 0.5-2(-2.5) cm long; peduncular bracts 3, ca. 1 mm long; receptacle subglobose to subpyriform, when dry ca. (0.8- )1 - 1.5 cm in diameter, hispidulous, at maturity red.

3a. Ficus pachycIada subsp. pachycIada

F. broussonetifolia Baker (1890).

Leafy twigs 2--6 mm thick. Lamina 4-14(-22) x 2-8(-14) cm; lower surface sparsely hispidulous and hirtellous (to subhirtellous); tertiary venation for the greater part scalariform, the smaller veins mostly very prominent beneath. Re­ ceptacle when dry up to 1.5 cm in diameter.

Distribution. Madagascar and probably also in the Comoro Islands (Grande Comore).

3b. Ficus pachycIada sub sp. arborea (Perrier) C.c. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 20 (1986a).

F. soroceoides Baker var. arborea Perrier (1928).

Leafy twigs 1.5-3 mm thick. Lamina 3- 8 x 2-6 cm; lower surface sparsely his­ pidulous and subhirtellous to puberulous; tertiary venation predominantly (loosely) reticulate, the smaller veins slightly prominent to plane beneath. Re­ ceptacle when dry up to 1 cm in diameter.

Distribution. Madagascar (Central to East). In forest.

4. Ficus lateriftora Vahl (fig. 16)

Vahl, Enum. 2: 197 (1805); Berg et al., FI. Mascareignes, Moracées: 5, fig. 2 (1985).

F. morifolia Lam. (1788), non Forssk. (1775).

Tree up to 12 m tall. Leafy twigs 1.5-4 mm thick, glabrous or scabrous with

66 .Fe. 4 5

Fig. 16. Ficus lateriflora Vahl. I, leafy twig with lig; 2, branch with ligs on short-shoots; 3, lig; 4-5, leaves. After Berg & van Heusden (1985, pI. 2).

minute rigid hairs. Leaves distichous; lamina ovate to oblong, sometimes slightly unequal-sided, 6-15(-21) x 2.5- 8(- 10) cm, subcoriaceous to chartaceous, apex acuminate to acute, base subcordate to rounded (in juvenile specimens to

67 cordate), margin crenate-dentate to almost entire (in juvenile plants often pinnately to palmately lobed to parted, sometimes almost laciniate); upper surface scabrous to scabridulous, lower surface scabridulous to smooth (in ju­ venile plants scabrous); lateral veins 5-9 pairs, basal pair branched reaching the margin below, at, or just above or somewhat bel ow the middle of the lamina, tertiary venation reticulate to partly (loosely) scalariform; petiole 1.5-3{-7) cm long, glabrous or scabrous with minute rigid hairs; stipules 0.3-1.5 cm long, glabrous or minutely puberulous, caducous. Figs solitary or sometimes in pairs in the leafaxils or below the leaves, or also on the older wood, and then mostly (fasciculate) on (smalI) spurs or on up to 5 cm long ramified branches; peduncle 0.7- 1.5 cm long, usually with a bract at its base; peduncular bracts (2-)3, ca. I mm long, scattered or almost in a whorl at the top of the peduncle, sometimes a single bract on the outer surface of the receptacle; receptacle subglobose to subpyriform, up to 5 mm long stipitate, when fresh ca. 1.5-2 cm, when dry ca. 1-1.5 cm in diameter, scabrous with sparse, minute, rigid hairs, at maturity yellow to red.

Distribution. Réunion. In forest, often along water courses; at altitudes up to 1400 m. The pollinator is Kradibia etiennei Wiebes. The parasitoid Sycoscapter gibbus Saunders was recorded from this fig.

5. Ficus asperifolia Miq. Miquel in Hooker, Niger FI.: 524 (1849); Berg et al., FI. Cameroun 28: 124, fig. 40 (1985); Berg et al. , FI. Trop. East Mrica, Moraceae: 53 (1989).

F acutifo/ia Huteh. (1915), F cnestrophylla Warb. (1904), F colpophylla Warb. (1904), F irumuensis De Wild. (1922), F pa/udicola Warb. (1904), F pendula Hiem (1900), non Link (1822), F sco/pophora Warb. (1904), F storthophylla Warb. (1904), F storthophylla var. cuneata De Wild. (1922), F urceolaris Hiem (1900), F urceo/aris var. bumbana Hiem (1900), F warburgii H.l. WinkIer (1908), F xiphophora Warb. (1904).

Shrub up to 5 m tall, often with whippy, straggling or subscandent branches. Leafy twigs 1-5 mm thick, white to brown hirtellous to hispid(ulous) to strig(ill)ose or almost glabrous. Leaves distichous, alternate; lamina elliptic to oblong to ovate to subobovate to (narrowly) lanceolate or sometirnes to linear, 3- 23 x 1.5-12 cm, usually ± unequal-sided, chartaceous to subcoriaceous, apex acuminate to caudate to acute or sometimes to obtuse, base acute to obtuse to rounded, margin dentate to irregularly pinnately lobed to fid or sometimes to subentire; upper surface scabrous, hispidulous to strig(ill)ose, lower surface hispid{ulous) to hirtellous, sometimes almost glabrous; lateral veins 3-10, in large leaves up to 13, or in very narrow leaves often more than 13 pairs, basal pair unbranched and reaching the margin below the middle of the lamina, tertiary venation partly scalariform; petiole 0.5-2 cm long, white to brown hirtel­ Jous to hispidulous to strigillose; stipules 0.3-0.6 cm long, puberulous to almost

68 glabrous, caducous. Figs 1-3(-5) together in the leafaxils or just below the leaves, sometimes on the older wood; peduncle up to 0.4 cm long; bracts 2-4, ca. I mm long scattered on the peduncle and 2-4 similar ones on the outer surface of the 'receptacle; receptacle (depressed) globose to obovoid, when fresh ca. 1-2 cm, when dry 0.5---1.2 cm in diameter, hispid(ulous), at maturity dark pink to red to orangè (or yellowish?).

Distribution. From N. Angola and NW. Zambia to W. Tanzania, W. Kenya, S. Sudan and Sénégal. Mostly along streams in areas with rain forest and drier types of evergreen forest (in Uganda also in wooded grassland). The species is very variabIe. In the eastern part of its range, the figs are short pedunculate. The two forms might merit recognition at the subspecific level. The species, occurring in areas with evergreen forest, can be recognized by the une­ qual-sided lamina. The other two frutescent species have leaves with a equal­ sided lamina and occur in savanna woodland areas. Also from this fig, as weil as from F. exasperata and F. capreifolia, Kradibia gestroi afrum (Wiebes) was recorded; the 'variety' urceo/aris has Kradibia hilli Wiebes as a polljnator.

6. Ficus capreüolia Delile (fig. 17) Delile, Ann. Sci. nat. Bot. (2 ) 20: 94 (1843); Berg et al., FI. Trop. East Africa, Moraceae: 53, fig . 17 (1989).

F. antithetophylla Miq. (1848), F. capreifo/ia var. ovatifo/ia Huteh. (1916), F. pa/ustris Sim (1909), F. pao/ii Pamp. (1915).

Shrub up to 5 m tall, usually with stiff branches. Leafy twigs 1-5 mm thick, puberulous or to hirsute. Leaves ± distinctly distichous and alternate or subop­ posite, or the leaves subverticillate; larnina subovate to oblong to lanceolate, sometimes elliptic, 2-15 x 1-5.5 cm, equal-sided, chartaceous, apex (sub)acute, obtuse, rounded, 3-lobed or 3-dentate, base rounded to obtuse to acute, margin subentire or more or less faintly crenate; both surfaces scabrous; lateral veins 5- 12 pairs, basal pair unbranched, reaching the margin bel ow the middle, tertiary vena ti on reticulate to partly loosely scalariform; petiole 0.2-1(-2.5) cm long, hispidulous tosubhispid; stipules 0.5-1 cm long, partly puberulous, mostly subpersistent. Figs solitary or in pairs in the leafaxils; peduncle 0.5-1.5 cm long; bracts 3, ca. 1-1.5 mm long, scattered on the peduncle or almost in a whorl; receptacle (at least when dry) up to 5 mm long stipitate, when fresh globose, 1.5---3 cm in diameter, when dry often pyriform, 1- 2.5 cm in diameter, his­ pidulous, at maturity green to pale yellow.

Distribution. From Narnibia and South Africa to Ethiopia and Somalia, and westwards to Sénégal. Along streams and in swamps in savanna woodland areas. The species is rather uniform throughout its range. It can usually be rec­ ognized by the occurrence of subopposite (or subverticillate) leaves. The stipules

69 7

~ - ;t. Fig. 17. Ficus capreifolia Delile. I, leafy twigs with figs; 2, 1eaf; 3-4, long-styled pistillate flowers; 5, short-styled pistillate flower; 6, staminate flower; 7, ostiole. After Aweke (1979, fig. 2). are often subpersistent. The apex of the lamina is often 3-dentate (or 3-1obes); the margin otherwise is often almost entire. Kradibia gestroi afrum (Wiebes) was recorded from F capreifolia.

70 7. Ficus pygmaea Hiern Hiem, Cat. Mr. PI. Welw. 4: 1009 (1900). F brevicula Hiem (1900).

Shrubs mostly Up to 1.5 m tall, sometimes up to 4 m. Leafy twigs 1.5-4 cm thick, hirtellous, when dry brown, solid periderm persistent. Leaves alternate, tending to distichous; lamina oblong to el1iptic to subovate to ovate, 1- 13.5 cm long, 0.5--6 cm broad, (almost) equal-sided, chartaceous, apex acute (to obtuse), base rounded to subacute or to subcordate, margin more or Ie ss irregularly serrate­ dentate to pinnately lobed or fid , sometimes subentire; upper surface scabrous with minute rigid hairs of different length, on the midrib (and lateral veins) hirtellous, lower surface hirtellous to subhirsute or to pubescent; lateral veins 3-9 pairs, basal pair unbranched and reaching the margin far below the middle, tertiary reticulate or in large leaves partly scalariform; petiole 0.1-0.8 cm long, hirtellous; stipules 0.2-0.6 cm long, (Ianceolate), subpersistent or caducous, (appressed)puberulous. Figs solitary in the leafaxils; peduncle 0.5-2 cm long; bracts 4-5, ca. 1-1.5 mm long, scattered from the peduncle up to the wall of the receptacle/ (on the peduncle 1 or 0, on the base of the receptacle 0, 1 or 2 bracts); receptacle subpyriform (obovoid) to ellipsoid to subglobose, (or to oblongoid), when fresh up to 3 cm, wh en dry 1- 1.5 cm in diameter, hispidulous, at maturity red.

Distribution. Angola, Zambia, Zimbabwe, Caprivi, Botswana, and Namibia. Along streams, on sandy soi!. F. pygmaea is confined almost entirely to the Kalahari Sand area and shows little or no overlap with the area of F. capreifolia. The leaves are always alternate and the stipules caducous. The margin of the lamina is usually distinctly serrate­ dentate.

8. Ficus bojeri Baker Baker, FI. Mauritius & Seychelles: 284 (1877); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 20 (1986a).

Tree up to 12 m tall. Leafy twigs 1-3 mm thick, usually very sparsely hispidulous and nearly smooth, periderm of older parts flaking off. Leaves (tending to) distichous, often subopposite; lamina oblong to elliptic or to (sub)ovate, (3.5-) 10-17(- 24) x (1.5- )4-6.5(- 10) cm, equal-sided or unequal-sided, chartaceous, apex (sub)acuminate, base acute to rounded (or subcordate), margin (sub)entire or crenate to dentate towards the apex, sometÏmes sublobate; both surfaces sca­ bridulous to scabrous, sparsely hispidulous; lateral veins 5-9 pairs, basal pair unbranched or (faintly) branched, reaching the margin bel ow or at the middle of the lamina, sometimes smaller basal veins present below the main basal pair, tertiary venation partly scalariform; petiole (0.5-)1- 5 cm long, sparsely

71 hispidulous, and usually nearly smooth; stipules 0.2--0.5 mm long, minutely pu­ berulous, caducous. Figs solitary (or in pairs) in the leafaxils or just below the leaves, or several together on (often ± cushion-shaped clusters of 0.5-1 cm long) spurs on the older wood down to the trunk, subsessile or pedunculate; peduncle (0.3- )0.5-1.5 cm long; peduncular bracts (ca.) 3, elliptic to ovate, ca. 1 mm long; receptacle (sub)globose, when dry ca. (0.5-)0.8-1.2 cm in diameter, hispidulous, at maturity red (?).

Distribution. Seychelles, Comoro Islands (Anjouan, Mayotte) and Madagascar (prov. Diego Suarez and Nossi-Bé). In forest (undergrowth) and in secondary growth, at low altitudes.

9. Ficus brachyclada Baker Baker, J. Linn. Soc., Bot. 20: 259 (1883); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 21 (1986a).

F. phanerophlebia Baker (1887), F. soroceoides Baker var. brachyclada (Baker) Perrier (1928), F. soroceoides Baker var. macrophlebia Perrier (1928).

Shrub or treelet up to 4 m tall. Leafy twigs 1.5-4 mm thick, brownish (or partly whitish) hispidulous to hispid with hairs the bases of which are ± strongly swol­ len (or hirtellous or sparsely hispidulous). Leaves (in spirals to) distichous, mostly subopposite; lamina mostly subobovate (to obovate) oblong, sometimes oblong to elliptic (or to lanceolate), (3.5- )10-17(- 26) x (1.5-)4.5-7.5 cm, equal­ sided, chartaceous, apex (sub)acuminate, base acute (to rounded), margin dent(icul)ate, sometimes pinnately lobed to parted; upper surface scabrous, sparsely hirtellous to hispidulous, lower surface scabridulous (to scabrous), densely hirtellous (or sparsely hispidulous); lateral veins (4-)6-10(-18) pairs, basal pair unbranched, reaching the margin below the middle of the lamina, tertiary venation tending to scalariform; glandular spots (for the greater part) on the base of the midrib beneath, conspicuous; petiole 0.4-1.5 or 1- 7 cm long, hispidulous; stipules 0.3- 1 cm long strig(ill)ose, or minutely puberulous, caducous. Figs solitary or in pairs in the leafaxils or just below the leaves or several together on up to 2 cm long spur-branchlets (the development of which al ready initiated in the leaf axils) on the older wood; peduncle 0.2-1 cm long; peduncular bracts (2 or) 3, 0.5-1 mm long or 2 or 3 bracts subtending the receptacle and then up to 3 mm long; receptacle globose, when dry 0.5-1 cm in diameter, hispidulous, at maturity red(dish) or red-brown.

Distribution. Madagascar (East-Central-North). In forest; at altitudes up to 1600 m. The pollinator is Kradibia cowani Saunders.

72 10. Ficus politoria Lam. Lamarek, EncycI. Bot. 2: 500 (1788), non Lour. (1790), non Moon (1824); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 22 (1986a). F. c1aoxyloides Baker (1883), F. longipes Baker (1883), F. soroceoides Baker (1883), F. soroceoides var. ambongensis Perrier (1928), F. soroceoides var. calcicola Perrier (1928), F. soroceoides var. longipes (Baker) Perrier (1928), F. soroceoides var. mananarensis Perrier (1928), F. soroceoides var. onivensis Perrier (1928), F. xiphocuspis Baker (1883).

Shrub or tree up to 8 m taB. Leafy twigs 0.5--25 mm thick, scabrous to smooth, hispidulous to subglabrous, periderm of older parts flaking off. Leaves (almost) distichous, some of them subopposite; lamina elliptic to oblong, obovate, subo­ bovate, lanceolate, suborbicular, or sublinear, 1.5-10(-16) x 1-4(-6.5) cm, often slightly unequal-sided, chartous to subcoriaceous, apex acuminate (to subacute), base acute to rounded, margin (towards the apex) irregularly crenate-dentate or subentire, sometimes the apex tridentate, in juvenile specimens lamina to pinnate1y lobed or even parted; upper surface scabrous, sparsely hispidulous, lower surface hispidulous and scabrous or sometimes on the veins also hirteBous to puberulous; lateral veins ~8(-10) pairs, basal pair unbranched or (faintly) branched, reaching up to the middle of the lamina, tertiary venation reticulate; petiole 0.3-1 cm long, sometimes up to 3 cm long, sparsely hispidulous, epidermis flaking off; stipules O.l-D.4 mm long, (sub)glabrous, caducous. Figs solitary or in pairs in the leafaxils or on previous season's growth, sometimes (?) also up to 4 together on up to 0.5 mm long spurs on the older wood; peduncIe 0.1-D.8 or sometimes up to 2.5 cm long; peduncular bracts 3-4, ca. 0.5 mm long; receptacIe (sub)globose, when dry ca. 0.5-D.8(-1) cm in diameter, hispidulous, at maturity red, red-brown or yell ow to red.

Distribution. Madagascar. In forest, often along streams, at aItitudes up to 1700 m. The pollinator is Kradibia saundersi Wiebes. Sycoscapter gibbus Saunders, a parasitoid, was recorded from this fig.

Ficus subgenus Sycomorus (Gasp.) Mildbr. & Burret (section Sycomorus) Trees (or shrubs), terrestrial, monoecious, sap milky. Leaves in spirals or tend­ ing to distichous, margin mostly den tate, crenate or repand, tertiary venation for the greater part, scalariform, glandular spots in the axils of (main) basal lateral veins or sometimes also in the axils of other lateral veins; stipules fully amplexicaul, free. Figs solitary (or in pairs) in the leafaxils or 1-3 together on unbranched or branched leafless branchlets on the older wood, often down to the trunk, pedunculate, basal bracts 3, lateral bracts absent, receptacIe (rather) large, ostiole circular and several (sometimes only 3-4) ostiolar bracts visible, only the lower ones descending, interfloral bracts absent among the pistillate flowers, staminate flowers near the ostiole, enveloped by 2 bract(eole)s, perianth

73 saccate, 3-lobed, sta mens 2 or 3; pistillate flowers: tepals 2-4-6, irregularly shaped, free or connate, often laciniate, glabrous, stigma 1, truncate to elongate, often (sub)infundibuliform; endocarp body often released; wall of the fig at fruit soft, red, orange or yellowish.

This description refers to subg. Sycomorus sensu Corner (1965), thus only com­ prising monoecious species, and not of the Sycomorus-group as indicated and discussed above (p. 49). Five species of this subgenus occur in continental Africa. The species are arborescent and most of them bear the figs on long, often branched, leafless branchlets on the older wood, in some species down to the base of the trunk. Eight species (including F. sycomorus) are known from Madagascar and neighbouring islands. One species, F. racemosa L., is confined to Asia. The figs are often (or usually?) filled with a watery sap during (part of) the interfloral phase. The pollinators belong to the genus Ceratosolen Mayr.

11. Ficus sycomorus L. (fig. 18) Linnaeus, sp. PI.: 1059 (1753); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 24 (1986a); Berg et al., FI. Trop. East Africa, Moraceae: 54 (1989).

F chanas Forssk. (1775), F coccu/ifolia Baker (1887), F coccu/ifolia subsp. saka/avarum (Baker) Perrier (1952), F coccu/ifolia var. saka/avarum (Baker) Perrier (1928), F comorensis Warb. (1894), F damarensis Engl. (1888)*, F gnapha/ocarpa (Miq.) A. Rich. (1851), F integrifo/ia Sim (1909), F scabra Sim (1909), non Forster f. (1786), nec. Willd. (1801), F saka/avarum Baker (1887), F lrachyphy /la (Miq.) Miq. (1867), Sycomorus anliquorum Gasp. (1845), S. gnaphalocarpa Miq.(1848)*, S. rigida Miq. (1848), S. trachyphy/la Miq. (1867)*.

Tree up to 15 m tall. Leafy twigs white puberulous, intermixed with much (> 10 x) longer hairs, periderm flaking off. Lamina elliptic to oblong to (sub)obovate, ovate or subcordiform, 1.5- 14 x 1- 10.5 cm, chartaceous, often brittie when dry, apex rounded to obtuse, base cordate to rounded, margin faintly crenate to subentire; upper surface scabrous, hispidulous, on the midrib puberulous to hirtellous or hirsute, lower surface on the veins puberuious to tomentellous, on the main veins intermixed with much (> 10 x) longer hairs; lateral veins 4-7 pairs, basal pair branched, reaching the margin bel ow to near the middle of the lamina, tertiary venation partly scalariform; petiole 0.5-3 cm long, puberulous, intermixed with much (> 10 x) longer hairs, epidermis flaking off; stipules 0.5--1.5 cm long, puberulous or partly subsericeous or subhirsute, caducous. Figs on up to 20 cm long, branched branchlets on the main branches or solitary (or in pairs) in the leafaxils or just below the leaves; peduncle 0.3-0.6 cm long, epidermis flaking off; basal bracts 2.5--3 mm long; receptacle globose to broadly obovoid to subpyriform, when dry ca. 1.5-2 cm or up to 5 cm (*) in diameter, sparsely puberulous, hirtellous or yellowish velutinous; wall when dry 1-2 or 5- 7 (*) mm thick.

74 ., .,

. 10

, ; ,"

9

Fig, 18. Ficus sycomorus L. I , leaftess branchlet with figs; 2, leafy twig; 3, stipules and petioles; 4-6, pistillate flowers; 7, staminate ftowers with bracteoles; 8, staminate flower in bud; 9-10, sta­ mens; 11, ostiole. After Aweke (1979, fig. 18).

75 Distribution. From Namibia, N. Botswana and South Africa extending to Ma­ dagascar, Sénégal and Cape Verde Islands ; also in the Arabian Peninsuia, and in cultivation for consumption of the figs in Egypt, Israel, and Syria. In woodland, near streams. In Namibia and Angola (also Botswana?) the specimens have the figs (sol­ itary) in the leafaxils or just below the leaves (subsp. gnapha/ocarpa (Miq.) c.c. Berg, 1980). This form (to which the synonyms indicated with an asterisk belong) also occurs in West Africa. In the rest of the distribution, the forms with figs in the leafaxils and with figs on leafless branchlets on the older wood occur side by side, making recognition of subspecies hardly worthwhile. The type collection of F. saka/avarum has figs that are slightly larger than normal, namely up to ca. 2.5 cm in diameter. These figs are bom on leafless branchlets on the older wood. In the collections Baron 4881, Blommers 21, Boiteau 347, Humbert 5071, and Perrier de la Bätrue 16585 (some ofthe features of which are marked with an asterisk (*) in the description above) the figs are much larger than normal and axillary. It is not quite dear whether a subspecific taxon based on F. saka/avarum can be maintained, as the presently available data suggest th at a gradual transition from the normal form with relatively small figs to the form with much larger figs occurs. It is also possible that the dimen­ sions of the figs merely depend on the place where the figs are formed on the plant. A study on the pollinating fig wasp, as also of the parasitoids of the genus Apocrypta by Uien berg (1985), indicated that different species of this genus are associated with the two forms of F. sycomorus: the nominate form has Cerato­ solen arabicus Mayr (and the 'cuckoo' C. ga/ili Wiebes) and Apocrypta /on­ gitarsus Mayr, while from the sakalavarum-form Ceratoso/en namorakensis (Risbec) and Apocrypta cercaria Uien berg are known. There were quite a number of parasitoids recorded from F. sycomorus, viz., Apocryptophagus gigas (Mayr), Eukoebelea sycomori Wiebes, [dames graci/e Wiebes, Sycophaga sycomori (L.) and S. va/entinae Grandi, [Apocrypta, see above], Sycoscapteridea bambeyi (Risbec), Ficomi/a gambiensis (Risbec), Syceu­ ry loma ficus Boucek, and Sycophila naso Boucek.

12. Ficus mucuso Ficalho Fica1ho, PI. Ut. Afr. Portug.: 270 (1884); Berg et al. , FI. Cameroun 28: 132, fig . 43 (1985); Berg et al. , FI. Trop. East Africa, Moraceae: 56 (1989).

F. cory/ifolia De Wild. & T. Durand (1901), F. eory/ifo/ia var. g/abreseens Warb. (1904), F. sidifolia Riem (1900).

Tree up to 30(-40) m taU, with tall trunk, often with buttresses, main branches ascending. Leafy twigs 3-8 mm truck, densely minutely white puberulous and with much longer wrute to dark brown hairs, especially on the nodes, periderm flaking off. Lamina suborbicular to subcordiform to eUiptic or sometimes to obovate, 6-17 x 4-15 cm, chartaceous to subcoriaceous, apex shortly acuminate to acute, base cordate, sometimes truncate, margin entire or crenate; upper

76 surface scabridulous to scabrous, sparsely brownish hirtellous to hirsute on the main veins, lower surface pube~ulous to hirtellous, on the main veins often densely puberulous and with long to dark brown hairs with an echinate base; lateral veins 3- 6 pairs, basal pair above or at the middle of the larnina, tertiary vena ti on for the greater part scalariform; petiole (1-)2- 9 cm long, densely minutely white puberulous and with much longer to dark brown hairs with an echinate base, epidermis flaking oIT; stipules 1-2 cm long, puberulous to sub­ sericeous, caducous. Figs on up to 30(-40) cm long, branched leafless branchlets on the main and less branches or also on the trunk; peduncle 1-2.5 cm long; basal bracts 3-5 mm long; receptacle (depressed) globose to obovoid, when fresh 2.5-4 cm, when dry 2-3 cm in diameter, puberulous; at maturity red to dark orange.

Distribution. From NW. Angola through Zaire to Ethiopia, Tanzania, and to Guinea-Bissau. In rain forest areas. This species shows close morphological affinities to F. sycomorus. It is, how­ ever, ecologically distinct; moreover, in contrast to F. sycomorus, it forms tall trunks. The pollinator is Ceratosolen arabicus Mayr, as in F. sycomorus, and also the 'cuckoo' is the same (C galili Wiebes). The parasitoid Apocrypta crypta UIen berg was described from F. mucuso.

13. Ficus SUf F orssk. Forsskäl, Fl. Aegypt.-Arab.: 180 (1775); Berg et al., Fl. Trop. East Africa, Moraceae: 56, fig. 18 (1989). F. beniensis De Wild. (1921), F. brassii Sabine (1824), F. capensis Thunb. (1786), F. capensis var. beniensis (De Wild.) Lebrun (1934), F. capensis var. guineensis (Miq.) Miq. (1867), F. capensis var. iluriensis (De Wild) Lebrun (1934), F. capensis var. mallolocarpa (Warb.) Mildbr. & Burret (1911), F. capensis var. mallolocarpa forma pubifolia Mildbr. & Burret (19\1), F. capensis var. oslio/ala (De Wild.) Lebrun (1934), F. capensis var. pubescens De Wild. & T. Durand (1901), F. capensis var. lrichoneura Warb. (1894), F. clelhrophylla Hiern (1900), F. erubescens Warb. (1904), F. gongoensis De Wild. (19\3a), F. guineensis (Miq.) Stapf (1906), F. iluriensis De Wild. (1921), F. kondeensis Warb. (1901), F. lichlensleinii Link (1822), F. mallolocarpa Warb. (1894), F. mallolocarpa var. Irichoneura (Warb.) A. Peter (1932), F. munsae Warb. (1904), F. oslio/ala De Wild. (19J4), F. oSlio/ala var. brevipeduncu/ala De Wild. (1914), F. panifica Delile (1843), F. p/aleiocarpa Warb. (1901), F. riparia A. Rich. (1851), F. slellu/ala Warb. var. g/abrescens Warb. (1904), F. sycomorus L. var. a/nea Hiern (1900), F. sycomorus var. prodigiosa Hiern (1900), F. sur var. erylhreae Fiori (19\Oa), F. sycomorus var. po/ybolrya Hiern (1900), F. thonningiana (Miq.) Miq. (1867), F. villosipes Warb. (1904), Sycomorus capensis (Thunb.) Miq. (1848), S capensis forma guineensis (Miq.) Miq. (1849, S guineensis Miq. (1848), S panifica Miq. (1848), S? riparia Miq. (1848), S. sur (Forssk.) Miq. (1849), S thon­ ningiana Miq. (1848).

Tree up to 25(-30) m tall. Leafy twigs 2-5 mm thick, white to yellowish (or brownish) puberulous to hirtellous to tomentose, partly hirsute or almost glabrous, periderm usually not flaking oIT. Lamina elliptic to ovate to subovate or to oblong, sometimes to suborbicular or to lanceolate, 4-20(-32) x 3-13(-16)

77 cm, chartaceous to coriaceous, apex acuminate to acute, base subacute to (sub)cordate, margin coarsely crenate-dentate to re pand to (sub)entire; upper surface smooth, sometimes scabrous, glabrous or puberulous on the proximal parts on the main veins, lower surface on the whole surface or only on the main veins puberulous to tomentose or glabrous; lateral veins (3- )5- 9 pairs, basal pair (faintly) branched, reaching the margin bel ow the middle of the lamina, tertiary venation for the greater part scalariform to al most reticulate; petiole 1.5- 9 cm long, puberulous, hirtellous, hirsute, subtomentose or glabrous, epidermis usually not flaking off; stipules 1-3.5 cm long, white to yellow subsericeous, pubescent, hirsute or almost glabrous, caducous. Figs on up to 50(- 150) cm long, branched leafless branchlets on the older wood, down to the trunk, occa­ sionally in the leafaxils just below the leaves; peduncle 0.5-2 cm long; basal bracts 2- 3 mm long; receptacle obovoid to subglobose, often more or less depressed globose, at least when dry often stipitate, when fresh 2-4 cm, when dry 0.5- 2.5 cm in diameter, sparsely white to yellowish puberulous to almost glabrous, to densely tomentose or to subvelutinous, at maturity red to dark orange.

Distribution. From Namibia and E. South Africa to Sénégal and to Ethiopia; also on the Cape Verde Islands, Sào Tomé, Fernando Po, Principe, and in Yemen. In wood land and moist (e.g. rain and montane) forest. The species is rather variabie in the indumentum of the leaves and figs (glabrous to densely pubescent) and in the shape and margin of the larnina. The pollinators are Ceratosolen capensis Grandi in East and South Africa, and C. silvestrianus Grandi in West Africa - in both West and East Africa also C. flabellatus Grandi is found in the figs. Parasitoids are Apocrypta guineensis Grandi, Sycophaga afJiicta Grandi and S. cyc/ostigma Waterston.

14. Ficus vogeliana (Miq.)

Miquel, Ann. Mus. Bot. Lugd.-Bat. 3: 295 (1 867); Berg et al ., Fl. Cameroun 28: 140, fig . 46 (1985); Berg et al. , Fl. Trop. East Africa, Moraceae: 58 (\989).

F fleuryi A. Chev. (1920), F seretii Lebrun & Boutique (\948, 1949), F stel/ulata Warb. (\894), F vogeliana var. latifolia Huteh. (191 6), Sycomorus vogeliana Miq. (1848).

Tree up to 20 m tall, with buttresses. Leafy twigs 3- 5 mm thick, minutely white puberulous and with much longer yellowish to white hairs, periderm flaking off. Lamina broadly elliptic to obovate to oblong or to subovate, 5- 22 x 2.5- 11 cm, apex acurninate, base cordate to subtruncate, margin coarsely dentate to sub­ entire; upper surface scabrous to scabridulous, sparsely, but on the main veins more densely white to yell ow hirtellous to hirsute, lower surface white to yell ow hirsute, often only on the main veins; lateral veins 5- 9 pairs, basal pair (faintly) branched reaching the margin mostly below, sometimes at or above the rniddle of the lamina, tertiary venation for the greater part scalariform; petiole 0.5-5.5 cm long, minutely white puberulous and with much longer white to yellowish

78 hairs, epidermis flaking ofT; stipules 1- 2 cm long, rninutely brownish to white puberulous, often also with much longer appressed hairs or sometimes glabrous, subpersistent. Figs on up to lOm long branched leafless branchlets arising from the base of the trunk (and then the figs in the litter underneath the tree) or on usually short branchlets up to the main branches; peduncle 0.5-1.5 cm long; basal bracts 3-5 mm long; receptacle mostly depressed globose, when fresh 2-3 cm, when dry 1- 2 cm in diameter, minutely puberulous to hirtellous, at maturity red to orange-red, often with yellowish to white spots.

Distribution. From Angola (Cabinda) to Uganda and Guinea. In areas with evergreen forest, in periodically inundated riverside and swamp forest. This species is geocarpic; the fig-bearing branch lets can be up to 10 m long and all or most of them are found at the base of the trunk. These branches are partly subterranean. The fig-bearing branchlets are sometimes also hanging down from the main branches. The pollinator is Ceratosolen acutatus Grandi.

15. Ficus vallis-choudae Delile Delile, Ann. Sci. nat. Bot. (2) 20: 94 (1843); Berg et al., FI. Cameroun 28: 142, fig. 47 (1985); Berg et al. , FI. Trop. East Africa, Moraceae: 58 (1989).

F. schweinfurthü Miq. (1867), F. vallis-choudae var. pubeseens A. Peter (1932), Sycomorus schim­ periana Miq. (1 848).

Tree up to 10(- 20) m tall. Leafy twigs 2- 10 mm thick, glabrous, sparsely white appressed-puberulous or sometimes white hirtellous to tomentose, periderm flaking ofT. Larnina ovate to cordiform to (sub)deltoid, 4-24(-36) x 3-24(- 30) cm, coriaceous to subcoriaceous, apex acute to subobtuse or very shortly acu­ mina te, base obtuse to truncate or to cordate, margin coarsely and obtusely den ta te to repand, sometimes subentire; upper surface smooth and glabrous or puberulous, sometimes hirtellous to pubescent, or occasionally scabrous; lateral veins 5- 8 pairs, basal pair branched reaching the margin at or above the rniddle of the lamina, tertiary venation for the greater part scalariform; petiole 2- 11(- 13.5) cm long, glabrous, white appressed-puberulous or sometimes white hirtel­ lous to tomentose, epidermis flaking ofT; stipules 1-3 cm long, in the lower part ciliolate, appressed-puberulous or subsericeous, caducous. Figs solitary in the leafaxils or just bel ow the leaves or occasionally on up to 30 cm long branchlets on the older wood; peduncle 0.2- 1.2 cm long; basal bracts ca 2 mm long; re­ ceptacle subglobose to obovoid, when fresh 3--6(- 10) cm, when dry 1- 5 cm in diameter, more or less densely white to yellowish puberulous to hirtellous or to tomentose, only tomentellous near the ostiole or glabrous, at maturity yellowish to orange with longitudinal orange to reddish stripes.

Distribution. Zambia, Malawi, Mozambique, Zimbabwe; extending to Ethiopia and Guinea.

79 In areas with woodland, often in periodically inundated places, at altitudes up to 1800 m. The pollinator is Ceratosolen megacephalus Grandi. Parasitoids are Apocrypta robusta Grandi and Sycophaga tenebrosa Grandi.

16. Ficus mauritiana Lam.

Lamarek, Encycl. Bot. 2: 499 (1788); Berg et al. , F1. Mascareignes, Moracées: 7, fig. 3 (1985).

F. ayersii Baker (1877), F. obtusata Link (1822), F. sapotoides Baker (1877), F. soraria Cordem. (1895) , F. terragena Bory (1804).

Shrubs or trees up to 10 m tall. Leafy twigs 3-5 mm thick, minutely appressed­ puberulous to yellow hirtellous, when dry the internodes often narrowed. Lamina ovate to elliptic or to oblong, sometimes to suborbicular, 6-15(-27) cm long, 3-8(-15) cm broad, subcoriaceous, apex acute to acuminate, base cordate to truncate or to rounded, margin repand to ± faintly crenate-dentate (in juvenile plants sometimes lobed); both surfaces (almost) glabrous or the lower surface hirsute to hirtellous to tomentose on the whole surface or only on the veins; lateral veins 4-6 pairs, basal pair branched, reaching the middle of the lamina above, at or below the middle, tertiary venation for the greater part (loosely) scalariform; petiole 1.5-13 cm long, puberulous to pubescent; stipules 0.5-2 cm long, usually only ciliolate, caducous or sometimes subpersistent. Figs solitary or in pairs on leafless, unbranched or branched, up to 70 cm long branches on the older wood, down to on the base of the trunk, sometimes the figs on normal leafy twigs; peduncle 0.5- 1.5 cm long; basal bracts 3 (or occasionally 2), ca. 3 mm long; receptacle subglobose to obovoid or to turbinate (or when dry some­ times almost discoid), when dry ca. 1.5-3.5 cm in diameter, puberulous, at ma­ turity brown to reddish.

Distribution. Mauritius and Réunion. In forest or thicket, mostly along water courses; at altitudes up to 1100 m. The pollinator is Ceratosolen coecus (Coquerel). Parasitoids are Apocryp­ tophagus explorator (Coquerel) and Apocrypta perplexa Coquerel.

17. Ficus tiliifolia Baker Baker, J. Linn. Soc., Bot. 21: 443 (1885); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 26 (1986a).

F. pulvinifera Baker (1887), F. sphaerophylla Baker (1885), F. tiliifolia var. ambrensis Perrier (1928), F. tiliifolia var. sphaerophylla (Baker) Perrier (1928), F. trichophlebia Baker (1885).

Tree up to 20 m tall. Leafy twigs 2-{5 mm thick, minutely puberulous or on the scars of the stipules white to yellowish hirtellous, periderm flaking ofT. Lamina cordiform to ovate to elliptic, (3-5-)6.5-16(-35) x (2- )3.5-15(-26) cm, chartaceous to subcoriaceous, apex shortly acuminate to subacute, base sub­ cordate to rounded, margin crenate-dent(icul)ate to coarsely crenate or sub-

80 entire; upper surface scabrous to smooth, sparsely hirtellous, lower surface hirtellous to puberulous on the (main) veins or throughout; lateral veins 5-7(-8) pairs, main basal pair branched, reaching the margin near the middle of the lamina, tertiary venation scalariform; petiole (1-)l.5-9.5(-13) cm long, sparsely minutely puberulous, epidermis fiaking off; stipules 0.5-2.5 cm long, whitish to yellowish(brown) subsericeous, hirtellous or subhirsute. Figs solitary in the leaf axils or just below the leaves, sometimes on up to 10 cm long branched leafiess branchlets on the older wood down to the trunk; peduncle 0.3-1 cm long; basal bracts ca. 2-3 mm long; receptacle subglobose, often ± depressed with a concave apex, when dry 1.5.2.5 cm in diameter, densely yellowish puberulous.

Distribution. Madagascar and Comoro Islands (Anjouan). In forest, at altitudes up to 1700 m. The pollinator is Ceratosolen stupefactus Wiebes.

18. Ficus torrentium Perrier Perrier de la Bàthie, Areh. Bull. Mens. 2: 149 (1928); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 27 (1986a).

Tree up to 20 m taU. Leafy twigs 3-5 mm thick, dark brown hirsute, with minute whitish hairs among the long brown hairs, periderm ± fiaking off. Lamina oblong to elliptic to (sub)obovate, 5-l.8 x 3- 7 cm, subcoriaceous to coriaceous, apex subacute to faintly and shortly acuminate, obtuse or rounded, base cordate to rounded, margin subentire; upper surface scabrous to scabridulous, sparsely white hirsute (to puberulous), lower surface rather densely white hirteUous to pubescent; lateral veins 4(-5) pairs, (main) basal pair (faintly) branched, reach­ ing the margin above the middle of the lamina, tertiary venation scalariform; petiole 1- 3.5 cm long, with long brown haies, intermixed with minute white hairs; stipules 0.5-1 mm long, white puberulous to tomentellous and with long, ± appressed brown hairs, caducous. Figs solitary in the leafaxils; peduncle 0.2--0.4 cm long; bracts l.5- 2 mm long; receptacle (depressed-)globose, when fresh 3--4 cm, when dry 1.8-3 cm in diameter, densely yeUowish puberulous and with sparse longer, dark brown haris, at maturity yellowish.

Distribution. Madagascar. Along streams, at altitudes up to 1100 m.

19. Ficus polyphlebia Baker Baker, J. Linn. Soc., Bot. 20: 262 (1883).

F. albidula Baker (J 887).

Shrub or tree up to 1O(?) m taU. Leafy twigs 2-4 mm thick, white hirtellous, white strigillose or sparsely minutely puberulous, periderm persistent, in the upper part of te internodes a few large lenticels. Lamina clliptic to oblong to

81 (sub)ovate, 2-12(-23) x 1-6(- 12) cm, subcoriaceous, apex subacute or (faintly) shortly acuminate, base cordate to rounded to obtuse, margin entire; upper surface smooth, sparsely (on the midrib more densely) hirtellous, lower surface sparsely to densely hirtellous (to puberulous) or subglabrous; lateral veins (3-)5- 8 pairs, basal pair unbranched (to faintly branched), running almost parallel to the margin and reaching it in or below the middle of the larnina, tertiary vena ti on partly scalariform; petiole 0.3- 3(- 3.5) cm long, white hirtellous, epidermis flak­ ing off; stipules 0.5-2 cm long, white strigose to hirtelJous or subglabrous, caducous. Figs solitary in the axils or on up to 15 cm long, unbranched, leafless branch lets on the ol der wood, subsessile or up to 0.4 cm long pedunculate; basal bracts broadly 2- 3 mrn long; receptacle (depressed-)globose, when dry ca. 2-3 cm in diameter, yellowish or white puberulous to hirtellous, at maturity red with white spots.

Distribution. Madagascar. In rocky places along streams; at altitudes up to 1200 m. The pollinator is Ceratosolen longimucro Wiebes, a parasitoid Apocrypta se­ toptera UIenberg.

20. Ficus botryoides Baker

Baker, J. Linn. Soc., Bot. 22: 523 (1887); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 28 (1986a).

F oxystipu/a Baker (1890), F stenoc/ada Baker (1890).

Tree up to 22 m talJ. Leafy twigs 2-2.5 mrn thick, minutely appressed-pu­ berulous (or glabrous), periderm ± flaking off. Lamina lanceolate to oblong or sublinear, (3-)7-18(- 24) x (0.5-)1.5-6(- 9) cm, (sub)coriaceous, apex acuminate to acute, base acute to obtuse (to rounded), margin (sub)entire; upper surface glabrous, lower surface glabrous or very sparsely appressed-puberulous on the main veins; lateral veins (5- )7- 10(- 15) pairs, basal pair unbranced (or faintly branched), reaching the margin below the rniddle of the larnina, tertiary venation reticulate; petiole (0.7-)1- 3(- 5) cm long, rninutely appressed-puberulous (or glabrous), epidermis flaking off; stipules (0.7-)1- 3 cm long, glabrous, ciliolate, or very sparsely appressed-puberulous, caducous. Figs on up to 5 cm long, branched branchlets (or in pairs or solitary on up to 30 cm long unbranched branch lets with long internodes and terminally with smallieaves) from the base of the trunk up to the main branches; peduncle (0.7-)1- 8 cm long, epidermis flaking off; basal bracts ca. 4 mm long, inside conspicuously veined; receptacle broadly ovoid to depressed-globose, when dry 1.2- 2.5 cm in diameter, minutely appressed-puberulous, at maturity red(-brown).

Distribution. Madagascar. In forest; at altitudes up to 1600 m. The pollinator is Ceratosolen blommersi Wiebes, a parasitoid Apocrypta po­ lyspina UIenberg.

82 21. Ficus trichoclada Baker Baker, 1. Linn. Soc., Bot. 22: 524 (1887).

F laurus Baill. (1895).

Shrub or tree up to 15 m tall. Leafy twigs 1--4 mm thick, white puberulous to tomentose, periderm flaking ofT. Lamina lanceolate to oblong (to subovate), (1.5- )5- 15(-20) x (1 - )1.5- 5.5(-6.5) cm, coriaceous, apex subacute to obtuse, base acute to obtuse (or rounded), margin entire; upper surface smooth, sparsely puberulous on the lower part of the midrib; lower surface puberulous (or on the midrib to hirtellous) or subglabrous; lateral veins (5-)8- 13 pairs, basal pair ± distinct, unbranched, reaching the margin (far) below the middle of the lamina, tertiary venation reticulate; petiole (0.5- )0.7- 2(-2.5) cm long, puberulous to sub­ tomentose, epidermis flaking ofT; stipules 0.5-1.2(-2) cm long, (sometimes sparsely) subsericeous to appressed-puberulous (or pubescent), caducous. Figs solitary in the leafaxils, or just below the leaves, or (sometirnes?) on up to ca. 15 cm long, unbranched or branched branchlets on the older wood, down to the trunk; peduncle 0 . ~ . 8 cm long, epidermis flaking ofT; basal bracts 1.2-2 mm long; receptacle depressed-globose to subpyriform, when dry 1.5-2.5 cm in di­ ameter, sometimes ± stipitate, puberulous to subtomentose.

Distribution. Madagascar. Along streams; at altitudes up to 1500 m.

22. Ficus karthalensis c.c. Berg (fig. 19) Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 30, fig. 1 (1986a).

Tree up to 20 m tall. Leafy twigs 2-3 mm thick, sparsely and minutely pu­ berulous to glabrous, periderm of the ol der parts ± flaking ofT. Lamina (sub)ovate to elliptic to oblong, (1.5-)4-15 x (1 - )2-9 cm, chartaceous to sub­ coriaceous, apex acuminate to acute, base (acute) to rounded to emarginate, margin entire; upper surface smooth and glabrous, lower surface sparsely and minutely puberuious on the ma in veins or glabrous; lateral veins (2-)4-6 pairs, (main) basal pair faintly branched, reaching the margin near the middle of the lamina, tertiary venation for the greater part scalariform; petiole (0.5- )1-3 cm long, sparsely and minutely puberulous to subglabrous, epidermis ± flaking ofT; stipules 0.5- 1.3 cm long, appressed-puberulous or subglabrous, caducous. Figs solitary in the leafaxils (fide Bernardi 11779: also cauliflorous); peduncle 0.3-0.4 cm long; basal bracts ca. I mm long; receptacle globose, when dry 0.8-1.2 cm in diameter, minutely puberulous; wall when dry 0.5- 1 mm thick; ostiole ca. 1.5 mm in diameter, ± sunken, and only 3 (or 4) ostiolar bracts visible.

Distribution. Comoro Islands (Anjouan and Moheli).

83 Fig. 19. Ficus kartluJlensis C.c. Berg. I, leafy twig with figs; 2, staminate ftower with bracteoles; 1-4, pistiUate ftowers. After Berg (1986a, pI. I).

Ficus subgenus Pharmacosycea Miq. section Oreosycea (Miq.) Corner Trees (or shrubs), terrestrial, monoecious, sap (scanty and) watery. Lenticels concentrated in the uppermost part of the internodes. Leaves in spirals to dis­ tichous, margin entire to dentate, (in some species when juvenile irregularly pinnately incised), tertiary venation often for the greater part scalariform, glan­ dular spot(s) at the base of the midrib beneath; stipules fully amplexicaul (but when juvenile semi-amplexicaul), free. Figs solitary or in pairs in the leafaxils, pedunculate, basal bracts 3, (normally) in a whorl, lateral bracts occasionally present; ostiole circular with 3 ostiolar bracts visible, only the lower ostiolar bracts descending; interfioral bracts lacking or present; starninate flowers near the ostiole, tepals 2-3(-7), connate, g1abrous, stamen 1, pistillode sometimes

84 present; pistillate flowers: tepals 2- 3(- 7), connate, glabrous, stigrnas 2, (sub)filiforrn; fruits achene-like; wall of the receptacle at fruit fum, yellow to orange.

This section comprises 40-50 species, most of them in Asia-Australasia. It has two closely related species in continental Africa, and two in Madagascar. Three of these four species have pinnately incised leaves in juvenile states. Tbe pollinators belong to the genus Do/ichoris HilI.

23. Ficus varüfolia Warb. Warburg, Ann. Mus. Congo (6) 1: 30, fig. 15 (1904); Berg et al., Fl. Cameroun 28: 145, fig. 48 (1985); Berg et al., Fl. Trop. East Africa, Moraceae: 59 (1989).

F. bongouanensis A. Chev. (1920, nomen; Aubréville, 1959, sub F. bangouanensis), F. inopinata Mildbr. (1953), F. sciarophylla Warb. (1904), F. zenkeri Mildbr. & BUITet (1911).

Tree up to 35 (or more?) m tall, with buttresses. Leafy twigs 2-3 mm thick, white puberulous or glabrous, when juvenile white hirsute to pubescent, when dry red-brown to blackish. Lamina oblong to elliptic to (sub)ovate, 5-20 x 2- 11.5 cm, when juvenile often larger, subcoriaceous, when juvenile to chartaceous, apex subacute to acuminate, base cordate to rounded, margin (sub)entire, when juvenile irregularly lobed to fid; both surfaces giabrous to sparsely puberulous, smooth or scabridulous, when juvenile white hirsute to hirtellous, often scabrous; lateral veins (5-)8-13 pairs, when juvenile often more pairs, tertiary venation for the greater part scalariforrn; petiole 1-5 cm long, sparsely to densely white puberulous; stipules 0.3-0.7 cm long, puberulous or partly hirsute, caducous. Figs solitary or in pairs in the leafaxils; peduncle 0.3-1 cm long; basal bracts ca. 2 mm long; receptacle globose to obovoid, when fresh ca. 2 cm, when dry ca. 1.5 cm in diameter, puberulous, at maturity yellow. StamÏnate and pistil­ late flowers with a 2-3-lobed to -fid perianth. Interfloral bracts absent.

Distribution. Angola (Cabinda); extending to Uganda, Sudan and Sierra Leone. In rain forest and drier types of evergreen forest; at altitudes below 1300 m. This species is very closely related to F. dicranostyla, from which it difTers in the greater number of lateral veins and a smooth upper surface of the lamina. The two taxa may prove to be distinct only at the subspecific level. The pollinator is Do/ichoris flabella ta Wie bes.

24. Ficus dicranostyla Mildbr. (fig. 20)

Mildbraed, Bot. Jahrb. Syst, Pfianzengesch. & Pfianzengeogr. (EngIer) 46: 204 (191Ib); Berg et al., Fl. Trop. East Africa, Moraceae: 59, fig . 19 (1989).

F. dicranostyla var. nitida Huteh. (1916), F. lynesii Lebrun (1935).

Tree up to 6(-20) m or shrub. Leafy twigs 2-5 mm thick, white pubescent to

85 3

2

JW Fig. 20. Ficus dicranostyla Mildbr. I, leafy twig with figs; i -4, figs ; 5-6, pistillate flowers; 7, pistiI; 8, staminate flower in bud and interfloral bract. After Aweke (1979, fi 'g. 4). puberulous, when dry pale brown to dark red-brown. Lamina elliptic to oblong to (sub)ovate, (2-)5- 20 x (1-)2=-9 cm, subcoriaceous to chartaceous, apex acu-

86 minate, sometimes subacute, base obtuse to acute or to (sub)cordate, margin (sub)entire; upper surface scabrous, hispidulous, on the main veins puberulous to hirtellous to pubescent, lower surface puberulous to hirtellous to pubescent; lateral veins 5-8(-9) pairs, tertiary venation partly scalariform; petiole 1-3.5 cm long, densely white pubescent to tomentose to hirtellous; stipules 0.5- 1.5 cm long, puberulous to pubescent, caducous. Figs solitary or in pairs in the leaf axils; peduncle 0.3-1 cm long; basal bracts 2-2.5 mm long; receptacle globose to obovoid, when fresh 1-2.5 cm, when dry 0.5-1.5 cm in diameter (at least when dry often shortly stipitate), puberulous to hispidulous, at maturity yellowish to pale orange. Staminate and pistillate flowers with a 2- 3-lobed to -fid perianth. Interfloral bracts absent.

Distribution. From Sénégal to S. Ethiopia, NE. Zaire and N. Uganda, also disjunctly in SE. Zaire and N. Zambia. In savanna woodland, often in rocky places; in N. Zambia often on termite mounds; at altitudes up to 1500 m. The material from the (smalI) southem partial area (Zambia and Shaba) is somewhat different from th at from the main areas of the species. It appears to be morphologically closer to F variifolia than the material from the large northem partial area. In some specimens the basal bracts are not arranged in a distinct whori, but scattered on the peduncle or scattered from the peduncle to the receptacle, sim­ ilar to arrangements as often occur in section Sycidium. The pollinator is Dolichoris flabella ta Wie bes (material from Zambia).

25. Ficus assimilis Baker Baker, J. Linn. Soc., Bot. 25: 345 (1890); Berg, Bull. Mus. natn. Hist. Nat. Paris (4) 8 (B): 32 (I 986a).

F. gualleriifolia Baker (1890).

Tree up to 25 m tall. Leafy twigs 1.5-3(-6) mm thick, glabrous, minutely pu­ berulous or sparsely to densely white hirtellous to pubescent, when dry usually blackish or dark brown. Lamina subovate to oblong, ovate to elliptic or lanceo­ late, 5.5- 17 x 2.5-6 cm, (sub)coriaceous, apex (faintly) acuminate to acute, base rounded to subcordate or to subacute, margin (sub)entire; upper surfce smooth to scabrous, glabrous or white hirtellous to pubescent on the midrib, lower surface glabrous or white hirtellous to subhirsute to pubescent on the (main) veins; lateral veins 7-14 pairs, tertiary venation partly scalariform or reticulate; petiole 1- 5 cm long, glabrous, hirtellous or pubescent; stipules 0.3- 1.5 cm long, white subsericeous or appressed-puberulous, caducous. Figs in pairs in the leaf axils or just below the leaves; peduncle 0.3- 1.3 cm long; bas al bracts 1-1.5 mm long; receptacle (sub)globose (to ellipsoid or obovoid), when dry (0.5- )1 - 1.5 cm in diameter, glabrous or minutely puberulous; 3- 5 ostiolar bracts visible. Sta­ minate flowers with 3- 8 tepals; pistillate flowers with 3- 8, free tepals. Interfloral bracts present or absent.

87 ~ Fig. 21. Ficus ampana c.c. Berg. I, leafy twig with fig; 2-3, ostiole; 4, stamen; 5, long-styled pistillate flower; 6, short-styled pistillate flower and interfloral bracts. After Berg (1986a, pI. 2).

Distribution. Madagascar. In forest; at altitudes up to 1000 m. The interfloral bracts occur both on the receptacle and the pedicels of both staminate and pistillate flowers with 3-5 tepals. If the flowers have more, up to 8 tepals, the interfloral bracts are absent on the receptacle and thepedicels. This difTerence suggests that the bracts are included in the perianth in the latter case.

26. Ficus ampana C.C. Berg (fig. 21)

Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 33, fig . 2 (1986a).

Shrub or treelet. Leafy twigs 1.5-2.5 mm thick, glabrous, when dry blackish. Lamina oblong to elliptic or to lanceolate, 3-7 x 1-4 cm, coriaceous, apex obtuse to rounded or very shortly and obtusely acuminate, base obtuse (to rounded), margin entire; both surfaces glabrous; lateral veins 6-8 pairs, tertiary venation reticulate; petiole 0.5-1.2 cm long, glabrous; stipules 0.3-0.5 cm long, glabrous or sparsely appressed-puberulous, caducous. Figs solitary (or in pairs) in the leaf axils; peduncle 0.3-D.6 cm long; basal bracts ca. I mm long; receptacle globose, when dry ca. 0.8 cm in diameter, glabrous; 3 (or 4) ostiolar bracts visible. Sta­ minate flowers with 4, free tepals; pistillate flowers with 4 (or 5), free tepals. Interfloral bracts few, sometimes on the pedicels (or sometimes lacking?).

Distribution. Madagascar.

Ficus subgenus Urostigma (Gasp.) Miq. section Urostig1lUl Trees or shrubs, usually terrestrial (or epilithic), sometimes herni-epiphytic, monoecious, sap milky. Leaves in spirals or sometimes subdistichous or subop­ posite, margin (sub)entire, tertiary venation reticulate, glandular spot at the base of the rnidrib beneath; stipules fully amplexicaul, free. Figs in pairs or sometimes more together in the leafaxils or just below the leaves, pedunculate or sessile, basal bracts 3, lateral bracts absent, ostiole circular with 3 ostiolar bracts visible, only the lower ostiolar bracts descending, interfloral bracts absent or present (but few) ; staminate flowers usually only near the ostiole, tepals 3-4, free; stamen I, pistillode sometimes present; pistillate flowers with tepals 3-4(- 7), free tepals, stigma I, (sub)filiform; fruit achene-like, reddish-brown; wall of the fig at fruit soft, red, pink or purplish.

This section comprises 20 species; 15 species in Asia-Australasia and one species each in Madagascar and the Mascarene Islands. Three species occur in continen­ tal Africa. Two of these species are trees or treelets, occurring in dry, often rocky places; the third species, F. verruculosa, in shrub occurring along streams.

88 1

5

3 The poUinators are species of the genus Platyscapa Motschoulsky.

27. Ficus ingens (Miq.) Miq. (fig. 22)

Miquel, Ann. Mus. Bot. Lugd.-Bat. 3: 288 (1867); Berg et al ., FI. Trop. East Africa, Moraceae: 60, fig. 20 (1989).

F caffra (Miq.) Miq. (1867), F caffra var. longipes Warb. (1906), F caffra var. natalensis Warb. (1906) , F caffra var. pubicarpa Warb. (1906), F caffra var. sambesiaca Warb. (1906), F ingens var. tomen/osa Hutch. (1925), F ingentoides Hutch. (1915), F katagumica Hutch. (1915), F kawuri Hutch. (1915), F magenjensis Sim (1909), F ovato-cordata De Wild. (1921), F pondoensis Warb. (1906) , F stuhlmanni Warb. var. glabrifolia Warb. (1894), F schimperiana A. Rich. (1851), Urostigma ingens Miq. (1847), U. caffrum Miq. (1849), U. xanthophyllum Miq. (1847), U. xanthopyhyllum var. ovato-cordatum Sonder (1850).

Tree up to 15 m taU. Leafy twigs 3-{5 mm thick, sparsely white to brownish pubescent to densely tomentellous or subvelutinous. Lamina ovate to subovate to eUiptic or to oblong, or occasionally lanceolate, (2.5-)5- 20 x (2- )3- 11 cm, coriaceous, apex (shortly) acuminate to acute, sometimes to obtuse, base cordate, occasionally truncate or obtuse, margin entire; both surfaces glabrous; lateral veins 8-11 pairs, the (main) basal pair more or less distinctly branched, almost straight, thus running not parallel to the margin, the others often furcate far from the margin; petiole 0.5--4 cm long, glabrous to ± densely white to brownish puberulous to subvelutinous; stipules 0.5-1 cm long, densely yellowish tomentose to subvelutinous or glabrous, caducous. Figs in pairs in the leafaxils or just below the leaves, subsessile or up to 0.5 mm long pedunculate; basal bracts ca. 2 mm long; receptacle (sub)globose, when fresh 1-2 cm, when dry 0.5- 1 cm in diameter, minutely white to brown puberulous or partly hirtellous, sometimes densely tomentose, or pubescent, at maturity whitish to pink or to pale to dark purple; wall when dry wrinkled.

Distribution. From Namibia and South Africa to Sénégal, Ethiopia and Somalia; also in the Sahara region and Yemen. In woodland, often in rocky places. The species is rather variabIe and sometimes difficult to tell apart from broad­ leaved forms of F cordata subsp. salicifolia. The pollinator is Platyscapa soraria Wie bes.

28. Ficus cordata Thunb. Thunberg, Diss. Fic.: 8, with plate (1786); Berg, Kew Bull. 43: 81 (1988). Tree up to 15(-35) m tall. Leafy twigs 2- 5 mm thick, densely white puberulous or pubescent to subglabrous. Lamina cordiform, ovate elliptic or to oblong, lanceolate, or sometimes (sub)obovate, 2- 17 x 1-{5(- 8) cm, coriaceous, apex acute to (faintly) acuminate or to obtuse (or rounded), base cordate to rounded to ob tu se (or subacute), margin entire; both surfaces glabrous; lateral veins 6--12 pairs, the bas al pair unbranched, usually curved and running almost parallel to

90 Fig. 22. Ficus ingens (Miq.) Miq. 1, leafy twig with figs; 2-4, pistillate flowers; 5, staminate nOWt,. , 6, stipules and petioles; 7, ostiole. After Aweke (1979, fig . 9). the margin, the others furcate not far from the margin, smaller veins beneath prominent and conspicuous; petiole 0.5-4(-6.5) cm long, glabrous; stipules 0.5- 1.5 cm long, glabrous or ciliolate. Figs mostly in pairs in the leafaxils or just below the leaves, or up to 2- 3 mm long spurs on older wood, subsessile or up to 0.3 cm long pedunculate; basal bracts ca. 1- 1.5 mm long, sometimes caducous; receptacIe (sub)globose, when fresh 0.5- 1 cm, wh en dry 0.5-0.8(-1) cm in diameter, maturing from green to whitish to dark purple or dark red; wall when dry usually smooth.

91 Three allopatric subspecies can be recognized. On the basis of morphology, distribution and ecology subsp. corclnta and subsp. salicifolia could be treated as distinct at the species level, if the West-African subsp. lecardii would not show intermediate features, making the situation rather complex.

28a. Ficus cordata subsp. cordata

F. conJata var. fleckii Warb. (\ 906), F. cordata var. marlothii Warb. (\ 906), F. cordata var. tristis (Kunth & Bouché) Warb. (1906), F. rupium Dinter (\909), F. tristis Kunth & Bouché (1847), F. welwitschii Warb. (\894), F. welwitschii var. beroensis Hiern (\900), Urostigma cordatum (Thunb.) Gasp. (1845), U. thunbergii Miq. (\847).

Leafy twigs usually ± densely puberulous to pubescent. Lamina usually cor­ diform to ovate to elliptic, (2-)4-9 x (1-)2-4.5 cm, apex acuminate to sub­ cordate, base cordate, truncate or rounded (to subacute); petiole 1-2(-6) cm long, mostly drying yellowish to brown; stipu1es 0.2-1 cm (on flush up to 5 cm) long, usually puberulous to pubescent. Figs sessile, sometimes subsessile.

Distribution. W. Angola, NW. Botswana, Namibia, and Sw. South Africa. In semi-desert, in rocky places, at altitudes up to 1500 m. Material from West Africa, part of which resembles subsp. cordata in the 1eaf shape andJor the sessile figs, is placed in subsp. lecardii. The latter subspecies shows a mixture of features found in the two other subspecies. The pollinator is Platyscapa desertorum Compton. A parasitoid described and named in this paper (p. 185) is Comptoniella vannoorti Wiebes.

28b. Ficus cordata subsp. lecardü (Warb.) e.C. Berg Berg, Kew Bull. 43: 81 (1988); Berg et al. , Fl. Cameroun 28: 162, fig. 50 (1985).

F. lecardii Warb. (1904), F. salicifolia Vahl. var. latifolia Huteh. (\916).

Leafy twigs glabrous to puberulous. Lamina ovate to subovate to oblong (or lanceolate), 2- 12 x 1.5- 8, apex acuminate to subacute, base cordate, rounded or obtuse (to subacute); petiole 0.8-4.5 cm long, often drying dark brown to blackish; stipules 0.5-1 cm long. Figs mostlyon peduncles up to 0.3 cm long, sometimes sessile.

Distribution. Sénégal to the Central Africa Republic. In savannas, often near rocks, at altitudes up to 1500 m.

28c. Ficus cordata subsp. salicifolia (Vahl) e.e. Berg Berg, Kew Bull. 43: 82 (1988); Berg et al., Fl. Trop. East Africa, Moraceae: 63 (1989).

F. eucalyptoides Battand. & Trabut (1912), F. indica Forssk. (1775), non L. (\753), F. pretoriae Burtt-Davy (1912), F. salicifolia Vahl (\790), F. salicifolia var. australis Warb. (\906), F. teloukat Battand. & Trabut (\912), Urostigma salicifolium (Vahl) Miq. (\847).

92 Leafy twigs usually minutely puberulous to glabrous. Larnina subovate to oblong to lanceolate, sometimes ovate or elliptic, 2- 17 x 1- 5.5 cm, apex acute to subacuminate (to rounded), base rounded, cordate or obtuse (to subacute); petiole 0.5-4(-6.5) cm long, usually drying yellowish to brown; stipules 0.5-1.5 cm long, glabrous or ciliolate. Figs usually on peduncles up to 0.3 cm long, som times subsessile.

Distribution. From SE. Botswana and NE. South Africa, to the southern part of the Arabian Peninsula, Socotra, Egypt, and Algeria. In woodland, often in rocky places, at altitudes up to 2700 m. The po1linator is Platyscapa awekei Wiebes. A parasitoid is Otitesella serrata Mayr.

29. Ficus verruculosa Warb. Warburg, Bot. Jahrb. Syst., Pflanzengsch. & Pflanzengeogr. (EngIer) 20: 166 (1894); Berg et al., FI. Carneroun 28: 154, fig. SI (1985); Berg et al. , FI. Trop East Africa, Moraceae: 63 (1989).

F praeruptorwn Hiem (1900), F verruculosa var. stipitata Mildbr. & Burret (191 la).

Shrub or treelet mostly up to 2.5 m, sometimes up to 5, occasionally up to 12 m tal\. Leafy twigs 1- 5 mm thick, glabrous or densely white hirtellous to sub­ tomentellous. Larnina oblong to lanceolate, sometimes elliptic, 3.5-10(-20) x 1.5-3.5(- 8.5) cm, coriaceous, apex subacute to obtuse, base obtuse to rounded to subcordate, margin entire; both surfaces glabrous; lateral veins (8-)10-16 pairs, the basal pair unbranched, curved and running almost parallel to the margin, beneath only the midrib prominent, the other veins often plane and inconspicuous; petiole 0.5-2(- 3) cm long, glabrous; stipules 0.5-3.5(-4) cm long, glabrous to densely white puberulous, caducous. Figs mostly in pairs or some­ times up to 4 together in the leafaxils or just below the leaves; peduncle 0.~.5(­ I) cm long; basal bracts ca. I mm long; receptacle (sub)globose (to ellipsoid), when fresh 0.5-2 cm, when dry 0.5-1(- 1.2) cm in diameter, glabrous or minutely puberulous, at maturity to dark-purple or to dark red; wall when dry often wrinkled.

Distribution. From Angola and South Africa (and Botswana?) to Uganda and Niger. In areas with woodland, along streams. The species is distinct from F. ingens and F. salicifolia in its frutescent habit and in its ecology. The pollinator is Platyscapa binghami Wiebes.

30. Ficus madagascariensis C. C. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 34 (l986a).

Shrub or (large) tree. Leafy twigs 2- 3 mm thick, glabrous or minutely pu-

93 berulous, periderm of older parts flaking off. Lamina oblong to lanceolate (or to elliptic) or (sub)ovate, 7-15 x 2.5--5.5 cm, (sub)coriaceous, apex long-acu­ minate, base obtuse to rounded or to subacute; both surfaces glabrous; lateral veins 7- 12 pairs, tertiary venation reticulate; petiole 0.5-5 cm long, glabrous; stipules 0.3-0.5 cm long, white (appressed-)puberulous, caducous. Figs in pairs in the leafaxils or just below the leaves, sessile; basal bracts ca. 1.5 mrn long; receptacle subglobose, when dry 0.4--0.6 cm in diameter, sparsely white pu­ berulous.

Distribution. Madagascar. Dry forest on xerophytic bush.

31. Ficus densifolia Miq. Mique1, Ann. Mus. Bot. Lugd.-Bat. 3: 218 (1867); Berg et al., Fl. Mascareignes, Moracées: 9 (1985).

F. lucens Cordem. (1895).

Tree up to 6 m taU. Leafy twigs 1.5- 5 mm thick, glabrous or minutely pu­ berulous, with large conspicuous lenticels. Lamina ovate to elliptIc, 3.5--12.5 cm long, 1.5--7 cm broad, coriaceous, apex shortly acuminate to subacute, base rounded to truncate to obtuse or to subcordate, margin entire; both surfaces glabrous; lateral veins 9- 12 pairs, tertiary venation reticulate; petiole 1-4.5 cm long, glabrous; stipules 0.5-1.5(-8) cm long, glabrous, caducous. Figs usually in pairs in the leafaxils or just bel ow the leaves, sessile; basal bracts 3-5 mrn long, often split; receptacle (sub)globose, when dry ca. 0.5--1 cm in diameter, glabrous.

Distribution. Réunion and Mauritius. At altitudes up to 1200 m. The pollinator is Platyscapa etiennei Wiebes. Parasitoids are Sycoryctes anceps Wiebes and S. comparabilis Wie bes.

Ficus subgenus Urostigma section Conosycea (Miq.) Corner In most features this section is similar to section Urostigma. The circular ostiole may have more than 3 ostiolar bracts visible. The fruits are whitish, mostly with a reddish mark; at maturity the wall of the fig can be pink, red, yellow to orange, or greenish.

This section comprises about 65 species, most of them in Asia-Australasia, two in Madagascar. The differences between the sections Urostigma and Conosycea appear to be small and it is questionable whether the two groups of species are distinct at the level of section. The pollinator of one of the African Conosycea figs is a species of Platyscapa, but th is is not the usual asociation in the Indo-Australian region.

94 32. Ficus menabeensis Perrier Perrier de la Bäthie, Not. Syst. 24: 167 (1952); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 36 (1986a).

F. pyrifolia Lam. var. amobongensis Perrier (1928), F. pyrifolia var. meridionalis Perrier (1928).

Tree up to 20 m tall, shrub, or epilithic scrambler, (often?) with aerial roots on the branches. Leafy twigs 2- 5 mm thick, white puberulous (or glabrous), often also with dark brown, pluricellular hairs, periderm of older parts mostly flaking off. Lamina oblong to lanceolate or to elliptic, sometimes sublinear or suborbi­ cular, (1 - )3- 20 x (0.5--)1.5--5.5 cm, coriaceous, apex obtuse, acute or apiculate, base acute to rounded; upper surface glabrous or sparsly puberulous on the lower half of the lamina or only on the midrib, lower surface densely puberulous to hirtellous to tomentellous to subglabrous, often also dark brown, pluricellular hairs present; lateral veins (3- )12- 22 pairs, especially in relatively narrow leaves connected by a slightly arcuate submarginal vein, tertiary venation reticulate; petiole 0.3-2 cm long, white puberulous to tomentellous to subglabrous; stipules 0.5--1.5(- 2.5) cm long, puberulous, caducous. Figs in pairs in the leafaxils, ses­ sile; basal bracts 2.5--3 mm long, coriaceous, often subcarinate; receptacle globose, when dry 0.5-0.8 cm in diameter, sparsely to densely white puberulous andJor with dark brown pluricellular hairs, at maturity reddish; 3 ostiolar bracts visible.

Distribution. Madagascar (South and South-West). In savannas, dunes, riv­ ersides, often in rocky places; at altitudes up to 1200 m. The pollinator is Platyscapa bergi Wiebes.

33. Ficus humbertü c.c. Berg (fig. 23) Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 37, fig . 3 (l986a).

Shrub up to 2 m tall. Leafy twigs 1.5- 2 mm thick, white puberulous. Lamina suborbicular to elliptic, 0.5--2 x 0.5--1.5 cm, coriaceous, apex rounded, obtuse, or apiculate, base rounded to truncate, margin entire; upper surface sparsely puberulous in the lower half of the lamina, lower surface (sparsely) minutely puberulous; lateral veins 3- 5 pairs, the middle ones often furcate far from the margin, tertiary venation reticulate; petiole 0.1-0.5 cm long, puberulous; stipules 0.2-0.5 cm long, puberulous, caducous. Figs in pairs (or solitary) in the leaf axils; peduncle 0.1-0.2 cm long; basal bracts ca. 1 mm long; receptacle globose, wh en dry 0.4--0.7 cm in diameter, minutely puberulous; 4 or 5 ostiolar bracts visible, these bracts small and in the lower part thickened.

Distribution. Madagascar.

95 2

1

6 Fig. 23. Ficus humbertii e.C. Berg. I, leafy twig; 2, leafy twig with figs; 3-4, ostiole; 5, staminate flower; 6-7, pistillate flowers with interfioral bracts. Mter Berg (1986a, pI. 3).

96 Ficus subgenus Urostigma section Galoglychia (Gasp.) End!. Trees, shrubs or sometimes lianas, hemi-epiphytic or terrestrial (or epilithic), sap milky. Leaves in spirals, sometimes almost distichous andJor subopposite, margin (sub)entire, tertiary venation scalariform to reticulate to predominantly parallel to the lateral veins, glandular spot at the base of the midrib beneath; stipules fully amplexicaul, usually free or (in some species) partly connate. Figs in the leafaxils or just bel ow the leaves or on (± peg-like) spurs on the lesser branches or down to the base of the trunk, pedunculate or sessile, basal bracts 2 or occasionally 3, lateral bracts absent; ostiole split-shaped (or triradiate), all ostiolar bracts descending, interfloral bracts present; staminate flowers disperse, tepals 2-4, free or connate, glabrous, stamen 1, pistillode usually absent; pistil­ late flowers: tepals 2-4, free or partly connate, glabrous, stigma(s) 1 (or 2); fruit achene-like or drupaceous and th en often releasing the endocarp body or the upper part forming a mucilaginous cap; wall of the fig at fruit soft to rather form, reddish, orange, yellow, greenish, purplish or brownish.

This section is confined to the African floristic region and comprises 72 species. Sixty-six of them belong to the flora of continental Africa, but three extend to Madagascar and neighbouring islands; six species of this group are confined to the Madagascar subregion. Six subsections can be recognized in section Ga­ loglychia (see Berg, 1986b). There are some uncertainties in this subdivision, which are discussed bel ow. The pollinators belong to the tribe Agaonini, different from those of the Blas­ tophagini, to which tri he all species belong that were mentioned above.

Ficus subgenus Urostigma section Galoglychia subsection Galoglychia . Trees hemi-epiphytic or secondarily terrestrial, evergreen, bark not flaking off, usually with conspicuous lenticels in longitudinal rows. Leafy twigs, stipules, leaves and figs, often pubescent to hirsute or villous. Leaves large to medium­ sized, oblong to lanceolate or to elliptic, coriaceous, tertiary venation (partly) scalariform to (loosely) reticulate; stipules relatively large, caducous. Figs in pairs in the leafaxils or more together and also just below the leaves, large to smalI, sessile, at maturity yellow to orange. Fruitlets with a pericarp becoming partly mucilaginous, endocarp body released and surrounded by a rather thin mucilaginous layer.

The subsection Galoglychia comprises three species in West and Central Africa; only F. lutea extends to East and South Africa, Madagascar, the Aldabra Islands, and the Seychelles. The subsection is associated with evergreen forest. The pollinators belong to the genus Allotriozoon Grandi.

97 34. Ficus saussureana De. De Candolle, Mem. Soc. Phys. Genève 9: 2, with plate (1841); Berg et al, FI. Cameroun 28: 209, fig. 74 (1985); Berg et al., FI. Trop. East Africa, Moraceae: 69 (1989).

F. dawei Huteh. (1915), F. eriobolryoides Kunth & Bouché (1847), F. eriobolryoides var. cail/ei Mildbr. & Burret (1911), F. eriobotryoides var. monbullensis (Warb.) Lebrun (1934), F. galac­ tophora Tenore (1851), F. monbuttensis Warb. (1904), F. principes Kunth & Bouché (1847), F. rubroreceptaculata De Wild. (1914), F. zobiaensis De Wild. (1914), Galoglychia saussureana (DC.) Gasp. (1844), G. tenoreana Gasp. (1844), Urostigma eriobotryoides (Kun th & Bouché) Miq. (1849), U. saussureanum (De.) Miq. (1847).

Tree up to 20 m tall, hemi-epiphytic or (secondarily?) terrestrial, with spreading crown. Leafy twigs 10-15 mm in diameter, puberuious and usually also white to yell ow pubescent to subhirsute, periderm flaking ofr. Lamina oblong to sub­ obovate to oblanceolate, (10-)15-50 x 3-17(-25) cm, coriaceous, apex acu­ minate, base acute to obtuse or to (sub)cordate, margin entire; upper surface glabrous, lower surface puberulous to hirtellous, on the main veins to sub­ hirsute; lateral veins 12- 20 pairs (in small laminas, Ie ss than 15 cm long, some­ times 10-11 pairs), tertiary venation partly scalariform; petiole 1-8(-16) cm long, 4-7 mm thick, epidermis flaking otT; stipules 1-8 cm, on flush up to 12 cm long, outside brown to yellowish pubescent, inside with long appressed yellow hairs, caducous or subpersistent. Figs in pairs, or sometimes up to 3 together in the leafaxils or just bel ow the leaves, subsessile, initially enclosed by up to I cm long greyish subvillous axillary calyptrate bud covers, these being caducous; basal bracts 7-15 mm long, persistent; receptacle subglobose, some­ times obovoid, when fresh 2-4 cm, when dry 1.5- 3 cm in diameter, yell ow to orange-brown subhirsute to villous, at maturity yellow to orange to reddish.

Distribution. From Guinea to S. Soudan, W. Kenya, and NW. Tanzania. In forest, particularly riverine forest; at altitudes up to 1600 m. Often planted. The polinator is Allotriozoon prodigiosum Grandi. Parasitoids are Philo­ trypesis finitimorum Wiebes, Sycoscapter montis Wiebes, Sycoryctes lomaensis Wiebes, Otitesella royi Wiebes and Camarothorax africanus (Wiebes).

35. Ficus chlamydocarpa Mildbr. & Burret

Mildbraed & Burret, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (Engier) 46: 244, fig. 5 (191Ib); Berg et al., FI. Cameroun 28: 212 (1985); Berg, Kew Bull. 43: 80 (1988).

Tree, often large, hemi-epiphytic or (secondarily?) terrestrial. Leafy twigs 0.5-- 1.5 cm thick, ± densely minutely puberulous, also with long yellow to brownish hairs on the scars of the stipules, or glabrous. Lamina oblong to elliptic to subobovate, 12-23 x 4-10.5 cm, coriaceous, apex (often faintly andJor shortly) acuminate, base cordate to rounded or sometimes obtuse; upper surface glabrous, lower surface densely minutely puberulous on the main veins andJor with longer (often soon disappearing) hairs on the midrib and proximal parts

98 of the lateral veins, or entirely glabrous; lateral veins 8- 14 pairs, tertiary vena­ tion partIy scalariform to loosely reticulate; petiole (1 - )2- 8 cm long, ca. 3 mm thick, glabrous or long white to yellow hairs, epidermis flaking ofT; stipules 0.5- 2.5 cm long, minutely puberulous andJor with sparse to dense long yellow hairs, or entirely glabrous, caducous. Figs solitary in the leafaxils, sessile or up to 0.8 cm long pedunculate; basal bracts ca. 5 or 15-25(- 35) mrn long, pu­ berulous, appressed-pubescent or glabrous, persistent; receptacle (depressed)­ globose to obovoid, when fresh 2.5-4 (or more?) cm, when dry 2-3.5 cm in diameter, glabrous or yellow pubescent, smooth to ± warty, at maturity yell ow to orange (or red?).

Three allopatric subspecies can be recognized.

35a. Ficus chlamydocarpa subsp. chlamydocarpa

F c1arencensis Mildbr. & Hutch. in Hutch. (1915). Plants ± hairy, at least on the stipules. Basal bracts 1.5-2.5(-3.5) cm long.

Distribution. In Cameroun and Bioko (= Femando Po). In forest; at altitudes between 1300 and 2000 m. The pollinator is Allotriozoon nigeriense Wiebes.

35b. Ficus chlamydocarpa subsp. fernandesiana (Huteh.) c.c. Berg Berg, Kew Bull. 43: 80 (1988).

F fernandesiana Hutch. in Exell (1956).

Plants entirely glabrous, also on the stipules. Basal bracts ca. 0.5 cm long.

Distribution. In Sào Tomé. At low altitudes. Probably introduced in Gabon (Libreville).

35c. Ficus chlamydocarpa subsp. latifolia (Huteh.) c.c. Berg Berg, Kew Bull. 43: 81 (1988).

F eriobo/ryoides Kunth & Bouché var. la/ifolia Hutch. (1916).

Plants ± hairy, at least on the stipules. Basal bracts ca. 0.5 cm long.

Distribution. In Annobon. At low altitudes.

36. Ficus lutea Vahl (fig. 24) Vahl, Enurn. 2: 185 (1805); Berg et al. , Fl. Cameroun 28: 206, fig. 73 (1985); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 38 (1986a); Berg et al., Fl. Trop. East Africa, Moraceae: 69 (1989).

F akaie De Wild. (1914), F apodocepha/a Baker (1885), F arimensis Britton (1922), F baronii Baker

99 Fig. 24. Ficus IUlea Vahl. 1, Leafy twig with figs; 2, 3, staminate fiowers; 4-5, pistillate flowers; 6, stipules. Mter Aweke (1979, fig. 22).

(1883), F. cabrae Warb. (1904), F. holstii Warb. (1894), F. incognila De Wtld. (1914), F. kaba De Wild. (1914), F. lanigera Warb. (1894), F. nau/arum Baker (1877), F. nekbudu Warb. (1904), F. neumannii Kunth & Bouché (1847), F. pachyclada Baker (1890), F. pseudo-vogelii A. Chev. (1902), F. quibeba Ficalho (1884), F. senegalensis Miq. (1867), F. subcalcarala Warb. & Schweinf. in Warb.

100 (1894), F. subca/carata var. vestito-bracteata (Warb.) Mildbr. & Burret (1911), F. trichosphaera Baker (1883), F. uti/is Sim (1909), F. verrucocarpa Warb. (1901), F. vestilO-bracteata Warb. (1894), F. vogelii (Miq.) Miq. (1867), F. vogelii var. pubicarpa Mildbr. & BUffet (1911), Urostigma /uteum (Vahl) Miq. (1847), U. neumanii (Kunth & Bouché) Miq. (1847), U. vogelii Miq. (1847).

Tree up to 20 m tall. Leafy twigs 5-12(-20) mm thick, puberulous, white to yellow tomentose to subhirsute to subvillous or glabrous, periderm flaking ofT. Lamina elliptic to oblong to (sub)ovate, sometimes (ob)lanceolate, 7-25(-45) x 3- 12(- 20) cm, coriaceous, apex shortly acuminate, base obtuse to acute or to subcordate, margin entire; upper surface glabrous or minutely puberulous on the midrib, lower surface sparsely (to densely) puberulous to hirtellous to (sub)tomentose, on the main veins to subhirsute or entirely glabrous; lateral veins (4-)6-8(- 10) pairs, tertiary vena ti on reticulate to partly scalariform; petiole (1-) 1. 5-13(-17) cm long, 2-4(-8) mm thick, glabrous or white pilose, epidermis flaking ofT; stipules 0.5- 2.5 cm, on flush up to 12 cm long, sparsely to densely white to yellowish puberulous to subsericeous, caducous. Figs in pairs or sometimes up to 4 together in the leafaxils or (just) below the leaves, sessile, initially enclosed by up to 0.5 cm long, white pubescent to subhirsute calyptrate bud covers; basal bracts 3-6 mm long, persistent; receptacle (sub)globose, when fresh 1-2.5(- 3.5?) cm, when dry 0.5- 1.5(-2.5) cm in diameter, puberulous, white pubescent or (partly) yellowish villous, maturity yell ow to orange or brownish.

Distribution. From N. Angola, N. Zambia, South Africa (Natal) to Ethiopia and Sénégal, also in the Cape Verde Islands, Bioko (Fernando Po), Madagascar, Comoro Islands, Aldabra Islands, and the Seychelles. In forest, often along rivers, marshes and sea coast; at alti tud es up to 1800 m. Often planted. In West Africa the species is rather uniform and generally has puberulous small figs (when fresh ca. 1-1.2 in diameter) and rather small leaves. More eastwards the species is more variabie. The figs are often larger (when fresh up to 2.5 cm in diameter. They may bear long hairs. The leaves are often also larger and may have up to 12 pairs of lateral veins. Therefore, F lutea and F saus­ sureana ean not be easily told apart in the eastern part of Central Africa. The species is also quite variabie in Madagascar and and other Indian Ocean islands. The pollinator is Allotriozoon heterandromorphum Grandi. Parasitoids are Crossogaster atratus Masi and C. si/vestrü Grandi, Philotrypesis selenitica Grandi and Otitesella africana Grandi.

Ficus subgenus Urostigma section Galoglychia subsection Platyphyllae (Mildbr. & Burret) c.c. Berg Berg, Proc. Kon. Ned. Akad. Wet. (C) 89: 122 (1986b).

Trees or shrubs, hemi-epiphytic, terrestrial or epilithic, often deciduous. Bark often exfoliating. Leafy twigs, stipules, leaves and figs often conspicuously hairy. Leaves often ovate, cordate or reniform, often with several pairs of basal lateral

lOl veins, lateral veins usually branched, leaf margin sometimes repand to coarsely crenate, tertiary venation often ± distinctly scalariform; petiole often relatively long, stipules often rather long, caducous. Figs medium-sized to large, mostly pedunculate, wall at maturity often soft. Fruitlets with a dry pericarp, sometimes the inner layers becoming mucilaginous and then the endocarp body released and surrounded by a rather thin mucilaginous layer.

Subsection Platyphyllae comprises 18 species and shows a concentration of species in eastem Africa. It is represented by five species in Madagascar and neighbouring islands. Most of the species of this subsection are associated with various types of savanna woodland, although in most cases growing in situations in some way protected against fire (gallery forest, rocky places, termite mounds). Only F. recurvata and F. jansii are apparently confined to rain forest conditions. F abutilifolia, F glumosa and F. tettensis are found on rocks; F. abutilifolia and F. tettensis are 'rock-splitters'. F. trichopoda is usually found in marshy areas; it forms stilt-roots or 'pillar-roots', apparently a feature that can be connected with its common habitat. Fourteen species of this subsection belong to the flora of the African con­ tinent. One of them extends to Madagascar and four species are confined to Madagascar and neighbouring islands. The subsection Platyphyl/ae is more clearly than the other subsections as­ sociated with relatively dry habitats. This is reflected in some morphological features: often relatively long petioles, ± mesophyllous leaves, a cordate to re­ niform larnina, dense indument on twigs, leaves, and/or figs, and ± exfoliating bark. Most of the species are associated with savanna woodland. They are often terrestrial or epilithic. Some of them (e.g. F. platyphylla) may be fire-resistent. Others are often protected against fire by their occurrence on rocks or on terrnite mounds. The position of F. jansii is not quite certain. It might belong to subsection Crassicostae. The pollinators belong to various genera, viz., Alfonsiella Waterston, Elisabethiella Grandi and Nigeriella Wiebes.

37. Ficus platyphyUa Oelile Delile, Cent. PI. Méroé: 62 (1 826); Berg et al. , FI. Cameroun 28: 165, fig . 55 (1985); Friis, Nord. J. Bot. 5: 333 (1985); Berg et al. , FI. Trop. East Africa, Moraceae: 64 (1989) .

F. kotschyana (Miq.) Miq. (1867), F. lateralis Warb. (1904), U. kotschyanum Miq. (1 847).

Tree up to 15 m tall, (secondarily?) terrestrial. Leafy twigs (5- )10-20 mm thick, densely white puberulous to velutinous, on the nodes pubescent, periderm flak­ ing ofT. Lamina ovate to elliptic or subovate to oblong, (8-)15-26 x (5-)10-19 cm, coriaceous, apex obtuse to acute, base cordate; margin entire to repand; upper surface glabrous or puberulous on the lower part of the rnidrib, lower

102 surface white puberulous to pubescent or only puberulous on the ma in veins; lateral veins 10-16 pairs, basal pair branched, reaching the margin (far) below the middle of the lamina, tertiary venation partly scalariform; petiole 4--10 cm long, 2-5 mm thick, puberulous to pubescent; stipules 0.8-2.5 (-3.5) cm long, densely white tomentose to puberulous to subvelutinous, caducous. Figs up to 5 together in the leafaxils or just below the leaves; peduncle 1- 2.5 cm long; basal bracts ca. 3 mm long, persistent; receptacle globose, when fresh 1-2 cm, when dry ca. 1(-1.5) cm in diameter, puberulous, warty, at maturity greenish(?).

Distribution. Sénégal to Somalia. In wooded grassland, often at rocky places; at altitudes up to 750 m. The pollinator is an unidentified species of Elisabethiel/a Grandi near E. pec­ tinata (Joseph); a parasitoid is Philocaenus barbatus Grandi.

38. Ficus bussei Mildbr. & Burret Mildbread & Burret, Bot. Jahrb. Syst., Pftanzengesch. & Pftanzengeogr. (EngIer) 46: 213 (l911b); Friis, Nord. 1. Bot. 5: 332 (1985); Berg et al., Fl. Trop. East Africa, Moraceae: 64 (1989).

F changuensis Mildbr. & Burret (1911), F changuensis var. somalensis Pamp. (1915), F fasciculata Warb. (1894), F somalensis (Pamp.) Chiov. (1929), F zambesiaca Huteh. (1915).

Tree up to 20 m high, (secondarily?) terrestrial. Leafy twigs 4--12 mm thick, sparsely (to densely) puberulous to hirtellous or glabrous. Lamina subovate to oblong, 5-24 x 3-9.5 cm, coriaceous (when dry often brittle), apex (sub)acute to obtuse, base cordate, margin entire to repand; upper surface glabrous or puberulous on the midrib, lower surface subglabrous or (rather) sparsely hirtel­ lous to puberulous; lateral veins (8-) 10-16 pairs, basal pair branched, reaching the margin far below the middle of the lamina, tertiary vena ti on partly ± loosely scalariform to almost reticulate; petiole 2- 8 cm long, 2-4 mm thick, (sparsely) puberulous to hirtellous; stipules 0.3- 1.2 cm, on flush up to 3 cm long, glabrous or at the base pubescent, caducous. Figs in pairs or solitary in the leafaxils; peduncle 1-2.5 cm long, recurvate; basal bracts ca. 3 mm long, persistent; re­ ceptacle (sub)globose (to ellipsoid), when fresh 2-3 cm, when dry 1- 1.5 cm in diameter, puberulous, smooth or ± warty, at maturity greenish (?).

Distribution. In Zambia, Malawi, Mozambique, Zimbabwe, and through the coastal regions of Tanzania and Kenya to Somalia. In lowland forest. The species shows morphological affinities with F platyphyl/a and more strongly with F recurvata. The two taxa may prove to be distinct only at the subspecific level.

39. Ficus recurvata De Wild. De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 200 (1913a); Berg et al., Fl. Cameroun 28: 168, fig . 57 (1985).

103 F bussei Mildbr. & BUITet var. longepetiolata Mildbr. in Mildbr. & BUITet (191Ia), F golungensis Huteh. (1915), F goliath A. Chev. (1909, nomen; Aubréville, 1959).

Tree up to more than 35 m tall, (secondarily?) terrestrial. Leafy twigs 5-10 mm thick, sparsely hirsute to glabrous. Larnina elliptic to oblong, (10-)15-28(-44) x 7- 15(- 21) cm, subcoriaceous, apex subacuminate to subacute or almost rounded, base cordate, margin subentire to coarsely crenate-dentate, especially towards the base; upper surface sparsely hirsute to hirtellous, lower surface hirsute on the midrib and the lateral veins, sparsely hirtellous to puberulous or glabrous on other parts (or the whole surface glabrous); lateral veins (8-)10-18 pairs, basal pair branched, reaching the margin far below the middle of the la mina, tertiary veins partly ± loosely scalariform; petiole (2- )5- 8(- 10) cm long, ca. 3 mm thick, hirsute; stipules 1-2.5 cm, on flush up to 4.5 cm long, (sub)glabrous, caducous. Figs in pairs in the axils of the leaves; peduncle 1.5-3.5 cm long, ca. 2 mm thick, hirtellous; basal bracts ca. 4 mm long, persistent; receptacle ellipsoid to subglobose, when dry 1.5-3 cm in diameter when fresh, green with cream spots, almost glabrous, at maturity greenish (?).

Distribution. From N. Angola to E. Zaire and to Cameroun; also in West Africa (lvory Coast). In evergreen (e.g. rain) forest; at low altitudes.

40. Ficus vasta Forssk. (fig. 25) Forsskäl, FI. Aegypt.-Arab.: 179 (1775); Aweke, Meded. Landbouwhogeschool Wageningen: 88, fig. 21 (1979); Berg et al., FI. Trop. East Africa, Moraceae: 64 (1989).

F callabatensis Warb. (1905), F dahro DeWe (1 843), F hararenis Warb. (1905), F rivae Warb. (1905) , F socotrana Balfour (1884), F vasta var. glabrescens Huteh. (1917), F vasta var. velutina Fiori (19IOa).

Tree up to 25 m tall, (secondarily?) terrestrial or hemi-epiphytic. Leafy twigs 6-15 mm thick, yellowish to whitish or to brownish hirsute to hirtellous, peri­ derm flaking off. Lamina cordiform to ovate to suborbicular to elliptic or to subreniform, (5-)8- 25(-35) x (2.5- )4-23(- 25) cm, coriaceous, apex rounded to obtuse or very shortly and bluntly acuminate, base cordate, margin (sub)entire; upper surface (rather) sparsely hirtellous to puberulous or almost glabrous, lower surface whitish subvelutinous or densely to sparsely hirtellous to pu­ berulous or almost glabrous; lateral veins 5-10 pairs, basal pair branched, reach­ ing the margin usually at or above the middle of the lamina, tertiary venation for the greater part scalariform; petiole (1.5- )3-12(-19) cm long, 1.5-4 mm thick, puberulous to hirtellous, epidermis flaking off; stipules 2- 5(- 8.5) cm long, whitish to yellowish to brownish subhirsute to subsericeous, caducous. Figs in pairs or solitary in the leafaxils or just below the leaves, subsessile or up to 0.6 cm long pedunculate; basal bracts 3.5-4.5 mm long, free parts caducous; re­ ceptacle subglobose to ellipsoid, wh en fresh 2-2.5 cm, when dry ca. 1-1.5 cm

104 Fig. 25. Ficus vasta Forssk. I, leafy twig with figs; 2, figs; 3--4, staminate flowers; 5, stamen; 6-10, pistillate flowers; 11 , stipules. After Aweke (1979, fig. 21). in diameter, white to yellowish velutinous to sparsely hirtellous, often ± warty, at maturity green with paler spots.

105 Distribution. N. Uganda, N. Kenya, S. Sudan, Ethiopia, and Somalia, also in the southem part of the Arabian Peninsuia and Socotra. In riverine vegetation, at altitudes up to 2000 m. The pollinator is Elisabethiella socotrensis (Mayr). Parasitoids are: Cros­ sogaster triformis Mayr, Sycoryctes coccothraustes Mayr and S. truncatus Mayr.

41. Ficus wakefieldii Huteh. Hutchinson, Kew Bull. (1915): 335, with plate (1915); Berg et al., F1. Trop. East Africa, Moraceae: 65 (1989).

Tree up to 25 m tall, (secondarily?) terrestrial, with a wide crown. Leafy twigs (3-)5-12 mm thick, with minute to short hairs, intermixed with much longer yellow to brownish hairs, periderm flaking ofT. Lamina cordiform to ovate to (broadly) elliptic to suborbicular to subreniform or sometimes to broadly obovate, 6-23 x 5-23 cm, (sub)coriaceous apex rounded or sometimes very shortly and bluntly acuminate, base cordate, margin (sub)entire; upper surface sparsely hirtellous to subhirsute, lower surface sparsely to densely hirtellous to puberulous, on the main veins yellow hirsute; lateral veins 5-8 pairs, (main) basal pair branched, reaching the margin at or just above the rniddle of the lamina, tertiary venation partly scalariform; petiole 2-5.5(-9) cm long, (1.5-)2- 4(- 5) mm thick, hirtellous to hirsute, epidermis flaking ofT; stipules 0.5-1.5 cm, on flush up to 4 cm long, yellow to brownish hirsute to subsericeous, caducous. Figs in pairs in the leafaxils, initially enclosed by up to 1.5 cm long ovoid hairy calyptrate bud covers, sessile; basal bracts 3-5 mm long, persistent; receptacle (sub)globose, when fresh (1.2-)1.5-2 cm, when dry (0.8-)1- 1.5 cm in diameter, densely white to yellow pubescent to (sub)hirsute or to sparsely hirtellous.

Distribution. Uganda, Kenya, Tanzania, and N. Zambia. Woodland, often on termite mounds or in rocky places; at altitudes up to 2000 m. The pollinator is Elisabethiella socotrensis (Mayr), as also for F. vasta.

42. Ficus glumosa Delile (fig. 26) Delile, Cent. PI. Méroé: 63 (1826); Berg et al., F1. Cameroun 28: 170 , fig. 58 (1985); Berg et al. , F!. Trop. East Africa, Moraceae: 65 (1989).

F. barbata Warb. (1894), F. durandiana Warb. (1904), F. fazokelensis (Miq.) Miq. (1867), F. glumosa var. glaberrima Martelli (1886), F. glumosa var. intermedia Martelli (1886), F. glumosa var. lanu­ ginosa Martelli (1886), F. glumosoides Huteh. (1915), F. gombariensis De Wild. (1913a), F. kitaba De Wild. (1914), F. montana Sim (1909), F. rehmannii Warb. (1906), F. rehmannii var. ovatifolia Warb. (1906), F. rehmannii var. vil/osa Warb. (1906), F. rubicunda (Miq.) Miq. (1867), F. rukwaensis Warb. (1901), F. sonderi Miq. (1867), Sycomorus hirsuta Sonder (1850), Urostigmafazokelense Miq. (1847) , U. glumosum (Delile) Miq. (1847), U. rubicundum Miq. (1847).

Tree up to lOm tall or shrubs, terrestrial, with spreading branches. Leafy twigs 2-8 mm thick, with dense short white hairs, especially on the nodes intermixed

106 3

6 g ó Fig. 26. Ficus glumosa Delile. I, leafy twig with figs; 2, leaf; 3, ostiole; 4, staminate fiower; 5--6, pistillate fiowers. After Aweke (1979, fig. 6). with much longer bright yellow (or sometimes to whitish) hairs or glabrous, periderm of older parts flaking off. Lamina oblong to (broadly) elliptic to (broadly) ovate, sometimes to obovate or suborbicular, 2- 14(-19) x 1.2-9.5(- 13) cm, subcoriaceous, apex shortly acuminate to subacute or to subobtuse, base

107 cordate, sometimes rounded, margin entire; upper surface puberulous to hirtel­ lous to subtomentose, or al most glabrous, lower surface subtomentose on the ma in veins often partly yellow hirsute or entirely glabrous; lateral veins 3-7(-8) pairs, basal pair (faintly) branched, reaching the margin usually (far) below, sometimes (in broad leaves) at or just above the middle of the lamina, tertiary vena ti on reticulate or tending to scalariform, vein-reticulum beneath incon­ spicuous; petiole 0.5-4(-8) cm long, 1-2(-3) mm thick, puberulous and partly also (sub)hirsute or sometimes glabrous; stipules 0.5-1.5 cm, on flush up to 4 cm long, at least partly or rather sparsely bright yellow (or sometimes almost whitish) hirsute to subsericeous, caducous. Figs in pairs in the leafaxils or down to ca. 1 m below the leaves, (sub)sessile or sometimes up to 0.3 cm long peduncu­ late; basal bracts ca. 3 mm long, persistent; receptacle globose to ellipsoid, when fresh ca. 1-1.5 cm, when dry 0.5-1 cm in diameter, densely tomentose to pu­ bescent to al most glabrous, at maturity orange to red, or pink, often with darker spots.

Distribution. From Namibia, Sw. Angola and South Africa to Ethiopa and the Sudan, and westwards to Sénégal; also in Yemen and Saudi Arabia. In savanna woodland, often in rocky places; at alti tud es up to 2000 m. The material from the southern part of the range usually has a rather dense indumentum on the leaves, etc., while almost glabrous specimens are common in the northern part of the range (Sénégal to Ethiopia). F glumosa can be confused with F stuhlmannii. In contrast with that species the reticulum of smaller veins is (almost) plane beneath and part of the hairs on the twigs is yellowish. The pollinator is Elisabethiel/a glumosae Wiebes.

43. Ficus stuhlmanoü Warb. Warburg, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 20: 161 (1894); Berg et al., F!. Trop. East Africa, Moraceae: 66 (1989).

F. dar-es-salaamii Huteh. (1916), F. homblei De Wild. (l9I3a), F. howardii Sim (1909).

Tree up to 10(- 15) m tall, (secondarily) terrestrial or hemi-epiphytic. Leafy twigs (2-)4-8 mm thick, (rather) densely white puberulous to hirtellous, on the nodes to pale yellow hirsute, periderm of older parts ± flaking off. Lamina oblong to elliptic to (sub)ovate or to (sub)obovate, sometimes to suborbicular, 2.5-18 x 1.8 cm, (sub)coriaceous, apex rounded to subacute or sometimes very shortly and bluntly acuminate, base cordate to rounded, margin (sub)entire; upper surface puberulous to hirtellous, lower surface densely hirtellous to sub­ tomentose on the veins; lateral veins (3- )4-7 pairs, basal pair not or faintly branched, usually reaching the margin (far) bel ow the middle of the lamina, tertiary venation reticulate, vein-reticulum prominent beneath; petiole 0.5-4 cm long; (1-)2-3 mm thick, puberulous to hirtellous; stipules 0.5-1.5 cm long, white to pale yellow subsericeous to subhirsute or puberulous, caducous. Figs in pairs

108 in the leafaxils sometimes also just below the leaves, (sub)sessile; basal bracts ca. 3 mm long; receptacle globose to ellipsoid, when fresh 1.5-2.2 cm, when dry (0.7-)1.2-1.8 cm in diameter, densely white pubescent to sparsely puberulous, at maturity pinkish or purplish.

Distribution. From Uganda and Kenya through Tanzania to Zambia, SE. Zaire (Shaba), Malawi, Mozambique, Zimbabwe, Swaziland, and South Africa. In woodland or dry evergreen forest, often on termÎte mounds; at altitudes from 500 to 2000 m. The species is rather uniform; it can be confused with F. glumosa (see above). The pollinator is Alfonsiella binghami Wiebes.

44. Ficus nigropunctata Mildbr. & Burret Mildbraed & Burret, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 46: 220, fig. 3 (191Ib); Berg et al. , Fl. Trop. East Africa, Moraceae: 67 (1989).

Shrub or tree up to 7 m tall, terrestrial or sometimes hemi-epiphytic. Leafy twigs 1.3 (-6) mm thick, puberulous to hirtellous to subtomentellous, bark of the older wood usually blackish and conspicuously lenticellate. Lamina oblong to elliptic to (sub)obovate or sometimes to ovate, 1-9.5 x 0.6-5.5 cm, chartaceous to sub­ coriaceous, apex shortly acuminate to subacute, base rounded to cordate, margin crenulate; upper surfacepuberulous to hirtellous to hispidulous, (when dry sometimes black punctate), lower surface puberulous to hirtellous on the veins; lateral veins 3- 5(-6) pairs, the basal pairs faintly branched, usually reach­ ing the margin (far) below the middle of the lamina, tertiary venation reticulate; petiole 0.3- 2 cm long, 0.5-1 mm thick; stipules 0.2-1.2 cm long, sparsely pu­ berulous to hirtellous to subtomentellous, caducous. Figs in pairs in tbe leafaxils or also on the older wood, sessile; basal bracts 2-2.5 mm long, persistent; re­ ceptacle (sub)globose, when fresh 1- 1.2 cm, wben dry 0.5-1 cm in diameter, puberulous to hirtellous, at maturity green with red spots or reddish.

Distribution. Zambia, Malawi, Mozambique, Zimbabwe, Botswana, Tanzania, and Kenya. In woodland; at altitudes up to 1300 m. This species resembles F. stuhlmannii, but its leaves are smaller and thinner and the figs are smaller. It can be easily recognized by the dried older wood, which is blackish with conspicuous lenticels. The pollinator is Alfonsiella bergi Wie bes.

45. Ficus tettensis Hutch. Hutchinson, Kew Bull. (1915): 341 (1915).

F smUlsii Verdoom (1935).

Tree up to 6 m high or shrub, mostly (hemi-)epilithic, with roots penetrating

109 rock-crevices. Leafy twigs 2-6 mm thick, whitish hirtellous to pubescent (with hairs of different length, but not falling into two very distinct categories), peri­ derm of ol der parts flaking off. Lamina subreniform to cordate, 1.5--11.5 x 2-12 cm, (sub)coriaceous to chartaceous, apex shortly to longly acuminate to rounded, base cordate (to emarginate), margin subentire or (irregularly) crenate; upper surface ± densely hirtellous to pubescent, often somewhat scabridulous, lower surface hirtellous to subtomentose on the veins; lateral veins 4--6 pairs, basal pair branched, reaching the margin above the middle of th er lamina, tertiary venation reticulate (or partly scalariform), vein-reticulum prominent beneath; petiole 0.5- 3(-5.5) cm long, 1-2 mm thick, hirtellous to pubescent; stipules 0.2-0.6 cm long, white to brownish pubescent (to subsericeous), caducous. Figs in pairs or solitary in the leafaxils, subsessile or up to 2 mm long pedunculate; basal bracts 2, 2-3 mm long, basally connate, the upper parts caducous; receptacle globose, when dry 0.5- 1 cm in diameter, densely white pubescent to puberulous.

Distribution. Mozambique, Zimbawe, Botswana, and South Africa (Transvaal). On rocks. The species mayalso occur on Socotra. A sterile collection (Balfour 451, Dec. 1883) resembles material of this species, and not material of any species known from that island. This species can be easily recognized by the hairy, subreniform to cordiform leaves. The pollinator is Nigeriella excavata Compton.

46. Ficus mueUeriana C. C. Berg Berg, Kew Bull. 43: 85, fig. 3 (1988).

Shrub. Leafy twigs 2.5- 3 mrn thick, densely white puberulous, when dry red­ brown. Larnina ovate, 4.5-9.5 x 3- 5 cm, coriaceous to subcoriaceous, apex acute to subacuminate, base cordate, margin entire; upper surface sparsely, on the midrib more densely white puberulous, lower surface on the midrib white puberulous, on the lateral veins to white hirtellous, on the smaller veins to white tomentose; lateral veins 5-7(- 8) pairs, basal pair branched, reaching the margin bel ow (or at) the middle of the lamina, tertiary venation reticulate, venation above obscure, beneath prominent; petiole (1.2- )3.5- 5.5 cm long, 1-1.5 mm thick, puberulous; stipules 1- 1.5 cm long, minutely puberulous, caducous. Figs in pairs in the 1eafaxils, subsessile; basal bracts 1.5--2 mm long, persistent; re­ ceptacle subglobose, when dry ca. 0.5 cm diameter, densely white puberulous.

Distribution. Mozambique (Manica e Sifala), only known from two collections.

47. Ficus abutilifolia (Miq.) Miq. Miquel, Ann. Mus. Bot. Lugd.-Bat. 3: 288 (1867); Berg et al., Fl. Cameroun 28: 162, fig. 52 (1985); Berg et al. , Fl. Trop. East Africa, Moraceae: 67 (1989).

110 F. discifera Warb. (1905), F. kerstingii Huteh. (1916), F. /edermannii Huteh. (1915), F. mittuensis Warb. (1904), F. picta Sim (1909), F. so/danella Warb. (1906), Urostigma abutilifolium Miq. (1847), U. cata/paefotium Miq. (1847).

Tree up to 15 m tall, terrestrial, often (hemi-)epilithic. Leafy twigs 6-10 mrn thick, glabrous or yellowish to white tomentose to tomentellous to puberulous, periderm often flaking ofT. Lamina cordiform to broadly ovate to subreniform, 6-9 x 5-20 cm, (sub)coriaceous, apex shortly acurrunate to subacute to obtuse or to rounded, base cordate, margin entire; upper surface glabrous or with sparse hairs on the main veins, lower surface puberulous to subtomentellous, sometimes only in the axils of the lateral veins, or occasionally entirely glabrous; lateral veins 7-9 pairs, basal pair branched, reaching the margin above or at the rruddle of the lamina, tertiary venation partly scalariform; petiole 2- 10(-18) cm long, 2-4 mrn thick, puberulous to glabrous; stipules 0.5-2 cm long, puberulous or glabrous, caducous. Figs in pairs or sometimes with 4 together in the leafaxils or just below the leaves; peduncle 0.3- 1.5 cm long; basal bracts 3- 3.5 mm long, free parts persistent or caducous; receptacle subglobose, obovoid or ellipsoid, when fresh 1.2-2 cm, when dry 0.5- 1.5 cm in diameter, sparsely minutely pu­ berulous, at maturity reddish with yellowish spots (or entirely yellowish?).

Distribution. From South Africa (Transvaal), Botswana and S. Zambia to Tanzania, and from N. Kenya and Somalia westwards to Guinea. Mostlyon rocks in areas with savanna woodland; at altitudes up to ca. 1000 m. The material from the southem part of the range is distinet from that of the northem part of the range (Guinea - Somalia) in the persistence of the basal bracts; moreover, the plants are (alrnost) glabrous, while often hairy in the northem part of the range. The leaves of F abutilifolia resembie those of F tettensis, but are larger (up to 17 x 18 cm as opposed to up 12 x 12 cm) and (sub)glabrous. Two species were recorded as pollinators, viz., Elisabethiella comptoni Wiebes and Nigeriella fusciceps Wiebes.

48. Ficus populifolia Vahl Vahl, Enum. 2: 181 (1805); Berg et al., F1. Carneroun 28: 160, fig. 53 (1985); Berg et al, F1. Trop. East Mrica, Moraceae: 68 (1989).

F. intermedia Delile (1826), F. popu/ifolia var. major Warb. (1905), F. popu/ifolia var. soma/ensis Warb. (1905), F. popu/ifolia var. taitensis Warb. (1905), F. religiosa Forssk. (1775), non L. (1753), Urostigma popu/ifolium (Vahl) Miq. (1847).

Tree up to 1O(-30?) m taU, terrestrial. Leafy twigs 3- 10 mrn thick, glabrous or white puberulous. Lamina cordiform to broadly ovate to (sub)reniform, 3-17 x 3-14 cm, (sub)coriaceous, apex (longly) acuminate, base cordate to truncate, margin entire to repand; both surface glabrous; lateral veins 6-11 pairs, basal pair branched, reaching the margin below or sometimes at the middle of the

lil lamina, tertiary venation; petiole 2.5-12 cm long, 1-2 mm thick, glabrous; stipules 1-4 cm long, glabrous or ciliolate, caducous. Figs in pairs or solitary in the leafaxils; peduncle 0.8- 2 cm long; basal bracts ca. 2 mm long, free parts caducous; receptacle obovoid to ellipsoid to subglobose, when fresh 1- 1.5 cm, when dry 0.5-1 cm in diameter, minutely puberulous to glabrous, when dry often with distinct ribs, at maturity green with red spots.

Distribution. From Ghana to Ethiopia, Kenya and N. Tanzania; also in Yemen. In savanna wood land, often in rocky places; at altitudes up to 1600 m.

49. Ficus jansii Boutique Boutique in Lebrun & Boutique, Bull. Jard. Bot. Brux. 19: 211 (1949); Berg et al., FI. Cameroun 28: 166, fig . 56 (1985).

Tree up to 7 m tall, terrestrial (?). Leafy twigs 6-10 mm thick, rather densely minutely puberulous to subglabrous. Lamina oblong to elliptic, 16-27 x 9-16 cm, coriaceous, apex rounded to subacuminate, base subacute, margin subentire (to repand?); both surfaces glabrous; lateral veins 10-14 pairs, loop-connected 3- 5 mm from the margin, the lower ones small, rnidrib usually not reaching the apex of the lamina, tertiary venation (almost) reticulate; petiole 3-5.5 cm long, 3-4 mm thick, glabrous; stipules 0.3- 1 cm long, glabrous, those of the tig-bear­ ing spurs white to brownish puberulous, caducous. Figs to 5 on short spurs in the leafaxils or just below the leaves; peduncle 1.5- 2 cm long; basal bracts (free parts) caducous, connate parts persistent, forming a di sc of 4-5 mm in diameter; receptacle subglobose, when dry 0.7- 1 cm in diameter, warty, puberulous, at maturity yeUowish (?).

Distribution. Known from Zaire and Cameroun. In rain forest; at altitudes up to 1500 m. This species may prove not to belong to subsection Platyphyllae, but related to F. adolfi-friderici in subsection Crassicostae.

50. Ficus trichopoda Baker Baker, J. Lino. Soc., Bot. 20: 261 (1883); Berg et al. , FI. Cameroun 28: 158, fig. 52 (1985); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 41 (l986a); Berg et al., FI. Trop. East Africa, Moraceae: 68 (1989). F. budduel/sis Huteh. (1915), F. cOl/geI/sis Eng!. (1886), F. congensis var. mollis Huteh. (1917), F. fiavovenia Warb. (1894), F. hippopotami Gerstner (1943), F. zuvalensis Sim (1909).

Shrub or tree up to 10(- 20) m taU, terrestrial, often with stilt-roots (or 'piUar­ roots') on the main branches. Leafy twigs 3- 7 mm thick, glabrous or white puberulous to hirtellous, when dry brown to blackish with conspieuous lenticels. Lamina broadly ovate to eUiptic, 6-20(- 28) x 4-12(-21) cm, coriaceous, apex shortly acuminate to obtuse, base obtuse to cordate, margin entire, ± revolute;

112 upper surface white puberulous to hirtellous on the main veins, glabrescent or entirely glabrous, lower surface densely white hirtellous to tomentellous, at least on the midrib, or sometimes glabrous; lateral veins 7- 11 pairs, basal pair branched, reaching the margin below or (in broad leaves) sometimes at the middle of the lamina, tertiary venation partly scalariform; petiole 2-4(-7) cm long, (1 - )2- 3 mm thick, glabrous or white puberulous to hirtellous; stipules 1.5-4.5 cm, on flush up to 8 cm long, densely white puberulous to hirtellous, caducous. Figs up to 4 together in the leafaxils; peduncle 0.5-1 cm long; basal bracts ca. 2 mm long, persistent; receptacle (sub)globose, when fresh 1-2 cm, when dry 0.5-1.5 cm in diameter, glabrous to densely white puberulous, ± warty, at maturity red to yellowish.

Distribution. From South Africa through eastern Africa and from N. Zambia to the Sudan and westwards to Sénégal; also in Madagascar. In savanna woodland, in marshy places, at altitudes up to 1200 m. This species often forms stilt-roots or 'pillar-roots' on the branches, a feature apparently related to its normal habitat. The pollinator is E/isabethiella bergi Wiebes (W. Africa) and its subspecies E. b. breviceps Wiebes (southern Africa and Madagascar).

51. Ficus grevei Baill. Baillon in Grandidier, Hist. Madag., Bot. Atlas 3: fig. 297 (1 895); Berg, Bull. Mus. natn. Hist nat. Paris (4) 8 (B): 42 (1986a).

Tree up to 20 (or more) m tall, (usually terrestrial?). Leafy twigs 2.5--6 mm thick, glabrous or minutely puberulous. Lamina elliptic to ovate to subcordiform, or sometimes oblong to subovate, 6-16(- 25) x 4.5-13.5(- 16.5) cm, coriaceous, apex obtusely and shortly acuminate, base cordate to rounded, margin entire; upper surface glabrous or puberulous on the midrib, lower surface (sparsely) puberulous to hirtellous on the veins; lateral veins 6-9 pairs, basal pair (faintly) branched or unbranched, reaching the margin below (or at) the middle of the lamina, tertiary venation reticulate; petiole 2.5-8 cm long, 2- 2.5(-3) mm thick, glabrous or minutely puberulous; stipules (0.5- )1--6 cm long, glabrous or minutely puberulous, caducous. Figs in pairs or solitary in the leafaxils or just below the leaves; peduncle 1-2 cm long, sometirnes cupula-like broadened at the apex; basal bracts caducous; receptacle (sub)globose, when dry (0.8- )1 - 1.7 cm in diameter, puberulous to hirtellous.

Distribution. Madagascar. In areas with dry forest, often near water.

52. Ficus rubra Vahl (fig. 27) Vah1, Enum. 2: 191 (1805); Berg et al., Fl. Mascareignes, Moracées: 11 , fig . 4 (1985); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 42 (l986a).

113 Fig. 27. Ficus rubra Vahl. I, leafy twig; 2, leafy twig with figs; 3-4, figs. After Berg & van Heusden (1985, pI. 4).

F. avi-avi Blurne (1825), F. cinerea Cordem. (1895), F. consimilis Baker (1877), F. pyrifolia Lam. (1788), non Burman (1768), Urosligma rubrum (Vahl) Miq. (1847).

114 Tree up to 8 m tall, hemi-epiphytic or (secondarily?) terrestrial. Leafy twigs 3-5 mm thick, glabrous or sometimes sparsely and minutely puberulous, when dry reddish-brown. Lamina ovate to subovate to (broadly) elliptic, 2.5- 10(-16) x 1.5-6.5(-8) cm, (sub)coriaceous, apex obtuse to subacute or to shortly acu­ minate, base rounded to truncate or to emarginate, margin entire; both surfaces glabrous; lateral veins 6-10(- 12) pairs, basal pair unbranched (or faintly branched) running almost parallel to the margin bel ow the middle of the lamina, tertiary venation reticulate; petiole 0.7-4.5(-7) cm long, ca. 2 mm thick, glabrous; stipules 0.3- 1.5 cm, on flush up to 10 cm long, glabrous, caducous. Figs in pairs in the leafaxils or just below the leaves; peduncle 0.2-0.8 cm long; bas al bracts ca. 2 mm long, persistent; receptacle globose (to ellipsoid), when dry 0.6-1 cm in diameter, glabrous or minutely puberulous, at maturity (dark) red to (dark) purple.

Distribution. N. Madagascar, Mascarene Islands, Seychelles, Aldbara Islands, and Comoro Islands. Often in rocky places. The pollinator is Nigeriella avico/a Wiebes. Parasitoids are Sycoscapter tibialis Wiebes, Sycoryctes cae/ebs Wiebes and Watshamiella fictitia Wiebes.

53. Ficus marmorata Baker Baker, J. Linn. Soc., Bot. 20: 257 (1883); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 43 (l986a).

Tree up to 15 m tall, (usually terrestriai?). Leafy twigs 1.5-3 mm thick, (rather) densely puberulous. Lamina elliptic to lanceolate or ovate to subovate, some­ times subovate, 2-8 x 1.2-4.5 cm, (sub)coriaceous, apex acumimate, base rounded to subcordate, margin entire; both surfaces glabrous; lateral veins (5-) 7- 10 pairs, basal pair unbranched (or faintly branched) running almost parallel to the margin bel ow the middle of the lamina, tertiary venation reticulate; petiole (0.8-)1.5-6 cm long, 0.5- 1.5 mm thick, glabrous; stipules 0.5-1.5 cm, on flush up to 4 cm long, (sub)glabrous, caducous. Figs in pairs in the leafaxils or just below the leaves; peduncle 0.2-0.5 cm long; basal bracts ca. 2.5 mm long, per­ sistent: receptacle globose, when dry 0.5-0.8 cm in diameter, white hirtellous to puberulous, sometimes ± warty.

Distribution. Madagascar. In forest.

54. Ficus bivalvata Perrier Perrier de la Bäthie, Arch. Bot. Bull. Mens. 2: 151 (1928); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (8): 44 (l986a).

Tree up to 30 m tall, (terrestrial?). Leafy twigs 2.5-4 mm thick, minutely pu­ berulous. Lamina oblong to elliptic, 4-9.5 x 2.5-5 cm, coriaceous, apex shortly acuminate, base rounded to truncate to subcordate, margin entire; both surfaces

115 glabrous; lateral veins 10-12 pairs, basal pair unbranched (or faintly branched) running almost parallel to the margin below the middle of the lamina, tertiary venation reticulate; petiole 0.5-1.6 cm long, ca. 2 mm thick, minutely pu­ berulous; stipules 0.5- 1 cm long, puberulous, caducous. Figs in pairs in the leaf axils, subsessile or up to 0.4 cm long pedunculate; basal bracts caducous, except for their basal parts which leave a disc; receptacle ovoid to ellipsoid, when dry ca. 2-2.5 x 1- 1.2 cm, puberulous to strigillose, ± warty.

Distribution. Madagascar. Only kl)own from the type collection made in sub­ montane (cloud?) forest at 1700 m.

Ficus subgenus Urostigma section Galoglychia subsection Chlamydodorae (Mildbr. & Burret) c.c. Berg Berg, Proc. Kon. Ned. Akad. Wet. (C) 89: 123 (1986b).

Trees or shrubs, hemi-epiphytic or (secondarily) terrestrial, sometimes epilithic, bark not exfoliating. Leafy twigs, stipules, leaves and figs mostly glabrous or minutely puberulous, sometimes densely pubescent. Leaves oblong to lanceolate to elliptic or to obtriangular, sometimes ovate or cordiform, shortly to longly petiolate, tertiary venation reticulate or tending to parallel to the lateral veins, margin entire; stipules smalI, mostly caducous. Figs small to medium-sized, in the leafaxils or just below the leaves, often initially in ± well-developed ca­ lyptrate bud covers, at maturity wall mostly soft and often reddish. Fruitlets with a dry pericarp, endacarp body not released.

Subsection Chlamydodorae comprises 13 species. It is somewhat better re­ presented in eastern Africa than in other parts of Africa. Two species accur in Madagascar and neighbouring islands and eJeven species belong to the flora of the African continent. Some species are ± distinctly associated with the rain forest (e.g., F. calyptrata and F. kamerunensis), other species with savanna wood land (e.g. , F. fischeri and F. faulkneriana). F. ilicina is adapted to semi-desert conditions in South-West Africa. F. thonningii is a transitional species with a broad ecological amplitude, as some forms occur in savanna woodland, others in various types of forest, including rain forest. The pollinators belong to the genera Alfonsiella Waterston and Elisabethiella Grandi.

55. Ficus calyptrata Vahl

Vah1, Enum. 2: 186 (1805); Berg et al. , F1. Cameroun 28: 190, fig. 64 (1985), sub F. mallolOides; Berg, Kew Bull. 43: 77 (1988).

F dja/onensis A. Chev. (1917), F ma//otoides Mildbr. & Hutch. in Hutch. (1915), Urostigma ca/yp­ traturn (Vahl) Miq. (1847).

116 Shrubs or trees up to 6 m tall, hemi-epiphytic. Leafy twigs 4--6 mm thick, minutely puberulous, (when dry) hollow, axillary calyptrate buds covers, leaving ± conspieuous triangular scars. Lamina broadly ovate to elliptic to cordate or to suborbicular, 6-22 x 5.5-17.5 cm, coriaceous to subcoriaceous, apex acu­ minate to obtuse to rounded, base deeply cordate to subcordate to truncate, margin entire; both surfaces glabrous; lateral veins 7-10 pairs, often furcate far from the margin, tertiary venation reticulate; petiole 1.5- 7.5 (-10) cm long, 1.5-3.5 mm thick, glabrous or sparsely puberulous; stipules 1- 3.5 mm long, outside (minutely) puberu10us to almost glabrous, inside at least in the lower part appressed-pubescent (to -puberu1ous), caducous to subpersistent. Figs in pairs in the axils ofthe leaves or often below the leaves on previous season's growth, sessile, initially enclosed by up to 1.5 cm long axillary calyptrate bud covers, being densely hairy inside and caducous; basal bracts 2- 2.5 long, persistent; receptacle more or less depressed globose, when fresh 2.5-3 cm, when dry ca. 1.5- 2 cm in diameter, puberulous, at maturity brownish (with orange spots), smelling after rotting fish, when dry mostly ± wrinkled.

Distribution. From Cameroun to Zaire, and in West Africa. In forest; at low altitudes. The pollinator is Elisabethiella pectinata (Joseph).

56. Ficus tischeri Mildbr. & Burret

Mildbraed & BUITet, Bot. Jahrb. Syst. , Pfianzengesch & Pfianzengeogr. (EngIer) 46: 227 (1911 b); Berg et al. , Fl. Trop. East Africa, Moraceae: 70 (1989). F kiloneura Hornby (1948).

Tree up to 15 m tall, hemi-epiphytic, often soon terrestrial, with a fiat-topped crown. Leafy twigs 4-10 mm thick, glabrous or puberulous. Lamina ovate to elliptic, (4-)6.5-17 x (3- )5- 11 cm, coriaceous, apex acuminate, sometimes sub­ acute or rounded, base cordate to truncate, sometimes rounded, margin entire; both surfaces glabrous; lateral veins 9- 15 pairs, tertiary venation predominantly parallel to the lateral veins to reticulate; petiole 2.5-10 cm long, (1-)1.5-3 mm thick, glabrous; stipules 0.3-0.8 cm long, prominent. glabrous or puberulous, caducous. Figs solitary or in pairs in the leafaxils; peduncle 0.8- 1.8 cm long; basal bracts 2- 2.5 mm long, caducous; receptacle globose, when fresh 1.5-2 cm, when dry 1.5-2 cm in diameter, glabrous or minutely (brown) puberulous, at maturity yellowish green, when dry mostly ± wrinkled.

Distribution. Angola (Huila), Zambia, Mozambique, Zimbabwe, Botswana, Namibia (fide Palgrave, 1977: 108), and Tanzania. In wood land; at altitudes up to 1500 m. F fischeri differs from related species in habit. It forms a rather tall trunk and has an obovate, rather fiat-topped crown. The leaves have long petioles and ovate laminas.

117 The pollinator is Elisabethiella platyscapa Wiebes.

57. Ficus amadiensis De Wild. De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 200 (19I3a).

F calotropis Lebrun & Toussaint (1948), F kitabalu Huteh. (1915), F ndola Mildbr. (1953).

Tree up to ca. 15 m tall, (secondarily?) terrestrial. Leafy twigs ca. 5--10 mm thick, white puberulous. Lamina oblong to elliptic to (sub)ovate or to lanceolate, sometimes (sub)obovate, (4-)7-17 x (2-)3-8.5 cm, (sub)coriaceous, apex obtuse to rounded, base obtuse to (sub)cordate, margin entire; both surfaces glabrous; lateral veins (8- )10--15 pairs, tertiary venation reticulate but towards the rnidrib tending to parallel to the lateral veins; petiole (1-)1.5-5.5(--6) cm long, 1.5--2.5 mm thick; stipules 0.5--1.5 cm long, outside glabrous, puberulous or partly pu­ bescent, inside in the lower part densely pubescent, subpersistent. Figs in pairs in the leafaxils, sessile, initially enclosed by up to ca. 1 cm long axillary ca­ lyptrate bud covers, splitting into two persistent scales, being outside pubescent to puberulous and inside white pubescent to villous; basal bracts 1.5-2 mm long, persistent; receptacle (sub)globose, when dry 1.2- 2 cm in diameter, sparsely (rninutely) puberulous, at maturity yell ow with red spots to entirely reddish, when dry wrinkled.

Distribution. Zaire, Rwanda, Uganda, Kenya, Tanzania, and Cameroun (from where a single collection is known). In woodland; at altitudes up to 2100 m.

58. Ficus craterostoma Mildbr. & Burret (fig. 28)

Mildbraed & Burret, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 46: 247 (191Ib); Berg et al. , FI. Cameroun 28: 182, fig . 62 (1985); Berg et al., FI. Trop. East Mrica, Moraceae: 70 (1989).

F anomani Huteh. (1915), F furcata Warb. var. angustifolia De Wild. (I9I3b), F luteola De Wild. (19I3a), F mutantifolia Huteh. (1915), F pilosula De Wild. (19I3a), F rubropunctata De Wild. (I9I3a), F ruwenzoriensis De Wild. (1921).

Tree up to 10 m taU or shrubs, hemi-epiphytic. Leafy twigs 2- 5 mm thick, glabrous or white puberulous to hirtellous. Leaves often subopposite; larnina (narrowly) obtriangular to (sub)obovate, or to oblong to elliptic, 3-8 x 2-4.5 cm, (sub)coriaceous, apex truncate to emarginate (to 2-lobed) or to obtuse, base acute to obtuse, margin entire; both surfaces glabrous; lateral veins 5- 10 pairs, rnidrib not reaching the apex of the lamina, tertiary venation reticulate to tend­ ing to parallel to the lateral veins; petiole 0.5--2 cm long, 1- 2 mm thick, glabrous; stipules ca. 0.5 cm long, glabrous or yeUowish to white puberulous, subpersistent or caducous. Figs in pairs in the leafaxils, sessile, initially enclosed by up to 1 cm long axillary calyptrate bud covers, splitting into two subpersistent or caducous parts, being pubescent inside; basal bracts 1-1.5 mm long, persistent;

11 8 Fig. 28. Ficus craterostoma Mildbr. & BUITet. 1-2, 1eafy twigs with figs; 3, pistillate fiower with interfioral bracts; 4, staminate fiower. After Berg et al ., 1985, pI. 62). receptacle globose to ellipsoid, when fresh 0.8-1.2 cm, when dry ca. 0.5 cm in diameter, glabrous or puberulous, at maturity reddish (or yellowish), when dry mostly slightly wrinkled.

Distribution. From Angola, N. Zambia and South Africa (Natal) to Uganda

119 and westwards to Sierra Leone. In evergreen (e.g. rain or gallery) forest; at altitudes up to 2100 m. The material from Angola described as F mutantifolia difTers in the narrow leaves and might merit recognition at subspecific level. F eraterostoma is often confused with F nata/ensis and F thonningii. It difTers from F natalensis in the sessile figs. In F eraterostoma as weIl as in F nata/ensis the leaves can be subopposite and on the same twig the petioles tend to be rather short and rather constant in length. The latter two features can be used to dis­ tinguish F eraterostoma from forms of F thonningii with sessile figs. In contrast to F thonningii, the young figs are enc10sed in up to I cm long calyptrate buds; the halves of the bud cover are often subpersistent. The pollinator is Alfonsiella mieha/oudi Wiebes, the same species as that of F lingua.

59. Ficus lingua De Wild. & T. Durand

De Wildeman & T. Durand, Ann. Mus. Congo (3) 2: 216 (1901); Berg et al ., Fl. Cameroun 28: 180 (1985); Berg, Kew Bull. 43: 85 (1988); Berg et al., Fl. Trop. East Africa, Moraceae: 72 (1989).

Tree up to 30 m tall, hemi-epiphytic or (secondarily?) terrestrial, often sublia­ nescent. Leafy twigs 1-4 mm thick, whitish or brownish puberulous. Leaves often subopposite; lamina oblanceolate to subobovate or narrowly obtriangular 0.5- 5 x 0.3-2(- 3) cm, (sub)coriaceous, apex obtuse to truncate to emarginate or to subacute, base cuneate ta obtuse, margin entire; both surfaces glabrous; lateral veins 5--8 pairs, rnidrib usually not reaching the apex of the larnina, tertiary venation reticulate to ± parallel to the lateral veins; petiole 0.2-0.8 cm long, 0.5--1 mm thick, glabrous; stipules 0.2-0.5 cm long, puberulous or only ciliolate, caducous or subpersistent. Figs in pairs in the leafaxils or just below the leaves, 0.2-0.5 cm long pedunculate or sometimes subsessile; basal bracts 1- 2 mm long, persistent or caducous; receptac1e globose, sometimes ta ellipsoid, when fresh ca. 0.5, when dry 0.3-0.4 cm in diameter, minutely puberulous, at maturity reddish or yellowish, when dry smooth or slightly wrinkled.

Twa allopatric subspecies can be recognized.

59a. Ficus lingua subsp. lingua

F buxifolia De Wild. (19\3b), F micropison Mildbr. (1953).

Leafy twigs with brownish hairs; stipules usually subpersistent. Pedunc1e usually glabrous; basal bracts caducous.

Distribution. From Uganda to Cameroun, and also in West Africa (lvory Coast). In forest; at altitudes up to 1200 m. The pollinator is Alfonsiella mieha/oudi Wiebes (see also with F eraterostoma).

120 59b. Ficus lingua subsp. depauperata (Sim) c.c. Berg Berg, Kew Bull. 43: 85 (1988).

F depauperata Sim (1909).

Leafy twigs with whitish hairs; stipules caducous. Pedunc1e usually minutely puberulous; basal bracts persistent.

Distribution. Malawi, Mozambique, Tanzania, and Kenya. In forest, coastal bushland and coral outcrops; at low aItitudes.

60. Ficus natalensis Hochst.

Hochstetter, Flora 28: 88 (1845); Berg et al., Fl. Cameroun 28: 184, fig. 63 (1985); Berg, Kew Bull. 43: 87 (1988); Berg et al., Fl. Trop. East Africa, Moraceae: 71 (1989).

Tree up to 30 m taU or shrub, hemi-epiphytic or terrestrial, sometimes lia­ nescent. Leafy twigs 2-5 mm thick, glabrous or sparse\y minute\y puberulous (or hirtellous). Leaves often subopposite; lamina oblong to elliptic to (sub)obovate to (broadly) obtriangular (or to lanceolate), 2.5-10 x 1-4.5 cm, (sub)coriaceous, apex (faintly and shortly) acuminate to obtuse to subacute to rounded or to emar­ ginate, base acute to obtuse, margin entire; both surfaces glabrous; lateral veins 6-13 pairs, midrib usually not reaching the apex of the lamina, tertiary venation reticulate to parallel to the lateral veins; petiole 0.2- 2(-3) cm long, 1-2(-2.5) mm thick, glabrous; stipules 0.2-1 cm long, glabrous or hairy, caducous. Figs in pairs in the leafaxils or sometimes also just below the leaves, initially enc10sed by smaU or sometimes up to 1.5 cm long (almost) glabrous axillary calyptrate bud covers, being caducous; pedunc1e 0.2- 1 cm long; basal bracts 2-2.5 mm long, caducous; receptac1e globose to ellipsoid to obovoid, when fresh ca. 1.5-2 cm, when dry 0.5-1.5 cm in diameter, (at least when dry often shortly stipitate), glabrous, at maturity reddish, orange or yellowish (to brown), when dry usually wrinkled and apex plane or slightly protruding.

Two allopatric subspecies can be recognized.

6Oa. Ficus natalensis subsp. natalensis

F. durbanii Warb. (1906), F. mammosa Lebrun (1934), F natalensis var. latifoUa Warb. (1906), F natalensis var. pedunculata Sim (1909), F. scutata Lebrun (1934), F subacuminata (De Wild.) Lebrun (1934), F variabilis De Wild. forma obtusifo/ia De Wild. (1921), F variabilis forma suba­ cuminata De Wild. (1921), F volkensii Warb. (1894), Urostigma natalense (Hochst.) Miq. (1847).

Tree or shrub. Leafy twigs glabrous or sparsely puberulous. Lamina oblong to elliptic to (sub)obovate, sometimes broadly obovate to obtriangular (or lanceo­ late), apex acuminate to rounded to emarginate, margin plane, end of the midrib usually near the apex of the lamina, lateral veins 6-13 pairs; stipules glabrous or sparse\y, minutely puberulous. Calytrate bud covers enc10sing the young figs

121 often conspicuous, sometimes up to I cm long. Figs when dry 0.8-1.5 cm in diameter; basal bracts caducous.

Oistribution. In South Africa (Natal), Mozambique, E. Zimbabwe, Malawi, N. Zambia (Copperhelt Province), Tanzania, Kenya, Uganda, and E. Zaire. In wet and dry forest, also in wood land (often in rocky places); at altitudes up 2200 m. Often planted. This taxon, often confused with F thonningii (and F craterostoma), can be distinguished by the caducous basal bracts and the (usually) glabrous stipules. Moreover, the leaves may he subopposite (normally altemate in F thonningit), the petioles are rather short and on the same twig fairly constant in length (in F thonningii mostly variabie in length), and the midrib rarely reaches the apex of the lamina (in F thonningii it usually does reach the apex). The pollinator is Elisabethiella socotrensis Mayr, but also Alfonsiella lon­ giscapa Joseph was recorded. In some other instances there may have been confusion over the status of the host tig: (natalensis-)group or species?

6Ob. Ficus natalensis subsp. leprieurü (Miq.) c.c. Berg Berg, Kew Bull. 43: 88 (1988).

F. brevipedicellata De Wild. (1921), F. chrysocerasus Warb. (1894), F. excentrica Warb. (1894), F. furcata Warb. (1894), F. leprieurii Miq. (1867), F. leprieurii var. intermedia Huteh. (1916), F. leprieurii var. sessilis Huteh. (1916), F. triangularis Warb. (1894).

Tree or shrub, sometimes lianescent. Leafy twigs glabrous, minutely puberulous or sometimes white to yellowish hirtellous. Lamina elliptic to oblong to broadly obovate to obtriangular, revolute, end of the midrib often far from the apex of the lamina, lateral veins (3- )6--10 pairs; stipules usually white to yellowish sub­ sericeous to hirtellous, at least at the base (leaving a fimbriate scar), sometimes puberulous or almost glabrous. Calyptrate bud covers enclosing the young tigs minute. Figs when dry 0.5-1 cm in diameter; basal bract caducous, occasionally subpersistent.

Oistribution. From N. Angola and NW. Zambia (NW) through Zaire and Sénégal. Often in periodically inundated riverine forest or in marshy areas, also in rain forest or in woodland; at altitudes up to 1200 m. This taxon differs from subsp. natalensis in the presence of white to yellowish hairs on the stipules and sometimes also on the leafy twigs. The leaves are more often broadly obovate (or even broadly obtriangular). Oue to the presence of hairs on the stipules, this subspecies can be confused with F thonningii; on the other hand, it is more easily distinguishable because of the differences in the shape and venation of the lamina. The pollinator is Alfonsiella fimbriata Waters ton, the same species as that of F kamerunensis. A parasitoid is Phagoblastus /iodontus Wiebes.

122 61. Ficus faulkneriana c.c. Berg Berg, Kew Bull. 43: 83, fig. 2 (1988); Berg et al. , Fl. Trop. East Africa, Moraceae: 72 (1989).

Tree up to 30 m tall, (secondarily?) terrestrial. Leafy twigs 1.5-3 mm thick, minutely puberulous. Leaves sometirnes nearly subopposite; lamina 1.5- 7 x 0.8- 3.5(-4.5) cm, (sub)coriaceous, apex rounded to obtuse, sornetimes very shortly and bluntly acuminate or emarginate, base rounded to cordulate or to obtuse, margin entire; both surfaces glabrous; lateral veins (3-)5-8 pairs, rnidrib not reaching the apex of the lamina, tertiary venation reticulate to parallel to the lateral veins; petiole 0.2-1(-1.5) cm long, 1- 1.5 mm thick, glabrous; stipules 0.2-0.6 mm long, sparsely puberulous or only ciliolate, often subpersistent. Figs in pairs in the leafaxils or just bel ow the leaves; peduncle (0.3- )0.5-1.5 cm long, sparsely minutely puberulous; basal bracts 1.5-2 mm long, persistent; receptacle (sub)globose to obovoid, when dry 0.4-0.8(-1.2) cm long, subglabrous, at ma­ turity reddish or yellowish, when dry smooth.

Distribution. Tanzania and Kenya. In coastal bush and wooded grassland; at low altitudes.

62. Ficus burtt-davyi Huteh. Hutchinson, Kew Bull. (1916): 232 (1916); van Greuning, S. Afr. J. Bot. 56, 614 (1990).

F. nata/ensis Hochst. var. minor (Sonder) Warb. (1906), F. nata/ensis var. puheru/a Warb. (1906), Urostigma nata/ense (Hochst.) Miq. var. minor Sonder (1850).

Tree up to 8 m tall, (scrambling) shrub or ± lianescent, hemi-epiphytic, hemi­ epilithic or terrestrial. Leafy twigs 1.5- 3 mm thick, (rather) densely, minutely puberulous, (wh en dry) hollow. Lamina elliptic to obovate or sometimes to oblong, ovate or suborbicular, (0.5-)1.5-10 x (0.3-)0.7-4(-5.5) cm, coriaceous, apex very shortly and bluntly (occasionally sharply) acuminate to subacute or to obtuse, base rounded and usually with a small noteh, about as wide as the petiole, margin entire, revolute, especially towards the base of the lamina; upper surface glabrous, lower surface glabrous or very sparsely, minutely puberulous on the midrib; lateral veins (3-)5-6(-7) pairs, midvein reaching the apex of the lamina, tertiary venation reticulate, the subquadrangular reticulum conspieuous beneath; petiole 0.4-1.5(-2.7) cm long, 1- 1.5 rnrn thick, (sparsely) minutely pu­ berulous; stipules 0.2-0.8 cm (on flush up to 1.5 cm) long, ciliolate, caducous. Figs in pairs in the leafaxils; peduncle 0.1-0.4(-0.7) cm long; basal bracts 1- 1.5 rnm long, basally connate, persistent; receptacle globose, when fresh ca. 1- 1.5 cm, when dry 0.4-1.2 cm in diameter, sparsely minutely puberulous, at rnaturity dark-brown to black, when dry smooth.

Distribution. Mozambique (Maputo) and South Mrica (Cape Province, Natal).

123 Coastal forest or scrub, in rocky places and riverbanks; at altitudes up to 1000 m. The pollinator is Elisabethiel/a baijnathi Wiebes. Parasitoids are Otitesel/a sesquianel/ata van Noort and 0. uluzi Compton.

63. Ficus ilicina (Sonder) Miq. Miquel, Ann. Mus. Bot. Lugd.-Bat. 3: 289 (1867).

F guerichiana Eng!. (1894), Urosligma ilicinum Sonder (1850).

Tree up to 6 m tall or shrub, sometimes lianescent, hemi-epilithic or terrestrial (?); sterns white. Leafy twigs 2-6 mm thick, glabrous or puberulous. Lamina oblong to lanceolate or to (sub)ovate or elliptic, 4-9 x 1.2-4 cm, (thickly) coriaceous, apex obtuse to rounded or to subacute, base obtuse to rounded (and with a small notch) or to subcordate, margin entire; upper surface glabrous, lower surface glabrous or sparsely puberulous on the midrib; lateral veins (3-)7- 10 pairs, tertiary venation reticulate to parallel to the lateral veins; petiole 0.2- 1.8 cm long, 1.5-2.5 mm thick, glabrous or sparsely puberulous; stipules 0.5-1 cm long, glabrous or puberulous, caducous. Figs 1-2(-4) in the leafaxils, peduncle 0.3-0.6 cm long; basal bracts ca. 2.5 mm long, minutely puberulous, persistent; receptacle (sub)globose, when fresh 1.5-2 cm, wh en dry 0.8- 1 cm in diameter, sparsely to densely puberulous, at maturity yellow, when dry smooth.

Distribution. Sw. Angola, Namibia, and South Africa (NW. Cape Province). In (semi-)desert, in rocky places, against rock faces; at altitudes 1300 m. The pollinator is Elisabethiel/a enriquesi (Grandi).

64. Ficus antandronarum (Perrier) C.c. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 45 (1986a).

F pyrifolia Lam. var. anlandronarum Perrier (1928).

Shrub or tree up to 20 m tall. Leafy twigs 2- 3 mm thick, glabrous or minutely puerulous. Lamina elliptic to oblong to (sub)obovate (or to suborbiculate), 2- 7 x 1.5- 5 cm, coriaceous, apex shortly acuminate to obtuse to subacute or to rounded, base acute to rounded, margin entire, often ± revolute; both surfaces glabrous; lateral veins 4-8 pairs, tertiary venation reticulate; petiole 0.3- 1.5(-2) cm long, 1- 1.5 mm thick, glabrous or minutely puberulous; stipules 0.2-0.5 cm long, glabrous or minutely puberulous, caducous. Figs in pairs in the leafaxils, subsessile or up to 0.4 mm long pedunculate; basal bracts 1.5- 2.5 mm long, persistent; receptacle when dry 0.4-0.5 m or 0.7-1.3 cm in diameter, glabrous, when dry smooth, often dark brown to blackish.

Two subspecies can be recognized.

124 64a. Ficus antandronarum subsp. antandronarum

F pyrifolia Lam. var. pedunculata Perrier (1928).

Lamina usually elliptic to obovate, sometirnes to suborbicular. Figs subsessile or up to 4 mrn long pedunculate; receptacle when dry (0.7-)0.8- 1.3 cm in diam­ eter. Staminate flowers pedicellate. Interfloral bracts numerous, up to 2 mm long.

Distribution. Madagascar (East to Centra]). In forest and along streams; at altitudes up to 1600 m.

64b. Ficus antandronarum subsp. bernardü c.c. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 46, fig. 4 (1986a).

Lamina usually oblong to subobovate. Figs 0.2-0.3 mm long pedunculate; re­ ceptacle when dry 0.4-0.5 cm in diameter. Starninate flowers sessile. Interfloral bracts (very) few, up to 1 mm long.

Distribution. Comoro Islands (Anjouan, Mayotte). In forest.

65. Ficus reflexa Thunb. Thunberg, Diss. Bot. Fic.: II (1786); Berg et al., Fl. Mascareignes, Moracées: II (1985); Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 46 (1986a).

Shrub or tree up to lOm tall, terrestrial, hemi-epilithic or hemi-epiphytic (?). Leafy twigs 0.5-4(-8) mm thick, glabrous or puberulous. Lamina oblong to elliptic to (sub)obovate to (ob)lanceolate, 1.5-12 x 1-5(-5.5) cm, coriaceous, apex acute to rounded or shortly acuminate, base acute to obtuse (and emar­ ginate with a notch narrower than the petiole), margin entire; both surfaces glabrous; lateral veins 5- 11 pairs, midrib often not reaching the apex ofthe lamina, tertiary venation reticulate; petiole 0.3-3(-4) cm long, 1- 2 mm thick, glabrous (or puberulous); stipules 0.5--1.5(-2) cm long, glabrous or sparsely and minutely pu­ berulous, usually subpersistent. Figs in pairs in the leaf .axils or just bel ow the leaves, sessile; basal bracts 1.5- 2 mm long, persistent; receptacle (sub)globose, when fresh 0.6-1(- 1.5) cm, when dry 0.5-0.8(-1.2) cm in diameter, glabrous, minutely puberulous or sometimes tomentose, at maturity pink, red(dish) to red-brown or to purplish, when dry smooth or slightly wrinkled.

This species appears to be closely related to the extremely variabie, continental (complex-)species F. thonningii. The difTerences between these two species are very smalI.

Three subspecies can recognized.

125 65a. Ficus reflexa subsp. reflexa F aggregala Vahl (1805), F melleri Baker (1883), F punclata Lam. (1788), non Thunb. (1786), F pyrifo/ia Lam. var. imerensis Perrier (1928), F pyrifo/ia var. melleri (Baker) Perrier (1928), F pyrifo/ia var. occidenlalis Perrier (1928), F pyrifo/ia var. lsaralananensis Perrier (1928), F lerebrata Willd. (1806), Urostigma aggregalum (Vahl) Miq. (1847), U. rejlexum (Thunb.) Miq. (1847), U. terebralum (Willd.) Miq. (1847).

Leafy twigs glabrous or puberulous. Lamina 1.5-12 x 1- 5(- 5.5) cm; lateral veins 5- 11 pairs; stipules 0.5- 1.2(-2) cm long, puberulous or only ciliolate, subper­ sistent. Figs sessile; receptacle when dry 0.5-0.8(-1.2) cm in diameter, glabrous, puberulous or tomentose.

Distribution. Madagascar and Mascarene Islands. In various types of forest; at altitudes up to 1400 m. The pollinator is Elisabethiel/a reflexa Wiebes. Parasitoids are Phagoblastus bouceki Wiebes, Philotrypesis cnephaea Wiebes, Sycoscapter cornutus Wiebes, Sycoryctes remus Wiebes, Camarothorax brevimucro Boucek and C. hemimucro Wiebes.

65b. Ficus reflexa subsp. secheUensis (Baker) C.c. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 49 (l986a).

F rubra Vahl var. sechellensis Baker (1877), F sechellarum Summerhayes (1928).

Leafy twigs glabrous. Lamina 4-17 x 1.3-4 cm; lateral veins (6-)8- 12 pairs; stipules 0.5-3 cm long, glabrous, caducous. Figs sessile; receptacle when dry 0.4-0.7 cm in diameter, glabrous.

Distribution. Seychelles.

65c. Ficus reflexa subsp. aldabrensis (Baker) c.c. Berg Berg, Bull. Mus. natn. Hist. nat. Paris (4) 8 (B): 49 (l986a).

Faldabrensis Baker (1894).

Leafy twigs glabrous or sometimes puberulous. Lamina 2- 8.5(- 11) x 1--4 cm; lateral veins 5-8(-9) pairs; stipules 0.7-0.8(-1.5) cm long, minutely appressed­ puberulous or only ciliolate, caducous. Figs subsessile or up to 0.15 cm long pedunculate; receptacle when dry 0.4-0.6 cm in diameter, glabrous.

Distribution. Comoro Islap..ds (Anjouan, Moyote) and Aldabra Islands. The pollinator is Elisabethiel/a reflexa Wiebes. A parasitoid is Camarothorax orientalis (Wiebes).

126 ,'\ I ' - :16 ( ) ~ " ",,! , ;.<. ' .:.~ ~! 7

4

Fig. 29. Ficus rhonningii BI. I, leafy twig with figs; 2-4, pistillate flowers; 5, staminate flower; 6--7, stamens; 8, ostiole. After Aweke (1979, pI. 8).

66. Ficus thonningü BI. (fig. 29) Blume, Rumphia 2: 17 (1836); Berg et al., Ft. Cameroun 28: 175, fig. 59 (1985); Berg et al., Ft. Trop. East Africa, Moraceae: 73 (1989).

E annabonesis Mildbr. & Huteh. in Huteh. (1915), Ebasarensis Mildbr. & Burret (1911), E be-

127 quaerlii De Wild. (1914), F. bongoensis Warb. (1894), F. brevipedice/lala De Wild. var. cuneala De Wild. (1922), F. burkei (Miq.) Miq. (1867), F. bulagensis De Wild. (1921), F. chlamydodora Warb. (1894), F. cognala N.E. Brown (1921), F. crassipedice/lala De Wild. (1922), F. crassipedicel/ala forma anguslifolia De Wild. (1922), F. crassipedicellata forma boonei De Wild. (1922), F. cras­ sipedice/lata forma cuneala De Wild. (1922), F. cyphocarpa Mildbr. in Mildbr. & Burret (1911), F. dekdekena (Miq.) A. Rich. (1851), F. dekdekena var. anguslifolia A. Peter (1932), F. dekdekena var. pubiceps Mildbr. & Burret (1911), F. din/eri Warb. (1906), F. dusenii Warb. (1894), F. erici-rosenii R.E. Fries (1914), F. eriocarpa Warb. (1901), F. galpinii Warb. (1906), F. goelzei Warb. (1900), F. hochsle/leri (Miq.) A. Rich. (1851), F. hochste/leri var. glabrior Miq. (1867), F. ileophyl/a Miq. (1867), F. kagerensis Lebrun & Toussaint (1948), F. mabifolia Warb. (1894), F. mammigera R.E. Fries (1924), F. medul/aris Warb. (1894), F. microcarpa Vahl (1805), non L. f. (1781), F. neriifolia A. Rich. (1851), non J.E. Sm. (1810), F. neurocarpa Lebrun & Toussaint (1948), F. persicifolia Warb. (1894), F. persicifolia var. angustifolia Warb. (1904), F. persicifolia var. glabripes Warb. (1904), F. pelersii Warb. (1894), F. phillipsii Burtt-Davy & Huteh. (1932), F. psilopoga Ficalho (1884), F. pubicosla Warb. ex De Wild. & T. Durand (1901) and Warb. (1904), F. rhodesiaca Mildbr. & Burret (1911), F. rokko Warb. & Schweinf. in Warb. (1894), F. ruficeps Warb. (1901), F rupicola Lebrun & Toussaint (1948), F. ruspolii Warb. (1905), F. schimperi (Miq.) A. Rich. (1851), F. schimperi var. hochstetteri (Miq.) Mildbr. & Burret (1911), F. schinziana Warb. (1906), F. spragueana Mildbr. & Burret (1911), F Ihonningii var. heterophyl/a A. Peter (1932), F. tropophylon Lebrun & Toussaint (1948), Uros­ ligma acrocarpum Miq. (1847), U. burkei Miq. (1847), U. dekdekena Miq. (1847), U. hochslelleri Miq. (1847), U. schimperi Miq. (1847), U. Ihonningii (BI.) Miq.

Tree up to 15(-30) m tall or shrub, terrestrial or hemi-epiphytic. Leafy twigs 1.5-8 mm thick, minutely puberulous to hirtellous or white to brown pubescent, at least on the scars of the stipules or sometimes entirely glabrous. Leaves occa­ sionally subopposite; lamina elliptic to (ob)lanceolate to (sub)obovate to (sub)ovate, (1.5-)3-18 x (1 - )1.5- 6(- 7) cm, (sub)coriaceous, apex acuminate to obtuse to rounded, base subacute to rounded or to (sub)cordate, often slightly unequal-sided, margin entire; upper surface glabrous or sparsely (on the midrib to rather densely) puberulous to pubescent, lower surface glabrous or sparesly to densely white (to brownish) puberulous to pubescent on the whole surface, on the main veins or only on the midrib; lateral veins (5-)7-12(- 16) pairs, midrib often reaching the apex of the lamina (even in leaves with a rounded apex), tertiary venation reticulate to parallel to the lateral veins; petiole (0.5-)1-4(-6) cm long, 1-2 mm thick, often (independent of the size of the lamina or the position of the leaf on the twig) variabIe in length on the same twig, glabrous, puberulous, hirtellous or pubescent; stipules 0.3- 1(-1.5) cm long, white to brown pubescent, puberulous or only ciliolate, caducous or (on flush) subpersistent. Figs in pairs in the leafaxils or sometimes also below the leaves, sessi1e or up to 1 cm long pedunculate; basal bracts 2-4 mm long, persistent; receptacle globose to ellipsoid, when fresh 0.5-1.5(- 2) cm, when dry ca. 0.4--1.2(-1.7) cm in diameter, glabrous or sparsely to densely white to brown puberulous or pu­ bescent, at maturity reddish, yellowish or brownish, wh en dry mostly smooth or slightly wrinkled and apex plane to strongly protruding.

~istribution. From South Africa, Botswana, and Namibia to Ethiopia and Sénégal, also in the Cape Verde Islands and the islands in the Gulf of Guinea. In savanna wood land and wet or dry forest; at altitudes up to 2500 m.

128 In the present concept F. thonningii is an extremely variabie species or a com­ plex of taxa which cannot be separated satisfactorily on morphological criteria. The variation occurs in the dimension and shape of the leaves, the presence, denseness and colour of the indument on various parts of the plant, the size of the figs, and the length of the petiole. It is also found in the size of the plants, the abundance of aerial roots and the predominance of a terrestrial or hemi­ epiphytic life form. Many morphological entities ('forms') can be distinguished. Several of these 'forms' are wide-spread, others have a more restricted distribu­ tion or occur disjunctly. DifTerences in ecology can be recognized. In certain regions the morphological entities can be easily distinguished and intermediate forms appear to be absent, which suggests the occurrence of reproductive isola­ tion. However, in other regions the difTerences between those en ti ties fade away. The major morphological extremes of the variation are treated as unformal entities ('forms') below. In some cases distribution and habitat preference can be added as attributes of these 'forms'. a. 'arcocarpa' (= typical) form Leafy twigs sparsely puberulous to pubescent to subglabrous. Lamina elliptic to oblong, drying brownish or greyish, usually less than JO cm long; midrib usually reaching the apex of the lamina; stipules only ciliolate to densely whitish or brownish puberulous. Figs sessi1e or subsessile; receptacle, when dry 0.5--D.6 cm in diameter; when dry apex nor or slightly protruding.

Distribution. From Sénégal to Ethiopia, Uganda, and Burundi, also in Bioko (=Fernando Po) and Sào Tomé. Mainly in wooded grassland, at altitudes up to 1200 m. b. 'rokko' form

Largely similar to the 'arcocarpa ' form, but the lamina often longer than 10 cm; midrib reaching the apex of the lamina or not. Figs usually sessile or subsessile, sometimes up to 0.3 cm long pedicellate; receptacle when dry ca. 0.8-1.2 cm in diameter; when dry apex not or slightly protruding or (in Zimbabwe) distinctly protruding.

Distribution. From Mali and Guinea to Zaire, Ethiopia, Uganda, and southwards to Angola, moreover through Tanzania to E. Zimbabwe, in Sào Tomé and the Cape Verde Islands. In wet and dry forest, at altitudes up to 2000 m. c. 'mammigera' form Leafy twigs, as weil as other parts, sparsely puberulous to almost glabrous. Lamina usually ca. 10 cm long; stipules caducous. Figs sessile or subsessile; re­ ceptacle when dry 0.5-1.2(-1.7) cm in diameter; when dry apex ± protruding.

129 Distribution. East and southem Africa. In riverine forest or sometimes in wooded grassland and bushland; at altitudes up to 2300 m. d. 'iteophyUa' form Leafy twigs ± densely whitish pubescent to subtomentose. Lamina eUiptic to oblong to (sub)obovate, sometimes to (ob)lanceolate, drying greyish; midrib usuaUy reaching the apex of the lamina; petiole relatively long and slender; stipules densely whitish hairy. Figs pedunculate; receptacle wh en dry usually 0.6-0.8 cm, sometimes 1.2-1.5 cm in diameter; peduncle and receptacle whitish pubescent to subtomentose; apex when dry not protruding.

Distribution: Sénégal to Zaire. Usually in savanna wood land, at low altitudes. e. 'burkei' form Leafy twigs mostly densely whitish (to pale brown) puberulous to pubescent, the same indumentum on the lower leaf-surface, the petiole and/or figs . Lamina ellip­ tic to obovate or oblong to subobovate, usually less than 10 cm long; midrib reaching the apex of the lainina or not; petiole relatively short and stout. Figs subsessile to pedunculate; receptacle when dry ± 0.4--1.1 cm. in diameter; apex when dry not or only slightly protruding.

Distribution. East and Southem Africa. In wooded grassland and thickets; at alti tud es between 900 and 1800 m. The associated insects were listed by Boucek et al. (1981): pollinator Elisabethiella stuckenbergi (Grandi), and also Alfonsiella brongersmai Wiebes and A. longiscapa Joseph were recorded; parasitoids Crossogaster odorans Wiebes, Phagoblastus barbarus Grandi, Philotrypesis parca Wiebes, Sycoscapter cornutus Wiebes, Sycoryctes hirtus Wiebes and S. remus Wiebes, Watshamiella alata Wiebes, Otitesella tsamvi Wiebes, Camarothorax brevimucro Boucek, C. equicollis Boucek and C. longimucro Boucek, Sycotetra serricornis Boucek, Eu­ rytoma ficusgallae Boucek, Ficomila curtivena Boucek and F. gambiensis (Risbec), Syceurytoma ficus Boucek, Sycophila flaviclava Boucek, S. kestraneura (Masi), S. modesta Boucek, S. naso Boucek, S. punctum Boucek and S. sessilis Boucek, Ormyrus flavipes Boucek, 0. subconicus Boucek and 0. watshami Boucek. f. 'neurocarpa' form Leafy twigs, as well as the petiole, ± densely puberulous to hirtellous. Lamina mostly glabrous, elliptic to oblong, up to 6.5 x 3-5 cm.; midrib usually not reaching the apex of the lamina; petiole short, up to 1(-1.5) cm. long, stout; stipules caducous. Figs on peduncles 0.2-0.5 cm. long; receptacle when dry 0.5- 0.8 cm. in diameter, sparsely to densely puberulous to pubescent; apex when dry not or slightly protruding.

130 Distribution. Tanzania, E. Zaire, Ruanda, and Uganda. In rocky places, thick­ ets; at altitudes between 1500 and 1800 m. g. 'persicifolia' form Leafy twigs usually densely brown puberulous to pubescent. Leaves drooping (?); lam in a mostly oblanceolate to oblong, usually less than 10 cm long, often drying brown; rnidrib reaching the apex of the larnina; petiole relatively long and slender; stipules often subpersistent. Figs sessile to shortly pedunculate; re­ ceptacle 0.54).8 cm in diameter, almost glabrous; apex when dry not or slightly protruding.

Distribution. Angola and Gabon, to the Central African Republic; Uganda, and N. Zambia. Generally in forest, along lakes or by rivers, rareJy in wooded grassland; at altitudes up to 1800 m. h. 'petersii' form LargeJy sirnilar to the 'persicifolia' form, but the lamina often elliptic to obovate, often drying greenish; stipules mostly caducous. Figs sessile or subsessile; re­ ceptacle wh en dry 0.8-1 cm, with dense brown indumentum; apex when dry not or only slightly protruding.

Distribution. East and southem Africa. In deciduous wood land and bushland, rocky places; at alti tud es up to 1400 m. Some of the insects listed with the burkei-form may (also) belong here. i. 'dekdekena' form Largely similar to the 'pelersii' form, but the lamina usually narrower (subo­ bovate to oblanceolate) and the figs sessile or pedunculate and almost glabrous.

Distribution. From Ethiopia to Ivory Coast and Bukina Faso. In wooded grassland, riverine and lake-sides; at altitudes between 1000 and 2100 m. j. 'hochstetteri' form Similar to the 'dekdekena' form, but the lower surface of the lamina hairy, usually more densely so on the midrib.

Distribution. From Ethiopia to Tanzania and Zaire. In riverine forest and wooded grassland; at altitudes up to 2200 m.

67. Ficus kamerunensis Mildbr. & Burret Mildbraed & Burret, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (Engier) 46: 258 (1911 b); Berg et al., Fl. Cameroun 28: 179, fig. 60 (1985).

131 Shrubs or trees, sometimes lianescent. Leafy twigs 1-5 mm thick, glabrous or with sparse to rather dense, brown to yellow, soft or stifT hairs. Leaves some­ times nearly subopposite; lamina elliptic to lanceolate or to subobovate, (1.5-)4- 11(-14) cm long, (0.5-)2-4(- 5) cm broad, chartaceous to subcoriaceous, when dry brown and folded, apex acuminate to acute, base acute to obtuse, margin entire, when dry ± strongly involute; both surfaces glabrous; lateral veins (5-)9- 12(14) pairs, tertiary venation for the greater part parallel to the lateral veins, therefore, primary lateral veins often difficult to distinguish form the well-devel­ oped secondary lateral veins; petiole 0.5- 2(- 3) cm long, 1-2 mm thick, glabrous; stipules 0.2--0.8 cm long, ± densely, brown to yellowish or whitish villose to velutinous, ± glabrescent, or (sub)glabrous, often subpersistent. Figs in pairs in the axils of the leaves, or on already leaftess parts of the twigs, sessile or some­ times up to 0.3(--0.5) cm long pedunculate; basal bracts ca. 3 mm long, the upper free and membranous parts often caducous; receptacle globose to obovoid, when fresh ca. 0.5-1, when dry 0.5--0.8 cm in diameter, glabrous, at maturity pinkish, reddish or reddish-brown, when dry more or less wrinkled; apex when dry slightly depressed to slightly prominent.

Distribution. From Sierra Leone to Ivory Coast and from Nigeria to Zaire and Burundi (and possibly extending to Ethiopia), also in Sào Tomé. In hurnid and sernideciduous forest; at low altitudes. This species is very closely related to F thonningii, and possibly just one of the 'forms' of the F thonningii-complex. It can Gust) be distinguished by the way the leaves are drying and by the usually subpersistent stipules. F kamerunensis is a species of humid forest. The 'form' of F thonningii commonly found in such forest in West and Central Africa has larger and broader leaves. However, in Central Africa F kamerunensis occurs also sympatrically with the narrow-leaved 'persicifolia' form of F thonningii. The pollinator is Alfonsiella fimbriata Waterston (see also F natalensis leprieuriz).

Ficus subgenus Urostigma section Galoglychia subsection Crassicostae (Mildbr. & Burret) c.c. Berg

Berg, Proc. Kon. Ned. Akad. Wet. (C) 89: 124 (1986b).

Trees hemi-epiphytic, evergreen, bark not exfoliating. Leafy twigs, stipules, leaves and figs glabrous or inconspicuously hairy. Leaves often rather small, oblong to elliptic or to lanceolate, coriaceous, mostly shortly petiolate, tertiary venation reticulate to pronouncedly parallel to the lateral veins; stipules small, caducous. Figs in pairs in the lead axils or several together on small spurs in the leafaxils and bel ow the leaves, small or medium-sized, pedunculate (or sessile), at maturity red (or greenish?). Fruitlets with a dry pericarp, endocarp body not released.

132 The subsection comprises eight species and is morphologically somewhat heterogeneous. F. adolfi-friderici, F. pseudomangifera, F. usambarensis, F. louisii, and (probably) F. leonensis constitute the kemel of this subsection. These species show c1ear affinities, in particular in the occurrence of the small fig-bearing short-shoots (spurs) in the leafaxils and also below the leaves, bearing up to 7 figs at the same time. The relationships of these four species to the other four species ranked under the subsection are not quite c1ear and certain. Considering the type of venation and the position of the figs (in pairs in the 1eafaxils) F. elasticoides and F. burretiana might belong to subsection Chlamydodorae, whereas F. oreodryadum might belong to subsection Cyathistipulae. Some characters, like those of fruitlets, to be studied on fresh material, were not avail­ able, as to propose relationships with some more accuracy. F. barteri which has been placed in this group (Berg et al., 1985) proved to a member of subsection Cyathistipulae, largely because of the characters of the fruitlets. F. jansii, (provisionally) placed in subsection Plathyphyl/ae, might prove to belong to subsection Crassicostae. Seven species of this subsection occur in Central and West Africa, only F. usambarensis is East-African. Some of the species appear to be rare. All members of this subsection are components of rain forest or montane forest. In distribution and ecology the subsection shows similarities with subsection Cya­ thistipulae. The pollinators belong to the genera Nigeriel/a Wiebes, Paragaon Joseph and Elisabethiel/a Grandi.

68. Ficus elasticoides De Wild. (fig. 30)

De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 302 (I9I3b); Berg et al, FI. Cameroun 28: 202, fig. 71 (1985).

Tree up to 40 m taU, hemi-epiphytic or (secondarily) terrestrial. Leafy twigs 2- 5 mm thick, minutely puberulous. Lamina oblong to eUiptic to subobovate, 6.5-15 x 2.5- 8.5 cm, coriaceous, apex shortly and ± sharply acuminate (to subacute), base acute to subobtuse, margin entire; both surfaces glabrous; lateral veins (12- ) 15- 20 pairs, 100p-cOnnected 2-3 mm from the margin, tertiary vena ti on paral1el to the lateral veins, therefore, primary and secondary lateral veins often hardly distinguishable; petiole 1.5-6 cm long, ca. 2- 3 mm thick, glabrous, when dry red-brown to purplish or blackish; stipules 1-1.5 cm long, glabrous, caducous. Figs usuaUy in pairs in the 1eafaxils; pedunc1e 0.5- 1 cm long; basal bracts 3- 7 mm long, basal1y connate, persistent or the uppermost membranous parts caducous; receptac1e globose, when dry ca. 1.5 cm in diameter, almost glabrous.

Distribution. From Angola to Cameroun, also in Ivory Coast. In rain forest; at low altitudes.

133 Fig. 30. Ficus elasticoides De Wild. Leafy twig with figs. Mter Berg et al. (1985, pI. 71).

The pollinator is Elisabethiella articulata (Joseph).

69. Ficus burretiana Huteh. Hutchinson, Kew Bull. (1915): 344 (1915); Berg et al., Ft. Cameroun 28: 204, fig. 72 (1985).

134 Tree or shrub, hemi-epiphytic. Leafy twigs ca. 1.5- 2 mm thick, glabrous. Lamina oblong to elliptic to subobovate, 5--8 x 2-4 cm, coriaceous, apex shortly and faintly acuminate, base acute to (sub)obtuse, margin entire; both surfaces glabrous; lateral veins ca. 10--15 pairs, loop-connected 1-2 mm from the margin, teriary venation predominantly parallel to the lateral veins; petiole 2.5--3.5 cm long, 1- 1.5 mm thick, glabrous; stipules 0.5--1 cm long, glabrous, caducous. Figs in pairs in the leafaxils; peduncle 0.7-1.5 cm long; basal bracts with the upper parts caducous, basal connate parts leaving a narrow collar; receptacle (sub)globose, when dry 0.6-0.8 cm in diameter, glabrous.

Distribution. In Cameroun, Gabon, Zaire, and the Central African Republic. In rain forest; at low altitudes. The pollinator is Elisabethiella longiscapa Wiebes.

70. Ficus oreodryadum Mildbr. Mildbraed in Mildbraed & Burret, Bot. Jahrb. Syst., Pftanzengesch. & Pflanzengeogr. (EngIer) 46: 240 (1911 b); Berg et al. , FI. Cameroun 28: 200, fig. 70 (1985); Berg et al, FI. Trop. East Africa, Moraceae: 82 (J 989).

Tree, often large, hemi-epiphytic. Leafy twigs 2.5-6 mm thick, almost glabrous. Lamina oblong to elliptic or to subovate, 9-18 x 3.5-6 cm, coriaceous, apex acuminate, base (sub)acute to obtuse, sometimes to subcordate, margin entire; both surfaces glabrous; lateral veins 8- 14 pairs, tertiary venation reticulate; petiole 2-5 cm long, 1.5- 2 mm thick, glabrous; stipules ca. 1.5 cm long, yellowish appressed-puberulous or glabrous, caducous. Figs in pairs in the leafaxils, sometÏrnes also just bel ow the leaves, sessile; basal bracts ca. 4 mm long, per­ sistent; receptacle subglobose to ellipsoid to ovoid, when fresh ca. 2 cm, when dry ca. 1.5 cm in diameter, almost glabrous, warty, at maturity orange to yellow.

Distribution. Uganda, Rwanda and Burundi to Cameroun, SE. Nigeria, and Bioko (= Fernando Po). In wet montane and submontane forest at altitudes between 1300 and 2500 m.

71. Ficus pseudomangifera Huteh. (fig. 31) Hutchinson, Kew Bull. (1915): 342 (1915); Berg et al. , FI. Cameroun 28: 192(bis), fig. 66 (1985); Berg et al. , FI. Trop. East Africa, Moraceae: 82 (1989).

F mangiferoides Huteh. (1915).

Tree up to 10 m tall, hemi-epiphytic. Leafy twigs 3-6 mm thick, glabrous or minutely brownish puberulous. Lamina oblong to lanceolate, sometimes to ellip­ tic, 8- 32 x 2-9 cm, coriaceous, apex (sharply) acuminate, base rounded to (sub)cordate or to acute, margin entire; both surfaces glabrous; lateral veins 14-27 pairs, tertiary venation parallel to the lateral veins to reticulate; petiole

135 -1) . .HOLEè Fig. 31. Ficus pseudomangifera Hutch. I, leafy twig with figs; 2, branch with figs. After Berg et al. (\985, pI. 66).

1-3 cm long, ca. 3 mm thick, glabrous; stipules 0.2-0.5 cm long, (rather) densely greyish to yellowish to brownish strigose to subsericeous. Figs up to 6 together on small spurs in the leafaxils or also just below the leaves; peduncle 0.3-0.8 cm long; bas al bracts 2-3 mm long, persistent; receptacle (sub)globose, when

136 fresh 0.6-1.2 cm, when dry 0.3--0.7 cm in diameter, minutely puberulous, at maturity orange to red.

Distribution. Sierra Leone, Ivory Coast, and from Cameroun to Uganda. In rain forest; at altitudes up to 1200 m. The pollinator is Nigeriella letouzeyi Wiebes.

72. Ficus usambarelL'iis Warb.

Warburg, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 20: 159 (1894); Berg et al., FI. Trop. East Africa, Moraceae: 82 (1989).

Tree up to 15 m tall, (secondarily?) terrestrial. Leafy twigs 5- 7 mm thick, pu­ berulous to almost glabrous. Lamina oblong to lanceolate, 9.5-14.5 x 4.5--6 cm, coriaceous, apex (sub)obtuse to rounded, base obtuse to rounded, margin entire; both surfaces glabrous; lateral veins 10--13 pairs, tertiary venation largely paral­ lel to the lateral veins; petiole 1.5-3.5 cm long, 2-2.5 mm thick, glabrous; stipules 0.3--0.5 cm long, white to yellowish puberulous, caducous. Figs 2- 7 together on small spurs in the leafaxils or also just bel ow the leaves; peduncle 0.8-1.7 cm long; basal bracts ca. 3 mm long, persistent; receptacle globose, when dry 0.8-1 cm in diameter, sparsely puberulous.

Distribution. Tanzania. In woodland; at altitudes up to 1000 m.

73. Ficus adolfi-friderici Mildbr.

Mildbraed in Mildbraed & Burret, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 46: 232 (191Ib); Berg et al., FI. Cameroun 28: 194, t. 67 (1985).

F aganophila Huteh. (1915).

Tree up to 15 m tall. Leafy twigs 2.5-6 cm thick, glabrous or minutely pu­ berulous. Lamina oblong to elliptic to subobovate, (4--)8-20 x (2-)4--9.5 cm, coriaceous, apex shortly and bluntly acuminate, base acute to rounded (to sub­ cordate), margin entire; both surfaces glabrous; lateral veins 8-12(-15) pairs, loop-connected 1-3 mm from the margin, tertiary venation predominantly parallel to the lateral veins; petiole 0.5-3.5 cm long, 1.5- 2.5 mm thick, glabrous; stipules 0.5-1.5 cm long, glabrous or minutely puberulous, caducous. Figs in pairs or up to 6 together on small spurs in the leafaxils or also below the leaves, sessile; basal bracts ca. 2.5 mm long, basally connate, often ± split, persistent; receptacle subglobose, when fresh ca. 1 cm, when dry ca. 0.5 cm in diameter, minutely puberulous.

Distribution. Zaire, Equatorial Guinea, Cameroun, and Nigeria. In forest; at low altitudes.

137 74. Ficus louisii Boutique & J. Léonard

Boutique & 1. Léonard, Bull. 1ard. Bot. Bmx. 19: 215 (1949); Berg et al., FI. Cameroun 28: 196, fig. 68 (1985).

Tree up to 40 m tal!, sometimes lianescent. Leafy twigs 2-3.5 mm thick, glabrous. Lamina oblong to el!iptic, 4-12 x 1--6(-8) cm, coriaceous, apex sharply to bluntly acuminate, base acute to obtuse or to rounded, margin entire; both surfaces glabrous; lateral veins 8- 12 pairs, loop-connected 1- 2 mm from the margin, tertiary vena ti on largeiy parallel to the lateral veins; petiole 0.8- 2(-5.5) cm long, ca. 1.5 mm thick, glabrous; stipules 0.5- 1 cm long, glabrous or sparsely white puberulous, caducous. Figs 1-4 on small spurs in the leafaxils or also (just) below the leaves, sessile or up to 0.5 cm long pedunculate; basal bracts 1-1.5 mm long, persistent; receptacle (sub)globose, when fresh ca. 0.6-0.8 cm, when dry 0.3-0.5 cm in diameter, glabrous, smooth or slightly warty.

Distribution. Zaire, Gabon, Cameroun, and Guinea. In forest; at low altitudes. The pollinator is Paragaon josephi Wiebes.

75. Ficus leonensis Huteh.

Hutchinson, Kew Bull. (1915): 338 (1915); Berg et al. , FI. Cameroun 28: 198, fig . 69 (1985).

Tree or climber. Leafy twigs 2- 7 mm thick, minutely puberulous, ± con­ spicuously lenticellate, when dry hollow. Lamina elliptic to oblong, 3- 16 x 2-9.5 cm, coriaceous, apex ± sharply acurninate, base cordate to subcordate or (to truncate), margin entire; both surfaces glabrous; lateral veins 8-10 pairs, loop­ connected 1- 3 mm from the margin; tertiary venation reticulate or partly paral­ lel to the lateral veins; petiole (0.5-)1-5.5 cm long, 1- 2 mm thick, glabrous, epidermis ftaking off; stipules 0.5-1.5 cm long, almost glabrous, caducous. Figs solitary, in pairs or sometimes up to 4 together (on minute spurs?) in the leaf axils or sometimes also below the leaves, sessile; basal bracts ca. 1.5 mm long, basally slightly connate, often split, persistent; receptacle ± depressed-globose, when dry ca. 0.5-0.6 cm in diameter, sparsely minutely puberulous.

Distribution. Guinea, Sierra Leone, Ivory Coast, and Cameroun. In forest. The species is only known by a few collections.

Ficus subgenus Urostigma section Galoglychia subsection Cyathistipulae (Mildbr. & Burret) c.c. Berg

Berg, Proc. Kon. Ned. Akad. Wet. (C) 89: 125 (1986b).

Trees or shrubs, hemi-epiphytic (or secondarily terrestrial), evergreen. Leafy twigs, stipules, leaves mostly glabrous or inconspicuously hairy. Leaves large to small, elliptic to oblong, often pandurate, coriaceous, tertiary venation reticulate;

138 stipules often large and/or subpersistent, and in some species usually connate. Figs in the leafaxils, large to medium-sized (or smalI), often sessile, at maturity wall rather soft, sometimes spongy, green, yellowish or purplish. Fruitlets with the upper part ofthe pericarp becoming mucilaginous, this part can swell considerably, endocarp body not released.

This subsection comprises 19 species, most of them components of evergreen forest in Central and West Africa. Only F scassellatii is confined to eastern Africa. The section is cIear-cut and characterized by (thickly) coriaceous, glabrous, shortly petiolate leaves, relatively large figs, and relatively long, often (sub)persistent stipules. The pollinators belong to the genus Agaon Dalman.

76. Ficus conraui Warb. Warburg, Ann. Mus. Congo (6) I: 25, fig. 11 (1904); Berg et al., FI. Cameroun 28: 234, fig . 84 (1985); Berg et al. , FI. Trop. East Africa, Moraceae: 83 (1989).

F praticola Mildbr. & Huteh. in Huteh. (1915), F sessilis De Wild. (I 9 l3a), F stipulifera Huteh. (1915).

Treelet or shrub, hemi-epiphytic, sometimes lianescent. Leafy twigs 2-4 mm thick, glabrous or puberulous. Larnina oblong to lanceolate to subobovate or to elliptic, (6-)10-20(-28) x (2- )3.5-7 cm, coriaceous, apex acuminate, base acute to rounded, margin entire; both surfaces glabrous; lateral veins 8-11 pairs, the lower ones relatively smalI, tertiary venation reticulate; petiole 0.8-4.5 cm long, 1- 2 mm thick, epidermis flaking off or not so; stipules (0.5- )1-2.5 cm long, basally connate, glabrous, (sub)persistent. Figs in pairs or solitary in the leaf axils, sessile; basal bracts 2-3 mm long, persistent; receptacIe subglobose to obovoid (or subpyriform), when fresh 2- 3 cm, wh en dry (0.5- )1.5- 2.5 cm in diameter, glabrous or subhispid, warty or smooth, at maturity greenish with reddish spots or warts or partly reddish; wall when dry 1.... 2 mm thick.

Distribution. From NW. Angola and SE. Zaire (Shaba) to NW. Tanzania, Uganda and Nigeria, and from Ivory Coast to Sierra Leone. In ra in forest and semi-deciduous forest. Sterile specimens of F conraui can be distinguished from F cyathistipula by the greater number of lateral veins (8-11 pairs instead of 5- 7(- 8) pairs). The figs are sessile, in contrast to those of F cyathistipula subsp. cyathistipula. The pollinator is Agaon kiellandi Wiebes (see also F cyathistipula prings­ heimiana and F densistipulata).

77. Ficus tesselata Warb. Warburg, Bot. Jahrb. Syst., Pfianzengesch. & Pfianzengeogr. (Engier) 20: 156 (1894).

F camptoneuroides Huteh. (1915), F winkleri Mildbr. & Burret (1911).

139 Tree up to 18 m tall or shrub, hemi-epiphytic. Leafy twigs 3-6 mm thick, glabrous, drying brown, on older twigs usually with rather large lenticels, con­ centrated in the uppermost part of the internodes. Lamina oblong to subobovate (or subovate), 9-25(- 30) x 4-9.5(-13) cm, (sub)coriaceous, apex acuminate, base acute to obtuse (or nearly rounded), margin entire; both surfaces glabrous; la­ teral veins 4-7(-8) pairs, tertiary venation reticulate; petiole 1-3(-6) cm long, 2-2.5(-4) mm thick, (in ol der leaves) epidermis flaking ofT; stipules (1-)1.5- 2(- 3) cm, on flush up to 6 cm long, mostly basally connate, glabrous or sparsely, minutely puberulous, persistent. Figs solitary or in pairs in the leafaxils, sessile; basal bracts ca. 2 mm long, persistent; receptacle globose (to obovoid), when fresh 2-2.5 cm, when dry (1.2-)1.5- 2 cm in diameter, usually yellowish hirtellous to hispid and surface ± scabrous and warty, at maturity (pale) green with yellow spots; wall when dry 2- 2.5 mm thick not (or hardly) spongy; apex of the re­ ceptacle ± protracted.

Distribution. From Sierra Leone to E. and S. Zaire and Rwanda. Forest, often in riverside forest; at altitudes up to 1900 m. The pollinator is Agaon laïense Wie bes.

78. Ficus pachyneura C. C. Berg Berg, Kew Bull. 43: 92, fig. 5 (1988).

Tree, hemi-epiphytic. Leafy twigs (2- )5- 9 mm thick, glabrous, with conspicuous lenticels on the uppermost part of the internodes. Lamina subobovate to oblanceolate, 9.5- 27 x 3.5-8 cm, coriaceous, apex acuminate, base acute, margin entire; both surfaces glabrous; lateral veins 6-8 pairs, the lower pairs weak, the middle pairs strong and furcate far from the margin, tertiary venation reticulate; petiole 1-2.5 cm long, 2-3.5 mm thick, glabrous, epidermis ± flaking ofT; stipules 1-2 cm long, (sub)glabrous, usually persistent. Figs solitary or up to 4 together in the leafaxils, sessile; basal bracts 2-2.5 mm long, persistent; receptacle globose, when dry ca. 0.8 cm in diameter, glabrous or white appressed-puberulous; wall wh en dry 0.5-1 mm thick; apex of receptacle ± pro­ minent to plane.

Distribution. Liberia, Ivory Coast and Ghana. In rain forest; at low altitudes. Only known by a few collections. This recalls F. camploneura by its small figs and small number of lateral veins, but difTers from the latter in the leaf shape, the thickness of the petiole and the leafy twigs, the length of the uppermost parts of the intern odes. In most of these difTerentiating characters F. pachyneura matches F. lesse/ala Warb. which can be distinguished by the larger figs (when dry (1.2-)1.5-2 cm in diameter) with yellow hairs, the leaf base, which is often obtuse to rounded or even subcordate, and also in the larger number of interfloral bracts and larger number of tepals

140 (five). This species was treated as F afT. camptoneuroides by Aubréville (1959: 82, fig. 15C).

79. Ficus ardisioides Warb. Warburg, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 20: 171 (1894); Berg et al ., FI. Cameroun 28: 238, fig . 85 (1985) sub F. camptoneura; Berg, Kew Bull. 43: 77 (1988).

Tree up to 8 m tall or shrub, (sometimes lianescent?), hemi-epiphytic. Leafy twigs 2-4 mm thick, minutely puberulous to glabrous, (when dry) often hollow. Lamina oblong to elliptic or to lanceolate, or subobovate, (1.5- )6-16(- 26) x (1 - )1.4--6.5(- 9.5) cm, (sub)coriaceous, apex acuminate to subcaudate, base acute to rounded, margin entire; both surfaces glabrous; lateral veins (3- )4-7 pairs, loop-connected up to 7 mm from the margin, basal pair weak, the first strong lateral veins arising 0.5- 2 cm above the base of the lamina, tertiary venation reticulate; petiole 0.5-1.5(-2) cm long, 1- 2 mm thick, glabrous, epidermis usually flaking ofT; stipules 0.2- 1 cm long, glabrous, or in the upper part cilio­ late, caducous or subpersistent. Figs in pairs or solitary in the leafaxils, sessile or up to 0.9 cm long pedunculate; basal bracts 1.5-2 mm long, persistent; re­ ceptacle when fresh ca. I cm, when dry 0.8-1 cm in diameter, puberulous, smooth or ± warty; wall when dry ca. 0.5 mm thick; apex of receptacle plane.

Two (entirely?) allopatric subspecies can be recognized.

79a. Ficus ardisioides subsp. ardisioides Figs on peduncles up 0.2--0.9 cm long.

Distribution. N. Zaire and the Central African Republic. In forest; at low al­ titudes.

79b. Ficus ardisioides subsp. camptoneura (Mildbr.) c.c. Berg Berg, Kew Bull. 43: 77 (1988).

F. arcuato-nervata Huteh. (1915), F. camptoneura Mildbr. in Mildbr. & Burret (1911), F. camp­ toneura var. angustifolia Mildbr. in Mildbr. & Burret (1911).

Figs sessile.

Distribution. From N.W. Zambia to E. Zaire and SE. Nigeria, possibly also in Ivory Coast. In forest; at low altitudes. Subspecies camptoneura has sessile figs as F conraui and F subcostata, but it can be distinguished by the smaller size of the figs (usually 1-1.5 cm as against usually 1.5-2.5 cm in diameter, wh en dry, in the other species) and the relatively short (usually up to ca. I cm long), caducous stipules; the stipules are up to 2.5 cm long and (sub)persistent in the other two species.

141 The pollinator is Agaon megalopon Wiebes.

80. Ficus preussii Warb.

Warburg, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 20: 156 (1894); Berg et al. , Ft. Cameroun 28: 242, fig. 87 (1985); Berg et al. , Ft. Trop. East Africa, Moraceae: 85 (1989).

F. lujae De Wild. (I 9 l3a).

Tree up to 10 m taU or shrub, hemi-epiphytic. Leafy twigs 8-10 mm thick, (densely) white strigose to puberulous, periderm flaking off. Lamina oblong to (sub)ovate, sometimes subpandurate, 19-26(-45) x 8- 13(- 17) cm, coriaceous, apex acuminate, base obtuse to subcordate, sometimes to cordate, margin entire; upper surface glabrous, lower surface glabrous or white to brownish hirtellous on the lower part of the midrib; lateral veins 6-8(-9) pairs, tertiary venation (sub)reticulate; petiole 2.5- 8.5 cm long, ca. 4 mm thick, epidermis flaking off; stipules ca. 2 cm long, free or basally connate, glabrous or sometimes white strigillose, persistent. Figs solitary or in pairs in the leafaxils, sessile or subsessile; basal bracts ca. 5 mm long, persistent; receptacIe globose, when fresh 2.5- 5 cm, in when dry 2-3 cm in diameter, rather densely yellowish to white strigillose to hirtellous, ± pronouncedly warty, at maturity greenish with yeUowish spots; wall when dry ca. 3-5 mm thick; apex up to 5 mm protruding.

Distribution. Uganda to Nigeria. In rain forest; at altitudes up to 1200 m.

81. Ficus abscondita c.c. Berg Berg, Adansonia (2) 20: 264, fig. I (1980); Berg et. al., Ft. Cameroun 28: 244, fig. 88 (1985).

Shrub up to 3 m tall, hemi-epiphytic (or hemi-epilithic), sometimes lianescent. Leafy twigs 5- 10 mm thick, glabrous or sparsely puberulous. Lamina obovate to oblanceolate, 28-40 x 8-15 cm, (sub)coriaceous to coriaceous, apex abruptly and more or less sharply acuminate, base acute to subobtuse to truncate or to cordate; margin entire; both surfaces glabrous; lateral pairs 6-8 pairs, loop­ connected 3- 8 mm from the margin, tertiary vena ti on reticulate; petiole 2-8 cm long, 3-4 mm thick, glabrous or puberulous, epidermis flaking off; stipules 2.5- 6.5 cm long, glabrous or sparsely puberulous, persistent. Figs in the leafaxils or just below the leaves, sessile, often (entirely) concealed by the persistent stipules; basal bracts ca. I mm long, persistent; receptacIe oblongoid to sub­ globose or ovoid, when fresh up to 2.5 cm, when dry 0.7-1.5 cm in diameter, when fresh up to 3 cm, when dry 1- 1.8 cm long, sparsely puberulous to hirtel­ lous; wall when dry ca. 0.5 mm thick; ostiole in a crateriform, or sometimes strongly swoUen, apex of the receptacIe.

Distribution. Cameroun and Gabon. In forest; at low altitudes.

142 82. Ficus cyathistipula Warb.

Warburg, Bot. Jahrb. Syst. , Pflanzengesch. & Pflanzengeogr. (Engier) 20: 173 (1894); Berg et al. , F!. Cameroun 28: 246, fig . 89 (1985) and 240, fig . 86 (1985) sub F. pringsheimiana; Berg, Kew Bull. 43: 82 (1988); Berg et al. , F!. Trop. East Africa, Moraceae: 83 (1989).

Tree up to 8 m tall, terrestrial or hemi-epiphytic. Leafy twigs 3- 5 mm thick, glabrous or white puberulous, periderm sometimes flaking ofT. Lamina oblanceolate to obovate, 6-20 x 3-7 cm, coriaceous apex acuminate, base acute to attenuate, margin entire; both surfaces glabrous or sometimes sparsely pu­ berulous at the base of the midrib beneath; lateral veins 5-7(-8) pairs, tertiary venation reticulate; petiole 1.5-4 cm long, 2-3 mm thick, glabrous or sparsely (brownish) puberulous; stipules 1.5- 2.5 cm (or 0.5-0.7 cm) long, partly connate, minutely white puberulous or almost glabrous, persistent. Figs solitary (or up to 3 together) in the leafaxils; sessile or 0.5- 2.5 cm long pedunculate; basal bracts ca. 4 mm long, persistent; receptacle globose to obovoid (and usually up to ca. 10 mm long stipitate) or pyriform, when fresh 3- 5 cm, when dry 2- 3 cm in diameter, often somewhat scabrous, at maturity pale green to pale yellow; wall when dry (2-)4-8 mm truck and spongy, smooth, warty or sometimes with up to 3 mm long protuberances; apex of receptacle (at least when dry) up to 10 rnm protruding.

The charactistic truck and spongy wall probably promotes dispersalof the figs by water.

Two subspecies can be recognized.

82a. Ficus cyathistipula subsp. cyathistipula

F. cal/escens Hiem (1900), F. nyanzensis Huteh. (1915), F rederi Huteh. (1915), F. rhynchocarpa Mildbr. & Burret (1911).

Figs pedunculate.

Distribution. From Ivory Coast to Angola, N. Zambia, Kenya, and Tanzania. In forest, often along water courses, sometimes on rocks; at altitudes up to 1800 m. The material from West Africa (Ivory Coast) with reiativeiy slender twigs, small leaves and short (0.5-0.7 cm long) stipules might represent another sub­ species. The pollinator is Agaon fasciatum Waters ton.

82b. Ficus cyathistipula subsp. pringsheimiana (Braun & K. Schum.) e.C. Berg Berg, Kew Bull. 43: 83 (1988).

F. pringsheimiana Braun & K. Schum. (1 889).

143 Figs sessile.

Distribution. Cameroun and Gabon. In forest; at low altitudes. The pollinator is Agaon kieflandi Wiebes (see also F. conraui and F. densisti­ pulata).

83. Ficus scasseUatii Pamp. Pampanini, Boll. Soc. Bot. Hal. (1915:1-2): 15 (1915); Friis, Nord. J. Bot. 5: 331 (1985); Berg, Kew Bull. 43: 94 (1988); Berg et al. F1. Trop. East Africa, Moraceae: 84 (1989).

Tree up to 50 m tall, hemi-epiphytic, secondarily terrestrial. Leafy twigs 3-8 mm thick, glabrous or minutely puberulous, periderm of older parts flaking ofT. Lamina oblong to (ob)lanceolate to (sub)obovate or to elliptic, (6-)10-20(- 28) x 3-8 cm, coriaceous, apex shortly and mostly bluntly acuminate to rounded, base (sub)acute, margin entire; both surfaces glabrous; lateral veins 8- 18 pairs, gradually becoming stronger towards the apex, tertiary venation reticulate; petiole 0.5- 2.5(- 3.5) cm long, 2-3 mm thick, glabrous; stipules 0.3-2 cm long, glabrous or minutely puberulous, caducous. Figs solitary or in pairs in the leaf axils; peduncle 0.5-1.5 cm long or (sub)sessile; basal bracts 3-5 mm long, per­ sistent; receptacle globose to ellipsoid, when fresh 4-4.5 cm, (at least when dry often shortly stipitate) when dry 1.2- 2(- 3) cm in diameter, (sparsely) minutely puberulous, at maturity green; wall when dry (1 - )4-5 mm thick, not spongy; apex of receptacle (at least when dry) up to 7 mm protruding.

F. scasselatii apears to be closely related to F. cyathistipula. It can be distin­ guished from the latter by the caducous stipules. Moreover, the wall of the figs is not spongy, as it is in F. cyathistipula.

Two subspecies can be recognized.

83a. Ficus scasseUatii subsp. scasseUatii

F. kirkii Hutch. (1915), F. michelsonii Boutique & J. Léonard (1949).

Lamina oblong to subobovate to elliptic, sometimes lanceolate to oblanceolate, up to 20 x 8 cm. Figs on peduncles 0.5-1.5 cm long; receptacle when dry 1.2-2 cm in diameter and the apex often ± protruding.

Distribution. In E. Zimbabwe, Malawi, Mozambique, Tanzania, Kenya, and S. Somalia, also (disjunctly?) in E. Zaire. In evergreen forest; at altitudes up to 1800 m.

83b. Ficus scasseUatii sub sp. thikaensis c.c. Berg Berg, Kew Bull. 43: 95 (1988).

144 Lamina mostly oblanceolate, up to 28 x 4.5 cm. Figs sessile or subsessile; re­ ceptacle when dry 2-3 cm in diameter and the apex hardly protruding.

Distribution. Kenya, only known from the surroundings of Thika. Subspecies thikaensis difTers from subsp. scassellatii in the sessile figs.

84. Ficus cyathistipuloides De Wild. (fig. 32) De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 194 (1913a); Berg et al., Fl. Cameroun 28: 248, fig. 90 (1985).

Tree, often large, hemi-epiphytic. Leafy twigs 1.5-3.5 mm thick, glabrous or (sparsely) puberulous to hirtellous, on the scars always densely white puberulous to shortly pubescent, lenticels usually conspicuous. Lamina oblong to elliptic or to lanceolate, 4.5-15.5 x 2-6 cm, coriaceous, apex shortly and bluntly acuminate (to subacute), base acute to obtuse or sometimes to rounded, margin entire; both surfaces glabrous; lateral veins 6-10 pairs, tertiary venation reticulate; petiole 0.5- 2 cm long, 1-2 mm thick, glabrous, epidermis usually flaking ofT in ring­ shaped flakes; stipules 0.5-2 cm long, puberulous to shortly pubescent, caducous or subpersistent. Figs solitary in the leafaxils; peduncle 0.3-0.9 cm long; basal bracts 3-5 mm long, persistent; receptacle subglobose to ellipsoid or to ovoid, when fresh ca. 2.5-4 cm, when dry 2-2.5 cm in diameter, 2-8 mm long stipitate, glabrous or the lower part sparsely puberulous; apex of receptacle very pro­ minent.

Distribution. Zaire, Gabon, and Ivory Coast. In rain forest; at low altitudes. The pollinator is Agaon obtusum Wiebes. A parasitoid is Sycoecus thauma­ stocnemus Waterston.

85. Ficus scott-eUiotii Mildbr. & Burret Mildbraed & Burret, Bot. Jahrb. Syst., Ptlanzengesch. & Ptlanzengeogr. (EngIer) 46: 234 (191Ib).

Tree up to 15 m tall, usually hemi-epiphytic. Leafy twigs (2-)3-6 mm thick, glabrous, when dry dark brown to blackish with conspicuous white lenticels, often arranged in ± distinct circular bands. Lamina oblong to subobovate, (6-)8 - 15 x (3-)3.5-7.5 cm, coriaceous, apex shortly and bluntly acuminate, base subacute to ob tu se (to rounded); both surfaces glabrous; lateral veins 7-9 pairs, tertiary venation reticulate; petiole 1.2-3 cm long, 1.5-2 mm thick, epidermis flaking ofT; stipules 0.6-1 cm long, minutely puberulous or only ciliolate, caducous. Figs solitary in the leafaxils; peduncle 0.3-0.5 cm long, ca. 2 mm thick; basal bracts ca. 3 mm long, persistent; receptacle globose, when fresh ca. 2-3 cm, when dry ca. 2-3 cm in diameter, sparsely minutely puberulous, at maturity greenish (?); wall when dry ca. 2 mm thick, ± wrinkled; apex of re­ ceptacle shortly apiculate to plane.

145 .D.H.OL E'r Fig. 32. Ficus cyathistipuloides De Wild. Leafy twig with figs. Mter Berg et al. (1985, pI. 90).

Distribution. Sénégal to Ivory Coast. In (rain) [orest; at low altitudes.

86. Ficus densistipulata De Wild.

De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 194 (1913a); Berg et al. , FI. Cameroun 28: 250, fig. 91 (1985); Berg et al., F1. Trop. East Mrica, Moraceae: 84 (1989).

F. coriacea De Wild. (1921), non Aiton (1789), F. ebolowensis Mi1dbr. & Huteh. in Huteh. (1915), F. namalalensis Huteh. (1915), F. slipitala Lebrun (1934).

146 Tree up to 10 m tall, hemi-epiphytic. Leafy twigs 2.5-5 mm thick, puberulous to hirtelJous. Lamina oblong to lanceolate to subobovate, 12-25 x 4-7(-11) cm, coriaceous, apex acurninate, base acute to rounded, margin entire; both surfaces glabrous; lateral veins 6-8 pairs, tertiary venation reticulate; petiole 1- 2.5(-5) cm long, 2-4 mm thick, epidermis flaking off; stipules 1.5-3 cm long, partly connate, sparsely puberulous, (sub)persistent. Figs solitary in pairs in the leaf axils, subsessile or up to 0.4 cm long pedunculate; basal bracts 2-3.5 mm long, persistent; receptacle subglobose, when fresh 1.3-2.2 cm, when dry 1- 1.5 cm in diameter, 4-9 mm long stipitate, sparsely puberulous to almost glabrous, smooth or warty, at maturity greenish with yellow spots or partly reddish; wall when dry 0.5-1 mm thick, nqt spongy; apex of receptacle almost plane.

Distribution. From N . Angola to Uganda and Cameroun. In rain forest; at altitudes up to 1250 m. This species is very similar to F. cyathistipula in its vegetative parts. Fertile material of F. densistipulata can be easily told apart by the smaller figs (1 - 1.5 instead of 2-3 cm in diameter when dry). They are always stipitate and the basal bracts are smaller (2- 3.5 as against ca. 4 mm long). The pollinator is Agaon kiellandi Wiebes (see also F. conraui and F. cyathi­ stipuia subsp. pringsheimiana).

87. Ficus subcostata De Wild. De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 197 (1913a); Berg et al., FI. Cameroun 28: 252, fig. 92 (1985).

Tree or shrub, hemi-epiphytic. Leafy twigs 4-7 mm thick, puberulous to almost hirtellous. Lamina ovate to subovate (or oblong to elliptic), 5-25 x 2- 9 cm, coriaceous, apex acurninate to subacute, base cordate to truncate (to obtuse), margin entire; both surfaces glabrous; lateral veins 6-9 pairs, loop-connected (1 - )3-4 mm from the margin; tertiary venation reticulate; petiole (0.5- )1-3(-3.5) cm long, 1- 2.5 mm thick, puberulous to almost hirtellous, epidermis flaking off or persistent; stipules 1-2.5 cm long, free or basally connate, almost glabrous, caducous or subpersistent. Figs in pairs or solitary in the leafaxils, sessile; basal bracts 2- 3 mm long, persistent; receptacle (sub)globose, when fresh ca. 2 cm when dry 1.3- 1.7 cm in diameter, ± scabrous with rather dense white rigid hairs, ± warty; walJ when dry ca. 1.5 mm thick; apex of receptacle slightly prominent.

Distribution. From N. Angola to Zaire and Gabon. In rain forest; at low al­ titudes. The pollinator is Agaon acutatum Wie bes.

88. Ficus barteri Sprague (fig. 33) Sprague, Gard. Chrono 33: 354 (1903); Berg et al., FI. Cameroun 28: 190, fig. 65 (1985); Berg et al, FI. Trap. East Africa, Moraceae: 83 (1989).

147 .D.X Ot..,. Fig. 33. Ficus barteri Sprague. I, leafy twig with figs; 2- 3, pistillate ftowers; 4, fruit; 5, fruit with mucilage 'cap'; 6, staminate ftower. After Berg et al. (1985, pI. 65).

F. laurentii Warb. (I904a), F. mildbraedii Hutch. (1915).

Tree up to 10 m taB or shrub, hemi-epiphytic. Leafy twigs 3-5 mm thick, glabrous. Lamina lanceolate to linear (or oblong to elliptic), (5.5-)10-18(- 30) x 1.5-3.5(- 7) cm, coriaceous, apex acuminate to subacute, base acute (to

148 rounded), margin entire; both surfaces glabrous; lateral veins 10-20 pairs, tertiary venation reticulate to parallel to the lateral veins; petiole 1-4.5 cm long, ca. 2 mm thick, glabrous; stipules (0.5-)1 - 2 cm long, glabrous, caducous. Figs in pairs in the leaf muis; peduncle (0.5-)1-2.5 cm long; basal bracts 1.5-2 mm long, caducous; receptacle globose, when fresh 1- 1.5 cm, when dry ca. 0.5-1 cm in diameter, glabrous, smooth to warty, at maturity yell ow to orange; apex of receptacle plane.

Distribution. From N. Zambia through Zaire to Uganda and westwards to Sierra Leone. In evergreen (ra in and gallery) forest; at altitudes up to 1200 m. This species can he distinguished from the other species of subsection Cyathi­ stipulae with pedunculate figs by its caducous stipules, small receptacle and greater number of lateral veins (up to 20 rather than at most 15 pairs as in the other species). The pollinator is Agaon paradoxum Dalman.

89. Ficus Iyrata Warb. Warburg, Bot. Jahrb. Syst. Pftanzengesch. & Pftanzengeogr. (EngIer) 20: 172 (1894); Berg et al., FI. Cameroun 28: 254, fig. 93 (1985).

Tree or shrubs, hemi-epiphytic or (secondarily) terrestrial. Leafy twigs 5-12 mm thick, rather densely minutely puberulous. Lamina broadly pandurate, 20-40 x 10-28 cm, coriaceous, apex shortly and bluntly acuminate, base more or Ie ss deeply cordate, margin entire, both surfaces glabrous; lateral veins 6-7 pairs, tertiary venation reticulate; petiole 1.5-5 cm long, 4-6 mm thick, glabrous; stipules 1-6 cm long, free, both surfaces minutely puberulous, persistent. Figs in pairs or solitary in the leafaxils, sessile or occasional1y up to 0.3 cm long pedunculate; basal bracts ca. 2.5 mm long, persistent; receptacle ± depressed globose, when fresh 4-6.5 cm, when dry ca. 3 cm in diameter and wrinkled, puberulous, at maturity greenish, often with paler spots; wall when dry wrinkled and apex of receptacle ± umbonate.

Distribution. From Sierra Leone to Cameroun. In rain forest; at low altitudes. The pollinator is Agaon spatulatum Wie bes.

90. Ficus sagittifolia Mildbr. & Burret Mildbraed & BUITel, Bot. Jahrb. Syst., Pftanzengesch. & Pftanzengeogr. (EngIer) 46: 241 (19 11 b); Berg et al., FI. Cameroun 28: 254, fig. 94 (1985).

Tree up to lOm tal1 or shrub, hemi-epiphytic. Leafy twigs 5-10 mm thick, densely to sparsely minutely puberulous. Lamina narrowly pandurate, 15-40 x 4-9.5 cm, coriaceous, apex more or less sharply acuminate to subacute, base ± deeply cordate, sometimes rounded; margin bel ow the constriction of the lamina; margin at the base usual1y ± distinctly coarsely crenate-dentate; both

149 surfaces glabrous; lateral veins 10-15 pairs, tertiary venation reticulate; petiole 1-4 cm long, 3-5 mm thick, glabrous; stipules 1-7 cm long, both surfaces ± densely minutely puberulous, persistent. Figs solitary or in pairs in the leafaxils, sessile; basal bracts ca. 5 mm long, persistent; receptacle subglobose to depressed globose, when fresh ca. 2.5 cm, when dry ca. 1.5 cm in diameter, at maturity greensish or partly purplish-red, with pale spots; wall when dry ± wrinkled; apex of receptacle plane.

Distribution. From Guinea to Cameroun. In rain forest; at low altitudes. The pollinator is Agaon c. cicatriferens Wiebes.

91. Ficus subsagittifolia c.c. Berg (fig. 34) Berg, Adansonia (2) 20: 268, fig. 3 (1980); Berg et al. , Fl. Cameroun 28: 256, fig. 95 (1985).

Shrub or treelet up to 5 m tall, hemi-epiphytic. Leafy twigs 7-15 mm thick, glabrous or rather sparsely hirtellous. Lamina subpandurate or lanceolate to oblong, 16-40 x 3-15 cm, coriaceous, apex more or less sharply acuminate, base ± deeply cordate to truncate; margin sometimes faintly repand below the con­ striction of the lamina; both surfaces glabrous or sparsely puberulous on the basal part of the midrib beneath; lateral veins 9- 15 pairs, tertiary venation reticulate; petiole 1- 5.5 cm long, J-Q mm thick, glabrous or (rather) sparsely hirtellous, epidermis flaking off; stipules 1-8 cm long, usually strigillose, per­ sistent. Figs solitary or in pairs in the leafaxils, sessile; basal bracts 3-5 mm long, persistent; receptacle often more or less depressed globose, when dry 2.5-3.5 cm in diameter, puberulous to hirtellous, at maturity green with white spots to dark maroon with pa Ie brown spots; when dry wall often wrinkled; apex of receptacle plane.

Distribution. In Cameroun and Gabon. In ra in forest; at low altitudes. The poUinator is Agaon cicatriferens multum Wiebes.

92. Ficus wildemaniana De Wild. & T. Durand

De Wildeman & T. Durand, Ann. Mus. Congo (3) 2: 217 (1901); Warburg, Ann. Mus. Congo (6) 1: 7 (1904); Berg et al. , Fl. Cameroun 28: 258, fig. 96 (1985); Berg et al. , Fl. Trop. East Africa, Moraceae: 85 (1989).

Tree up to 20 m taU or shrub, hemi-epiphytic. Leafy twigs 10--20 cm thick, glabrous or brownish puberulous. Lamina lanceolate to oblong to subobovate, often (sub)pandurate, (15- )25-60 x 6--25 cm, coriaceous apex acuminate to sub­ acute, base acute or subattenuate and cordulate, margin entire; both surfaces glabrous; lateral veins 8-14 pairs, tertiary venation partly scalariform; petiole 1.5-8 cm long, 4-10 mm thick, glabrous; stipules 0.3-1 cm long, glabrous, caducous. Figs solitary or in pairs in the leafaxils, sessile; basal bracts 4-10 mm long, persistent; receptacle (depressed-)globose, when fresh 2.5-4.5 cm, when

150 Fig. 34. Ficus subsagittifolia C.C. Berg. I, leafy twig with figs; 2, staminate flower and stamen; 3, pistillate flower. After Berg (1980, pI. 3). dry 2.5-4 cm in diameter, yell ow hirtellous to strigillose; wall when dry 1-1.5 mm thick, wrinkled; apex of receptac1e plane.

151 Distribution. From Uganda to Cameroun. In rain forest; at altitudes up to 1100 m. The poUinator is Agaon gabonense Wiebes.

93. Ficus oresbia C.e. Berg Berg, Adansonia (2) 20: 266 (1980); Berg et al., F1. Cameroun 28: 260, fig. 97 (1985).

Tree. Leafy twigs 6-10 mm thick, white puberulous with partly retrorse hairs. Lamina subobovate to oblanceolate, 15-25 x 5-9 cm, coriaceous, apex shortly acuminate, base subacute to obtuse, margin entire; both surfaces glabrous; lateral veins 7-8 pairs, departing the midrib in sharp angles, often furcate far from the margin, loop-connected 2- 3 mm from the margin, tertiary venation reticulate; petiole 1.5-5 cm long, 2-3 mm thick, glabrous, epidermis sometimes flaking off at the lower part; stipules 0.5- 2 cm long, sparsely to densely white to pale yeUow puberulous, partly with retrorse hairs, caducous. Figs in pairs or solitary in the leafaxils, sessile; basal bracts 8-10 mm long, persistent; receptacle (sub)globose, when fresh 2-2.5 cm, when dry ca. 2 cm in diameter, puberulous, at maturity purplish; wall when dry wrinkled; apex of receptacle (at least when dry) ± umbonate.

Distribution. Cameroun. In forest. Only known from the type coUection, col­ lected at 1900 m.

94. Ficus crassicosta Warb. Warburg, Ann. Mus. Congo. (6) 1: 11 (1904a).

F. epiphytica De Wild. (1913a),

Tree up. to 10 m taU, hemi-epiphytic. Leafy twigs 2-5 mm thick, glabrous or minutely puberulous. Lamina lanceolate to oblong, tending to pandurate, 10-17 x 2- 5 cm, coriaceous, apex (sub)acuminate, base cordate; both surfaces glabrous; lateral veins 10-15 pairs, tertiary venation reticulate; petiole 0.5-1.5 cm long, 2-2.5 mm thick, glabrous, epidermis flaking off; stipules 0.3-0.8 cm long, minutely puberulous (or glabrous), caducous. Figs solitary in the leafaxils; peduncle 0.3-0.8 cm long; basal bracts, 3-4 mm long, persistent; receptacle when fresh ca. 4 cm, when dry 2.5- 3 cm in diameter; waU when dry 2-3 mm thick, wrinkled; apex of receptacle (at least when dry) ± umbonate.

Distribution. Zaire. In forest; at low altitudes.

Ficus subgenus Urostigma section Galoglychia subsection Caulocarpae (Mildbr. & BUITet) e.C. Berg

Berg, Proc. Kon. Ned. Akad. Wet. (C) 89: 125 (1986b).

152 Trees, shrubs or lianes, mostly hemi-epiphytic, mostly evergreen, bark smooth, not flaking ofT. Leafy twigs, stipules, leaves and figs glabrous or minutely pu­ berulous. Leaves coriaceous (to chartaceous), oblong to elliptic or ovate to cor­ diform, shortly or longly petiolate, margin entire; stipules mostly smal! and caducous. Figs medium-sized to large, in most species on spurs on the older wood, in most species pedunculate, wal! at maturity often soft, mostly greenish. Fruitlets with the pericarp becoming partly mucilaginous, endocarp body released and surrounded by a mucilaginous layer which can swell considerably.

The subsection Caulocarpae comprises 11 species, a single species of which (F po/ita) extends to Madagascar. It does not show a distinct concentration of species in one or another part of Africa. Most species are associated with wet to rather dry evergreen forest. A few species (e.g. F sansibarica) extend to savanna woodland. The subsection is char­ acterized by the occurrence of (up to 10(-15) cm long) leafless (peg-shaped, curved or straight, to cushion-shaped) branchlets (spurs), in most species bear­ ing one to several (up to 10) medium-sized to large figs at the same time. F ovata is inserted in this subsection on the basis ofsimilarities in the fruitlets. It deviates from the other spcies of the subsection in several features: the figs are not borne on spurs on the older wood, but (often) subsessile in the axils of the leaves andthe leaves are often rather densely hairy beneath. The young figs are enelosed in conspicuous calyptrate bud covers. For the identification of the species of this subsection the presence of leafy twigs, figs and spurs is necessary. The pollinators belong to the genus Courtella KiefTer.

95. Ficus ottoniifolia (Miq.) Miq. Miquel, Ann. Mus. Bot. Lugd.-Bat. 3: 288 (1867); Berg et al., Fl. Cameroun 28: 217, fig. 77 (1985); Michaloud et al., Proc. Kon. Ned. Akad. Wet. (C) 88: 93 (1985); Berg, Kew Bull. 43: 88 (1988); Berg et al., Fl. Trop. East Africa, Moraceae: 76 (1989).

Tree up to 15 m tall or sometimes shrub or liana, terrestrial or hemi-epiphytic. Leafy twigs 2-5(-10) mm thick, minutely puberulous or glabrous. Lamina ellip­ tic to oblong to (sub)ovate or (sub)obovate, 6.5-15(- 22) x 3-7(-9) cm, (sub)­ coriaceous (or chartaceous), apex acuminate to subcaudate, base acute to rounded to subcordate, margin entire; both surfaces glabrous; lateral veins (3-) 4-15 pairs, tertiary venation reticulate; petiole 1.5-9 cm long (0.5-)1.5-2 mm thick, glabrous; stipules 0.2-0.8 cm, on flush up to 4 cm long, sparsely pu­ berulous or glabrous, caducous. Figs up to 4(-10) together on up to 1.5(-4) cm long spurs on the older wood; bud scales of the spurs (alrnost) glabrous; pedunele 0.8-2.5 cm long, 1- 1.5 mm thick; basal bracts 2-3 mrn long, free parts caducous or (sub)persistent; receptaele ellipsoid to (sub)globose, when fresh 1.5- 4 cm, when dry 0.7-2.5 cm in diameter, puberulous or almost glabrous, at ma-

153 turity greenish to pa Ie orange or brownish, with pale green to whitish spots; wall when dry ca. 7 mm thick, not or hardly wrinkled.

Five subspecies can be recognized. Subspecies ulugurensis occurs disjunctly in East Africa. The area of subsp. ottoniifolia overlaps those of the West African subsp. multinervia and the Central-African subsp. lucanda. The ranges of subsp. macrosyce and subsp. lucanda might overlap.

95a. Ficus ottoniifolia subsp. ottoniifolia

F. buettneri Warb. (1894), F. buellneri var. globicarpa Mildbr. & Burret (1911), F. dewevrei De Wild. & T. Durand (1901), F. dewevreoides De Wild. (1921), F. ea/aensis De Wild. (l9I3a), F. fasciculiflora Huteh. (1915), F. gilletii Warb. (1904), F. maculosa Huteh. (1915), F. polybraclea De Wild. & T. Durand (1901), F. pynaerlii De Wild. (1913a), F. umangiensis De Wild. (1913a), F. umagiensis var. laurentii De Wild. (19I3a), F. viridimaculala De Wild. (I913a), Urostigma ottonii­ folium Miq. (1848).

Tree, treelet or sometimes liana. Leafy twigs usually drying pale brown. Lamina coriaceous to subcoriaceous, 8- 22 x 3-9 cm, usually broadest in the middle, apex acuminate, base usually rounded to obtuse, som times subacute or emar­ ginate; lateral veins 6-10(- 11) pairs; petiole 2- 13.5 cm long, 1- 2 mm truck. Peduncle 1-2.5 cm long, 1-2 mm thick. Basal bracts caducous. Receptacle el­ lipsoid to subglobose, when fresh 2-2.5 cm, wh en dry 1- 1.5 cm in diameter, at maturity pale orange-green to pale yell ow with paler green to white spots.

Distribution. From Uganda to Sierra Leone (or to Sénégal?), also in Bioko (= Femando Po). In ± open forest, often in secondary vegetation and in gallery forest; at alti tud es up to 1600 m. The pollinator is CourtelIa camerunensis (Wiebes) (see also subspp. macrosyce and ulugurensis), but in Gabon also C. gabonensis Wiebes is a pollinator.

95b. Ficus ottoniifolia subsp. multinervia c.c. Berg Berg, Kew Bull. 43: 89 (1988).

Tree, treelet or sometimes liana. Leafy twigs usually pale brown wh en dry. Lamina (4-)10-21 x (1.2-)4-6.5 cm, usually broadest above the middle, coriaceous to subcoriaceous, apex acuminate (or rounded), base acute (to obtuse); lateral veins (4-)10- 15 pairs; petiole (1 - )2- 11 cm long, I- U mm truck. Peduncle 1- 2 cm long, 1-1.5 mm thick. Basal bracts caducous. Receptacle el­ lipsoid, when dry 1-1.2 cm in diameter.

Distribution. West Africa, from Sénégal to Ivory Coast. In more or less open (places in) forest; at low altitudes. The pollinator is CourtelIa gabonensis Wiebes.

154 95c. Ficus ottoniifolia subsp. lucanda (Ficalho) C.C. Berg Berg, Kew Bull. 43: 90 (1988).

F. /ongepeduncu/ata De Wild. (l913a), F. /ucanda Ficalho (1884), F. slerculioides Warb. (1894).

Tree or treelet. Leafy twigs usually pale brown when dry. Lamina 4.5-15 x 2 cm, mostly broadest at or below the middle, (sub)coriaceous, apex acuminate to subcaudate, base acute to obtuse, sometime alrnost rounded; lateral veins (3-)4--8 pairs; petiole 1.5-7 cm long. Peduncle 0.5-2.5 cm long, 1- 1.5 mm thick. Basal bracts caducous. Receptacle ellipsoid to subglobose, when fresh 1.5-2 cm, when dry 0.8- 1.2(-1.5) cm in diameter.

Distribution. From Angola through Zaïre to U ganda and NW. Tanzania, and to Gabon. In wet to rather dry forest; at altitudes up to 1500. Subspecies lucanda difTers from subsp. macrosyce in the smaller figs (0.7- 1.5 as aginst 2-2.5 cm, in diameter, wh en dry) and from subsp. ulugurensis in the caducous basal bracts and the smaller number of lateral veins (3- 8 instead of 8-14 pairs). The pollinator is Cour/elia scobinifera (Waterston). A parasitoid is Seres levis Waterston.

95d. Ficus ottoniifolia subsp. macrosyce C.c. Berg Berg, Kew Bull. 43: 90, fig . 4 (1988).

Liana or treelet, sometimes a tree. Lamina (4--)8-16 x (1.2-)3- 7 cm, chartaceous, usually broadest in the middle, apex shortly to long-acuminate, sometimes rounded, base emarginate (to subcordate); lateral veins (4--)6-8 pairs; petiole (0.8- )1.5-7 cm long, 0.5- 1 mm thick. Peduncle 1-1.5(- 2) cm long, ca. 2 mm thick. Basal bracts caducous. Receptacle ellipsoid to subglobose, wh en fresh 2.5-4 cm, when dry ca. 2- 2.5 cm in diameter, at maturity dark olivaceous or red-brown with pa Ie green to white spots.

Distribution. In NE. Angola, N. Zambia, and southern Zaïre. In gallery forest. Subspecies macrosyce is usually lianescent and differs from the .other sub­ species in the larger figs (2-2.5 as against 0.7-1.8 cm in diameter, when dry). The pollinator is CourtelIa camerunensis (Wiebes), as also in subspp. otto­ niifolia and ulugurensis.

95e. Ficus ottoniifolia subsp. ulugurensis (Mildbr. &_Burret) c.c. Berg Berg, Kew Bull. 43: 92 (1988).

F. modesta F. White (1990), F. scheffleri Mildbr. & Burret (1911), F. u/ugurensis Mildbr. & BUfret (1911).

155 Tree up to 10 m tall or shrub. Leafy twigs usually dark brown (to blackish) when dry. Lamina 7.5-15 x 3-7 cm, (sub)coriaceous, apex shortly acuminate, base (subacute to) rounded to subcordate; lateral veins 8- 12(-14) pairs; petiole 1.5- 4(-5.5) cm long, 1- 1.5 mm thick. Basal bracts (sub)persistent. Receptacle el­ lipsoid to subglobose, when fresh 2-3 cm, when dry (1-)1.2-2 cm in diameter, at maturity yellow with paler spots.

Distribution. In Kenya, Tanzania and Malawi. In rain forest, gallery forest, dry forest transitional to wood land, and in coastal bushland; at altitudes up to 1500 m. Subspecies ulugurensis differs from other subspecies in the (sub)persistent basal bracts. Moreover, the leafy twigs are usually dark brown when dry instead of pale brown as in the other subspecies. The pollinator is CourtelIa camerunensis (Wiebes), as in subspp. ottoniifolia and macrosyce.

96. Ficus tremula Warb.

Warburg, Bot. Jahrb. Syst., Pfianzengesch. & Pfianzengeogr. (EngJer) 20: 171 (1894); Berg et al., FI. Cameroun 28: 224, fig. 79 (1985) sub F. kimuenzensis; Berg, Kew Bull. 43: 95 (1988); Berg et al., FI. Trop. East Africa, Moraceae: 77 (1989).

Tree up to 10 m tall or shrub, sometimes lianescent, hemi-epiphytic or (sec­ ondarily?) terrestrial. Leafy twigs 1- 3(- 5) mm thick, sparsely minutely pu­ berulous. Lamina oblong to subobovate to elliptic to (sub)ovate, 2.5-8 x 0.7- 3 cm, subcoriacous to chartaceous, apex subacute to ratber faintly acuminate, base rounded to emarginate, margin entire; both surfaces glabrous or the midrib puberulous beneath; lateral veins 5- 9(-12) pairs; tertiary venation reticulate; petiole 0.7-3(-4) cm long, ca. 0.5(-1) mm thick; stipules 0.2-1, on flush up to 3 cm long, glabrous, caducous. Figs 1-6 together on up to 2 cm long (curved) spurs on the ol der wood; bud scales of the spurs (sub)glabrous; peduncle 0.5-2 cm long, 0.5-7 mm thick; basal bracts ca. 3 mm long, free parts caducous, sometimes subpersistent; receptacle subglobose to ellipsoid, when fresh 2- 2.5(- 3?) cm, when dry 1- 1.5(-1.8) cm in diameter, puberuious or subglabrous, at maturity greenish(?); wall when dry 0.5-1 mm thick, not or hardly wrinkled, sometimes slightly stipitate.

F. tremuia resembles F. ottoniifola in the common presence of ellipsoid figs. In eastem Africa, where both F. tremuia and F. ottoniifolia subsp. ulugurensis occur, the former can be recognized by the smaller leaves (2.5-7.5 as against 7.5-15 cm long). Moreover, the leafy twigs of F. ottoniifolia subsp. ulugurensis usually are dry dark brown, those of F. tremuia subsp. tremuia pale brown.

Three allopatric subspecies can be recognized.

156 96a. Ficus tremula subsp. tremula

F. pulvinata Warb. (1894).

Tree or sometimes a climber. Twigs usually drying yellowish or greyish. Lamina mostly drying brown above and greenish beneath, base rounded to emarginate; lateral veins 5-8 pairs; petiole up to 3(-4) cm long, ca. 0.5(-1) mm thick. Re­ ceptacle ellipsoid to subglobose, when dry 1-1.5 cm in diameter.

Distribution. From South Africa (Natal) through Mozambique to Kenya. In lowland dry evergreen forest and coastal bush land; at low altitudes. The pollinator is CourtelIa wardi Compton.

96b. Ficus tremula subsp. kimuenzensis (Warb.) c.c. Berg Berg, Kew Bull. 43: 96 (1988).

F. kimuenzensis Warb. (1904), F. rudens Huteh. (1915).

Treelet or often a climber. Twigs drying dark red-brown to blackish. Lamina .drying brownish or greenish and without a colour contrast between upper and lower surface, base cordate to rounded; lateral veins 5-9 pairs; petiole (0.5-)1- 3.5(- 7) cm long, 0.5-1 mm thick. Receptacle ellipsoid to globose, when dry 1.2-1 .5 cm in diameter.

Distribution. From N. Angola through western Zaire to SE. Nigeria. In rain forest; at low altitudes. The two lowland subspecies can be easily told apart to the colour of the dried twigs: yellowish in subsp. tremu/a and dark brown to blackish in sub sp. ki­ muenzensis. The dried leaves of subsp. tremu/a are more or less dark coloured above and pa Ie beneath; those of subsp. kimuenzensis do not show a pronounced colour contrast.

96c. Ficus tremula subsp. acuta (De Wild.) c.c. Berg Berg, Kew Bull. 43: 96 (1988).

F. acula De Wild. (1921).

Tree or often a liana up to 15 m taU. Twigs drying dark brown to blackish, older wood with warty lenticels. Lamina drying brownish without astrong colour contrast between the upper and lower surface, base obtuse to rounded (to sub­ cordate); lateral veins 6--9(-12) pairs; petiole 1- 3(-4) cm long, 1-1.5 mm thick. Receptacle ellipsoid to subglobose, when dry 1.2- 1.5(- 1.8) cm in diameter.

Distribution. In E. Zaïre, Rwanda, Burundi and W. Kenya. In (sub)montane forest, at altitudes between 1600 and 2300 m.

157 97. Ficus artocarpoides Warb. Warburg, Ann. Mus. Congo (6) I: 23, fig. 23 (1904); Berg et al., Fl. Cameroun 28: 216, fig. 76 (1985); Berg et al., Fl. Trop. East Africa, Moraceae: 78 (1989).

F demeusei Warb. (1904), F elegans (Miq.) Miq. (1867), non Hasskarl (1844), F inkansuensis Warb. (1904), F kisantuensis Warb.(1904), Urostigma elegans Miq. (1848).

Tree up to lOm tall, hemi-epiphytic. Leafy twigs 2-4 mm thick, minutely pu­ berulous. Lamina oblong to subobovate or to (ob)lanceolate, 6-21 x 2-6 cm, coriaceous, apex subacute to obtuse (to rounded, base obtuse to acute, margin entire; upper surface glabrous, lower surface minutely puberulous on the main veins; lateral veins 10-16 pairs, tertiary vena ti on reticulate to parallel to the lateral veins; petiole 1- 3.5 on long, (1-)2 rnm thick, glabrous or minutely pu­ berulous; stipules 0.3-4 cm long, puberulous or glabrous, caducous. Figs up to 5 together on up to 4 cm long and ca. I cm thick (curved) spurs on the main and les ser branches (to just below the leaves); bud scales of the spurs densely puberulous; peduncIe 1.5-3.5 cm long; basal bracts with the free parts caducous; receptacIe (sub)globose, when fresh 3-4 cm, when dry 2- 3.5 cm in diameter, minutely puberulous, at maturity greenish to purplish or yellow to orange; wall when dry 1-1.5 mm thick and wrinkled.

Distribution. From Angola to Uganda and Ivory Coast. In evergreen (rain and gallery) forest; at altitudes up to 1200 m. The material from the southern and eastern part of the species range has relatively narrow and more thickly coriaceous leaves with numerous lateral veins; towards the western extension of its range, the leaves become broader and thinner and they have fewer lateral veins. The pollinator is CourtelIa penicula (Wiebes), but also CourtelIa hladikae (Wiebes) was recorded.

98. Ficus polita Vahl Vahl, Enum. 2: 182 (1805); Berg et al., Fl. Cameroun 28: 227, fig. 81 (1985); Berg, Kew Bull. 43: 93 (1988); Berg, et al. , Fl. Trop. East Africa, Moraceae: 80 (1989).

Tree up to 15(-40) m tall, hemi-epiphytic or (secondarily) terrestrial. Leafy twigs 2- 5 mm thick, glabrous or minutely yellowish puberulous, lenticels often con­ spicuous. Lamina ovate to almost elliptic or oblong, 5-15(- 24) x 3.5- 10(-15) cm, (sub)coriaceous, (at least the midrib beneath and also the petiole often drying blackish), apex acuminate, base cordate to truncate to rounded, some­ times to subacute, margin entire; both surfaces glabrous; lateral veins 3- 5, 5-8(- 9), or (9-)10-12 pairs, tertiary venation partly scalariform to reticulate; petiole 2-12 cm long, 1-2 mm thick, glabrous; stipules 0.5- 2 cm long, glabrous, caducous. Figs 1-4 together on up to 3 cm long spurs on the older wood; bud scales of the spurs (almost) glabrous; peduncIe, 0.8-2 cm long; basal bracts 3- 5

158 mm long, coriaceous, persistent; receptacle globose to obovoid (at least when dry often shortly stipitate), when fresh 2--4(or more?) cm, when dry 1.5--4 cm in diameter, whitish puberulous, at maturity greenish to purplish; wall when dry 2- 3 mm thick, wrinkled.

Two subspecies can be recognized.

98a. Ficus polita subsp. polita (fig. 35) F barombiensis Warb. (1904), F megapoda Baker (1885), F niamniamensis Warb. (1904), F podophylla Baker (1 88 5), F syringifolia Warb. (1894), non Kunth & Bouché (1 847), F umbrosa Sim (1909), Uros/igma poli/urn (Vahl) Miq. (1 847).

Lamina ovate to elliptic; lateral veins 3-5 or 5- 8(- 9) pairs. Peduncle 1- 2 cm long. Receptacle when fresh (2- )3--4 cm, when dry 2--4 cm in diameter.

Distribution. From Sénégal to Uganda, southwards to N. Angola and to South Africa (Natal), also in Madagascar. In evergreen (rain and gallery) forest; at altitudes up to 1200 m. In S. Malawi and Mozambique this subspecies is often represented by a form with relatively small leaves having 3- 5 pairs of lateral veins and also often a rounded base. In West and Central Africa th is species can be easily confused with F. umbella/a. The pollinator is Cour/elia bekiliensis (Risbec): the nominate subspecies in Madagascar, subsp. bispinosa (Wiebes) on the continent of Africa.

98b. Ficus polita subsp. brevipedunculata c.c. Berg Berg, Kew Bull. 43: 93 (1988).

Lamina oblong to subovate or ovate; lateral veins (8-)10-12 pairs. Peduncle 0.8- 1.2 cm long. Receptacle when fresh 2- 2.5, when dry 1. 5-2 cm in diameter.

Distribution. In W. Zambia, N. Malawi, and Tanzania. In forest; at altitudes above 1500 m. This subspecies is only known from a few collections, but it is morphologically distinct from subsp. polita in the greater number of lateral veins, the shorter peduncle, and probably also in having smaller figs.

99. Ficus bizanae Hutch. & Burtt-Davy Hutchinson & Burtt-Davy, Kew Bull. (1921 ): 52 (1921); van Greuning, S. Afr. J. Bot. 56, 624 (1990).

Tree up to 12 m tall, (secondarily?) terrestrial. Leafy twigs 2--4 mm thick, glabrous or minutely puberulous, with relatively many lenticels. Lamina elliptic to oblong (to (sub)ovate), 4--13 x 2-7.5 cm, (sub)coriaceous, apex (faintly)

159 J) . )10U~ Fig. 35. Ficus po/ila Vahl subsp. po/ila. I, leafy twig; 2, short-shoot with figs. MIer Berg et al. (1985, pI. 81). acuminate, base acute to subattenuate, sometimes to rounded, margin entire; bath surfaces glabrous; lateral veins 3-11 pairs, tertiary venation reticulate; petiole 1.2-5.5(-8.5) cm long, 1-1.5 mm thick, glabrous; stipules 0.2-0.5 cm, on flush up to 3 cm long, glabrous or minutely puberulous, caducous. Figs on up to 1.5 cm long, unbranched or branched spurs; bud scales of the spurs minutely

160 appressed puberulous; peduncle 0.7- 2.5 cm long, 2- 2.5 mm thick, rninutely pu­ berulous; basal bracts 3-5 mm long, persistent; receptacle eUipsoid to sub­ globose to ovoid, when fresh 2.5-4.5 cm, wh en dry (1.8-)2-3 cm in diameter, rather densely puberulous to glabrous, at maturity yeUow-green; waU when dry ca. 3-7 mm thick, hardly wrinkled, ± warty, and apex of receptacle slightly protruding.

Distribution. South Africa (Natal and Transkei). In riverine forest; at low al­ titudes. This species shows close affinities to F. polita.

100. Ficus chirindensis C. C. Berg Berg, Kew Bull. 43: 78, fig. 1 (1988); Berg et al., FI. Trop. East Africa, Moraceae: 79 (1989).

Tree up to 35 m taU, (secondarily?) terrestrial. Leafy twigs 2-4(-6) mm thick, rninutely puberulous. Larnina oblong to eUiptic to (sub)ovate, (4-)6--12(-16) x (2.5--)3-5.5(-7.5) cm, subcoriaceous to coriaceous, apex (sub)acuminate, base cordate to trunctate (to rounded or obtuse), margin entire; upper surface minutely puberulous on the midrib or glabrous, lower surface minutely pu­ berulous on the main veins, sometimes glabrous; lateral veins (6--)8- 12 pairs, tertiary venation reticulate; petiole (1.5- )2-4(-6) cm long, 1.5-2 mm thick, rninutely puberulous or glabrous; stipules 0.3--0.5 cm (on flush up to 4 cm) long, outside with appressed (on long stipules ± patent) white hairs, caducous. Figs up to 3 together on up to 2 cm long spurs on the older wood, down to the main branches; bud scales (almost) glabrous; peduncle (1-)1.5--2(-4) cm long; basal bracts ca. 2.5(-3) mm long, chartaceous, caducous; receptacle (sub)globose, when fresh 2.5-4 cm, when dry 1.5--3 cm in diameter, minutely puberulous, at maturity greenish to yeUowish with brown spots; waU when dry ca. 2 mm thick, wrinkled.

Distribution. Malawi, Mozambique (Tete), E. Zimbabwe, Tanazania, Kenya, and E. Zaire. In submontane evergreen forest, at altitudes between 1000 and 1600 m. This species shows similarities to both F. sansibarica and F. ottoniifolia subsp. ulugurensis and it is difficult to distinguish herbarium material. Most helpful is the presence of short appressed white hairs on the stipules. The stipules of F. chirindensis are glabrous and those in F. sansibarica subsp. sansibarica are dis­ tinctly ciliolate. The lower leaf surface in F. chirindensis is often puberulous, while glabrous in F. ottoniifolia subsp. ulugurensis. The poUinator is CourtelIa malawi Wiebes.

101. Ficus sansibarica Warb. Warburg, Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 20: 171 (1894); Berg et al, FI. Carneroun 28: 220, fig. 78 (1985) sub F macrosperma; Berg, Kew Bull. 43: 93 (1988); Berg et al., FI. Trop. East Africa, Moraceae: 79 (1989).

161 Tree up to 20(-40) m tall, hemi-epiphytic or (secondarily?) terrestrial. Leafy twigs 2-5 mm thick, glabrous or sparsely minutely puberulous. Lamina oblong to lanceolate or to elliptic or (sub)ovate, 4.5-13(-24) x 2-6(- 11.5) cm, coriaceous to subcoriaceous, (at least the midrib beneath (and the petiole) usually drying (red-)brown, apex acuminate to subacute or to subobtuse (to rounded), base rounded to (sub)cordate to subacute, margin entire; both surfaces glabrous; lateral veins 5- 10(- 14) pairs, (in adult specimens) loop-con­ nected 1-2 mm from the margin, tertiary venation (predorninantly) reticulate; petiole (0.8-)2-5.5(-8) cm long, 1-2(-3) mm thick, glabrous, usually drying (red-) brown; stipules 0.1 - 1.5 cm, on flush up to 4.5 cm long, glabrous or ciliolate, caducous or on flush subpersistent. Figs 2-4 together on short up to ca. 2(- 5) cm long, branched and finally more or less cushion-shaped or on up to 15 cm long straight peg-like or sometimes curved spurs on the ma in or also the lesser branches; bud scales of the spurs densely puberulous; peduncle 1.2- 2.5(-5) cm long; basal bracts 3-5 mm long, their free parts caducous, occasionally subper­ sistent; receptacle (sub)globose, when fresh 2-6(-10) cm when dry 1.5-3(-6) cm in diameter, brownish to yellowish puberulous, at maturity greenish or partly purplish; wall when fresh 5- 10 mm, when dry 2-4(-5) mm thick and ± wrinkled or sometimes smooth.

Two allopatric subspecies can be recognized.

IOla. Ficus sansibarica subsp. sansibarica F. delagoensis Sim (1909), F. langenburgii Warb. (1901).

Larnina mostly oblong to lanceolate, often less than 10 cm long; lateral veins 5-10(-14) pairs; stipules ciliolate. Fig-bearing spurs up to 3.5(-5) cm long.

Distribution. From South Africa to SE. Zambia and through Malawi to Tanzania and Kenya. In forest and coastal bushland; at altitudes up to 900 m. This subspecies can be distinguished from subsp. macrosperma by its ciliolate stipules. Moreover, the spurs are short (mostly up to 5 cm long), becoming much longer (up to 10, or sometimes up to 15 cm long) in subsp. macrosperma. The pollinator is CourtelIa armata (Wiebes).

IOlb. Ficus sansibarica subsp. macrosperma (Mildbr. & Burret) e.C. Berg Berg, Kew Bull. 43: 94 (1988).

F. brachylepis Hiern (1900), F. buntingii Huteh. (1915), F. gossweileri Huteh. (1915), F. macro­ sperma Mildbr. & Burret (1911), F. mangenotii A. Chev. (1949), F. ugandensis Huteh. (1915).

Lamina oblong to lanceolate and often less than 10 cm long or elliptic to oblong or to (sub)ovate and then often longer than 10 cm; lateral veins 8-12 pairs; stipules glabrous. Fig-bearing spurs up to 10(- 15) cm long.

162 Distribution. From N. Angola and N. Zambia to Uganda and Guinea-Bissau. In · ra in forest and drier types of evergreen forest to savanna woodland; at al­ titudes up to 1200 m. In the southern part of the area of the subspecies the leaves are usually larger and broader than in West Africa. The pollinator is CourtelIa armata (Wiebes).

102. Ficus dryepondtiana De Wild.

De Wildeman, Repert. Spec. Nov. Regni Veg. 12: 303 (1913b); Berg et al., Fl. Cameroun 28: 222, fig. 78 (1985).

Tree or shrub, hemi-epiphytic. Leafy twigs 1-3 mm thick, glabrous or minutely puberulous, when dry hollow. Lamina oblong to subovate (or ovate) or to sub­ obovate, 6-16 x 2.5-6(-7) cm, chartaceous to subcoriaceous, (drying (dark) brown), apex acuminate to subacute, base acute to truncate to subcordate (to cordate), margin entire; both surfaces glabrous or lower surface minutely pu­ berulous on the main veins; lateral veins 5- 8 pairs, loop-connected 3- 8( 10) mm from the margin, often with a short straight part, departing the midrib at angles to 90° (!), tertiary venation reticulate; petiole (1 - )1.56--4.5 cm long, ca. 1.5 mm thick, glabrous or sparsely puberulous, (often drying blackish); stipules 0.3-2 cm long, glabrous or sparsely minutely puberulous, caducous or on Bush sometimes subpersistent. Figs on cushion-shaped or up to 3 cm long peg-like spurs on the main branches (and on the trunk?); bud scales of the spurs yellow shortIy ve­ lutinous to puberulous; peduncle 1- 3 cm long; basal bracts with their upper free parts caducous; receptacle globose, when fresh 4--5 cm, when dry 3-4.5 cm in diameter, yell ow to brownish puberulous to hirtellous, rather densely so around the ostiole, at maturity green with small white spots; wall when dry 2-3 mm thick, ± wrinkled.

Distribution. Zaire, the Central African Republic, Congo, Cameroun, and Gabon. In lowland rain forest and gallery forest. This species is very closely related to F sansibarica. The two taxa might even be conspecific as they are similar in many characters. But the leaves of adult specimens are distinct. However, saplings of F sansibarica have sometimes leaves somewhat reminiscent of those of F dryepondtiana. F dryepondtiana may only represent a form that retained traits of the juvenile leaf. The pollinator is Cour/elia sylviae Wiebes.

103. Ficus umbeUata Vahl Vahl, Enum. 2: 182 (1805); Berg et al., FI . Cameroun 28: 224, fig. 80 (1985).

F mega/odisca Warb. (1904), F po/ira Vahl var. persicicarpa Huteh. (1916), F tubercu/osa Hiern (1900).

163 Tree up to 10 (or more?) m tall, hemi-epiphytic or terrestrial. Leafy twigs 3-11 mm thick, glabrous, lenticels (rather) conspicuous, ± concentrated in rings bel ow the nodes. Lamina broadly ovate to broadly elliptic (to suborbicular), 11-26 x 7.5-21 cm, coriaceous, apex shortly acuminate, base truncate to sub­ cordate (to cordate), margin entire; both surfaces glabrous; lateral veins 7-12 pairs, often furcate far from the margin, tertiary venation partly scalariform; petiole 3-12.5 cm long, 2-4 mm thick, glabrous; stipules 0.3-0.5 cm long, glabrous, caducous. Figs on up to 1 cm long spurs on the older wood; bud scales usually ± densely minutely puberulous, when dry ± wrinkled; peduncle 0.2-0.9 cm long; basal bracts ca. 4 mm long, persistent; receptacle (sub)globose, when fresh ca. 3 cm, when dry 2- 2.5 cm in diameter, minutely puberulous, at maturity yellowish-green with brighter spots, wall when dry 1-2 mm thick, wrinkled.

Distribution. From NW. Angola to the Central African Republic and westwards to Sénégal. In savanna woodland, gallery forest and rain forest; at low altitudes. This species with broadly ovate to broadly elliptic leaves can be confused with F. polita, and juvenile forms of it possibly also with F. bubu. The figs in F. umbella ta and F. bubu are rather similar. F. umbella ta can be distinguished from F. bubu in the colour of the dried leaves (brown in F. umbella ta and greenish in F. bubu) and the bud scales of the spurs (puberulous and ± wrinkled when dry in F. umbella ta and sublepidote when dry in F. bubu). The pollinator is CourtelIa medleri (Wiebes).

104. Ficus bobu Warb. Warburg, Ann. Mus. Congo (6) 1: 3, fig. 8 (1904); Berg et al, Fl. Carneroun 28: 228, fig. 82 (1985); Berg et al, Fl. trop. East Africa, Moraceae: 81 (1989).

F. kyimbilensis Mildbr. (1953), F. pachypleura Warb. (1904a).

Tree up to 20(-30) m tall, hemi-epiphytic or often terrestrial; bark pale green to whitish. Leafy twigs 6-12 cm thick, glabrous or minutely puberulous, peri­ derm often flaking ofT. Lamina elliptic or sometimes to oblong or to suborbi­ cular, 12- 30 x 6-23 cm, coriaceous, apex (very) shortly acuminate to almost rounded, base obtuse to rounded or (especially in large leaves) to cordate, margin entire; both surfaces glabrous; lateral veins 6-8(- 9) pairs, often furcate far from the margin; tertiary venation partly scalariform; petiole 3.5-11(-16) cm long, 2-5 mm thick, glabrous; stipules 0.3-0.5 cm, on flush up to 4 cm long, glabrous or partly puberulous, caducous. Figs on short (often almost spine-like) spurs on the main branches and on the trunk; bud scales of the spurs minutely puberulous, mostly apiculate, when dry sublepidote; peduncle 0.7-1 cm long; basal bracts, 4-5 mm long, persistent; receptacle globose, when fresh ca. 3 cm, when dry ca. 2.5 cm in diameter, glabrous or minutely puberulous, at maturity brownish; wall when dry ca. 2 mm thick and wrinkled.

164 Distribution. From Ivory Coast (or from Sénégal?) to Kenya, and southwards to W. Angola and South Africa (Natal) In various types of forest, often in secondary growth ; at altitudes up to 1200 m .. The species is rather rare (or has been overlooked). It is usually terrestrial and forms rather tall trunks with a whitish bark. The pollinator is CourtelIa michaloudi (Wiebes).

105. Ficus ovata Vahl (fig. 36)

Vahl, Enum. 2: 185 (1805); Berg et al., FI. Cameroun 28: 230, fig. 83 (1985); Berg et al. , Fl. Trop. East Africa, Moraceae: 81 (1989).

F. a/ricana Kunth & Bouché (1847), F. asymmelrica Huteh. (1915), F. brachypoda Huteh. (1915), F. buchneri Warb. (1894), F. johnslonii Stapf (1906), F. megaphylla Warb. (1904), F. megaphylla var. glabra Warb. (1904), F. oClomelifolia Warb. (1904), F. ovala var. oClomelifolia (Warb.) Mildbr. & BUITet (1911), F. pseudo-elaslica Hiern (1900), F. sapinii De Wild. (1913b), F. speclabilis Kunth & Bouché (1847), F. luberculosa Hiern var. elliplica Hiern (1900), F. vohsenii Warb. (1894), Urosligma ovalum (Vahl) Miq. (1847).

Tree up to 10(-25) m tall, hemi-epiphytic or terrestrial, sometimes shrubs or lianescent. Leafy twigs 6-12 mm thick, densely white to yellowish puberulous to pubescent or to almost glabrous, with distinct scars of the axillary bud covers. Lamina ovate to elliptic to subovate or to oblong, (5-)9-31 x (3.5-)6-20 cm, coriaceous, apex acuminate, base cordate to trunctae to obtuse or subacute, margin entire; upper surface glabrous or sparsely puberulous in the lower part of the lamina, lower surface sparsely to densely white (to brownish) puberulous to hirtellous or glabrous; lateral veins 10-14 pairs; tertiary venation partly scalariform; petiole 3-10(-13) cm long, 2-4 mm thick, puberulous, hirtellous or glabrous; stipules 0.3- 1 cm long, glabrous or puberulous to pubescent, caducous. Figs solitary (or in pairs) in the leafaxils, just below the leaves, or sometimes (also) on the older wood, initially enclosed by an up to 2.5 cm long axillary capyptrate bud covers, these being caducous; peduncle up to 0.5(-7) cm long, (3-)4-6 mm thick; basal bracts ca. 3-4 mm long, persistent; receptacle ovoid to ellipsoid, sometimes subglobose or to obovoid, when fresh 3-5 x 2.5- 4.5 cm, when dry 1-4 x 1- 3 cm, puberulous to pubescent, at maturity greenish; wall when dry ca. 1.5-2 mm thick, smooth.

Distribution. From Sénégal to S. Ethiopia and Kenya, southwards to N. Angola, N. Zambia, Malawi, and Mozambique. In savanna woodland, forest edges, riverside forest, secondary forest; at altitudes up to 2100 m. Often planted. The species is often terrestrial. In contrast to the other species of subsection Caulocarpae the figs are not borne on spurs on the older wood, but rather in their leafaxils or just below the leaves. The pollinator is CourtelIa hamifera Kieffer: the nominate subspecies in W. Africa, the sub sp. modesta (Wiebes) in Uganda. Parasitoids are Seres a. armipes Waterston (in W. Africa) and S. a. breviceps Wiebes (Uganda), Philotrypesis africana Grandi and Philosycus monstruosus (Grandi).

165 Fig. 36. Ficus ovata Vahl. I, leafy twig with figs; 2, pistillate fiower; 3, staminate fiower. After Berg et al . (1985, pI. 83).

INTRODUCED SPECIES Several species have been introduced from Asia and cultivated as ornamentals in Africa, viz., F. altissima Bl., F. benjamina L., F. benghalensis L., F. carica L. [pollinator: Blastophaga psenes (L.)], F. elastica Roxb., F. microcarpa Linn. f. [pollinator Parapristina verticillata (Waterston)], F. obliqua Forst. f., F. pumila

166 L., F religiosa L. [pollinator Platyscapa quadraticeps (Mayr)], F vs. sagittata Vahl, F virens Ait.

REFERENCE

Corner, EJ.H. - Check-list of Ficus in Asia and Australasia, with keys to identification. Gdns' Bull. Sing. 21, 1- 186 (1965).

167

IV. The African fig wasps

Introduction For a long time the pollinating tig wasps were known as the Agaonidae, but in an important publication of recent date, Boucek (1988: 156-157) treated them as the nominate subfamily of the Agaonidae sensu lato, in which he also included wasps parasitic to the symbiosis, hitherto classified with the Pteromalidae and the Torymidae (see table 7). Much as there may he good taxonomie reasons for the new classitication when judged by the general Chalcidologist, for the tig wasp specialist there is no gain. To the contrary: the co-ordination of all groups men­ tioned within the Agaonidae is rather crude, as it does not give any indication

Table 7. Recent classifications of fig wasps. The Eurytomidae (in all systems considered a separate family) and Orrnyridae (by Bou~ek et al., 1981, c1assified as a subfamily of the Torymidae) are not included.

Bou~ek, 1988 Bou~ek et al. , 1981 Wiebes, 1981

AGAONIDAE s.1. 6. Agaoninae AGAONIDAE AGAONIDAE

TORYMIDAE 3. Sycoecinae (not placed) Sycoecinae

TORYMIDAE Sycophaginae Sycophaginae 5. Sycophaginae Sycophagini Sycophagini 4. Sycoryctinae Apocryptini Apocryptini Apocryptini

TORYMIDAE or PTEROMALIDAE Sycoryctini Sycoryctinae Sycoryctinae Philotrypesini Philotrypesini Sycoryctini Sycoryctini

PTEROMALIDAE PTEROMALIDAE 2. Otitesellinae Otitesellinae Otitesellinae

I. Epichrysomallinae EpichrysomaIlinae EpichrysomaIlinae

169 of the unmistakable relationships that exist between e.g., the Otitesellinae and the Epichrysomallinae, and it conceals the obvious difTerence between e.g., the pair of Sycoscapter and Watshamiella (a.o. in the old group Sycoryctini), and Philotrypesis (Philotrypesini). Although 1 follow Boucek in treating most groups as subfamilies of the Agaonidae s.l., I maintain their division into tribes (as in Wiebes, 1981). A number of the African parasitoids were described by J. Risbec, in a series of papers (Risbec, 1951a - 1957) and, although some were aUocated to their proper genus (Wiebes, 1970), for many there remains incertitude as to their identity. A tentative key to all groups, and a survey of the species known, are given below. Most of the generalized pictures of the genera were taken from a survey of the fig insects from New Guinea (Wiebes, 1982a). Some of the host records mentioned in the list of species, were conc1uded from their association with an Agaonid, the host of which is known. In all, the recent fig-names (Berg, 1990b) are used. Most of the following key is taken from Boucek et al. (1981: 159-164) and Wiebes (1981: 544-548).

KEY TO THE GENERA OF AFRICAN FIG WASPS

1. Females (fully winged) . . ' . .. , . , ...... 2 - Males (most are apterous, but some may have wings) .. .. , . .. . . ' . .. , 22

2. Gaster with a distinct tail formed either by the ovipositor and its valves, or also by extremely narrowed last one or two tergites . , . . . . , . . . . , ' ...... 3 - Gaster without such a slender tail (Comptonie/la has a short extension of the gaster!); ovipositor and its valves hardly exserted . . . , ...... , ...... 12

3. Gastral tail formed by one or two ultimate urotergites and the valves of the ovipositor (S YCORYCTINAE) . , ... . , ...... , .... , ... . , ...... ,. 4 - Last tergite short, tai! formed by the ovipositor and its valves . ... , ...... 7

4. Tail fonned by two ultimate (eighth and ninth) urotergites and the ovipositor and its valves (Philotrypesini) ...... , .. . , .. , ...... Phi/otrypesis - Tai! fonned by the last (ninth) urotergite and short ovipositor-valves (Sycoryctini; Sycoscapteridea, to which two African species were referred, could not be fitted into the key) . ... , . . . , . . , .. , . , . , . .. . , . . ' ...... , .... , . 5

5, Stigmal vein produced into a prolonged or wide, 'boot-like' stigma. Fore wing with stout setae in the margino-stigmal angle. Epistomal margin with an acute prolongation , . . . , . , . . , ...... , . , ...... , . . , , , . . . , , . , . . . Sycoscapter - Stigma more normal. Fore wing at most with some stout setae (along the veins or in the margino-stigmal angle), but it may be pilose all over. Epistomal margin straight or with a median lobe , , , . . , . . . , . , . . , . . . . , , ' ...... , . ' . . , ., 6

6. Epistomal margin straight. Antenna with three ring-segments . , , . . . . Walshamie/la - Epistomal margin with a median lobe. Antenna with two ring-segments .. Sycoryctes

7, Gastral segments ventrally keeled (Apocryptini) .. . . , , . . ' ...... Apocrypla - Gastral segments more normal, not keeled ...... 8

170 8. Marginal vein thickened (and longer than short postmarginal) ...... SYCOECINAE - Marginal vein normal, slender ...... 9 9. Antennal scape strongly broadened, the third antennal segment bearing an appendage (however short) ...... AGAONINAE - Antennal scape not broadened, the third segment simple (Sycophagini; as '[dames' is questionable and Dynatogmus is not weil known, the key that follows is restricted to the three other genera. Some difTerential characters are borrowed from Boucek, 1988: 8) ...... 10 10. Fore tibia with a dorso-apical crest of two or three teeth, the hind tibia with an antiaxial row of live to eight teeth ...... Sycophoga - Fore tibia without such terminal teeth and also the hind tibia devoid of the antiaxial row of teeth ...... II 11. Pronotum medially more than 0.7S of the length of the mesoscutum; clypeus trans­ verse, weil delimited, but supraclypeal area not defined; on thorax at least scutellar square yellowish ...... Eukoebelea - Pronotum medially much shorter than scutum; clypeus and supraclypeal area not dis­ tinctly separated, but both weil defined on sides; thorax sometimes dark-coloured · ...... Apocryptophagus 12. (Also 00) Gaster with at least some coarse sculpture; marginal vein very long, at least seven times as long as short stigmal vein; antenna 13-segmented, with two anelli (ORMYRIDAE) ...... Ormyrus - (00: go on to 16) Gaster without any coarse sculpture, either smooth or very finely reticulate; marginal vein never so much longer than the stigmal vein ...... 13

13. Body with metallic gloss, very finely reticulate; apex of gaster slightly curved ventrad (OTITESELLlNAE) ...... 14 - Body non-metallic, frequently either almost smooth or without distinct sculpture on thorax and mostly also on head; apex of gaster often curving dorsad, never ventrad 16 14. Pronotum only visible from above as two lateral parts; antenna with three anelli ... . · ...... Otitesella - Pronotum much wider, distinctly visible from above; antenna with one or two anelli IS IS. Gaster ending in a short 'taiI', formed by the ninth urotergite and the ovipositor-valves; antenna with one anellus; pronotum almost as long as the mesoscutum ...... · ...... CompIOnielIa - Ninth urotergite and the ovipositor-valves not elongated; antenna with two anelli; pronotum collar-like ...... Philosycus 16. Thorax (and head) with distinct sculpture and pubescence; stigmal vein usually at acute angle or short marginal vein distinctly expanded (EURYTOMIDAE) ...... 17 - Body without distinct sculpture, although often microscopically alutaceous; thorax always with pubescence reduced to pairs of bristles; marginal vein not so short or, if so, then stigmal vein arising at right angle (EPICHRYSOMALLINAE) ...... 20 17. Gena posteriorly with strong and distinctly raised carina extending up to the tempIe; thorax densely umbilicately punctured ...... 18 - Gena posteriorly with an edge which is blunt, or at most indistinctly carinate in middle or at mandible; umbilicate punctures, if present on thorax, usually not very dense . .. 19

171 18. Marginal vein distinctly swollen; gaster with a distinct petiole which has anteriorly a transverse crest; fore coxa antero-ventra1ly flat; genal carina smoothly joining mouth margin; body usually extensively rufous, in darkest specimens at least ventrally . . . . . · ...... Syceurytoma - Marginal vein normal, slender; gaster in Ç> virtually sessile; fore coxa on antero-ventral surface with strong oblique carina; genal carina with a tooth at base of mandible; body black, legs heyond coxae usually orange ...... Eurytoma 19. Mesostemal part of thorax with distinct though not high median projection excavated from the sides; marginal vein in Ç> not longer than stigmal which is usually distinctly curved; wing sometimes without fuscous spot (macula) ...... Ficomi/a - Mesostemum regularly sloping, posteriorly with median hairy depression (at mid co­ xae); marginal vein always with fuscous macula and frequently longer than the stigmal vein ...... Sycophi/a 20. Tarsi 4-segmented ...... 21 - Tarsi 5-segmented. Notauli groove-like, complete or nearly so (some 0 0) ...... · ...... Camarothorax

21. Notauli incomplete; male mandibles not falcate; gaster subsessiie ...... Sycotetra - Notauli complete; male mandibles falcate; gaster petiolate ...... Odontofroggatia 22. Apterous males ...... 23 - Half or fully winged ...... 33 23. The last four or five gastral segments tubularly lengthened; antenna reduced to five to seven segments ...... AGAONINAE - Last gastral segments not tubularly lengthened; antenna mostly consisting of more segments (if in doubt, take this alternative) ...... 24 24. Gaster ending in two gill-like structures (actually, outgrowths of the spiracular peritre­ mata). For Africa two genera are known in the male sex, but they cannot he further separated. Not all extralimital genera (some of which might occur in Africa) key out here ...... Sycophagini - The spiracular peritremata may he large, but they are not exserted as outgrowths . . . 25 25. Body rather long and slender; antenna consisting of only three segments (Apocryptini) · ...... Apocrypta - Body more robust; antenna consisting of ten or more segments . . 26

27. Antenna ten-segmented (not counting the anelli) (OTITESELLINAE) 28 - Antenna consisting of eleven or twelve segments (SYCORYCTINAE) . 30 28. Tarsi tetrarnerous; antennal toruli situated in the upper half of the face; mesonotum separate ...... 29 - Tarsi pentamerous; antennal toruli situated close to the epistomal margin; mesonotum fused with the metanotum and propodeurn ...... Comptonie//a

29. Antennal scape large, dilated ...... Phi/osycus - Antennal scape more slender, if a bit expanded, not dilated ...... Otitese//a 30. Two basal segments of the hind tarsus strongly expanded, hearing long setae; hyposto- mal margin deeply incised hetween sublateral lobes (Philotrypesini) . . . . Phi/otrypesis - Hind tarsus at most with fust segment expanded, without long setae; hypostomal margin moderately emarginate or straight (three genera of the Sycoryctini) ...... 31

172 31 . Thorax with two terga in between the pronotum and the propodeum: most probably, these represent the meso- and metanotum ...... Sycoscapleridea - Thorax at most with three separate terga, viz., the pronotum, mesonotum (may be coalescent with the next), and the fused metanotum-propodeum...... 32

32. Basitarsus of hind leg not expanded. Last segment of all tarsi inflated . . . Sycoscapler - Basitarsus of hind leg expanded. Last segment of all tarsi normaI...... Sycoryctes 33. Fore wing wholly pilose, narrow; marginal vein rather long and postmarginal fully two times as long as the stigmal; antennal flagellum not or hardly longer than scapus (one genus of the Sycoryctini) ...... Watshamiella - Venation different; wing at base extensively bare; flagellum longer...... 34

34. Marginal vein thickened and a1most two times as long as stigmal . ... SYCOECINAE - Marginal vein not so long and thickened; head different. Go back to ...... 12

AFRICAN GROUPS AND SPECIES

Subfamily AGAONINAE

This is the group of fig pollinators, to be more fully treated in the next chapter.

Subfamily SYCOECINAE (fig. 37)

Because of their similar habits, the Sycoecinae form a parallel to the Agaoninae: for oviposition, the females enter the fig receptacIe through the narrow ostiole, which may explain their resemblance in the armature of the mandibles andJor fore tibiae. They do, however, not pollinate (but see Newton & Lomo, 1979, who concIuded otherwise). Grandi (1955: 102) published a key to the genera (Syco­ phagini, in his sense, pro parte). A recent review of Crossogaster and Pha­ goblastus was given by Wiebes (in Boucek et al., 1981: 168- 170). Before long, the group will be revised by Simon van Noort, at the time at Rhodes University, to whom I sent all material available to me. The following genera and species were described from Africa.

Crossogaster Mayr Mayr, Verh . zool.-bot. Ges. Wien 35: 189-192 (1885); Wiebes in Boucek et al., Tijdschr. Ent. 124: 169-170 (1981).

African species of Crossogasler: atratus Masi - CrossogasIer alralus Masi, 1917: 125--126 (S! only, A1dabra Isl., ex Ficus lU/ea Vah1). odorans Wiebes - CrossogasIer odorans Wiebes in Boucek et al. , 1981: 170--173 (Zambia, Zim­ babwe, ex Ficus lhonningii 81.). silvestrii Grandi - Crossogasler silvestrii Grandi, 1916a: 253- 265 (Sénégal, ex Ficus sp. [host of Allolriozoon helerandromorphum, i.e., F. lulea Vah1]). trifonnis Mayr - CrossogasIer Iriformis Mayr, 1885: 192 ([the apterous male exc1uded], Socotra, ex Ficus vasla Forssk.).

173 Fig. 37. Female of Phagoblastus barbarus Grandi.

Phagoblastus Grandi Grandi, Boll. Ist. Ent. Univ. Bologna 21 : lOO--JOI (1955); Wiebes in Boucek et al., Tijdschr. Ent. 124: 169-170 (1981 ).

African species of Phagoblastus: barbarus Grandi - Phagoblastus barbarus Grandi, 1955: 102-106 ( '? only, South Africa, ex Ficus thonningii BI.); Wiebes in Boucek et al., 1981: 174-175 (Zambia, Zimbabwe, ex Ficus thonningii BI.). bouceki Wiebes - Phagoblastus bouceki Wiebes, 1981 : 554-555 (Réunion, ex Ficus reflexa Thunb.). liodontus Wiebes - Phagoblastus /iodontus Wiebes, 1979b: 397-400 (Gabon, ex Ficus natalensis leprieurii (Miq.) Berg).

Philocaenus Grandi Grandi, Boll. Lab. Ist. Ent. Bologna 19: 38-39 (1952).

African species of Philocaenus: barbatus Grandi - Philocaenus barbatus Grandi, 1952: 40-45 (Sénégal, ex Ficus platyphy /la DeL).

Seres Waterston Waterston, Ent. monthly Mag. (3) 5: 275-277 (1919); Wiebes, Zool. Meded. Leiden 37: 234 (1961).

African species of Seres: armipes Waterston - Seres armipes Waterston, 1914: 276-277 ('? only, Ghana); Grandi, 1952: 34-38 ('? only, Sénégal, ex Ficus ovala Vahl).

174 subspecies breviceps - Seres armipes breviceps Wiebes, 1961 : 233 ('i' only, Uganda, ex Ficus ovata Vahl). levis Waterston - Seres levis Waterston, I 920a: 135--136 ('i' only, Uganda, ex Ficus olloniifolia lucanda (Ficalho) Berg). longicephalus Risbec - Seres longicephalus Risbec, 1951a: 381-383 ('i' only, Sénégal, ex Ficus gna­ phalocarpa [most certainly wrong]).

Sycoecus Waterston Waterston, Bull. ent. Res. 5: 253 (1914).

African species of Sycoecus: thaumastocnemus Waterston ~ Sycoecus thaumastocnemus Waterston, 1914: 253-256 ('i' only, Uganda, ex Ficus sp. [host of Agaonfasciatum, i.e., F c. cyathistipula Warb.]).

Subfamily SYCOPHAGINAE Tribe Sycophagini (fig. 38) The contents ofthis group have changed some times: see Boucek (1988: 186--187) for a discussion. Several genera occur in the Indo-Australian region; five are known from Africa, but their delimitation is very unsatisfactory.

Apocryptophagus Ashmead Ashmead, Mem. Carnegie Mus. I: 238, 366 (l904a); Wiebes, Ann. Soc. ent. France (N.S.) 17: 555 (1981). Synonym: Parakoebelea Joseph, Ann. Soc. ent. France 125: 97 (1957). This is the group of robust insects, in the female sex different from Eukoebelea

Fig. 38. Female and male of the Sycophaginae, after Wiebes (1982, fig . 10).

175 by the shorter ovipositor-valves. The genus is based on the African Chalcis explorator Coquerel, but some species from India that Joseph descrihed as Pa­ rakoebelea have to he included as weIl. There is a cataloque by Wiebes (1981). Three African species are known, the host figs belong to the subgenus Sycomo­ rus. Except perhaps by their size, the males cannot he difTerentiated from those of Sycophaga. From his observations on an unnamed Papuan species, Godfray (1988: 286) concluded it was responsible for transforming anormal gall (caused by Ceratosolen) into a large gall.

African species of Apocryptophagus: explorator (Coquerel) - Cha/cis explorator Coquerel, 1855: 426--427 ['i') and Apocrypta paradoxa Coquerel, 1855: 367-369 (0) (Réunion, ex Ficus mauritiana Lam.). Apocryptophagus explora­ tor - Wiebes, 1981 : 555 (Réunion, ex Ficus mauritiana Lam.). gigas (Mayr) - Eukoebelea gigas Mayr, 1906: 164-165 (0 only, Vemen, ex Ficus sycomorus L.). Parakoebelea gigas - Wiebes, 1968a: 318 (Ethiopia, Kenya, Tanganyika, ex Ficus sycomorus L.).? Otitesella gnaphalocarpae Risbec, 1951a: 333 ('i', Sénégal, ex Ficus gnaphalocarpa [= F sycomorus)). Risbec recorded several samples, which may form a mixtum of species (Wiebes, 1970: 9). Also Wiebes (1969a: 402) recorded the species from Sénégal. spec. Compton & McLaren - Apocryptophagus spec. Compton & McLaren, 1989: 58 (South Africa, ex Ficus sur Forssk.).

Dynatogmus Mayr

Mayr, Wien. ent. Ztg. 25: 181-182 (1906). This genus remained unclassified and its type species unrecognised, until Boucek (1988: 187) stated that it certainly helongs to the Sycophaginae 'and may he the same genus as wh at is known in America wrongly as Neosycophila'. There was one species named, which remains unknown to me: robustll'i Mayr - Dynatogmus robustus Mayr, 1906: 182 ('i' only, Cameroon, ex Ficus sp.). The sample from Cameroon ('Aus von Herrn Dr. Yngve Sjöstedt in Kamerun gesarnmelten Fei­ gen') also contained Ceratosolen acutatus Mayr (1906: 154), and it could thus have been ex Ficus vogeliana (Miq.) Miq. Also Sycophaga vicina Mayr was recorded from what probably was the same sample.

Eukoebelea Ashmead

Ashmead, Proc. ent. Soc. Wash. 6: 126 (l904b; replacement name for Koebelea Ashmead, 1904a, Mem. Carnegie Mus. I: 238). Some Sycophaginae, very close to Apocryptophagus and Sycophaga in the female sex, but with males lacking the peculiar appendages at the gastral tergites (Boucek, 1988: 188), are known as Eukoebelea. Although it is questionable whether this name will apply to all, it is used here for three African species associated with figs of subgenus Sycomorus.

African species of Eukoebelea (all known in the 'i' sex only): camerunensis Mayr - Eukoebelea camerunensis Mayr, 1906: 165 (Cameroon, ex Ficus sp.). Tbe host may have been Ficus vogeliana (Miq.) Miq.

176 sycomori Wiebes - Eukoebelea sycomori Wiebes, 1968a: 315, 317 (Tanganyika, Kenya, ex Ficus sycomorus L.). The species was recorded from F sycomorus in Ethiopia by Wiebes (in Aweke, 1979: 101), in Sénégal by Wiebes (1969a: 402), and in Namibia by Wharton et al. (1980: 479). spec. Compton & Nefdt, 1988: 105 (Cameroon, ex Ficus vallis-choudae De!.). spec. Wiebes - Eukoebelea spec. Wiebes, 1981 : 556 (Réunion, ex Ficus mauritiana Lam.).

Idarnes Walker

Walker, Ann. Mag. nat. Hist. 12: 47 (1843); Gordh, Univ. Kansas Sci. Bull. 50: 389-455 (1975). Although ldarnes probably is an Arnerican genus of fig insects associated with figs of section Americana Miq., the name was used also for one African species, distinguished in the females sex by being very slender and by the long ovipositor­ valves.

African species of [dames: gracile Wiebes - [dames gracile Wiebes, 1968a: 318- 319 ( 9 only, Kenya, ex Ficus sp. (probably, F sycomorus L.]).

Sycophaga Westwood (fig. 39)

Westwood, Trans. ent. Soc. Lond. 2: 222 (1840); Grandi, Boll. Lab. Zoo!. Portici 10: 221- 236 (1916a). In the subfamily, this is the only genus the females of which oviposit from inside the syconium: they have characteristic teeth in the dorso-apical comb of the fore

•.f.\

Fig. 39. Sycophaga sycomori (L.), female and male, after G alil (1968, fig. 6).

177 tibia and in the antiaxial row of the hind tibia, which probably are used when traversing the scales of the fig-ostiole. Galil et al. (1970) discussed the efTects of Sycophaga sycomori on the structure and development of the syconia of Ficus sycomorus. There are twelve nominal species known, all African, but - judging from the host figs that are assumed to have na more than one species of Sycophaga each - actually there may be only five or six. Grandi (19l6a: 234-236) provided a key to six species, containing all known males.

African species of Sycophaga: affticta Grandi - Sycophaga afflicla Grandi, 1916a: 250-252 (0 only, Guinea, ex Ficus sur Forssk.). ludging from the host-fig, S. ajJlicla may be the male of S. cyclosligma Waterston. callani Grandi - Sycophaga cal/ani Grandi, 1955: 98-100 (S' only, Zimbabwe, ex Ficus 'rhodesiaca Warb.'). F rhodesiaca Mildbr. & Burret is a synonym of F Ihonningii BI. (Berg, 1990: 168), but this is probably not the host of S. cal/ani. It may have been Ficus sycomorus L., which is the host of Ceraloso/en arabicus Mayr (as C. myersi Grandi recorded from the same sample as S. callam). cyclostigma Waterston - Sycophaga cyc/osligma Waterston, 1916: 126-129 (S' only, Zimbabwe, ex Ficus sp.); Baijnath & Ramcharun, 1983: 883-888, Baijnath et al., 1986: 348-349 (South Africa, ex Ficus sur Forssk.); Compton & McLaren, 1989: 58 (South Africa, ex Ficus sur Forssk.). See also under S. ajJlicla. depressa Risbec - Sycophaga depressa Risbec, 1956: 189-191 (S' only, Madagascar). gigantea Grandi - Sycophaga gigantea Grandi, 1916a: 243-246 (0 only, Eritrea, Cameroon, ex Ficus sp.). The samples from Eritrea (from Ghinda and Keren, leg. G. Rossetti, 1913) are those from which also Ceraloso/en arabicus Mayr was recorded (Grandi, 1917: 25) and th us could have been ex Ficus sycomorus L. The differences between S. giganlea and S. sycomori are mainly those of size, and for that reason S. gigantea could be Apocryptophagus? insularis Grandi - Sycophaga insu/aris Grandi, 1916a: 236-239 (S' only, Cape Verde Is., ex Ficus sp.). silvestrii Grandi - Sycophaga si/veslrii Grandi, 1916a: 239- 243 (Senegal, ex Ficus jerruginea Desf.'). F ferruginea Desf. is a synonym of the Australian Ficus rubiginosa Desf. ex Vent. (Corner, 1965: 26). From the African sample also Ceralosolen si/veslrianus Grandi was described, which was recorded from Ficus sur Forssk. as weil as (with some doubt) from F vallis-choudae Del. Apocrypla robusla, from the same sample, later proved to be associated with Ficus val/is­ choudae. Risbec's record (195Ia: 386-389) belongs to S. valentifUJe Grandi. sycomori (L.) - Cynips sycomori Linnaeus, 1758: 554 (Egypt, ex Ficus sycomorus L.). Sycophaga sycomori - Grandi, 1917: 30-39 (Eritrea, ex Ficus sycomorus L.). Sycophaga crassipes West­ wood, 1840: 222-223 (Egypt, ex Ficus sp.). The species was recorded from Greece (Waterston, 1916: 128), Syria, Israel, Egypt, Eritrea (see Grandi, 1917: 39, and for Ethiopia: Wiebes in Aweke, 1979: 101), Kenya and Tanganyika (Galil & Eisikowitch, 1968a: 262; Wiebes, 1968a: 312), Sénégal (Wiebes, 1969a: 402) and Namibia (Wharton et al. , 1980: 479). tenebrosa Grandi - Sycophaga si/veslrii tenebrosa Grandi, 1916a: 235, 1917: 39-42 (Uganda, ex Ficus sp.). As this species evidently is associated with Ceralosolen megacepha/us Grandi, the host may have been Ficus val/is-choudae Del. valentinae Grandi - Sycophaga valentinae Grandi, 1952: 32- 34 (S' only, Sénégal, ex Ficus gnapha­ /ocarpa Steud. [= F sycomorus L.I). Sycophaga si/veslrii - Risbec, 19151a: 386-389 (Senegal, ex Ficus gnapha/ocarpa). vicina Mayr - Sycophaga vicina Mayr, 1906: 163 (0 only, Cameroon, ex Ficus sp.). The host fig may have been Ficus vogeliana (Miq.) Miq. viduata Grandi - Sycophaga viduala Grandi, 1916a: 248- 250 (0 only, Cameroon, ex Ficus sp.).

178 Fig. 40. Female and male of Apocrypta, after Wiebes (1982, fig. 12).

Tribe Apocryptini

Apocrypta Coquerel (fig. 40) Coquerel, Rev. Mag. Zool. (2) 7: 369- 371 (1855); Grandi, BoU. Lab. Zool. Portici 10: 264-273 (1916a; 0 only); UIenberg, Proc. Kon. Ned. Akad. Wet. (C) 86: 63-94 (1983); UIenberg & van Pelt in UIenberg, Verh. Kon. Ned. Akad. Wet. (2) 83: 41-148 (1985). Ulenberg (1985: 41-148, key to the African species: 97-106) revised the genus which, next to Philotrypesis, now is the best known of the fig parasitoids. Some nominal species of Apocrypta had to remain incertae sedis, among which the African Apocrypta longitarsus imbecil/us Grandi and Apocrypta regalis Grandi, both from an unidentified Cameroon fig. Tbe nine known African species have a strict specific symbiosis with fig species of subgenus Sycomorus. UIenberg & Nübel (1982: 607-613) described tbe oviposition behaviour of Apocrypta robusta Grandi. Uien berg (1985: 149- 171) and later, Wiebes (l989b: 253), compared the classifications of Sycomorus, Ceratosolen and Apocrypta, which proved only partially to agree.

African species of Apocrypta: brachycephala Grandi - Apocrypta /ongitarsus brachycepha/a Grandi, 1916: 277-279 (0 only, Ca­ meroon, ex Ficus sp.). Apocrypta brachycepha/a - UIenberg, in UIenberg & van Pelt, in Ulen­ berg·, 1985: 129-132 (Ivory Coast, Nigeria, ex Ficus voge/iana (Miq.) Miq.).

• further, this correct, but rather cumbersome way of citation will be restricted to Uien berg, 1985.

179 cercaria UIenberg - Apocypta cercaria UIenberg, 1985: 109- 112 (Madagascar, ex Ficus cocculifo/a Baker var. saka/avarum Baker [= var. of F. sycomorus L.D. crypta UIenberg - Apocrypta cryp/a Uien berg, 1985: 124-126 (lvory Coast, Nigeria, Tanganyika, ex Ficus mucuso Welw. ex Ficalho). It is worthy of note that, although the ot her symbionts of Ficus mucuso are found to be identical to those of F. sycomorus, UIenberg could distinguish between the Apocrypta-species. guineensis Grandi - Apocrypta guineensis Grandi, 1916a: 282- 285 ( 0 only, Guinea, ex Ficus sp.); UIenberg, 1985: 121- 123 (Ivory Coast, Nigeria, Zambia, South Africa, ex Ficus sur Forssk.). imbecillus Grandi - Apocrypta /ongitarus imbeci//us Grandi, 1916a: 279-280 (0 only, Cameroon, ex Ficus sp.). longitarsus Mayr - Apocrypta /ongitarsus Mayr, 1906: 163- 164 (0 only, Cameroon, ex Ficus sp.); Grandi, 1916a: 274-276 (Eritrea, Sénégal, ex Ficus sycomorus L.): UIen berg, 1985: 118-121 (lsrael, ex Ficus sycomorus L.). Apocrypta minima Risbec, 1951a: 389-390 (0 only, Sénégal, ex Ficus gnapha/ocarpa [= F. sycomorus L.D. Apocryptophagus bambeyi Risbec, 1951a: 318 ('" only, Sénégal, ex Ficus gnapha/ocarpa [= F. sycomorus L.D. This species was also recorded from Ethiopia (Wiebes in Aweke, 1979: lOl), Kenya and Tanganyika (Galil & Eisikowitch, 1968a: 262-263), Sénégal (Wiebes, 1969a: 402) and Namibia (Wharton et al. , 1980: 479). perplexa Coquerel - Apocrypta perp/exa Coquerel, 1855: 369-371 (0 only, Réunion, ex Ficus mau­ ritiana Lam.); Wiebes, 1981 : 557 (Réunion, ex Ficus mauritiana Lam.); UIenberg, 1985: 106- 108 (same material). polyspina UIenberg - Apocrypta po/yspina UIenberg, 1985: ll2- ll5 (Madagascar, ex Ficus bo/ry­ oides Baker [det. c.c. Berg; not F. po/yph/ebia as in UIenberg, 1985]) regalis Grandi - Apocrypta rega/is Grandi, 1916a: 281 - 283 (0 only, Cameroon, ex Ficus sp.). robusta Grandi - Apocrypta /ongitarsus robusta Grandi, 1916a: 280-281 (0 only, Sénégal, ex Ficus ferruginea Desf. [see remark under Sycophaga si/vestril1). Apocrypta robusta - UIenberg, 1985: 126-129 (Ivory Coast, ex Ficus va//is-choudae DeL). setoptera UIenberg - Apocrypta se/op/era Uien berg, 1985: 115-118 (Madagascar, ex Ficus po/y­ ph/ebia Baker [det C.c. Berg, not F. /orrenrium Perrier as in Uien berg, 1985]).

Subfamily SYCORYCTINAE This group contains most genera forrnerly c\assified with the Torymidae andJor Pteromalidae, characterized in the femaie sex by the long extruded ovipositor­ valves. Boucek (1988: 180) differentiated it from the Sycophaginae by the undi­ vided femaie scutellum, which in the Sycophaginae bear conspieuous sublateral grooves. He took the Sycoryctini (ovipositor and its valves covered by the ex­ tended last tergite) and the Philotrypesini (ovipositor and its valves for a great part covered by the last two tergites) together, but here the oid division is maintained. Consequently, there are two tribes, viz., Philotrypesini and Syco­ ryctini.

Tribe Philotrypesini

Philotrypesis Förster (fig. 41) Förster, Verh. naturh. Ver. preuss. Rheinl. 35: 59- 60 (1878); Grandi, Boll. Lab. Zool. Portici 15: 33- 190 (192Ib); Wiebes in Boucek et al., Tijdschr. Ent. 124: 175 (1981). This genus is immediately recognized by the tubularly lenghtened eighth and ninth urotergites of the female, and by the deeply emarginate hypostomal edge

180 Fig. 41. Male and female of Philotrypesis, after Wiebes (1982, fig. 9). of the male. There are eight known African species, associated with figs of sec­ tion Ficus and Galoglychia (subsections Galoglychia, Chlamydodorae and Cau­ locarpae), but there are certainly many more (e.g., the Philotrypesis-species recorded from Ficus burtt-davyi by Baijnath & Ramcharun, 1988: 229, 231 ; and Compton & Nefdt, 1988: 102- 105); moreover, I expect Philotrypesis in African Sycidium, as it is also known from e.g., Indian Ficus exasperata. Grandi (1921b) gave a monograph of the genus; Wiebes (in Boucek et al., 1981 : 175) published a key and discussed most African species.

African species of Philotrypesis: africana Grandi - Philotrypesis a/ricana Grandi, 1921b: 162- 173 (Guinea and Nigeria, ex Ficus sp.). Judging from the Agaonine from the same sample, i.e., CourtelIa hamifera, the host fig is Ficus ovata Vahl. cnephaea Wiebes - Philotrypesis cnephaea Wiebes, 1981: 558- 559 (Réunion, ex Ficus reflexa Thunb.). erythraea Grandi - Phi/otrypesis ery thraea Grandi, 1921b: 118-128 (Eritrea, ex Ficus sp.). It prob­ ably is a symbiont of E/isabethiella allotriozoonoides (Grandi): host fig not known. finitimorum Wiebes - Philotrypesis finitimorum Wiebes, 1971: 373-378 (Sierra Leone, ex Ficus saussureana De.). longicomis Grandi - Philotrypesis /ongicornis Grandi, 1921b: 173-180 (Guinea, ex Ficus sp.). palmata Joseph - Philotrypesis pa/mata Joseph, 1954: 66-77 (India, ex Ficus palmata Forssk.). Recorded from Ficus palmata in Ethiopia by Wiebes (in Aweke, 1979: 100). Note that there is confusion over the pollinators of F. palmata and F. carica; see under B/astophaga psenes (Agaoninae, no. I). The three species P. caricae (L.) (from Ficus carica L.), P. erythraea and P. pa/mata are very close, if not specifically similar. parca Wiebes - Philotrypesis parca Wiebes in Bouèek et al., 1981 : 176-179 (Zambia, Zimbabwe, ex Ficus thonningii BI.).

181 selenitica Grandi - Philotrypesis selenitica Grandi, 1921b: 158-161 (Guinea, ex Ficus lutea Vahl). spec. Compton & Nefdt, 1988: 102- 105 (South Africa, ex Ficus burtt-davyi Hutch.).

Trihe Sycoryctini (fig. 42)

Boucek (1988: 183- 184) reconsidered the generic names used in this group and found it to be impossible to correlate the characters of the two sexes. He con­ cluded th at the subgroups 'are to he regarded as of mere specific (or species­ group) value' and synonymized almost all under Sycoscapter. I experienced sim­ ilar difficulties with the association of male and female characters of four genera

Fig. 42. Male and fema1e of the Sycoryctini, after Wiebes (1982, fig. 4).

182 (Wiebes, 1966b: 173), but - if only for heuristic reasons - prefer to keep them apart on characters of the one or the other sex. If a species, or a species-group, need not be difTerentiated from other such taxa in every possible character, why should it be clearly differentiated in both sexes? For Africa there are five generic names, viz., Grandimyia, Sycoryctes, Syco­ scapter, Sycoscapteridea and Watshamie/la, and 23 nominal species, for fifteen of whjch the host is not known.

Sycoscapter Saunders Saunders in Westwood, Trans. ent. Soc. Lond. for 1883: 34-35 (1883); Wiebes, Ent. Ber. Amst. 38: 186 (l978b); Wiebes in Boucek et al., Tijdschr. Ent. 124: 180 (1981). Grandimyia Risbec, Ann. Soc. ent. France 124: 148- 149 (1956). In the female fore wing, the stigmal vein is remarkably wide or elongate; in the male, the antenna has eleven free segments and the hind basitarsus is not ex­ panded, but (normally) the last segment of all tarsi is infiated. One species de­ scribed in Sycoscapter, i.e., S. anceps Wiebes, on renewed inspection has too many aberrant characters to maintain it in the genus: it is now reallocated to Sycoryctes. Wiebes (in Boucek et al., 1981: 180) published a key to three species and Wiebes (1981: 563-566) added two, but four species named by Risbec (1951a, b, 1956) were not included.

African species of Sycoscapter: cadenati (Risbec) - Goniogaster cadenati Risbec, 1951a: 325- 327 (S? only, Sénégal, ex Ficus sp.). See Wiebes (1970: 10) for a diseussion. comutu<; Wie bes - Sycoscapter comutus Wiebes in Boueek et al., 1981 : 180-182 (Zambia, Zim­ babwe, ex Ficus thonningii BI.); Wie bes, 1981: 564-565 (Réunion, ex Ficus reflexa Thunb.). gibbus Saunders - Sycoscapter gibbus Saunders in Westwood, 1883: 25 (S? only, Madagascar, ex Ficus politoria Lam.); Wiebes, 1978b: 187- 189 (Madagascar, ex Ficus politoria Lam., and Réunion, ex Ficus lateriflora Vahl). There were stated to exist small dilTerences between the two samples (Wiebes, 1978b: 188), without taxonomie eonsequences. montis Wiebes - Sycoscapter montis Wiebes, 1971 : 369- 373 (Sierra Léone, ex Ficus saussureana oe.). niger (Risbec) - [dames '(= Koebela)' nigra Risbec, 1951b: 1123- 1125 (S? only, Ivory Coast, ex Ficus sp.). The 0 is not recognizable, it may be Apocrypta (Wiebes, 1970: 12). senegalemis (Risbec) - Sycoryctes senegalensis Risbec, 1951a: 329- 333 (S? only, Sénégal, ex Ficus sp.). tananarivemis (Risbee) - Grandimyia tananarivensis Risbee, 1956: 149- 151 (S? only, Madagasear). Referred to Sycoscapter by Wie bes (1970: 12). tibialis Wiebes - Sycoscapter tibialis Wiebes, 1981: 563-564 (Réunion, ex Ficus rubra Vahl). spee. Wiebes - Sycoscapter spee. Wiebes, 1969a: 402 (S? only, Sénégal, ex Ficus gnaphalocarpa Steud. [= F. sycomorus L.D.·

Sycoscapteridea Ashmead Ashmead, Mem. Carnegie Mus. 1: 239 (1904); Wiebes, Tijdschr. Ent. 110: 428 (1967). An important character is found in the male thorax, both the mesonotum and metanotum of which are visible dorsally. Two species, described by llisbec

183 (1951a, b) in Goniogaster Mayr, were reallocated in Sycoscapteridea by Wiebes (1970: 10).

African species of Sycoscapteridea: bambeyi (Risbec) - Goniogaster bambeyi Risbec, 1951a: 328- 329 (C( only, Sénégal, ex Ficus gnap­ ha/ocarpa). ?Sycoscapteridea spec. Wiebes, I 969a: 401 (0 only, Sénégal, ex Ficus gnapha/o­ carpa Steud. [= F. sycomorus L.D. spec. Risbec - Goniagaster cadenati Risbec, 1951b: 1126 (including a?, Ivory Coast, ex Ficus close to F. gnapha/ocarpa). This is not the same as Goniogaster cadenati Risbec (I95Ia), the C( of which was referred to Sycoscapter and the 0 to Sycoryctes. See Wiebes (1970: 10).

Sycoryctes Mayr

Mayr, Verh. zool.-bot. Ges. Wien 35: 211-214 (1885); Wiebes in Bouëek et al., Tijdschr. Ent. 124: 182- 183 (1981). Some ten African species were described in, or reallocated to Sycoryctes. Wiebes (in Boucek et al., 1981: 183) published a key to the males of seven species.

African species of Sycoryctes: anceps (Wiebes) nov.comb. - Sycoscapter anceps Wiebes, 1981: 565-566 (Réunion, ex Ficus densi­ folia Miq.). This species could not be maintained in Sycoscapter, but, actually, the placing in Sycoryctes is not quite satisfactory either. The male, except for the norrnal last tarsal segment (not infiated), does resembIe Sycoscapter. caelebs Wiebes - Sycoryctes cae/ebs Wiebes, 1975: 231 - 233 (0 only, AJdabra Isl., ex Ficus rubra Forssk.; Wiebes, 1981 : 56&-567 (Réunion, ex Ficus densifolia Miq.). coccothraustes Mayr - Sycoryctes coccothraustes Mayr, 1885: 217- 218 (Socotra, ex Ficus vasta Vahl). For a discussion of the host record, see Wiebes (I977b: 210). comparabilis Wiebes - Sycoryctes comparabilis Wie bes, 1981 : 567 (C( only, Réunion, ex Ficus den­ sifo/ia Miq.). dagatiguyi (Risbec) - Apocryptophagus dagatiguyi Risbec, 1951a: 331-332 (S' only, Ivory Coast, ex Ficus sp.). hirtus Wiebes - Sycoryctes hirtus Wiebes in Boucek et al., 1981 : 186 (Zambia, Zimbabwe, ex Ficus thonningii BI.). lomaensis Wiebes - Sycoryctes /omaensis Wiebes, 1971 : 367- 369 (Sierra Léone, ex Ficus saussureana De.). remus Wiebes - Sycoryctes remus Wiebes in Bouëek et al., 1981: 18:>- 185 (Zambia, Zimbabwe, ex Ficus thonningii 81.); Wiebes, 1981: 566 (Réunion, ex Ficus rejiexa Thunb.). sebertianus Masi - Sycoryctes sebertianus Masi, 1917: 126 (C( only, SeychelIes Is.). sjoestedti Mayr - Sycoryctes sjoestedti Mayr, 1906: 167- 168 (Carneroon, ex Ficus sp.). truncatus Mayr - Sycoryctes truncatus Mayr, 1885: 218 (0 only, Socotra, ex Ficus vasta Forssk.). See a1so under S. coccothraustes. spec. Risbec - Goniogaster cadenati Risbec, 1951a: 327-328 (only the 0, Sénégal, ex Ficus gnapha­ /ocarpa) . Wiebes (1970: 10) noted: 'the description of the male reminds one of Sycoryctes'. This is not the same species as the female, which was referred to Sycoscapter, and also not the same as G. cadenati Risbec (195Ib: 1156), which could have been Sycoscapteridea.

Watshamiella Wiebes

Wiebes in Bouëek et al., Tijdschr. Ent. 124: 18&-189 (1981). This genus was erected for W a/ata, the male of which is fully winged (but tbis

184 proved also the case in a species of Philotrypesis). There appeared to be many more species, in Africa (Wiebes, 1981: 560-563) as weil as in the Indo-Australian region (Boucek, 1988: 182). There are three known African species.

African species of Watshamiel/a: alata Wiebes - Walshamiella a/ala Wiebes in Boueek et al., 1981: 189- 191 (Zimbabwe, ex Ficus Ihanningii BI.). fictitia Wiebes - Watshamiel/aficlitia Wiebes, 1981 : 562- 563 (Réunion, ex Ficus rubra Vahl). lucem Wiebes - Walshamiel/a /ucens Wiebes, 1981 : 560-561 (Réunion, ex Ficus densifa/ia Miq.).

Subfamily OTITESELLINAE There are three African genera, but many more are known from the Indo­ Australian region. In the world catalogue (Wiebes, 1967) many of the African forms are not included, because they were not yet known. Otitesella Westwood was known, but there were several species added; Philosycus Wiebes was not, aIthough indicated as an unnamed group (Wiebes, 1967: 436, 437); Com­ ptoniella, finally, is an addition now made: it is aberrant by the female 'tail' and the male pentamerous tarsi.

ComptonieUa Wiebes, gen. nov. (fig. 43) Diagnosis - Female with elongated ninth urotergite and ovipositor-valves, to­ gether forming a short tail; antenna with only one anellus; the pronotum is almost as long as the mesoscutum. Male with pentamerous tarsi; toruli situated anteriorly; antenna with one, very narrow anellus; meso-, metanotum and pro­ podeum fused; sm all wing-rernnants.

The new genus is named to Dr. S.G. Compton, in recognition of bis important contributions to our knowledge of the fig insects.

Type-species: Comptoniella vannoorti spec. nov.

ComptonieUa vannoorti Wie bes, spec. nov. Female. Length ca. 1.8 mmo The colour is metallic black, the tibiae and tarsi are yellowish. The head (fig. b) is distinctly longer than wide across the compound eyes (0.8); these eyes are approximately as long as the cheek; there are three ocelli. The epistomal margin has a median lobe. The mandible has two teeth, each with a gland; the labial pal pus consists of one segment, the maxillary of two (1 : 4). The situation of the antennal toruli is rather low in the face, i.e., just on the imaginary line connecting the lower rim of the compound eyes. The antennal formula is 1115(3); the scape is almost three times as long as the pedicel (11 : 4). The flagellum has 1- 9 sensilla per segment, the proximal: three, the second and fourth (both shorter than the other segments): one or two, the third and fifth:

185 o

d Fig. 43. Comptoniella vannoorti spec. nov. a, male head and thorax, dorsal aspect; b, outline of female head; c, female hypopygium (h), eighth urotergite with stigma (s), ninth urotergite with pygostyle (p), and valve (v) of the ovipositor, lateral aspect, semidiagrammatical; d, male genitalia, ventral aspect. five or six, and the sixth to eighth (together forming the club): six (on the eighth segment) to ni ne (on the seventh); all segments bear long setae. There is a wide pronoturn, almost as long as the mesoscutum; the fore wing (2 : 1) is l.3 mm long, the submarginal, marginal, stigmal, and postmarginal veins are approximately in ratio 13 : 5 : 4 : 5, the disc is almost hyaline, a fringe is present on the apical third. The hind wing (4 : 1) is l.0 mm long; it has a normal fringe. The fore tarsi are approxirnately in ratio 6 : 3 : 4 : 3 : 7; the mid tarsi 10 : 6 : 5 : 4 : 6; the hind tarsi 10 : 6 : 5 : 5 : 6.

186 The hypopygium has a short spine, much like that of 0. tsamvi Wiebes (in Boucek et al., 1981, fig. 108). The e1ongation of the gaster (fig. c) is for a small part formed by the eighth urotergite (the base of the 'tail', bearing the subcircu­ lar spiracular peritremata), but mostly by the ninth (bearing the pygostyles) and the ovipositor-valves (morphologically, third valvifers), approximately in length-ratio 2 : 2 : 3.

Male. Length of the head (without mandibles) and thorax 0.6 - 0.95 mm: there is a rather large variation - for the description a robust male of 0.9 mm length was used. The colour is yellowish, the larger specimens have the head brown. The head (fig. a) is distinctly shorter than wide (ca. 0.85), and it is not at all depressed above the epistomal edge. The eyes are about as long as the cheek. The toruli are situated in the anterior-most part of the head, close together and close to the epistomal margin, which is widely incised medially. The mandible is ca. half as long as the head, and it has three apical and subapical teeth, but only one gland; there is one axial, basal tooth with a gland. The antennal for­ mula is 1115(2), the anellus is very narrow, only visible ventrally; the scape is a bit expanded, much less so than in Philosycus; the pedicel is ca. two-fifths of the length of the scape; the first funicular segment is wide and large, the fol­ lowing are much shorter; the club is divided (2 : I). The thorax (fig. a) consists of two separate terga, viz., the large pronoturn, ca. I 1/2 times as wide as long, and the fused meso-, metanotum and propodeurn, together shorter than the pronotum (2 : 3); there are small wing-remnants. The tarsi of all legs are pentamerous, with the segments of the fore tarsus approxi­ mately in ratio 3 : 2 : I : 2 : 6; the mid tarsi 2 : I : I : 2 : 4; the hind tarsi 3 : I : I : 2 : 5. The claspers of the genitalia (fig. d) bear four claws.

Type-material.- Holotype (female), nine female and twelve male paratypes, South Africa, N. Cape Augrabies Falls Nat. Pk., ex Ficus c. corrlata Thunb., leg. S.G. Compton, 5.xi.1988, no. C.75.

Derivation.- The species is named to Simon van Noort, Grahamstown who, together with Dr. Compton, gave a revision of African Otitesella. The above description is made so as to allow direct comparison with the descriptions by van Noort & Compton (1988).

OtiteseUa Westwood Westwood, Trans. ent. Soc. Lond. for 1883: 39-40 (1883); Grandi, Boll. Lab. Zool. Portici 16: 12-34 (1922); Wiebes, Tijdschr. Ent. 110: 420-421 , 436--437 (1967); van Noort & Compton, Proc. Kon. Ned. Akad. Wet. (C) 91: 425--427 (1988). There are two species-groups, viz., the 0. africana-group (all, African species, associated with figs of section Galoglychia, subsections Galoglychia and Platy­ phyllae) and the 0. digitata-group (associated with figs of section Urostigma,

187 mainly Indo-Australian; one species known from Africa, i.e., 0. serrata). Grandi (1922) discussed the comparative morphology of Otitesel/a and relatives; Wiebes (1967) provided a catalogue; Van Noort & Compton (1988) published a key to the species.

African species of O/itesella: africana Grandi - O/i/esel/a africana Grandi, 1921a: 310--311 (Guinea, ex Ficus lu/ea Vahl); Risbec, 1956: 148 (Madagascar, ex Ficus sp.). epicarioides Grandi - O/itesella epicarioides Grandi, 1921a: 310-311 (Eritrea, ex Ficus sp.). gnaphalocarpae Risbec - O/itesella gnaphalocarpae Risbec, 1951a: 332-335 (0 only, Sénégal, ex Ficus gnaphalocarpa (probably wrong, Wiebes, 1970: 10, nota)). royi Wiebes - O/itesella royi Wiebes, 1971: 378-379 (Sierra Léone, ex Ficus saussureana DC.). serrata Mayr - O/itesella serra/a Mayr, 1885: 210--211 (0 only, Socotra, ex Ficus corda/a salicifolia (VahI) Berg). This species runs to no. 4 in the key (van Noort & Compton, 1988: 427), but cannot be keyed out further. sesquianeUata van Noort - O/i/esella sesquianel/a/a van Noort in van Noort & Compton, 1988: 41~23 (South Africa, ex Ficus burll-davyi Huteh.). tsamvi Wiebes - O/itesella /samvi Wiebes in Boucek et al., 1981 : 191- 194 (Zambia, Zimbabwe, ex Ficus /honningii BI.). u1uzi Compton - O/itesella uluzi Compton in van Noort & Compton, 1988: 42~24 (South Africa, ex Ficus burtt-davyi Huteh.). spec. Wiebes - Otitesel/a spec. Wiebes, 1981 : 567 (Réunion, ex Ficus reflexa Thunb.).

Philosycus Wiebes

Wiebes, Ann. Mus. Roy. Afr. centr., in 8°, Zool. 175: 439 (1969b).

The male has a large, dilated antennal scape and an oversized head; the female is much like those of Otitesella, but the pronotum is distinctly visible from above. For only one species the host fig is known, it belongs to the Galoglychia-section Caulocarpae.

African species of Philosycus: cadenati (Risbec) - Colotreclmus cadenati Risbec, 1951a: 287- 289 (9 only, Sénégal, ex Ficus sp.). coUaris Wiebes - Philosycus col/aris Wiebes, 1969b: 440--443 (Ivory Coast, ex Ficus sp.). monstruosus (Grandi) - Sycobiella monstruosa Grandi, 1921 a: 311-312 (0 only, Guinea); Risbec, 1951a: 325-336 (0 only, Sénégal, ex Ficus sp.); Grandi, 1952: 45 (0 only, Sénégal, ex Ficus ova/a Vahl).

Subfamily EPICHRYSOMALLINAE This group has wandered a lot, from Torymidae to Pteromabdae (Boucek m Boucek et al., 1981: 165-166) and recently (Boucek, 1988), Agaonidae.

Camarothorax Mayr (fig. 44) Mayr, Wien. ent. Ztg. 25: 182- 183 (1906); Boucek in Boucek et al., Tijdschr. Ent. 124: 194, 196 (1981).

188 Figs. 44 (left), 45 (middle) and 46 (right). Males of Camarothorax hemimucro Wiebes (after Wiebes, 1981, fig. 64), OdontoJroggatia galili Wiebes (after Wiebes, 1980, fig. I) and Sycophila hilla Wats­ ham (after Watsham, 1977, fig. 6).

This is what formerly was known in Africa as Sycobia. Boucek (in Boucek et al., 1981: 163) published a key to three species, and compared two others (p. 200, 203). Boucek (1988: 170) suggested that the genus, originally described from the Indo-Australian region, should be more restricted than in his treatment of 1981: the African species may not belong. There are three host figs known, belonging to the subsections Galoglychia and Platyphyllae.

African species of Camarothorax: africanus (Wiebes) - Sycobia aJricana Wiebes, 1971: 381-382 (Sierra Léone, ex Ficus saussureana De.). brevimucro Boucek - Camarothorax brevimucro Boucek in Boucek et al., 1981: 202-204 (Zim­ babwe, South Africa, ex Ficus thonningii BI.); Wiebes, 1981: 569 (Réunion, ex Ficus reflexa Thunb.).

189 equicollis Boucek - Camarolhorax equicollis Boueek in Boucek et al., 1981: 197- 200 (Uganda, Zimbabwe, South Africa, ex Ficus Ihonningii BI.). heminlUcro Wiebes - Camarolhorax hemimucro Wiebes, 1981: 567-569 (Réunion, ex Ficus rejlexa Thunb.). imerinell'iis (Risbec) - Cal/imomus imerinensis Risbec, 1956: 153 (Madagascar). longimucro Boucek - Camarolhorax longimucro Boucek in Boucek et al., 1981 : 20 1-203 (Zim­ babwe, ex Ficus thonningii BI.). orientalis (Wiebes) - Sycobia orientalis Wiebes, 1975: 234 (A1dabra Isl., ex Ficus rejlexa Thunb.).

Odontofroggatia Ishii (fig. 45)

Ishii, Kontyu 8: 95 (1934); Wiebes, Zool. Meded. Leiden 56: 1-6 (1980).

As the host fig of all known species, i.e., Ficus microcarpa Linn.f., was intro­ duced into Africa and is expected to spread, the genus is included in this list. One species is known from Israel: galili Wiebes - Odontofroggatia galili Wiebes, 1980: 3- 5 (Israe1, ex Ficus microcarpa Linn.f.).

Sycotetra Boucek

Boucek in Boucek et al. , Tijdschr. Ent. 124: 195-196 (1981).

African species of Sycoletra: serricornis Boucek - Sycotetra serricomis Boucek in Boueek et al., 1981: 196-197 (Zimbabwe, ex Ficus thonningii BI.).

Family EURYTOMIDAE Ten years ago, Boucek (in Boucek et al., 1981: 204-205) wrote: 'Figs [-] rep re sent an ecological niche virtually identical to many plant galls and as such attract many species of gall-inhabiting Eurytomidae'. He discussed four genera, and keyed out (on p. 161-163) alltogether ten species, which are listed below.

Eurytoma Illiger

Illiger, Mag. f. Insectenkunde 6: 192 (1807); Boucek in Boucek et al., Tijdschr. Ent. 124: 205 (1981).

African species of Eurytoma (associated with figs): ficusgallae Boueek - Eurytoma ficusgal/ae Boucek in Boucek et al. , 1981: 205-208 (Zimbabwe, ex Ficus thonningii BI.).

Ficomila Boucek

Boucek in Boucek et al, Tijdschr. Ent. 124: 211 - 212 (1981).

African species of Ficomila: curtivena Boueek - Ficomila curlivena Boucek in Boucek et al., 1981 : 212- 213 (2 only, Zimbabwe & South West Africa, ex Ficus thonningii BI.).

190 gambielL'iis (Risbec) - Deca/oma gambiense Risbec, 1955a: 561-564 (Gambia); Boucek in Boueek et ai., 1981 : 213-214 (Gambia, ex Ficus sycomorus L.; Zimbabwe, South Africa, ex Ficus /honningii BI.).

Syceurytoma Boucek

Bouéek in Boucek et al., Tijdschr. Ent. 124: 208- 109 (1981).

African species of Syceury/oma: fic~ Boueek - Syceury/oma ficus Boueek in Boucek et al., 198\: 209-211 (Uganda, ex stem galls on Ficus sycomorus L.; Zimbabwe, ex Ficus /honningii BI.; South Africa).

Sycophila Walker (fig. 46)

Walker, Notes Chalc.: 63 (1871); Boucek in Bouéek et al., Tijdschr. Ent. 124: 214--215 (1981).

Except for those reared from figs, listed below, Boucek listed eight other African species (p. 214-215).

African species of Sycophila (associated with figs): ftaviclava Boucek - Sycophila jiavic/ava Boueek in Bouéek et al., 1981 : 222-223 (Zimbabwe, ex Ficus /honningii BI.). kestraneura (Masi) - Decatoma kestraneura Masi, 1917: 138-139 ('i' on1y, Seychelles); Bouéek in Boucek et al., 1981: 215-216 (Uganda, ex Ficus sp.; Zimbabwe, South West Africa, ex Ficus thonningii BI.; South Africa). modesta Bouéek - Sycophila modes/a Bouéek in Boucek et ai., 1981: 218-220 ('i' on1y, Zimbabwe, ex Ficus thonningii BI.). naso Boucek - Sycophila naso Boucek in Bouéek et al., 1981: 220--222 (Uganda, ex stem galls on Ficus sycomorus L.; Zimbabwe, ex galls on Ficus thonningii BI.; South West Africa, ex Ficus thonningii BI.). punctum Boucek - Sycophila punctum Boucek in Bouéek et ai., 1981: 216-218 (Zimbabe, ex Ficus /honningii BI.). sessilis Boueek - Sycophila sessilis Boucek in Boucek et al ., 1981: 218 ('i' on1y, Zimbabwe, ex Ficus thonningii BI.; Sou th Africa).

Farnily ORMYRIDAE

Ormyrus Westwood (fig. 47)

Westwood, Phil. Mag. (3) I: 127 (1832); Bouéek in Boucek et ai., Tijdschr. Ent. 124: 223-224 (1981).

Three species of Ormyrus were rea red from figs in Africa: they were described and keyed out (p.160-161) by Boucek (in Boucek et al., 1981), who also listed eight other African species (p. 224).

African species of Ormyrus (associated with figs): ftavipes Bouéek - Ormyrus jiavipes Bouéek in Boucek et al., 1981: 226-228 (Zimbabe, ex Ficus thonningii BI.).

191 Fig. 47. Female of Ormyrus watshami Boucek, after Boueek (in Boucek et al. , 1981 , fig . 157).

subconicus Boucek - OrmynJs subconicus Boucek in Boueek et al., 1981: 228- 229 (Zimbabwe, ex Ficus thonningii BI.). watshami Boucek - Ormyrus watshami Boucek in Boueek et al., 1981 : 224--226 (Zimbabwe, ex galls on Ficus thonningii BI.).

REFERENCES (only those not mentioned in the bibliography) Ashmead, W.H. - Classification of the Chalcid Bies or the superfamily Chalcidoidea, with descrip­ tions of new species in the Carnegie museum, collected in South America by Herbert H. Smith. Mem. Carnegie Mus. I, I-x, 225-551 (1904a). Ashmead, W.H. - New generic names in the Chalcidoidea. Proc. ent. Soc. Wash. 6, 126 (1904b). Boueek, Z. - Australasian Chalcidoidea (Hymenoptera). c.A.B. International, Wellingford, V .K ., pp. 1- 832 (1988). Corner, E.J.H. - Check-list of Ficus in Asia and Australasia, with keys to identification. Gdns' Bull. Singapore 21, 1- 186 (1965). Förster, A. - Kleine Monographien parasitischer Hymenopteren. Verh. naturh. Ver. preuss. Rheinl. 35, 42-84 (1878). Galil, J. - An ancient technique for ripening sycomore fruit in East-Mediterranean countries. Econ. Botany 22, 178- 190 (1968). Galil, J., R. Dulberger & D. Rosen - The efTects of Sycophaga sycomori L. on the structure and development of the synconia of Ficus sycomorus L. New Phytol. 69, 103-111 (1970). Godfray, H.C.J. - Virginity in haplodiploid populations: a study on fig wasps. Ecol. Entom. 13, 183- 291 (1988). Gordh, G. - The comparative external morphology and systematics of the neotropical parasitic fig wasp genus [dames (Hymenoptera: Torymidae). Univ. Kansas Bull. 50, 389-455 (1975). IlIiger, K. - Vergleichung der Gattungen der Hautflügler Piezata Fabr. Mag. Insektenkunde 6, 189- 199 (1807).

192 Ishü, T. - Fig chalcidoids of Japan. Kontyu 8, 84-100 (1984). Joseph, K.J. - Contributions to our knowledge of fig insects (Cha1cidoidea: parasitic Hymenop­ tera) from India vii. Descriptions of fifteen new and records of some old species. Ann. Soc. ent. France 125, 97-133 (1957). Walker, F. - Descriptions of Chalcidites discovered in St. Vincent's Isle by the Rev. Lansdown Guilding. Ann. Mag. nat. Hist. 12, 46-49 (1843). Walker, F. - Notes on Chalcidiae. Chalcididae, Leucospidae, Agaonidae, Eucharitidae, Perilampi­ dae, Ormyridae, Encyrtidae: 55-70 (1871). Westwood, J.O. - Description of several new British forms among the parasitic hymenopterous insects. Lond. Edinb. Dubl. Phil. Mag. (3) 1, 127- 129 (1832). Wiebes, J.T. - Bomean fig wasps from Ficus stupenda Mique1 (Hymenoptera, Chalcidoidea). Tijd­ schr. Ent. 109, 163--192 (1966b). Wiebes, J.T. - The genus Odontofroggatia Ishü (Hymenoptera, Pterornalidae, Epichrysomallinae). Zool. Meded. Leiden 56, l...{i (1980). Wiebes, J.T. - Fig wasps (Hymenoptera). In: J.L. Gressitt (ed.), Biogeography and ecology of New Guinea. Monogr. Biol. (w. Junk) 42, 735-755 (1982a).

193 v. The pollinators (Agaoninae)

Diagnosis

The Agaoninae can he shortly characterized as follows. The female mandibles are situated undemeath the head or slightly in front of it; the mandible has an appendage bearing transverse lamellae, or rows of fine teeth or small crenulations. The head is often longer than wide across the com­ pound eyes; mostly there are three ocelli. The face medially has a broad channel or depression, in which the antennal scapes may fit. The antennal scape is dis­ tinctly widened; the third antennal segment is produced apically into a point. In many species, the mesostemum has pollen pockets; the fore coxa may have a 'corbicula'. The fore tibia bears a dorso-apical comb of teeth; the bind tibia has axial and antiaxial teeth; all tarsi are pentamerous. The male is apterous. The eyes are small or vestigial; ocelli are absent. The antennae are situated in separate grooves on either si de of a medial prorninence, or in a common groove in the frontal part of the head; the number of segments is reduced. The legs have shortened spiny tibiae; the tarsal segments are often reduced. The gaster ends in a tube with the genitalia, often bearing small claspers with claws. The colour is yellowish.

Introduced species Two extra-limital species are included, because they were introduced or are known to occur at the borders of the continent, viz., Parapristina verticillata (Waters ton) (no. 20), recently found in Morocco (G. Michaloud, 1989, personal communication) and on the isle of Malta (M.A. Thake, 1990, personal com­ munication), and Platyscapa quadraticeps (Mayr) (no. 25), which for some dec­ ades already, is known to occur in Israel (Galil & Eisikowitch, 1968c). The host figs, Ficus microcarpa Linn. f. and F religiosa L., respectively, are common lane or park trees and the wasps - for some reason or another - seem to be easily dispersed. McKey (1989: 664--668) recently discussed the conditions of human­ aided range extensions of fig-insects, and listed many in stances; he could not yet know, of course, of our African P verticillata. See also Ramirez & Montero (1988).

195 Classification For the Agaoninae, as well as for Ficus, there is agreement over their mono­ phyly, although Corner (1978: 405) speculated on the parallel evolution of the groups recognized within Ficus, from some pre-Ficus not pollinated by fig wasps. The pollinators can be divided into two subgroups (Wiebes, 1973), mainly char­ acterized by features of the female; the males are more difficult to divide. Al­ though the genera seem well-defined, especially when the morphology of both sexes is taken into consideration, straight key-characters (without exceptions) are difficult to find. Notabie exceptions are E/isabethiella articulata (no. 31) and Agaon megalopon (no. 58), the females of which have two ocelli instead of three (Elisabethiella) and zero (Agaon), respectively. The two key out with most spe­ cies of Alfonsiella, from which they difTer jn the rod-like sensilla of the antenna (E. articulata, vs. long and flexible sensilJa in Alfonsiella and Agaon) and in the funicular segments of the antenna being more than twice as long as wide (A. megalopon, vs. subquadrate in Alfonsiella and Elisabethiella). Nigeriella fus­ ciceps (no. 46) has sensilla chaetica as well as sensilla linearia in the female antennal funicle, while the other species of Nigeriella have only sensilla linearia. In the males, Platyscapa soraria (no. 27) is exceptional in the genus in th at all dorsal sclerites of the thorax are separate, but it is recognizable from Do/ichoris flabella ta (no. 19), with which it keys out, by the nurnber of antennal anelli (one or two, vs. three). If the exceptions are left out, the key is rather simpie.

KEY TO THE GENERA (figs. 48, 49)

I. Females ...... 2 - Males ...... 15 2. Mandibular appendage fused witb tbe body of tbe mandible; it bears ventral lamellae (fig. 48, a). Third antennal segment witb aseparation between tbe main part and tbe produced apex (fig. 48 , b: 3a and 3b), whicb itself may be divided into two parts (3c); tbe pedicel (2) with axial spines (B1astophagini) ...... 3 - Mandibular appendage truly appended to tbe mandible (fig. 49, b); in most species it bears ventral rows of fine teeth or small crenulations. Third antennal segment more simple (fig. 49, a): the produced apex, if at all prominent, not separate from the main part of the segment; tbe pedicel without axial spines (Agaonini)...... 8

~ Fig. 48. B1astophagini - a, P/alyscapa bergi Wiebes, female mandible and appendage in ventral view, after Wiebes (1986b, fig. 40); b, B/aslophaga psenes (L.), female antenna in axial view: apex of scape (I), pedicel (2), third segment (3; a, proxirnal, b, medial, and c, distal portion) and fourth segment (4), after Grandi (1929, fig. iv, 2); c-h, dorsal aspect of male bead and thorax (except for c and g, outline only): c, B/aslophaga psenes, composed after Grandi (1929, figs . xxviii, I and xix, 3 and 5); d, Kradibia cowani Saunders, after Wiebes (1981, fig . 11); e, Ceratoso/en capensis Grandi, composed after Grandi (1955, figs. iii, I and 4); f, Do/ichoris fiabellala Wiebes, after Wiebes (1979a, fig. 8); g, Paraprislina verlici//ala (Waterston), composed from Grandi (1927, figs. vii, 3 and 6); b, Platyscapa etiennei Wiebes, after Wiebes (1981, fig. 8).

196 a

c

. . , ",. \' .. . { I' ====~= ! .. ,.-._ _....

J . !~cf.4 ... ~( . I

197 -7 Fig. 49. Agaonini - a, Alfonsiella brongersmai Wiebes, proximal segments of female antenna, after Wiebes (1972b, fig . 14); b, Elisabethiel/a soeotrensis (Mayr), female mandible and appendage in ventral view, after Wiebes, 1989a, fig. 27); c-i, dorsal aspect of male head and thorax (except for c and h, outline only): c, Allotriozoon prodigiosum Grandi, composed after Grandi (1916a, figs . xix, I and xx, 2); d, Elisabethiel/a reflexa Wiebes, after Wiebes, 1981 , fig. 9); e, Nigeriel/a avieola Wiebes, after Wiebes (1981 , fig. 10); f, Alfonsiella brongersmai, after Wiebes (1972b, fig. 8); g, Paragaon josephi Wiebes, after Wiebes (1986b, fig. 15); h, Agaon spatula/um Wiebes (a1so part of abdominal tergum), partly after Wiebes (1989c, fig. 12); i, CourtelIa wardi Compton, after Wiebes & Compton (1990, fig. 27).

3. Spiracular peritremata of the eighth urotergite large and gouge-Iike, with distinct la- teral extensions ...... Cera/osolen - Spiracular peritremata of the eighth urotergite smalI, subcircular...... 4 4. Mesosternum with pollen pockets .... . 5 - Mesosternum without pollen pockets . .. . . Blastophaga psenes (no. I) 5. Fore tibia with two or three teeth in the dorso-apical combo Fore coxa with a corbicula 6 - Fore tibia with four to six teeth in the dorso-apical comb (figs. 52, e and 1). Fore coxa without a corbicula ...... Kradibia 6. Antenna consisting of ten segments ànd the maxilla with a bacilliform process (as in figs. 52, g and 54, g) ...... Doliehoris flabella/a (no. 19) - Antenna consisting of eleven segments, occasionally ten, but then the max.illa without a bacilliform process ...... 7

7. Antennal segments, from the fifth onwards, more and more cup-shaped, bearing one row of long sensilla chaetica th at project over the apical edge of their segment for their total length ...... Parapristina vertieillata (no. 20) - If the sensilla of the eighth to e!eventh segments strongly project over the apex of the segment, those of the fifth to seventh segments do not project for more than a third or half of their length ...... Platyseapa

8. Three ocelli present ...... 9 - Two (lateral) ocelli present ...... Alfonsiella* (pars) - No ocelli ...... \3 9. Funicular segments of antenna with sensilla linearia ...... 10 - Funicular segments of antenna with both sensilla linearia and chaetica . · ...... Nigeriella fuseieeps (no. 46) - Funicular segments of antenna with sensilla chaetica ...... 11 10. Mandibular appendage with ridges, laterally produced into teeth . · ...... Nigeriella (pars: nos. 45, 47, 48) - Mandibular appendage with rows of many fine teeth or denticulations ...... · ...... Elisabethiella (pars: nos. 33, 34, 38-44)

* Antennal funicular segments two or more times as wide as long. Antennal sensilla long and ftexible ...... Agaon megalopon (no. 58) Antennal funicular segments subquadrate. Antennal sensilla rod-like ...... Elisabethiel/a artieulata (no. 31) Antennal funicular segments subquadrate. Antennal sensilla long and ftexible ...... Alfonsiella (pars: nos. 49, 50, 53- 55)

198 , , ~' "

199 11. Mandible unidentate ...... Paragaon - Mandible witb two or more teeth...... 12

12. Eye half as long as the cheek, or shorter ...... CourtelIa - Eye (mostly much) more than half as long as tbe cheek ...... · ...... Elisabethiella (pars: 32, 35-37)

13. FunicuJar segments of antenna with sensiUa linearia ...... Allotriozoon - FunicuJar segments of antenna with sensiUa chaetica ...... 14

14. Funicular antennal segments more than twice as long as wide. Postmarginal vein of fore wing longer than stigmal ...... Agaon - Funicular antennal segments subquadrate. Postmarginal vein of fore wing half as long as stigmal ...... Alfonsiella (pars: nos. 51 , 52) IS. All tarsi, incJuding the fore tarsi, essentially pentamerous (wbich may be seen in the dorsal subdivisions) ...... 16 - F ore tarsi bi- or trimerous ...... 19

16. All thoracic terga distinctly separate (fig. 48, e) ...... Ceratosolen - Mesonoturn, metanoturn, and often (dorsally) also the propodeum, fused ...... 17

17. Antennae situated in a common, anterior groove (fig. 49, d) ...... Elisabethiella - Antennae situated in separate sockets...... 18

18. Head pear-shaped (fig. 49, f) ...... Alfonsiella - Head more subquadrate (fig. 49, e) ...... Nigeriella 19. Also the rnid and bind tarsi oligomerous . 20 - Mid and hind tarsi pentamerous ...... 24 20. Mid leg reduced, at most with three tarsal segments. Thoracic terga separate (fig. 48, d) ...... Kradibia - Mid leg not reduced; the rnid and bind tarsi tetramerous. Tboracical terga fused or free 21

21. All thoracical terga separate (fig. 49, c) ...... Allotriozoon - At least the meso- and metanotum fused dorsally . 22 22. Propodeum for tbe greater part dorsally separate ...... 23 - Also the propodeum dorsally fused with the meso- and metanotum (fig. 48, g) . · ...... Parapristina verticillata (no. 20) 23. Antennae, separately, borne in an anterior groove, in between lateral elevations. An- tenna with two or three anelli. Head and thorax, fig . 49, h ...... Agaon - Antennal sockets widely separate, the head without lateral elevations. Antenna with one anellus. Head and thorax, fig. 49, g ...... Paragaon josephi (no. 57) 24. Metanotum dorsally visible as two separate, sublateral sclerites (fig. 48, c) ...... · ...... Blastophaga psenes (no. I) - Metanotum, if separate dorsally, not divided ...... 25 25. Metanotum and propodeum separate dorsally (fig. 48, f)* ...... · ...... Do/ichoris flabella ta (no. 19) - Metanoturn and propodeurn for the greater part fused dorsally ...... 26

* Tbe fore tibia has three dorsaI teeth. Platyscapa soraria (no. 27), exceptional in that all thoracical terga are separate, has one or indistinctly two teeth in the dorso-apical comb of the fore tibia.

200 26. Head subquadrate or wider than long, also the pronotum transverse (fig. 48, h) ...... Pla/yscapa - Head oblong, also the pronotum longer than wide (fig. 49, i) ...... Cour/elia

AFRICAN GENERA AND SPECIES Blastophaga Gravenhorst (fig. 50)

Gravenhorst, Uebers. Arb. Veränd. Schles. Ges. vaterl. Cult. iJ. (1826): 23- 24 (1827, Blas/opha­ gus); Gravenhorst, Beitr. Ent., Breslau 1: 27-33 (1829, BlaslOphaga, validated 1964 by ICZN Opinion 694: China, 1964).

There is only one African species known, i.e., B. psenes, from figs of the sub­ genus Ficus, section Ficus. Many more species occur in Indo-Australian region, and also these are associated with figs of the section Ficus.

Fig. 50. Blas/ophaga psenes (L.), female and male, after Grandi (1929, figs. i and xxxi, I, respec­ tively).

201 1. Blastophaga psenes (Linnaeus)

Linnaeus, Syst. Nat., edit. lOa, I: 554, no. 13 (1758, Cynips); Grandi, Boll. Lab. Ent. Bologna 2: 1- 147 (1929).

The female head is shorter than wide across the compound eyes (0.9), and the eye is a bit over two times as long as the cheek. There are three ocelli. The antenna has eleven segments, the fifth to tenth of which bear one row of long sensilla linearia; those on the ten th are shorter. The mandible is bidentate at the apex, and it bears an appendage with five or six ventral lamellae. The thorax lacks pollen pockets and there are no coxa I corbiculae. The veins of the fore wing are complete, the marginal, stigmal and postmarginal veins are subequal in length. The fore tibia has bidentate dorsal and ventral complexes of teeth. The spiracular peritremata of the eighth urotergite are smalI, subcir­ cular. The total length (head, thorax and gaster) is ca. 1.8 mm; the ovipositor valves are about one-quarter of the length of the gaster. The colour is dark brown.

The male head is wider than long (1.1); the eye is as long as the cheek, and nearly one-fifth of the length of the head. The head bears many small spines, directed backward. The antennae are situated in a common anterior groove, and they have five distinct segments: the scape is as long as the combined lengths of the pedicel and the two anelli, and it is approximately as long as the club. The thorax has four distinct dorsal sclerites, viz. , the pronotum, wider than long (1.3); the mesonotum, about two-thirds of the length of the pronotum; the metanotum, visible as two lateral parts; and the propodeurn, as large as the mesonotum. The propodeal spiracles are smalI, circular, occupying about one­ third of the lateral margin of the propodeum. The fore tibia has a tridentate dorsal crest and two ventral teeth; the tarsus is bimerous; the mid and hind tarsi have five segments. The genitalia are simp Ie. The length of the head and thorax is ca. 1 mmo The colour is yellow-brown.

The host fig in Africa is Ficus pa/mala Forssk. (Ethiopia). In the Mediterranean region and in Lesser Asia, B. psenes is known from F carica L. , as also in India. Joseph (1954: 401-408) described B. vaidi Joseph from F pa/mala in India, but the differential characters are not clear (Wiebes & Compton, 1990: 206). In the key by Mani (1989: 30) the length of the (female) bind metatarsus relative to that of the following (two) segments is used to differentiate B. psenes and B. vaidi. For the one the metatarsus is said to be 'equal to 2.5 of following segments combined', and for the other 'equal to 2 or less than 2 following segments com­ bined', which is in contradiction with Joseph (1953: 404, B. vaidl) 'metatarsus longer than the two following segments combined' (as also repeated by Mani himself, in the description on p. 35). Waterston (1920a: 128) recorded B. psenes from the introduced F carica in Transvaal; Wiebes (in Aweke, 1979: 100) re­ corded B. vaidi from Ficus pa/mala in Etbiopia.

202 Galil & Neeman (1977) studied the pollen transport in B. psenes from F carica in Israel: it is carried in intersegmental and pleural invaginations in the gaster; pollination is passive.

Kradibia Saunders (fig. 51) Saunders, Trans. ent. Soc. Lond. (1883): 20-25 (1883); Wiebes, Zool. Meded. Leiden 53: 165--180 (1978a); Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 206-209 (1990).

The female head is shorter than wide across the compound eyes (0.8--0.9); the eye is mostly two times as long as the cheek or longer (one exception). There are three ocelli. The antenna has ten segments, from the sixth onwards with one to three rows of long sensilla linearia. The mandible is bidentate at the apex, and it bears an appendage with four to seven ventral lamellae. In some species the maxilla bears a bacilliform process.

Fig. 51. Kradibia cowani Saunders, female and male, after Saunders (1883, pI. üi figs . 33 and 32, respectively).

203 The thorax has pollen pockets, but the fore coxae have no corbicula. "(he postmarginal vein of the fore wing is longer than the stigmal vein, even if only little. The fore tibia has four to six dorso-apical teeth. The spiracular peritremata of the eighth urotergite are very smalI, subcircular. The totallength (head, thorax and gaster) is ca. 1.5 rnm (there is little variation between the species); the ovipositor valves are about one-quarter of the length of the gaster. The colour is dark brown.

The male head is wider than long, if only little in some species; the eye is over twice as long as the cheek, and one-fifth to one quarter of the length of the head. The head has small spines, directed backward. The antennae are situated in a shallow common anterior groove, and they have five distinct segments, viz., the scape, the pedicel, a (sometimes very short) anellus, the funicle and the club. The thorax has four distinct dorsal sclerites, viz., the pronotum, approxi­ mately as long as the other scIerites combined (but in one species, distinctly shorter); the mesonotum, rather short; the metanotum, in most species visible as two lateral parts, but these may be connected in the rniddle; and the pro­ podeum, in general twice as wide as long. The propodeal spiracles occupy almost the entire lateral margin of the propodeum. The fore tibia has a comb of eight unequal dorso-apical teeth and two ventrais; the tarsus is bimerous (trimerous in one species). The mid leg is reduced in most species, but more complete in others (and then the tarsus is trimerous); the hind tarsus is pentamerous. The genitalia have claspers with two to four claws. The length of the head and thorax is I mm, or somewhat less, but the species vary little. The colour is yellow-brown.

There are five African species known, next to a number of Indo-Australian forms; the host figs belong to the section Sycidium.

KEY TO THE SPECIES (fig. 52)

I. Female antennal segments, from the sixth onwards, three times as long as wide, with three rows of sensiUa. Male fore tarsus three-segmented; the mid leg is rather complete (as in fig. a). Hind tibia (fig. c) with some spines a10ng the dorsal margin . . 3. K. hilli . Female antennal segments shorter (fig. d), at most two times as long as wide. Male fore tarsus two·segmented ...... 2 2. Female fore tibia with a comb of six teeth (fig. e), a1temately large and smaU. Male mid leg (fig. b) reduced; the hind tibia with a dorsaI row of spines ...... 3 - Female fore tibia with a comb of five unequal teeth (fig. 1), or four (some times with a small auxilliary). Male mid leg either not reduced, or hind tibia without dorsaI spines 4 3. Female mandibular appendage with four ventral lamellae (fig. g), the maxilla with a bacilliform process. Compound eye twice as long as the cheek (fig. h) . The male cannot he distinguished from the following species ...... 4. K. cowani • Female mandibular appendage with six ventral lameUae; the maxilla without abaciUi· form process. Compound eye more than twice as long as the cheek . . . 5. K. saundersi

204 Fig. 52. Details of Kradibia. a, K. gestroi (Grandi), male mid leg, after Wiebes (1969c, fig. 30); b, K. cowani Saunders, male mid leg, after Wiebes (l978a, fig. 18); c, K. hilli Wiebes, male hind leg, after Wiebes (1978a, fig. 31); d, K. cowani, third to seventh segments of fernale antenna, after Wiebes (I 978a, fig. 23); e-f, comb of female fore tibia: e, K. saundersi Wiebes, and f, K. etiennei Wiebes, after Wiebes & Compton (1990, figs. 2 and 3, respectively); g- h, K. cowani: g, female mouth-parts, and h, female head, after Wiebes (1978a, figs. 24 and 25, respectively).

4. Female compound eye distinctly shorter than twiee the length of the cheek. Maxilla with a bacilliform process. Fore tibia with a comb of five unequal teeth. Male mid leg reduced; the hind tibia without distinct dorsal spines along the dorsal margin ...... 6. K. etiennei - Female compound eye over two times as long as the cheek. Maxilla without a bacilli­ form process. Fore tibia with a comb of four teeth, sometimes with an auxilliary. Male mid leg slender, but rather complete; the hind tibia with some dorsaI spines ...... 2. K. gestroi afrum

2. Kradibia gestroi afrum (Wiebes)

Wiebes, Ann. Mus. Roy. Afr. eentr., in 8°, Zool. 175: 460 (1969c, Liporrhopalum gestroi afrum); Wiebes, Zool. Meded. Leiden 53: 176-177 (1978a).

The female compound eye is rather long relative to the cheek (2.4). The antennal segments have one irregular row of sensilla. The mandibular appendage has four ventral lamellae; the maxilla does not bear a bacilliform process. The fore tibia has a dorso-apical comb of four teeth, and also an auxilliary may be present.

205 The antenna of the male has a very short anellus. The fore tarsus is bimerous; the mid leg is rather complete; the hind tibia has spines along the dorsal margin. The genital claspers bear four claws.

The host fig is Ficus exasperata Vahl (Ivory Coast, Ghana, Zimbabwe), from which fig it was described in Ceylon by Grandi (1916a, b). The African form is slightly different from the Singhalese one, and it was distinguished as subspecies afrum. The species was also recorded from Sénégal (host tig unknown) and from Ficus asperifolia Miq. (Ghana, Togo, Cameroun, Gabon; see Verkerke, 1987) and from Ficus capreifolia Delile (Ethiopia). The population dynamics of K. gestroi in Indian F exasperata were studied by Balakrishnan Nair & Abdurahiman (1984).

3. Kradibia hilli Wiebes Wiebes, Zool. Meded. Leiden 53: 177- 179 (1978a).

The female compound eye is distinctly longer than twice the length of the cheek (2.3). The antennal segments are th ree times as long as wide, and they bear three rows of long sensilla. The mandibular appendage has five large and two smaller transverse lamellae; the maxilla bears a short bacilliform process. The fore tibia has six dorso-apical teeth.

The male eye is rather large, and the antennal anellus is long compared to that of K. gestroi, i.e., one-tifth of the funicular segment. The fore tarsus is trimerous; the mid leg is rather complete; the hind tibia has some spines along the dorsal margin. The genital claspers bear two claws.

The name of the host fig was given as Ficus urceolaris Riem. (which is a form of F asperifolia) (Uganda). The species was also recorded from Tan­ ganyika.

4. Kradibia cowani Saunders Saunders, Trans. ent. Soc. Lond. (1883): 20-25 (1883); Wiebes, Zool. Meded. Leiden 53: 173- 176 (1978a, pro parte).

The female compound eye is twice as long as the cheek. The antennal segments have two rows of long sensilla. The mandibular appendage has four distinct ventral lamellae; the maxilla be ars a bacilliform process. The fore tibia has a dorso-apical comb of six teeth, altemately large and small.

The male is much like that of K. gestroi, but the mid leg is reduced to two short segments. The hind tibia bears some spines along the dorsal margin. The genital claspers bear three claws.

206 The host fig is Ficus brachyclada Baker (Madagascar).

5. Kradibia saundersi Wiebes

Wiebes in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 208 (1990).

The species is much like K. cowani, but the female compound eye is longer relative to the cheek (2.3) and the mandibular appendage has six ventral lamel­ lae. The maxilla has no bacilliform process.

The host fig is Ficus politaria Lam. (Madagascar).

6. Kradibia etiennei Wiebes

Wiebes in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 208-209 (1990).

The female compound eye is distinctly shorter than twice the length of the cheek (1.75). The mandibular appendage has six ventral lamellae, as in K. saundersi, and the maxilla has a bacilliform process, as in K. cowani. The fore tibia has a comb of five unequal teeth.

The male mid leg is reduced; the hind tibia has no distinct spines along the dorsal margin. The genital claspers bear three claws.

The host fig is Ficus lateriflora Vahl (Réunion).

Ceratosolen Mayr (fig. 53) Coquerel, Rev. Mag. Zool. (2) 7: 367, 422 (1855, Sycocrypta); Mayr, Verh . zool.-bot. Ges. Wien 35: 164 (1885, Ceratoso/en, subgenus of B/astophaga; validated 1963 by ICZN Opinion 682: China, 1963); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 251-266 (1989b).

The female head is in most species not quite as long as wide across the compound eyes (0.85-0.95), in a few species as long as wide, or a bit longer (1.05- 1.10); the eye is approximately equal in length to the cheek (1 - 1.2), or distinctly longer: ca. two times (1 .85- 2.2), or up to three or five times and then strongly buIging. There are three ocelli. The last two antennal segments (nos. 10-11), in four species the last three (nos. 9- 11), are fused, and from the sixth or seventh onwards they bear one or two rows of sensilla linearia, in two species three; in one (other) species these segments are ftabellate. The mandible is bidentate at the apex, and it bears an appendage with four to eight ventral lamellae. In most species, the maxilla bears a bacilliform process. The thorax has large pollen pockets, but the fore coxae have no corbicula. The postmarginal vein of the fore wing is distinctly longer than the stigmal, in most species approximately two times as long, but in two species these veins are sub­ equal in length. The fore tibia has four or five dorso-apical teeth, but in two

207 Fig. 53. Ceratosolen coecus (Coquerel), female and male, after Wiebes (1981, figs . 5 and 3, respec­ tively.

species there ase three. The spiracuJar peritremata of the eighth urotergite are large and elongate, gouge-like. Tbe total length (head, thorax and gaster) is ca. 2 mm, a few species are smaller (1.5 mm) or larger (up to almost 3 mm); the ovipositor valves are approx­ imately as long as the gaster or a bit longer (1 - 1.1), in a few shorter (0.7) or distinctly longer (1.3-1.5).

The male head is longer than wide (1.3-1.5); the eye is small in most species, approximately as long as the cheek, but in four species the males are blind. The antennae are situated in grooves at either side of a trilobate process of the epi­ stomal margin; they have four or five segments: the anellus may he lacking (and then there are only four segments), or it may he disc-like (fig. h), subquadrate (fig. i), or this segment may he longer than wide, even up to twice as long as wide (fig. j).

208 The thO.rax mO.stly has a IO.ng prO.nO.tum (i.e., as IO.ng as the cO.mbined lengths O.f mesQ.- and metanO.tum), a transverse mesO.nO.tum; the metanO.tum is incO.m­ pletely separated frO.m the prO.pO.deum, which may be acute O.r blunt at the apex, and bears large, sublateral spiracular peritremata. The fO.re tibia bears a dO.rsal cO.mb O.f three teeth; all tarsi are pentamerous. In many species the genitalia bear claspers with fO.ur tO. many claws; in fO.ur species there are nO. claspers. The length O.f the head and thO.rax in mO.st species is ca. 1.5 mmo The cO.lO.ur is yellO.w-brown.

There are twelve African species knO.wn; the hO.st figs belO.ng tO. the subgenus Sycomorus. IndO.-Australian representatives O.f this group O.f Ceratosolen are assO.ciated with a Sycomorus-fig and with figs O.f sectiO.n Neomorphe.

KEY TO THE SPECIES (females; fO.r the males see table 8) (fig. 54)

I. Last three segments of the female antenna (nos. 9-11) fused, or shaped so as to form a distinct club (fig. a). Antennal segments, from the sixth onwards, with one row of sensilla ...... 2 - Only the last two segments of the female antenna (nos. 10 & 11) form a distinct club (fig. c). Antennal segments with two or three rows of sensilla ...... 5 2. Valves of the female ovipositor distinctly shorter than the gaster (0.7) ...... 10. C. acutatus - Valves of the female ovipositor longer than the gaster, if only little (1.1 - 1.5) ...... 3

3. Female head large, longer than wide across the compound eyes (1.15); the eye distinctly shorter than the cheek (0.85). Mandibular appendage with six to eight ventral lamellae ...... 18. C. megacephalus - Female head smaller, approximately as long as wide across the compound eyes (0.95- 1.10; fig. b); the eye slightly (1.1) or distinctly (2.2) longer than the cheek. Mandibular appendage with four to six ventral lamellae ...... 4

4. Female compound eye slightly longer than the cheek (1.1). Mandibular appendage with six ventral lamellae. Fore tibia with five teeth in the dorso-apical combo Hypopygium with sclerotized lateral patches at either side of the base of the spine, which bears no hyaline setae (fig. I) ...... 14. C. galili - Female compound eye more than twice as long as the cheek (2.2). Mandibular ap­ pendage with four or five ventrallamellae. Fore tibia with a dorso-apical comb of four teeth. Hypopygium without sclerotized lateral patches, with a transverse row of hyaline setae (fig. e) ...... 11. C. capensis 5. Compound eye almost as long as the cheek (1.0-1.2). Maxilla without a bacilliform process (present in the other species, e.g., fig . g) ...... 6 - Compound eye approximately twice as long as the cheek (1.85 - 2.1) ...... 8 - Compound eye three or five times as long as the cheek ...... 10 6. Mandibular appendage with four ventral lamellae. Ovipositor valves distinctly longer than the gaster (1.35). Spine of the hypopygium more than three times as long as wide at the base ...... 9. C. namorakensis - Mandibular appendage with five ventral lamellae. Ovipositor valves shorter than the gaster. Hypopygial spine little longer than wide at the base ...... 7

209 7. Sixth and seventh antennal segments flabellate (fig. d). Fore tibia with four teeth in the dorso-apical comb ...... 12. C. ftnbellatus - Antennal segments of normal shape (fig. c). Fore tibia with a dorso-apical comb of five teeth ...... 13. C. si/vestrianus 8. Compound eye less than two times as long as the cheek (1.85). Antennal segments (nos. 7 - 9) with two rows of sensilla. Hypopygial spine 1.3 times as long as wide at the base ...... 16. C. b/ommersi - Compound eye approximately twice as long as the cheek (2.0-2.1). Antennal segments with three rows of sensilla. Hypopygial spine two to three times as long as wide at the base ...... 9 9. Fore tibia with four teeth in the dorso-apical combo Hypopygial spine three times as long as its width at the base ...... 17. C. /ongimucro - Fore tibia with three teeth in the dorso-apical combo Hypopygial spine not quite 2'/, times as long as wide basally ...... 15. C. coecus 10. Compound eye approximately three times as long as the cheek. Fore tibia with four or five teeth in the dorso-apical combo Stigmal and postmarginal veins of the fore wing subequal in length ...... 7. C. arabicus

Fig. 54. Details of Ceratoso/en. a- b, C. capensis Grandi: a, female antenna, and b, head, after Grandi (1955, figs . ii, 2 and I, respectively); c-

210 Table 8. Some characters of the males of Ceralosolen. no. Eyes Lengthlwidth Propodeum Number present of anellus of c1aws genit.

7. C. arabicus ...... + .. · 1.4 . · blunt ...... 4 8. C. stupefactus ...... + . . · 1.3 . · blunt ...... - 9. C. namorakensis .. . . + . .2.0 . · blunt ...... - 10. C. acutatus ...... + . . 0.5 . · acute ...... 4 11 . C. capensis ...... 0.5 ...... blunt ...... 5 12. C. flabellatus ...... + . . 1.0 ...... · acute...... 6 13. C. silvestrianus .. . + ...... 0.5 . . . . . · acute...... 6 14. C. galili ...... 0.5 .. . · blunt ...... 4 15. C. coecus · acute . . . . . 4 16. C. blommersi ...... + . . · 1.0 . . · blunt ...... 17. C. longimucro . . . + . . 0.5 .. · blunt ...... 18. C. megacephalus . .. - · 1.0 . . · acute ...... 4

- Compound eye five times as long as the cheek. Fore tibia with three teeth in the dorso-apical combo Postmarginal vein of the fore wing more than twice as long as the stigma I (2.25) ...... 8. C. stupefaclus

7. Ceratosolen arabicus Mayr Mayr, Wien. ent. Ztg. 25: 154 (1906); Grandi, Boll. Soc. ent. Ital. 48: 15-25 (1917). Synonyrns: Ceratosolen myersi Grandi, Boll. Ist. Ent. Univ. Bologna 21: 92- 97 (1955); C. longicornis Joseph, Proc. ent. Soc. Lond. (B) 28: 34-36 (1959); C. aelleni Grandi, Boll. Ist. Ent. Univ. Bologna 27: 71 - 75 (1964).

The female compound eye is th ree times as long as the cheek. The antennal segments bear two rows of sensilla; the sixth is more than three times as long as wide; the ten th and eleventh are fused in a club. The mandibular appendage bears five ventral lamellae; the maxilla has a bacilliform procesS. The stigma1 and postmarginal veins of the fore wing are subequal in length. The fore tibia has four or five teeth in the dorso-apical combo The spine of the hypopygium is as long as wide basally. The ovipositor valves are a bit longer than the gaster (1.1). The colour is yellow-brown: in comparison with the dark C. gatiti, which may occur in the same receptacles, C. arabicus is the 'blond' one.

The male eye is smalI, but clearly visible. In the antenna the anellus is almost 1 1 / 2 times as long as wide (1.4), and the third and fourth segments are subequal in length. The propodeum is blunt at the apex. The claspers of the genitalia bear four small claws.

The host figs are Ficus sycomorus Linn. (Sénégal, Guinea, Ivory Coast, Nigeria, Cameroon, Chad, Yemen, Ethiopia, Kenya, Tanganyika, Zaïre, Malawi, Zim-

211 babwe, Namibia, South Mrica, Madagascar), and Ficus mucuso Ficalho (Ivory Coast, Nigeria, Uganda): for a distribution map, see Wiebes, 1989b, fig. 1. In part of its range, C. arabicus occurs together with C. galili (no. 14), in the same fig-receptacles. The pollination ecology of C. arabicus was discussed by Galil & Eisikowitch (1969a, 1974); see also Wharton, Tilson & Tilson (1980).

8. Ceratosolen stupefactus Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet (C) 92: 264-265 (1989b).

The female compound eye is very large, five times as long as the cheek. The antennal segments bear two rows of sensilla; the sixth is approximately two times as long as wide; the tenth and eleventh segments are fused in a club. The man­ dibular appendage bears four ventral lamellae; the maxilla has a short bacil­ liform process. The postmarginal vein of the fore wing is more than two times as long as the stigmal (2.25). The fore tibia has a comb of three dorso-apical teeth. The spi ne ofthe hypopygium is 3'/2times as long as wide basally. The total length is ca. 1.5 mm; the ovipositor valves are 1'/2 times as long as the gaster. The colour is rather uniform light brown.

The male eye is smalI. The antennal anellus is somewhat longer than wide (5 : 4); the lengths of the fourth and fifth segments are in the ratio 5 : 6. The pro­ podeum is blunt at the apex. The genitalia do not bear claws.

The host fig is Ficus liliifolia Baker (Madagascar).

9. Ceratosolen namorakensis Risbec Risbec, Ann. Soc. ent. France 124: 186-188 (1956); Wiebes, Zool. Meded. Leiden 45: 6 (1970); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 263 (1989b).

The female head is longer than wide across the compound eyes (1.15); the eye is a bit longer than the cheek (1.2). The antennal segments bear two rows of sensilla; the tenth and eleventh are fused in a club. The mandibular appendage bears four ventrallamellae; the maxilla does not have a bacilliform process. The postmarginal vein is 1.65 times the length of the stigmal. The fore tibia has four teeth in the dorso-apical combo The spine of the hypopygium is very long: four times as long as wide basally. The ovipositor valves are 1.35 times as long as the gaster. The colour is brown.

The male eye is small. The third segment of the antenna is as long as the fourth, the fifth is longer (ratio 11 : 11 : 14). The propodeum is blunt at the apex. The genitalia do not bear claws.

The host fig is the form of Ficus sycomorus that was described as F. sakalavarum Baker (Madagascar).

212 10. Ceratosolen acutatus Mayr Mayr, Wien. ent. Ztg. 25: 154 (1906); Grandi, Boli. Lab. Zool. Portici 10: 176-182 (1916a).

The female compound eye is twice as long as the cheek. The antennal segments bear one row of sensilla; the ninth to eleventh are shaped so as to form a club. The mandibular appendage bears six or seven ventral lamellae; the maxilla has a bacilliform process. The stigmal and postmarginal veins of the fore wing are subequal in length. The fore tibia has five teeth in the dorso-apical combo The spine of the hypopygium is very short. The totallength is ca. 1.7 mm; the valves of the ovipositor are distinctly shorter than the gaster (0.7). The colour is dark brown.

The male eye is distinct, approximately as long as the cheek. The antennal anel­ lus has the shape of a disco The propodeum is acute at the apex. The genital claspers bear four large claws.

The host fig is Ficus vogeliana (Miq.) Miq. (Ivory Co ast, Cameroon, Nigeria).

11. Ceratosolen capensis Grandi

Grandi, Boli. Ist. Ent. Univ. Bologna 21 : 87-92 (1955).

The female compound eye is more than twice as long as the cheek (2.2). The antennal segments bear one row of sensilla; the ninth to eleventh are shaped so as to form a club. The mandibular appendage bears four or five ventrallamellae; the maxilla has a bacilliform process. The postmarginal vein of the fore wing 1 is ca. 1 / 2 times as long as the stigma!. The fore tibia has a comb of four dorso­ apical teeth. The spine of the hypopygium is rather wide and short, and it bears a transverse row of hyaline setae. The total length is ca. 1.7 mm; the valves of the ovipositor are about as long as the gaster. The colour is dark brown.

The male is blind. The antennal anellus is disc-like; the fourth and fifth segments are subequal in length. The propodeurn is blunt at the apex. The genital claspers bear five claws.

The host fig is Ficus sur Forssk. (Ethiopia, Uganda, Kenya, Tanganyika, Zam­ bia, Zimbabwe, South Africa): for a distribution map see Wiebes, 1989b, fig. 11 (the record from Ivory Coast is false!). In the northem part of its range, C. capensis occurs together with C. jlabellatus (no. 12). The pollination ecology was described by Baijnath & Ramcharun (1983); Compton & Robertson (1988) pub­ lished a note on the interaction of C. capensis with ants (in this connection, see also Thomas, 1988).

213 12. Ceratosolen f1abeUatus Grandi Grandi, Boll. Lab. Zool. Portici 10: 153- 160 (I916a). Synonym: C. dagatiguyi Risbec, Bull. Ist. franç. Afr. noire 13: 1126-1128 (195Ib).

The female eye is as long as the cheek. The fifth and sixth antennal segments are flabellate; the segments bear two rows of sensilla. The mandibular appen­ dage bears five ventral lamellae; the maxilla does not have a bacilliform process. The postmarginal vein of the fore wing is twice as long as the stigma!. The fore tibia bears a dorso-apical comb of four teeth. The hypopygial spi ne is a bit over twice as long as wide basally (30 : 13). The valves of the ovipositor are almost as long as the gaster (0.95). The colour is dark brown.

The male eye is distinct. The antennal anellus is as long as wide; the fourth and fifth segments are subequal in length. The propodeum is acute at the apex. The genital claspers bear six large claws.

The host fig is Ficus sur Forssk. (Guinea, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Ethiopia, Uganda, Kenya), in which the species occurs together with C. capensis (no. 11) or C. silvestrianus (no. 13); see Wiebes, 1989b, fig. 11, for a distributional map.

13. Ceratosolen silvestrianus Grandi Grandi, Boll. Lab. Zool. Portici 10: 160--166 (1916a).

The female head is as long as wide across the compound eyes; the eye is as long as the cheek. The antennal segments bear two rows of sensilla: the tenth and eleventh are fused in a club. The mandibular appendage has five ventral lamel­ lae; the maxilla does not have a bacilliform process. The postmarginal vein of the fore wing is twice as long as the stigma!. The fore tibia has a dorso-apical comb of five teeth. The hypopygial spine is almost twice as long as wide basally. The valves of the ovipositor are almost as long as the gaster (0.95). The colour is brown.

The male eye is smal!. The antennal anellus is half as long as it is wide; the fourth and fifth segments are subequal in length. The propodeum is acute at the apex. The genital claspers bear ca. six small claws.

But for the difference in the female antennae, and some other, small differences, C. silvestrianus is very similar to C. flabellatus. With some doubt, it was recorded by Wiebes (1989b: 264) from Ficus sur Forssk., in which it now appears to be anormal inhabitant in West Africa, next to C. flabellatus (no. 12), and from Ficus vallis-choudae Delile (probably wrong; it has C. megacephalus (no. 18) as the normal symbiont). It is known from Guinea, Sénégal, Ivory Coast and Ni­ gena.

214 14. Ceratosolen galili Wiebes Wiebes, Ent. Ber. Arnst. 24: 188-191 (1964).

The female compound eye is as long as wide across the compound eyes; the eye is subequal in length to the cheek. The anten na I segments bear one row of sen­ silla; the ninth to eleventh are shaped so as to form a club. The mandibular appendage has six ventral lamellae; the maxilla bears a bacilliform process. The postmarginal vein of the fore wing is al most twice as long as the stigmal (1.85). The fore tibia has five teeth in the dorso-apical combo The hypopygium has a short and wide spine, which shows heavily sclerotized lateral patches. The valves of the ovipositor are one quarter longer than the gaster. The colour is dark brown.

The male is blind. The antennal anellus is half as long as it is wide; the fourth and fifth segments are subequal in length. The propodeum is blunt at the apex. The genital claspers bear four claws.

The host figs are Ficus sycomorus Linn. (Sénégal, Ivory Coast, Nigeria, Zaïre, South Africa, and in East Africa: Ethiopia, Kenya, Tanganyika, see the map in Wiebes, 1968a, fig. 1) and Ficus mucuso (lvory Co ast, Ghana, Nigeria, Uganda). C. galili, which behaves as a cuckoo (ovipositing in the flowers without pollinat­ ing them, see Galil & Eisikowitch, 1969a), occurs together with C. arabicus (no. 7), which is the pollinator. For biological notes, see Compton et al. (1991).

15. Ceratosolen coecus (CoquereI) Coquerel, Rev. Mag. Zool. (2) 7: 422--426 (1855, Sycocrypta). Synonym: Ceratosolen emmerezi Grandi, Boll. Soc. ent. Ital. 56: 81-87 (1924).

The female head is longer than wide across the compound eyes (l.05); the eye is more than twice as long as the cheek (2.1). The antennal segments bear three rows of sensilla; the tenth and eleventh are fused in a club. The mandibular appendage bears four or five ventral lamellae; the maxilJa has a bacilliform process. The postmarginal vein of the fore wing is twice as long as the stigmal. The fore tibia has three teeth in the dorso-apical combo The hypopygial spine is more than twice as long as wide basally (2.3). The totallength is ca. 1.7 mm; the valves of the ovipositor are approximately as long as the gaster. The colour is dark brown.

The male is blind. The antenna has only four segments, i.e. , an anellus is absent; the penultima te and ultimate segments are subequal in length. The propodeum is acute at the apex. The genital claspers bear some four small claws.

The host fig is Ficus mauritiana Lam. (Réunion, Mauritius).

215 16. Ceratosolen blommersi Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 257-259 (1989b).

The female compound eye is almost twice as long as the cheek (1.85). The an­ tennal segments have two rows of sensilla; the tenth and eleventh together form a club. The mandibular appendage bears five ventral lamellae; the maxilla has a bacilliform process. The postmarginal vein of the fore wing is almost twice as long as the stigmal (1.8). The fore tibia has four teeth in the dorso-apical combo The hypopygium has a rather short spine (1.3). The total length is ca. 1.8 mm; the ovipositor valves are shorter than the gaster (0.8). The colour is yellow­ brown.

The male eye is smal!. The antennal anellus is almost as long as wide; the fourth segment is much shorter than the fifth (0.6). The propodeum is blunt at the apex. The genitalia have no claws.

The host fig is Ficus botryoides Baker (Madagascar).

17. Ceratosolen longimucro Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 262 (1989b).

The female compound eye is twice as long as the cheek. The antennal segments have three rows of sensilla; the tenth and eleventh are fused in a club. The mandibular appendage bears five ventrallamellae; the maxilla has a bacilliform process. The postmarginal vein of the fore wing is almost twice as long as the stigmal (1.85). The fore tibia has four teeth in the dorso-apical combo The hypo­ pygial spine is long: three times as long as wide basally. The valves of the ovipo­ sitor are a bit longer than the gaster (1.1). The colour is brown.

The male eye is smal!. The antennal anellus is half as long as wide; the fourth segment is distinctly shorter than the fifth (0.6). The propodeum is blunt at the apex. The genitalia have no claws.

The host fig is Ficus polyphlebia Baker (Madagascar).

18. Ceratosolen megacephalus Grandi

Grandi, Boll. Soc. ent. Ital. 48: 25-30 (1917). Synonym: Ceralosolen carayoni Grandi, Boli. Ist. Ent. Univ. Bologna 26: 231 - 238 (1963a).

The female head is large, longer than wide across the compound eyes (1.1); the eye is shorter than the cheek (0.85). The antennal segments have one row of sensilla; the ninth to eleventh segments are fused in a club. The mandibular appendage bears six to eight ventral lamellae; the maxilla has a bacilliform process. The

216 postmarginal vein of the fore wing is almost twice as long as the stigmal (\ .9). The fore tibia has four teeth in the dorso-apical combo The hypopygium has a rather short spine, not much longer than wide at the base. The total length is ca. 2.8 mm; the valves of the ovipositor are equal in length to the gaster or longer (1 - 1.5). The colour is yellow-brown.

The male is blind. The antennal anellus is approximately as long as wide; the fifth segment is four-fifths of the length of the fourth. The propodeum is acute at the apex. The genital c1aspers bear four c1aws. The totallength is ca. 2 mmo

The host tig is Ficus vallis-choudae DeWe (Ivory Coast, Togo, Cameroon, Ethio­ pia, Kenya, Burundi, Uganda, Tanganyika, Zambia, Malawi).

Dolicboris Hill (fig. 55)

Hili, Zool. Verh. Leiden 89: 38-39 (1967a).

There is only one Mrican species, i.e., D. flabella ta Wiebes, from figs of the subgenus Pharmacosycea, section Oreosycea. Some of the characters of this one species (e.g., the flabellate female antenna with long sensilla chaetica, and the structure of the male meso- and metanotum) are exceptional in the genus, which is mainly Indo-Australian and always is associated with figs of the section Ore­ osycea.

19. Dolicboris flabeUata Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 82: 184-186 (1979a).

Fig. 55. Do/ichoris ftabellata Wiebes, male and female.

217 The female head is shorter than wide across the compound eyes (0.75); the longitudinal diameter of the eye is twice as long as the cheek. There are three ocelli. The antenna consists of ten segments; the fourth to sixth each have one whorl of long sensilla, the seventh to ninth, moreover, have one or two apical sensilla ànd a long flabellum, also bearing sensilla; the ten th is shaped as a flabellum. The mandible is bidentate at the apex; the appendage has five ventral lamellae; the maxilla has a bacilliform process, which has a free apical segment. The thorax has pollen pockets and the fore coxae have a corbicula. In the fore wing, the postmarginal vein is obsolete. The fore tibia has two dorso-apical teeth. The stigmatal peritremata of the eighth urotergite are small, subcircular. The hypopygium has a rather short, blunt spine, with two (or only one) trans­ verse rows of hyaline setae. The total length (head, thorax and gaster) is ca. 1.6 mm; the valves of the ovipositor are over one mm long, i.e., twice as long as the gaster. The colour is dark brown.

The male head is sligthly wider than long; the eyes are rather small. The antenna consists of four segments, the third of which is anelliform. The separation of the meso- and metanotum is visible only laterally, the me­ tanotum (through which the metapleura are faintly visible) and the propodeum are fully separate; the spiracular peritremata are half as long as the propodeurn. The fore tibia has three teeth in the dorso-apical comb; the tarsus is bimerous; the tarsi of the mid and hind legs are pentamerous. The genital claspers have twelve to fifteen small claws. The length (head and thorax) is ca. 0.9 mmo The colour is yellowish.

The host figs are Ficus variifolia Warb. (Gabon) and Ficus dicranostyla Mildbr. (Zambia). The host relations were discussed by Wiebes & Compton (1990: 209). The species is also known from Sénégal and Ethiopia.

Parapristina Hill (fig. 56)

Hili, Zool. Verh. Leiden 89: 31 - 32 (l967a, subgenus of Blastophaga).

There is only one species in this Indo-Australian group, that is treated as a separate genus rather than as a subgenus of Blastophaga. The species was intro­ duced into Africa.

20. Parapristina verticillata (Waterston)

Waterston, Bull. ent. Res. 12: 38-40 (Eupristina, 1921a); Grandi, Boll. Lab. Zool. Portici 20: 183- 185 (I927); Hili, Zool. Verh. 89: 80--93 (Parapristina, 1967a). Synonym: Euprista okinavensis Ishii, Kontyu 8: 85- 86 (1934). The female head is shorter than wide across the compound eyes (0.8); the longi­ tudinal diameter of the eye is subequal to the length of the cheek. There are three

218 - """'"""""'::;;_.!!lir---__

Fig. 56. Parapristina verticillata (Waterston), male and female.

ocelli. The antenna consists of eleven segments; the fifth to tenth are more or 1 less cup-like, with sensilla chaetica in one apical whorl, which are 2 - 2 / 2 times as long as the segment; the eleventh is inversely pyriform in shape. The mandible is bidentate at the apex; the appendage has ca. ten ventrallamellae, the fust two of which are tooth-like. The labium and maxillae are sirnple. The thorax has pollen pockets and the fore coxae have a corbicula. In the fore wing, only the submarginal and the marginal veins are developed, although a trace of the stigmal may be seen. The fore tibia bears two dorso-apical teeth. The stigmatal peritremata of the eighth urotergite are smalI, subcircular. The hypo­ pygium has a short, acute spine. The totallength (head, thorax and gaster) is 1.0-1.2 mrn; the ovipositor valves

219 are longer than the gaster (1.25). The colour is dark brown, the legs are yel­ lowish.

The male head is wider than long (1.25); the eyes are quite large, positioned anteriorly, near the attachment of the mandibles. Antennal grooves separate; the antenna consists of four segments, the third is anelliform. The thoracical terga are fused dorsally, but laterally some separation is pre­ sent between the metanotum and the propodeum; the spiracles of the propo­ deum are dorsal in position, sm all and circular in shape. The fore tibia has three dorso-apical teeth; the tarsus is incompletely trimerous; the mid and bind tarsi are tetramerous. The genitalia are simpIe. The length (head and thorax) is 0.5-0.6 mmo The colour is yellowish.

The host fig is Ficus microcarpa Linn.f. (Morocco).

Platyscapa Motschoulsky (fig. 57) Motschoulsky, Bull. Soc. Natural. Moscou 36: 48 (1864); Wiebes, Neth. J. Zool. 27: 209-223 (l977b); Wiebes & Abdurahllnan, Proc. Kon. Ned. Akad. Wet. (C) 83: 195-207 (1980); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: table 5 (1986b).

With one exception, the female head is shorter than wide (0.85-0.95); the lon­ gitudinal diameter of the compound eye is (sometimes much) longer than the cheek (one exception: shorter). There are three ocelli. The antenna normally has eleven segments, but two species have ten; the fifth to (tenth or) eleventh seg­ ments have the sensilla, which in some species project beyond the edge of the segment, usually in one row (one exception). The mandible is bidentate at the apex; the appendage has seven to ten ventral lamellae, the proximal one to four of wbich may he medially produced into a tooth. The labium and maxillae are simpie. The thorax has pollen pockets and the fore coxae have a corbicula. The vena­ tion of the fore wing is complete or it may be reduced beyond the submarginal vein. The fore tibia has two or three dorso-apical teeth. The hind femur has a ventral concavity, in which fits the tibia. The stigmatal peritremata of the eighth urotergite are small and circular. The hypopygium usually has a (sometimes very) short, blunt spine, but it may be longer and sharp. The total length (head, thorax and gaster) is ca. 1.5 mm; the ovipositor valves are slightly longer than the gaster to 1'/2times as long. The colour is dark.

The length of the male head varies from somewhat shorter (0.9, one species 0.6) to somewhat longer (1.1) than the width; the length of the eye is one-sixth to one-quarter of the length of the head. An anterior depression usually reaches to about one-third of the length of the head, but it may reach to about one-half of this length. The antenna has only one, or two short anelli, two anelli of inequal length, or three short, subequal anelli.

220 Fig. 57. Platyscapa quadraticeps (Mayr), male and female, after Galil & Eisikowitch (1968c, figs. 2 and I, respectively).

The pronotum is transverse in shape. The meso- and metanotum may be completely free, or fused, the metanotum and the propodeum are completely free, or partially (in the middle) fused. The spiracular peritremata of the pro­ podeum are smalI, subcircular. The fore tibia usually has two teeth in the dorso­ apical comb (three in one species); the tarsus is bimerous, but (incompletely) pentamerous in one species. The armature of the hind tibia consists of two bidentate teeth, but the antiaxial tooth may be tricuspidate. The genitalia are simpie. The length (head and thorax) is 0.7-1.0 mmo The colour is yellowish.

There are six African species known (and eight Indo-Australian species, one of which - P. quadraticeps - was introduced into Israel). The host figs belong to the sections Urostigma and Conosycea of the subgenus Urostigma.

221 KEY TO THE SPECIES (fig. 58)

I. Females 2 - Males . 8

2. Veins in the fore wing reduced beyond the apex of the submarginal (fig. a) . 3 - Fore wing with complete venation . 6

3. Antenna consisting of ten segments . 4 - Antenna consisting of e1even segments ...... 5

4. Epistomal margin distinctly trilobate (fig. b). All veins of the fore wing beyond the submarginal, obsolete. Hypopygium with a sharp spine, which has a transverse row of hyaline setae at half length ...... 21. P. awekei - Epistomal margin almost straight, but the median lobe rather prominent. Only the stigmal vein obsolete. Hypopygium with a short, blunt spine, lacking the transverse row of setae ...... 22. P. bingharni

5. Fore wing with a striking, dark, stigmal macula (fig.c) ...... 23. P. etiennei - Fore wing without such a striking macula ...... 24. P. desertorurn

6. Antennal sensilla on segments 7 - 9 in more than one complete whorl (fig. d) ...... 27. P. soraria

~Z) d

G(--:. .) Fig. 58. Details of Platyscapa. a-b, P. awekei Wiebes: a, veins of fore wing, and b, female head, after Wiebes (l977b, figs. 16 and 19, respectively); c, P. eliermei Wiebes, wings, after Wiebes (1981, fig . 2); d, P. soraria Wiebes, female antenna, after Wiebes & Abdurabiman (1980, fig. 36); e, P. qundraticeps (Mayr), female antenna, after Grandi (1923, fig. 2); f- h, male antenna of: f, P. soraria, g, P. quadraticeps, after Grandi (l928b, fig. vii, 2) and h, P. awekei, after Wiebes & Abdurahiman (1980, fig. 17); i- j, male head: i, P. awekei, after Wiebes & Abdurahiman (1980, fig. 14) and j. P. etiennei, after Wiebes (1977b, fig. 36); k, P. binghami Wiebes, male antenna, after Wiebes & Abdu­ rahiman (1980, fig. 19).

222 - Antennal sensilla in one whorl ...... 7 7. Head over three times as long as the longitudinal diameter of the compound eye; the length of the eye as long as the cheek. Antennal sensilla, especially those of the distal segments, strongly projecting (fig.e) ...... 25. P quadraticeps - Head a little over twice as long as the longitudinal diameter of the compound eye (2.25); the eye I '/2 times as long as the cheek. Antennal sensilla with the distal parts only slightly projecting...... 26. P bergi 8. Antenna with two short anelli (fig. f), often indistinguishable as if there is only one. Armature of the hind tibia consisting of a tricuspidate antiaxial tooth and a bidentate axial ...... 9 - Antenna with two anelli of different length, or with three short, subequal anelli. Arma- ture of the hind tibia consisting of two bidentate teeth ...... 10 9. Head a bit longer than wide. Metanotum separate from the propodeum ...... 27. P sararia - Head much shorter than wide (5 : 8). Metanotum and propodeum separate only la­ terally, fused medially ...... 24. P desertorum

10. Antenna with two inequal anelli (fig. g). Mesonotum and metanotum completely sep­ arate...... 25. P quadraticeps - Antenna with three short anelli (figs. h, k). Mesonotum and metanotum fused, but some separation may be apparent laterally ...... 11 11. Fore tibia (indistinctly) pentamerous ...... 26. P bergi - Fore tibia bimerous ...... 12 12. Antennal groove rather wide at front, toruli situated to the sides of the head (fig. i) 13 - Antennal groove more shallow, toruli situated more close together (fig. j) ...... 23. P etiennei

13. Antennal scape 2'/2 times as long as wide (fig. k) ...... 22. P binghami - Antennal scape more slender, almost 3'/2 times as long as wide (fig. h) .. 21. P awekei

21. Platyscapa awekei Wiebes

Wiebes, Neth. J. Zool. 27: 215-217 (1977b); Wiebes & Abdurahiman, Proc. Kon. Ned. Akad. Wet. (C) 83: 199- 200 (1980).

The female compound eye is rather large, three times as long as the cheek. The epistomal margin is trilobate. The antenna has only ten segments, the fifth to tenth of which each have one row of long sensilla. The mandibular appendage bears seven ventrallamellae, the proximal three of which are produced medially. The submarginal vein of the fore wing is complete, but the marginal and post­ marginal veins are obsolete; the stigmal vein is only partIy visible. The fore tibia has a dorso-apical comb of two teeth, one ventral and a spur. The spine of the hypopygium is not very long, but sharp; it has a transverse row of hyaline setae at half length.

The length of the male eye is one-quarter of the length of the head; the anterior depression reaches to about half the length of the head. The antennal toruli are situated at the sides of the head; the antenna has a rather slender scape (length

223 : width, almost 3.5) and three anelli. The meso- and metanotum are completely fused. The hind tibia has a bifurcate axial tooth and a tricuspidate antiaxial.

The host tig is Ficus cordata subspec. salicifolia (Vahl) Berg (Ethiopia, South Mrica). It is of interest to note that the nominate form of F cOf{iata Thunb. harbours a distinct species, i.e. , P desertorum (no. 24).

22. Platyscapa binghami Wiebes

Wiebes in Wiebes & Abdurahiman, Proc. Kon. Ned. Akad. Wet. (C) 83: 201 - 202 (1980).

The female compound eye is twice as long as the cheek. The epistomal margin is almost straight, but the median lobe is rather prominent. The antenna, man­ dible and legs are much like those of P awekei, but the axial spur of the bind tibia is simple, not bifurcate. The veins of the fore wing are complete, but for the obsolete stigmal. The spine of the hypopygium is short and blunt.

The male is much like that of P awekei, but the anterior depression is shallower and the scape is more robust, i.e., about 2'/2 times as long as wide.

The host tig is Ficus verruculosa Warb. (Zambia).

23. Platyscapa etiennei Wiebes Wiebes, Neth. J. Zool. 27: 217-220 (1977b).

The female compound eye is not much longer than the cheek (1.2). The epistomal margin has indistinct lateral lobes and a more distinct median prorninence. The antenna has eleven segments, with sensilla much like those of P awekei. The mandibular appendage bears ten or eleven ventral lamellae, the proximal two of which are produced medially. The fore wing has a striking, dark macula; the veins beyond the submarginal are obsolete. The dorso-apical comb of the fore tibia has two teeth. The spine of the hypopygium is short and blunt.

The length of the male eye is one-sixth of the length of the head; the anterior depression reaches to about one-third of the length of the head. Most characters are much as in P awekei, but for the situation of the antennal toruli, which are more close together, and both tee th of the hind tibia are bidentatelbifurcate.

The host tig is Ficus densifolia Miq. (Réunion).

24. Platyscapa desertorum Compton Compton in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 211 - 212 (1990).

The female compound eye is 1'/2 times as long as the cheek. The epistomal

224 margin has distinct lateral and median projections. The antenna, man di bie and also the wing-venation are much like in P etiennei, but there is no large, dark wing-macula. The fore tibia has three teeth in the dorso-apical combo The spine of the hypopygium is distinct, but not very long.

The male head is rather short relative to its width (0.6); the length of the eye is one-quarter of that of the head, and also the anterior depression reaches to about one-quarter of the length of the head. The antenna has only one anellus. The pronotum is wide, with concave lateral margins; the meso- and metanotum are completely fused . The fore tibia has three dorso-apical teeth; the hind tibia has a tricuspidate antiaxial tooth.

The host fig is Ficus C. cordata Thunb. (South Africa). See the remark under P awekei (no. 21).

25. Platyscapa quadraticeps (Mayr)

Mayr, Verh. zool.-bot. Ges. Wien 35: 176-177 (1885, Blastophaga); Grandi, Bull. ent. Res. 13: 295- 297 (1923); Grandi, Boll. Lab. Ent. Bologna I: 123-126 (I 928b); Wiebes, Neth. J. Zool. 27: 214 (I 977b, full bibliography). Synonym: Blastophaga arnofliana Joseph, Agra Univ. J. Res. (Sci.) 2: 270-277 (1953).

The female compound eye is shorter than the cheek (0 .9). The epistomal margin shows a feebie angular median prominence and two rounded lateral projections. The antenna has eleven segments, the fifth to seventh of which bear long sensilla, and those of the eighth to eleventh are longer still, projecting to a distance equal to once or twice the length of the segment. The mandibular appendage bears seven ventral lamellae, the first of which has a tooth-like appearance. The fore wing has a complete venation. The fore tibia has three teeth in the dorso-apical combo The hypopygium has a distinct spine, but it is not very long.

The male head is subquadrate; the length of the eye is one-third of that of the head; the anterior depression reaches to about one-third of the length of the head. The antenna has two, inequal anelli. The meso- and metanotum are com­ pletely separate. The hind tibia has two bidentate teeth.

The host fig is Ficus religiosa L. (India, Ceylon, Malaya; introduced into Israel). Pollination was studied by several authors e.g. , GaW & Snitzer-Pastemak (1970) - see Wiebes (l977a: 221) for a summary.

26. Platyscapa bergi Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 347-351 (I 986b). The female head is longer than wide across the compound eyes (1.15); the eye 1 is 1/ 2 times as long as the cheek. The epi stomaI margin has rather indistinct

225 lobes. The antenna, mandible and fore tibia are much like those of P. awekei, but the antennal sensilla are shorter. The venation of the fore wing is much like that of P. quadraticeps, but the postmarginal vein is rather short, i.e., half as long as the stigmal, versus subequal in length. The hypopygium has a short, blunt spine.

The male head is longer than wide (1.15); the length of the eye is one-quarter the length of the head and the anterior depression reaches to about half the length of the head. The antenna has three anelli. Dorsally, all terga of the thorax beyond the pronotum are fused. The fore tarsus is indistinctly pentamerous. Both teeth of the hind tibia are bidentate.

The host fig is Ficus menabeensis Perrier (Madagascar).

27. Platyscapa soraria Wiebes Wiebes in Wiebes & Abdurahiman, Proc. Kon. Ned. Akad. Wet. (C) 83: 206 (1980).

The female compound eye is almost twice as long as the cheek (1.85). The epi­ stoma I margin is almost straight, with a weak median lobe. The antenna has eleven segments, the distal of which, from the seventh onwards, bear two to three rows of sensilla. The mandible, fore tibia and the venation of the fore wing are much like those of P. bergi. The hypopygium has a distinct spine, which is not very sharp.

The male head is longer than wide (1.1) and the length of the eye is about one-quarter of the length of the head; the anterior depression reaches to about one-third of the length of the head. There are two anelli in the antenna. The terga of the thorax are completely separate. The hind tibia has a bidentate axial tooth and a tricuspidate antiaxial.

The host fig is Ficus ingens (Miq.) Miq. (Ivory Coast, Ethiopia, Malawi, Zambia, South Africa). Barker (1985) gave evidence of a volatile compound in F ingens attracting P. soraria.

Allotriozoon Grandi (fig. 59) Grandi, BoU. Lab. Zool. Portici 10: 182-191 (l916a); Wiebes, Zool. Meded. Leiden 48: 137-138 (I 974b).

The female head mostly is longer than wide across the compound eyes (1.3), but it may be as long, or even shorter than wide (0.9); the longitudinal diameter of the compound eye equals the length of the cheek or is distinctly longer (up to 2.5). There are no ocelli. The antennal toruli are situated close to the epistomal margin, which has a characteristic clypeal process. The antenna consists of

226 Fig. 59. Allolriozoon prodigiosum Grandi, female and male. eleven free segments; the third segment is simpie, with a blunt apex; the fourth is short, sometimes transverse; the fifth to eleventh bear shortly elongate sensilla linearia in two or three rows. The mandible is rather robust, with one distinct apical tooth which, as weil as the mandible itself, may be minutely dentate along the oral margin; the appendage has a great number (ca. twenty to over sixty) of ventral rows of sm all crenulations (ten to ca. forty). The maxillo-labial com­ plex is elongate, simpie, with a tuft of long distal setae. The thorax has pollen pockets, but the fore coxae have no corbicula. The venation of the fore wing is complete; the postmarginal vein is short. The fore tibia has two or three dorso-apical teetb. Tbe hind tibia has a bidentate ventral tooth. The stigmata I peritremata of the eighth urotergite are very large and elongate. The hypopygium has an acute spine.

227 The total length (head, thorax and gaster) is ca. 2 to 3 mm; the ovipositor valves are about as long as the gaster. The colour is brown.

The male head is more than 1'/2 times as long as wide (1.55--1.6); there are no eyes. The antennal toruli are situated close to the epistomal margin, on either si de of the median process, which bears long setae. The antennae consist of four segments. The thoracical terga are all free: the pronotum is a bit wider than long (0.9) or subquadrate; the mesonotum is 1.3-1.5 times as wide as long, the posterior angles appear produced and reach the metanotum, the two dorsal parts of which are situated laterad of the propodeurn. The propodeal spiracles are almost half as long as the propodeurn, and they are dorsal in position. The fore tibia has a dorso-apical hook, more or Ie ss distinctly bilobate; the tarsus is trimerous; the mid and bind tibiae may have some spines, but no teeth, the tarsi are tetra­ rnerous. The genitalia are simpie, without claspers. The length of the head and thorax is ca. 1.5-2 mmo The colour is yellowish.

There are three species known as pollinators of the fig-species of the subsection Galoglychia.

KEY TO THE SPECIES (fig. 60)

I. Oral margin of the female mandible, and thaI of its apical tooth, finely dentate (fig. b). The number of ventral rows of crenulations on the mandibular appendage is less than thirty, the crenulations number up to ten. Male antenna ca. half as long as the head, the segments are not elongate (fig. f) ...... 30. A. heterandromorphum - Oral margin of the female mandible and that of its apical tooth, smooth (fig. a). The number of rows on the mandibular appendage is larger, as is the number of crenula- tions per row. Male antenna as long as the head, the segments are elongate (fig. g) .. 2 2. Between forty and fifty ventral rows of crenulations on the female mandibular ap­ pendage, the number of crenulations per row are up 10 twenty. Tooth of the bind tibia short, triangular (fig. j ). The male of this species and that of the next are very similar and cannot be distinguished ...... 28. A. prodigiosum - Over sixty rows on the female mandibular appendage, the number of crenulations per row are ca. forty. Tooth of the hind tibia very large, evenly bifurcate (fig. k) ...... 29. A. nigeriense

28. AUotriozoon prodigiosum Grandi Grandi, Boll. Lab. Zool. Portici 10: 191- 200 (l916a); Wiebes, Zool. Meded. Leiden 48: 142- 243 (1974b).

The female head is approximately 1.3 times as long as wide between the com­ pound eyes; the eye is much shorter than the cheek (0.4--0.5). There is some varia ti on in the shape of the head, in which the eye may be longer relative to the length of the cheek than mentioned above (0.7). The oral margin of the man di bie and that of its apical tooth are smooth; the appendage is six times as

228 "

'. " "'"! : 9 . :':1, ,\ Nl klIt

Fig. 60. Details of Allotriozoon. a, A. prodigiosum Grandi, female mandible and base of appendage, after Grandi (I916a, fig. xviii , 3); b, A. heterandromorphum Grandi, female mandible and appen­ dage, after Wiebes (I 974b, fig. 58); c, A. nigeriense Wiebes, female antenna, after Wiebes (I 974b, fig . 48); d-e, female head of d, A. heterandromorphum, and e, A. prodigiosum, after Wiebes, 1974b (figs. 55 and 59, respectively); f, A. heterandromorphum, male antenna, after Grandi (1916a, fig. xxiii, 2); g, A, prodigiosum, male head, after Grandi (1916a, fig. xix, I); h-i, female fore tibia of h, A. nigeriense (after Wiebes, 1974b, fig. 51), and i, A. heterandromorphum; j-k, armature of female hind tibia of j, A. prodigiosum and k, A. nigeriense (after Wiebes, 1974b, fig . 53).

long as wide, and it bears ca. forty to fifty ventral rows of ab out twenty small crenulations (sixteen in one sample). The fore tibia is short, with a bidentate process dorso-apically; the hind tibia has a short, triangular antiaxial tooth. This is the smallest of the species of Allotriozoon: the totallength is ca. 2- 2.5 mmo

The male head is strongly depressed; the antennae are characteristically e1on­ gate, with subcylindrical segrnents. The mandible is falcate. The segments of the legs are elongate; the femora bear stout dorsal spines.

The host fig is Ficus saussureana De. (Guinea, Sierra Leone, Ivory Coast, Ghana, Nigeria, Zaïre, Uganda).

229 29. Allotriozoon nigerieme Wiebes

Wiebes, Zool. Meded. Leiden 48: 138-139 (1974b).

The female head is approximately 1.3 times as long as wide across the compound eyes; the cheek is twice as long as the eye. The oral margin of the mandi bIe is smooth and the tooth is blunt as if wom; the appendage is four times as long as wide, with over sixty vent ral rows of about forty fine crenulations. The fore tibia has two dorso-apical teeth; the hind tibia has an evenly bifurcate, ventral tooth. The total length is ca. 3 mmo

The male is similar to that of A. prodigiosum.

The species was described from a female specimen caught at light (Nigeria), but later (Wiebes & Compton, 1990: 212) the host was found to be Ficus chlamy­ docarpa Mildbr. & Burret (Carneroon).

30. AUotriozoon heterandromorphum Grandi

Grandi, Boll. Lab. Zool. Portici 10: 191 , 200-206 (1916a); Wiebes, Zool. Meded. Leiden 48: 139- 142 (1974b). Synonyms: Allotriozoon seychellense Masi, Novit. Zool. 24: 123 (1917); Blastophaga wanei Risbec, Mém. Inst. franç. Afr. noire 13: 385-386 (195Ia, male only); Blastophaga dnkarensis Risbec, Bull. Inst. franç. Afr. norre 16: 536-538 (1954a).

The female head is approximately as long as wide across the compund eyes, which are equal in length to the cheek, or a bit longer (1.1). There is a rather large variation in the shape of the head, as the length-width ratio may vary from 0.9 to 1.3 (or even 1.7 in specimens from the Seychelles Is.), and the eye may be shorter relative to the length of the cheek than mentioned (0.75). The oral margin of the mandible and th at of its apical tooth are finely den tate ; the number of ventral rows on the mandibular appendage (which is ca. seven times as long as wide) varies from ca. twenty to thirty (in specimens from the Seychelles Is.) and the rows have ca. ten fine crenulations. The fore tibia has three minute dorso-apical teeth; the rund tibia has a short, triangular antiaxial tooth. The total length is ca. 3 mmo

The male he ad is not depressed; the antennae are robust and not at all elongate; the scape is relatively flat. The mandible is rather massive. The segments of the legs are more robust than in A. prodigiosum, not at all elongate, and the femora have no stout, dorsal spines.

The host fig is Ficus lu tea Vahl (Sénégal, Guinea, Ivory Coast, Ghana, Nigeria, Zaïre, Kenya, Comores Is., Seychelles Is., Madagascar). Newton & Lomo (1977) published a note on the possible role of the pollen load on the flight of the female insect; Newton & Lomo (1979) discussed the pollination of Ficus lutea in Ghana.

230 ElisabethieUa Grandi (fig. 61)

Grandi, Boll. Lab. Ent. Bologna 1: 68-69 (1928a, subgenus of Blastophaga): Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 117-136 (1989a). The female head is approximately as long as wide across the compound eyes (0.85- 1.3). There are three ocelli (one exception with only two). The antenna consists of eleven free segments; the scape is aften wide (or only twice as long as wide); it is prolonged apically in one group of species; the pedicel is rotundate, or distinctly angular; the third segment is simpie; the fifth to eleventh bear long flexible, or rod-like sensilla chaetica, or long sensilla linearia, in one row per segment (one exception with more rows). The mandible in most species is bi-

Fig. 61. Elisabethiella baijnathi Wiebes, male and E. stuckenbergi (Grandi), female.

231 dentate, it has a number of ventral ridges (that in some species may be produced into teeth); the appendage has many rows of sm all teeth or fine crenulations. The labium and maxillae are simpIe. The thorax has pollen pockets and the fore coxae have a corbicula. The vena­ tion of the fore wing is complete. The fore tibia has a dorso-apical comb of three teeth (two in two species). The hind tibia has a bidentate ventral tooth, but it is tricuspidate in two species. The stigmatal peritremata of the eighth urotergite are large, or medium-sized in most species. The spine of the hypopygium is not very long. The total length (head, thorax and gaster) is 1.3-2.1 mm; The ovipositor valves may be only as long as the gaster, but mostly they are longer: up to twice as long as the gaster.

The male head is approximately as long as wide (0.9- 1.1); the longitudinal diam­ eter of the eye is ca. one-sixth of the length of the head, or a bit longer (one-fifth) or shorter (one-eighth), and the eye is situated close to the epistomal margin. The antenna is five-segmented, consisting of the scape, ped icel , two anelli, and 1 a club, which may be ca. two times as long as wide, or (almost) 2 / 2, or exception­ ally three times. The thorax has a characteristic shape (fig. 49, d) in that it narrows posterad, i.e., the propodeum is much narrower than the mesonotum, and also narrower than the metanotum; these terga are fused dorsally. The spiracular peri trema ta of the propodeum may be small (one-quarter or one-third of the length of the propodeurn: fig. h), medium-sized (ca. one-half: fig. 49, d), or larger (three- or four-fifths: fig. g). The fore tibia has three large dorso-apical teeth (sometimes with a smaller in between the first and the second) and two in the ventral angle; the tarsal segments may be partially fused. The hind tibia has a bicuspidate antiaxial crest and a bidentate, axial tooth. The disc of the tibiae may bear spines; all tarsi are essentially pentamerous. The genitalia are simpIe. The total length of the head and thorax is ca. 1 mm (0.8-1.3). The colour is yellowish.

There are fourteen species (one with two subspecies), which pollinate figs belong­ ing to the subsections Platyphyllae, Chlamydodorae and Crassicostae. For two species the species of host fig is not known. There are three species-groups which, as well as the species in the female sex, can be identified with the following key. The males are much more difficult to identify.

KEY TO THE SPECIES (females; for the males, see table 9) (fig. 62)

I. Antennal scape elongate, prolonged beyond the point of attachment of the pedicel (fig. a). Mandibular appendage with thirty or (many) more rows of denticulations (group of E. articu/ata) ...... 2 - Antennal scape not or scarcely prolonged apically (figs. b, f). Mandibular appendage with less than thirty rows of denticulations (fig. 49, b) ...... 7

232 Fig. 62. Details of Elisabe/hiella. a-b, base of female antenna: a, E. ar/ieulata (Joseph), after Wiebes (l989a, fig. I) and b, E. s/uekenbergi (Grandi), after Grandi (1955, fig. i, 2); c, E. b. bergi Wiebes, female mandible, after Wiebes (l989a, fig. 13); d, E. pla/yseapa Wiebes, female antennal club, after Wiebes (I 989a, fig. 7); e, E. b . .bergi, female head, after Wiebes (1989a, fig. 11); f, E. soco/rensis (Mayr), female antenna (without tip), after Grandi (1928b, fig . xxiii, 2); g-h, male meso-, metano­ turn and propodeurn of: g, E. ar/ieula/a, after Wiebes (I 986b, fig. 2) and h, E. baijnathi Wiebes, after Wiebes (1989a, fig. 21).

2. Fifth to eleventh antennal segments with long sensilla chaetica (fig. a) . 3 - Fifth to eleventh antennal segments with sensilla linearia (fig. b) .... 4

3. Two ocelli (the median one obsolete). Compound eye longer than the cheek. Mandi- bular appendage a little over three times as long as wide ...... 31. E. arlicula/a - Three ocelli. Compound eye shorter than the cheek. Mandibular appendage over seven times as long as wide ...... 32. E. hilli

4. Fifth to eleventh segments of the antenna with two or three rows of sensilla (fig. d) ...... 33. E. platyseapa - Fifth to eleventh segments of the antenna with one row of sensilla ...... 5

5. Compound eye approximately half as long as the cheek, i.e., the face rather long (fig. e). Mandible with a great many ventral ridges (fig. c) ...... 34. E. bergi - Compound eye longer than the cheek. Face shorter. Mandible with less (ca. ten) ventral ridges ...... 6

6. Mandibular appendage with almost fifty rows consisting of ca. thirty denticulations ...... 35. E. peelina/a - Mandibular appendage with no more than thirty rows consisting of ca. twenty den- ticulations ...... 36. E. longiseapa

233 7. Antennal pedicel rotundate or sub-ovate in outline (fig. b) (group of E. enriquesl) . .. 8 - Antennal pedicel longer than wide, and more angular in outline (fig. f) (group of E. allo/riozoonoides) ...... II

8. Fifth to eleventh antennal segments with long sensilla chaetica ..... 37. E. baijna/hi - These segments with (long) sensiUa linearia ...... 9 9. Compound eye longer than the cheek. Mandibular appendage with rows consisting of ca. fifteen small teeth ...... 38. E. enriquesi - Compound eye as long as the cheek. Mandibular appendage with rows consisting of ca. ten teeth ...... 10 10. Antennal scape distinctly expanded dorsad (fig. b). Spiracular peritremata of the eighth urotergite small, subcircular ...... 39. E. sfuckenbergi - Antennal scape not at all expanded. Spiracular peritremata of the eighth urotergite rather large ...... 40. E. g/umosae 11. Mandible with the ventral ridges produced into distinet teeth (fig. 49, b) ...... 12 - Mandible bidentate, the ventral ridges not so distinctly produced into teeth...... 13 12. Length of the female (head, thorax and gaster) almost 2 mmo The mandibular teeth strong, four in number ...... 43. E. socofrensis - Length of the female less than 1.5 mmo The mandibular teeth less distinct...... 42. E. reflexa 13. . Spiracular peritremata of the eighth urotergite very large, occupying almost the whole width of the gaster. Valves of the ovipositor rather short (Iittle longer than the gaster) ...... 41. E. comp/oni - Spiracular peritremata of the eighth urotergite of medium size. Valves of the ovipositor almost twice as long as the gaster ...... 44. E. allo/riozoonoides

Table 9. Some characters of the males of E/isabethie//a. no. Eyel Uw Fore Spir.! pron.! head ant. tarsi prop. rest club part. thorax fused

31. E. articulata . 1/8 . 3.0 · + . 5/8 0/5 32. E. hilli ... . 116 . 2.4 · + . 113 0.9 33. E. platyscapa l iS · 2.3 · + . 1/3 1.0 34. E. b. bergi . . · 1/6 . 2.5 1/4 .0.8 35. E. pectinata. · 1/6 · 2.5 · + . 1/2 1.0 36. E. longiscapa · 118 . 3.0 · + . 1/2 . 0.5 37. E. baijnathi . 1/5 · 1.8 1/4 . 0.7 38. E. enriquesi . 1/7 . 2.5 · + . 2/5 . 0.7 39. E. stuckenbergi liS . 2.2 1/3 0.9 40. E. glumosae . 1/5 . 2.0 · +. 2/5 0.8 41. E. comptoni . . l iS . 2.0 · + . 4/5 0.8 42. E. reflexa . . . 1/6 . 2.1 · + . 1/2 1.0 43. E. socotrensis . 1/6 · 2.5 · + . 3/5 1.0 44. E. allotriozoonoides . liS . 2.4 · + . 3/5 1.0

234 31. ElisabethieUa articuIata (Joseph)

Joseph, Proc. ent. Soc. London (B) 28: 32 (1959, Blastophaga); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 338 (1986b).

The female head is a little longer than wide across the compound eyes (1.05); the longitudinal diameter of the eye is much longer than the cheek (1.75). There are two (lateral) ocelli. The antennal scape is shortly prolonged apically, so as to cover the basal third of the ovoid pedicel in axial view; the sixth to eleventh segments bear one row of long, rod-like sensilla chaetica. The mandible has two apical teeth and ca. fourteen ventral ridges; the appendage is a little over three times as long as wide, and it has ca. forty rows of 25 small crenulations. The postmarginal vein of the fore wing is twice as long as the stigma\. The spiracular peritremata of the eighth urotergite are large, ovoid. The spine of the hypo­ pygium is not very long, blunt. The total length is ca. 1.8 mm; the ovipositor valves are as long as the gaster. The colour is yellowish.

The male head is longer than wide (1.1); the eye is small (one-eighth of the length of the head). The antennal club is three times as long as wide. The pronoturn is half as long as the other terga combined. The spiracular peritremata are five­ eights of the length of the propodeum. The fore tarsi are partially fused. The total length is 1.25 mmo

The species was described from females caught in a Berlese trap (Guinea) and it was collected at light (Ivory Coast); later the host fig was found to be Ficus e/asticoides Wild. (Gabon).

32. ElisabethieUa hilli Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 122- 124 (1989a).

The female head is a little longer than wide across the compound eyes (1.1); the longitudinal diameter of the eye is three-fifths of the length of the cheek. The antennal scape is prolonged, covering more than half of the axial aspect of the long pedicel, which has an angular prominence at half length; the fifth to eleventh segments bear one row of long, flexible sensilla chaetica. The mandible has two distinct apical teeth and ca. fifteen vent ral ridges; the appendage is over seven times as long as wide, and it has ca. fifty rows of up to twelve fine crenula­ tions. The postmarginal vein of the fore wing is twice as long as the stigma\. The fore tibia has two distinct teeth in the dorso-apical combo The spiracular peri­ tremata of the eighth urotergite are of medium size, ovoid. The hypopygium has a short, blunt spine. The totallength is ca. 2.1 mm; the ovipositor valves are a little longer than the gaster (1.1). The colour is brown.

The male head is shorter than wide (0.9). The eye is one-sixtb of the length of

235 the head. The antennal club is 2.4 times as long as wide. The pronotum is almost as long as the other terga combined (0.9). The spiracular peritremata are one­ quarter to one-third of the length of the propodeum. The fore tarsi are partially fused. The total length is 1.3 mmo

The specific identity of the host fig is unknown (Uganda).

33. ElisabethieUa platyscapa Wie bes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 124-125 (I989a).

The female head is as long as wide across the compound eyes; the longitudinal diameter of the eye is shorter than the cheek (0.85). The antennal scape is rather wide, prolonged so as to cover the axial view of only the basal third of the sub-ovate pedicel, which is stalked antiaxially; the fifth to eleventh segments have two to three rows of sensilla linearia. The mandible has two apical teeth and eleven ventral ridges; the appendage is three times as long as wide, and it has ca. forty rows of up to ca. thirty crenulations. The postmarginal vein of the 1 fore wing is 1 / 2 times as long as the stigma!. The spiracular peritremata of the eighth urotergite are of medium size, ovoid. The hypopygium has a short, but acute spine. The totallength is ca. 1.6 mm; the ovipositor valves are almost twice as long as the gaster. The colour is yellow-brown.

The male is rather similar to that of E. hi/li (see table 9).

The host fig is Ficus fischeri Mildbr. & Burret (Zambia).

34a. Elisabethiella b. bergi Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 125-126 (I989a).

The female head is 1.3 times as long as wide across the compound eyes; the longitudinal diameter of the eye is half as long as the cheek. The antennal scape is prolonged so as to cover the greater part of the pedicel in axial view; the pedicel is twice as long as wide; the fifth to eleventh segments have one row of sensilla linearia. The mandible has one distinct apical tooth and ca. thirty ventral 1 ridges, like a was hing board; the appendage is 3 / 2 times as long as wide, and it has ca. fifty rows of ca. 35 fine crenulations. The postmarginal vein of the fore 1 wing is approximately 1 / 2 times as long as the stigma!. The antiaxial tooth of the hind tibia is tricuspidate. The spiracular peritremata of the eighth urotergite are rather large, ovoid. The hypopygium has a short, but acute spine. The total length is ca. 1.75 mm; the head (0.55 mm) is longer than the thorax (0.6); the valves of the ovipositor are somewhat longer than the gaster (1.15). The colour is brown.

The male head is al most as long as wide (0.95). The eye is one-sixth of the length

236 of the head. The antennal scape is 2'/2 times as long as wide. The pronotum is distinctly shorter than the other terga combined (0.8). The spiracular peritre­ mata are one-quarter of the length of the propodeurn. The fore tarsi are free. The total length is 1.1 mmo

The host fig is Ficus trichopoda Baker (Ivory Coast).

34b. Elisabethiella bergi breviceps Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 126 (1989a).

The female head is only one-tenth longer than wide across the compound eyes; the longitudinal diameter of the eye is more than half as long as the cheek (0.6). The antennal pedicel is wider than in the nominate subspecies and the prolonga­ tion of the scape covers only a third of it in axial view. The mandible has ca. twenty ventral ridges; the appendage has ca. 45 rows of ca. thirty fine crenula­ tions.

The male is similar to that of E. b. bergi.

The host fig, also of this subspecies, is Ficus trichopoda Baker (Madagascar, Zambia, Malawi).

35. Elisabethiella pectinata (Joseph) Joseph, Proc. ent. Soc. London (B) 28: 33- 34 (1959, Blastophaga); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 126--127 (1989a).

The female head is a little longer than wide across the compound eyes (1.05); the longitudinal diameter of the eye is 1'/4 times as long as the cheek. The an­ ten na I scape is prolonged to the apex of the pedicel, which is twice as long as wide; the fifth to eleventh segments bear one row of rod-like sensilla chaetica, projecting beyond the distal margin of the segment for the greater part of their length. The mandible has two distinct apical teeth and ca. twelve ventral ridges; the appendage is ca. five times as long as wide, and it has almost fifty rows of ca. twenty fine crenulations. The postmarginal vein of the fore wing is over twice as long as the stigmal (2.1). The spiracular peritremata of the eighth urotergite are large, subcircular. The hypopygium has a sort, acute spine. The total length is ca. 2.1 mm; the valves of the ovipositor are a little longer than the gaster (1.1). The colour is yellow-brown.

The male he ad is as long as wide. The eye is one-sixth of the length of the head. 1 The antennal club is 2 / 2 times as long as wide. The pronotum is as long as the other terga combined. The spiracular peritremata are half as long as the pro­ podeum. The fore tarsi are partially fused. The total length is 1.4 mmo

237 The species was described from females caught in a Berlese trap (Guinea); the host fig is Ficus calyptrata Vahl (lvory Coast).

36. ElisabethieUa longiscapa Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 339-340 (1986b).

The female head is as long as wide across the compound eyes; the longitudinal diameter of the eye is 1.7 times the length of the cheek. The prolongation of the antennal scape almost covers the pedicel in axial view; the pedicel is rather wide; the fifth to eleventh segments bear one row of rod-like sensilla chaetica, project­ ing beyond the distal margin of the segment for approximately one-third of their length. The mandible has one apical tooth and a smaller subapical, and eight ventral ridges; the appendage is four times as long as wide, and it has ca. thirty rows of ca. twenty fine crenulations. The postmarginal vein of the fore wing is 1 1/ 2 times as long as the stigma!. The spiracular peritremata of the eighth uro­ tergite are of medium size, subcircular. The hypopygium has an acute spine, which is not very long. The totallength is ca. 1.4 mm; the valves of the ovipositor are 1.4 times as long as the gaster. The colour is light brown.

The male is much like that of E. articulata, but the spiracular peritremata are half as long as the propodeum. The total length is 1.1 mm.

The host fig is Ficus burretiana Hutch. (Gabon).

37. ElisabethieUa baijnathi Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 127, 129 (1989a).

The female head is distinctly shorter than wide across the compound eyes (0.85); the longitudinal diameter of the eye is almost one-third longer than the cheek (1.3). The antennal scape is three times as long as wide, not prolonged apically; the pedicel is longer than wide (1.15); the fifth to eleventh segments have one row of long, ftexible sensilJa chaetica. The mandible has two teeth and four ventral ridges; the appendage is almost six times as long as wide, and it bears fifteen ventral rows of mostly only two hook-like teeth. The postmarginal vein of the fore wing is very short (one-eighth of the stigmaI). The fore tibia has two teeth in the dorso-apical comb; the antiaxial tooth of the hind tibia is tricu­ spidate. The spiracular peritremata of the eighth urotergite are of medium size, ovoid. The spine of the hypopygium is long and acute. The total length is ca. 1 1.4 rnm; the valves of the ovipositor are 1 / 2 times as long as the gaster. The colour is light brown.

The male head is shorter than wide (0.9). The eye is one-fifth of the length of the head. The antennal club is not quite twice as long as wide (1.8). The pro-

238 notum is two-thirds (0.7) of the length of the other terga combined. The spira­ cular peritremata are one-quarter of the length of the propodeum. The fore tarsi are free. The total length is 0.9 mmo

The host fig is Ficus burtt-davyi Hutch. (Sou th Africa), the reproductive biology of which and its Chalcid symbiosis was described by Baijnath & Ramcharun (1988) and Compton & Nefdt (1990).

38. ElisabethieUa enriquesi (Grandi) Grandi, BoU. Lab. Zool. Portici 10: 129- 133 (1916a, Blastophaga); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 130 (1989a).

The female head is shorter than wide across the compound eyes (0.85); the longitudinal diameter of the eye is a little longer than the cheek (1.1). The apex of the antennal scape is not prolonged; the pedicel is as wide as long, rotundate; the fifth to eleventh segments bear one row of long sensilla linearia. The mand i­ bie has two teeth and about ten ventral ridges; the appendage is four times as long as wide, and it bears ca. twenty ventral rows of up to fifteen small teeth. The postmarginal vein of the fore wing is a bit longer than the stigmal (1.25). The antiaxial tooth of the hind tibia is tricuspidate. The spiracular peritremata of the eighth urotergite are of medium size, ovoid. The spine of the hypopygium is not very long, blunt. The totallength is ca. 1.3 mm; the valves of the ovipositor are twice as long as the gaster. The colour is blackish brown.

The male head is slightly longer than wide (1.05). The eye is one-seventh of the 1 length of the head. The antennal club is 2 / 2 times as long as wide. The pronotum is two-thirds (0.7) of the length of the other terga combined. The spiracular peritremata are two-fifths of the length of the propodeurn. The fore tarsi are partially fused. The total length is 1.1 mmo

The species was described from an unknown fig (Angola); the host fig is now known to be Ficus ilicina (Sonder) Miq. (South West Africa).

39. ElisabethieUa stuckenbergi (Grandi) Grandi, BoU. Ist. Ent. Univ. Bologna 21 : 85--87 (1955, Blastophaga); Wiebes in Boueek et al ., Tijdschr. Ent. 124: 165--167 (1981).

The female head is as long as wide across the compound eyes; the eye is as long as the cheek. The antennal scape is expanded dorsad, but not prolonged, three times as long as the rotundate pedicel; the fifth to eleventh segments have one row of long sensilla linearia. The mandible has two apical teeth and five ventral ridges; the appendage is four times as wide as long, and it bears ca. twenty rows of ten to twelve small teeth. The postmarginal vein of the fore wing is twice as long as the stigma!. The spiracular peritremata of the eighth urotergite are smalI,

239 subcircular. The hypopygium has a rather short spine with a blunt apex. The total length is ca. 1.4 mm; the valves of the ovipositor are two times as long as the gaster, as long as the body. The colour is dark brown to blackish.

The male head is as long as wide. The eye is one-fifth of the length of the head. The antennal club is more than two times as long as wide (2.2). The pronotum is almost as long as the other terga combined (0.9). The spiracular peritremata are one-third of the length of the propodeum. The fore tarsi are free . The total length is 0.9 mmo

The host fig is Ficus thonningii BI. (Cameroon, Kenya, Zambia, Zimbabwe, South Africa) and it was recorded from a putative hybrid of F. thonningii and F. natalensis Hochst. (South Africa, Ramcharun et al., 1990). From F. thonningii also two species of Alfonsiella were recorded, viz., no. 53, A. brongersmai (Zam­ bia, Zimbabwe) and no. 54, A. longiscapa (Zambia).

40. ElisabethieUa glumosae Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 131- 133 (l989a).

The female head is almost as long as wide across the compound eyes (0.95); the longitudinal diameter of the eye is as long as the cheek. The antennal scape is not prolonged, slender, 2'/2 times as long as the subovate pedicel; the fifth to eleventh segments have one row of long sensilla linearia. The mandible has one apical tooth and two subapicals and six or seven ventral ridges; the appendage is five times as long as wide, and it has up to seventeen rows of ca. ten small teeth. The postmarginal vein of the fore wing is almost 1'/2times as long as the stigmal (1.35). The peritremata of the eighth urotergite are rather large, ovoid. The hypopygium has a spine that is short and not very acute. The total length is ca. 1.5 mm; the valves of the ovipositor are 1'/2 times as long as the gaster. The colour is brown.

The male head is longer than wide (l.l5). The eye is one-fifth of the length of the head. The antennal club is two times as long as wide. The pronotum is distinctly shorter than the other terga combined (0.8). The spiracular peritre­ mata are two-fifths of the length of the propodeurn. The fore tarsi are partially fused. The totallength is 0.9 mmo

The host fig is Ficus glumosa Del. (Carneroon, Ethiopia, Tanganyika, Zambia).

41. ElisabethieUa comptoni Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 133 (1989a).

The female head is shorter than wide across the compound eyes (0.9); the longi-

240 tudinal diameter of the eye is longer than the cheek (1.15). The antennal scape 1 is slender, 2 / 2 times as long as wide, only very little prolonged apically; the pedicel is angular in shape, its length is one-third of that of the scape; the fifth to eleventh segments have one row of long sensilla linearia. The mandible has two apical teeth and eight ventral ridges; the appendage is five times as long as wide, and it bears almost twenty rows of up to ni ne small teeth. The postmar­ ginal vein of the fore wing is twice as long as the stigma!. The spiracular peri­ tremata of the eigthth urotergite are very large, occupying almost the whole width of the gaster. The spine of the hypopygium is not very long, blunt. The totallength is ca. 2.1 mm; the valves of the ovipositor are somewhat longer than the gaster (1.1). The colour is brown.

The male head is slightly longer than wide (1.05). The eye is one-fifth of the length of the head. The antennal club is two times as long as wide. The pronotum is distinctly shorter than the other terga combined (0.8). Tbe spiracular peri­ tremata are more than four-fifths of tbe length of the propodeum. The fore tarsi are partially fused. The total length is 1.5 mmo

The host fig is Ficus abutilifolia (Miq.) Miq. (Malawi). From this species of fig also a species of Nigeriella was recorded, i.e., no. 46, N. fusciceps (Nigeria).

42. Elisabethiella reflexa Wie bes Wiebes, Zool. Meded. Leiden 49: 226-229 (1975).

The female head is longer than wide across the compound eyes (1.2); the longitudinal diameter of the eye is two-thirds of the length of the cheek. The antennal scape is not prolonged apically, it is twice as long as wide and also twice as long as the angular pedicel; the fifth to eleventh segments have one row of long sensilla linearia, which distinctly project beyond the distal margin of the segment. The mandible has two large teeth and six ventral ridges (some a bit 1 produced); the appendage is 6 / 2 times as long as wide, and it bears up to 25 ventral rows of up to eight sm all teeth. The postmarginal vein of the fore wing is one-third longer than the stigmal (1.3). The spiracular peritremata of the eighth urotergite are smalI, ovoid. The spine of the hypopygium is rather short, blunt. The totallength is ca. 1.0-1.3 mm (the larger dimensions are from speci­ mens collected in Réunion, and these have the larger number of rows and teeth 1 on the mandibular appendage); the valves of the ovipositor are almost 1/ 2 times as long as the gaster (1.4). The colour is brown.

The male head is as long as wide. The eye is one-sixth of the length of the bead. The antennal club is slightly more than two times as long as wide. The pronotum is almost as long as the ot her terga combined. The spiracular peritremata are half as long as the propodeurn. The fore tarsi are partially fused. The totallength is 0.8 mmo

241 The host fig is Ficus reflexa Thunb. (Aldabra Is!., Réunion, Madagascar).

43. ElisabethieUa socotrensis (Mayr)

Mayr, Verh. zool.-bot. Ges. Wien 35: 175--176 (1885, Blastophaga); Grandi, Boll. Lab. Ent. Bo- 10gna 1: 159- 163 (1928b). Synonym: Blastophaga dyscritus Waterston, Trans. ent. Soc. London (1921): 417-418 (1921).

The female head is little longer than wide across the compund eyes (1.15); the longitudinal diameter of the eye is as long as the cheek. The antennal scape is 1 almost 2 / 2 times as long as wide (2.4) and li2 times as long as the angular pedicel; the fifth to eleventh segments have one row of long sensilla linaria. The mandible has an apical tooth and three of the seven ventral ridges are produced into teeth; the appendage is five times as long as wide, hearing ca. slightly over twenty rows (21-23) of eight small teeth. The postmarginal vein of the fore wing is 1.65 of the length of the stigma!. The spiracular peritremata of the eighth urotergite are large, ovoid. The spine of the hypopygium is not very long, acute. The total 1 length is ca. 2 mm; the valves of the ovipositor are ca. 1/ 2 times as long as the gaster. The colour is brown.

The male head is as long as wide. The eye is one-sixth of the length of the head. 1 The antennal club is 2 / 2 times as long as wide. The pronotum is as long as the other terga combined. The spiracular peritremata are three-fifths of the length of the propodeum. The fore tarsi are partially fused. The totallength is 1.2 mmo

The original host fig probably was Ficus vasta Forssk. (Socotra), from which it was also recorded in Ethiopia; the species was, however, also recorded from Ficus wakefieldii Hutch. (Kenya, Zambia), and from Ficus 'natalensis', which may be only an indication of the group (South Africa; see Ramcharun et a!., 1990). It is also known from Uganda.

44. ElisabethieUa aUotriozoonoides (Grand i)

Grandi, Boll. Lab. Zool. Portici 10: 126, 128 (1916a, Blastophaga); Grandi, Boll. Lab. ent. Hal. 48: 4-15 (1917).

The female head is a little longer than wide across the compound eyes (1 .05); the longitudinal diameter of the eye is shorter than the cheek (0.85). The an­ 1 tennal scape is two times as long as wide, not prolonged apically, 2 / 2 times as long as the angular pedicel; the fifth to eleventh segments bear one row of long sensilla linearia. The mandible has two apical teeth and ten to twelve ventral ridges (not produced into teeth); the appendage is five times as long as wide, and it hears 25-28 rows of ca. twelve small teeth. The postmarginal vein of the fore 1 wing is 1 /2 times as long as the stigma!. The spiracular peritremata of the eighth urotergite are of medium size, ovoid. The spine of the hypopygium is not very

242 long, acute. The totallength is ca. 1.6 mm; the valves of the ovipositor are almost twice as long as the gaster. The colour is brown.

The male is rather similar to that of E. socotrensis (see table 9).

The species was described from an unnamed fig collected in Eritrea, and later it was recorded from Ficus 'natalensis' (Uganda), probably referring to a group rather than to a species of fig.

NigerieUa Wiebes (fig. 63) Wiebes, Zool. Meded. Leiden. 48: 36 (1974a); Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 212-213 (1990).

The female head is about as long as wide across the compound eyes, or clearly longer (to 1.5); the longitudinal diameter of the eye is distinctly shorter than the cheek (0.6-0.75). There are three ocelli. The antenna consists of eleven free seg­ ments; the fifth to eleventh segments bear one row of long sensilla linearia and in one species, also many long, flexible sensilla chaetica; the three apical seg­ ments are shaped so as to form a club. The mandible has one apical tooth and

Fig. 63. Nigeriella letouzeyi Wiebes, male, and N. fusciceps Wiebes, female.

243 a numher of ventral lamellae, some of which may be extended into teeth. The labium and maxillae are simpIe, they hear apical and subapical setae. The thorax has large pollen pockets, and the fore coxae have a corbicula. The venation of the fore wing is complete. The fore tibia has two dorso-apical teeth and one ventral; the hind tibia has a tricuspidate antiaxial tooth and a slender axia\. The stigmatal peritremata of the eighth urotergite are large, ovoid. The spine of the hypopygium is of medium length. The totallength (head, thorax and gaster) is ca. 1.5 mm (1.2-1.7).

The male head is as long as wide, or slightly longer (1.1); the eyes are large: one-quarter of the length of the head. The antennae are borne in separate sock­ ets. The mesonotum and metanotum are fused as is, incompletely, also the pro­ podeum. All tarsi are pentamerous. The genitalia are simpIe. The total length (head and thorax) is ca. 1 mm (0.9- 1.2). The colour is yel­ lowish brown, the head may he darker.

There are four species, pollinators of some figs of the Galoglychïa subsections Platyphyllae (three species) and Crassicostae (one). The species can he identified by the following key.

KEY TO THE SPECIES (fig. 64)

I. Female antennal flagellum with long, flexible sensilla chaetica, next to a few sensilla linearia (fig. a). Male head with dorsal prominences next to the antennal toruli (fig. b) ...... 46. N. fusciceps - Female antennal flagellum with sensiUa linearia only. Male head simpie...... 2

2. Female mandible with ca. twenty ventral ridges, the appendage with 25 lamellae that, distally, become distinctly denticulate. Male anten na with two ane\li (fig. d) ...... 48. N. /etouzeyi - Female mandible with ca. five ventral ridges, the appendage with less than twenty lamellae, each obliterated medially and produced into teeth at the sides. Male antenna with one or three anelli ...... 3

3. Width of the female mandibular appendage 0.4 of its length, with eleven ventral lamel- lae (fig. f). Male antenna with three anelli (as in fig. c) ...... 47. N. avico/a - Female mandibular appendage more e\ongate (widthllength 0.3), with seventeen ven- tral lamellae. Male antenna with one anellus (fig. e) ...... 45. N. excavala

45. NigerieUa excavata Compton

Compton in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 213- 215 (1990).

The female head is clearly longer than wide across the compound eyes (l.5); the longitudinal diameter of the eye is much shorter than the cheek (0.6). The ventral projection of the antennal scape is prominent; the appendage of the third seg­ ment is hardly developed; the fifth to eleventh segments bear one row of long sensilla linearia. The man di bie has four complete ventrallamellae, two of which

244 Fig. 64. Details of Nigeriella and Alfonsiella. a- f, Nigeriella. a--{;, N. fusciceps Wiebes: a, base of female antenna, b, male head, and c, male antenna, after Wiebes (1974a, figs. 12, 25, and 22, respectively); d, N. letouzeyi Wiebes, male antenna, after Wiebes (1974a, fig . 29); e, N. excavata Compton, male anten na, after Wiebes & Compton (1990, fig. 22); f, N. avicola Wiebes, femaIe mandible and appendage, after Wiebes (1975, fig. 15). g-h, Alfonsiella, femaIe antennaI scape of: g, A. longiscapa Joseph, and h, A. brollgersmai Wiebes, after Wiebes (1972b, figs. 25 and 14, respectively).

are extended into teeth; the width of the appendage is 0.3 of its length, it has seventeen ventral lamellae, each obliterated medially and developed into teeth 1 at the sides. The postmarginal vein of the fore wing is 1/ 2 times as long as the stigma!. The dorso-apical teeth of tJie fore tibia are elongate. The spine of the hypopygium is acute. The totallength is ca. 1.5 mm; the valves of the ovipositor are as long as the gaster. The colour is black; the scape and the legs are paler.

The male head is slightly longer than wide (1.1), the cheeks project considerably in front of the epistomal margin. Tlie antenna is indistinctly four-segmented, consisting of a scape that is almost three times as long as its widest part, a pedicel and one anellus in length-ratio 2: I, îogether as long as the funnel-shaped club. The mandible is elongate and falcate. The pronotum is wider than long (1 .35). The propodeal spiracles are less than half the leng th of the lateral margins of the propodeum. The total length is ca. I mmo

245 The host tig is Ficus tettensis Hutch. (South Africa).

46. NigerieUa fusciceps Wiebes Wiebes, Zool. Meded. Leiden 48: 36-39 (1974a).

The female head is as long as wide across the compound eyes; the longitudinal diameter of the eye is three-quarters of the length of the cheek. Tbe ventral projection of the antennal scape is very prominent; the pedicel is sub-circular and it has its axial surfaee expanded; the appendage of the third segment is long and slender; the fifth to eleventh segments bear three sensilla linearia and many long, flexible sensilla chaetica. The mandible has ca. ten ventral ridges; the width of the appendage is 0.3 of its length, it has ca. twenty ventral ridges, each consisting 1 of two teeth. The postmarginal vein of the fore wing is more than 1/ 2 times as long as the stigmal (1.6). The dorso-apical comb of the fore tibia has two short teeth and a smaller one in between. The spine of the hypopygium is blunt at the apex. The total length is ca. 1.7 mm; the valves of the ovipositor are as long as the gaster. The colour is blackish brown.

The male head is as long as wide, with stout dorsal prominenees next to the antennal toruli. The antenna consists of a large scape, narrowly stalked, a pe­ dieel that is one-third ofthe length ofthe scape, three anelli, and a club consisting 1 of three segments. The mandible is elongate and falcate. The pronotum is 1 /2 times as wide as long. The propodeal spiracles are circular, one-quarter of the length of the propodeum. The total length is ca. 1.2 mmo

Tbe host tig is Ficus abutilifolia (Miq.) Miq. (Nigeria). Tbis species of tig was also recorded as the host of Elisabethiel/a comptoni (no. 41).

47. NigerieUa avicola Wiebes Wiebes, Zool. Meded. Leiden 49: 229-231 (1975).

Tbe female head is longer than wide across the compound eyes (1.2); the longi­ tudinal diameter of the eye is distinctly shorter than the cheek (0.7). Tbe ventral projection of the antennal scape is prominent; the appendage of the third seg­ ment is blunt; the fifth to eleventh segments bear one row of sensilla linearia. Tbe mandible has six ventral lamellae; the width of the appendage is 0.4 of its length, it bears eleven ventral ridges axially produeed into a distinct tooth. The postmarginal vein of the fore wing is shorter than the stigmal (0.85). Tbe fore tibia bears two short dorsal teeth. Tbe spine of the hypopygium is blunt at the apex. The totallength is ca. 1.5 mm; the valves of the ovipositor are slightly longer than the gaster (1.2). Tbe colour is yellowish brown.

The male head is about as long as wide. Tbe antenna consists of scape, pedieel,

246 three anelli, and a two-segmented club. The mandible is short and rather robust. The pronotum is about as wide as long. The propodeal spiracles are circular, ca. one-third of the length of the propodeurn. The total length is ca. 0.9 mmo

The host fig is Ficus rubra Vahl (Aldabra Is1. , Réunion).

48. NigerieUa letouzeyi Wiebes Wiebes, Zool. Meded. Leiden 48: 39-41 (1974a); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 338 (1986b).

The female head is longer than wide across the compound eyes (1.1); the longi­ tudinal diameter of the eye is distinctly shorter than the cheek (0.7). The ventral projection of the antennal scape is not very prominent; the appendage of the third segment is short; the fifth to eleventh segments bear one row of sensilla linearia. The mandible has ca. twenty ventral lamellae; the width of the ap­ pendage is 0.2 of its length, it has ca. 25 ventral lamellae that, distally, become distinctly denticulate. The postmarginal vein of the fore wing is distinctly longer than the stigmal (1.3). The dorsal crest of the fore tibia consist of two, widely spaced teeth. The spine of the hypopygium is blunt at the apex. The totallength 1 is 1.2 mm; the valves of the ovipositor are 1/ 2 times as long as the gaster. The colour is brown.

The male head is as long as wide. The antenna consists of a short scape, pedicel, two anelli, and a three-segrnented club. The mandible is short and rather robust. The pronotum is about as wide as long. The propodeal spiracles are sublateral in position, ca. one-quarter of the length of the propodeurn. The total length is ca. 0.9 mmo

The host fig is Ficus pseudomangifera Hutch. (Carneroon, Gabon).

AlfonsieUa Waterston (fig. 65)

Waterston, Ent. monthly Mag. (3) 6: 198 (1920b); Wiebes, Zool. Meded. Leiden 47: 321 ~330 (1972b); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 91 : 429-436 (1988).

1 The fe male he ad is approximately as long, or 1 / 2 times as long as wide across the compound eyes; the longitudinal diameter of the eye is 1.3 to many (up to five) times as long as the cheek. There are no oceUi, or only the laterals are present. The antenna has eleven segments; the scape is with or without an apical prolongation; the third segment is simpie, without an apical appendage; the funicular sensiJla chaetica are long and flexible. Essentially, the mandible has two apical teeth (the subapical one may be inconspicuous); the mandibular ap­ pendage has hook-like ridges, or rows of fine crenulations. The labium and maxillae are simpie. The thorax has pollen pockets, and the fore coxae have a corbicula. The

247 Fig. 65. Alfonsiella brongersmai Wiebes, male and female, after Wiebes in Boucek et al. (1981, figs. 13 and 14, respectively). vena ti on of the fore wing is complete: usually, the postmarginal vein is half as long as the stigmal, but it is al most as long in one species, and twice as long in another. The fore tibia has two dorso-apical teeth and one ventral; the bind tibia has a bidentate antiaxial tooth and a slender axial. The spiracular peritremata of the eighth urotergite are small and subcircular. The hypopygium gradually narrows in an acute spine. The totallength (head, thorax and gaster) is ca. 1.3-1.8 mm; the valves of the ovipositor are as long as the gaster, in one species longer (1.25). The colour is light brown.

The male head is as long as its maximum width in one species, but usually longer (l.15-1.25); the head is rather wide in front Gust behind the eyes, smaller than the maximum width), or the head is more distinctly narrowing frontad. The compound eyes are prominent. The antennae are borne in separate sockets; they consist of a scape (slender in some, more robust in other species), a pedicel and three subquadrate anelli, and a divided club. The mandible is characteristically faJcate, with one tooth and one gland. The thorax consists of two subequal, dorsal sclerites, viz., the pronoturn, and the fused meso-, metanotum and propodeurn. The propodeum is indicated in two ear-like lobes, with large spiracular peritremata. The fore tibia has two

248 dorso-apical hook-like teeth and one ventral (or indistinctly two); the tarsus is oligomerous. The rnid leg is strikingly darker than the other legs and the femur is setose; the tarsus is pentamerous, as in the hind tarsus. The hind tibia has two ventral spurs. The gaster is short; the genitalia lack claspers. The total length (head and thorax) is ca. 1.0-1.2 mmo The colour is yellow­ brown, the mid leg is darker.

There are seven species, associated with some figs of the Galoglychia subsections Platyphyllae and Chlamydodorae. The specificity of the relationship has been doubted for some time, and in some species still is. For the identification the following key may be used.

KEY TO THE SPECIES (females; for the males, see tab Ie 10) (fig. 64)

I. Three ocelli present. Head distinctly shorter than wide (0.85). A species of E/isabe- thiella that resembles Alfonsiella ...... 37. E. baijnathi - No ocelli, or only the two laterals present. Head almost as long as wide, or longer (0.95-1.5) ...... 2

2. Head distinctly longer than wide (almost 1.5). Antennal scape with an apica1 projection (fig. g). Mandibular appendage with 35 - 40 hook-like ridges and fine crenulations . . 3 - Head almost as long as wide, or scarcely longer than wide (0.95 - 1.1). Antennal scape without the projection (fig. h). Mandibular appendage with fifteen to twenty hook-like ridges ...... 4 3. Longitudinal diameter of the eye 1.3 times the length of the cheek. Apica1 projection of the antennal scape wide; the pedicel rather slender: almost three times as long as wide. Mandibular appendage with about forty ridges ...... 55. A. nata/ensis - Longitudinal diameter of the eye 1.5 times as long as the cheek. Apical projection of the antennal scape narrower; the pedicel wider: about two times as long as wide. Man­ dibular appendage with about 35 ridges ...... 54. A. /ongiscapa 4. Lateral ocelli present ...... 5 - No ocelli present. The longitudinal diameter of the eye three or four times as long as the cheek ...... 7 5. Longitudinal diameter of the eye two or three times as long as the cheek ...... 6 - Longitudinal diameter of the eye five times as long as wide. Mandibular appendage with seventeen ridges ...... 50. A. bergi 6. Longitudinal diameter of the eye two times as long as the cheek. Mandibular ap- pendage with twenty ridges ...... 53. A. brongersmai - Longitudinal diameter of the eye three times as long as the cheek. Mandibular ap- pendage with fourteen ridges ...... 49. A. binghami 7. Mandibular appendage with about 15 - 18 hook-like ridges (consisting of one tooth) ...... 51. A. fimbriata - Mandibular appendage with more than one tooth (or crenulations) in the (18) ridges ...... 52. A. micha/oudi

49. AIfonsieUa binghami Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 91: 432-434 (1988).

249 The female head is slightly longer than wide across the compound eyes (l.05); the eye is almost three times as long as the cheek (2.75). There are two lateral ocelli. The slender antennal scape is not prolonged. The mandible has six ventral ridges; the appendage has fourteen hook-like ridges. The total length is ca. 1.8 mm; the ovipositor valves are longer than the gaster (1.25).

The male head is longer than wide (1.25), rather wide in front. The antennal scape is si end er.

The host fig is Ficus stuhlmannii Warb. (Tanganyika, Zambia, South Africa).

50. AlfonsieUa bergi Wie bes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 91 : 432 (1988).

The female head is approximately as long as wide across the compound eyes; the eye is five times as long as the cheek. There are two lateral ocelli. The antennal scape is slender, without prolongation. The mandible has six ventral ridges; the appendage has seventeen hook-like ridges. The total length is ca. 1.4 mm; the valves of the ovipositor are as long as the gaster.

The male head is rather short, approximately as long as wide, rather wide in front. The antennal scape is slender.

The host fig is Ficus nigropunctata Mildbr. & Burret (Zambia).

51. AlfonsieUa funbriata Waterston

Waterston, Ent. monthly Mag. (3) 6: 198-200 (1920b); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 91 : 434--435 (1988).

Table 10. Some characters of the males of Alfonsiella. no. Length/ Head Antennal max. scape width narrow wider head in In front front

49. A. binghami ...... 1.25 · + . .. · slender 50. A. bergi ...... 1.00 · + .. · slender 51. A. fimbriata ...... 1.1 5 · + .. . · slender 52. A. michaloudi ...... 1.25 · +. · robust 53. A. brongersmai ...... 1.15 · + . . . · robust 54. A. longiscapa ...... 1.25 · + . · robust 55. A. natalensis ...... 1.25 · + . · robust

250 The female head is approximately as long as wide across the compound eyes; 1 the eye is 3 / 2 times as long as the cheek. There are no ocelli. The antennal scape is slender, not prolonged. The mandible has eight ventral ridges; the appendage has eighteen hook-like ridges. The total length is ca. 1.3 mm; the valves of the ovipositor are slightly longer than the gaster (1.1).

The male head is little longer than wide (1.15), rather wide in front. The antennal scape is slender.

The species was described from Tanganyika, and later it was recorded from Guinea (Berlese trap) and Ivory Coast. The host fig is Ficus natalensis subsp. leprieurii (Miq.) Berg (Ivory Coast, Cameroon, Gabon), but it was also rea red from the sycones of Ficus kamerunensis Mildbr. & BUITet (lvory Coast , Gabon).

52. AlfonsieUa michaloudi Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 91 : 435-436 (1988).

The female head is little shorter than wide across the compound eyes (0.95); the eye is three times as long as the cheek. There are no ocelli. The anten na has a slender scape, without prolongation. The mandible has six ventral ridges; the appendage has eighteen ridges, most of which consist of more than one (up to three) crenulations. The total length is ca. 1.4 mm; the valves of the ovipositor are as long as the gaster.

The male head is longer than wide (1.25), narrowing frontad. The antennal scape is robust.

There are two host figs, viz., Ficus craterostoma Mildbr. & BUITet and Ficus l. lingua De Wild. & Durand (both, Gabon).

53. AlfonsieUa brongersmai Wie bes Wiebes, Zool. Meded. Leiden 47: 326 (\972b).

The female head is a bit longer than wide (1.1); the eye is twice as long as the cheek. There are two lateral ocelli. The antennal scape is slender, without pro­ longation. The mandible has eight ventral ridges; the appendage has twenty hook-like ridges. The postmarginal vein is twice as long as the stigma\. The total length is ca. 1.8 mm; the valves of the ovipositor are as long as the gaster.

The male head is little longer than wide (1.15), rather wide in front. The antennal scape is robust.

The host fig is Ficus thonningii B\. (Zambia, Zimbabwe, Kenya) and it was

251 caught at light in Nigeria. From F. thonningii also Elisabethiel/a stuckenbergi (no. 39) and Alfonsiel/a longiscapa (no. 54) were recorded.

54. Alfonsiella longiscapa J oseph Joseph, Proc. ent. Soc. London (8) 28: 30 (1959); Wiebes, Zool. Meded. Leiden 47: 328--330 (l972b).

The female head is distinctly longer than wide (1.5); the eye is 1'/2-2 times as long as the cheek. There are two lateral ocelli. The antennal scape is prolonged. The mandible has ca. ten ventral ridges; the appendage has ca. 35 rows of many small crenulations. The postmarginal vein is two-thirds of the length of the stigma\. The total length is ca. 1.4 mm; the ovipositor valves are almost as long as the gaster (0.9).

The male head is longer than wide (1.25), distinctly narrowmg frontad. The antennal scape is robust.

The host-relations are not weil understood. A. longiscapa was recorded from Ficus n. natalensis Hochst. (Kenya, Malawi, Zambia) and Ficus thonningii BI. (Zambia; from this fig also no. 39, Elisabethiel/a stuckenbergi, and no. 53, Alfon­ siella brongersmai, were recorded). The species was caught in Guinea (Berlese trap), Ivory Coast, and Nigeria.

55. Alfonsiella natalensis Wiebes Wiebes, Zool. Meded. Leiden 47: 326-327 (l972b).

The female head is almost } '/2times as long as wide across the compound eyes; the eye is longer than the cheek (1.3). There are two lateral ocelli. The antennal scape is very wide, prolonged apically. The mandible is much like that of A. longiscapa; the appendage has ca. forty ventral ridges of small crenulations. The postmarginal vein of the fore wing is as long as the stigma I. The total length is ca. 2 mm; the valves of the ovipositor are as long as the gaster.

The male head is longer than wide (1.25), narrowing frontad. The antennal scape is robust.

The bost relations are not weU understood. Tbe species was recorded from Ficus 'natalensis' (Uganda, Kenya), but this might indicate a group of figs rather than a species.

Paragaon Joseph (fig. 66) Joseph, Proc. ent. Soc. London (8) 28: 36 (1959).

252 Fig. 66. Paragaon josephi Wiebes, male and female.

The female head is longer than wide across the compound eyes (1.25); the lon­ gitudinal diameter of the eye is longer than the cheek in one species (1.3), but much shorter in the other (0.3). There are three ocelli. The antenna has eleven segments; the scape has a st rong ventral prominence and a hyaline dorsal ridge; 1 the pedicel is 1/ 2 times as long as wide; the third segment has a long seta on the apex of the short, blunt appendage; the fifth to tenth sements have one row of long sensilla chaetica, the eleventh has two. The mandible has one apical tooth; its appendage is long and bears ca. twenty ventral rows of teeth or crenulations. The labium and maxillae are simpie. The thorax has pollen pockets and the fore coxae have a corbicula. The vena­ tion of the fore wing is complete. The fore tibia has two teeth in the dorso-apical comb; the hind tibia has two bidentate teeth. The stigmatal peritremata of the eighth urotergite are very smalI. The hypopygium has a long spine.

253 The total length (head, thorax and gaster) is ca. 1-1.3 mm; the valves of the ovipositor are 1'/2times as long as the gaster. The colour is brown.

The male is known for P josephi only.

There are two species, for one of which the host-fig is known: it belongs to the Galoglychia-subsection Crassicostae.

KEY TO THE SPECIES (females only)

I. Compound eye 1.3 times as long as the cheek; the rows on the mandibular appendage consist of two teeth. Postmarginal vein of the fore wing about half as long as tbe stigmal ...... 56. P perplexum - Compound eye one-third of the length of the cheek; the rows on tbe mandibular ap­ pendage con sist of five to seven fine crenulations. Postmarginal vein of the fore wing one-third of the lengtb of the stigmal ...... 57. P josephi

56. Paragaon perplexum Joseph Joseph, Proc. ent. Soc. London (B) 28: 36 (1959); Wiebes, Ent. Ber. Amst. 36: 127 (1976).

The female head has a compound eye that is longer than the cheek (1.3). The mandibular appendage is about ten times as long as wide and its ventral ridges consist of two teeth. The postmarginal vein of the fore wing fades out at half the length of the stigma!. The total length is ca. 1.35 mmo

The male is not known.

The species was collected in a Berlese-trap (Guinea) and at light (Nigeria).

57. Paragaon josephi Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 340-342 (1986b).

The female compound eye is much shorter than the cheek (0.3). The mandibular appendage is seven times as long as wide and its ventral ridges consist of five to seven fine crenulations. The length of the postmarginal vein of the fore ~ing is one-third of that of the stigma!. The totallength is ca. I mmo

The male head is longer than wide (1.25). The antennae are borne in separate sockets. The eyes are smalI, not visible from above, situated laterad and distad of the antennal sockets. The antenna consists of four segments, viz., a large scape, not quite twice as long as wide (1.7) and four times as long as the pedicel, one anellus, slightly wider than long, and a club. The mandibles are stout and blunt; the labium and maxillae are reduced to one wide lobe. The thorax has the pronotum about as long as the combined meso- and me­ tanotum and propodeurn; the sub-circular peritremata of the propodeal stigmata

254 occupy not quite half the length of the propodeum (0.4). The fore tibia has two large dorsal and two ventral apical teeth, the tarsus is bimerous; the mid tarsus is tetrarnerous; the hind tibia has one dorsal tooth and one ventral, and an antiaxial crest of two teeth, the tarsus is tetrarnerous. All tarsi bear some ventral, conical spines. The genitalia are simpie. The length (head and thorax) is ca. 0.9 mmo The colour is light brown.

The host fig is Ficus louisii Bout. & Léon. (Gabon).

Agaon Dalman (fig. 67) Dalman, Svensk. Vetensk. Acad. Handl. (1818): 69 (1818); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 395--407 (1989c).

Fig. 67. Agaon balio/urn Wiebes, female, and A. taïense Wiebes, male, after Wiebes (1989c, fig . 6).

255 The female head is long, usually with the cheeks distinctly longer than the lon­ gitudinal diameter of the compound eye, but not with the species with large, buiging eyes. There are no ocelli (one exception: only the laterals present). The c1ypeus has an oblong patch of many setae, or only a few. The antennae, consist­ ing of e1even free segments, bear rows of long, flexible sensilla chaetica on the funicular segments. The mandibles have strong teeth and ventral ridges, some of which are developed into teeth; the mandibular appendage is long, it bears over thirty ventral rows of small teeth or crenulations. The labium and maxillae each have (sub)apical setae. The thorax has pollen pockets and the fore coxae have a corbicula. The vena­ tion of the fore wing is complete, and the postmarginal vein is distinctly longer than the stigma\. The legs are long and slender, and the apical teeth of the fore tibia and the antiaxial spur of the hind tibia are smal\. The stigmatal peritremata of the eighth urotergite are ovoid, of medium size. The hypopygium has a long, acute spine. The total length (head, thorax and gaster) is ca. 2-3 mm; the valves of the ovipositor are longer than the gaster. The colour is yellowish.

The male is robust, but it may be rather narrow when seen from above. The antennae are, separately, borne in an anterior groove, in between lateral eleva­ tions, and they are composed of the scape, pedicel, three anelli (two in one species), and a club. The mandibles are robust. The thorax has a long pronotum, approximately as long as the other segments combined; the propodeum has circular (fig. a) or oblong spiracles (fig. b) of varying size. The fore tarsus is bimerous (one exception: trirnerous); the mid and hind tarsi of most species have four segments. The first gastral segment is in a number of species heavily sclerotized dorsally, showing subcircular pores (fig. b), and a dorsal slit in some (fig. a). The genitalia are simpie. The total length of the head and thorax is ca. 1-2 mmo The colour is light brown.

There are eleven species, associated with figs of subsection Cyathistipulae. They can be distinguished by the following key. Nos. 64-68 form a complex of species c10sely allied to A. paradoxum.

KEY TO THE SPECIES (females; for the males see table 11) (fig. 68).

I. Compound eyes large and buIging, their longitudinal diameter 11/2- 2 times as long as 2 the cheek ...... - Compound eyes much smaller, even if slightly bulging, their longitudinal diameter 1/2- 1 times as long as the cheek ...... 3

2. Compound eyes 11/2 times as long as the cheek; two lateral ocelli present...... 58. A. megalopon - Compound eyes two times as long as the cheek; no ocelli ...... 59. A. kiellandi

256 Fig. 68 . Details of Agaon. a, A. gabonense Wiebes, male thorax and fust gastral segrnent, after Wiebes (1989c, fig. 17); b, e - f, A. spatulaturn Wiebes: b, male thorax and fust gastral tergite, e, pedice1 and appendage of third segrnent of female antenna, and f, female mandible, after Wiebes (I989c, fig . 12, and 1968b, figs. 21 and 19, respectively); c-d, c1ypeal part of female face, of: c, A. taïense Wiebes, and d, A. c. cicatriferens Wiebes, after Wiebes (1989c, figs. 5 and 15, respectively); g, A. obtusurn Wiebes, female mandible.

3. Clypeus with a short longitudinal row, or only a pair, of setae (fig. c) . 4 - Clypeus with a patch of setae (fig. d) ...... 5

4. Number of ventral rows on mandibular appendage ca. forty, consisting of twelve small teeth. Eye as long as the cheek ...... 60 . A. taïense - Number of ventral rows on mandibular appendage ca. fifty, consisting of ca. fifteen small teeth. Eye shorter, half as long as the cheek ...... 68. A. baliolurn

5. Number of small teeth or crenulations in the ventral rows on the mandibular ap- pendage ca. 15-20. Eye most\y .iS long as the cheek, but it may be shorter ...... 6 - Number of small teeth or crenulations in the ventral rows on the mandibular ap- pendage larger, 25-40. Eye shorter, at most three-quarters of the length of the cheek 7

6. Mandibular appendage with 45-50 rows of twenty small teeth or crenulations ... . . · ...... 64. A. fasciaturn - Number of rows ca. 55, with 15-20 small teeth or crenulations ...... · ...... 61a. A. c. cicatriferens - Number of rows ca. 60, with 15- 20 small teeth or crenulations ...... · ...... 65. A. paradox urn - Number of rows ca. 70, with ca. twenty small teeth or crenulations ...... · ...... 61 b. A. cicatriferens mul/urn

7. Number of small teeth or crenulations in the ventral rows on the mandibular ap­ pendage ca. 25 ...... 62. A. gabonense - Number of small teeth or crenulations in the ventral rows on the mandibular ap- pendage thirty to forty ...... 8

257 8. Process of third antennal segment very long and wide, spatulate, reaching far beyond the apex of the fourth segment (fig. e) ...... 63. A. spatulaturn - Process of third antennal segment much shorter, at most reaching the apical edge of the fourth segment ...... 9

9. Mandibular appendage with ca. 70 rows consisting of thirty small teeth or crenula- tions; the mandible has (four) norrnal teeth (as in fig. f) ...... 66. A. acutatum - Mandibular appendage with ca. 75 rows consisting of forty or more small teeth or crenulations; the teeth of the mandible characteristically blunt (fig. g) ...... 67. A. obtusum

58. Agaon megalopon Wie bes Wiebes, Ent. Ber. Amst. 36: 124-126 (1976).

The female head is longer than wide across the compound eyes (1.25), which are 1 buIging and very large: their longitudinal diameter is 1 / 2 times as long as the cheek. Two lateral ocelIi are present. The clypeus has only a few setae. The mandible has ca. mne ventral ridges, the apical-most of which are produced into sharp teeth; the appendage bears ca. 35 rows of up to ten fine crenulations. The postmarginal vein of the fore wing is two times as long as the stigmal. The total length is ca. 2 mm; the valves of the ovipositor are 1.1 times as long as the gaster.

The male head is rather wide, but still distinctly longer than wide (1.25); the eyes are very large. The antennal anelli number three; the club is divided at two-thirds of its length. The thorax is rather wide, compared with the other species; the propodeal spiracles are large, circular. The fore tarsus has three distinct seg­ ments. The total length is ca. 1 mmo

The host fig is Ficus ardisoides sub sp. camptoneura (Mildbr.) Berg (Nigeria).

Table 11. Some characters of the males of Agaon (that of no. 68, A. baliolum, is not known). no. Ant. Metan. First Propod.spiracles Fore anelli prop.: gastr. tarsi pores terg. small large with: circ. circ. obI.

58. A. megalopon . .3 + .3 59. A. kiellandi · 3 + .2 60. A. taïense · 3 . + . .2 61. A. cicatriferens .2 · scar .+ .2 62. A. gabonense · 3 · scar + .2 63. A. spatulaturn . · 3 .+ · pores . .+ .2 64. A. fasciaturn . · 3. + · pores . + .2 65. A. paradoxum . · 3 · pores . + .2 66. A. acutatum . · 3 .+ · pores . + .2 67. A. obtusum .3 . + · pores . + .2

258 59. Agaon kieUandi Wiebes Wiebes, Zool. Meded. Leiden 48: 124-127 (1974b).

The fe male head is longer than wide across the compound eyes (1.25), which are buiging and very large: their longitudinal diameter is two times as long as the cheek. The clypeus has oniy a few setae. The mandible has seven ventral ridges, only one of which is produced into a recognizable tooth; the appendage is long and narrow, and it bears ca. thirty rows of seven small crenulations. The post­ marginal vein of the fore wing is four times as long as the stigma!. The .total length is ca. 2.5 mm; the valves of the ovipositor as long as the gaster.

The male head is almost twice as long as wide, when seen from above (1.7). There are th ree antennal anelli; the club is divided at two-thirds of its length. The thorax has circular propodeal stigmata, which are small (one-quarter in length of the propodeurn). The fore tarsus has two segments. The totallength is ca. 1.5 mmo

The species was described from an unknown species of fig (Tanganyika) and later recorded from Ficus conraui Warb. (Carneroons, Gabon), Ficus cyathi­ stipula subsp. pringsheimiana (Braun & Schum.) Berg (Gabon) and Ficus densi­ stipulata De Wild. (Gabon).

60. Agaon taïense Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 399-401 (I 989c).

1 The female head is almast 1 / 2 times as long as wide across the compound eyes (1.35); the longitudinal diameter of the eye is subequal to the length of the cheek. The clypeus has only a short longitudinal row of setae. The mandible has one apical toa th and fifteen ventral ridges, five of which are produced into teeth; the appendage has forty rows of up to ten to twelve small teeth. The postmarginal vein of the fore wing is three times as long as the stigma!. The total length is ca. 2.3 mm; the valves of the ovipositor are 1.4 times as long as the gaster.

The male head is very narrow when seen from above, not quite three times as long as wide (2.8). There are three antennal anelli. The propodeal spiracles are large, oblong (0.3),covering 0.85 of the length of the propodeum. The fore tarsus has two segments. The total length is ca. 1.5 mmo

The host fig is Ficus tesselata Warb. (Liberia).

61a. Agaon c. cicakÜerens Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 401 (I 989c).

259 1 The female head is not quite 1 / 2 times as long as wide across the compound eyes (l.4); the longitudinal diameter of the eye is somewhat shorter than the cheek (0.8). The clypeus has an oblong patch of setae. The mandible is much as in A. taïense, but the appendage has 55 rows of ca. twenty small teeth. The postmar­ 1 ginal vein of the fore wing is almost 3 / 2 times as long as the stigmal (3.4). The total length is ca. 2.8 mm; the valves of the ovipositor are 1.1 times as long as the gaster.

1 The male head is 1 /2 times as long as wide. There are two short anelli in the antenna. The fore tarsus has two segments. The propodeal spiracles are oblong, and they occupy the whole length of the propodeurn. The fust gastral tergite has an oblong scar of three-quarters of its length. The totallength is ca. 1.7 mm.

The host tig is Ficus sagittifolia Mildbr. & Burret (Ivory Coast).

61b. Agaon cicatriferens multum Wiebes Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 401 (1989c).

The only difTerence with the nominate subspecies is found in the number of rows in the female mandibular appendage, which is seventy (vs. 55).

The host tig is Ficus subsagittifolia Berg (Gabon).

62. Agaon gabonense Wie bes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 402-403 (1989c).

1 The female head is approximately 1 / 2 times as long as wide across the compound eyes (1.55); the longitudinal diameter of the eye is half as long as the cheek. The clypeus has an oblong patch of setae. The mandible has six axial teeth, all exten­ sions of ventral ridges, and one at the antiaxial si de (level with the second axial); the appendage has about eighty rows of 25 small teeth or crenulations. The postmarginal vein of the fore wing is 2.6 times as long as the stigma!. The total length is ca. 2.5 mm (of which the head takes 0.8 mm); the valves of the ovipo­ sitor are 1.3 times as long as the gaster.

1 The male head is almost 1 / 2 times as long as wide (l.4). There are three antennal anelli. The fore tarsus has two segments. The propodeal spiracles are sm all and circular. The fust gastral tergite has an oblong scar of almost all of its length. The total length is ca. 1.4 mmo

The host tig is Ficus wildemaniana Warb. (Gabon).

260 63. Agaon spatuIatum Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 71 : 352- 353 (1968b).

The female head is 1.7 times as long as wide across the compound eyes; the longitudinal diameter of the eye is less than half as long as the cheek (0.45). The c1ypeus has an oblong patch of setae. The antenna has a very wide and long appendage of the third segment, it is spatulate and reaches far beyond the apex of the fourth segment. The mandible is much as in A. gabonense; the appendage has ca. 65 rows of 35 small teeth and crenulations. The postmarginal vein of the fore wing is 2.7 times as long as the stigma\. The totallength is ca. 2.5 mm; the valves of the ovipositor are 1.3 times as long as the gaster.

The male head is 1.3 times as long as wide. There are three anelli in the antenna. There are two segments in the fore tarsus. The propodeal spiracles are oblong (2.4), almost half as long as the propodeum. The thorax (meso- and metanotum and the propodeum) bear many subcircular pores; the first gastral tergite also has dorsal pores. The total length is ca. 1.6 mmo

The host fig is Ficus lyrata Warb. (Ivory Coast); the species was caught at light in Nigeria and Zaïre.

64. Agaon fasciatum Waters ton

Waterston, Bull. ent. Res. 5: 249-253 (1914); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 71 : 350-351 (l968b, Agaon paradoxum, male only). Synonym: Agaon tridentatum Joseph, Proc. ent. Soc. London (B) 28: 29-30 (1959).

The female head is 1.5-1.74 times as long as wide across the compound eyes; the longitudinal diameter of the eye almost as long as the cheek (0.9). The c1ypeus has an oblong patch of setae. The man di bIe bears three large axial teeth and one smaller (and one smal\, antiaxial tooth, level with the second axial) and ca. twenty ventral ridges; the appendage has 45-50 rows of 15-20 small teeth. The 1 postmarginal vein of the fore wing is 2 /2 times as long as the stigma\. The total length is ca. 3 mm; the valves of the ovipositor are almost as long as the gaster (0.75-1).

The male head is 1.2 times as long as wide. The antenna has three anelli. The fore tarsus has two segments. The propodeal spiracles are smalI, circular. The meso-, metanotum, propodeum and the first gastral tergite bear many dorsal pores. The total length is ca. 1.5 mmo

The host fig is Ficus c. cyathistipula Warb. (Cameroons, Gabon, Uganda, Zam­ bia); the species was caught in a Berlese apparatus in Guinea. Wiebes (1968b and 1974b) synonymized A. fasciaturn with A. paradoxurn Oalman, but revived it in 1989(c: 405).

261 65. Agaon paradoxum Dalman Dalman, Svensk. Vetensk. Acad. Handl. (1818): 69- 72 (1818); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 71: 346-350 (I 968b, femaJe only). Synonym: BlaslOphaga villiersi Risbec, Bull. Inst. franç. Mr. noire (A) 16: 538-540 (1954a).

The female head is almost twice as long as wide across the compound eyes (1.85); the longitudinal diameter of the eye is more than half as long as the cheek (0.6; to 0.8?). The clypeus has an oblong patch of setae. The mandible is much as described for A. fasciaturn; the appendage has ca. 60 rows of 15- 20 small teeth. The postmarginal vein of the fore wing is four times as long as the stigma!. The totallength is ca. 3 mm; the valves of the ovipositor are slightly longer than the gaster (l.l).

The male is much like th at of A. fascia turn , but the dorsal pores are virtually lacking.

The host fig is Ficus barteri Sprague (Gabon); the species was described from Sierra Leone, and it was caught at light in Sénégal and Ivory Coast.

66. Agaon acutatum Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 406 (l989c).

The species is much like A. fasciaturn, but the female compound eye is three­ quarters of the length of the cheek, and the mandibular appendage bears seventy rows consisting of thirty small teeth or crenulations. The male has less dorsal pores.

The host fig is Ficus subcostata De Wild. (Gabon).

67. Agaon obtusum Wiebes

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 92: 406 (1989c).

The species is much like A . fasciaturn, but the female compound eye is half as long as the cheek, the mandibular teeth are characteristically obtuse, and the appendage bears up to 75 rows of forty or more small teeth or crenulations. The male has Ie ss dorsal pores.

The host fig is Ficus cyathistipuloides De Wild. (Ivory Co ast, Gabon).

68. Agaon baliolum Wie bes Wiebes, Zool. Meded. Leiden 48: 128 (1 974b).

1 The female head is more thán 1 / 2 times as long as wide across the compound

262 eyes (1.65); the longitudinal diameter of the eye is about half as long as the cheek (0.55). The clypeus has a short transverse row of setae. The mandible has six prominent teeth, extensions of ventral ridges, and there are as many smaller ridges in between; the appendage bears ca. fifty rows of about fifteen small teeth. The postmarginal vein of the fore wing is two times as long as the stigma\. The total length is ca. 2 mm; the valves of the ovipositor are 1.25 times as long as the gaster.

The male is unknown.

The host fig is not exactly known. The species was recorded from Nigeria, ex Ficus winkleri Mildbr. & Burret (= F. tesselata Warb.), from which Agaon taïense (no. 60) is known.

CourteUa Kleffer (fig. 69) KiefTer, Ann. Soc. ent. France 80: 464 (1911); Wiebes, Zool. Meded. Leiden 48: 128- \37 (1974b, group of Agaon hamiferum); Michaloud, Michaloud-PeUetier, Wiebes & Berg, Proc. Kon. Ned. Akad. Wet. (C) 88: \05-112 (\985); Wiebes, Proc., Kon. Ned. Akad. Wet. (C) 89: 247 table 3 (\986b).

The female head is distinctly longer than wide across the compound eyes (1.2-2); the longitudinal diameter of the eye is approximately half as long as the cheek (0.3-0.55). There are three ocelli. The anten na has eleven free segments; the third segments has an appendage that usually is short; the funicular segments bear sensilla chaetica, mostly long and ftexible. The mandible either has one apical tooth and a number of ventral ridges, which in some species give the impression of a washing-board, or the ventral ridges are extended into teeth along the stoma1 margin, and th en may number up to eight. The appendage is two to four times as long as wide, and it bears twenty to fifty (in one species even sixty to seventy) rows of twenty to more than eighty small teeth or fine crenulations. The labium and maxillae each have a number of (sub)apical setae. The thorax has pollen pockets and the fore coxae have a corbicula. The vena­ ti on of the fore wing is complete, and the postmarginal vein in a few species is as long as the stigmal, but mostly it is distinctly longer: (almost) two to almost four times. The stigmatal peritremata of the eighth urotergite are of medium size, oblong. The totallength (head, thorax and gaster) is ca. 2- 3 mm; the ovipositor valves are subequal in length to the gaster, but in two species they are distinctly longer (1.5-1.75). The colour is brown or dark brown.

The male is blind in two species, but in most there are eyes, which may be large (ca. one-fifth of the length of the head; fig . b), of medium size (ca. one-sixth or one-seventh; fig. a), small (one-eighth to one-tenth), or very small (the head more than twenty-five times as long as the eye). The antennae are borne in a narrow

263 Fig. 69. Cour/elia michaloudi (Wiebes), female, and C. sylviae Wiebes, male.

groove, but in most species they are situated more separateJy, in toruli at the front of the head (in one species at the end of outgrowths), and their position may he transverse (fig. a) or porrect (fig. b). There are two (fig. a) or three anelli (fig. b), and the club may he divided. The thorax has an oblong pronotum, and the meso- and metanotum and the propodeum are fused dorsally, but for a short lateral separation of the pro­ podeum; the spiracular peritremata usually occupy one-third of the length of the propodeum, but they may be shorter (0.15) or longer (0.5-0.85) in some species.

264 Table 12. Some characters of the males of Courtefla (that of no. 75, C. scobinifera, is not known). no. Eye, Ant.c1ub Ant. Ant. Spir.! size: divided anelli scape: prop.

69. C. harnifera . · medium · + . . 3 · porrect . · 0.33 70. C. bekiliensis · small . · + . .3 · porrect . .0.5 71. C. malawi · small . · + . .3 · porrect . .0.33 n.c. gabonensis . · large · + . · 3 · porrect . · 0.33 73. C. arrnata · small . · + . .3 · porrect . · 0.375 74. C. carnerunensis . · large · + . . 3 · porrect . .0.33 76. C. sylviae · very small . · 3 · transv . 0.85 77. C. penicula · medium .2 · transv .0.75 78. C. wardi. · large · + . . 2 · transv . 0.33 79. C. hladikae · medium .2 · transv .0.33 80. C. michaloudi . · very small . .3 · *) .0.5 81. C. medleri . · absent · 3 · **) · 0.15

*) from medial sockets **) from lateral outgrowths

The fore tarsus is bimerous, the other tarsi are pentamerous. The genitalia are simpie. The totallength (head and thorax) is ca. 1.2-3 mm. The colour is light brown.

There are thirteen species and two subspecies, associated with figs of subsection Caulocarpae. They can be distinguished by the following key.

KEY TO THE SPECIES (females; for the males see table 12) (fig. 70).

I. Mandible with two or more apical teeth andlor up to seven prominent teeth along the stomal edge (e.g., fig. c); there are less than ten ventral ridges ...... 2 - Mandible with one apical tooth and none along the stomal edge (e.g., fig. d); there are ten or more ventral ridges ...... 8

2. Head relatively short, less than 1.3 times as long as wide across the compound eyes 3 - Head longer, 1'/2 or more times as long as wide across the compound eyes ...... 4 3. Mandibular appendage 3'/4 times as long as wide, with twenty rows of ca. twenty teeth. Postmarginal vein twice as long as stigmal ...... 78. C. wardi - Mandibular appendage almost three times as long as wide, with 35 rows of ca. forty fine crenulations. Postmarginal vein more than 3'j, times as long as stigmal ...... " ...... 76. C. sylviae

4. Head twice as long as wide across the compound eyes, which are one-third of the length of the cheek. Mandible with ca. seven strong teeth along the stomal edge...... 73. C. armata - Head 1.5 to 1.65 times as long as wide across the compound eyes, which are two-fifths of the length of the cheek. Mandible with one to three teeth next to the apical one . . 5

5. Fore tarsus with axial, conical spines (fig. e) ...... 69. C. hamifera - Fore tarsus without axiaI, conical spines ...... 6

265 c

/ / I I \ \

Fig. 70. Details of Cour/ella. a- b, male head of: a, C. penicu/a (Wiebes), and b, C. camerunensis (Wiebes), after Michaloud et al. (1985, figs. 23 and 24, respectively); c--

6. Antennal scape flat at the antiaxial surface, with a simple hyaline margin; the third segment, including the appendage, equal in length to the pedicel. Mandible with two apical teeth. Postmarginal vein twice as long as stigmal ...... 72. C. gabonensis

266 - Dorsal margin of antennal scape tumed outward, like the culT of a wide sleeve; the third segment, including the appendage, twice as long as the pedicel. Mandible with three apical teeth. Postmarginal vein as long as stigmal ...... 7

7. Mandibular appendage with ca. forty rows of 25-30 small teeth . . . 70. C. bekiliensis - Mandibular appendage with ca. thirty rows of 25-30 small teeth ..... 71. C. malawi 8. Mandible with 10 - IS ventral ridges; the appendage with ventral rows consisting of less than fifty small teeth or crenulations. Antennal scape not expanded (e.g., fig. h) 9 - Mandible with more than 25 ventral ridges; the appendage with ventral rows consisting of eighty or more fine teeth or crenulations. Antennal scape expanded (e.g. , fig. i). .. 12 9. Antennal sensilla short and rather stifT, subequal in length to the segment or shorter (fig. h) ...... 77. C. penicu/a - Antennal sensilla long and flexible, much longer than the segment...... 10

10. Mandibular appendage four times as long as wide, with ca. fifty rows of ca. fifty fine crenulations ...... 80. C. micha/oudi 1 - Mandibular appendage 2-2 /2 times as long as wide, with less than thirty rows of not more than forty fine crenulations ...... 11 11. Compound eye rather short, two-fifths of the length of the cheek. Mandibular ap- pendage with 25-30 rows of ca. forty fine crenulations ...... 74. C. camerunensis - Compound eye longer, more than half as long as the cheek (0.55). Mandibular ap­ pendage with ca. twenty rows of 20 - 30 fine teeth or crenulations ...... 75. C. scobinifera 12. Antennal pedicel enlarged in a peg-like extension (fig. i). Mandibular appendage with 75-80 ventral ridges. FOUT segments of the fore tarsus bear conical spines ...... 81. C. med/eri Antennal pedicel not extended. Mandibular appendage with 25-30 ventral ridges. Fore tarsus without conical spines ...... 79. C. hladikae

69a. CourtelIa h. hamifera Kleffer KielTer, Ann. Soc. ent. France 80: 466 (1911); Grandi, Boll. Lab. Zool. Portici 10: 207-221 (1916a, Agaon ?paradoxum).

1 The female head is more than 1/ 2 times as long as wide across the compound eyes (1.67); the longitudinal diameter of the eye is Ie ss than half the length of the cheek (0.4). The mandible has four teeth; the appendage is four times as long as wide, and it bears ca. forty rows of ca. thirty fine crenulations. The postmar­ ginal vein of the fore wing is approximately 1.7 times as long as the stigma!. The fore tarsus bears axial, conical spines, which on the first segment number ca. thirty. The hypopygium has a sharp, but short spine. The totallength is ca. 2.5 mm; the valves of the ovipositor are as long asthe gaster, or slightly longer.

The male eye is of medium size. The antennal scapes are porrect; there are three anelli; the club is divided. The stigmatal peritremata occupy one-third of the length of the propodeum. The total length is ca. 1.5-2 mmo

267 The species was described from Togo. The host tig is Ficus ovala Vahl (Sénégal, Ivory Coast).

69b. CourteUa hamifera modesta (Wiebes)

Wiebes, Zool. Meded. Leiden 37: 231 - 233 (1961 , Agaon paradoxum modes/um); Copland, King & Hili, J. Ent. (Lond.) (A) 48: 25- 35 (1973, Agaon paradoxum).

This subspecies difTers from the nominate form in the armature of the female fore tarsus, which has only ten axial, conical spines on the fust segment. The male is more slender than in subsp. hamifera.

The host tig is Ficus ovala Vahl (Uganda). Copland et a!. (1973) described the female reproductive system.

70a. CourteUa b. bekiliensis (Risbec) Risbec, Ann. Soc. ent. France 124: 191- 194 (1956, Pleistodontes); Wiebes, Zool. Meded. Leiden 45: 2- 3 (1970, Agaon).

1 The female head is not quite 1 / 2 times as long as wide across the compound eyes; the longitudinal diameter of the eye is half as long as the cheek. The mandible has three or four teeth and approximately eight ventral ridges (and some smaller ridges in between the larger); the appendage is three times as long as wide, and it bears thirty rows of 30-35 small teeth. The postmarginal vein of the fore wing is as long as the stigma!. There is a dark patch on the base of the stigma!. The hypopygium has a short, blunt spine. The totallength is ca. 3 mm. ; the ovipo­ sitor valves are as long as the gaster.

The male eye is smalI. The antennal scapes are porrect; there are three anelli; the club is divided. The spiracular peritremata occupy half of the length of the propodeum. The total length is ca. 1.7 mmo

The host tig is Ficus p. polila Vahl (Madagascar).

70b. CourteUa bekiliensis bispinosa (Wiebes)

Wiebes, Ann. Mus. Roy. Afr. centr., in 8°, Zool. 175: 453-454 (l969c, Agaon bispinosum).

This subspecies difTers from the nominate form in the length of the female com­ pound eye relative to that of the cheek (0.4 vs. 0.5) and in the length of the antennal pedicel (lengthlwidth, ca. 3, vs. 5); the mandibular appendage bears ca. forty (vs. thirty) rows of 25 - 30 sm all teeth.

The host tig is Ficus p. polila Vahl (Sénégal, Ivory Coast, Nigeria).

268 71. CourteUa malawi Wiebes Wiebes in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 217-218 (1990). The female is similar to that of C. b. bispinosa (no. 70b), but the mandibular appendage bears ca. fifty rows (vs. forty) of 15-30 small teeth. The totallength is ca. 2.4 mm; the ovipositor valves are slightly shorter than the gaster (0.9). The male is more distinct, in that the thoracical dorsum has longitudinal striae, ending in a series of pits on the propodeurn. The spiracular peritremata occupy little more than one-third of the length of the propodeum. The total length is ca. 1.2 mmo

The host tig is Ficus chirindensis Berg (Malawi).

72. CourteUa gabonensis Wie bes

Wiebes in Micbaloud et al., Proc. Kon. Ned. Akad. Wet. (C) 88: 107-109 (1985).

1 The female head is 1 / 2 times as long as wide across the compound eyes; the longitudinal diameter of the eye is less than half as long as the cheek (0.4). The mandible has two apical teeth, and ventral ridges as in C. bekiliensis; the ap­ 1 pendage is 2 / 2 times as long as wide, and it bears 25-30 rows of ca. twenty small teeth. The postmarginal vein of the fore wing is two times as long as the stigma!. The hypopygium has a short, but sharp spine. The total length is ca. 2.1 mm; the ovipositor valves are slightly shorter than the gaster (0.9).

The male eye is large. The antennal scapes are porrect; there are three anelli; the club is divided. The spiracular peritremata occupy one-third of the length of the propodeum. The total length is ca. 1.4 mmo

The host tig is Ficus o. ottoniifolia (Miq.) Miq. (Ivory Coast, Gabon). In Gabon, the species may share the tig syconium with C. camerunensis (no. 74), see Mi­ chaloud (1982).

73. CourteUa armata (Wiebes)

Wiebes, Zool. Meded. Leiden 48: 134-136 (I 974b, Agaon armatum); Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 89: 343-344 (l986b).

The female head is two times as long as wide across the compound eyes; the longitudinal diameter of the eye is one-third of the length of the cheek. The antennal sensilla are rather stiff, subequal in length to their segment. The man­ dible has eight strong teeth along the stomal edge; the appendage is three times as long as wide, and it bears ca. 40-45 rows of thirty fine crenulations. The postmarginal vein of the fore wing is subequal to the stigma!. The spine of the hypopygium is rather acute. The total length is ca. 3.2 mm; the valves of the ovipositor are as long as the gaster.

269 The male eye is smal\. The antennal scapes are porrect; there are three anelli; the club is divided. The spiracular peritremata occupy distinctly less than half of the length of the propodeum (0.375). The totallength is ca. 1.6 mmo

The host fig is Ficus sansibarica Warb. (both subspecies) (Ivory Coast, Nigeria, Zambia).

74. CourteUa camerunensis (Wie bes)

Wiebes, Zool. Meded. Leiden 48: 130-132 (l974b, Agaon).

1 The female head is 1 / 2 times as long as wide across the compound eyes; the longitudinal diameter of the eye is less than half as long as the cheek (0.4). The mandible has one apical tooth and ca. fifteen ventral ridges; the appendage is 1 2 / 2 times as long as wide, and it bears 25- 30 rows of forty fine crenulations. The postmarginal vein of the fore wing is two times as long as the stigma\. The spi ne of the hypopygium is short and blunt. The total length is ca. 2.1 mm; the valves of the ovipositor are as long as the gaster.

The male eye is large. The antennal scapes are porrect; there are three anelli; the club is divided. The spiracular peritremata occupy one-third of the length of the propodeum .. The total length is ca. 1.4 mmo

The host fig is Ficus ottoniifolia: subspecies otttoniifolia (Miq.) Miq. (Cameroon, Gabon), macrosyce Berg (Zambia) and ulugurensis (Mildbr. & Burret) Berg (Kenya). See also under C. gabonensis (no. 72).

75. CourteUa scobinifera (Waters ton)

Waterston, Trans. ent. Soc. London (1920): 132 (l920a, Agaon scobiniferum).

The female head is 11/4 times as long as wide across the compound eyes; the longitudinal diameter of the eye is somewhat more than half as long as the cheek (0.55). The mandible has one apical tooth and ten to twelve ventral ridges; the appendage is two times as long as wide, and it bears twenty rows of 20-30 small teeth. The wings and the hypopygium were not described. The totallength is ca. 2.25 mm; the valves of the ovipositor are a little shorter than the gaster.

The male is not known.

The host fig is Ficus ottoniifolia subsp. lucanda (Ficalho) Berg (Uganda).

76. CourteUa sylviae Wiebes Wiebes, Proc. Kon., Ned. Akad. Wet. (C) 89: 344-346 (1986b).

270 The female head is 1.35 times as long as wide across the compound eyes; the longitudinal diameter of the eye is Ie ss than half as long as the cheek (0.4). The mandible has one apical tooth and a row of prominent tooth-like lobes along the stoma 1 edge, and there are ca. ten ventral ridges; the appendage is not quite three times as long as wide, and it bears ca. 35 rows of forty fine crenulations. The postmarginal vein of the fore wing is 3.65 times as long as the stigmal. The spine of the hypopygium is rather long and sharp. The total length is ca. 3 mm; the valves of the ovipositor are much longer than the gaster (1.75).

The male eye is very small. The antennal scapes are transverse; there are three anelJj; the club is not divided. The spiracular peritremata are very large and occupy the greater part of the propodeum (0.85). The total length is ca. 3 mmo

The host fig is Ficus dryepondtiana De Wild. (Gabon).

77. CourteUa penicula (Wiebes)

Wiebes, Zool. Meded. Leiden 48: 133- 134 (I974b, Agaon peniculum).

1 The female head 1/ 2 times as long as wide across the compound eyes; the longi­ tudinal diameter of the eye is one-third of the length of the cheek. The antennal sensilla are rather stiff, subequal to the length of their segment or shorter. The mandible has one apical tooth and ca. ten ventral ridges; the appendage is three times as long as wide, and it bears ca. 35 rows of 40 - 45 fine crenulations. The 1 postmarginal vein of the fore wing is 3 / 2 times as long as the stigmal. The hypo­ pygium has a short and blunt spine. The totallength is ca. 3 mm; the valves of the ovipositor are a little shorter than the gaster (0.9).

The male eye is of medium size. The antennal scapes are transverse; there are two anelli; the club is not divided. The spiracular peritremata occupy the greater part of the length of the propodeum (0.75), and they are ventro-Iateral in posi­ tion. The totallength is ca. 1.8 mmo

The species was described from Ghana and Uganda; the host fig is Ficus arto­ carpoides Warb. (lvory Coast, Gabon). From this species of fig also C. hladikae (no. 79) was recorded (Michaloud et al. , 1985: 112).

78. CourteUa wardi Compton Compton in Wiebes & Compton, Proc. Kon. Ned. Akad. Wet. 93: 215-217 (1990).

The female head is longer than wide across the compound eyes, but less than in most other species (1.2); the longitudinal diameter of the eye is half as long as the cheek. The man di bie has eight teeth, most of which are extensions of ventral ridges; the appendage is somewhat more than three times as long as wide

271 (3.25), and it bears twenty rows of ca. twenty small teeth. The postmarginal vein of the fore wing is two times as long as the stigmal. The spine of the hypopygium is short and blunt. The total length is ca. 2 mm; the valves of the ovipositor are slightly shorter than the gaster (0.9).

The male eye is large. The antennal scapes are transverse; there are two anelli; the club is divided. The spiracular peritremata occupy one-third of the length of the propodeum. The totallength is ca. 1.4 mmo

The host fig is Ficus t. tremuia Warb. (South Africa).

79. CourteUa hladikae (Wiebes)

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 82: 394--397 (1979b, Agaon).

The female head is 1.35 times as long as wide across the compound eyes; the longitudinal diameter of the eye is less tban half as long as the cheek (0.4). The antennal scape is extended at the apex, but the pedicel is simpie. Tbe mandible has the shape of a was hing board, it ends in one apical tooth and bears 25-30 ventral ridges; tbe appendage is almost two times as long as wide, and it bears ca. 45 rows of eighty fine crenulations. Tbe postmarginal vein of tbe fore wing is two times as long as the stigmal. The spine of the hypopygium is short, but sharp. The total length is ca. 3 mm; the valves of the ovipositor are as long as the gaster.

The male eye is of medium size. The antennal scapes are transverse; there are two ane11i; the club is not divided. The spiracular peritremata occupy one-tbird of the length of the propodeum. The totallength is ca. 1.7 mmo

The host fig is Ficus artocarpoides Warb. (Ivory Co ast, Cameroon, Nigeria, Gabon). From th is species of fig, also C. penicula (no. 77) was recorded.

80. CourteUa michaloudi (Wiebes)

Wiebes, Proc. Kon. Ned. Akad. Wet. (C) 82: 392- 394 (1979b, Agaon).

The female had is 1.35 times as long as wide across the compound eyes; the longitudinal diameter of the eye is 0.4 of the length of the cheek (0.4). The apex of the antennal scape is not extended and the pedicel is simpie. The mandible has one apical tooth and ca. fifteen ventral ridges; the appendage is four times as long as wide, and it bears ca. fifty rows of ca. fifty fine cenulations. The postmarginal vein of the fore wing is two times as long as the stigmal. The spine of the hypopygium is short and blunt. The total length is ca. 3 mm; the valves of the ovipositor are shorter than the gaster (0.85).

272 The male eye is very smalI. The antennae are borne in sockets, close together in the front of the head; there are three anelli; the club is not divided. The falcate mandibles are situated on outgrowths of the head. The spiracular peritremata occupy half of the length of the propodeum. The total length is 2.2 mmo

The host fig is Ficus bubu Warb. (Gabon).

81. CourteUa medleri (Wiebes)

Wiebes, Ent. Ber. Amst. 32: 122- 124 (1972a, Agaon).

The female head is 1.65 times as long as wide across the compound eyes; the longitudinal diameter of the eye is one-third of the length of the cheek. The antennal scape is extended apicad and also the pedicel has a dorsal, spatulate outgrowth almost as long as the third segment. The mandible has the shape of a washing board, it ends in one apical tooth and bears ca. 75-80 ventral ridges; 1 the appendage is 2 /2 times as long as wide, and it bears ca. sixty to seventy rows of ca. eighty fine crenulations. The postmarginal vein of the fore wing is two times as long as the stigma!. The hypopygium has a rather acute spine. The total 1 length is ca. 3 mm.; the ovipositor-valves are 1/ 2 times as long as the gaster.

The male is blind. The antennal sockets are situated on outgrowths of the head, which also bear the falcate mandibles. The antennal scape is spatulate; there are three anelli; the club is not divided. The spiracular peritremata occupy one-sixth to one-seventh of the length of the propodeum. The totallength is ca. 2.5 mmo

The host fig is Ficus umbelIata Vahl (Sierra Leone, Togo, Nigeria).

REFERENCES (only those not mentioned in the bibliography)

Balakrishnan Nair, P. & U.e. Abdurahiman - Population dynarnics of the tig wasp Kradibia ges/roi (Grand i) (Hyrnenoptera Chalcidoidea, Agaonidae) from Ficus exasperata Vahl. Proc. Kon. Ned. Akad. Wet. (C) 87: 365-375 (1984). China, W.E. - Cera/osolen Mayr, 1885 (Insecta, Hymenoptera): validated under the plenary powers. Bull. zool. Nomencl. 20: 414-415 (1963). China, W.E. - Cynips caricae Linnaeus, 1762 (Insecta, Hyrnenoptera): validated under the plenary powers. Bull. zool. Nomencl. 21: 31-31 (1964). Corner, EJ.H. - Ficus dammaropsis and the multibracteate species of Ficus sect. Sycocarpus. Phil. Trans. R. Soc. London (B) 281 (982): 372-406 (1978). GaliI, J. & D. Eisikowitch - On the pollination ecology of Ficus re/igiosa in lsrael. Phytomorphol­ ogy 18: 356-363 (1968 c). Galil, J. & G. Neeman - Pollen transfer and pollination in the common tig (Ficus carica L.). New Phytol. 79: 163- 171 (1977). Galil, 1. & Y. Snitzer-Pasternak - Pollination in Ficus religiosa L. as connected with the structure and mode of action of the pollen pockets of Blastophaga quadra/iceps Mayr. New Phytol. 69: 775-784 (1970). Grandi, G. - Contributo alla conoscenza degli Agaonini (Hymenoptera, Chalcididae) di Ceylon e dell'lndia. Boli. Lab. Zool. Portici 11: 183-234 (1916b).

273 Grandi, G. - Identification of some fig-insects (Hymenoptera) in the British Museum (Natural History). Bull. ent. Res. 13: 295-299 (1923). Grandi, G. - Imenotteri sicofili raccolti dal Prof. F. Silvestri nell'lndocina, nella Cina, nella pe­ nisola Malacca ed a Formosa. Boll. Lab. Zool. Portici 20: 169- 188 (1927). Grandi, G. - Studio morfologico e biologico della Blastophaga psenes (L.). 2a edizione riveduta. Boll. Lab. Ent. Bologna 2: 1- 147 (1929). Gravenhorst, J.L.c. - AlIgemeiner Bericht der Entomologischen Section. Uebers. Arb. Veränd. Schles. Ges. vaterl. Cult. i. J. (1826): 23- 24 (1827). Gravenhorst, J.L.c. - Diquisitio de Cynipe psene auctorum, et decsriptio Blastophagae, novi Hy­ menopterorum generis. Beitr. Ent., Breslau I: 27- 33 (1829). Hili, D.S. - Fig-wasps (Chalcidoidea) of Hong Kong. i. Agaonidae. Zool. Verh. Leiden 89: I-SS (\967a). lshii, T. - Fig Chalcidoids of Japan. Kontyu 8: 84-100 (1934). Joseph, KJ. - Contributions to our knowledge of fig insects (Chalcidoidea: parasitic Hymenoptera (from India. ivo Descriptions of three new and records of four known species of Agaonini. Agra Univ. J. Res. (Sci.) 2: 267-384 (1953). Joseph, K .l. - Contributions to our knowledge of fig insects (Chalcidoidea: parasitic Hymenop­ tera) from India. vi. On six species of Agaoninae. Agra Univ. J. Res. (Sci .) 3: 401-416 (1954). Linnaeus, C. - V. Hymenoptera. Syst. Natur., ed. 10a, I: 553-583 (1758). Mani, M.S. - The fauna of India and the adjacent countries. Chalcidoidea (Hymenoptera). Part i. Madras, Zool. Survey India, pp. 1-1067 (1989). McKey, D. - Population biology of figs: applications for conservation. Experientia 45: 661-{i73 (1989). Motschoulsky, V. De - Essai d'un catalogue des insectes de I'ile Ceylan. Bull. Soc. Natur. Moscou 36 (2): 1- 153 (1864). RarnÎrez B. , W. & J. Montero S. - Ficus microcarpa L. , F. benjamina L. and other species introduced in the New World, their pollinators (Agaonidae) and other lig wasps. Rev. Biol. Trop. 36: 441-446 (1988). Waterston, 1. - On some Bornean fig insects (Agaonidae - Hymenoptera Chalcidoidea). Bull. ent. Res. 12: 35-40 (192la). Wiebes, J.T. - Phylogenetic specificity of figs and wasps. In N.RM. Brantjes (Ed.), Pollination and dispersal, Nijmegen, pp. 21-25 (1973). Wiebes, J.T. - A short history offig wasp research. Gdns' Bull. Singapore 29: 207-232 (1977a).

274 VI. Bibliography, figs and tig insects of Africa

Andrews, F.W. - The flowering plants of tbe Anglo-Egyptian Sudan 2. Arnroatb, viii + 485 pp. (Ficus: 261 - 272) (1952). Arozarena Doblado, R. - Aportacion de la superfamilia Chalcidoidea a la lucba biologica en la isla de Tenerifa (Hym. Chalcidoidea). Vieraea 7: 191-200 (1978). Aubréville, A. - La flore forestière de la Cote d'Ivoire (ed.2) 1. Rogent-sur-Marne, 369 pp. (Ficus: 68- 98) (1959). Audru, J. - Quelques figuiers d'Afrique de I'Ouest. Genre Ficus, Moracées. Inst. d'Elev. Méd. Vét. Pays Trop. 10. Maisons-Alfort, 149 pp. (1982). Aweke, G. - Revision of the genus Ficus L. (Moraceae) in Etbiopia. Meded. Landbouwhogeschool Wageningen 79-3, 1- 115 (1979). Baer, J.G. - Mésaventure d'un parasitologiste. Rev. Suisse Zoo1.61 , 484 (1954). Baez, M. & G . Ortega - Lista preliminar de los Himenópteros de las Islas Canarias. Bol. Asoc. esp. Entom. 2, 185-189 (1978). Baijnath, H., S. Naicker & S. Ramchuran - The interaction of figs and wasps. In: B.E. Juniper & T.R.E. Southwood, lnsects and the plant surface, 348- 349 (1986). Baijnath, H. & S. Ramcburan - Aspects of pollination and floral development in Ficus capensis Tbunb. Bothalia 14, 883-888 (1983). Baijnath, H. & S. Ramchuran - Reproductive biology and Chalcid symbiosis in Ficus burtt-davyi (Moraceae). In: P. Goldblatt & P.P. Lowry 11, Modern systematic studies in African botany. Monogr. Syst. Bot. Missouri Bot. Gard. 25, 227- 235 (1988). Baillon, H. in G. Grandidier - Histoire pbysique, naturelle et politique de Madagascar. Histoire naturelle des plantes. Atlas 3. Paris, pI. 261-411 (Ficus, pI. 296--304) (1895). Baker, J.G. - Flora of Mauritius & tbe Seychelles. London, 19 + L + 557 pp. (Ficus: 283- 286) (1877). Baker, J.G. - Contributions to the flora of Madagscar. Part i. Polypetalae. 1. Linn. Soc., Bot. 20, 87-304 (Ficus: 257- 262 (1883). Baker, J.G. - Further contributions to tbe flora of Madagascar. Second and final part. J. Linn. Soc. , Bot. 21, 407-455 (Ficus: 443-445) (1885). Baker, J.G. - Further contributions to the flora of Madagascar. J. Linn. Soc., Bot. 22, 441-537 (Ficus: 521-524) (1887). Baker, J.G. - Furtber contributions to the flora of Madagascar. J. Linn. Soc., Bot. 25, 211 - 306 (1889) , 307-483 (Ficus: 345- 346) (1890). Baker, J.G. - Flora of Aldabra islands. Bull. Misc. Inf. Kew (1894), 146--151 (Ficus: 151) (1894). Balfour, LB. - Diagnoses plantarum novarum Pbanerogamarurn Socotrensium etc. Pars tertia. Proc. Roy. Soc. Edinburgh 12, 76--98 (Ficus: 96) (1884). Barker, N.P. - Evidence of a volatile attractant in Ficlls ingens (Moraceae). Bothalia 15, 607-611 (1985). Battandier, J.-A. & L. Trabut - Contribution à la flore du pays de Touaregs. Bull. Soc. Bot. France 58, 623-629, 669-677 (Ficlls: 628, 674-676) (1912). Berg, e.C. - Three new African Ficus species and a comrnent on FicllS gnapha/ocarpa (Moraceae). Adansonia (2) 20, 263- 272(1980).

275 Berg, e.C. - The reinstatement of the West African Ficus lutea VahI (Moraceae). Kew Bull. 36, 597-598 (1981). Berg, C.e. - Moraceae. In: J.F. Brunei, P. Hiepko & H.Schulz, Flore analytic du Togo - Phanérogames. Englera 4, 355-366 (1984b). Berg, e.e. - The Ficus species (Moraceae) of Madagascar and the Comoro Islands. Bull. Mus. natn. Hist. nat. Paris (4) 8 (B) [= Adansonia I], 17-55 (I 986a). Berg, C.C. - Subdivision of Ficus subg. Urostigma sect. Galoglychia (Moraceae). Proc. Kon. Ned. Akad. Wet. (C) 89, 121- 127 (I 986b). Berg, C.C. - New taxa and combinations in Ficus (Moraceae) of Africa. Kew Bull. 43, 77-97 (1988). Berg, e.C. - Distribution of African taxa of Ficus (Moraceae). Mitt. Inst. Allg. Bot. Hamburg 23a: 401-405 (199Ob). Berg, e.e. - Annotated check-list of the Ficus species of the African floristic region, with special reference and a key to the taxa of southem Africa. Kirkia 13: 253-291 (I 99Oc). Berg, e.e. & E.C.H. van Heusden - Moraceae. Flore des Mascareignes 164, 21 pp. (1985). Berg, C.e. & M.E.E. Hijman - Moraceae. Flora of East Africa Il (Ficus, pp. 43-86) (1989). Berg, C.e. & M.E.E. Hijman - Flora Zambesiaca (1991 , in press). Berg, e.e., M.E.E. Hijman & J.e.A. Weerden burg - Moracées (incl. Cécropiacées). Flore du Ga­ bon 26, 276 pp. (1984). Berg, C.e., M.E.E. Hijman & J.e.A. Weerdenburg - Moracées (incl. Cécropiacées). Flore du Ca­ meroun 28, 296 pp. (1985). Berg, C.e., G. Michaloud, G . Couturier, D. Lachaise & L. Tsacas - La plante-hote: Ficus spp., Moracées. Courrier de l'Ecotrop, Num. spéc., 11-18 (1984). Berhaut, J. - Flore illustrée du Sénégal. Dakar, 636 pp. (Ficus: 419-495) (1979). Blume, e.L. - Bijdragen tot de flora van Nederlandsch Indië. Batavia, x + 1169 pp. (Ficus: 456-479) (1825-1827). B1ume, C.L. - Rumphia 2, Leiden, viii + 176 pp. (1836). Bory, 1.B.G.M. - Voyage dans les quatres principales les des mers d'Afrique 2. Paris, iv + 431 pp. (1804). Boucek, Z. - A study of the non-podagrine Torymidae with enlarged hind femora, with a key to the African genera (Hymenoptera). J. ent. Soc. sth. Afr. 41: 91 - 134 (1978). Boucek, Z., A. Watsham & J.T. Wiebes - The tig wasp fauna of the receptacles of Ficus thonningii (Hymenoptera, Chalcidoidea). Tijdschr. Ent. 124, 149- 233 (l98\). Boutique, R. - Une espéce nouvelle de Ficus du Congo Beige. Bull. Jard. Bot. Brux. 19, 211 - 212 (1949). Boutique, R. & J. Léonard - Notuiae Systematicae ii. Ficus nouveaux du Congo BeIge. Bull. Jard. Bot. Brux. 19, 215- 218 (1949). Breitwisch, R. - Frugivores at a fruiting Ficus vine in a southem Cameroon tropical wet forest. Biotropica IS, 125-128 (1983). Braun, J. & K. Schumann - Ficus pringsheimiana. In: 1. Braun, Botanischer Bericht über die Flora von Kamerun. Mitt. aus den Deutschen Schutzgeb. 2, 141-176 (1889). Britton, N.L.- Studies on West Indian plants - x. 60 undescribed species from Trinidad. Bull. Torrey Bot. Club 48, 327- 340 (Ficus: 330) (1922). Brown, N .E. - New plants from Tropical and Sou th Africa collected by Archdeacon F.A. Rogers. Bull. Misc. Inf. Kew (1921), 298- 301 (Ficus: 297) (1921). Burtt Davy, J. - Notes on the genus Ficus (Toum). Linn. Trans. Roy. Soc. S. Afr. Il, 365-368 (1912). Burtt Davy, J. - A manual of the flowering plants and fems of the Transvaal [?] with Swaziland. London-New York-Toronto, 2, xxiii + 273-529 (Ficus: 438-442) (1932). Carmin, J. & D. Scheinkin - The fauna of Palestinian plants: 2. Ficus sycomorus Linné. Buil. Soc. Roy. Ent. Egypte, N.S. IS, 164-187 (1932). Chevalier, A. - Botanique coloniale - sur quelques plantes à caoutchouc de la cote occidentale d'Afrique. Compt. Rend. hebd. Séanc. Acad. Sci. 135, 441-443 (1902).

276 Chevalier, A. - Les végétaux utiles de I'Afrique seientifiques et agronomiques 5, 315 pp. (Ficus: 262- 263) (1909). Chevalier, A. - Études sur la flore de I'Mrique centrale Française. Paris, 1, xü + 451 pp. (Ficus: 284-291) (1913). Chevalier, A. - Novitates florae Mricanae. Urticaceae. Bull. Soc. Bot. France (Mém. 8), 297- 302 (Ficus: 300-301) (1917). Chevalier, A. - Exploration botanique de I'Mrique Occidentale Française. Paris, 1, xüi + 798 pp. (Ficus: 593--606) (1920). Chevalier, A. - Sur deux Ficus africaines présentant de curieuses particularités biologiques. Rev. Int. Bot. Appl. Agr. Trop. 15, 456-459 (1935). Chevalier, A. - Vn Ficus nouveau d'Mrique Occidentale. Rev. Int. Bot. Appl. Agr. Trop. 29, 246-249, t. II (1949). Chiovenda, E. - Flora Somala 1. Roma, xvi + 436 pp. (Ficus: 312-313) (1929). Compton, R.H. - Flora of Swaziland. J. South Mr. Bot., Suppl. 11, 1--648 (Ficus: 175-176) (1976). Compton, S.G. - A collapse of host specificity in some Mrican lig wasps. S. Mr. J. Sci. 86, 39-40 (1990). Compton, S.G. & R .G .E. Baker - Figs and lig wasps. In: R.G.E. Baker, K. Richards & C.A. Rimes (eds.), Final report of the Huil University Cameroun Expedition. Mise. Publ. Huil Vniv. Dept. Geog. Mise. Ser. 30: 52--64 (1986). Compton, S.G., K.C. Holton, V.K. Rashbrook, S. van Noort, S.L. Vincent & A.B. Ware - Studies on Ceratosolen galili, a non-pollinating Agaonid lig wasp. Biotropica 23, 188-194 (1991). Compton, S.G . & F.A.C. McLaren - Respiratory adaptations in some male lig wasps. Proc. Kon. Ned. Akad. Wet. (C) 92, 57-71 (1989). Compton, S.G. & R. Nefdt - Extra-long ovipositors in Chalcid wasps: some examples and obser­ vations. Antenna 12, 102- 105 (1988). Compton, S.G. & R.J.C. Nefdt - The ligs and lig wasps of Ficus burtt-davyi. Mitt. Inst. AlIg. Bot. Hamburg 23a: 441-450 (1990). Copland, M.J.W. & P.E. King - The structure of the female reproductive system in the EurytoI)lÏ­ dae (Cha1cidoidea: Hymenoptera). J. Zool. (London) 166: 185-212 (1972). Copland, M.J.W., P.E. King & D.S. Hili - The structure of the female reproductive system in the Agaonidae (Cha1cidoidea, Hymenoptera). J. Ent. (London) (A) 48,25- 35 (1973). Coquere1, Ch. - Deseription de parasites anormaux d'un liguier de I'île de Bourbon. Rev. Mag. Zool. (2) 7, 365-371,422-427 (1855). Cordemoy, E. Jacob de - Flore de I'île de la Réunion. Paris, xxvii + 547 pp. (Ficus: 270-274) (1895). Couturier, G . & D. Lachaise - Le peuplement d'insectes en forêt de Taï (Cöte d'lvoire): l. Modi­ lications du peuplement dues à I'intervention hurnaine; 11. Les insectes associés aux liguiers en föret de Taï: Ie témoignage d'une longue histoire commune. Bull. Soc. ent. France 87, 384-388 (1983). Dale, l.R. & P.J. Greenway - Kenya trees and shrubs. Nairobi-London, xxvii + 654 pp. (Ficus: 312-324) (1961). Dalrnan, J.w. - Nägra nya Genera och Species af Insektar. KongI. Vetensk. Acad. Handl., Stock­ holm 39, 69-80 [+ TiIlägg och Rätte1ser, without page-nurnber: addition) (1818). Decaisne, J. - Énurnération des plantes recueillies par M. Bové dans les deux Arabies, la Palestine, la Syrie et I'Égypt. Florula Sinaica. Ann. Sci. Nat. Bot. (2) 2, 239-270 (Ficus: 242- 243) (1834). De Candolle, A.-P. - Deseription d'une nouvelle espèce de liguier (Ficus saussureana). Mem. Soc. Phys. & Hist. nat. Genève 9, 65-73 (1841). Delile, A. Raffenau - Note §ur que1ques plantes nouvelles d' Abyssinie. Ann. Sci. Nat. Bot. (2) 20, 88- 95 (1843). DeWe, A. Raffenau - Centurie de plantes d'Mrique du voyage à Méroé. Paris, 112 pp. (Ficus: 62--64) (1826).

277 De Wildeman, É.AJ. - Über einige neue Ficus-arten aus dem belgischen Kongo. Repert. Spec. Nov. Regni Veg. 12, 193- 200 (1913a). De Wildeman, É.AJ. - Über einige neue Ficus-arten aus den belgischen Kongo ii. Repert. Spec. Nov. Regni Veg. 12, 302-304 (l913b). De Wildeman, É.A.J. - Notes sur les Ficus de la Flore du Congo Beige. Bull. Soc. Roy. Belg. 52, 197- 236 (1914). De Wildeman, É.AJ. - Ficus. Ann. Soc. Sci. Brux. 40, 277-283 (1921). De Wildeman, É.A.J. - Plantae Bequaertianae 1. Paris, iii + 593 pp. (Ficus: 328- 351) (1921- 1922). De Wildeman, É.AJ. & Th. Durand - Reliquiae Dewevreanae. Ann. Mus. Congo (3) 1 (2), 1-291 (Ficus: 215-217) (1901). Dinter, K. - Deutsch-Südwest-Afrika. Flora. Forst- und Landwirtschaftliche Fragmente. Leipzig, viii + 191 pp (1909). Dowsett-Lemaire, F & F White - New and noteworthy plants from the evergreen forests of Malawi. Bull. Jard. Bot. Natn. Belg. 60, 73-110 (1990). Drummond, R.B. - A list of trees, shrubs and woody c1imbers indigenous or naturalised in Rho­ desia. Kirkia 10: 229- 285 (1975). Eggeling, WJ. & l.R. Dale - The indigenous trees of the Uganda Protectorate, ed. 2. Entebbe­ London, 12 + xxx + 491 pp (Ficus: 237- 262) (1952). Engler, G.H.A. - Beiträge zur Flora des Congogebietes gesammelt von Dr. Naumann auf der Expedition S.M.S. 'Gazelle'. Bot. Jahrb. Syst., Pfianzengesch. & Pfianzengeogr. (EngIer) 8, 59- 68 (Ficus: 59-60) (1886). Engier, G.H.A. - Plantae Marothianae. Ein Beitrag zur Kenntniss der Flora Südafrikas. Bot. Jahrb. Syst., Pfianzengesch. & Pfianzengeogr. (EngIer) 10, 1- 50 (Ficus: 5) (1888). Engier, G.H.A. - Plantae GÜrichianae. Ein Beitrag zur Kenntniss der Flora von Deutschsüdwestafrika . Bot. Jahrb. Syst., Pflanzengesch. & Pflanzengeogr. (EngIer) 19, 129-152 (Ficus: 130) (1894). Engier, G.H.A . - Die Pfianzenwelt Afrikas insondere seiner tropischen Gebiete. Charakterpflanzen Afrikas 2. Die dikotyledonen Angiospermen. Casuarinaceae bis Dichapetalaceae. In: G.G.H. Engler & O. Drude, Die Vegetation der Erde 9 (3, I). Leipzig, vi + 869 pp. (Ficus: 34-51) (1915). ExelI, A.W. - Catalogue of vascular plants of S. Tomé (with Principe and Annabon). London, xi + 428 pp. (Ficus: 306-308) (1944). Ficalho, FM.e. de - Plantas uteis da Africa Portugueza. Lisboa, 279 pp. (Ficus: 269-271) (1884). Fiori, A. - I sicomori dell'Eritrea. Bull. R. Soc. Toscana Orticult. 35, 162- 167 (19IOa). Fiori, A. - Boschi epiante legnose deU'Eritrea. Agric. Colon. 4 (8), 365- 386 (191Ob). Forsskäl, P. - Flora aegyptiaco-arabica. Hauniae, 32 + cxxvi + 220 pp. (Ficus: 269-271) (1775). Fosberg, FR. & S.A. Renvoize - The flora of Aldabra and neighbouring islands. Kew Bull., Add. Ser. 7, 358 pp. (Ficus: 272-273) (1980). Fries, R.E. - Wissenschaftliche Ergebnisse der Schwedischen Rhodesia-Kongo-Expedition 1911- 1912. Band J. Botanische Untersuchungen. Heft 1, Pteridophyta und Choripetalae, 1-184 (Ficus: 14-17) (1914). Friis, l. - Notes on Somalian species of Ficus (Moraceae). Nordic J. Bot. 5: 331-333 (1985). Friis, l. - Moraceae. In l. Hedberg & S. Edwards. Flora of Ethiopia 3. Addis Ababa-Asmara­ Uppsala, Ixxi + 659 pp. (Ficus: 278-299) (1989). Friis, l. - Distribution pattems of Ficus in Tropical N.E. Africa. Mitt. lnst. AUg. Bot. Hamburg 23a: 407-424 (1990). Galil, J. - [The sycomore tree in the civilisation of the Middle East]. Teva Wearetz 8, 306-318, 338-355 (in Hebrew) (1966). Galil, J. - Sycomore wasps from ancient Egyptian tombs. Israel J. Ent.' 2, 1-10 (1967). GaliI, J. - An ancient technique for ripening sycomore fruit in East-Mediterranean countries. Econ. Bot. 22, 178- 190 (1968). GaliI, J. - Sycomore fig . In: CRC Handbook of Fruit Set and Development, pp. 449-460 (1986).

278 Galil, J. - Adjustment of developmental spans in organisms participating in fig symbioses. Mitt. Inst. AlIg. Bot. Hamburg 23a: 433--440 (1990). Galil, J. & D. Eisikowitch - On the pollination ecology of Ficus sycomorus in East Africa. Ecology 49, 259- 269 (1968a). Galil, J. & D. Eisikowitch - Flowering cydes and fruit types of Ficus sycomorus in Israel. New Phytol. 67, 745-758 (I 968b). Galil, 1. & D. Eisikowitch - Further studies on the pollination ecology of Ficus sycomorus L. (Hymenoptera, Chalcidoidea, Agaonidae). Tijdschr. Ent. 112, 1-13 (1969a). Galil, J. & D. Eisikowitch - Note on pollen transport, pollination and protection of ovaries in Ficus sy comorus. New Phytol. 68, 1243-1244 (l969b). Galil, J. & D. Eisikowitch - Further studies on the pollination ecology in Ficus sycomorus ii. Pocket filling and emptying by Ceratosolen arabicus Mayr. New Phytol. 73, 515- 528 (1974). Galil, J., M. Stein & A. Horowitz - On the origin of the sycomore fig (Ficus sycomorus L.) in tbe Middle East. Gdns' Bull., Singapore 29, 191- 205 (1977). Gasparrini, G. - Nova genera, quae super nonnullis Fici speciebus struebat. Napoli, 11 pp. (1844). Gasparrini, G. - Ricerche suil a natura del Caprifico, e del Ficus e sulla caprificazione. Napoli, i + 96 pp. + 18 tab. (1845). Gautier-Hion, A. & G. Michaloud - Figs: are they keystone resources for frugivorous vertebrates throughout the tropics? A test in Gabon. Ecology 70, 1826-1833 (1989). Geerling, C. - Guide de terrain des ligneux Sahéliens et Soudano-Guinéens. 340 pp. (Ficus: 228- 241) (1982). Gerstner, FJ. - Ficus hippopotami. In: W.F Barker, FJ. Gestner, R.S. Adamson & R.H. Comp­ ton, Plantae novae Africanae series XX. J. South Afr. Bot. 9, 145- 163 (1943). Ghesquière, J. - Contribution a I'étude des Hyménoptères du Congo beige x. Nouvelles dénomina­ tions pour quelques genres de Chalcidoidea et Mymaroidea. Rev. Zool. Bot. Afr. 39, 367-371 (1946). Grandi, G. - Gli Agaonini (Hymenoptera Chalcididae) raccolti nell'Africa occidentale dal Prof. F Silvestri. Boll. Lab. Zool. Portici 10, 121- 286 (l916a). Grandi, G. - Contributo alla conscenza degli Agaonini (Hymenoptera Chalcididae) dell'Eritrea e dell'Uganda. Bull. Soc. ent. Ital. 48, 3-42 (1917). Grandi, G. - Diagnosi preliminari di Imenotteri dei Fichi. Ann. Mus. Stor. nat. Genova (3) 9, 304-316 (l92la). Grandi, G. - Ricerche sul Gen. Philotrypesis Först. (Hymenoptera - Chalcididae). Boll. Lab. Zool. Portici 15, 33- 190 (192lb). Grandi, G . - Ricostruzione e morfologia comparata dei generi Otitesella Westw., Sycobiella Westw. ed affini (Hymenoptera - Chalcididae). Boll. Lab. Zool. Portici 16, 3- 58 (1922). Grandi, G . - Un nuovo Agaonino dell'isole Maurizio. Boll. Soc. ent. Ital. 56, 81-87 (1924). Grandi, G. - Due specie di Blastophaga delle isole Figi ed istituzione di un nuova sottogenere. Boll. Lab. Ent. Bologna I, 65- 70 (1928a). Grandi, G. - Revisione critica degli Agaonidi descritti da Gustavo Mayr e catalogo ragionato delle specie fino ad oggi descritte di tutto il mondo. Boll. Lab. Ent. Bologna I , 107-235 (I 928b). Grandi, G. - Monografia del gen. Philotrypesis Först. Boll. Lab. Ent. Bologna 3, 1-181 (1930). Grandi, G. - Insetti dei fichi senegalesi. Boll. Ist. Ent. Univ. Bologna 19, 31-45 (1952). Grandi, G. - Insetti dei fichi (Hymenoptera - Chalcididae) dell'Africa australe. Boli. Ist. Ent. Univ. Bologna 21 , 85-106 (1955). Grandi, G. - Una nuova specie di Ceratosolen Mayr dell'Africa occidentale (Hymenoptera Chal­ cidoidea). Boli. Ist. Ent. Univ. Bologna 26, 231 - 238 (1963a). Grandi, G. - Catalogo ragionato degli Agaonidi del monde descritti fino a oggi (6a edizione). Boll Ist. Ent. Univ. Bologna 26, 319-373 (I 963b). Grandi, G . - Descrizione del maschio di un Ceratosolen Mayr (Hymenoptera - Chalcidoidea - Agaonidae) rinvenuto in Africa (Costa d'Avorio) fra il pelame di un Megachirottero (Hypsigna­ thus monstruosus Allen). Boll. Ist. Ent. Univ. Bologna 27, 71 - 75 (1964).

279 Greuning, J.v. van - 'n Morfologiese studie van die genus Ficus L. in suidelike Afrika. Thesis, Univ. Pretoria, iii + 265 pp. (1982). Greuning, 1.v. van - A synopsis of the genus Ficus (Moraceae) in southem Africa. S. Afr. J. Bot. 56, 599--630 (1990). Guy, P.R. - Notes on the host species of epiphytic ligs (Ficus spp.) on the ftood-plain of the Mana Pools Game Reserve. Kirkia 10, 559- 562 (1977). Hasselquist, F. - Iter Palaestinum. Elle Resa till Heiige Landet förrattad ifran Ár 1749 til 1752, med Beskrifningar, Rön, Anmarkningar ofver de markvardigaste Naturalier. Stockholm, xvi + 619 pp. (1757). Hasselquist, F. - Reise nach Pälästina in den Jahren von 1749 bis 1752. Auf Befehl Ihro Majestät der Königinn von Schweden herausgegeben von Carl Linneaus. Rostock, 17 + 606 pp. (1762). Hiem, WP. - Catalogue of the African plants collected by Dr. Friedrich Welwitseh 4. London, 785--1035 (Ficus: 996--10 19) (1900). Hochstetter, C.F. - Moreae. In F. Krauss, Pftanzen des Cap- und Natal-Landes, gesammelt und zusammengestelt vomn Dr. Ferdinand Krauss. (Fortsetzung). Flora 28 (6), 88 (1845). Homby, AJ.W. - A new species of Ficus from the borders of Moçambique and Nyasaland. Bo­ thalia 4, 1007-1008 (1948). Hutchinson, J. - New tropical African species of Ficus. Bull. Mise. Inf. Kew, 313-344 (1915). Hutchinson, J. - Ficus Linn. In: David Prain, Flora of Tropical Africa VI (2), pp. 78-192 (1916), 193- 215 (1917). Hutchinson, J. & J. Burtt Davy - Ficus burll-davyi. In: J. Burlt Davy, iii. New or noteworthy South African plants (i). Bull. Mise. Inf. Kew (no. 2), 49- 52 (1921). Hutchinson, 1. - Ficus. In: WT. Thiselton-Dyer, Flora Capensis 5 (2). London, viii + 606 pp. (Ficus: 523-541) (1920-1925). Joseph, K.J. - On a collection of lig insects (Chalcidoidea: Agaontidae) from French Guinea. Proc. R. ent. Soc. London (B) 28, 29- 36 (1959). Keay, R.WJ. - Flora of Tropical West Africa, ed. 2, 1. London, viii + 628 pp. (Ficus: 6()(kj11, 1958) (1954-1958). Keay, R.WJ. - Trees of Nigeria. Clarendon Press, Oxford, viii + 476 pp. (Ficus 288-298) (1989). KiefTer, J.-J. - Deseriptions de deux nouveaux Chalcidites (Hym.). Ann. Soc. ent. France 80, 463- 466 (1911). Kunth, C.S. & c. Bouché, In: Kunth, C.S. - Enumeratio synoptica Ficus specierum cum novarum turn cognitarum Horti Regii Botanici Berolinensis. Index Seminum Horto Botanico Berolinensi 1846, 14-22 (1847). Lamarck, J.B.A.P.M. de - Encyclopédie méthodique. Botanique 2. Paris, 774 pp. (Ficus: 489- 5(0) (1788). Lebrun, J. - Les espéces congolaises du genre Ficus L. Mém. Inst. Roy. Colon. Beige, Sect. Sci. Nat. et Med., in 8°, 3: 1-79 (1934). Lebrun, J. - Les Figuiers du Congo. Rev. Int. Bot. Appl. Agr. Trop. 15, 453-456 (1935a). Lebrun, J. - Note complémentaire sur les Ficus du Congo. Bull. Scéances, Inst. Roy. Col. Beige 6, 494-497 (1935b). Lebrun, J. & R . Boutique - Ficus. Flore du Congo Beige et du Ruanda-Urundi, 1: 98-174 (1948). Lebrun, J. & R. Boutique - Un Ficus nouveau pour Ie Congo Beige. Bull. Jard. Bot. Brux. 19, 213- 214 (1949). Lebrun, 1. & L. Toussaint - Ficus. In: J. Lebrun, A. Taton & L. Toussaint, Exploration du Parc National de la Kagera. Mission J. Lebrun (1937- 38), 39-44 (1948). Link, H.F. - Enumeratio plantarum hortii regii boatnici Berolinensis altera 2. Berolini, iv + 478 pp. (Ficus : 448-451) (1822). Martenelli, U. - Florula Bogonensis. Firenze, vii + 169 pp. (Ficus: 75--78) (1886). Masi, L. - Chalcidoidea ofthe Seychelles Islands. Novit. Zool. 24,121- 230 (1917). Masi, L. - Missione biologica Sagan - Omo diretta dal prof. Edoardo Zavattari (1939). Hymenop­ tera Chalcidoidea. Ann. Mus. Civ. Stor. nat. Genova 65, 173- 210 (1951).

280 Mayer, P. - Zur Naturgeschichte der Feigeninsekten. Mitt. zool. Sta. Neapel 3, 551-590 (1882). Mayr, G. - Feigeninsecten. Verh. zool.-bot. Ges. Wien 35, 147- 250 (1885). Mayr, G. - Neue Feigen-Insekten. Wien. ent. Ztg. 25, 153-187 (1906). Medler, l.T. - Insects of Nigeria; check list and bibliography. Mem. Amer. Ent. Inst. 30, 1- 919 (1980). Michaloud, G. - Figuiers tropicaux et pollinisation. Film, réalisation A.R. Devez, production Service du Film de Recherche Scientifique, Paris (1982). Michaloud, G. - Fighting in fig wasps. Trends in Ecology and Evolution 3, 77 (1988a). Michaloud, G. - Aspects de la reproduction des figuiers monoiques en forêt équatoriale africaine. Thèse d'Universlte du Languedoc, 256 pp. (I 988b). Michaloud, G . & S. Michaloud-Pelletier - Ficus hérni-épiphytes (Moraceae) et arbres supports. Biotropica 19, 125-136 (1987). Michaloud, G ., S. Michaloud-Pelletier, l.T. Wiebes & c.c. Berg - The co-occurrence of two pol­ linating species of tig wasp and one species of fig. Proc. Kon. Ned. Akad. Wet. (C) 88, 93-119 (1985). Mildbraed, 1. - Neue Ficus-Arten aus dem tropischen Afrika. Mitt. Bot. Gart. Berlin-Dahlem 1 (I), 126--128 (1953). Mildbraed, l. & M. Burret - Ficus. In: l . Mildbraed, Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907- 1908. Band 2, 182- 189 (l91Ia). Mildbraed, l . & M. Burret - Die Afrikanischen Arten der Gattung Ficus Linn. Bot. lahrb. Syst., Pflanzengesch. & Pflanzengeogr. (Engier) 46, 163-269 (I911b) [also indicated as 1911 ; it was published prior to the paper mentioned above as 191Ia!). MiqueJ, FA.G. - Prodromus Monographiae Ficuum. London l . Bot. 6, 514-588, t. 21 - 23 (I847); 7, 588, t. 2 {I 848). Miquel, FA.G. - Over de Afrikaansche Vijge-bomen. Verh. Eerste Kl. Kon. Ned. Inst. Wetensch. Amst. (3) 1, 111- 150, t. 1- 5 (1849). MiqueJ, FA.G. - Ficus. In: W.J. Hooker, Niger Flora, 520-525 (I849). Miquel, FA.G. - Annotationes de Ficus speciebus. Ann. Mus. Bot. Lugd.-Bat. 3, 260-315 {I 867). Nefdt, Roy - Fig wasps and their parasitoids. Rostrum (Newsletter Ent. Soc. Southem Africa) 17, [8] (1987). Nefdt, R.l.C. & S.G. Compton - Parasitoid pressure as a possible factor influencing ovipositor length in Agaonid fig wasps. Proc. 6th Ent. Congr., Ent. Soc. Southem Africa (50th Ann.), Stellen bosch, 102 (1987). Newton, L.E. & A. Lomo - ReJationship between pollen load and flight in agaonid wasps. Ent. Ber. Amst. 37, 72- 73 {I 977). Newton, L.E. & A. Lomo - The pollination of Ficus vage/ii in Ghana. Bot. l. Linn. Soc. 78, 21-30 (I979). Noort, S. van & S.G. Compton - Two new species of Otitesella (Hymenoptera, Chalcidoidea, Pteromalidae) from Ficus burtt-davyi. Proc. Kon. Ned. Akad. Wet. (C) 91 : 419-427 (1988). Noort, S. van, A.B. Ware & S.G. Compton - Pollinator-specific volatile attractants reJeased from the figs of Ficus burtt-davyi. S. Afr. l . Sci. 85, 323-324 (1989). Palgrave, K. Coates - Trees of Southem Africa. Cape Town-lohannesburg, 958 pp. (Ficus: 102- 119) (1977). Palmer, E. & N. Pitman - Trees of South Africa l. Cape Town, xv + 703 pp. (Ficus: 439-483) (1972). Pampanini, R. - Missione scientifica Stefanini-Paoli nella Somalia settentrionale (1913) - Contri­ buto all conoscenza della flora Somalia. Bull. Soc. Bot. Ital. 1915 (1-2), 12- 18 (Ficus: 14-15) (1915). Perrier de la Bäthie, H. - Le Ficus de Madagascar. Arch. Bot., Bull. Mens. 2 (8-9), 137- 180 (1928). Perrier de la Bäthie, H. - Un Ficus nouveau de Madagascar. Not. Syst. 14, 167 (1951). Perrier de la Bäthie, H. - Ficus Tourn. ex L. In: H. Humbert, Flore de Madagascar et des Comores (Plantes vasculaires) 55e Familie - Moracées. Paris, 40-71 (1952).

281 Peter, A. - Flora von Deutseh-Ostafrika 2 (I). Repert. Spec. Nov. Regni Veg., Beihefte 40 (2, I), 1- 272, incl. Deseriptiones plantarum novarum Africanae, 1-112 (Ficus: 85-112) (1932). RalTenau-Delile, A. - see Delile, A. RalTenau Ramcharun, S., H. Baijnath & J.v. Van Greuning - Some aspects of the reproductive biology of the Ficus natalensis complex in Southem Africa. Mitt. Inst. AIlg. Bot. Hamburg 23a: 451-455 (1990). Richard, A. - Tentamen Florae Abyssinicae, 518 pp. (Ficus: 265-272) (\851). Risbec, J. - Les Chalcidoïdes d'A.O.F Mém. Inst. franç. Afr. noire 13, 1-409 (195Ia). Risbec, J. - Chalcidoïdes d'A.O.F Bull. Inst. franç. Afr. noire 13, IJ 10-1130 (1951 b). Risbec, J. - Chalcidoïdes et Proctotrupoïdes de I'Afrique occidentale française (3e supplément). Bull. Inst. franç. Afr. noire 16, 524-552 (1954a). Risbec, J. - Chalcidoïdes et Proctotrupides de l'Afrique tropicale française (4e supplément). Bull. Inst. franç. Afr. noire 16, 1035-1092 (1954b). Risbec, J. - Chalcidoïdes et Proctotrupides africaines. Bull. Inst. franç. Afr. noire (A) 17: 533-580 ( 1955a). Risbec, J. - Chalcidoïdes du Kenya (Mission de I'Omo). Mém. Mus. natn. Hist. nat. Paris (A) 8, 153- 175 (1955b). Risbec, J. - Torymidae et Agaonidae de Madagasear (Hym. Chalcidoidea). Ann. Soc. ent. France 124, 147- 194 (1956). Risbec, J. - Chalcidoïdes et Proctotrupides de l'Afrique occidentale française (Se supplément). Bull. Inst. franç. Afr.noire 19, 228- 267 (1957). Roxburgh, W. - Flora Indica, ed. 1832, 3. Calcutta-London, viii + 875 pp. (1832). Sabine, J. - Some account of the edible fruits of Sierra Leone. Trans. Hort. Soc. London 5, 439-466 (Ficus: 448-449) (1824). Saunders, S.S. - On the ha bits and affinities of Apocrypla and Sycophaga, of the Hymenopterous family Agaonidae, with description of a new species of Apocrypta from the figs of Ficus Sycornori of Egypt. Trans. ent. Soc. London for 1878,313- 320 (1878). Saunders, S.S. - Descriptions of three new genera and species of fig insects allied to Blaslophaga from Calcutta, Australia and Madagascar; with notes on their parasites and on the affinities of the respective races. Trans. ent. Soc. London for 1883, 1- 27 (\883). Schumacher, Fe. - Beskrivelse af Guineiske Planter som ere fundne af Danske botanikere, issaer af Etatsraad Thonning. Kjöbenbavn, 466 pp (Ficus: 25-28) (1827). Sim, T.R. - Forest Flora and Forest Resources of Portuguese East Africa. Aberdeen, vi + 166 pp., 100 pI. (Ficus: 98- 102, pI. 87-95) (1909). Sonder, W. - Beiträge zur Flora von Südafrika. Linnaea 23, 1-138 (Ficus: 136-138) (1850). Sprague, R. - New or notewortby plants. Gard. Chrono 33, 354-356 (Ficus: 354) (1903). Stapf, o. - Appendix ivo List of tbe known plants of Liberia. In: H. Jobnston, Liberia 2, 570-668 (1906). Surnmerhayes, V.S. - New plants from tbe Seycbelles. Bull. Mise. Inf. Kew (1928), 38-394 (Ficus: 393) (1928). Tenore, M. - Decrizione del Ficus galactophora Ten. (Galactodendron seciosurn Cels.). Atti ReaJe Accad. Sci. Sez. Soc. Reale Borbon 6, 157- 163, with plate (1851). Tbunberg, e.P. - Dissertationes botanicae: D.d. Ficus genus (E. Gedner). Upsaliae, 16 pp (1786). Troupin, G. - Flore du Rwanda. Spermatopbytes, 1. Tervuren, xiii + 413 pp. (Ficus: 135- 148) (1978). Troupin, G . - Flore des plantes Iigneuses du Rwanda. Tervuren, xi + 747 pp. (Ficus: 434-450) (1982). UIen berg, S.A. - Morphological deseription of Apocrypta perplexa Coquerel, the type-species of the genus (fig wasp parasites; Hymenoptera, Chalcidoidea, Torymidae). Proc. Kon. Ned. Akad. Wet. (C) 86, 63-94 (1983). UIen berg, S.A. - The systematics of the fig wasp parasites of the genus Apocrypta Coquerel. Verh . Kon. Ned. Akad. Wet. , 2de reeks 83, 1- 176 (1985).

282 UIenberg, S.A. & B.K. Nübel - The oviposition of Apocrypta Coquerel (tig wasp parasites; Hy- menoptera, Chalcidoidea, Torymidae). Proc. Kon. Ned. Akad. Wet. (C) 85, 607--613 (1982). VahJ, M.H. - Symbolae Botanicae 1. Hauniae, viii + 85 pp. (Ficus: 82-84) (1790). Vahl, M .H. - Enumeratio Plantarum 2. Hauniae, viii + 423 pp. (Ficus: 181-204) (1805). Verdoom, I.e. - Ficus smutsii. In: Notes on the flora of Southem Africa, vi. Bul!. Mise. lof. Kew (1935), 205 (1935). Verkerke, W. - Anatomy of Ficus olloniifolia (Moraceae) syconia and its role in the tig-tig wasp symbiosis. Proc. Kon. Ned. Akad. Wet. (C) 89, 443-469 (1986). Verkerke, W. - Ovule dimorphism in Ficus asperifolia Mique!. Acta bot. Neer!. 36, 121-124 (1987a). Verkerke, W. - Syconial anatomy of Ficus asperifolia (Moraceae), a gynodioecious tropical fig. Proc. Kon. Ned. Akad. Wet. (C) 90, 461-492 (l987b). Verkerke, W. - Flower development in Ficus sur Forsskal (Moraceae). Proc. Kon. Ned. Akad. Wet. (C) 91 , 175- 195 (1988a). Verkerke, W. - Functional anatomy of Ficus asperifolia, a gynodioecious tropical lig. Acta Bot. Neer!. 37: 324 (l988b). Verkerke, W. - Sycone morphology and its influence on the flower structure of Ficus sur (Mora­ ceae). Proc. Kon. Ned. Akad. Wet. (C) 91,319-344 (1988c). Verkerke, W. - Fig anatomy and reproductive biology of African Ficus species (Moraceae). Mitt. lost. Allg. Bot. Hamburg 23a: 427-431 (1990). Warburg, O. - Moraceae africanae ii. Ficus. Bot. Jahrb. Syst., Pflanzengeseh. & Pflanzengeogr. (Engier) 20, 152-175 (1894). Warburg, o. - Ficus goetzei. In: G .H.A. Engier, Berichte über die botanisehen Ergebnisse der Nyassa-See und Kinga-Gebirgs-Expedition. iii. Die von W. Goetze und Dr. Stuhlman im Ulugu­ rugebirge, sowie von W. Goetze in der Kisaki- und Khutu-Steppe und in Uhehe gesarnmeIten Pflanzen. Bot. Jahrb. Syst., P/lanzengeseh. & Pflanzengeogr. (Engier) 28, 378 (1900). Warburg, O. - Ficus. In: G.H.A. Engier, Berichte über die botanischen Ergebnisse der Nyassa-See und Kinga-gebirgs-Expedition ivo Die von W. Goetze am Rukwa-See und Nyassa-See, sowie in den zwisehen beiden Seen gelegenen Gebirgsländem, insbesondere dem Kinga-gebirge gesamml­ ten Pflanzen, nebst einigen Nachträge zu Bericht iii. Bot. Jahrb. Syst., pflanzengesch. & Pflan­ zengeogr. (Engier) 30, 292-295 (1901). Warburg, O. - In: O. Warburg & É.A.J. de Wildeman - Les Ficus de la flore de l'État Indépendant du Congo. Ann. Mus. Congo Bot. (6) I , 1-36, t. 1- 17 (1904). Warburg, O. - Die Gattung Ficus im nichttropisehen Vorderasien. In: I. Urban & P. Graebner, Festsehrift zur Feier des siebzigsten Geburtstages des Herm Professor Dr. Paul Ascherson, 14. Jun. 1904, lii + 568 pp. (Ficus: 364--370) (1904b [at first omitted: 1904a is still indicated in the text as 1904]). Warburg, O. - Generis Ficus species et varietates novae africanae. BotJahrb. Syst., Pflanzengesch. & Pflanzengogr. (EngIer) 36,211- 212 (1905). Warburg, O. - Moraceae. Die südafrikanisehen Arten der Gattung Ficus. Vierteljahrsehr. Natur- forsch . Ges. Zürich 51 , 132-143 (1906). Waser, P. - The fecund tig. Africana 6 (&): 23- 25 (1977). Waterston, J. - Notes on African Chalcidoidea i. Bull. ent. Res. 5, 249-258 (1914). Waterston, J. - Notes on African Chalcidoidea v. Bull. ent. Res. 7, 123- 132 (1916). Waterston, J. - A new Sycophagine (Agaonidae - Chalcidoidea) genus and species from the Gold Coast. Ent. monthly Mag. (3) 5, 274-277 (1919). Waterston, J. - Notes on tig insects, induding deseriptions of three new species and a new Blasto­ phagine genus. Trans. ent. Soc. London (1920), 128-136 (1920a). Waterston, J. - A new Blastophagine genus and species from E. Africa (Hymenoptera - Chalc­ idoidea). Ent. month1y Mag. (3) 6, 197- 200 (I 920b). Waterston, J. - On a new African lig insect (Blastophaga dyscritus, sp. n.). Trans. ent. Soc. London (1921),417-418 (l921b).

283 Watsham, A. - A new species of Sycophila Walker (Hymenoptera: Eurytomidae) from Rhodesia. 1. ent. Soc. sth. Afr. 40, 247- 250 (l977a). Watsham, A. - Fig insects: an ecological study of Cha1cids in F. capensis and F. natalensis in Rhodesia, Proc. 2nd Ent. Congr. ent. Soc. sth. Afr., 40-41 (1977b). Watsham, A. - The ecology of fig insects. Proc. 3rd Ent. Congr. ent. Soc. sth. Afr., 14 (1981). Watsham, A. - How to study the fascinating ecology of the fig tree. The Zimbabwe Science News 17, 82-87 (1983). Westwood, 1.0. - Description of several new British forms among parasitic Hymenopterous in­ sects. Lond. Edinb. Dubl. Phil. Mag. (3) 1: 127-129 (1832). Westwood, 1.0. - On caprification as practised upon the figs in the south of Europe and the Levant, with descriptions of the insects employed for that purpose; and observations upon the Agaon paradoxum of Dalman. Trans. ent. Soc. London 2, 214--224 (1840). Westwood, 1.0. - Descriptions of the insects infesting the seeds of Ficus sycomorus and carica. Trans. ent. Soc. London (1882), 47-60 (1882). Westwood, I.O. - Further descriptions of insects infesting figs. Trans. ent. Soc. London (1883), 29-47 (1883). Wharton, R.A., I.W. Tilson & . R.L. Tilson - Asynchrony in a wild population of Ficus sycomorus L. S. Afr. I. Sci. 76, 478-480 (1980). White, F. - The vegetation of Africa. National Resources Research 20, Unesco, Paris (1983). White, M.A. - Forest flora of northern Rhodesia. Oxford Univ. Press, Oxford, xxvi + 455 pp. (Ficus: 26--35) (1962). Wiebes, 1.T. - On the variability of Agaon paradoxum (Dalman) Grandi and Seres armipes Water­ ston, with remarks on other African Agaonidae (Hymenoptera, Chalcidoidea). Zool. Meded. Leiden 37, 231 - 240 (1961). Wiebes, 1.T. - Fig wasps from Israeli Ficus sycomorus and related East African species (Hymenop­ tera, Cha1cidoidea) 1. Agaonidae. Ent. Ber. Amst. 24, 187- 191 (1964). Wiebes, 1.T. - Provisional host catalogue of fig wasps (Hymenoptera, Cha1cidoidea). Zool. Verh. 83, 3-44 (1966). Wiebes, 1.T. - Redescription of Sycophaginae from Ceylon and India, with designation of lectoty­ pes, and a worId catalogue of the Otitesellini (Hymenoptera Cha1cidoidea, Torymidae). Tijdschr. Ent. 110, 399-442 (1967). Wiebes, 1.T. - Fig wasps from Israeli Ficus sycomorus and related East African species (Hymenop­ tera, Cha1cidoidea) 2. Agaonidae (conc1uded) and Sycophaginae. Zool. Meded. Leiden 42, 307- 320 (l968a). Wiebes, 1.T. - Species of Agaon from Congo (Kinshasa), with notes on synonymy (Hymenoptera, Cha1cidoidea). Proc. Kon. Ned. Akad. Wet. (C) 71 , 346--355 (1968b). Wiebes, 1.T. - Preliminary report on a collection of fig insects (Hymenoptera Cha1cidoidea) from Ficus gnaphalocarpa. Mém. Inst. fond . Afr. noire 84, 401-402 (l969a). Wiebes, 1.T. - Philosycus, a new genus of fig wasps allied to Otilesella Westwood (Hymenoptera Chalcidoidea, Torymidae). Ann. Mus. Roy. Afr. centr., in 8°, Zool. 175,439-445 (1969b). Wiebes, 1.T. - Hymenoptera Agaonidae, with an introductory chapter on West African fig wasps. Ann. Mus. Roy. Afr. centr., in 8°, Zool. 175,449-464 (1969c). Wiebes, 1.T. - Revision of the Agaonidae described by 1. Risbec, and notes on their Torymid symbionts (Hymenoptera, Cha1cidoidea). Zool. Meded. Leiden 45, 1- 16 (1970). Wiebes, 1.T. - A new record of Allotriozoon prodigiosum Grandi, and description of its symbionts (Hymenoptera, Cha1cidoidea). Mém. Inst. fond . Afr. noire 86, 367-383 (1971). Wiebes, 1.T. - A new species of Agaon from Nigeria (Hymenoptera, Cha1cidoidea). Ent. Ber. Amst. 32, 122- 124 (l972a). Wiebes, 1.T. - The genus Alfonsiella Waterston (Hymenoptera, Cha1cidoidea, Agaonidae). Zool. Meded. Leiden 47, 321-330 (l972b). Wiebes, J.T. - Nigeriella, a new genus of West African fig wasps allied to Elisabethiella Grandi (Hymenoptera Cha1cidoidea, Agaonidae). Zool. Meded. Leiden 48, 29-42 (1974a).

284 Wiebes, 1.T. - Species of Agaon Dalman and Allotriozoon Grandi from Africa and Malagasy (Hymenoptera Chalcidoidea, Agaonidae). Zool. Meded. Leiden 48, 123-143 (l974b). Wiebes, 1.T. - Fig insects from Aldabra (Hymenoptera, Chalcidoidea). Zool. Meded. Leiden 49, 225-236 (1975). Wiebes, 1.T. - A new species of Agaon from Nigeria, and some additional records (Hymenoptera, Chalcidoidea, Agaonidae). Ent. Ber. Amst. 36, 124-127 (1976). Wiebes, 1.T. - Agaonid tig wasp from Ficus salicifolia Vahl and some related species of the genus Platyscapa Motschoulsky (Hym., Chalc.). Neth. I . Zool. 27, 209- 223 (I 977b). Wiebes, 1.T. - Tbe genus Kradibia Saunders and an addition to Ceratosolen Mayr (Hymenoptera Chalcidoidea, Agaonidae). Zool. Meded. Leiden 53, 165-184 (1978a). Wiebes, 1.T. - Redescription of Sycoscapter gibbus Saunders, a parasitic tig wasp from Madagascar and Réunion (Hymenoptera, Chalcidoidea, Torymidae). Ent. Ber. Amst. 38, 185-189 (I 978b). Wiebes, 1.T. - The tig wasp genus Do/ichoris Hili (Hymenoptera Chalcidoidea, Agaonidae). Proc. Kon. Ned. Akad. Wet. (C) 82, 181- 196 (1979a). Wiebes, 1.T. - Fig wasps from Gabon: new species of Agaon (Agaonidae) and Phagoblastus (To­ rymidae) (Hymenoptera ChaJcidoidea). Proc. Kon. Ned. Akad. Wet. (C) 82, 391-400 (l979b). Wiebes, l .T. - Tbe tig insects of la Réunion (Hymenoptera, Chalcidoidea). Ann. Soc. ent. France (N.S.) 17, 543-570 (1981). Wiebes, l .T. - The status of tig and tig-insect research in the eighties. Courrier de l'Ecotrop, Num. spéc., 55-63 (1984). Wiebes, l.T. - The association of tigs and tig-insects. Rev. Zool. afr. I()O: 63-71 (1986a). Wiebes, 1.T. - Agaonidae (Hymenoptera ChaJcidoidea) and Ficus (Moraceae): tig wasps and their tigs, 1. Proc. Kon. Ned. Akad. Wet. (C) 89, 335-355 (1986b). Wiebes, l .T. - Agaonidae (Hymenoptera Chalcidoidea) and Ficus (Moraceae): tig wasps and their tigs, II (A/fonsiella). Proc. Kon. Ned. Akad. Wet. (C) 91 , 429-436 (1988). Wiebes, l.T. - Agaonidae (Hymenoptera Chalcidoidea) and Ficus (Moraceae): tig wasps and their tigs, III (Elisabethiella). Proc. Kon. Ned. Akad. Wet. (C) 92, 117-136 (l989a). Wiebes, 1.T. - Agaonidae (Hymenoptera Chalcidoidea) and Ficus (Moraceae): tig wasps and their tigs, IV (African Ceratoso/en). Proc. Kon. Ned. Akad. Wet. (C) 92,251- 266 (1989b). Wiebes, 1.T. - Agaonidae (Hymenoptera ChaJcidoidea) and Ficus (Moraceae): tig wasps and their tigs, V (Agaon). Proc. Kon. Ned. Akad. Wet. (C) 92, 395-407 (1989c). Wiebes, 1.T. - African tigs and their pollinators - a brief overview. Mitt. Inst. AUg. Bot. Hamburg 23a: 425-426 (1990). Wiebes, l.T. & U.c. Abdurahiman - Additional notes on P/atyscapa Motschoulsky (Hymenoptera Chalcidoidea, Agaonidae). Proc. Kon. Ned. Akad. Wet. (C) 83, 195- 207 (1980). Wiebes, l.T. & S.G. Compton - Agaonidae (Hymenoptera Chalcidoidea) and Ficus (Moraceae): tig wasps and their tigs, VI (Africa concluded). Proc. Kon. Ned. Akad. Wet. 93: 203-222 (1990). Willdenow, C.L. -Détermination de quelques nouvelles espéces de tiguier, et observations générales sur Ie genre. Mém. Acad. Roy. Sci. BeUes-Lettres (1798),91- 104 (1798). Willdenow, c.L. - Species plantarurn 4. Berolini, 1157 pp. (Ficus: 1131 - 1153) (1805-1806). Williams, R.D. - The useful and omamental plants of Zanzibar and Pemba. Zanzibar, ix + 497 pp. (Ficus: 261 - 265) (1949). Winkier, H.l. - Neue Kameruner Phanerogamen aus verschiedenen Familien. Bot. lahrb. Syst., Pflanzengesch. & Pflanzengeogr. (Engier) 41, 275-286 (Ficus: 276) (1908).

285

Index

Accepted names are in roman, synonyms in italics; some principal page references are in bold print. For the lig names see p. 291.

Scientific names of animals (and Bacteria)

Acari 42 baliolum 38, 255, 257, 258, 262 acutatum (A.) 147, 258, 262 bambeyi (A.) 180 acutatus (C.) 79, 176, 209,211,213 bambeyi (G.) 184 ae/leni 211 barnbeyi (S.) 76, 184 affiicta 78, 178 barbarus 130, 174 africana (0.) lOl , 187, 188 barbatus 130, 174 africana (P.) 165, 181 bekiliensis 159, 265, 267, 268 africana (S.) 189 bergi (A.) 39, 109, 249, 250 africana (T.) 42 bergi (E.) 39, 113, 233, 234, 236, 237 africanus (C.) 98 , 189 bergi (P.) 95, 196, 223, 225, 226 afrum 65, 69, 70, 205, 206 binghami (A.) 39, 109, 249, 250 Agaon 34, 35, 36,37, 42, 139, 196, 200, 255, binghami (P.) 93 , 222, 223, 224 268, 270, 272, 273 bispinosa (C.) 159, 266,268, 269 Agaonidae 169, 170, 188 bispinosum ( A.) 268 Agaoninae 169, 171,172, 173,195 Blastophaga 34, 35, 36, 37, 62, 201 , 207, 218, Agaonini 97, 196 225, 231 , 235, 237, 239, 242 alata 130, 184, 185 Blastophagini 97, 196 Alfonsiella 27, 35, 37, 38, 102, 116, 196, 198, Blastophagus 201 200,247 blommersi 82, 210, 211, 216 Allotriozoon 34, 35, 37, 97, 200, 226, 229 bouceki 126, 174 allotriozoonoides 181 , 234, 242 brachycephaJa 179 anceps 94, 183, 184 breviceps (E.) 113, 237 Apocrypta 42, 76, 170,177, 179, 183 breviceps (S.) 165, 175 Apocryptini 169, 170, 172, 179 brevimucro 126, 130, 189 Apocryptophagus 171 , 175, 176, 178 brongersmai 39, 130, 198,240,245,248, 249, arabicus 25, 26, 28, 37, 38, 76, 77, 178, 210, 250, 251 , 252 211 , 212, 215 cadenati (G.) 183, 184 armata (C) 162, 163, 265, 269 cadenati (P.) 188 armatum (A.) 269 cadenati (S.) 183 armipes 165, 174, 175 caelebs 115, 184 arnottiana 225 callani 178 articulata 39, 134, 196, 198,232, 233, 234, 235, Camarothorax 172, 188, 189 238 camerunensis (E.) 176 atratus lOl , 173 camerunensis (C.) 26,27,28, 37, 38, 154,155, avicola 39, 115, 198, 244, 245, 246 156, 265, 266, 267, 269, 270 awekei 93 , 222, 223, 224, 225 capensis 26, 37, 38, 78, 196, 209, 210, 211 , 213, baijnathi 26, 39, 124, 231, 233, 234, 238, 249 214

287 carayoni 216 Eupristina 218 caricae 181 Eurytoma 172, 190 Cecidomyidae 41 Eurytomidae 169, 171 , 190 Ceratosolen 34,35,37, 41 , 74, 176, 179, 198, excavata 39, 110, 244 200, 207, 209 explorator 80, 176 cercaria 76, 180 fascia turn 143, 175, 257, 258, 261, 262 Cercopithecinae 31 Ficiomyia 41 Cha/cis 176 Ficomila 172, 190 cicatriferens 150, 257, 258, 259 fictitia 11 5, 185 cnephaea 126, 181 ficus 76, 130, 191 coccothraustes 106, 184 ficusgallae 130, 190 coecus 80,208,210,211 , 215 fima 41 collaris 188 fimbriata 38, 39, 122, 132, 249,250 C%trechnus 188 finitimorum 98, 181 comparabilis 94, 184 flabelIata (D.) 38, 85, 87, 196, 198, 200, 217 comptoni 38, 39, 111 , 234, 240, 246 flabellatus (C.) 26, 37, 38, 78, 210, 211, 213, Comptoniella 170, 171 , 172, 185, 186 214 comutus 126, 130, 183 flaviclava 130, 191 CourtelIa 34,35, 36,37,42, 153,200,201, 263 flavipes 130, 191 cowani 72, 196, 203, 204, 205, 206, 207 Formicidae 31 , 42 crassipes 178 froggatti 24 Crossogaster 173 fusciceps 26,38,39,111,196,198,241,243, crypta 77, 180 244, 245, 246 Curculionidae 41 curtivena 130, 190 gabonense (A.) 152,257,258, 260 cyclostigma 78, 178 gabonensis (C.) 26, 27, 28, 37, 38, 154, 265, Cynips 178 266, 269 galili (C) 26,37,38, 76,77,209, 210,211 , 215 dagatiguyi ( C) 214 galili (0.) 189, 190 dagatiguyi (S.) 184 gambiense (D.) 191 dakarensis 230 gambiensis (F.) 76, 130, 191 Decatoma 191 gestroi 26, 38, 42, 65, 69, 70, 205, 206 dentifer 27 gibbus 68, 73, 183 depressa 178 gigantea 178 desertorurn 92, 222, 223, 224 gigas 76, 176 Diptera 41 glumosae 39, 108, 234, 240 Dolichoris 34, 35, 36, 37, 85, 217 gnapha/ocarpae (A.) 176 Drosophilidae 41 gnaphalocarpae (0.) 188 Dynatogmus 171, 176 Goniogaster 184 dyscritus 242 gracile 76, 177 Elisabethiella 35, 36, 37, 38, 102, 116, 133, Grandimyia 183 196, 198,200, 231 guineensis 78, 180 emmerezi 215 gustatus 31 enriquesi 39, 124, 234, 239 hamifera (C.) 165, 181 ,265, 266, 267, 268 epicarioides 188 hamiferurn ( A.) 263 Epichrysomallinae 169, 170, 171 , 188 hemirnucro 126, 189, 190 equicollis 130, 190 heterandromorphum 26, 101, 173,228,229, erythraea 181 230 estherae 26 Heteroptera 31 etiennei (K.) 68, 205, 207 hewitti 26 l"tiennei (P.) 94, 196, 222, 223, 224, 225 hiUa (S.) 189 Eukoebelea 171,175,176, 177 hilli (E.) 232, 234, 235, 236 Euprista 218

288 hilli (K.) 38, 69, 204, 405, 206 narnorakensis 38, 76, 209, 210, 211, 212 hirtus 130, 184 naso 76, 130, 191 hladikae 38, 158, 265, 267,272 natalensis 38, 249, 250, 252 Homoptera 31 Nematoda 41 Hypsignathus 31 Neosycophila 176 niger 183 Idames 42, 171 , 177 Nigeriella 35, 36, 37, 38, 102, 133, 196, 198, imbecillus 179,180 200,243 imerinensis 190 nigeriense 99, 228, 229, 230 insularis 178 nipponica 26 Ixonotus 31 nigra 183 josephi 39, 138, 198, 200, 253, 254 obtusum 145, 257, 258, 262 kestraneura 130, 191 Odontofroggatia 172, 189, 190 kiellandi 38, 139, 144, 147, 256, 258,259 odorans 130, 173 Koebelea 176 Oecophylla 42 KIadibia 34, 35,36, 37, 65 , 198, 200, 271 , 203 okinavensis 218 orientalis 126, 190 Lepidoptera 41 Ormyridae 169, 171 , 191 letouzeyi 39, 137, 243, 244, 245, 247 Ormyrus 171 , 191, 192 levis 155, 175 Otitesella 171,172,185, 187, 188 Iiodontus 122, 174 Otitesellinae 42, 169, 170, 171,172, 185 Liporrhopalwn 205 Lissocephala 41 palmata 181 lomaensis 98, 184 paradoxa (Ap.) 176 longicephalus 175 paradoxum (Ag.) 149,256,257,258, 261 , 262, longicornis (C) 211 267, 268 longicornis (P.) 181 Paragaon 35, 36, 37, 38, 133, 200, 252 longirnucro (Ca.) 130, 190 Parakoebelea 175, 176 longirnucro (Ce.) 82, 210, 211, 216 Parapristina 36, 198, 218 longinoda 42 Parasitodiplogaster 42 longiscapa (A.) 38,39, 122, 130, 240, 245, 249, parca 130, 181 250,252 pectinata 39, 103, 117, 233, 234, 237 longiscapa (E.) 39, 135, 233, 234, 238 penicula (c.) 38, 158, 215, 265, 266,267, 271 , longitarsus 76, 179, 180 272 lucens 185 peniculwn (A.) 271 perplexa (A.) 80, 110 malawi 161 , 265, 267, 269 perplexum (P.) 254 medleri 164,265, 266, 267, 273 Phagoblastus 173, 174 megacephalus 38, 80, 178,209,211 ,214, 216 Philocaenus 174 megalopon 142, 196, 198, 256, 258 Philosycus 171 , 172, 185,187,188 Megaselia 41 Philotrypesini 169, 170, 172, 180 michaloudi (A.) 38, 39, 120, 249, 250, 251 Philotrypesis 42, 170,172, 179, 180, 181 , 185 michaloudi (C.) 165, 264, 265, 267, 272 Phoridae 41 minima 180 pilosa 42 modesta (C.) 165, 268 Platyscapa 34, 35, 36, 37, 90, 94, 98, 201, 220 modesta (8.) 130, 191 platyscapa 39, 118,233,234, 236 modeslwn (A.) 268 Pleistodontes 36, 268 monslruosa (S.) 188 polyspina 82, 180 monstruosus (H.) 31 prodigiosum 98, 198,227,228, 229, 230 monstruosus (p.) 165, 188 psenes 26, 29, 38,41, 63, 166, 181 , 196, 198, montis 98, 183 200, 201 , 202,203 multum 150, 257, 260 Pteromalidae 169, 180, 188 myersi 178, 211 punctum 130, 191

289 quadraticeps 26,41, 167, 195,221 , 222,223, sycomori (S.) 76, 177, 178 225,226 Sycophaga 42, 171 , 176,177, 178 Sycophaginae 169, 175, 180 reflexa 39, 126, 198, 234,241 Sycophagini 169, 171 , 172, 175 regalis 179, 180 Sycophila 172, 189, 191 remus 126, 130, 184 sycophilon 42 rennellensis 42 Sycoryctes 170, 173, 183, 184 robusta (A.) 80, 178, 180 Sycoryctinae 169, 170, 172, 180 robustus (D.) 176 Sycoryctini 169, 170,172, 173, 180, 182 royi 98 , 188 Sycotetra 172, 190 saundersi 73 , 204, 205, 207 Sycoscapter 170, 173, 182, 183, 184 scobinifera (C.) 37, 38, 155, 267,270 Sycoscapteridea 170, 173, 183, 184 scobiniferum (A.) 270 sylviae 163,264, 265, 266, 270 sebertianus 184 taïense 38, 140, 255, 257, 258, 259 selenitica 101, 182 tananarivensis 183 senegalensis 183 Tarsonemella 42 Seres 42, 174 Tarsonemidae 42 serrata 93 , 188 tenebrosa 80, 178 Serratia 42 Tetrapus 36, 40 serricornis 130, 190 thaumastocnemus 145, 175 sesquianellata 124, 188 tibialis 115, 183 sessilis 130, 191 tonduzi 26 setoptera 82, 180 Torymidae 169, 180, 188 seyche/lense 230 lridentatum 261 silvestrianus 26, 38, 78, 178, 210, 211 , 214 trifornlls 106, 173 silvestrii (C.) 101 , 173 truncatus 106, 184 silvestrii (S.) 178, 180 tsamvi 130, 187, 188 sjoestedti 184 socotrensis 38, 39,106,122, 198, 233, 234,242 uluzi 124, 188 soraria 24, 90, 196, 200, 222, 223, 226 vaidi 202 spatulaturn 149, 198, 257, 258, 261 valentinae 76, 178 Staphylinidae 41 vannoorti 92, 285, 286 Stathmopoda 41 verticillata 166, 195, 196, 198, 200, 218, 219 stuckenbergi 38, 39, 42, 130, 231 , 233, 234, vicina 176, 178 239, 252 viduata 178 stupefactus 81 , 211 , 212 vi/liersi 262 subconicus 130, 192 Syceurytoma 172, 191 wanei 230 Sycobia 189 wardi 157, 198, 265, 271 Sycocrypla 207, 215 watshami 130, 192 Sycoecinae 42, 169, 171 , 173 Watshamiella 170, 173, 183, 184,185 Sycoecus 42, 175 Zaprionus 41 sycomori (E.) 76, 177

290 Scientific tig names (c.q., Moraceae)

ADENOSPERMA Corner 34 assimilis Baker 54, 87 asynvnetrica Huteh. 165 AMERICANA Miq. 16,34, 35 avi-avi Blume 114 CAULOCARPAE (Mildbr. & Burret) Berg 14, ayersii Baker 80 31 , 34, 35, 36, 37, 42, 152, 153, 181 , 188, 265 barbata Warb. 106 C HLAMYDODORAE (Mildbr. & Burret) Berg barombiensis Warb. 159 34, 39, 116, 133, 181, 232, 249 baronii Baker 99 barteri Sprague 59, 133, 147, 148, 262 C ARICEAE Corner 62 basarensis Mildbr. & Burret 127 CONOSYCEA (Miq.) Corner 32, 34,35,36, 47 , benghalensis L. 13, 166 48, 54, 94, 221 beniensis De Wild. 77 benjamina L. 166 CRASSICOSTAE (Mildbr. & Burret) Berg 34, bequaertii De Wild. 127 37, 39, 102, 112, 132, 133, 232, 244, 254 bivalvata Perrier 61 , 155 CYATHISTIPULAE (Mildbr. & Burret) Berg bizanae Huteh. & Burtt Davy 49, 61, 159 14, 31,34,35,36, 37,42, 133, 138,256 bojeri Baker 53 , 71 bongoensis Warb. 128 FICUS L. 9, 10, 11, 12, 13, 14, 16, 18,22,30, bongouan( ouJensis A. Chev. 85 31, 33, 34, 35, 36, 37,40, 62, 181, 201 botryoides Baker 54, 82, 180,216 abscondita C.C. Berg 55, 142 brachyclada Baker 52, 53 , 72, 207 abutilifolia (Miq.) Miq. 13, 38, 39,50, 57,58, brachylepis Hiern 162 102, 110, 111 ,241 , 246 brachypoda Huteh. 165 acuta De Wild. 157 brassii Sabine 77 acutifoha Huteh. 68 brevicula Hiern 71 adolfi-friderici Mildbr. 14, 58, 60, 112, 133, brevipedice/lata De Wild. 122 137 - var. cuneata De Wild. 128 africana Kunth & Bouché 165 broussonetiifoha Baker 66 aganophila Huteh. 137 bubu Warb. 61, 164,273 aggregata Vahl 126 buchneri Warb. 165 akaie De Wild. 99 budduensis Huteh. 112 albidula Baker 81 buettneri Warb. 154 aldabrensis Baker 126 - var. globicarpa Mildbr. & Burret 154 altissirna Blume 166 bllntingii Huteh. 162 amadiensis De Wild. 14, 49, 55, 58, 118 bllrkei (Miq.) Miq. 128 ampana c.c. Berg 54,88 burretiana Mildbr. & Huteh. 39, 59, 133, 134, annabonensis Mildbr. & Huteh. 127 238 anomani Huteh. 118 burtt-davyi Huteh. 17, 25,26,39, 49, 60, 123, antandronarum (Perrier) c.c. Berg 62, 124 181 , 182, 188, 239 - subsp. antandronarum 125 bussei Mildbr. & Burret 49, 58, 60, 103 - subsp. bernardii c.c. Berg 125 - var. longepetiolata Mildbr. 104 antithetophylla Miq. 69 butagensis De Wild. 128 apodocephala Baker 99 buxifolia De Wild. 120 arcuato-nervata Huteh. 141 cabrae Warb. 100 ardisioides Warb. 56, 59, 60, 141 caffra (Miq.) Miq. 90 - subsp. ardisioides 141 - var. longipes Warb. 90 - subsp. camptoneura (Mildbr.) c.c. - var. natalensis Warb. 90 Berg 141 , 258 - var. pubicarpa Warb. 90 arimensis Britton 99 - var. sambesiaca Warb. 90 artocarpoides Warb. 38, 61, 158, 27 1, 272 callabatensis Warb. 104 asperifolia Miq. 21,23,26,38,50, 52, 68,206 ca/lescens Hiern 143

291 calotropis Lebrun & Toussaint 118 - var. marlo/hii Warb. 92 ca1yptrata Vahl 13, 14, 17, 39,58, 116,238 - var. Iris/is (Kunth & Bouché) Warb. 92 camptoneura Mildbr. 140, 141 coriacea De Wild. 146 - var. angustifolia Mildbr. & Burret 141 corylifolia De Wild. & T. Durand 76 camptoneuroides Huteh. 139 - var. glabrescens Warb. 76 capensis Thunb. 77 costaricana (Liebm.) Miq. 26 - var. beniensis (De Wild.) Lebrun 77 crassicosta Warb. 55, 60, 152 - var. guineensis (Miq.) Miq. 77 crassipedicel/a/a Warb. 128 - var. ituriensis (De Wild.) Lebrun 77 - forma angus/ifolia De Wild. 128 - var. mal/otocarpa (Warb.) Mildbr. & - forma boonei De Wild. 128 Burret 77 - forma cuneata De Wild. 128 - -forma pubifolia Mildbr. & Burret 77 craterostoma Mi1dbr. & Burret 14, 38, 39, 49, - var. ostiolata (De Wild.) Lebrun 77 56, 58, 59, 118, 119, 120, 122,251 - var. pubescens De Wild. & T. Durand 77 cyathistipula Warb. 17, 31,49,50,56, 139, - var. trichoneura Warb. 77 143, 144, 147, 175 capreifolia Delile 38, 50, 52, 69, 70, 71 , 206 - subsp. cyathistipula 139, 143,261 - var. ovatifolia Huteh. 69 - subsp. pringsheimiana (Braun & K . Schurn.) caricaL. 17,22, 26, 32,38,42,62,63,166, 181, e.e. Berg 38, 139, 143, 147,259 202,203 cyathistipuloides De Wild. 13, 16, 60, 145, caricoides Roxb. 63 146,262 chanas Forssk. 74 cyphocarpa Mildbr. 128 changuensis Mildbr. & BUITet 103 dahro Delile 104 - var. somalensis Pamp. 103 damarensis Eng!. 74 chirindensis e.C. Berg 49, 61, 161, 269 dar-es-salaamii Huteh. 108 cWamydocarpa Mildbr. & Burret 57, 58, 98, dawei Huteh. 98 230 dekdekena (Miq.) A. Rich. 128 - subsp. chlamydocarpa 50, 99 - var. angustifolia A. Peter 128 - subsp. femandesiana (Huteh.) C.C. Berg 99 - var. pubiceps Mildbr. & Burret 128 - subsp. latifolia (Huteh.) e.e. Berg 99 delagoensis Sim 162 chlamydodora Warb. 128 demeusei Warb. 158 chrysocerasus Warb. 122 densifo1ia Miq. 50, 55, 56, 94, 184, 185, 224 cinerea Cordem. 114 densistipulata De Wild. 17, 38, 55, 139, 144, c/aoxyloides Baker 73 146, 147, 259 c/arellcensis Mildbr. & Burret 99 depauperata Sim 121 c/ethrophyl/a Hiem 77 dewevrei De Wild. & T. Durand 154 cnestrophyl/a Warb. 68 dewevreoides De Wild. 154 cocculifolia Baker 74 dicranostyla Mildbr. 32, 38, 54, 85, 86, 218 - subsp. sakalavarum (Baker) Perrier 74, 180 - var. nitida Huteh. 85 - var. sakalavarum (Baker) Perrier 74 din/eri Warb. 128 cognata N.E. Brown 128 discifera Warb. 111 colpophyl/a Warb. 68 djalonensis A. Chev. 116 comorensis Warb. 74 dryepondtiana De Wild. 61,163,271 congensis Eng!. 112 durandiana Warb. 106 - var. mol/is Huteh. 112 durbanii Warb. 121 conraui Warb. 38, 55, 56, 139, 141, 144, 147, dusenii Warb. 128 259 consimilis Baker 114 ealaensis De Wild. 154 cordata Thunb. 54, 90 ebolowensis Mildbr. & Huteh. 146 - subsp. cordata 49, 92, 187, 224, 225 e1astica Roxb. 35, 166 - subsp. lecardü (Warb.) e.C. Berg 49, 92 e1asticoides De Wild. 13, 39, 59, 60, 133, 134, - subsp. salicifolia (Vahl) C.e. Berg 50, 90, 92, 235 188,224 elegans (Miq.) Miq. 158 - var. fieckii Warb. 92 epiphytica De Wild. 152

292 erecta Thunb. 26 - var. glabrior Miq. 128 erici-rosenii R.E. Fries 128 holstii Warb. 100 eriobotryoides Kunth & Bouché 98 homblei De Wild. 108 - var. cail/ei Mildbr. & BUITel 98 howardii Sim 108 - var. latifolia Huteh. 99 humbertü c.c. Berg 13, 54, 95, 96 - var. monbuttensis (Warb.) Lebrun 98 ilicina (Sonder) Miq. 39, 50, 60, 116, 124,239 eriocarpa Warb. 128 incognita De Wild. 100 erubescens Warb. 77 indica Forssk. 92 eucalyptoides Battand. & Trabut 92 ingens (Miq.) Miq. 24,50,54,90,91,93,226 exasperata Vahl 38,48,52,65,69, 181,206 - var. tomentosa Huteh. 90 excentrica Warb. 122 ingentoides Huteh. 90 fasciculata Warb. 103 inkansuensis Warb. 158 fasciculiflora Huteh. 154 inopinata Mildbr. 85 faulkneriana C.c. Berg 56, 60, 116, 123 integrifolia Sim 74 fazokelensis (Miq.) Miq. 106 intermedia DeliIe III fernandesiana Huteh. 99 irumuensis De Wild. 68 ferruginea Desf. 178, 180 iteophylla Miq. 128 tischeri Mildbr. & Burret 39, 59, 116, 117, 236 ituriensis De Wild. 77 tistulosa Reinw. ex Blume 26 jansii Boutique 61 , 102, 112, 133 flavovenia Warb. 112 johnstonii Stapf 165 fleuryi A. Chev. 78 forsskalii Vahl 63 kaba De Wild. 100 furcata Warb. 122 kagerensis Lebrun & Toussaint 128 - var. angustifolia De Wild. 118 kamerunensis Mildbr. & BUITet 38,39,56,57, 59, 116, 131, 132, 251 galactophora Tenore 98 karthalensis C.C. Berg 50, 53, 54, 83, 84 galpinii Warb. 128 katagumica Huteh. 90 gil/etii Warb. 154 kawuri Huteh. 90 glumosa Delile 39, 57, 59, 102, 106, 107, 108, kerstingii Huteh. III 109,240 kiloneura Homby 117 - var. glaberrima Martelli 106 kimuenzensis Warb. 157 - var. intermedia Martelli 106 kirkii Huteh. 144 - var. lanuginosa Martelli 106 kisantuensis Warb. 158 glumosoides Huteh. 106 kitaba De Wild. 106 gnaplzalocarpa (Miq.) A. Rich. 74, 175, 176, kitabalu Huteh. 118 178, 180, 184, 188 kondeensis Warb. 77 goetzei Warb. 128 kotschyana (Miq.) Miq. 102 goliath A. Chev. 104 kyimbilensis Mildbr. 164 golungensis Huteh. 104 gombariensis De Wild. 106 langenburgii Warb. 162 gongoensis De Wild. 77 lanigera Warb. 100 gossweileri Huteh. 162 laieralis Warb. 102 grevei Bail!. 62, 113 laterifiora Vahl 50, 53, 66, 67, 207 guatteriifolia Baker 87 laurenlii Warb. 148 guerichiana Warb. 124 laurus Bail!. 83 guineensis (Miq.) Stapf 77 lecardii Warb. 92 ledermannii Huteh. III hararensis Warb. 104 leonensis Huteh. 58, 133, 138 hemsleyana Stand!. 26 leprieurii Miq. 122 hippopotami Gerstner 112 - var. intermedia Huteh. 122 hispida LinnJ. 42 - var. sessilis Huteh. 122 hispidioides S. Moore 27 Iichtensteinii Link 77 hochstetteri (Miq.) A. Rich. 128

293 lingua De Wild. & T. Durand 13, 15, 38, 39, muelleriana c.c. Berg 50, 58, 110 49, 56, 59, 60, 120 munsae Warb. 77 - subsp. depauperata (Sim) C.c. Berg 121 mu/antifolia Huteh. 118, 120 - subsp. lingua 120, 251 namalalensis Huteh. 146 longepeduncula/a De Wild. 155 natalensis Hochst. 38, 39, 49, 59, 120, 121, longipes Baker 73 240, 242, 243, 252 louisii Boutique & J. Léonard 39, 58,60, 133, - subsp. leprieurii (Miq.) c.c. Berg 38,49, 50, 138, 255 122, 132, 174, 251 lucanda Ficalho 155 - subsp. natalensis 38, 121, 122, 252 lucens Cordem. 94 - var. latifolia Warb. 121 /ujae De Wild. 142 - var. minor (Sonder) Warb. 123 lutea Vahl 26,31,32, 41,50, 57,58, 62, 97, 99, - var. peduncu/ala Sim 121 100, 101 , 173, 182, 188, 230 - var. puberula Warb. 123 lu/eola De Wild. 118 nautarum Baker 100 /ynesii Lebrun 85 ndola Mildbr. 118 Iyrata Warb. 49, 55, 149, 261 nekbudu Warb. 100 mabifolia Warb. 128 neriifolia A. Rich. 128 macrophylla Desf. ex Vent. 24, 26, 28 neumannii Kunth & Bouché 100 macrosperma Mildbr. & Burret 162 neurocarpa Lebrun & Toussaint 128 macu/osa Huteh. 154 niamniamensis Warb. 159 madagascariensis c.c. Berg 55, 93 nigropunctata Mildbr. & Burret 39, 58, 109, magenjensis Sim 90 250 mall%carpa Warb. 77 nyanzensis Huteh. 143 - var. Irichoneura (Warb.) A. Peter 77 ob/usala Link 80 mallolOides Mildbr. & Huteh. 116 oc/omelifolia Warb. 165 mammigera R.E. Fries 128 oreodryadum Mildbr. 50, 59, 133, 135 mammosa Lebrun 121 oresbia c.c. Berg 50, 59, 152 mangenotii A. Chev. 162 osliola/a De Wild. 77 mangiferoides Huteh. 135 - var. brevipedunculata De Wild. 77 marmorata Baker 62, 115 ottoniifolia (Miq.) Miq. 13, 19, 22,25,26,27, mauritiana Lam. 50, 53,54,80, 176, 177, 180, 28,37,38,49, 61, 153, 156 215 - subsp. lucanda (Ficalho) c.c. Berg 37,154, medullaris Warb. 128 155, 175, 270 megalodisca Warb. 163 - subsp. macrosyce c.c. Berg 37, 49, 154, 155, megaphylla Warb. 165 156, 270 - var. glabra Warb. 165 - subsp. multinervia C.C. Berg 37, 49, 154 megapoda Baker 159 - subsp. ottoniifolia 37, 154, 155, 156, 269, melleri Baker 126 270 menabeensis Perrier 54, 95, 226 - subsp. ulugurensis (Mildbr. & Burret) c.c. michelsonii Boutique & J. Léonard 144 Berg 37, 61 , 154, 155, 156, 161 , 270 microcarpa Linn.f. 22, 166, 190, 195, 220 ovata Vahl 14, 57,153, 165, 174,175,181 , 188, microcarpa Vahl 128 268 micropison Mildbr. 120 - var. oClOme/ifolia (Warb.) Mildbr. & mildbraedii Huteh. 148 Burret 165 mittuensis Warb. 111 ovato-cordata De Wild. 90 modes/a F. White 155 oxys/ipula Baker 82 monbullensis Warb. 98 monlana Warb. 106 pachyclada Baker (1887) 52, 53 , 63 , 65 morifolia Forssk. 63 - subsp. arborea (Perrier) c.c. Berg 66 morifolia Lam. 66 - subsp. pachyclada 66 mucuso Ficalho 13, 32, 37, 38, 53, 76, 180, pachyc/ada Baker (1890) 100 212, 215 pachyneura c.c. Berg 49, 56, 140

294 pachypleura Warb. 164 pulvinata Warb. 157 paimata Forssk. 10, 19,31,32,38,48,49, 62, pulvinifera Baker 80 63, 64, 181, 202 pumila L. 32, 166 - var. morifolia (Forssk.) Warb. 63 punctata Lam. 126 - var. petitiana (A. Rieh.) Fiori 63 punctifera Warb. 65 - var. pseudocarica (Miq.) Warb. 63 pygmaea Hiem 50, 52, 71 - var. somalensis Warb. 63 pynaertii De Wild. 154 - var. slipitata Warb. 63 pyrifolia Lam. 114 paludicola Warb. 68 - var. ambongensis Perrier 95 paluslris Sim 69 - var. anlandronarum Perrier 124 panifica Delile 77 - var. irnerinensis Perrier 126 paolii Pamp. 69 - var. meIleri (Baker) Perrier 126 pendula Hiem 68 - var. meridionalis PeITier 95 persicifolia Warb. 128 - var. occiden talis Perrier 126 - var. anguslifolia Warb. 128 - var. pedunculata Perrier 125 - var. glabripes Warb. 128 - var. Isaralananensis Perrier 126 pelersii Warb. 128 quibeba Ficalho 100 petitiana A. Rieh. 63 phanerophlebia Baker 72 racemosa L. 26, 48, 74 phil/ipsii Burtt Davy & Huteh. 128 recurvata De Wild. 49, 58, 60, 102, 103 picla Sim III rederi Huteh. 143 pilosuIa De Wild. 118 reflexa Thunb. 39,62, 125, 174, 181 , 188, 189, plateiocarpa Warb. 77 190, 242 platyphylla Delile 13,39,49, 58, 60, 102, 103, - subsp. aidabrensis (Baker) e.C. Berg 126 174 - subsp. reflex a 126 podophylla Baker 159 - subsp. sechellensis (Baker) e.e. Berg 51, poli ta Vahl 50, 61 , 62, 153, 158, 164 126 - subsp. brevipeduneulata e.e. Berg 50, 159 rehmannii Warb. 106 - subsp. polita 159, 160, 268 - var. ovátifolia Warb. 106 - var. persicicarpa Huteh. 163 .c. var. vil/osa Warb. 106 politoria Lam. 53, 73, 207 religiosa F orssk. 1iI polybraclea De Wild. & T. Durand 154 religiosa L. 13, 26, 28, 167, 195, 225 polyphlebia Baker 54, 81 , 180, 216 rhodesiaca Mildbr. & BUITet 128, 178 pondoensis Warb. 90 rhynchocarpa Mildbr. & Huteh. 143 populifolia Vahl 49, 59, 111 riparia (Miq.) A. Rieh. 77 - var. major Warb. III rivae Warb. 104 - var. somalensis Warb. 1i1 rokko Warb. & Schweinf. 128 - var. taÎ/ensis Warb. 1i1 rubicunda (Miq.) Miq. 106 praeruplorum Hiem 93 rubiginosa Desf. ex Vent. 178 praticola Mildbr. & Huteh. 139 rubra Vahl 39, 62, 113, 114, 184, 185, 247 preloriae Burtt Davy 92 - var. sechellensis Baker 126 preussii Warb. 56, 142 rllbropunctata De Wild. 11 8 principes Kunth & Bouehé 98 rubroreceptaculata De Wild. 98 pringsheimiana Braun & K. Schum. 143 rudens Huteh. 157 pseudocarica Miq. 63 rllficeps Warb. 128 - var. tomenlOsa A. Rieh. 63 rukwaensis Warb. 106 pseudo-elastica Hiem 165 rupicola Lebrun & Toussaint 128 pseudomangifera Huteh. 39,61, 133, 135, 136, rupium Dinter 92 247 ruspolii Warb. 128 pseudosycomorus Deene. 63 ruwenzoriensis De Wild. 11 8 pseudo-vogelii A. Chev. 100 sagittata Vahl 167 psilopoga Fiealho 128 sagittifolia Mildbr. & Burret 49, 55, 149, 260 pubicosla De Wild. & T. Durand 128

295 saka/avarum Baker 38, 74, 76, 212 spragueana Mildbr. & Burret 128 salieifo/ia Vahl 92, 93 stel/u/ala Warb. 78 - var. austra/is Warb. 92 - var. g/abreseens Warb. 77 - var. latifolia Huteh. 92 stenoclada Baker 82 sansibarica Warb. 15,49, 61, 153, 162, 163, sterculioides Warb. 155 270 slipitata Lebrun 146 - subsp. macrosperma (Mildbr. & Burret) c.c. Slipu/ifera Huteh. 139 Berg 14,49, 162 storlhophylla Warb. 68 - - subsp. sansibarica 162 - var. cuneala De Wild. 68 sapinii De Wild. 165 stuhlmannii Warb. 39, 57, 108, 109, 250 sapotoides Baker 80 - var. g/abrifolia Warb. 90 saussureana DC. 31,55,57, 58, 98, lOl , 181, subacuminala (De Wild.) Lebrun 121 184, 188, 189, 229 subca/earala Warb. 100 seabra Sim 74 - var. vestito-braeteala (Warb.) Mildbr. & seabra Willd. 65 Burret lOl scassellatii Pamp. 49, 59,60, 139, 144 subcostata De Wild. 56,59, 141 , 147,262 - subsp. scasseUatii 144 subsagittifolia C.c. Berg 55, ISO, 151, 260 - subsp. thikaensis c.c. Berg 144 sur Forssk. 20, 21, 26, 28, 32, 37, 38, 42, 49, sehefJleri Mildbr. & Burret 155 50,53,77, 176, 177, 178, 180,213, 214 sehimperi (Miq.) A. Rich. 128 - var. erythreae Fiori 77 - var. hoehstetteri (Miq.) Mildbr. & sycomorus L. 13, 15, 22,23,26,28,29,32,37, Burret 128 38,48,50,53, 54, 74, 75,76, 77,176,177, 178, sehimperiana A. Rich. 90 180, 184, 191 , 211 , 212, 215 sehinziana Warb. 128 - subsp. gnapha/oearpa (Miq.) c.c. Ber~ 49, sehweinfurthii Miq. 79 76 sciarophyl/a Warb. 85 - var. a/nea Hiern 77 seo/pophora Warb. 68 - var. po/ybolrya Hiern 77 scott-elliotii Mildbr. & Burret 49, 60, 145 - var. prodigiosa Hiern 77 scutata Lebrun 121 syringifolia Warb. 159 seehellarum Surnmerhayes 126 le/oukat Battand. & Trabut 92 senega/ensis Miq. 100 lerebrala Willd. 126 serelii Lebrun & Boutique 78 terragena Bory 80 serrata Forssk. 65 tesselata Warb. 38, 56, 139, 259, 263 sessilis De Wild. 139 tettensis Huteh. 13, 39,50,57, 102, 109, lil, sidifolia Hiern 76 246 silieea Sim 65 Ihonningiana (Miq.) Miq. 77 smutsii Verdoorn 109 thonningii BI. 28,32,38, 39,42,49,50, 56,57, socotrana Balfour 104 58, 59, 60, 116, 120, 122, 125, 127, 129, 132, so/danella Warb. IiI 173,174, 178, 181 , 184, 185, 188, 190,191, soma/ensis (pamp.) Chiov. 103 192, 240, 251, 252 sonderi Miq. 106 - var. heterophylla A. Peter 128 soraria Cordem. 80 - arcocarpa form 129 soroeeoides Baker 73 - burkei form 130, 131 - var. arborea Perrier 66 - dekdekena form 131 - var. ambongensis Perrier 73 - hochstetteri form 131 - var. braehyclada (Baker) Perrier.72 - iteophylla form 130 - var. ea/cico/a Perrier 73 - mammigera form 129 - var. /ongipes (Baker) Perrier 73 - neurocarpa form 130 - var. macroph/ebia Perrier 72 - persicifolia form 131 , 132 - var. mananarensis Perrier 73 - petersii form 131 - var. onivensis Perrier 73 - rokko form 129 speclabilis Kunth & Bouché 165 tiliifolia Baker 54, 80, 212 sphaerophyl/a Baker 80

296 - var. ambrensis Perrier 80 vo/kensii Warb. 121 - var. sphaerophy/la (Baker) Perrier 80 wakefieldii Huteh. 38, 39, 57, 106, 242 torrentium Perrier 53, 81, 180 warburgii H.l. Winkler 68 trachyphy/la (Miq.) Miq. 74 we/witschii Warb. 92 tremuIa Warb. 49, 61, 156 - var. beroensis Hiern 92 - subsp. acuta (De Wild.) e.C. Berg 50, 157 wildemaniana De Wild. & T. Durand 13,55, - subsp. kimuenzensis (Warb.) e.e. Berg 157 59, ISO, 260 - subsp. tremuIa 156, 157, 272 wink/eri Mildbr. & BUfret 139, 263 triangu/aris Warb. 122 trichoclada Baker 54, 83 xiphocuspis Baker 73 trichoph/ebia Baker 80 xiphophora Warb. 68 trichopoda Baker 13, 39, 50, 58, 60, 62, 102, zambesiaca Huteh. 103 112, 237 zenkeri Mi1dbr. & BUfret 85 trichosphaera Baker 10 I zobiaensis De Wild. 98 Irislis Kunth & Bouché 92 zuva/ensis Sim 112 Iropophylon Lebrun & Toussaint 128 tubercu/osa Hiern 163 GALOGLYCHIA (Gasp.) End!. 16, 18, 19,29, - var. elliptica Hiern 165 31,32, 34, 35,36,37,39,42,47,48, 50,51, ugandensis Huteh. 162 97, 181 , 187, 189,228 u/ugurensis Mildbr. & Burret 155 saussureana (De.) Gasp. 98 umangiensis De Wild. 154 lenoreana Gasp. 98 - var. /aurenlii De Wild. 154 umbelIata Vahl 61 , 163, 164,273 KALOSYCE (Miq.) Corner 34, 36 umbrosa Sim 159 LEUCOGYNE Corner 34, 35 urceo/aris Hiern 38, 68, 69, 206 - var. bumbana Hiern 68 MALVANTHERA Corner 34, 35, 36, 39 usambarensis Warb. 49, 61 , 133, 137 NEOMORPHE King 34, 35, 36, 49, 209 utilis Sim lOl OREOSYCEA (Miq.) Corner 13, 18,30,32,34, vallis-choudae DeWe 38, 53, 79, 177, 178, 180, 35,36, 37,48,52,84,217 214,217 - var. pubeseens A. Peter 79 PHARMACOSYCEA Miq. 12,13, 14,15, 16, 19, variabilis De Wild. 31, 34, 35, 36, 39, 84, 217 - forma obtusifolia De Wild. 121 PLATYPHYLLAE (Mi1dbr. & Burret) Berg 34, - forma subanuninala De Wild. 121 37, 39, 50, 101, 102, 112, 133, 187, 189,232, variifolia Warb. 13, 32, 38, 54, 85, 87, 218 244, 249 vasta Forssk. 38,39, 49, 57,104, 105, 173, 184, 242 RHIZOCLADUS End!. 34, 36 - var. g/abrescens Huteh. 104 SINOSYCIDIUM Corner 34, 35 - var. ve/utina Fiori 104 verrucocarpa Warb. lOl SPARATTOSYCE Bureau 9 verruculosa Warb. 13, 50, 54, 88, 93, 224 STILPNOPHYLLUM End!. 34 - var. slipitata Mildbr. & Burret 93 vesliw-bracleala Warb. lOl SYCIDIUM Miq. 10, 12, 13, 14, 15, 16, 17, 18, villosipes Warb. 77 19, 25,31 , 32, 34, 35,36,37,40,47,48, 52, virens Ait. 167 63,65,87, 181 , 204 viridimacu/ata De Wild. 154 SYCOCARPUS Miq. 14, 34, 35, 36, 49 voge1iana (Miq.) Miq. 14,50, 53,78, 176, 178, 179, 213 SYCOMORUS (Gasp.) Mildbr. & Burret 12, - var. /atifolia Huteh. 78 13, 14, 15, 16, 17, 18, 19,30,31, 32,34,35, vogelii (Miq.) Miq. lOl 36, 37, 42, 48,49, 52, 53,73, 74, 179, 209 - var. pubicarpa Mildbr. & Burret lOl anliquorum Gasp. 74 vohsenii Warb. 165

297 capensis (Thunb.) Miq. 77 galac/ophorurn (Tenore) Miq. 98 - forma guineensis (Miq.) Miq. 77 glurnosum (Delile) Miq. 106 guineensis Miq. 77 hochs/etleri Miq. 128 gnaphalocarpa Miq. 74 ilicinwn Sonder 124 hirsu/a Sonder 106 ingens Miq. 90 panifica (Delile) Miq. 77 ko/schyanurn Miq. 102 rigida Miq. 74 lutewn (Vahl) Miq. 101 riparia Miq. 77 na/alense (Hochst.) Miq. 121 schimperiana Miq. 79 - var. minor Sonder 123 sur (Forssk.) Miq. 77 neumannii (Kunth & Bouché) Miq. 101 /honningiana Miq. 77 ol/oniifolium Miq. 154 /rachyphylla Miq. 74 ova/urn (VahJ) Miq. 165 vogeliana Miq. 78 poli/urn (Vahl) Miq. 159 populifolium (Vahl) Miq. 111 UROSTIGMA (Gasp.) Miq. 11 , 12, 13, 14, 15, principes (Kunth & Bouché) Miq. 98 18, 19,30,32,34, 35, 36, 37, 47,48,52, 54, rubicUfldurn Miq. 106 88, 94, 187, 221 rej/exwn (Thunb.) Miq. 126 abutilifoliurn Miq. 1I1 rubrum (Vahl) Miq. 114 acrocarpurn Miq. 128 salicifolium (VahJ) Miq. 92 aggrega/um (Vahl) Miq. 126 saussureanurn (DC.) Miq. 98 burkei Miq. 128 schimperi Miq. 128 caffrurn Miq. 90 /erebra/urn (Willd.) Miq. 126 calyp/ra/urn (Vahl) Miq. 116 /honningii (BI.) Miq. 128 ca/alpaefolium Miq. 111 tllUflbergii Miq. 92 corda/um (Thunb.) Gasp. 92 vogelii Miq. 101 dekdekena Miq. 128 xantllOphyllurn Miq. 90 elegans Miq. 158 - var. ovato-corda/urn Sonder 90 eriobo/ryoides (Kunth & Bouché) Miq. 98 fazokelense Miq. 106

298