Metamorphosis Occ. Supp. 4 Complete.Pdf

LEPIDOPTERISTS’ SOCIETY OF AFRICA

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EDITORIAL Editor: Doug Kroon Scientific advisers: Mark Williams, Martin Krüger, Rolf Oberprieler, Stephen Henning, Henk Geertsema, Alan Gardiner, Dick Vane-Wright, Axel Hausmann.

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Covers: Panoramic view of Kaya Muhaka forest edge; inset: side view of Charaxes lasti September 2000 METAMORPHOSIS Occasional Supplement 4 1 ______PRELIMINARY SURVEY ON BUTTERFLIES AND MOTHS AND THEIR HABITATS IN TWO KAYA FORESTS OF THE KENYA COAST INGO LEHMANN a and ESTHER KIOKO b

a Uferweg 30, 23996 Bad Kleinen, Germany b Department of Invertebrate Zoology, National Museums of Kenya, P.O. Box 40658, Nairobi, Kenya

ABSTRACT:The forest stratification, plant, butterfly and larger moth diversity of Kaya Muhaka and Kaya Kinondo (Kenya, Coast, Kwale District) are described. Both Kayas have old secondary forest patches and are certainly not unmodified relics. The Lepidopteran fauna includes species not previously recorded from Kenya or from coastal Kenya and was diverse in Kaya Muhaka. The Shimba Hills were found to be the potential source for the butterfly faunas of both Kayas. Together, all three areas support c. 30% of the total Kenyan butterfly fauna. 23% are coastal endemics and only 1.6% of species have a central and western Africa distribution. In contrast, for example 18–21% of plant species with a central and western Africa (Guineo-Congolian) distribution occur in Kaya Muhaka. Butterfly conservation priority is recommended all habitats in the Shimba Hills and for nearby coastal forests.

INTRODUCTION

Objective The main aim of the 1994 and 1996–1998 field work was to study as intensively as possible two coastal forests1 south of Mombasa, namely Kaya Muhaka and Kaya Kinondo, with the hope of throwing some light on the butterfly and moth fauna and their habitats. For both Kayas, no equivalent survey exists. Most of the information and data presented here is based on our report (Lehmann & Kioko, 1998). All information collected is not yet completely analyzed. Therefore, the objective of this article is to give a preliminary survey of the Lepidoptera fauna and their habitats, namely the forest stratification and floristic composition. In addition to our report a comparison with the butterfly fauna of the Shimba Hills will be done here and is based on our assessment of the data published by Sevastopulo (1973/1974). The Shimba Hills were chosen for comparison because they are close to both Kayas, constitute the largest coastal forest block in Kenya south of Mombasa and are a potential source area.

Distributions of butterflies and moths in the lowland forests of the Afrotropical region The Afrotropical Region (Crosskey & White 1977) has a butterfly fauna that consists of 301 genera and of some 3607 known species (Ackery et al. 1995). Of these c. 56% are restricted to forests (de Jong & Congdon 1993).

1 In this article the term “forest“ will be applied according to White (1983) for a continuous stand of trees with a canopy that varies in height from 10 m or more. The term “coastal forest“ will be used after Hawthorne (1993) for forest in the coast which is “ ... the land over the sedimentary (and intrusive volcanic) rocks of the coastal plains and plateau, to the east of the exposed basement complex land.“

2 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Largely based on the publication by Carcasson (1964), Ackery et al. (1995) stated, that within this region, the greatest density of butterfly species occurs in the forests below 1500 m. These lowland forests are “very much poorer“ in species in eastern Africa, although their butterfly fauna shows “unmistakable affinities“ with the species rich western lowland forest fauna. The impoverishment of the eastern forests “must have been due to their extreme fragmentation and reduction during interpluvial phases ... and to the tenuous and shortlived character of recent connections with the western forest reservoir.“ The occurrence of numerous endemic species and distinct subspecies in the lowland forests of eastern Africa suggests that broad based links have only been possible in the remote past. 400 Forest butterfly species, of which 79 species are endemic, inhabit the “eastern African coastal forests“ (Burgess et al. 1998). For the Lepidoptera of the Afrotropical region major eco-geographic sub-regions were proposed by Carcasson (1964). The Sylvan sub-region, Lowland forest division, includes the Eastern sub-division (Coastal forest zone) which comprises all the lowland forest to the east of the eastern tablelands, from Kenya to Zululand (Natal, South Africa). In Kenya, lowland forests occur mainly along the coast and in western Kenya. Larsen (1991) stated that “870 or so“ butterfly species occur in Kenya and of these, c. 400 species could be found in the Kakamega area (western Kenya) alone. Only “a small group of species“ is found in all the forest zones from Senegal to coastal Tanzania and Kenya. But “a very large proportion“ of the coastal forest species are endemic to the East African coast, including few limited to the Kenya and/or Tanzania coast, and the bulk of these endemics have close relatives in the equitorial forests. Larsen mentioned that it would appear that there has been contact between forests of western Africa and the coastal forests at various times in the Miocene or early Pliocene, as well as at a more recent date, probably through southern Tanzania and Malawi. Lovett (1993) also discussed the links between the forests of western and eastern Africa and pointed out that they existed possibly only during wet periods and that the relatively stable and warm climate of the Indian Ocean during the last 2.3 Myrallowing the survival of western (Guineo-Congolian) relict endemic species. Collins & Larsen (1996) stated that “some 30 species have been added to the Kenyan list“. Congdon & Collins (1998) mentioned “approximate[ly] 895“ butterfly species for Kenya and listed 25 endemic species. Their figures indicate that Kenya has a substantially lower species richness compared with Tanzania where c.1370 species including 121 endemics occur and Uganda with c. 1242 species and 34 endemics. There is little knowledge about the distributions of moth species in the lowland forests of the Afrotropical region.

Coastal Forests: their Status and Significance The coastal areas of East Africa and the western Indian Ocean contain some of the world’s richest ecosystems and provide valueable resources for over 30 million people, but the intensity and nature of human activities increasingly threatens their productivity and biological diversity (Lindén 1993; Richmond 1998). This is partly dfue to a high steady population increase (Goliber 1985). Today, the coastal forests of eastern Africa are a refuge for a wide variety of plant species (Brenan 1978; White 1983; Hawthorne 1993) and an important, highly threatened centre of endemism for plants, mammals, birds, reptiles, frogs, butterflies, snails and millipedes (Burgess et al. 1998).

September 2000 METAMORPHOSIS Occasional Supplement 4 3 ______Kenya2 has the most diverse forests in East Africa (IUCN/WCMC, 1992). Its coastal forests support the highest diversities of flora (Beentje 1988; Robertson & Luke 1993; Wass 1995) and fauna (Bennun 1995; Waiyaki 1995; Bennun & Njoroge 1999). The flora is part of the Zanzibar-Inhambane regional mosaic (Moll & white 1978; White 1983). Coastal forests occupy c. 660 km2 in Kenya of which the largest is Arabuko-Sokoke with 370 km2, but the majority is smaller than 5 km2 (Burgess et al. 1998). They are located within 30-50 km inland from the Indian Ocean – except for those within the forest belt of the Tana River floodplain (Andrews et al. 1975; Hughes 1985, 1990) - and are isolated from each other by non-forest habitats. Many patches of forest north of the Pangani River (Tanzania) once enclosed Kaya or Kaya-like clearings once (Hawthorne 1993). Kaya is a Mijikenda word for their small central residential villages protected by the surrounding forest (Spear 1978), but has several meanings today (Robertson & Luke 1993). The Mijikenda are nine closely related but distinct peoples (the Kauma, Giriama, Chonyi, Jibana, Kambe, Ribe, Rabai, Durma and Digo). They came from Sing-waya in southern Somalia hinterland at the turn of the 17th century and settled initially at the southern Kenya coast, but left their Kayas between the 1830’s and 1870’s (Spear, 1978). Today, few Kayas are still used for ceremonies or as burial groves (Robertson & Luke 1993). Several have already declined in size (Hunt et al. 1981). Based on our assessment of the data in Robertson & Luke (1993), there are 70 or so Kayas and Sacred Groves in Kenya (some covered by coastal forest), including 46 with an estimated size of c. 31 km2. 23 Kayas and Sacred Groves or c. 6.8 km2 are gazetted as National Monuments (NM) since 1992. They are protected and managed by the National Museums of Kenya (NMK) and their Coastal Forest Conservation Unit (CFCU) in collaboration with the local community (the Committee of Elders) and the district administration.

Classification of Kenyan Coastal Forests A number of different classifications and terminologies have been proposed for the coastal forests of Kenya (Table 1).

MATERIAL AND METHODS

Study area Location and size Kaya Muhaka (also called Kaya Kambe3 or Kaya Mwadabara) is situated close to Muhaka village which is about 32 km south of Mombasa (Kenya, Coast, Kwale District). The forest lies c. 2.5 km northwest of the village Mwabungu (at the Mombasa-Lungalunga road) and

2 The estimates of closed forest area in Kenya vary, depending on definition (World Bank 1987; FAO 1988, 1999; Beentje 1990). The World Bank’s (1988) estimate of natural forest and plantations is 13.870 km2 or c. 2.4% of Kenya’s total available land. FAO’s (1988) data describe broadleaved closed canopy forest, coniferous forest and bamboo of 11.050 km2 or c. 2% of the country. Beentje’s estimate is 5856 km2 for evergreen forest. FAO’s (1999) estimate of natural forest, plantations and bamboo is 12.920 km2 or c. 2.3% of the land area. 3 Another forest also called Kaya Kambe is located in Kilifi District north of Mombasa and was studied by Hunt et al. (1981). 4 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Table 1. Spectrum of classifications including comments of authors on Kaya forests and the Shimba Hills forests. Author Classification/ Terminology Comments Dale (1939) “Lowland Evergreen Rain-forest“; Considered large parts of the Shimba “Evergreen Dry-Forest“ Hills and Kaya Muhaka as lowland evergreen rain forest one type of a “mixed forest“ type with minor variations between both. Moomaw (1960) “Combretum schumanii – Cassipou- “Sterculia-Chlorophora-Memecylon rea euryoides lowland dry forest on lowland rain forest“ in wetter parts coral rag“; “Cynometra – Manilkara (included are the Kayas and Shimba (Sokoke) lowland dry forest“; Hills remnants). “Brachystegia – “Manilkara – Diospyros lowland dry Afzelia lowland woodland“ on slopes forest“; “Sterculia – Chlorophora/ of the Shimba Hills. Forests near Kaya Memecylon lowland rain forest“; Kinondo, namely at Jadini and Diani, “Brachystegia – Afzelia lowland were considered as lowland dry forest woodland“ on coral rag. Birch (1963) “Wet Evergreen forest climax“ Considered two forests near Kaya Kinondo, namely at Jadini and Shimoni, as wet and dry variants of a “Wet Evergreen forest climax“ Lucas (1968) “Lowland forest with wetter and Considered the Shimba Hills as drier types“ “typical lowland rain forest“ Greenway (1973) “Lowland Rain Forest“; “Lowland “Lowland Rain Forest“ and “Lowland Dry Evergreen Forest“; “Deciduous Dry Evergreen Forest“ occur in the Forest“ Shimba Hills. Deciduous forest locally in moist coastal districts (some Brachystegia woodland types). Hunt et al. (1981) “Semi-deciduous forest“ Considered Kaya Kambe and Kaya Ribe (north of Mombasa) as semideciduous with “wetter and drier forest elements“. White (1983) “Zanzibar-Inhambane* Z-I lowland rain forest and undifferentiated forest“ (with wetter “Transitional rain forest“ do not occur and drier types); “Zanzibar- along the Kenyan coast. Treated forest Inhambane scrub forest“; “Zanzibar- other than rain forest as Z-I Inhambane transitional woodland“ undifferentiated forest. Z-I transitional woodland locally in the Shimba Hills. Schmidt (1991) “Tropical ombrophilous alluvial Described four forest formations in forest“; “Tropical semi-deciduous the Shimba Hills National Reserve. lowland forest“; “Tropical evergreen Forests consisting mainly of evergreen seasonal lowland forest“; “Forest trees occur in wet eastern and plantations“ southeastern escarpments and on top of the Kwale plateau. Robertson & Luke sensu White (1983) Mentioned wetter and drier types of (1993) Kaya forests. Treated Kaya Muhaka as a wetter type. Stated that possibly the wet valleys in southeastern Shimba Hills belong to White’s “Transitional rain forest“. Hawthorne (1993) “Moist forest“, “Dry forest“ Stated for the “Moistest“ coastal forests, “mixed Moist forest“ or “Lowland Rain Forest“ are convenient. terms September 2000 METAMORPHOSIS Occasional Supplement 4 5 ______c. 5.5 km inland from the Indian Ocean (in a direct line) at 45 m a.s.l.4 (UTM 37MEF56235). It is gazetted as a NM since 1992 and is one of the largest Kaya forests in Kwale District. Robertson & Luke (1993) estimated a forest size of 150 ha. Our study area lies in the northeastern and central part of the forest and represents c. 30 ha. The Kaya is isolated by non-forest habitats, namely “Zanzibar-Inhambane secondary grassland and wooded grassland“ sensu White (1983), chiefly Hyphaene compressa grassland (locally modified by fire), Lantana camara thicket and orchards of Cocos, Anacardium and Mangifera. The Shimba Hills6 are located c. 15 km northwest of Kaya Muhaka. Kaya Kinondo (also called Kaya Ngalaani) is a coral rag forest, situated c. 7 km southeastern of Kaya Muhaka (Fig. 6). It lies c. 5 km north of Chale Point on the road from Diani to Chale Point, c. 100 m inland from the Indian Ocean at 5-10 m a.s.l.7 (UTM 37MEF6015). It is gazetted as a NM since 1992. Robertson & Luke (1993) estimated a forest size of 30 ha8. Our study area (c. 7 ha ) lies in the central and southern part of the Kaya. It is isolated by non forest habitat, namely “Zanzibar-Inhambane evergreen and semi- evergreen bushland and thicket“ sensu White (1983) and Hyphaene compressa grassland. The Shimba Hills are located c.23 km northwest of Kaya Kinondo.

Historical and recent human influences and disturbance Hollis (1909) stated that felling of trees in or near the Kayas is forbidden and that people go many miles to fetch their building poles and firewood. In contrast, human influences on the Kaya forests were reported by Spear (1978), who stated that e.g., the Digo collected copal - the dried resin of Hymenaea verrucosa - from the forest and supplied timber to main ports south of Mombasa. The Digo established Kaya Muhaka and Kaya Kinondo. The latter was their second “main Kaya“9, the most important Kaya after Kaya Kwale. Kaya Kinondo gave rise to Kaya Muhaka, probably by 1700. The Digo began to abandon centralized living in all their Kayas during the 1830s and 1840s. Kaya Kinondo was used for ceremonies up to ten years ago (Robertson & Luke 1993). Human influences can still be seen. Several graves are located within our study area, honoured as sacred place (Mzee Mnyenze, pers. comm. 1996) where clearance of undergrowth probably occurred. On the Kaya side of the road

4 Jaetzold & Schmidt (1983) mentioned an altitude of 60 m. According to Robertson & Luke (1993) the altitude is 45 m. [a.s.l. = above sea level] 5 The precise locality was communicated by Universal Transverse Mercator Grid (UTM) to reference to nearest 1.000 m derived from the Tactical Pilotage Chart Kenya, Tanzania 1: 500,000 (Series TPC, Edition I-GSGS, Sheet M-5CG, Ministry of Defence, United Kingdom, 1974). 6 According to Rodgers (1993) the forest patches in the Shimba Hills National Reserve and Forest Reserve constitute 20 km2 in 200 km2 woodland.Wass (1995) mentioned for the Shimba Hills “9 000 ha forest cover” 7 Kokwaro (1982) and Robertson & Luke (1993) mentioned an altitude of 5 m. As result of our own studies we think that the altitude is more or less 5 m near the Indian Ocean but higher, perhaps up to 10 m, in undulating parts further inland. 8 Other forest patches near Kinondo, namely Kaya Mvumoni (on Chale Island) and Kaya Magawoni (Timbwa) were most probably founded by people originated from Kaya Kinondo (Luke, pers. Comm. 2000) but are not included here. 9 Each “main Kaya“ had a main ritual symbol, its fingo (usually a pot full of different medicines), which was supposedly brought from Singwaya. Most important ceremonies can only be performed in main Kayas (Spear, 1978). 6 METAMORPHOSIS Occasional Supplement 4 September 2000 ______forest was cleared in early 1991 (Luke, pers. Comm. 1999). Collecting of fuelwood or cut stumps has not been seen in our study area. In contrast, “many timber trees“ were poached from Kaya Muhaka before 1992 (Robertson & Luke 1993) and three trees of Scorodophloeus fischeri (10-12 m in length) were poached in our study area in early 1996 (I.L., pers. observation). Stumps remaining after pole cutting occur locally. Collecting of fuelwood by local people is still allowed and common. Graves were not found. A Kaya Elder reported of a sacred place used for occasional ceremonies (Mzee Mwakaniki, pers. Comm. 1996).

Climate Michieka et al. (1978) mentioned a mean daily temperature range for SE coastal Kenya of 7-8°C which is larger than the difference between the monthly range of 3–4°C. According to Sombroek et al. (1980) both Kayas fall in the agro-climatic zone“III-1“ (semi-humid with fairly hot to very hot temperatures 24–30°C. The mean annual temperature within the Zanzibar-Inhambane regional mosaic is “more or less 26°C“ north of the Zambezi River (Moll & White 1978). Michieka et al. (1978) mentioned 26.1°C for coastal areas SE Kenya calculated from 13 stations. The temperatures vary during the hotter dry season (December to March) and cooler transitional season (July to mid September). Jätzold & Schmidt (1983) reported of a mean maximum of 32.7 °C in March and a mean minimum of 20.3 °C in July and August in Mombasa (24 years of records). They also published rainfall figures (having at least 10 years of records) for the areas in which Kaya Muhaka and Kaya Kinondo are located. In Kaya Muhaka rainfall exceeded 984 mm in 6 out of 10 years. The average annual rainfall of 1129 mm (23 years of records) with 132 mm during the drier months December to March (February is the driest month), 568 mm during the long rains April to June, 172 mm in July and August and 257 mm during the short rains from September–November. Rainfall data for Kaya Kinondo appear to be unceertain. The only figure available can be derived from two maps published by Jaetzold & Schmidt (1983) showing a 60% reliability of rainfall in 6 out of 10 years of 600–700 mm from March to September and less than 100 mm from November to December. Kaya Muhaka receives in the same periods 700-800 mm and 100-150 mm respectively.

Moll & White (1978) stated that within the Zanzibar-Inhambane regional mosaic the relative humidity is also high throughout the dry season. White (1983) concluded that the dry season is less severe and no month is absolutely dry. For example Schmidt (1991) recorded the relative air humidity throughout one year in a forest gap in Longomagandi Forest (Shimba Hills) and found the lowest value of 40% in February (the driest month) above 90% in August and 90–100% in May and November.

The northern and southern monsoons have a marked effect on air temperature, rainfall and winds. From November to March the northeastern monsoon wind or (Kaskazi) prevails and is usually steady, light, about 5m/sec, and veers north-westerly. From June to September the southeastern monsoon wind or (Kusi) prevails, veers to the west and south-westerly and reaching an average of 9 m/sec (Richmond 1998).

Physiography and Soils Caswell & Baker (1953) divided the coastal belt into four physiographic units. They treated the area of Kaya Kinondo as “Coast Plain“ and that of Kaya Muhaka as“ Foot Plateau“ (the latter is partly covered by Magarini sands of Middle Pliocene age). September 2000 METAMORPHOSIS Occasional Supplement 4 7 ______Michieka et al. (1978) treated the area of Kinondo and Muhaka as “coastal plain“ which is of Pleistocene age, flat to gently undulating, not more than 50 m a.s.l. at its western limit (5–10 km inland from the coastline) and is composed of the fossil coral reef on the seafront and of Kilindini sands further inland. Kaya Muhaka is situated on lagoonal deposits and subrecent marine deposits (Kilindini Sands). The soils are complex, very deep (> 120 cm), of varying drainage condition and colour, texture and salinity (albic and ferralic Arenosols; orthic Ferralsols; gleyic Luvisols to Acrisols and sodic Planosols; vertigleyic Luvisols and pellic Vertisols, sodic phase). Kaya Kinondo is situated on coral limestone (fossil coral reef) with sand admixtures. The soils are shallow (0-50 cm), well drained, dark brown to dark reddish brown, extremely rocky, sandy clay loam to sandy clay (Lithosols and ferralic Cambisols, lithic phase).

Time and Duration of research Field work was done in the dry season as well as in the rainy season over a total period of 17 weeks, usually daily for seven hours, in the following months: February, 1994 (two weeks), early January to mid February, 1996 (five weeks), early June to mid July, 1997 (five weeks) and January to early February, 1998 (five weeks).

Sampling Sites for Description of Forest stratification, Floristic composition and Lepidoptera Forest stratification and Floristic composition In each study area, permanent marked sample sites were established in the forest at least 50 m from the forest edge. Distance between sites was between 100-250 m, except for two sites in each Kaya that were only only 50 m apart. Sample sites were chosen subjectively and a choice of sample sites depended mainly on the representativeness of forest structure (high forest, gaps, thickets of shrubs/trees). Each sample site is 625 m2 in size (25 x 25 m). Ten sample sites or 6250 m2 (0.625 ha) were established in Kaya Muhaka and five or 3125 m2 (0.3125 ha) in Kaya Kinondo. The sum of all sample sites “per Kaya“ (6250 m2 and 3125 m2) is defined as sample area. Literature used for analysing stratification of individuals of trees and shrubs – some of which with a single main-stem will be termed here “treelets“ (Du Rietz 1931) - were Davis & Richards (1933/1934), Richards (1939, 1963, 1996), Grubb et al. (1963), Brünig (1970) for description of crown shapes and Halle et al. (1978). The term “stratum“ will be applied here for a layer or set of tree crowns and shrubs between certain (arbitrary) limits of height. Five strata were defined arbitrarily as A ( > 22 m), B (22-14 m), C (<14-5 m), D (<5-2 m) and E (< 2 m). The term “ emergent“ will be used for trees of stratum A“rising head and shoulders“ above their neighbours (Richards 1996). Trees of the A and B strata will be termed here as “overstorey trees“ and of the C stratum as “understorey trees“. Gaps in probably depleted forest patches and filled with impenetrable tangles of lianes, shrubs/ treelets will be termed here “holes“ (similar [to usage by] Richards 1939). One transect of 450 m2 (75 x 6 m) was marked out in each study area (in the central part of each Kaya) in 1998 to construct a profile diagram showing the stratification and the habitat of Lepidoptera. The distance was measured between all tree, shrub and liana specimens > 5 m tall and are rooting on each sample site. All specimens were tentatively determined, numbered and then figured on a piece of paper by drawing in their position of stem and approximate shape of crown. All specimens  3 cm diameter at breast height (d.b.h.) were measured in regard to their total height, height to lowest larger branch (at least c. 5 cm in diameter at base for trees > 5 m tall), lower limit of crown and width of crown (in north-south and east-west 8 METAMORPHOSIS Occasional Supplement 4 September 2000 ______direction), d.b.h. and diameter at base of trunk (DAB). On each transect the same measurements were undertaken for all individuals > 1 m tall. Plants < 3 cm d.b.h. were also measured in regard to their total height and DAB on three sample sites (plants < 1 m height were omitted), which include one gap and two sample sites of closed forest per Kaya. Heights > 2.5 m were measured with a pocket-sized Suunto clinometer, heights < 2.5 m with a folding rule. Dead standing and fallen trees were measured (height/length, d.b.h. and/or DAB) per sample site and transect. All pieces of dead wood lying on the ground were counted and measured when their length was  0.5 m and their midpoints of length were at least 5 cm in diameter. Their degee of decomposition was visually estimated and grouped into four classes according to Albrecht (1991). Five plots, each 4 m2 (2 x 2 m), were established in the central, NE, NW, SW, SE part on each sample site. Each plot had a subplot of 0.25 m2 (0.5 x 0.5 m)in the central part to measure litter height, leaf litter dry weight (g/m2) and pH values in January, 1998. Leaves (without fruits, flowers or twigs) were dried in a cupboard at an elevated temperature for 72 hours; pH values of soil samples from 10 cm and 30 cm depth were measured with a Checker pH electrode.

Lepidoptera The butterfly and moth fauna was studied by netting in the study area. Baits (a mixture of well-ripened banana, orange, mango and paw-paw fruit) were put either on the ground or in a trap10 . Every day, baits were put on each sample site at the same place on the ground in half shade. Bait-traps were suspended in the same tree. 2-5 m above the ground. The number of bait-trapping sites in each Kaya was enlarged from four in 1996 to five in 1997-1998. Bait-traps were emptied twice (every morning and afternoon). Macroheterocera, termed here “larger moths“, were collected. Butterflies were determined, noted, marked with nail polish according to Brussard’s (1971) modification of Ehrlich and Davidson’s (1960) 1-2-4-7 system and released. A portable ultra-violet light-trap with a 15 watt lamp was used for light-catching of larger moths by hanging the lamp at the forest edge, 2 m above ground, from 7.30 to 11.30. Seven nights of trapping in each Kaya were done. Two light-catching sites, c. 150 m apart, were chosen. In Kaya Muhaka both were located in the northeasternpart and in Kaya Kinondo in the southwestern part. Collected specimens were killed with potassium cyanide, pinned, prepared and labelled. Observations done in the field and the number of specimens per species were noted and later summarized for each butterfly and larger moth species (Lehmann & Kioko 1998 Appendix I & II).

Sampling for species identification We tentatively determined all plants (including tree saplings and seedlings) on each sample site and transect at genus and/or species level. Samples were collected, pressed and later determined by Quentin Luke either at the NMK or in the field. Literature used for identifications and nomenclature of plants were Willis (1980), Haines & Lye (1983), Swinscow & Krog (1988), Johns (1991), “Flora of Tropical East Africa“ (chiefly Polhill) and Beentje (1994).

10 Bait-traps were self constructed in Germany. The basic design involved making a long mosquito- net tube which was tied on top. Two 30 cm wooden sewing hoops at the top and the bottom of the cylinder of netting were used to keep its tubular shape. Lightweight wooden squares were used for the bottom platform at the base of trap. A male screw in the middle of this platform hold the separable small plastic dish containing the bait. September 2000 METAMORPHOSIS Occasional Supplement 4 9 ______Most butterfly species were determined in the field, and a few were pinned and prepared for identification. Literature used for idendification and nomenclature of butterflies were Henning (1989), Kielland (1990), Larsen (1991), Ackery et al. (1995) and Congdon & Collins (1998). Larger moths were collected, pinned and prepared for identification. Their determination was done at The Natural History Museum (BMNH), London. The collections at the NMK, Nairobi, were used for comparison of identifications done at the BMNH. Literature used for a preliminary identification was Pinhey (1956, 1975). The nomenclature of larger moth species follows those found in the collections of the BMNH as well as Carcasson (1976), Fletcher & Nye (1982), Poole (1989), Goodger & Watson (1995), Häuser & Boppré (1997) and Scoble (1999).

Patterns of plant species dominance The dominant species was first determined on each sample site for each stratum (A-D, E omitted). The most dominant species was then determined for the sample area for the same strata. The dominance for trees is usually defined as basal area (Mueller-Dombois & Ellenberg, 1974). The basal area is /4 (d2) and was computed for each tree and shrub specimen with  3 cm d.b.h. Climbers (lianes) were omitted. Their basal area is negligible on the sample sites11. The basal area is here the sum of trunks at d.b.h. level, or above buttresses. If a specimen branched below dbh, the diameter of each stem 3 cm d.b.h. or over was measured separately. A community with numerous dominant tree species is here termed “mixed forest“ and those with a single strongly dominant species a “single-dominant forest“. Some dominant tree species will be treated here as “pioneer“ or “climax (non-pioneer)“ species based on a definition by Swaine & Whitmore (1988). They defined a pioneer as a species with seeds that can only germinate in gaps in the forest canopy open to the sky and in which full sunlight impinges at ground level for at least part of the day. Climax species are those whose seeds can germinate under forest shade (very rarely in full sun as well) and seedlings can establish and survive there and grow.

RESULTS AND DISCUSSION

Classification of Forest As of the different rainfall figures and different patterns of dominance presented here, Kaya Muhaka will be classified as a wetter type and Kaya Kinondo as a drier type of the Zanzibar- Inhambane undifferentiated forest. Both will be treated as a mixed lowland forest (semideciduous lowland forest).

Vertical Stratification Kaya Muhaka: The stratification is heterogeneous with usually five or four strata in closed forest and three or two strata in gaps and“holes“. Palms were only found at the forest edge but rarely in strata A or B (Borassus aethiopum, Cocos nucifera, Hyphaene compressa, Hyphaene coriacea and Elaeis guineensis). Larger epiphytes were not seen in strata A and B and rarely in strata C,D and E. Ferns are rare.

11 Unesco/UNEP/FAO (1978) stated that the basal area of climbers is in general negligible, contrary to Gentry (1982). 10 METAMORPHOSIS Occasional Supplement 4 September 2000 ______

lianes are lianes

. Lying dead dead Lying .

and and lianes occur

species species occur.

Comments

Ficus Ficus

stemmed, wood stemmed,

e erse in plant species. No

Thick rare. Microliches common and obscure the smooth are bark trunks. of very Strnglin locally. Cauliflorous Lianes are canopies common. are bound Locally, together by lian Species rarely occur old and (?) patches in Intact stratum. the A secondary forest with high plant diversityspecies occur. Div ground the on mosses Pole rare. are length) m 5 (≥ stems cutting is locally common.

) or

arely arely

Antiaris Antiaris

Branching Branching

shaped or

and and narrow,

Julbernardia

Some species

)

or or tall

25 m diameter, rarely diameter, m 25

≤

small

some some bent with stems arch

Scorodophloeus Scorodophloeus fischeri

Garcinia Garcinia volkensii

spreading, spreading,

3 m from the trunk, upwards rarely upwardsrarely trunk, the from m 3

broad and

spherical. spherical. Some trunks buttressed,

are are

tipulata

Branching Branching begins low down the trunk.

Gigasiphon macrosiphon

Crown/Trunk regeneration features or

wns are usually long and narrow,

5 5 m in diameter,

1.5 m.

Crowns wide Crowns in lateral composite contact, spreading umbrella ≥ Crowns are less broad and more rounded, 7 m in Buttresses diameter, are uncommon and smaller rarely than in stratum A. in lateral contact. Crowns ≤ over. rare. are Buttresses C broad (e.g. beginsdown lowthe stem. ( abundant is trees of Regeneration magnis rare ( show an lack apparent of regeneration ( toxicaria

are

Many Many

(tallest (tallest

species species

A dense

any

strata strata A and

Gaps Gaps and

) are locally

Scorodophloeus Scorodophloeus

gaps common are

, patchy patchy ,

700 700 m

sedges, sedges, grass, shrubs/

. Evergreen species

Characteristics

36 36 m). Some species are

Few senescent trees. senescent Few

(250

Standing dead stems are rare. rare. are dead stems Standing

iscontinuous ot dense indefinite ery

Discontinuous,never completely but absent. Large scattered through and/or the forest tall trees 32 trees trees. deciduous. senescent Few D Several species have dispersion a clumped (e.g. fischeri N more abundant than in B. V are “holes“ evergreen. common. or absent either developed, Unevenly more or species less continuous. are growth evergreen. of gaps.young intreelets, trees

Description of the stratification of the forest in the study area of MuhakaKaya studyof forest area in the the of stratification the Description

B C

A D

Table 2. Stratum

September 2000 METAMORPHOSIS Occasional Supplement 4 11 ______

). ).

and and

Ficus Ficus

re re rare,

Sorindeia Sorindeia

Stadmania Stadmania

Ficus

sp.) sp.) occurs on

)

discus fraxinifolius

). Lianes a

d forest patches (< 1 000 Ramalina

dominant forest patches

Lecanio

Comments Some species grow bare on almost coral oppositifolia (e.g. Microliches and mosses onlocallytrunks. occur Single with rather even height and high (e.g. occur density stem madagascariensis locallylianes are common. Some species grow on bare coral (e.g. Cauliflorous species occur. spp. and common. lianes Some are bylianes.bound together canopies locally are Stratum Encephalartos A occurs with species trunks ≤ hildebrandtii 2 some m tall, rare. on disturbe bare coral m in less Diverse in plant mosses on the ground. A macro species. No lichen ( in (≥ 5 stems m coral. dead Lying length) are rare. Cut stumps polenot seen. were cutting or

- )

≥

12 m;

Antiaris

Stadmania Stadmania

rrow, some

sp. sp. A FTEA).

Kinondo

25 25 m in diameter,

) ) Branching begins

≤

5 m, upwards rarely upwards m, 5

s abundant ( s abundant

spherical. Some trunks

t with stems arch over,

Milicia excelsa

Diphasia

Chytranthus Chytranthus obliquinervis

Sorindeia madagascariensis

spreading,

sp. A) or very rare (

). ). Some species show an apparent

Drypetes Drypetes natalensis

Crown/Trunk Regeneration Features or Crowns wide heavy, rarely in lateral shaped or contact, composite umbrella 1 spreading buttressed, 2 m. Crowns less broad, more rounded, 7 rarely ≤ 5 in m diameter, long, na bent and tapering ( Crowns are occasionally in lateral overbuttressedSome trunks coral. especially contact. be to tend or rounded small, rather are Crowns tall and narrow, are ≤ 5 m in diameter, rarely up to 7 m, often ben (e.g. rare. are low Buttresses downtrunk. the Crowns are usually long and narrow, rarely broad (e.g. down oftenlowtheBranching stem. begins i trees Regeneration of oppositifolia Diphasia toxicaria ( lack of regeneration

33 m)

Zamioculcas Zamioculcas

ttered through

. Many species are

). ). Many species

700 m

rgreen.

his stratum is not dense. Many

Characteristics Discontinuous or absent. Large and/or tall trees sca the forest (tallest trees 31 Some species not were Senescent trees seen. are deciduous. Variable, locally closed or very open. Species tend to evergreen be more and are represented usually in not stratum A. occur. trees senescent Few T species are evergreen and rarely represented in strata A rare. Standing dead are trees and B. absent. locally and indefinite Very Number of plant individuals and species variable. Gaps 500 usually eve Unevenly absent. A developed, dense growth of grass in trees young and treelets shrubs/ often gaps. Few species grow on bare coral zamiifolia (e.g. evergreen.

Description of the stratification of the forest in the study area of Kaya studyof forest area in the thethe Description of stratification

B C

A D

Stratum

Table 3:

12 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Kaya Kinondo: The stratification is heterogeneuos with locally only two strata in closed forest (e.g., D/E). “Holes“ were not seen. Palms were only found at the forest edge, rarely in strata A or B (Cocos nucifera, Hyphaene compressa, Hyphaene coriacea). Noteworthy is a baobab (Adansonia digitata) older than 50 years (Mzee Mnyenze, pers. comm. 2000) in stratum A and a large Cycus thouarsii probably older than 600 years (Luke, pers. comm. 2000) in stratum C. Larger epiphytes are rare in all strata. Ferns were not seen.

A comparison of four strata in Kaya Muhaka with those of Kaya Kinondo is shown in Table 4. The mean crown height of all strata is similar. The highest basal area occurs in Kaya Muhaka in stratum A. Kaya Kinondo has a considerably higher number of individuals and basal areas in strata B and C. The total basal area and number of individuals per ha is similar between both Kayas, although Kaya Kinondo has many coral outcroppings where growth of trees and shrubs is limited on bare, pinnacled coral (it was expected that Kaya Kinondo has less individuals per ha due to these coral outcroppings than Kaya Muhaka).

Table 4. Comparison of strata of Kaya Muhaka (1) and Kaya Kinondo (2)

Mean crown Number of individuals Basal area Stratum Kaya height (m) (≥ 3 cm d.b.h.) per ha (m2 ha-1 )

A 1 27.9 61 21.9 2 27.9 32 18.2 B 1 17.8 54 10.4 2 17.5 118 14.2 C 1 8.3 104 4.3 2 9.8 253 9.3 D 1 3.4 293 0.5 2 3.5 99 0.2 Total 1 512 37.1 2 502 41.9

The “above-ground“ structure of a forest stand can be described by a height-diameter curve which shows the relation between height and d.b.h. A height-diameter curve is represented for the forest stand of the study area of Kaya Muhaka and Kaya Kinondo in Figures 1 and 2. Both curves are similar and indicate generally, that for a given d.b.h. a number of very different total heights is possible and vice versa. Trees of stratum A have usually a dbh of > 35 cm. Few emergents as well as trees of > 100 cm d.b.h. occur in both Kayas. In Kaya Muhaka, the d.b.h. of emergents ranges from c. 50 to 140 cm and their height from 32 to 36 m. In Kaya Kinondo, they are between 31 and 33 m tall with a d.b.h. ranging from c. 58 to 107 cm. This appears to be noteworthy for a forest on coral rag. The separation between stratum C and D which is very sharply defined by d.b.h. in Kaya Kinondo is remarkable.

September 2000 METAMORPHOSIS Occasional Supplement 4 13 ______

Figure 1. Height and d.b.h. for 212 trees in the study area of Kaya Muhaka. Dots indicate trees, black dots represent large emergents (solid line is Y= 1.48801*X  0.65200).

Height (m) Height 15

0 0 15 30 45 60 75 90 105 120 135 150 165 180 d.b.h. (cm)

Figure 2. Height and d.b.h. for 193 trees in the study area of Kaya Kinondo. Dots indicate trees, black dots represent large emergents (solid line is Y= 2.18361*X  0.54956).

14 METAMORPHOSIS Occasional Supplement 4 September 2000 ______

Key: 1 Parkia filicoidea; 2 Drypetes reticulata; 4 Sorindeia madagascariensis; 5 Julbernardia magnistipulata; 9 Dracaena usambarensis; I 0 Drypetes natalensis; 11 Lecaniodiscus fraxinifolius; 13 Pseudobersama mossambicensis; 18 Pancovia golungensis ?; 20 Antiaris toxicaria; 21 Hymenaea verrucosa; 24 Mkilua fragrans; 25 Craibia brevicaudata; 26 Cynometra suaheliensis; 35 Asteranthe asterias; 39 Acalypha neptunica; 41 Uvariodendron kirkii; 42 Garcinia volkensii; 58 Dichapetalum ruhlandii; 59 Grandidiera bolvinii; 60 Salacia madagascariensis; 64 Saba comorensis; 68 Synepalum subverticillata; 78 Garcinia livingstonei; 92 Landolphia watsoniana; I 0 I Mildbraedia carpinifolia; 121 Streblus usambarensis; U.=Unidentified; Dt.=Standing dead stem: Dw.=Dead wood.

Figure 3. Profile diagram of a wetter mixed lowland forest (Kaya Muhaka, January, 1998). The strip of forest is 7 5 m long and 6 m wide. Plants < 1 m omitted. The contours of the crown projections below were determined in the field by I.L. taking a position vertically under the extremities of branches. Five strata are present. Stratum E is the only stratum that is more or less continuous. The profile plot runs along a forest patch with some disturbance in stratum E (cut stumps, left), through a "hole" (centre) to a relatively intact, probably old secondary forest patch (right), with standing and fallen dead stems and a high plant species diversity. The forest patch (right) was found to be a wetter site and is the habitat of the rare butterfly species Euxanthe tiberius ssp. tiberius.

September 2000 METAMORPHOSIS Occasional Supplement 4 15 ______

Key: 4 Sorindeia madagascariensis; l 0 Drypetes natalensis; l 8 Pancovia golungensis; 35 Asteranthe asterias; 41 Uvariodendron kirkii; 44 Terminalia catappa; 45 Diphasia sp. A of FTEA; 46 Trichilia emetica; 48 Diospyros ferrea; 49 Mallotus oppositifolius; 50 Stadmania oppositifolia; 53 Milicia excelsa; 54 Tricalysia sp.; 55 Cycas thouarsii; 96 Cola clavata; l 32 Grevea eggelingii; U.=Unidentified; Dt.=Dead standing stem; Dw.=Dead wood.

Figure 4. Profile diagram of a drier mixed lowland forest on coral rag (Kaya Kinondo, January, 1998). The cropping out of coral rag is densely stippled. The strip of forest is 75 m long and 6 m wide. Plants < l m omitted. The contours of the crown projections below were determined in the field by I.L. taking a position vertically under the extremities of branches. The profile plot showing five strata, all are discontinuous. Stratum E is locally absent. Noteworthy are light-demanding species (centre) growing close together in stratum A (Milicia excelsa, Terminalia catappa and Trichilia emetica) and in stratum D (a group of Mallotus oppositifolius), indicating that once there has been a large gap, probably caused by human disturbance. The c. 600 years old Cycas thouarsii is remarkable (left). The forest patch (left) is the habitat of the butterfly species Sallya amulia rosa and Iolaus mermis.

16 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Floristic Composition

Kaya Muhaka: According to the CFCU database (February, 2000) 76 families with 324 species occur. In our sample area one more family and probably seven more species were found. Hence, 77 families with 331 species occur - which is c. 4.7% of all plant species known for Kenya or c. 11 % of all species estimated for the Zanzibar-Inhambane regional mosaic3. 24 Families were recorded in strata A-D. Of these 24 families, 15 (62.5%) are represented by only a single species, three families containing two species whereas six containing three or more species (Table 5). The 24 families contain 62 species (including 10 unidentified species) with 19 overstorey and 20 understorey trees, 22 shrubs/treelets and one liana. On three sample sites or 0.1875 ha, 36 families were recorded in strata A-E, including 14 families restricted to stratum E. Of the 36 families, 19 (53%) containing a single species, ten containing two species and seven containing three or more species (Table 6). 73 Species were recorded including 36 restricted to stratum E.

Kaya Kinondo: According to the CFCU database (February, 2000) 63 families with 187 species occur. In our sample area one more family and probably five more species were found. Hence, 64 families with 192 species occur - which is c. 2.7% of all plant species known for Kenya or c. 6.4% of all species estimated for the Zanzibar-Inhambane regional mosaic. 16 Families were recorded in strata A-D. Of the 16 families, 12 (75%) are represented by only a single species, two families contain two species and two families contain three species with no family containing more than three species (Table 5). The 16 families contain 25 species (including three unidentified species) with 11 overstorey and 4 understorey trees, 9 shrubs/treelets and one liana. The overstorey trees include Diphasia sp. A of FTEA4, an endangered species, only known from Kaya Kinondo (Kokwaro 1982; Beentje 1994; Luke, pers. comm. 1999). On three sample sites or 0.1875 ha, 26 families were recorded in strata A-E including 12 families restricted to stratum E. Of the 26 families, 18 (69.2%) containing a single species,

3 White (l983) mentioned that the Zanzibar-Inhambane regional mosaic has "about 3 000 species of which at least several hundred are endemic". Bennun & Njoroge (l 999) stated that in Kenya "7 000 plant species have so far been recorded ". 4 Diphasia sp. A of FTEA was found to be a climax species. Seedlings and young plants (≤:5: 3 cm d.b.h.) survive and grow under forest shade as well as in gaps (where they tend to grow faster). Some of the marked leaves were still alive after 14 months and appeared to be insusceptible to herbivory (no caterpillars were found). A characteristic feature of the petiole in leaves is one or two swollen pulvini, probably responsible for adjusting leaf positions. In young plants some trifoliolate, occasionally 1-2-foliolate, leaves had an almost erect position to sun-light but two or three different positions of leaflets in the same leaf also occur. Leaves which hang down during day-time were in a more erect position from late afternoon until next morning. In young plants some older as well as developing leaves were shed during the dry season (normally the two outer leaflets first, then the middle leaf, later the petiole) (I.L., pers. observations). The adjusting of leaf positions as described here also supports our idea that it is a climax species and not a pioneer species.

September 2000 METAMORPHOSIS Occasional Supplement 4 17 ______five contain two species and three contain three or more species (Table 6). 41 species were recorded including 21 restricted to stratum E.

Table 5. Plant families of strata A-D and number of species per family with ≥ 3 cm d.b.h. in the sample area of Kaya Muhaka and Kaya Kinondo (unidentified species omitted).

Kaya Plant families Number of species per family

Muhaka Anacardiaceae, Apocynaceae, Bignoniaceae, Connaraceae, Dichapetalaceae, Dracaenaceae, Flacourtiaceae, Guttiferae, Papilionaceae, Rhamnaceae, Sterculiaceae, Tiliaceae, Ulmaceae, Verbenaceae, Violaceae 1 Meliaceae, Mimosaceae, Rubiaceae 2 Annonaceae, Sapotaceae 3 Moraceae, Sapindaceae 5 Euphorbiaceae 7 Caesalpiniaceae 8 Kinondo Anacardiaceae, Apocynaceae, Bombacaceae, Combretaceae, Ebenaceae, Meliaceae, Rhamnaceae, Rubiaceae, Rutaceae, Sterculiaceae, Tiliaceae, Violaceae 1 Annonaceae, Moraceae 2 Euphorbiaceae, Sapindaceae 3

Table 5 shows that the sample area of Kaya Muhaka has a considerably higher number of families and species in strata A-D. 12 families of Kaya Kinondo are shared with Kaya Muhaka, with nine exhibiting similar species numbers. Four families of Kaya Kinondo were not found in Kaya Muhaka. In Kaya Muhaka, the Caesalpinaceae and Sapotaceae are relatively rich in species but were not represented in Kaya Kinondo (see also Table 6). 18 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Table 6. Plant families of strata A-E and number of species per family in Kaya Muhaka and Kaya Kinondo on 0.1875 ha.

Number of species Kaya Plant families per family Muhaka Anacardiaceae, Asclepiadaceae, Bignoniaceae, Celastraceae, Commelinaceae, Cyperaceae, Davalliaceae, Dracaenaceae, Gramineae, Loganiaceae, Malvaceae, Melastomataceae, Meliaceae, Mimosaceae, Papilionaceae, Passifloraceae, Thymeleaceae, Tiliaceae, Violaceae 1

Acanthaceae, Connaraceae, Dichapetalaceae, Flacourtiaceae, Guttiferae, Moraceae, Sapotaceae, Sterculiaceae, Verbenaceae, Vitaceae 2

Apocynaceae, Ebenaceae 3

Rubiaceae, Caesalpinaceae 4

Annonaceae 5

Sapindaceae 6

Euphorbiaceae 9 Kinondo Anacardiaceae, Araceae, Celastraceae, Combretaceae, Connaraceae, Ebenaceae, Erythroxylaceae, Flacourtiaceae, Gramineae, Meliaceae, Rutaceae, Sterculiaceae, Thymeleaceae, Tiliaceae, Verbenaceae, Violaceae, Vitaceae, Zamiaceae 1

Apocynaceae, Guttiferae, Moraceae, Rhamnaceae, Rubiaceae 2

Euphorbiaceae 3

Annonaceae, Sapindaceae 5

The results obtained from Table 6 provide some lessons. First, it is worth noting that both Kayas contain species-rich plant communities on relatively small sample sites. Only three sample sites represent nearly half of all families and c. 22% of all species recorded in each Kaya. Second, on three sample sites more than 50% of the families were only represented by a single species in both Kayas. Third, more than 50% of the families of Kaya Kinondo are shared with Kaya Muhaka. The Annonaceae and Sapindaceae bear a numerical resemblance to Kaya Muhaka where two species of the Annonaceae and four species of the Sapindaceae were recorded. The Euphorbiaceae contains fewer species but all occur in Kaya Muhaka, too.

September 2000 METAMORPHOSIS Occasional Supplement 4 19 ______

If we assume that the simplest measure of species diversity is a count of the number of species (MacArthur 1965), stratum A has the lowest and stratum E the highest plant species diversity in both Kayas whereas Kaya Muhaka has a considerably greater species diversity in strata C, D and E (Table 7).

Table 7. Number of plant species in five strata in Kaya Muhaka (1) and Kaya Kinondo (2) on 0.1875 ha.

Number of plant species per stratum Kaya Site code A B C D E Total 1 1 3 3 7 11 43 47 1 3 1 3 7 13 39 41 1 4 2 2 6 8 24 31 Total 6 6 17 23 68 73

2 2 3 4 5 5 29 33 2 3 1 2 6 1 19 22 2 5 0 3 4 2 12 14 Total 3 7 9 8 38 41

Patterns of Plant Species Dominance and Chorological Categories The pattern of dominance (Table 8) provides several results: First, both Kayas can be termed "mixed forest" as a number of dominant species occur. Second, Kaya Muhaka will be treated here as a "wetter" and Kaya Kinondo as a "drier" forest type. Dominant species of a Moist forest type and of a Dry forest type occur in both Kayas. According to Hawthorne (1993) both forest types refer to the physiognomy and not necessarily to the water availability; e.g., Antiaris toxicaria is a Moist forest species; Rinorea elliptica, Synsepalum brevipes are Moist Maritime-Riverine species; Parkia filicoidea is a Maritime-Riverine species; Sorindeia madagascariensis, Uvariodendron kirkii are Dry Maritime-Riverine species; Acalypha neptunica var. neptunica, Drypetes reticulata, Julbernardia magnistipulata, Scorodophloeus fischeri are Dry forest species. We used dominant overstorey tree species to define here a forest type. The NE of Kaya Muhaka is dominated by Dry forest species in strata AIB and hence, it is a Dry forest type. The central part, where dominance of Dry forest species becomes local, is more of a Moist forest type with dominance of Moist forest species in strata A/B. In Kaya Kinondo, a scattered dominance of Moist forest species and a strongly local dominance of Dry Maritime-Riverine species occur. The latter tend to form "single- dominant forest" patches, indicating a drier forest type. Hawthorne (1993) stated ".. as the forest becomes moister there is less of a tendency towards local dominance ... " Third, strata A-D are heterogeneous over short distances (c. 50-250 m) since there is no pair of sample sites that has exactly the same combination of dominant species in strata A-D. Only few sample sites have the same combination in two strata (AID; AIB; BIC); including three sample sites in the NE of Kaya Muhaka dominated by Julbernardia magnistipulata (which was found to be a climax species) in stratum A and Acalypha neptunica var. neptunica in stratum D and two sample sites in the central part where Scorodophloeus fischeri was 20 METAMORPHOSIS Occasional Supplement 4 September 2000 ______dominant in strata A/B. In Kaya Kinondo, two sample sites are strongly dominated by Sorindeia madagascariensis and Drypetes natalensis in strata B/C. Vertically, dominant Dry and Moist forest species occur on two sample sites in both Kayas. Finally, three dominant species (25%) are shared with Kaya Muhaka, six species (50%) were not found in Kaya Muhaka and three were found in Kaya Muhaka but were not dominant. The most dominant species are different and include Dry forest species in strata A/B/D in Kaya Muhaka; Moist forest species in stratum A and Dry Maritime-Riverine species in B (C ?) in Kaya Kinondo; Moist Maritime-Riverine species in stratum C in Kaya Muhaka and in stratum D in Kaya Kinondo.

Table 8. Dominant species of ten sample sites and the most dominant species of the sample area in strata A-D on 0.625 ha in Kaya Muhaka and on 0.3125 ha in Kaya Kinondo. For the most dominant species the basal area (m2) and the percentage of the total basal area per stratum of the sample area are given.

Kaya Stratum Dominant Species Most Dominant Species Muhaka A Antiaris toxicaria, Julbernardia magnistipulata, Scorodophloeus fischeri Parkia filicoidea, Scorodophloeus fischeri 3.3 m2 or 24.1 %

B Cynometra suaheliensis, Cynometra webberi, Scorodophloeus fischeri Julbernardia magnistipulata, Scorodophloeus 1.76 m2 or 27.2% fischeri, Sorindeia madagascariensis, Synsepalum brevipes

C Craibia brevicaudata, Drypetes natalensis, Drypetes Synsepalum brevipes reticulata, Fernandoa magnifica, Julbernardia 0.59 m2 or 22.1 % magnistipulata, Scorodophloeus fischeri, Sorindeia madagascariensis, Synsepalum brevipes

D Acalypha neptunica var. neptunica, Cynometra Acalypha neptunica var. suaheliensis, Julbernardia magnistipulata, neptunica 0.056 m2 or Mildbraedia carpinifolia var. carpinifolia, Mkilua 16.7% fragrans, Streblus usambarensis Kinondo A Adansonia digitata, Antiaris toxicaria, Trichilia Antiaris toxicaria emetica 2.16 m2 or 37.9%

B Diospyros ferrea, Diphasia sp. A, Sorindeia Sorindeia madagascariensis madagascariensis 3.27 m2 or73.5%

C Diphasia sp. A, Drypetes natalensis, Sorindeia Drypetes natalensis 0.65 madagascariensis, Terminalia catappa m2 or 22.2%

D Grewia plagiophylla, Polysphaeria parvifolia, Rinorea elliptica 0.009 Rinorea elliptica, Uvariodendron kirkii m2 or 15%

Hawthorne (1993) determined the global distribution of plant species of the East African coastal forests and summarized eight chorological categories which are: "Coastal" (endemic to the Indian Ocean coastal belt or extending only a short distance from it); "Eastern" (extending e.g., along rivers to the Lake Victoria Regional mosaic; elsewhere in eastern central Africa; in or near the Rift Valley); "Oceanic" (as Coastal or Eastern but also on Indian September 2000 METAMORPHOSIS Occasional Supplement 4 21 ______Ocean is lands, excluding offshore islands); "Zambesian" (widespread in Zambesian but not in Sudanian, Somali-Masai nor Guineo-Congolian region); "GuineoCongolian" (common in at least Congolian region, including species extending to Madagascar); "Wide African" (widespread, but not previous categories, including afromontane and savanna species, excluding species well established in the GuineoCongolian region); "Tropical" (species found beyond Indian Ocean islands);"Unknown" (e.g., genera needing revision). Hawthorne assigned plant species to each category and listed a sample of taxa. We compared his sample of taxa with our sample of 62 respectively 25 species ~ 3 cm d.b.h. (Table 9) as well as with our sample of strata A- E (Table 10).

In spite of the problems posed by comparing data of different sample area sizes in Table 9, six of his chorological categories occur in both Kayas which are: "Coastal", "Eastern", "Oceanic '', "Gui11eo-Congolian ", "Wide African" and "Unknown " (the latter consists here mainly of species not listed by Hawthorne). The following percentage of species ≥ 3 cm d.b.h. belonging to these categories (Kaya Muhaka =1; Kaya Kinondo =2): (1) 33.9% Coastal, 21 % Guineo-Congolian, 9.7% Eastern, 6.4% Wide African, 1.6% Oceanic, 27.4% Unknown; (2) 24% Coastal, 16% Eastern, 12% Guineo-Congolian, 8% Wide African, 4% Oceanic, 36% Unknown (Table 9)

Table 9. Number of plant species ≥ 3 cm d.b.h. assigned to chorological categories found on 0.625 ha in Kaya Muhaka (1)and on 0.3125 ha in Kaya Kinondo (2).

Number of plant species Category Kaya Overstorey Understorey Shrubs Liana Total trees trees Treelets

Coastal 1 7 7 7 0 21 2 2 2 2 0 6

Eastern 1 3 2 1 0 6 2 3 0 1 0 4

Oceanic 1 1 0 0 0 1 2 0 0 1 0 1

Guineo-Congolian 1 5 4 3 1 13 2 1 1 0 1 3

Wide African 1 0 1 3 0 4 2 1 0 1 0 2

Unknown 1 3 6 8 0 17 2 4 1 4 0 9

Total 1 19 20 22 1 62 2 11 4 9 1 25 22 METAMORPHOSIS Occasional Supplement 4 September 2000 ______A slightly different figure is obtained if species are compared recorded in strata A-E on three sample sites: (1) 30.1% Coastal, 17.8% Guineo-Congolian, 9.6% Eastern, 9.6% Wide African, 2.8% Oceanic, 30.1 % Unknown; (2) 29.3% Coastal, 12.2% Eastern, 9.8% Wide African, 7.3% Guineo-Congolian, 7.3% Oceanic, 2.4% Zambesian, 31.7% Unknown (Table 10).

Table 10. Number of plant species assigned to chorological categories found in strata A-E in Kaya Muhaka (1) and Kaya Kinondo (2) on 0.1875 ha.

Number of plant species per stratum Category Kaya A B C D E Total

Coastal 1 2 1 4 9 18 22 2 1 2 4 2 11 12

Eastern 1 1 3 5 4 8 7 2 0 3 2 1 5 5

Oceanic 1 0 0 0 0 2 2 2 0 0 0 1 4 3

Guineo-Congolian 1 3 2 3 4 10 13 2 1 0 0 1 3 3

Zambesian 1 0 0 0 0 0 0 2 0 0 0 0 1 1

Wide African 1 0 0 0 2 6 7 2 1 0 0 1 4 4

Unknown 1 0 0 5 4 24 22 2 0 2 3 2 10 13

Total 1 6 6 17 23 68 73 2 3 7 9 8 38 41

Despite the number of species with an unknown category, the figures of Table 9 and 10 indicate that chorological relationships are complex. In both Kayas the number of coastal endemic species is high, possibly between 30 and 34% in Kaya Muhaka and between 24 and 30% in Kaya Kinondo. The higher number of coastal endemic overstorey tree species is remarkable in Kaya Muhaka. It is different [from] that [of] the "Guineo-Congolian" category [which] is well represented in all five strata of Kaya Muhaka whereas in Kaya Kinondo the "Eastern" and "Oceanic" categories appear to be more distributed. The "Wide African " and "Zambesian" categories are slightly higher in Kaya Kinondo; the latter category was not recorded in Kaya Muhaka and was represented by only one E stratum species in Kaya Kinondo.

September 2000 METAMORPHOSIS Occasional Supplement 4 23 ______The Butterfly Fauna

Kava Muhaka: 112 species of butterflies have been recorded from the study area (c. 30 ha) which represent 12.5% of the total known Kenyan butterfly fauna. 88 species (79%) of those were found in the sample area (0.625 ha). About 20% of the species were attracted to bait- traps (Figs 6 & 7). Sample site l (an old secondary fore st patch) and 3 (a tree-fall gap) represent highest diversities in plants (47 and 41 species) and butterflies (between 20-28 species or c. 50% of the total number found in the sample area per year in 1996-1998). These figures indicate a high diversity of butterfly species in the study area as well as in the sample area where two sample sites have a remarkable high plant and butterfly species diversity. We assume that our list is incomplete since some species were expected but were not recorded because they have the reputation for staying at tree-top level and rarely descend (e.g., Pseudathyma lucretioides lucretioides) .

In the study area, 76 species were recorded in 1996 and 80 species in 1998 during the dry season. The lowest number of species (62) was found during the rainy season of 1997. This was partly due to species not recorded during the rains (e.g., Graphium kirbyi, Appias lasti lasti, Pseudacraea eurytus ssp. conradti) and it might be also a consequence of the El Niňo phase that resulted in wetter than normal conditions. In contrast, the total number of individuals recorded in 1996/ 1998 was similar (649/639), but was considerably greater in 1997 (993) mainly due to Bicyclus safitza safitza with 280 individuals and Charaxes lasti ssp. lasti with 215 individuals.

Six species recorded (Nymphalidae) are rare in Kenya (based on Larsen 1991): Acraea aubyni, Acraea zonata, Charaxes pythodoris ssp. nesaea, Euxanthe tiberius ssp. tiberius, Hypolimnas usambara and Pseudacraea eurytus ssp. conradti. All were found to be very local. Additionally, three species were locally recorded and are listed by Larsen (1991) as Kenyan endemics: Acraea matuapa (Nymphalidae), Baliochila minima and Baliochila latimarginata (Lycaenidae). According to Ackery et al. (1995) only Acraea matuapa and Baliochila minima are limited to coastal Kenya. Congdon & Collins (1998) listed the 25 Kenyan endemic species and did not include any of these three species. Following their list, no Kenyan endemic was recorded. However, according to Larsen (1991) and Ackery et al. (1995), Acraea aubyni, Euxanthe tiberius ssp. tiberius and Hypolimnas usambara are endemic to coastal forests in Kenya and Tanzania (the latter two species occur inland to the Usambara Mountains) and Acraea matuapa and both Baliochila species are endemic to coastal areas of Kenya and Tanzania. Hence, there are no local (Kenyan) endemics in Kaya Muhaka but at least three species that are rare and endemic on a larger scale (Kenya and Tanzania). Based on material seen in The Natural History Museum (BMNH, London), in the National Museums of Kenya (NMK, Nairobi), in the Museum für Naturkunde (Berlin) and of Steve Collins (Nairobi) remarkable records are: (1) Lycaenidae: Anthene liodes (Hewitson) only one specimen was collected on 11.vii. 1997, I.L. leg. & det. According to Larsen (1991) it "... seems very scarce in Kenya ..." but is often common in the main equatorial rainforest zone. Kenyan specimens in the BMNH and NMK are all from western Kenya (Busia, Kakamega Forest, Kitale). The only specimens from coastal Kenya (Shimba Hills) were seen in the collections of Steve Collins, who stated the species is "common in the Shimba Hills " (Collins, pers. comm. 2000). Kielland (1990) mentioned it is " ... not very common ... " in Tanzania and occurs also in the east from the Usambaras to the Uzungwa scarp. 24 METAMORPHOSIS Occasional Supplement 4 September 2000 ______(2) Nymphalidae: Charaxes acuminatus Thurau was first recorded near sea-level. Six specimens were found (Lehmann & Kioko 1998, Appendix I), one male was collected on 13. ii. 1996, I. L. leg. & det. It appears to be similar to the Kenyan endemic ssp. shimbanus van Someren. Henning (1989) emphasized that ssp. shimbanus is being found throughout the high forests of the Shimba Hills. Larsen (1991) stated that this species is " . . . essentially a montane butterfly ... " and that "the Shimba Hills population lives at unusually low levels ... and is only found in the highest level forest ...". Ackery et al. (1995) mentioned that the species descends on the Kenya coast to 300 m a.s.l. Specimens from coastal Kenya seen in the BMNH and NMK are from the Shimba Hills and from Mrima Hill (Kwale District).

Kaya Kinondo: 45 species of butterflies, including 12 (27%) not found in Kaya Muhaka, have been recorded from the study area (c. 7 ha) . This are 5% of the total known Kenyan butterfly fauna. 30 species (67%) were found in the sample area (0.3125 ha). About 28% of the species were attracted to bait-traps. Sample site 2 and 3 (both old secondary forest patches) represent highest diversities in plants (33 and 22 species) and butterflies (between 5-19 species or c. 50% of the total number found in the sample area per year 1996-1998). These figures indicate a relatively high diversity of butterfly species in the study area as well as in the sample area where two sample sites have a high plant and butterfly species diversity. Compared to Kaya Muhaka the butterfly diversity is lower. We assume that our list is incomplete for the same reasons as in Kaya Muhaka.

In the study area 21 species were recorded in 1996 and 31 species in 1998 during the dry season. Again, the lowest number of species (15) was found during the rainy season of 1997. This was partly due to species not recorded during the rains (e.g., Amauris ochlea, Bicyclus campinus) and it might be also a consequence of the El Nino phase. The number of individuals found in 199611998 was not similar (75/189) and was low in 1997 (66). The number of species was higher during the dry seasons in both Kayas whereas the number of individuals appeared to be lower in Kaya Kinondo, probably due to low population densities of many species. For example, the only species found in Kaya Kinondo that is rare in Kenya is Hypolimnas usambara. It appears that its population is small since only two specimens were seen in 1996-1998 whereas in Kaya Muhaka 19 specimens were recorded. Bicyclus campinus is common in Kaya Muhaka where 141 specimens were recorded in 1996-1998, including only nine in 1997. In Kaya Kinondo, 34 specimens were recorded in 1996-1998 and no specimen was seen in 1997.

Among the recorded species there is no Kenyan endemic but Iolaus mermis, Graphium kirbyi and Hypolimnas usambara are endemic to Kenya and Tanzania. Iolaus mermis is listed as a Kenyan endemic, restricted to coastal forests of Kenya and NE Tanzania, by Larsen (1991). He stated it " ... may be found sparingly ... in the remaining coastal forests". Ackery et al. (1995) stated it occurs in coastal forests of Kenya/Tanzania, inland to Meru in Kenya and Amani in Tanzania. Kielland (1990) also gave two localities from further inland Tanzania. Graphium kirbyi is endemic to the Kenyan and Tanzanian coast (inland to Morogoro) and "... sometimes common, though apparently rare in the Shimba Hills " (Larsen 1991). Ackery et al. (1995) stated that it is restricted to lowland forests in Kenya/Tanzania.

September 2000 METAMORPHOSIS Occasional Supplement 4 25 ______Remarkable records are: (1) Nymphalidae: Sallya amulia rosa (Hewitson). Two specimens (fresh, not worn) were seen in the lower canopy of the same Drypetes natalensis tree (larval food plant?) on 12- 15. vii. 1997. I.L. found one specimen in the NMK (labelled: Diani 15 m. S. Mombasa K.C. April 1953 N. Mitton). Larsen (1991) stated, he located specimens from Diani in the BMNH, " ... probably migrants ... since it is certainly not a resident ... just penetrating the southern Kenya coast." Ackery et al. (1995) did not mention Kenya but eastern Tanzania as the range of distribution. (2) Nymphalidae: Charaxes acuminatus Thurau was recorded again near sea-level. Three specimens (probably ssp. shimbanus van Someren) were found (Lehmann & Kioko 1998, Appendix I).

Comparison with the Butterfly Fauna of the Shimba Hills The comparison with the butterfly fauna of the Shimba Hills is based on our assessment of the data published by Sevastopulo (1973/1974). Our results show that in the Shimba Hills5, Kaya Muhaka and Kaya Kinondo a total of 262 species of butterflies have been recorded, comprising 29.3 % of the total known Kenyan butterfly fauna. Five families are represented in all three areas (the Papilionidae, Pieridae, Lycaenidae, Nymphalidae and Hesperiidae). Table 11 shows that the Papilionidae are less represented per area in percentage of the total (6-13%), but in the Shimba Hills comprise 14 species and in Kaya Muhaka 9 species which are 52% and 33% of the Kenyan species of this family. The Pieridae constitute 10% of the total Kenyan fauna (Larsen 1991). Hence, they are well represented in all three areas (11- 15%). In the Shimba Hills, the Pieridae comprise 35 species or c. 40 % of all Kenyan species of this family. The Lycaenidae are underrepresented in all three areas (≤ 21 %) since 31 % of the total Kenyan species belong to this family. The Nymphalidae constitute a large number of species: on the basis that 39% of the total butterfly fauna in Kenya are Nymphalidae, the family is over-represented, especially in Kaya Kinondo with 58% of the total number of species. The Hesperiidae are less well distributed in both Kayas based on the figure that 17% of the total Kenyan butterfly fauna belong to this family. Of the 262 species, 138 (52.7%) have been recorded only in the Shimba Hills, 11 (4.2%) only in Kaya Muhaka, 4 (1.5%) only in Kaya Kinondo and I species only in both Kayas. 68 species of Kaya Muhaka were not found in Kaya Kinondo but in the Shimba Hills and 8 species of Kaya Kinondo were not found in Kaya Muhaka but in the Shimba Hills. 32 species (12.2 %) have been found in all three areas. Hence, 100 species of Kaya Muhaka and 40 species of Kaya. Kinondo have also been recorded in the Shimba Hills. These figures indicate that the Shimba Hills are the potential species source for the butterfly fauna of both Kayas.

Biogeographical Patterns of the Butterfly Faunas If the butterfly species are assigned to biogeographical elements as determined by Larsen (1991), 11 elements occur in the Shimba Hills, 9 in Kaya Muhaka and 7 in Kaya Kinondo (Special habitats omitted) (Table 12).

5 Sevastopulo (1973/ 1974) listed 248 butterfly species but two species were omitted here (Lycaenidae: Deudorix diopolis Hewitson (= wardi Mabille) and Euchrysops dolorosa (Trimen) since neither occur in Kenya (Larsen 1991; Ackery et al. 1995; Collins & Larsen 1996). Of the 246 butterfly species, 20 species are rare in Kenya (based on Larsen 1991). 26 METAMORPHOSIS Occasional Supplement 4 September 2000 ______Table 11. Butterfly families, their number of species and percentage of total number of species in the Shimba Hills, Kaya Muhaka and Kaya Kinondo and total number of species per family.

Number of butterfly species per family and area Shimba Hills Kaya Muhaka Kaya Kinondo Total Number Family code Number % Number % Number % 1 14 6 9 8 6 13 14 2 35 14 17 15 5 11 37 3 53 21 18 16 5 11 57 4 105 43 55 49 26 58 108 5 39 16 13 12 3 7 46 Total 246 100 112 100 45 100 262

Table 12. Biogeographical elements of butterfly species of three coastal forests. Number of species per area, percentage of the total per area, total number of species of all three areas and percentage for each element are given.

Number of species per area Shimba Hills Kaya Muhaka Kaya Kinondo Totals Biogeographical element Number % Number % Number Number % General distributions 29 12 19 17 12 27 31 12

All forest zones, incl. coast 36 15 20 18 7 16 36 14

Main forest zone 1 0.4 1 1 0 0 2 0.8

Equatorial forest zone 2 0.8 0 0 0 0 2 0.8

Coastal forest zone 56 23 35 31 14 31 60 23

Transitional species 13 5 6 5 3 7 13 5

All Africa open formations 36 15 12 11 2 4 38 14.5

Sudanian open formations 3 1 0 0 0 0 3 1

Somali open formations 7 3 1 1 0 0 7 2.7

Zambesian open formations 43 17 9 8 3 7 45 17

Montane distributions 1 0.4 1 1 1 2 1 0.4

Special habitats 2 0.8 0 0 1 2 2 0.8

Unknown 17 7 8 7 2 4 22 8

Total 246 100 112 100 45 100 262 100

Table 12 indicates that the bulk of all species belong to the coastal forest zone, including a number of endemic species. Endemics of all three areas include e.g., Graphium kirbyi,

September 2000 METAMORPHOSIS Occasional Supplement 4 27 ______Pentila tropicalis, Acraea satis and Hypolimnas usambara. Endemics not recorded in Kaya Kinondo but in the Shimba Hills and Kaya Muhaka include e.g., Graphium colonna, Appias lasti lasti, Baliochila minima, Teriomima micra, Acraea matuapa, Acraea rabbaiae mombasae, Acraea zonata, Charaxes lasti ssp. lasti, Charaxes violetta ssp. maritimus, Euxanthe tiberius ssp. tiberius and Physcaeneura leda. Endemics not found in Kaya Muhaka but in the Shimba Hills and Kaya Kinondo include Iolaus lalos lalos. Among the species of the Shimba Hills is Spialia kituina, listed by Congdon & Collins (1998) as a Kenyan endemic. Another species of the Shimba Hills, Neptis rogersi, is limited to coastal areas in Kenya according to Larsen (1991) and Ackery et al. (1995) but is not listed as a Kenyan endemic by Congdon & Collins (1998).

The Shimba Hills have the highest number and Kaya Kinondo the lowest number of endemics. All three areas have a number of species occurring in all forest zones, stretching from Senegal to Tanzania and Kenya. Species from the main forest zone, stretching from Senegal to western Kenya, are under-represented since only one species was found in the Shimba Hills (Fresna nyassae) and one in Kaya Muhaka (Anthene liodes). Species from the equatorial forest zone, stretching from the Congo basin to western Kenya, are also underrepresented with two species only recorded in the Shimba Hills (Sarangesa maculata, Neptis kiriakoffi). One montane forest species (Charaxes acuminatus) occurs in the Shimba Hills and in both Kayas. A large proportion of the total number of species recorded in each area are inhabitants of forests, not able to penetrate into savannah vegetation: 39% in the Shimba Hills, 50% in Kaya Muhaka and 47% in Kaya Kinondo.

Noteworthy is the large proportion of the Zambesian element in the Shimba Hills with many of the species represented by the Pieridae and Lycaenidae. Larsen (1991) mentioned that the Zambesian fauna has produced a rich variety of Pieridae and Acraeinae, many of which reach Kenya, in addition to those of the forest zone. 27% of all species recorded in Kaya Kinondo have "General distributions", containing the most common and widespread African butterflies.

The Larger Moth Fauna

Kaya Muhaka: 12 families and 165 species of larger moths have been recorded in the study area, of which 16% are still unidentified. About 85% of the species were attracted to light on two light-catching sites in the understorey. Since the range of the lamp was small and few light-catching sessions were undertaken it appears that the number of species is remarkable, indicating a species richness for a relatively small area of understorey vegetation. Holloway (1989) stated that the majority of species in Old World rain forests appear to fly in or above the canopy. Kaya Muhaka is not a rain forest but the situation is probably similar. If so, our list is far from complete even for the small area where light catching was done since probably few canopy species were recorded. 15% of the species were seen by day in the lower strata and/or were attracted to bait traps. Species attracted to bait-traps include 24 Noctuidae (Lehmann & Kioko 1998, Appendix II) in particular Giria pectinicornis and species of the genus Anomis, Gracilodes and Eudocima as well as the Arctiidae Secusio drucei. Species seen by day include some Noctuidae that will be called here "litter species" since they were often resting on or hiding among the litter e.g., Achaea lienardi, Amyna punctum, Mocis conveniens and Attonda 28 METAMORPHOSIS Occasional Supplement 4 September 2000 ______adspersa (Pinhey 1978, called Amyna punctum a "grassland or bush species"). The Noctuidae Digama africana was often resting together with Zamarada rufilinearia (Geometridae) in the lower canopy of two Drypetes reticulata trees, growing on wetter sample sites. The only Kenyan record of Zamarada rufilinearia was found in Fletcher (1974, p. 169: "Kenya: Mombasa (van Someren), 1 male "). Some species appear to have a central and western Africa distribution e.g., Caryonopera breviramia, Metaleptina nigribasis, Pseudogiria polita and Rougeotiana xanthoperas.

From the data in Pinhey (1975), Sevastopulo (1979, 1982) and Dall’Asta (1997) and from the material seen in the BMNH, NMK and in the Museum fur Naturkunde, species not previously recorded from Kenya (1) or from coastal Kenya (2)6 include: (1): Lymantriidae: Lymantria leucerythra Collenette, at light on 17.i.1998, I.L. leg. & det. The BMNH has 4 specimens: from Zomba, Mt. Mlanje (Nyasaland), Kilossa and Jinja (Uganda). The NMK has got specimens from Amani (Usambara Mountains), probably the nearest records to Kaya Muhaka. Noctuidae: Grammodes congesta Berio, at light on 26.vi.1997, I.L. leg. & det. The BMNH has got specimens from Senegal, Gold Coast, Belgian Congo, Madagascar. Nearest records to Kaya Muhaka are labelled: Tanganyika Terr. Lindi, 14. ii. 1925 W.E. Cutler, B.M. 1925- 202; Usambara N.L. Tanganyika 2650 ft. Jan. 1922 T.A. Barns; Uganda: Turkana Prov. v. 1934 D.R. Buxton. The latter record might be a Kenyan specimen. Noctuidae: Metaleptina nigribasis Holland, one female was attracted to a bait-trap on 24. vi. 1997, I.L. leg. & det. The specimens in the BMNH are all from West Africa (Sierra Leone, Liberia, Ivory Coast, Gold Coast, South Nigeria, Cameroon, Belgian Congo, Gabon). Two specimens in Berlin are from Cameroon. Noctuidae: Caryonopera breviramia Hampson, at light on 26. vi. 1997, I.L. leg. & det. The BMNH has specimens from West Africa (French Guinea, Cameroon, Gabon) and from West Kivu. (2): Lymantriidae: Euproctis rufopunctata Walker, at light on 17.i.1998, LL. leg. & det. The BMNH has two Kenyan specimens from Mt. Elgon (April 1933, T.H.E. Jackson B.M. 1935- 177), several from southern Africa (Natal, Nyasaland, Pondoland) and one from Lake Kivu, Rugege Forest. The Museum fur Naturkunde has got two specimens from Transvaal and the NMK has several from Amani, the nearest records to Kaya Muhaka. Notodontidae: Eurystaura griseitincta Hampson, at light on 09. vii. 1997, I.L. leg. & det. The NMK has got specimens from Ngong, Nairobi, the Malaba Forest (near Kakamega) and from Uganda (Kibale, Jinja, Mabira Forest and Bugoma Forest). The BMNH has two specimens from Durban, Natal.

Kaya Kinondo: The general impression is that the diversity of larger moths is much lower than in Kaya Muhaka.

Although intensive fieldwork was done only 6 families and 64 species have been recorded; 10% of them are still unidentified. 15% of the species were recorded at the lamp in the understorey but all light-catching sessions were disappointing, partly due to on-shore winds. 70% of the species were recorded by day and include some "litter species" e.g. Achaea lienardi, Amyna octo, Amyna punctum, Attonda adspersa, Oglasa nana and Thyas arcifera

6 The term "coastal" will be used here for the coastal zone as defined by Carcasson (1964). Species not previously recorded from Kenya or from coastal Kenya were represented by a single specimen. The specimens will be preserved in the NMK. September 2000 METAMORPHOSIS Occasional Supplement 4 29 ______(Noctuidae). The Noctuidae Erebus walkeri was often hiding in or among coral rag. 15% were attracted to bait-traps (only Noctuidae). Again, species of the genus Anomis were well represented. Few species appear to have a central and western Africa distribution e.g., Anomis punctulata and Facidina semifimbria. The Geometridae Racotis breijeri Prout was first recorded from Kenya. One male was resting on a trunk of Trichilia emetica on 23.i.1996, I.L. leg. & det. The BMNH has specimens mainly from Madagascar (Diego Suarez) and a paratype, labelled: Griffin H. Jan. 1915 H.G. Breijer, L.B. Prout Coll. B.M. 1939-643 (from Transvaal Mus.?); no locality. Scoble (1999) gives the locality of the holotype which is: South Africa, Transvaal, Nelspruit.

The Families of the Larger Moth Fauna Table 13 indicates that the Noctuidae and Geometridae are the most diverse families in the understorey; the Lymantriidae come third in Kaya Muhaka, the Arctiidae in Kaya Kinondo. The Metarbelidae were recorded only in Kaya Muhaka; Dall’Asta (1997) reported that he did not find this family in his sample of the Kakamega Forest (Western Kenya) but that specimens were well represented in samples of the Bossematié Forest (eastern Ivory Coast).

Table 13. Larger moth families, their number of species and percentage of total number of species in Kaya Muhaka and Kaya Kinondo and total number of species per family.

Number of Larger Moth Species per Family and Area

Kaya Muhaka Kaya Kinondo Total Family Number % Number % Number % Limacodidae 1 0.7 0 0 1 0.6 Metarbelidae 4 3.0 0 0 4 2.4 Epiplemidae 2 1.4 1 1.7 2 1.2 Geometridae 27 19.4 11 19.0 35 21.l Saturniidae 2 1.4 0 0 2 1.2 Lasiocampidae 1 0.7 0 0 1 0.6 Sphingidae 5 3.6 1 1.7 5 3.0 Arctiidae 9 6.5 6 10.4 12 7.2 Ctenuchidae 1 0.7 0 0 1 0.6 Lymantriidae 10 7.2 1 1.7 10 6.0 Notodontidae 3 2.2 0 0 3 1.9 Noctuidae 74 53.2 38 65.5 90 54.2 Total 139 100 58 100 166 100

30 METAMORPHOSIS Occasional Supplement 4 September 2000 ______CONCLUSIONS

We follow the theory of island biogeography by MacArthur & Wilson (1967) who implied that the theory could also be applied to continental habitat islands, and Hawthorne (1993) who stated that the two main causes of variation in any community are logistic/historical factors and habitat availability; other factors (e.g., disturbance) play a subordinate role.

1. Logistic/historical factors. Plant and Lepidoptera species with a central and western Africa distribution occur in Kaya Muhaka and Kaya Kinondo, indicating that both forest islands were connected to forests of central and western Africa once. The percentage of such Lepidoptera species is low and does not correspond with the higher percentage of plant species with a central and western Africa (Guineo-Congolian) distribution. In contrast, the high proportion of endemic coastal plant species seems to correlate with the high proportion of endemic coastal butterfly species. Based on the data presented in Tables 9, 10 and 12 the percentage of endemic plant species appears to give a direct prediction of the percentage of endemic butterfly species (Table 14).

Table 14. The percentage of plant species and Lepidoptera with a central and western Africa distribution compared to coastal endemic species in Kaya Muhaka (I) and Kaya Kinondo (2).

Percentage of Species per Area Distribution Kaya Plants Butterflies Larger Moths Central and western Africa 1 18–21 1 2–5 2 7–12 0 1 Coastal (eastern Africa) 1 30–34 31 ? 2 24–30 31 ?

For the Shimba Hills, the percentage of forest butterfly species with a central and western Africa distribution is also low (1.2%) and does not correspond with the larger forest size; the percentage of coastal endemics is high (23%). Since in general, endemic species are an indication of a longer period of divergence in isolation (MacArthur & Wilson 1967), the high percentage of endemics indicate that all three areas are long isolated. For example, the endemic tree Diphasia sp. A of FTEA (currently only known from Kaya Kinondo) is either a relict or a product of recent local speciation. The latter explanation is probably false since Burgess et al. (1998) stated ‟for the plants there is ... little evidence of recent evolution in the coastal forests” and endemics exhibiting ‟single-site endemism” are best interpreted as relicts. If so, Diphasia sp. A is a relict and its population must be long isolated, making its dispersal impossible. Burgess et al. (1998) pointed out that coastal forests are usually of progressively younger ages the nearer they are to the present shoreline. If so, Kaya Kinondo is a younger forest site than Kaya Muhaka. Hence, its connection to the western forests was perhaps over a shorter period, allowing not many western species to immigrate, before it became isolated. The younger age could explain the lower percentage of plant and Lepidoptera species with a central and western Africa distribution. Colonisations of island colonists, as described by MacArthur & Wilson (1967) as being predominantly r strategists, dispersive species with high levels of reproduction and a short generation time, have probably contributed to the higher proportion of plant species with a "Wide African" (8-10%) and "Oceanic " (4-7%) September 2000 METAMORPHOSIS Occasional Supplement 4 31 ______distribution and to the higher proportion of ecologically widespread butterfly species with ‟General distributions” (27%) and to larger moth genera that contain chiefly species which are mobile, widespread ecological generalists (e.g., Noctuidae: Amyna, Anomis, Spodoptera). If a higher proportion of the (non-endemic) Lepidoptera in Kaya Kinondo are widespread ecological generalists, then it has acquired this fauna through the spread of mobile species throughout the Afrotropical region and perhaps beyond. This is less obvious for plant and Lepidoptera species in Kaya Muhaka. For example, its diverse noctuid fauna appears to contain fewer ecological generalists. This leads to the conclusion that dominant and in part endemic plant species of the Caesalpiniaceae/ Mimosaceae (=Leguminosae; not recorded in the sample area of Kaya Kinondo) support the diversity of the Noctuidae, as Leguminosae are ‟particularly favoured” as host-plants by the Noctuidae (Holloway 1989). The most probable source area for both Kayas are the Shimba Hills. The fact that nearly all the butterfly species of each Kaya occur in the Shimba Hills suggest that the latter are the potential source area. The Shimba Hills (the largest tract) contain 100 butterfly species (89%) of Kaya Muhaka (the medium tract), the medium tract contain 33 butterfly species (73%) of Kaya Kinondo (the smallest tract), and the largest tract contain 40 species (89%) of the smallest tract. As in all tracts the proportion of forest butterfly species is high, including one montane forest species, especially the forests in the Shimba Hills (and less its other vegetation types) are the potential source area. Variations in butterfly species composition between all three areas occurred as the (forest) communities were apart, diverging through differences in patterns of local extinctions and immigrations, as described for islands in general by MacArthur & Wilson (1967). Their description that larger islands usually have more species and that species numbers decrease with increasing distance from a source area, agrees with our results as 246 butterfly species occur in the largest tract, 112 in the medium tract and 45 in the smallest tract and since species numbers of plants and larger moths are higher in Kaya Muhaka than in Kaya Kinondo. Perhaps all three areas lost e.g., butterfly species in a particular sequence as Kaya Kinondo became isolated first and species disappeared faster due to its small size and greatest distance to the source area. Forman et al. (1976) stated that island size is generally found to be most or nearly most important as a predictor of species numbers; Shafer (1990) discussed this in more detail. However, in contrast to Kaya Muhaka, Kaya Kinondo’s isolation, small size and greater distance to the source area probably caused more extinctions of Lepidoptera species but fewer extinctions of plant species as its flora is diverse and does not correspond with the lower diversity of the butterfly and larger moth fauna. Vane-Wright· (l978) stated ‟it is not apparent that island butterfly faunas are directly responsive to the ... diversity of their floras”. Kaya Kinondo is one example. 2. Habitat availability. The physical environment is complex and patchy and determines the range of habitats. We conclude from the rainfall figures that Kaya Kinondo receives an average annual rainfall of ~ 1000 mm, which is c. 200 mm less than for Kaya Muhaka, showing a noteworthy variation within c. 7 km. Water availability appears also to be different in each Kaya. In Kaya Muhaka, the central part is more of a Moist forest type. The impression is that dominant Moist forest species in strata NB correspond with wetter habitats (probably with higher groundwater availability). The NE part is more of a Dry forest type and sites also appear to be drier. Generally, dominant species indicate a patchy distribution of Dry and Moist forest types usually within 100-250 m. In Kaya Kinondo, rainwater is limited. Dry Maritime-Riverine species are more dominant e.g., Sorindeia madagascariensis which is an example for a species that might indicate a good supply with

32 METAMORPHOSIS Occasional Supplement 4 September 2000 ______groundwater locally (as it is one of the major tree species in the groundwater-fed Tana River floodplain forests, Andrews et al. 1975; Hughes 1984). A local groundwater supply maybe supports the local dominance of Moist forest species. Dominant Dry and Moist forest species sometimes occur together in strata A-D in both Kayas. However, Kaya Kinondo is drier and hence, Lepidoptera species that tend to favour wetter habitats were not recorded but occur in Kaya Muhaka. Euxanthe tiberius ssp. tiberius, Pseudacraea eurytus ssp. conradti and Zamarada rufilinearia were found to be sensitive to, and thus recognise, small changes that define wetter habitats. This exclusiveness of species inhabiting wetter and not adjacent drier habitats in Kaya Muhaka is an example of what MacArthur (1965) termed ‟between habitat” diversity. We assume that moistness is more available for Lepidoptera in Kaya Muhaka than in Kaya Kinondo during the dry season as the butterfly diversity and the total numbers of individuals were higher in Kaya Muhaka during the dry seasons of 1996/ 1998. Our assumption agrees with results of Janzen & Schoener (1968) who studied insect abundance and diversity in three adjacent forest habitats of increasing moistness in Costa Rica during the dry season and found that numbers of species and numbers of individuals were increasing with moistness. The effect of increasing moistness in the rainy season is not clear yet as the numbers of butterfly species were decreasing in both Kayas and numbers of individuals were increasing only in Kaya Muhaka in 1997 (due to two abundant species). No data from larger moths can be presented yet. 3. Human disturbance. For a long time, the coast has had a relatively large population of agricultural people. Schmidt (1989) pointed out that many forests in East Africa have been exploited by man for at least 2000 years. We assume that occupational periods and forest exploitations assisted with the isolation of Kaya Muhaka and Kaya Kinondo and affected their present vertical stratification since they are neither located in remote nor inaccessible areas. We conclude that both Kayas have old secondary forest patches and are certainly not unmodified relicts: First, evidence is provided by dominant A stratum species which show an apparent lack of adequate regeneration (based on their d.b.h. size-class distribution, Lehmann & Kioko 1998) e.g., Antiaris toxicaria, Parkia filicoidea, Trichilia emetica. These usually emergent species appear to be light-demanding pioneers. They grow only in gaps due to wind throws or disturbance. Gaps caused by disturbance seem to have occurred in a patchy pattern in both Kayas maybe 100-150 years ago as pioneers only represented in stratum A show sometimes a clumped dispersion of > 40 m in diameter. This is more than the length of an emergent. Hence, it is unlikely that such large gaps were caused by wind throws (Figure 4). If so, these patches are old secondary forest. This conclusion agrees with Richards (1973) who stated that the reason for the lack of regeneration of dominant emergent tree species in Africa seems to be that most supposedly primary African forests are in fact secondary. Second, if disturbance occurred it contributed locally to the discontinuous A and B strata. But it is not easy to be sure if the largely discontinuous A and B strata are due to disturbance; Richards (1996) pointed out that the more open A and B strata in African forests might also reflect severe seasonal drought of the climate. Third, tree growth is more limited in Kaya Kinondo (coral outcroppings). Hence, the basal area and number of individuals per ha should be higher in Kaya Muhaka. This was not found and must be due to more human disturbance in Kaya Muhaka. Less disturbed patches occur (chiefly < 1000 m2) maybe for some decades. They are dominated in stratum A by climax species that also occur (not necessarily dominant) in other strata in the same patch e.g. Diphasia sp. A, Julbernardia magnistipulata. Single dominant

September 2000 METAMORPHOSIS Occasional Supplement 4 33 ______forest patches in Kaya Kinondo could also be less disturbed as Hawthorne (1993) stated ‟local dominance ... is common in undisturbed stands of Dry forest”.

CONSERVATION Robertson & Luke (1993) stressed the need to introduce or enhance sustainable management practices for Kenyan coastal forests. Meanwhile, impressive conservation measures have been done by the CFCU (e.g., to disperse and plant rare and/or endemic plant species from a CFCU nursery). As human pressure is evident and future habitat loss is likely in all three areas considered here, we recommend that all habitats in the Shimba Hills should become a priority area in Kenya for butterfly conservation as it represents at least 27.5% of the total Kenyan butterfly fauna, including 20 rare species, one Kenyan endemic and as 61 % are non-forest-dependent species (the forests have been identified as one priority forest area for biodiversity conservation to support populations of sufficient size to have long-term viability, Wass 1995). The moth fauna is probably also diverse. All habitats are a potential source for the Lepidoptera faunas of nearby forests. Hence, priority areas should be larger forest islands near the Shimba Hills (e.g., Kaya Muhaka) to raise each island's immigration and because extinctions are more likely on smaller islands and on islands more distant from the source area as predicted by MacArthur & Wilson (1967). The edges near the forests should be protected to maximize both size (to increase species number) and habitat availability. A zone for complete protection should be marked out to protect the quality habitats (= less disturbed patches). In Kaya Muhaka, this zone should include c. 50 ha of the NE and central part where collecting of fuelwood should be not allowed. Kaya Kinondo is of less importance for Lepidoptera conservation but it is after Diani Forest and Shimoni Forest, one of only three remaining "larger" forests (≥ 30 ha) growing on coral rag on the Kenya coast. Their complete protection is important for the representativeness of coral rag forests as they are so rare. Finally, as all coastal forests have potential value for Lepidoptera conservation, an inventory on Lepidoptera is needed (beginning in larger forests) to find priority areas where species diversity and endemism are high.

ACKNOWLEDGEMENTS We are very grateful to Dr Richard Bagine for his guidance with the research in Kenya; to Quentin Luke for his important help in identifying plant species and for valuable criticism on the draft manuscript; to the CFCU for their immense help and logistical support; to our companions Saidi Ali Chidzinga and Saidi Hamadi Matata for valuable fieldwork; to the Kaya Elders Mzee Hamissi Kassim, Mzee Ali Abdullah Mnyenze and Mzee Omari Bakara Mwakaniki for allowing us access to their land. I.L. is very grateful to Professor Dr Lutz Kobes and Professor Dr Achim Dohrenbusch for their guidance with the research; to David Carter, Martin Honey and Geoff Martin for their help in identifying larger moths in the BMNH; to Hilary Sommerlatte and Steve Collins for valuable criticism on the draft manuscript; to Dr Malte Sommerlatte and his wife as well as to Grit Hecht, John Jones and Heike Langenheim for their support, encouragement and generous hospitality; to Dr Wanja Kinuthia and Dr Koen Maes for their advice and support of the research through the NMK. The fieldwork was carried out under a research permit from the Office of the President (Kenya). The authors gratefully acknowledge financial support provided by the African Butterfly Research Institute (A.B.R.I.), Nairobi (Kenya) towards the printing of colour Figures. 34 METAMORPHOSIS Occasional Supplement 4 September 2000 ______REFERENCES ACKERY, P.R., SMITH, C.R. & VANE-WRIGHT, R.I. (Eds.) 1995. Carcasson’s African Butterflies: an annotated catalogue of the Papilionoidea and Hesperioidea of the Afrotropical region. The Natural History Museum, London. ALBRECHT, L. 199 l. Die Bedeutung des toten Holzes im Wald. Forstwissenschaftliches Centralblatt 110: 106-113. ANDREWS, P., GROVES, C.P. & HORNE, J.F.M. 1975. Ecology of the lower Tana River flood plain (Kenya). Journal of the East Africa Natural History Society and National Museum 151: 1-3l. BEENTJE, H.J. 1988. Atlas of the rare trees of Kenya. Utafiti (Occasional Papers of the National Museums of Kenya) 1 (3): 71 -123. BEENTJE, H.J. 1990. The forests of Kenya. Mitteilungen aus dem lnstitut fiir Allgemeine Botanik in Hamburg 23a: 265-286. BEENTJE, H.J. 1994. Kenya trees, shrubs and lianas. National Museums of Kenya, Nairobi. BENNUN, L. 1995. Arabuko again ... Swara (Magazine of the East African Wild Life Society) 18 (5): 22-23. BENNUN, L. & NJOROGE, P. 1999. Important Bird Areas in Kenya. East Africa Natural History Society, Nairobi. BIRCH, W.R. 1963. Observations on the littoral and coral vegetation of the Kenya coast. Journal of Ecology 51: 603-615. BRENAN, J.P.M. 1978. Some aspects of the phytogeography of tropical Africa. Annals of the Missouri Botanical Garden 65: 437-478. BRÜNIG, E.F. 1970. Stand structure, physiognomy and environmental factors in some lowland forests in Sarawak. Tropical Ecology 11 (1): 26-43. BRUSSARD, P.F. 197 l. Field techniques for investigations of population structure in a ‟ubiquitous” butterfly. Journal of the Lepidopterists' Society 25 (1): 22-29. BURGESS, N. D., CLARKE, G.P. & RODGERS, W.A. 1998. Coastal forests of eastern Africa: status, endemism patterns and their potential causes. Biological Journal of the Linnaean Society of London 64: 337-367. CARCASSON, R.H. 1964. A preliminary survey of the zoogeography of African butterflies. East African Wildlife Journal 2: 122-157. CARCASSON, R.H. 1976. Revised catalogue of the African Sphingidae (Lepidoptera) with descriptions of the East African species. Second edition, E.W. Classey, Faringdon, UK. CASWELL, P. V. & BAKER, B.H. 1953. Geology of the Mombasa-Kwale area (report no. 24). Geological Survey of Kenya. The Government Printer, Nairobi. COLLINS, S.C. & LARSEN, T.B. 1996. Butterflies new to Kenya since 1991, updating, and corrections. In: T.B. Larsen The butterflies of Kenya and their natural history, Oxford University Press, Oxford, pp. 491-500. CONGDON, C. & COLLINS, S.C. 1998. Kielland’s Butterflies of Tanzania Supplement. A.B.R.I. - Lambillionea, Nairobi and Tervuren. CROSSKEY, R.W. & WHITE, G.B. 1977. The Afrotropical Region - A recommended term in zoogeography. Journal of Natural History 11: 541-544. DALE, I.R. 1939. The woody vegetation of the Coast Province of Kenya. Imperial Forestry Institute Paperno. 18: 1-28. DALL'ASTA, U. 1997. Moths collected in the Kakamega forest (eastern Kenya) and a possible use of moths to characterise tropical forests. Metamorphosis Supplement 3: 98-104. September 2000 METAMORPHOSIS Occasional Supplement 4 35 ______DAVIS, T.A.W. & RICHARDS, P.W. 1933/1934. The vegetation of Moraballi Creek, British Guiana: an ecological study of a limited area of tropical rain forest {parts I and II). Journal of Ecology 21: 350-384, 22: 106-155. DE JONG, R. & CONGDON, T.C.E. 1993. The montane butterflies of the eastern Afrotropics. In: J.C. Lovett & S.K. Wasser (Eds.) Biogeography and ecology of the rain forests of eastern Africa, Cambridge University Press, Cambridge, pp. 133-172. DU RIETZ, G.E. 1931. Life-forms of terrestrial flowering plants. Acta Phytogeographica Suecica III, Uppsala. EHRLICH, P.R. & DAVIDSON, S.E. 1960. Techniques for capture-recapture studies of Lepidoptera populations. Journal of the Lepidopterists ' Society 14 (4): 227-229. FAO (1988). An Interim Report on the State of Forest Resources in the Developing Countries. FAO, Rome. FAO (1999). State of the World's Forests 1999. FAO, Rome. FLETCHER, D.S. 1974. A revision of the old world genus Zamarada (Lepidoptera: Geometridae). Bulletin of the British Museum (Natural History), (Entomology), Supplement 22: 1-498, London. FLETCHER, D.S. & NYE, I.W.B. 1982. The Generic Names of Moths of the World. Volume 4: Bombycoidea, Mimallonoidea, Sphingoidea, Castnioidea, Cossoidea, Zygaenoidea and Sesioidea. Reprint 1995. The Natural History Museum, London. FORMAN, R.T.T., GALLI, A.E. & LECK, C.F. 1976. Forest Size and Avian Diversity in New Jersey Woodlots with Some Land Use Implications. Oecologia (Berl.) 26: 1-8. GENTRY, A.H. 1982. Patterns of Neotropical plant species diversity. Evolutionary Biology 15: 1-84. GOLIBER, T.J.1985. Sub-Saharan Africa: Population Pressures on Development. Population Bulletin 40(1):1 -46. GOODGER, D.T. & WATSON, A. 1995. The Afrotropical Tiger-Moths: an illustrated catalogue, with generic diagnoses and species distribution, of the Afrotropical Arctiinae (Lepidoptera: Arctiidae). Apollo Books, Stenstrup. GREENWAY, P.J. 1973. A classification of the vegetation of East Africa. Kirkia 9: 1-68. GRUBB, P.J., LLOYD, J.R., PENNINGTON, T.D. & WHITMORE, T.C. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy and floristics. Journal of Ecology 51: 567-601. HAINES, R.W. & LYE, K.A. 1983. The sedges and rushes of East Africa. East African Natural History Society, Nairobi. HALLÉ, F., OLDEMAN, R.A.A. & TOMLINSON, P.B. 1978. Tropical Trees and Forests: an architectural analysis. Springer-Verlag, Berlin. HÄUSER, C.L. & BOPPRÉ, M. 1997. A revision of the Afrotropical taxa of the genus Amerila Walker (Lepidoptera: Arctiidae). Systematic Entomology 22: 1-44. HAWTHORNE, W.D. 1993. East African coastal forest botany. In: J.C. Lovett & S.K. Wasser (Eds.) Biogeography and ecology of the rain forests of eastern Africa, Cambridge University Press, Cambridge, pp.57-99. HENNING, S.F. 1989. The Charaxinae Butterflies of Africa. Aloe Books, Johannesburg. HOLLIS, A.C. 1909. A note on the graves of the Wa-Nyika. Man 9: 145. HOLLOWAY, J.D. 1989. Moths. In: H. Lieth & M.J.A. Werger (Eds.) Tropical Rain Forest Ecosystems: biogeographical and ecological studies, Elsevier, Amsterdam, pp. 437-453. HUGHES, F.M.R. 1984. A comment on the impact of development schemes on the floodplain forests of the Tana River of Kenya. The Geographical Journal 150 (2): 230-244. HUGHES, F.M.R. 1990. The influence of flooding regimes on forest distribution and composition in the Tana River floodplain, Kenya. Journal of Applied Ecology 27: 475-491. 36 METAMORPHOSIS Occasional Supplement 4 September 2000 ______HUNT, K.J., HAWTHORNE, W.D., RUSSELL, A. & JONES, A. 1981. Kaya: an ethnobotanical perspective. Report of the Oxford Ethnobotanical expedition to Kenya, January-June 1981. Unpublished report. IUCN/WCMC 1992. The Conservation atlas of tropical forests. Africa. IUCN, Gland and Macmillan, London. JAETZOLD, R. & SCHMIDT, H. 1983. East Kenya (volume II, part C). In: Ministry of Agriculture, Kenya, in Cooperation with the German Agricultural Team (GA T) of the German Agency for Technical Cooperation (GTZ) Fann Management Handbook of Kenya, Erhart GmbH, Trier. JANZEN, D.H. & SCHOENER, T.W. 1968. Differences in insect abundance and diversity between wetter and drier sites during a tropical dry season. Ecology 49 (1): 96-110. JOHNS, R.J. 1991. Pteridophytes of Tropical East Africa. Royal Botanic Gardens, Kew. KIELLAND, J. 1990. Butterflies of Tanzania. Hill House, Melbourne and London. KOKWARO, J.O. 1982. Rutaceae. In: R.M. Polhill (Ed.) Flora of Tropical East Africa, Balkema, Rotterdam. LARSEN, T.B. 1991. The butterflies of Kenya and their natural history. Oxford University Press, Oxford. LEHMANN, I. & KIOKO, E. 1998. Notes on butterflies and moths and their habitats in two Kaya forests (Kenya, Coast, Kwale District). Report submitted to the Office of the President (Kenya) and to the National Museums of Kenya. Unpublished report. LINDÉN, O. 1993. Resolution on Integrated Coastal Zone Management in East Africa signed in Arusha, Tanzania. Ambio 22 (6): 408-409. LOVETT, J.C. 1993. Climatic history and forest distribution in eastern Africa. In: J.C. Lovett & S.K. Wasser (Eds.) Biogeography and ecology of the rain forests of eastern Africa, Cambridge University Press, Cambridge, pp. 23-29. LUCAS, G.L. 1968. Kenya. In: I. Hedberg & O. Hedberg (Eds.) Conservation of vegetation in Africa south of the Sahara, Acta Phytogeographica Suecica 54: 152-159. MACARTHUR, R.H. 1965. Patterns of species diversity. Biological Reviews 40: 510-533. MACARTHUR, R.H. & WILSON, E.O. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, New Jersey. MICHIEKA, D.O., VAN DER POUW, BJ.A. & VLEESHOUWER, J.J. 1978. Soils of the Kwale-Mombasa-Lungalunga area. Reconnaissance Soil Survey Report no. R3. In: Ministry of Agriculture and National Agricultural Laboratories, Kenya Soil Survey, Nairobi. MOLL, E.J. & WHITE, F. 1978. The Indian Ocean coastal belt. In: M.J.A. Werger (Ed.) Biogeography and Ecology of Southern Africa, W. Junk, The Hague, pp. 561-598. MOOMAW, J.C. 1960. A study of the plant ecology of the coast region of Kenya Colony, British East Africa. The Government Printer, Nairobi. MUELLER-DOMBOIS, D. & ELLENBERG, H. 1974. Aims and methods of vegetation ecology. John Wiley & Sons, New York. PINHEY, E.C.G. 1956. The Emperor Moths of Eastern Africa. Journal of the East Africa Natural History Society 23 No. 1 (98): 1-62. PINHEY, E.C.G. 1975. Moths of Southern Africa. Tafelberg Publishers, Cape Town. PINHEY, E.C.G. 1978. Lepidoptera. In: M.J.A. Werger (Ed.) Biogeography and Ecology of Southern Africa, W. Junk, The Hague, pp. 763-773. POLHILL, R.M. (current Ed.) 1949 - . Flora a/Tropical East Africa. Balkema, Rotterdam. POOLE, R.W. 1989. Noctuidae (parts 1-3). In: J.B. Heppner (Ed.) Lepidopterorum catalogus (New Series), Fascicle 118, E.J. Brill, Leiden.

September 2000 METAMORPHOSIS Occasional Supplement 4 37 ______RICHARDS, P.W. 1939. Ecological studies on the rain forest of southern Nigeria. I. The structure and floristic composition of the primary forest. Journal of Ecology 27: 1-61. RICHARDS, P.W. 1963. Ecological notes on West African vegetation. II. Lowland forest of the Southern Bakundu Forest Reserve. Journal of Ecology 51: 123-1 49. RICHARDS, P.W. 1973. Africa, the "Odd Man Out''. In: BJ. Meggers, E.S. Ayensu & W.D. Duckworth (Eds.) Tropical Forest Ecosystems in Africa and South America: a Comparative Review, Smithsonian Institution Press, Washington, pp. 21-26. RICHARDS, P.W. 1996. The tropical rain forest: an ecological study. Second edition, Cambridge University Press, Cambridge. RICHMOND, M.D. (Ed.) 1998. A guide to the seashores of eastern Africa and the Western Indian Ocean islands. Sida/Department for Research Cooperation, SAREC. ROBERTSON, S.A. & LUKE, W.R.Q. 1993. Kenya coastal forests. Report of the NMKIWWF Coast Forest Survey. Unpublished report. World Wide Fund for Nature, Nairobi. RODGERS, W.A. 1993. The conservation of the forest resources of eastern Africa: past influences, present practices and future needs. In: J.C. Lovett & S.K. Wasser (Eds.) Biogeography and ecology of the rain forests of eastern Africa, Cambridge University Press, Cambridge, pp. 283-331. SCHMIDT, P.R. 1989. Early exploitation and settlement in the Usambara Mountains. In: A.C. Hamilton & R. Bensted-Smith (Eds.) Forest Conservation in the East Usambara Mountains, Tanzania, IUCN, Gland and Cambridge, UK, pp. 75-78. SCHMIDT, R. 1991. Ecology of a tropical lowland rain forest: plant communities, soil characteristics and nutrient relations of the forests in the Shimba Hills National Reserve, Kenya. Dissertationes Botanicae 179, Gebriider Borntraeger, Berlin-Stuttgart. SHAFER, C.L. 1990. Nature reserves: island theory and conservation practice. Smithsonian Institution Press, Washington and London. SCOBLE, M.J. (Ed.) 1999. Geometrid moths of the world: a catalogue (Lepidoptera, Geometridae). The Natural History Museum, London. SEVASTOPULO, D.G. 1973/1974. The butterflies of the Shimba Hills. Entomologist's Record and Journal of Variation 85: 263 -266, 86: 18-23, 86: 131 - 136. SEVASTOPULO, D.G. 1979. Notes on Heterocera of Mombasa and Neighbourhood (Kenya). Entomologist's Monthly Magazine 115: 181-197. SEVASTOPULO, D.G. 1982. Supplementary notes on Heterocera of Mombasa and Neighbourhood (Kenya). Entomologist's Monthly Magazine 118: 87-92. SOMBROEK, W.G., BRAUN, H.M.H. & VAN DER POUW, BJ.A. 1982. Exploratory soil map and agro-climatic zone map of Kenya, 1980, scale 1: 1,000,000. Exploratory Soil Survey Report no. E1. In: Ministry of Agriculture and National Agricultural Laboratories, Kenya Soil Survey, Nairobi. SPEAR, T.T. 1978. The Kaya complex: a history of the Mijikenda peoples of the Kenya coast to 1900. Kenya Literature Bureau, Nairobi. SWAINE, M.D. & WHITMORE, T.C. 1988. On the definition of ecological species groups in tropical rain forests. Vegetation 75: 8 l -86. SWINSCOW, T.D.V. & KROG, H. 1988. Macrolichens of East Africa. British Museum (Natural History), London. UNESCO/UNEP/FAO 1978. Tropical forest ecosystems: a state-of-knowledge report. Unesco, Paris. VANE-WRIGHT, R.I. 1978. Ecological and behavioural origins of diversity in butterflies. In: L.A. Mound & N. Waloff (Eds.) Diversity of insect faunas. Symposium of the Royal Entomological Society of London 9: 56-70. 38 METAMORPHOSIS Occasional Supplement 4 September 2000 ______WAIYAKI, E.M. 1995. Effects of forest fragmentation, isolation and structure on the richness and abundance of bird communities in major coastal forests of south coast, Kenya. A thesis submitted for partial fulfilment of a degree in Master of Science in Conservation Biology. University of Kent, UK. WASS, P. (Ed.) 1995. Kenya’s Indigenous Forests: Status, Management and Conservation. IUCN, Gland and Cambridge, UK. WHITE, F. 1983. The vegetation of Africa: a descriptive memoir to accompany the Unesco/AETFAT/UNSO vegetation map of Africa. Natural Resources Research XX. Unesco, Paris. WILLIS, J.C. 1980. A Dictionary of the Flowering Plants and Ferns. Cambridge University Press, Cambridge. WORLD BANK, 1988. Forestry Sector Review: Kenya (volume!). World Bank, Washington, DC.

Addendum Unfortunately the book The Coastal Forests of eastern Africa by N.D. Burgess & G.P. Clarke (Eds.) was still in press when this article was completed and it was not possible to check the data in the book against our data.

Note In the Appendices which follow it should be noted that the genus Freyeria is now considered to be Chilades. Furthermore the genus Spindasis on the lists should arguably be replaced by Apharitis or the older name generic designation Cigaritis.

The genus Nola has been transferred to the Noctuidae.

September 2000 METAMORPHOSIS Occasional Supplement 4 39 ______APPENDIX I

Plants recorded in the sample areas of Kaya Muhaka (0.625 ha) and Kaya Kinondo (0.3125 ha). Nomenclature and hierarchical ordering of families follow Beentje (1994) and CFCU database (Feb. 2000). Families or species marked ‘*’ are new records (= species not included in the CFCU database). Species marked ‘?’ are new records but are not yet confirmed by CFCU. Rare species in bold (after CFCU database). U =Unidentified. Categories after Hawthorne (1993): A= widespread African; E =Eastern; E(n) =Eastern (north); E(v) = Eastern (around Lake Victoria); E(z) = Eastern (reaching Zambesian); GC= Guineo-Congolian; GC+ = Guineo- Congolian and beyond; O = Oceanic; OM =Oceanic (Madagascar); Z = Zambesian; ! =Coastal endemic; !(s) =Coastal endemic (southwards); !!= restricted within coast; !!!= very restricted within coast; ? = Unknown.

Kaya Muhaka Family Plant Species and Subspecies Stratum Category

Angiosperms, Acanthaceae Monothecium aristatum T. Anders. E ? Dicotyledons Whitfeldia elongata (Beauv.) C.B.Cl. E GC + Anacardiaceae Sorindeia madagascariensis DC. A,B,C,D,E E(z) Annonaceae Asteranthe asterias (S. Moore) Engl.& Diels ssp. asterias C,D,E ! Mkilua fragrans Verde. C,D,E !! Uvaria lucida Benth. ssp. lucida E ? Uvariodendron kirkii Verde. D,E ! Xylopia parviflora (A. Rich.) Benth. E A Apocynaceae Ancylobotrys petersiana (Kl.) Pierre E ? Landolphia watsoniana Romburg E ? Saba comorensis (Bojer) Pichon C,D,E GC + Asclepiadaceae Tylophora sp.? nov. E ? Bignoniaceae Fernandoa magnifica Seem. C,D,E E(z) Caesalpiniaceae Cynometra suaheliensis (Taub.) Bak.f. A,B,C,D ! Cynometra webberi Bak.f. B,C,D ! Dialium holtzii Harms C,D,E ! Erythrophleum suaveolens (Guill. & Perr.) Brenan A,E GC Gigasiphon macrosiphon (Harms) Brenan A,E !! Hymenaea verrucosa Gaertn. A,E O Julbernardia magnistipulata (Harms) Troupin A,B,C,D,E ! Scorodophloeus fischeri (Taub.) J. Leonard A,B,C,D,E ! Celastraceae Salacia madagascariensis (Lam.) DC. E OM Connaraceae Agelaea pentagyna (Lam.) Baill. E ? Ellipanthus madagascariensis (Schellenb.) Keraudren C,E ! Dichapetalaceae Dichapetalum ruhlandii Engl. D,E E Tapura fischeri Engl. E GC Ebenaceae Diospyros abyssinica (Hiern) F. White ssp. Abyssinica E A Diospyros greenwayi F. White E E(z) Diospyros kabuyeana F. White E ! Euphorbiaceae ? Acalypha fruticosa Forssk. * D ? Acalypha neptunica Muell. Arg. var. neptunica C,D,E GC Alchornea laxiflora (Benth.) Pax & K. Hoffm. D,E A Antidesma venosum Tul. E A Drypetes natalensis (Harv.) Hutch. var. leiogyna Brenan B,C,D,E E Drypetes reticulata Pax C,D,E ! (s) Mallotus oppositifolius (Geisel.) Muell. Arg. D GC + Mildbraedia carpinifolia (Pax) Hutch. var. carpinifolia D ! ? Oldfieldia somalensis (Chiov.) Milne-Redh. * E ! Tragia furialis Bojer E ? Flacourtiaceae Dovyalis macrocalyx (Oliv.) Warb. E ? Grandidiera boivinii Jaub. D,E ! Guttiferae Garcinia livingstonei T. Anders. D A Garcinia volkensii Engl. E ? Loganiaceae Strychnos panganensis Gilg E ! Malvaceae Gossypioides kirkii (Mast.) J.B. Hutch. E ! Melastomataceae Memecylon sansibaricum Taub. var. sansibaricum E ?

40 METAMORPHOSIS Occasional Supplement 4 September 2000 ______

Kaya Muhaka Family Plant Species and Subspecies Stratum Category Angiosperms, Meliaceae Pseudobersama mossambicensis (Sim) Verde. C,D,E ! Dicotyledons Trichilia emetica Yahl D A Mimosaceae Newtonia paucijuga (Harms) Brenan A,B ! Parkia filicoidea Oliv. A,E GC Moraceae Antiaris toxicaria Leschenault A,B GC Dorstenia kameruniana Engl. D GC Ficus sp. D ? Streblus usambarensis (Engl.) C.C. Berg D ? Trilepisium madagascariensis DC. B GC Papilionaceae Craibia brevicaudata (Vatke) Dunn ssp. brevicaudata C,D,E ? Passifloraceae Schlechterina mitostemmatoides Harms E ! Rhamnaceae ? Lasiodiscus mildbraedii Engl. ssp. ferrugineus (Verde.) Faden * D ! Rubiaceae Chazaliella abrupta (Hiern) Petit & Verde. var. abrupta E GC Keetia zanzibarica (Klotzsch) Bridson ssp. zanzibarica E ? Polysphaeria parvifolia Hiern C,D,E E (n) Vangueria randii S. Moore ssp. acuminata Verde. D,E ? Sapindaceae Blighia unijugata Bak. D GC + Chytranthus obliquinervis Engl. C,D,E ! Lecaniodiscus fraxinifolius Bak. ssp. vaughanii (Dunkley) Friis B,C,D,E E (v) Lepisanthes senegalensis (Poir.) Leenh. E A Majidea zanguebarica Oliv. A,E ! Pancovia golungensis (Hiern) Exell & Mendonca C,D,E A Paullinia pinnata L. E ? Sapotaceae Synsepalum brevipes (Bak.) Pennington B,C,D,E GC ? Synsepalum msolo (Engl.) Pennington * D GC Synsepalum subverticillata (E.A. Bruce) Pennington D,E !! Sterculiaceae Cola minor Brenan D,E ! Nesogordonia holtzii (Engl.) Capuron * E ! Thymeleaceae Synaptolepis kirkii Oliv. E ! Tiliaceae Grewia plagiophylla K. Schum. C,D,E ? Ulmaceae* Celtis mildbraedii Engl. * C GC Verbenaceae Clerodendrum incisum Klotzsch E ? Lantana camara L. * D,E ? Violaceae Rinorea ilicifolia (Oliv.) O. Ktze. var. ilicifolia E GC Rinorea squamosa (Tul.) Baill. ssp. kaessneri (Engl.) Grey-Wilson D ! Vitaceae Cissus sciaphila Gilg E ? Cissus sylvicola Masinde & L.E. Newton E ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. C ? U. U. E ? U. U. C ? U. U. D ? U. U. C ? U. U. D ? U. U. C ? U. U. D ? U. U. B ? U. U. A ? U. U. A ? Angiosperms, Commelinaceae Commelina sp. E ? Monocotyledons Cyperaceae Kyllinga cartilaginea K. Schum. E ? Dracaenaceae Dracaena usambarensis Engl. C GC Gramineae Panicum sp. E ? Pteridophytes Davalliaceae Davallia denticulata (Burm. f.) Kuhn var. denticulata E ? Total 38 88 (unidentified species omitted)

September 2000 METAMORPHOSIS Occasional Supplement 4 41 ______Kaya Kinondo Family Plant Species and Subspecies Stratum Category Gymnosperms Zamiaceae Encephalartos hildebrandtii A. Br. & Bouche var. E hildebrandtii Angiosperms, Anacardiaceae Sorindeia madagascariensis DC. B,C,E E (z) Dicotyledons Annonaceae Asteranthe asterias (S. Moore) Engl. & Diels ssp. asterias C,D,E ! Monanthotaxis fornicata (Baill.) Verde. E ! Ophrypetalum odoratum Diels E ! ? Polyalthia stuhlmannii (Engl.) Verde. * E ! Uvariodendron kirkii Verde. D,E ! Apocynaceae Hunteria zeylanica (Retz.) Gardn. E O Saba comorensis (Bojer) Pichon D,E GC + Bombacaceae* Adansonia digitata L. * A ? Celastraceae Salacia elegans Oliv. E ? Combretaceae Terminalia catappa L. A,C,E ? Connaraceae Agelaea pentagyna (Lam.) Baill. E ? Ebenaceae Diospyros ferrea (Willd.) Bakh. B,C,E ? Erythroxylaceae Erythroxylum emarginatum Thonn. E A Euphorbiaceae Drypetes natalensis (Harv.) Hutch. var. leiogyna Brenan B,C,D,E E Drypetes reticulata Pax B,C ! (s) Mallotus oppositifolius (Geisel.) Muell. Arg. C,D,E GC + Flacourtiaceae Ludia mauritiana Gmelin E O Guttiferae Calophyllum inophyllum L. E ? Garcinia livingstonei T. Anders E A Meliaceae Trichilia emetica Yahl A,E A Montiniaceae Grevea eggelingii Milne-Redh. var. keniensis Verde. E !! (= Grevea madagascariensis Baill. ssp. keniensis Verde.) Moraceae Antiaris toxicaria Leschenault A,E GC Ficus sp. B,C ? Oleaceae Chionanthus battiscombei (Hutch.) Stearn E ? Rhamnaceae Lasiodiscus pervillei Baill. ssp. pervillei D,E ? Ziziphus robertsoniana Beentje E ? Rubiaceae Polysphaeria parvifolia Hiern D,E E (n) ? Tricalysia pollens Hiern * E ? Rutaceae Diphasia sp. A of FTEA A,B,C,D,E !!! Sapindaceae Chytranthus obliquinervis Engl. C,E ! Lecaniodiscus fraxinifolius Bak. ssp. vaughanii (Dunkley) Friis E E (v) Majidea zanguebarica Oliv. E ! Pancovia golungensis (Hiern) Exell & Mendonca D,E A Stadmania oppositifolia Pair. ssp. oppositifolia A,B,E E (z) Sterculiaceae ? Cola minor Brenan* D,E ! Thymeleaceae Synaptolepis kirkii Oliv. E ! Tiliaceae Grewia plagiophylla K. Schum. D,E ? Verbenaceae Premna hildebrandtii Guerke E ? Violaceae Rinorea elliptica (Oliv.) O. Ktze. D,E O Vitaceae Cissus sylvicola Masinde & L. E. Newton B,C ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. E ? U. U. D ? U. U. D ? U. U. C ? Angiosperms, Araceae Zamioculcas zamiifolia (Ladd.) Engl. • E Z Monocotyledons Gramineae Oplismenus burmannii (Retz) P. Beauv. E ? Total 29 44 (unidentified species omitted)

42 METAMORPHOSIS Occasional Supplement 4 September 2000 ______APPENDIX II

Species list of butterflies recorded in the Shimba Hills (SH) by Sevastopulo (1973/1974) and in our study areas of Kaya Muhaka (KM) c. 30 ha and Kaya Kinondo (KK) c. 7 ha. Hierarchical ordering and nomenclature follow Larsen (1991) and Ackery et al. (1995). Rare species (in bold) and categories (1-8) after Larsen (1991): 1 = General distributions; 2a = All forest zones, incl. coast; 2b = Main forest zone; 2e = Equatorial forest zone; 2g = Coastal forest zone; 3 = Transitional species; 4a = All Africa open formations; 4b = Sudanian open formations; 4c = Somali open formations; 4d = Zambesian open formations; 5 = Montane distributions; 8 = Special habitats; ? = Unknown. 9 = Kenyan endemic species (after Congdon & Collins 1998).

Superfamily Family Butterfly Species and Subspecies SH KM KK Category Papilionoidea Papilionidae Papilio dardanus Brown X X X 2a Papilio constantinus Ward X 0 X 3 Papilio nireus lyaeus Doubleday X X X 2a; 3 Papilio ophidicephalus Oberthür X X X 2g Papilio demodocus Esper X X 0 1 Graphium angolanus (Goeze) X X 0 4a Graphium philonoe (Ward) X 0 0 ? Graphium leonidas Fabricius X 0 0 3 Graphium kirbyi (Hewitson) X X X 2g Graphium colonna (Ward) X X 0 2g Graphium polistratus (Grose-Smith) X 0 0 2g Graphium policenes (Cramer) X X X 2a; 3 Graphium antheus (Cramer) X X 0 2a; 3 Graphium porthaon (Hewitson) X 0 0 2g; 3 Subtotal 14 9 6 Pieridae Catopsilia florella (Fabricius) X X X 1 Eurema hecabe solifera (Butler) X X 0 1 ? Eurema floricola orientis (Butler) 0 X 0 ? Eurema brigitta brigitta (Stoll) X X X 1 Pinacopteryx eriphia melanarge (Butler) X 0 0 4a Nepheronia argia (Fabricius) X X 0 2a Nepheronia thalassina (Boisduval) X X 0 2a; 3 Nepheronia buquetii buquetii (Boisduval) X X 0 4a Eronia cleodora Hübner X X 0 4a Eronia leda (Boisduval) X 0 0 4c Colotis protomedia (Klug) X 0 0 4c Colotis ione (Godart) X 0 X ? Colotis regina (Trimen) X 0 0 ? Colotis hetaera (Gerstaecker) X 0 0 4d Colotis danae eupompe (Klug) X X X 4a Colotis aurora (Cramer) X 0 0 4b Colotis auxo (Lucas) X 0 0 4d Colotis antevippe (Boisduval) X 0 0 4a Colotis euippe omphale (Godart) X X 0 ? Colotis daira (Klug) X 0 0 4a Colotis amata (Fabricius) X 0 0 ? Colotis evagore (Klug) X 0 0 4a Colotis eris (Klug) X 0 0 4a Belenois aurota aurota (Fabricius) X X X 4a Belenois creona (Cramer) X 0 0 4a Belenois gidica (Godart) X 0 0 4a Belenois thysa (Hopffer) X X 0 4d Dixeia orbona vidua (Butler) 0 X 0 4a Dixeia charina (Boisduval) X 0 0 4d Dixeia spilleri (Spiller) X 0 0 3; 4d Appias epaphia orbona (Boisduval) X X 0 3 Appias lasti lasti (Grose-Smith) X X 0 2g Appias sabina phoebe (Butler) X 0 0 2a Leptosia alcesta inalcesta Bernardi X X 0 2a Mylothris agathina (Cramer) X X 0 1 Mylothris rueppelli rhodesiana Riley X 0 0 3; 4d Pontia glauconome Klug X 0 0 ? Subtotal 35 17 5 Lycaenidae Alaena picata Sharpe X 0 0 4d Pentila tropicalis (Boisduval) X X X 2g Pentila rogersi rogersi (Druce) X 0 0 2g Ornipholidotos peucetia peuceda (Grose-Smith) X 0 0 2g; 3

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Superfamily Family Butterfly Species and Subspecies SH KM KK Category Papilionoidea Lycaenidae Teriomima subpunctata Kirby X X 0 2g Teriomima micra (Grose-Smith) X X 0 2g Baliochila hildegarda (Kirby) X 0 X 2g; 3 Baliochila minima (Hawker-Smith) X X 0 2g Baliochila latimarginata (Hawker-Smith) 0 X 0 2g Deloneura ochrascens ochrascens Neave X 0 0 4d Aslauga purpurascens Holland X 0 0 4d Lachnocnema bibulus (Fabricius) X 0 0 ? Spindasis nyassae (Butler) X 0 0 4d Spindasis victoriae (Butler) X X 0 4d Spindasis apelles (Oberthür) X 0 0 4d Spindasis homeyeri (Dewitz) X 0 0 4d Chloroselas pseudozeritis tytleri Riley X 0 0 4d Axiocerses harpax ugandana Clench X 0 0 4b Axiocerses amanga (Westwood) X 0 0 4a Axiocerses punicea (Grose-Smith) X X 0 2g Iolaus diametra (Karsch) X 0 0 4d Iolaus silanus silanus Grose-Smith X X 0 2g Iolaus mermis (Druce) 0 0 X 2g Iolaus pallene (Wallengren) X 0 0 4d Iolaus lalos lalos (Druce) X 0 X 2g; 4d Hemiolaus caeculus littoralis (Stempffer) X 0 0 2g Hypolycaena philippus philippus (Fabricius) X X 0 4a Hypolycaena buxtoni rogersi Bethune-Baker X 0 0 4d Leptomyrina hirundo (Wallengren) X 0 0 4d Deudorix caerulea obscurata Trimen X X 0 4a Deudorix antalus (Hopffer) X 0 0 1; 4a Deudorix diocles Hewitson X 0 0 2g Deudorix dinochares Grose-Smith X 0 0 4d Deudorix dariaves Hewitson X 0 0 2g Deudorix lorisona (Hewitson) X X 0 4a Anthene lasti (Grose-Smith and Kirby) X X 0 2g Anthene liodes (Hewitson) 0 X 0 2b Anthene lunulata (Trimen) X 0 0 4a Anthene amarah amarah (Guérin-Méneville) X 0 0 4a Anthene kersteni (Gerstaecker) X X 0 2g Cupidopsis jobates jobates (Hopffer) X 0 0 4a Cupidopsis cissus (Godart) X 0 0 8 Pseudonacaduba sichela sichela (Wallengren) X 0 0 4a Lampides boeticus (Linnaeus) X 0 0 1 Cacyreus lingeus (Stoll) X 0 0 1 Leptotes pirithous (Linnaeus) X 0 0 1 Zizeeria knysna (Trimen) X 0 0 1 Zizula hylax (Fabricius) X X 0 1 Actizera lucida (Trimen) X 0 0 ? Azanus jesous (Guérin-Méneville) X 0 0 4a Azanus mirza (Plötz) X 0 0 4a Eicochrysops hippocrates (Fabricius) X 0 X 8 Euchrysops malathana (Boisduval) X 0 0 4a Euchrysops osiris (Hopffer) 0 0 0 4a Euchrysops barkeri (Trimen) X X 0 4d Lepidochrysops peculiaris peculiaris (Rogenhofer) X 0 0 4d Freyeria trochylus (Freyer) X 0 0 4a Subtotal 53 18 5 Nymphalidae Libythea labdaca laius Trimen X X 0 2a Danaus chrysippus (Linnaeus) X X X 1 Tirumala petiverana (Doubleday) X X X 1 Amauris niavius dominicanus Trimen X X X 1 Amauris ochlea ochlea (Boisduval) X X X 2g Melanitis leda (Linnaeus) X X X 1; 3 Gnophodes betsimena diversa (Butler) X 0 0 2a Bicyclus ena (Hewitson) 0 0 X 4d Bicyclus campina ocelligera (Strand) X X X 4d Bicyclus anynana anynana (Butler) X 0 0 4d Bicyclus safitza safitza (Westwood) X X X 1; 3

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Superfamily Family Butterfly Species and Subspecies SH KM KK Category Papilionoidea Nymphalidae Henotesia perspicua (Trimen) X X 0 4d Ypthima asterope asterope (Klug) X X 0 4a Physcaeneura leda (Gerstaecker) X X 0 2g Charaxes varanes vologeses (Mabille) X X 0 1 Charaxes acuminatus Thurau X X X 5 Charaxes candiope candiope (Godart) X 0 X 1 Charaxes protoclea azota (Hewitson) X 0 0 2a; 3 Charaxes lasti lasti Grose-Smith X X 0 2g Charaxes jasius saturnus Butler X 0 0 ? Charaxes castor flavifasciatus Butler X 0 0 1 Charaxes brutus alcyone Stoneham X X X 3 Charaxes bohemani Felder and Felder X 0 0 4d Charaxes violetta maritima van Someren X X 0 2g Charaxes cithaeron kennethi Poulton X X 0 4d Charaxes pythodoris nesaea Grose-Smith X X 0 2a Charaxes etesipe tavetensis Rothschild X 0 0 2a; 3 Charaxes jahlusa kenyensis Joicey and Talbot X 0 0 4d Charaxes ethalion littoralis van Someren X 0 0 4d Charaxes viola picta van Someren and Jackson X 0 0 4b Charaxes contrarius van Someren X X 0 2g Charaxes guderiana rabaiensis Poulton X 0 0 4d Charaxes pleione oriens Plantrou X 0 0 2a Charaxes zoolina zoolina (Westwood) X 0 0 4d Euxanthe wakefieldi (Ward) X X X 2g Euxanthe tiberius tiberius Grose-Smith X X 0 2g Euriphene achlys (Hopffer) X X 0 2g Bebearia chriemhilda (Staudinger) X 0 0 2g Bebearia orientis orientis (Karsch) X X X 3 Euphaedra neophron littoralis Talbot X X X 2g Euphaedra orientalis Rothschild X 0 0 2g Hamanumida daedalus (Fabricius) X 0 0 4a Aterica galene theophane Hopffer X X 0 2a Harma theobene blassi (Weymer) X 0 0 2a Cymothoe coranus Grose-Smith X X X 2g Euptera pluto kinugnana (Grose-Smith) X X X 2g Pseudathyma lucretioides lucretioides X 0 0 2g Pseudacraea eurytus conradti Oberthür X X 0 2a Pseudacraea lucretia expansa (Butler) X X X 2a Pseudacraea boisduvalii trimeni Butler X X 0 ? Neptis saclava marpessa Hopffer X 0 0 1 Neptis kiriakoffi Overlaet X 0 0 2e Neptis alta Overlaet X 0 0 ? Neptis rogersi Eltringham X 0 0 2g Neptis trigonophora trigonophora Butler X 0 0 2g Neptis goochi Trimen X X 0 2g Cyrestis camillus sublineata Lathy X 0 0 2a Sallya garega (Karsch) X 0 0 ? Sallya amulia rosa (Hewitson) 0 0 X ? Byblia ilithyia (Drury) X 0 0 4a Byblia anvatara acheloia (Wallengren) X X 0 4a Neptidopsis fulgurata platyptera Rothschild and Jordan X 0 0 2g Eurytela dryope angulata Aurivillius X X X 1 Hypolimnas misippus (Linnaeus) X X X 1; 4a Hypolimnas deceptor deceptor (Trimen) X 0 0 2g Hypolimnas anthedon wahlbergi (Wallengren) X 0 0 2a Hypolimnas usambara (Ward) X X X 2g Salamis parhassus (Drury) X X X 2a Salamis anacardii (Linnaeus) X X 0 2a; 3 Salamis cacta amaniensis Vosseler X X 0 ? Junonia orithya madagascariensis Guenée X 0 0 4a Junonia oenone oenone (Linnaeus) X X X 1 Junonia hierta cebrene Trimen X 0 0 4a Junonia natalica natalica (Felder and Felder) X X 0 3; 4d Junonia terea elgiva Hewitson X X 0 2a; 3

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Superfamily Family Butterfly Species and Subspecies SH KM KK Category Papilionoidea Nymphalidae Junonia antilope (Feisthamel) X 0 0 4d Catacroptera cloanthe cloanthe (Stoll) X 0 0 4a Cynthia cardui (Linnaeus) X 0 0 4a Lachnoptera ayresii Trimen X X 0 2g Phalanta phalantha aethiopica (Rothschild and Jordan) 0 X X 1 Phalanta eurytis eurytis (Doubleday) X 0 X 2a Acraea aubyni Eltringham X X 0 2g Acraea encedon encedon (Linnaeus) X 0 0 1; 4a Acraea esebria esebria Hewitson X X 0 3; 4d Acraea eponina (Cramer) X X 0 1 Acraea petraea Boisduval X 0 0 2g Acraea egina areca Mabille X 0 0 4a Acraea braesia Godman X X 0 4c Acraea equatorialis anaemia Eltringham X 0 0 4c Acraea oncaea Hopffer X 0 0 4d Acraea pudorella pudorella Aurivillius X 0 0 4d Acraea natalica Boisduval X 0 0 3; 4d Acraea zonata Hewitson X X 0 2g; 4d Acraea rabbaiae mombasae Grose-Smith X X 0 2g Acraea satis Ward X X X 2g Acraea zetes acara Hewitson X 0 0 3; 4a Acraea chilo chilo Godman X 0 0 4c Acraea anemosa Hewitson X 0 0 3; 4d Acraea boopis ama Pierre X X 0 2g Acraea quirina rosa Eltringham X X 0 2a Acraea cuva cuva Grose-Smith X 0 0 2g Acraea insignis insignis Distant X 0 0 4d Acraea neobule neobule Doubleday X 0 0 1; 4a Acraea matuapa Grose-Smith X X 0 2g Acraea adrasta adrasta Weymer X 0 0 2g Acraea aganice Hewitson X X 0 3; 4d Acraea epaea epitellus Staudinger X 0 0 2a Pardopsis punctatissima (Boisduval) X X 0 ? Subtotal 105 55 26 Hesperioidea Hesperiidae Coeliades libeon (Druce) X 0 0 2a Coeliades anchises anchises (Gerstaecker) X X 0 4d Coeliades forestan forestan (Stoll) X 0 0 1 Coeliades pisistratus (Fabricius) X 0 0 1 Coeliades sejuncta (Mabille and Vuillot) X 0 0 2g Celaenorrhinus galenus (Fabricius) X X 0 2a Tagiades flesus (Fabricius) X X 0 2a Eagris sabadius ochreana Lathy X 0 0 4d Eagris nottoana nottoana (Wallengren) X 0 X 2g Sarangesa motozi (Wallengren) X 0 X 4d Sarangesa maculata (Mabille) X 0 0 2e Caprona pillaana Wallengren X 0 0 4d Netrobalane canopus (Trimen) X 0 0 4d Abantis paradisea (Butler) X 0 0 4d Spialia kituina (Karsch) X 0 0 4c; 9 Spialia spio (Linnaeus) 0 X 0 7 Spialia diomus diomus (Hopffer) X 0 0 4a Spialia confusa obscura Evans X 0 0 4d Spialia dromus (Plötz) X 0 0 4a Spialia zebra bifida (Higgins) X 0 0 4c Gomalia elma elma (Trimen) X 0 0 1 Astictopterus stellata stellata (Mabille) X X 0 2g Ampittia capenas capenas (Hewitson) X 0 0 4d Gorgyra subflavidus Holland X 0 0 2g Gorgyra diva Evans X 0 0 ? Gorgyra johnstoni (Butler) X 0 0 2g Pardaleodes incerta (Snellen) X X X 2a Teniorhinus herilus (Hopffer) X X 0 2g Acada biseriata (Mabille) X 0 0 4d Parosmodes morantii morantii (Trimen) 0 X 0 ?

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Superfamily Family Butterfly Species and Subspecies SH KM KK Category Hesperioidea Hesperiidae Paracleros biguttulus (Mabille) X 0 0 2a Acleros ploetzi Mabille X 0 0 2a Acleros mackenii (Trimen) X 0 0 2a Sernalea arela (Mabille) X 0 0 2a Andronymus neander neander (Plötz) X 0 0 ? Andronymus caesar philander (Hopffer) X X 0 2a Zophopetes nobilior (Holland) 0 X 0 ? Artitropa reducta Aurivillius X 0 0 ? Artitropa erinnys radiata Riley X 0 0 4d Fresna nyassae (Hewitson) X 0 0 2b Pelopidas thrax inconspicua (Bertoloni) 0 X 0 1 Borbo fatuellus fatuellus (Hopffer) X 0 0 2a Borbo lugens (Hopffer) X X 0 4d Borbo detecta (Trimen) 0 X 0 4d Borbo ferruginea ferruginea (Aurivillius) X 0 0 2g Borbo borbonica borbonica (Boisduval) X 0 0 4a Subtotal 39 13 3 TOTAL 246 112 45

September 2000 METAMORPHOSIS Occasional Supplement 4 47 ______

APPENDIX III

Species list of larger moths recorded in the study areas of Kaya Muhaka (KM) c. 30 ha and Kaya Kinondo (KK) c. 7 ha. Nomenclature follows those of the collections in the BMNH (London) as well as Carcasson (1976), Fletcher & Nye (1982), Poole (1989), Goodger & Watson (1995), Häuser & Boppré (1997) and Scoble (1999).

Superfamily Family Larger Moth Species KM KK Cossoidea Limacodidae Micraphe haematoessa (Hampson) X 0 Metarbelidae Metarbela ?dialeuca Hampson X 0 Metarbela haberlandorum Lehmann X 0 Salagena ?irrorata Le Cerf X 0 ? Salagena sp. nov. X 0 Subtotal 5 0 Geometroidea Epiplemidae Epiplema barbara Warren X 0 Epiplema dohertyi (Warren) [Leucoplema] X X Geometridae Allochrostes biornata Prout X 0 Antharmostes papilio papilio Prout X 0 Chiasmia feraliata (Guenée) 0 X Cleora munda (Warren) X 0 Colocleora divisaria divisaria (Walker) X 0 Comibaena esmeralda (Warren) X 0 Comibaena rufitornus Prout X 0 Erastria albosignata albosignata (Walker) X 0 Ereunetea reussi reussi Gaede X 0 Isoplenia trisinuata Warren X X Isturgia catalaunaria (Guenée) 0 X Isturgia supergressa (Prout) 0 X Metallochlora grisea Prout 0 X Mixocera viridans Prout X 0 Omizodes rubrifasciata (Butler) X X Paraptychodes tenuis Butler X X Pingasa rhadamaria alterata (Walker) 0 X Pitthea trifasciata Dewitz X 0 Racotis apodosima Prout X 0 Racotis breijeri (Prout) 0 X Racotis squalida squalida (Butler) X 0 Scopula addictaria (Walker) X 0 Scopula atricapilla atricapilla Prout X 0 Scopula cassioides Prout X 0 Scopula donovani (Distant) 0 X Scopula lactaria (Walker) X 0 Scopula minorata minorata (Boisduval) X 0 Scopula ?ossicolor Warren X 0 Traminda neptunaria (Guenée) X X Traminda obversata obversata (Walker) 0 X Traminda vividaria (Walker) X 0 Zamarada euphrosyne Oberthür X 0 Zamarada sp. nov. X 0 Zamarada ?plana denticincta Hampson X 0 Zamarada rufilinearia Swinhoe X 0 Subtotal 29 12 Bombycoidea Saturniidae Decachorda aspersa Bouvier [? subsp.] X 0 Gonimbrasia zambesina (Walker) [=Imbrasia] X 0 Lasiocampidae Beralade continua Aurivillius X 0 Subtotal 3 0 Sphingoidea Sphingidae Acherontia atropos (Linnaeus) X 0 Centroctena imitans (Butler) X 0 Neopolyptychus compar septentrionalis Carcasson X 0 Praedora marshalli tropicalis Rothschild and Jordan X 0 Temnora marginata (Walker) X X Subtotal 5 1 Noctuoidea Arctiidae Amerila phaedra Weymer X X Amphicallia bellatrix (Dalman) 0 X Argina amanda (Boisduval) 0 X Argina astrea (Drury) 0 X

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Superfamily Family Larger Moth Species KM KK Noctuoidea Arctiidae Asura anticraspeda Hampson X X Asura ichorina Butler X 0 Eilema distigmata Hampson X 0 Eyralpenus melanocera (Hampson) X 0 ? Metarctia rufescens Walker X X ? Nola chionea Hampson X 0 Nyctemera restrictum (Butler) X 0 Secusio drucei Rothschild X 0 Ctenuchidae Syntomis phoenica Hampson X 0 Lymantriidae Cropera testacea Walker X 0 Dasychira callipepla Collenette X 0 Dasychira extorta (Distant)[= Laelia] X X Euproctis rufopunctata (Walker) X 0 Euproctis torrida Distant X 0 Hemerophanes xanthopa Collenette X 0 Leucoma parva (Plötz) X 0 Lymantria leucerythra Collenette X 0 Marblepsis macrocera Sharpe X 0 Stracena bananae (Butler) X 0 Notodontidae Eurystaura griseitincta Hampson X 0 Paracleapa psecas Druce X 0 ? Peratodonta extensa Gaede X 0 Noctuidae Aburina sobrina Möschler X 0 Acantholipes trimeni Felder and Rogenhofer X X Achaea catella Guenée 0 X Achaea dasybasis Hampson X 0 Achaea lienardi (Boisduval) X X Alelimma pallicostalis Hampson X X Amyna octo (Guenée) 0 X Amyna punctum (Fabricius) X X Anomis flava (Fabricius) X X Anomis leona (Schaus [ & Clements]) X 0 Anomis punctulata (Holland) 0 X Anomis sabulifera (Guenée) X X Anomis simulatrix (Walker) X X Anua gonoptera (Hampson) [Ophiusa] X 0 ? Anua sp. nov. X 0 Asota speciosa (Drury) X X Attonda adspersa (Felder and Rogenhofer) X X ? Baniana sp. nov. X 0 Blenina quadripuncta Hampson X X Caligatus angasii Wing X 0 Caryonopera breviramia Hampson X 0 Cyligramma limacina (Guérin-Méneville) X X Digama africana Swinhoe X 0 Diparopsis castanea Hampson X 0 Dysgonia conjunctura (Walker) X 0 Enispa flavitincta Hampson X 0 Entomogramma pardus Guenée X 0 Ercheia subsignata (Walker) X X Erebus walkeri (Butler) X X Eudocima divitiosa (Walker) X 0 Eudocima materna (Linnaeus) X 0 Eutelia amatrix Walker X 0 Eutelia musicalis Berio X 0 Facidia vacillans (Walker) X X Facidina semifimbria (Walker) 0 X Fodina embolophora Hampson X 0 Geniascota patagiata Hampson X 0 Giria pectinicornis (Bethune-Baker) X 0 Gracilodes caffra Guenée X X Gracilodes nysa Guenée X 0

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Superfamily Family Larger Moth Species KM KK Noctuoidea Noctuidae Grammodes congesta Berio X 0 Heliophisma xanthoptera (Hampson) X 0 Hypena glyptalis Mabille X 0 Hypena masurialis Guenée X 0 Hypena varialis Walker X 0 Lacera alope (Cramer) 0 X Lamprolopha melanephra Hampson X 0 Lophocrama phoenicochlora Hampson X 0 Lophoptera litigiosa (Boisduval) X 0 Lophotavia globulipes (Walker) X X Marathyssa cuneala (Saalmüller) X 0 Marcipa insulata (Walker) X 0 Marcipa mediana Hampson X 0 Marcipa pyramidalis (Hampson) X X Maxera marchalii Boisduval X 0 Mazuca strigicincta Walker X 0 Metaleptina nigribasis Holland X 0 Mocis conveniens (Walker) X X Mocis mayeri (Boisduval) X 0 Oglasa nana (Walker) 0 X Oraesia emarginata (Fabricius) X 0 Ozarba perplexa Saalmüller 0 X Plecoptera aspila (Hampson) X X Plecoptera flavilinea Hampson 0 X Plecoptera rufirena (Hampson) 0 X Plusiodonta commoda Walker X 0 Pseudogiria polita Berio X 0 Pteronycta fasciata Fawcett X 0 Radara subcupralis (Walker) X 0 Rhanidophora agrippa Druce X 0 Rhanidophora cinctigutta (Walker) X 0 Rhesala moestalis (Walker) 0 X Rhynchina ?leucodonta Hampson X 0 Rhynchina revolutalis (Zeller) X 0 Rhynchina sp. 0 X Rougeotiana xanthoperas (Hampson) X 0 Serrodes partita (Fabricius) X 0 Simplicia extinctalis (Zeller) X 0 Soloe tripunctata Druce X X Spodoptera exempta (Walker) 0 X Spodoptera littoralis (Boisduval) X 0 Tatorinia fumipennis (Felder and Rogenhofer) X 0 Tavia instruens Walker 0 X Tavia nycterina (Boisduval) X X Thyas arcifera (Hampson) 0 X Thyatirina achatina (Weymer) X 0 Trichopalpina sp. X 0 Trigonodes hyppasia (Cramer) 0 X Ugia amaponda (Felder and Rogenhofer) X X ? Ugia sp. nov. 0 X Subtotal 97 45 TOTAL 139 58

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Figure 5. Forest edge of Kaya Muhaka, NE part, February 1994. In the foreground, note Hyphaene compressa grassland; in the background there are trees of the discontinuous B and C strata. The smaller tree in the centre is Dichrostachys cinerea, the grassland left from it is invaded by Lantana camara thicket in stratum E.

Figure 6. View of canopy of Kaya Kinondo, NE part, from c. 7 m on an elevation, February 2000. Note the very open B stratum and two deciduous trees bare of leaves. Several trees look like emergents but stratum A is absent.

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Figure 7. A male of Charaxes lasti ssp. lasti recovers after being marked in Kaya Muhaka, June 1997. Note the two markings with red nail polish in the apex of the wings. The subspecies is common in Kaya Muhaka and is endemic to Kenya and northern Tanzania coastal areas inland to the Usambara Mountains.

Figure 8. A female of Aterica galene ssp. theophane being attracted to a fresh orange put on the ground in Kaya Muhaka, February 1998. The subspecies flies close to the forest ground and was rarely attracted to our bait-traps (Photographs by Ingo Lehmann).

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