• General introduction
• Scope of the database
• Features of the application
• Using the application
  
  • Browsing Plant and Fruit Data
  • Multiple-entry Key to the Plants
  • Browsing Insects Reared from Fruits
• Notes on distributional data for Kenyan plants and insects
  
  • Geographical distribution
  • Temporal distribution
• A note on fruit type and fruit size
• Abbreviations found in plant descriptions
• The photographs
  
  • Photography of fruits
  • Fruits pictured on splash page
  • Photography of insects
  • Copyright of photographs
• Some useful sources
• Acknowledgments
• About the authors

General Introduction

This CD contains a runtime version of a Microsoft Access program enabling users to examine the data and images from a 5-year study (1999-2004) of the wild fruits of Kenya and their associated insect fauna, specifically those that use fruits in their breeding cycles. The program will run on Windows-based computers and does not require that Access is loaded on the computer. It will also run on a Mac running Windows.

Although the primary focus of our study was on fruits as larval habitats of fruit flies (Tephritidae), particularly those of economic importance, many other insects exploit fruits. And while there is a large body of information on the associations of insects with the leafy and woody parts of plants, there is little data available on the guild of insects exploiting fruits, either as pulp or seed feeders, or as parasitoids of species from these two groups. This is especially true for Africa. In addition to tephritids, by including in the CD records of the many other insect groups feeding within fruits we hoped to provide the baseline information necessary for a fuller appreciation of the possibilities available for studying insect-fruit interactions.

We hope that the CD will also have value when used as a field guide to the fruits of Kenya and other parts of eastern Africa. Fruits are an important food for many species of animals, including primates, bats, and birds, and ornithologists and mammalogists should benefit from using this CD. Of course, fruits are the vessels in which seeds develop, and plant species have evolved many strategies for protecting and dispersing their seeds. Many edible fruits are brightly colored (often red, orange, or yellow) and easy to locate. For these fruits, whose bright colors attract the attention of fruit-eating animals, seed dispersal often results from fruit consumption. Seeds then pass through the vertebrate gut and are deposited in feces, or are swallowed and regurgitated; either way at some distance from parent plants. Duller fruits (brown, black) are more difficult to see and are less likely to be fed upon by diurnal (daylight-active) animals. These fruits are often eaten by nocturnal fruit bats and other night-active mammals (e.g. bush babies, palm civets). Fruit bats lack the highly developed sonar of their insect-feeding cousins and, accordingly, they are not particularly agile fliers in habitats with dense foliage. Plant taxa that have caulicarpous fruits (i.e. fruits developing on inflorescences that appear on older wood [trunk or limbs]) present food that is easily accessed by fruit bats. The most familiar examples of this are the figs, but cauliflory has evolved in many other plant families (e.g. Annonaceae), particularly in the tropics, where fruit bats are common. For the generalist interested in natural history, knowledge of the fruits will add to the pleasures of the outdoor life, and we hope the CD appeals to a wide audience. Plants are easier to identify when flowering or fruiting and, while flowering specimens are the cornerstone of plant taxonomy, fruits are often available when flowers are not. These two features of plants complement each other, greatly expanding the season when plants may be readily identified in the field.

Scope of the database

The plant database includes 928 plant taxa (mostly species; some subspecies and varieties) representing 118 familes. Nearly all species are native to Kenya, but some exotic, naturalized (e.g. guava) and cultivated (e.g. mango) species are also included. Our collecting efforts, particularly during the earlier stages of the project, targeted likely hosts of frugivorous tephritids and were, therefore, biased toward sampling fleshy fruits. Many important plant taxa, including most of the Asteraceae, Burseraceae, and Fabaceae, contain few species with fleshy fruits, and were less frequently collected. Additionally, we concentrated mostly on woody plants which normally produce larger numbers of fruits than do herbaceous plants. Though by no means exhaustive, coverage of woody plants was substantial, representing about 40% of all the species (and a much higher percentage of those with fleshy fruits) treated in Kenya Trees Shrubs and Lianas (Beentje 1994), the “bible” for identification of Kenyan woody plants. Certain other groups were also sampled extensively, including the herbaceous Cucurbitaceae and Vitaceae, and the parasitic Loranthaceae (mistletoes). Finally, because certain fruit fly groups consume flower parts (e.g., of Asteraceae) and not fruits, while others may eat flower and fruit parts (e.g., of Acanthaceae), representatives of these plant families were also included in our sampling.

Collections were made in many habitat types, the majority in forested areas, from the coastal forest mosaic in the east to Mt. Elgon on the western border, from the southern forests to Marsabit Mountain in the north, and at elevations from sea level to 3125 m.

We reared at least 672 insect species from the fruit collections. In order of decreasing diversity, these were 273 Lepidoptera, 220 Hymenoptera, 145 Diptera and 34 Coleoptera species. Poorly preserved specimens (mostly moths) that could not be definitively identified or assigned to a specific morphospecies were not included in the total. Tephritid fruit flies, with 123 species, and tortricid and pyralid moths, with 87 and 43 species respectively, were the most common frugivores.

Features of the application

1. Plants can be browsed by family or genus to generate species lists. Users can then select individual species to view a description of the plant, a list of taxonomic and biogeographical features, images of fruit and plant specimens, a list of insect taxa reared from the fruit, and images of those insects (where available). With few exceptions, plant names and authors correspond to those found in The African Flowering Plant Database (http://www.ville-ge.ch/cjb/bd/africa/index.php).
2. A multiple-entry key to plant species, based on characters of both the fruit and associated leafy specimen, will enable users to identify a large proportion of the fruiting woody and herbaceous plants of Kenya. Photographs of fruits and associated leaf and stem specimens provide a visual check of the result(s) generated by the key.
3. Insects reared from collected fruits may also be browsed. Insects are arranged by order (Diptera, Lepidoptera, Coleoptera, and Hymenoptera). Under each order, selecting an insect family name (or genus name for the Tephritidae and subfamily name for the ichneumonoid wasps) generates a list of insect species. Clicking on an individual species produces a list of plant species from which that insect has been reared. For most species this is a list of host fruits, but in the case of a parasitoid indicates that it has emerged from another insect feeding inside that fruit. Again, where available, images of the selected insect species are also displayed. Also indicated are the geographical areas where we sampled fruits that produced the insect species.
4. An illustrated glossary of botanical terms can be used as a primary teaching tool, or referred to when using the multiple-entry key to the plants to call up definitions and images of plant structures and features. Double-clicking on a term in the key will bring up its definition in the glossary. Photographic images illustrating a term are accessed by clicking on the “images” button to the right of each definition. Some self-evident terms are not illustrated. A general, alphabetically arranged glossary is available as are category-related glossaries (e.g. terms related to leaf margin characteristics, fruit shapes, etc.).

Using the application

The application is organized in page format. Each main page is selected by clicking on one of the tabs at the top of the screen. The main data-related pages corresponding to the tabs are;

• Browse Plant and Fruit Data
• Multiple-entry Key to the Plants
• Browse Insects Reared from Fruits

Additionally, there are tabs directing you to;

• Illustrated glossaries of plant terms
• Maps - illustrating the KTSL and UKWF geographical regions (see A note on information regarding the distribution of Kenyan plants).

Browsing Plant and Fruit Data

When using the “Browse Plant and Fruit Data” page, data pages for individual plant species are generated by clicking on that species’ name. A plant data page contains a description of the plant, expandable photographs (single or double click on a thumbnail image to expand it), and data boxes displaying information on the characteristics and geographical distribution of the plant, as well as its fruiting phenology, the latter determined empirically during this project. Exotic species are indicated as such in the first line of a plant’s description. A list of insects reared from the fruit is also provided. The availability of images for a listed insect is indicated (yes/no) to the right of the species name. When available, these images can be seen by clicking on the insect species’ name.

Species page for Acokanthera oppositifolia.

Multiple-entry Key to the Plants

Users select one or more values for each of several character states (e.g. fruit color, leaf type, etc.) by clicking on the appropriate entries. All selections within a character box are “or”, not “and”, selections, making the key inclusive, not exclusive. Say, for example, you have made a collection from a single plant species with compound leaves that are trifoliolate (three leaflets per leaf), and whose fruits include both red and yellow ones. You would select both colors in the Fruit color box. If the fruit from your plant is described in the literature as being yellow only, selecting red in addition to yellow will not eliminate the correct plant from the list generated by your query, because you have included yellow. Additionally, clicking on trifoliolate in the leaf type box will not exclude the correct plant from the results even if it is described in the literature as having 3 or 5 leaflets. Submitting a query will return a list of one or more plant names. Selecting a plant name from the list brings up the data page for that species.

Example of character-state selections using the multiple-entry key.

Results page generated by the query.

It is not necessary to select a value for each character state. While not selecting a value for one or more character states diminishes the strength of your search, you may very well have entered enough values for other characters to identify your plant or to narrow the possibilities considerably. For example, you may not know whether your plant contains latex. Just leave out that character state.

In the key, we have tried to keep technical terms to a minimum to enable non-specialists to use the key. Thus, for example, when examining the leaf or leaflet edge or margin, the term “with sharp or rounded teeth” includes margins more technically described as “dentate”, “serrate” or “crenate”. Many terms in the key are represented in the illustrated glossaries. Double-clicking on a term will call up the definition and one or more images illustrating it. In putting together the glossary we relied on several sources, particularly The Cambridge Illustrated Glossary of Botanical Terms (Hickey & King 2000).

Browsing Insects Reared from Fruits

When using the “Browse Insects Reared from Fruit” page, selecting an insect species generates a page with information on that species, including a list of fruit species from which that insect was reared. Clicking on any fruit species in the list will generate the data page for that plant. To return to where you were on the “Browse Insects Reared from Fruit” page, simply click on “Browse Insects Reared from Fruit” on the tab panel at the top of the page.

Browse Insects page with Ceratitis fruit flies selected.

Page with list of Ceratitis species, and information and image for the default species (the first in the alphabetically sorted list).

Illustrated glossaries of plant terms

An illustrated glossary of botanical terms can be used as a primary teaching tool, or referred to when using the multiple-entry key to the plants to call up definitions and images of plant structures and features. Double-clicking on a term in the key will bring up its definition in the glossary. Photographic images illustrating a term are accessed by clicking on the “images” button to the right of each definition. Some self-evident terms are not illustrated. A general, alphabetically arranged glossary is available as are category-related glossaries (e.g. terms related to leaf margin characteristics, fruit shapes, etc.).

The Glossary default page with the top (blue letters) button selected.

The alphabetically-ordered Glossary page.

The same glossary page with "Ellipsoid (fruit shape)" selected.

Notes on distributional data for Kenyan plants and insects

Geographical distribution

The primary single-volume sources for identification of Kenyan plants are Kenya Trees, Shrubs and Lianas (KTSL) (Beentje 1994) and Upland Kenya Wildflowers (UKWF) (Agnew & Agnew 1994). In each, a separate scheme is used to illustrate the distribution of the plants considered. Like its precursor, Dale and Greenway’s (1961) Kenya Trees and Shrubs (KTS), KTSL has as its scope Kenyan woody plants but increases its coverage by including lianas, which were not covered in the former. KTSL follows the distribution map format of the multi-volume Flora of Tropical East Africa (FTEA), in which the country is divided into 7 large divisions (K1-K7). UKWF, in contrast, only includes herbaceous plants found in the uplands of Kenya (essentially the southwest quarter of the country, although many of the listed plants have wider distributions). In UKWF, an abbreviated version of actual place names serves to identify each of 20 “vegetation districts”. When using the CD, the data page for each plant species contains information on its distribution as either one or more of the K divisions of KTSL or of the “vegetation districts” of UKWF. When necessary, users can click on the “Maps” tab to see which parts of the country these divisions refer to.

Maps - illustrating the KTSL and UKWF geographical regions

The known altitudinal range for each plant is also provided on the plant data page. In the CD, distributional data (including altitude) of individual plant species sometimes differs from that published in FTEA and UKWF. These changes reflect observations made during the current project.

Distributional data is also provided indicating general areas where we collected specific fruit species during the project. Additionally, areas where fruits were collected that produced particular insect species are indicated on the Browse Insects pages. Areas were defined as follows.

Temporal distribution

The monthly distribution of flowering among woody plant species is taken from KTSL, and indicated with roman numerals. These appear at the end of a plant description. Bracketed roman numerals indicate months of peak flowering. An asterisk (*) indicates that the author (of KTSL) saw less than a dozen specimens of a species. As previously stated, monthly fruit phenology, based on data from the present study is given for each species on its data page.

A note on fruit type and fruit size

We have tried to be as accurate as possible when assigning a fruit type to a species. For some fleshy fruits, however, based on the reference literature we used (mainly UKWF, KTSL and FTEA [see explanation of abbreviations below]), it was sometimes not possible to decide between a drupe and a berry. In these cases, both are listed on a plant species’ page and selecting either in the multiple-entry key will lead to the correct plant.

Fruit size is one of the characteristics of individual plant species that appear on its data page. It is also one of the characters found in the multiple-entry key. Fruit size is a continuous variable, and to keep things manageable, we defined 5 size classes (<10mm, 10-15mm, 15-25mm, 25-40mm, and >40mm), based on the length of the fruit along its longest dimension. Often, more than one fruit size class is listed on a fruit data page, each class separated by a semicolon. For example, on the data page for the cucurbit Oreosyce africana (Cucurbitaceae) two size classes are indicated, 10-15mm; 15-25mm. This simply means that mature fruits of this species occur in sizes that fall within the limits of the two size classes, not necessarily that they range from 10 to 25mm (in fact such a case would be rare). A species with fruits from 12-17mm would fall in these classes, as would a species whose fruits range from 10-24mm.

Abbreviations found in plant descriptions

The photographs

Photography of fruits

With few exceptions, digital images of freshly collected fruits and, in most cases, associated stem and leaf specimens were made in the field. Throughout the project I used either a Fujifilm 2.8 megapixel digital camera, a Nikon Coolpix 4300 digital camera, or a Canon IS3S digital camera to capture images of fruits. Flash was avoided whenever possible to limit high contrast, shadowy images, and to capture natural coloration. In most cases, plant specimens were collected with a long-arm pruner and arranged on the ground with associated fruits. Images of a few infraspecific taxa (subspecies, varieties) are also included. Multiple images are usually provided, including wide-angle to close-up views of plant specimen, fruit, and flowers (when present). A writing pen was placed in the field of view to provide scale. The diameter of the pen barrel is approximately 10 mm. Photographs of entire plants were seldom made, because of the difficulty of controlling contrast and avoiding confusing backgrounds. In all, 2224 images are included representing 866 of the 928 plant taxa collected.

Fruits pictured on splash page and CD cover (left to right)

Photography of insects

Because this was a fruit fly project we concentrated on capturing images of live adult tephritids. Photographs are provided for 111 tephritid species. Usually, examples of both sexes are available, and this is always the case for sexually dimorphic species. Tephritidae, and representatives of several other insect taxa that were reared from fruits, were held with water and a sugar/yeast diet for 3-4 days. Thereafter they were starved for 24 h. They were then transferred to a 20 x 15 x 2.5 cm Plexiglas (Perspex) cage with a removable pane of glass through which photographs were taken. The dull, non-reflective underside of an avocado leaf was generally used as background. Sugar syrup was dotted on the veins of the leaf. Feeding insects remained stationary and were easily photographed. Kodachrome 64 slide film was used throughout. The photographic setup consisted of a CANON F1 camera body and a 100 mm macro lenses, to which a “12x” Raynox Microexplorer supplemental lens was attached. Two weak flashes mounted at either ends of a Lepp Macrobracket provided illumination.

There are also many photographs of pinned adult Hymenoptera and Lepidoptera, and of the genitalia of some moths. We also provide representative examples of most of the other insect taxa. In all, 614 images are included representing 422 of the insect species we reared.

Copyright of photographs

All photographs in this CD remain copyright of the individual photographers - Robert Copeland, Karie Darrow and Koen Maes.

Some useful sources

The African Flowering Plant Database. http://www.ville-ge.ch/cjb/bd/africa/index.php.

Agnew, A. D. Q. & S. Agnew. 1994. Upland Kenya Wild flowers. A Flora of the Ferns and Herbaceous Flowering Plants of Upland Kenya, 2^nd edition. East Africa Natural History Society. Nairobi, Kenya. 374 p.

Beentje, H. 1994. Kenya Trees, Shrubs and Lianas. National Museums of Kenya. Nairobi, Kenya. 722 p.

Copeland R. S., R. A. Wharton, Q. Luke and M. De Meyer. 2002. Indigenous hosts of Ceratitis capitata (Diptera:Tephritidae) in Kenya. Ann. Entomol. Soc. Amer. 95: 672-694. (Includes details on collection and rearing methodologies for the fruits included in this CD).

Dale, I. R. and P. J. Greenway. 1961. Kenya Trees and Shrubs. Buchanan’s Kenya Estates Limited. Nairobi. 654 p.

Heywood, V. H. 1993. Flowering Plants of the World, 2^nd Ed. B. T. Batsford, London. 336 p.

Hickey, M. & C. King. 2000. The Cambridge Illustrated Glossary of Botanical Terms. Cambridge University Press. Cambridge. 208 p.

Pijl, L.Van Der. 1969. Principles of Dispersal in Higher Plants. Springer-Verlag, Berlin. 154 p.

Acknowledgments

We are indebted to the many individuals and organizations who contributed to making the CD. Foremost among these was Glenn Sequeira who was responsible for all the software programming. The East African Natural History Society (EANHS) gave us permission to use the plant descriptions and the map of vegetation districts from Upland Kenya Wild Flowers. The National Museums of Kenya (NMK) did the same for the map illustrating the K1-K7 distribution zones of Kenya and for plant descriptions from Kenya Trees, Shrubs and Lianas. We thank the current editor of the Flora of Tropical East Africa series (FTEA), Henk Beentje, for allowing us to use descriptions of the small number of plants not described in these two books. Millicent Okumu and Peris Machera were instrumental in attending to the laboratory handling of fruit collections and the rearing of insects from them. Rose Justus, Asha Nassor, and Augustus Hare did an excellent job providing similar services at our coastal rearing facility. Wilberforce Okeka, Mathias Ngonyo, Saidi Chidzinga, and Juliet Muriuki were interpid fruit finders and collectors. Karie Darrow of the National Museum of Natural History, Washington DC made the dissections of Lepidoptera genitalia. The superb images found in the CD of these and many of the adult moths are a result of her much appreciated technical skills. Koen Maes also photographed some of the Lepidoptera. Matt Buffington, Systematic Entomology Laboratory, USDA/ARS, provided invaluable training and help with photography of the microhymenoptera, and the quality of these images is due largely to his interest and support. We thank Hans Herren, the Director, International Centre of Insect Physiology and Ecology ICIPE), for providing office space, an indispensable field vehicle, and continuing support. Dr. Richard Bagine of the Kenya Wildlife Service was very helpful in granting us permission to sample fruits within Kenyan national parks.

Accurate identification of host plants and the various insect species that exploit them was basic to developing our database and assigning names to specimens and photographs. We greatly appreciate the interest and commitment of the following systematists and their respective institutions; Marc de Meyer, Museum of Central Africa, Tervuren (Tephritidae); Ian White, Natural History Museum, London (Tephritidae); Amnon Freidberg, Tel Aviv University (Tephritidae); the following scientists, all of Systematic Entomology Laboratory, USDA/ARS; David Adamski (Coleophoridae, Stathmopodidae, Gelechioidea); John Brown (Tortricidae), Matt Buffington (Figitidae), Mike Gates (Eurytomidae), Eric Grissel (Torymidae), Al Norrbom (Tephritidae), Chris Thompson (Syrphidae), and Norm Woodley (Stratiomyidae, Tachinidae); Scott Miller (Geometridae, Noctuidae, Nolidae, Pyralidae, Yponomeutidae) and Wayne Mathis (Heleomyzidae) both of the National Museum of Natural History, Washington DC; Koen Maes, National Museums of Kenya (Crambidae, Pyralidae); Chris Lyall, Natural History Museum, London (Curculionidae); Alexander G. Kirejtshuk, National Academy of Sciences, St. Petersburg (Nitidulidae).

This work was supported in part by USAID grant no. PCE-G-00-98-0048-00 (in collaboration with the International Centre of Insect Physiology and Ecology and its African Fruit Fly Initiative) and in part by USDA/CSREES/IFAFS grant no. 00-52103-9651, both to R. Wharton, Texas A&M University.

The authors

Robert S. Copeland is Visiting Scientist and Director of African field activities for the Hydrilla Project at the International Centre of Insect Physiology and Ecology, Nairobi Kenya. He is currently based in Jinja, Uganda. Email: rcopeland@icipe.org or hunteriazeylanica@yahoo.com.

W.R. Quentin Luke is a Research Associate of the National Museums of Kenya and is currently the East African Field Conservationist for Fairchild Tropical Botanic Garden, Miami. He is also the Focal Point of the East African Plant Redlisting Authority and an alternate member for Africa on the CITES Plants Committee. Based in Nairobi, he can be reached on quentin.luke@swiftkenya.com.

Robert A. Wharton is a Professor in the Department of Entomology at Texas A&M University where he teaches and conducts research on systematics of parasitic wasps, with emphasis on species used in biological control. Email: rawbaw2@tamu.edu.