Journal of East African Natural History 94(1): 121–163 (2005) LEPIDOPTERA DIVERSITY, FLORISTIC COMPOSITION AND STRUCTURE OF THREE KAYA FORESTS ON THE SOUTH COAST OF KENYA Ingo Lehmann Turnerweg 09, 23970 Wismar, Germany ingo.lehmann@freenet.de Esther Kioko Department of Invertebrate Zoology, National Museums of Kenya P.O. Box 40658, Nairobi, Kenya ekioko@icipe.org ABSTRACT Three isolated lowland coastal forest patches in Kwale District, namely Kaya Muhaka, Kaya Kinondo and Kaya Diani are classified here as “Wetter mixed semi-deciduous forest”, “Groundwater forest on coral rag” and “Maritime scrub forest”. Although they are sacred to the Digo people, different rates of disturbance were assessed. Kaya Kinondo, which represents a rare forest type along the Kenya coast, is undisturbed at least since the authors began their studies in 1994. Floristic diversity and endemism are high in all Kayas. Lepidoptera diversity is low in Kaya Kinondo, showing that an undisturbed forest does not automatically have a rich Lepidoptera fauna and that the latter does not always respond to a diverse flora. With 352 species, Lepidoptera diversity and endemism is high in Kaya Muhaka. This includes species with a western and central Africa distribution, as well as the Kenyan endemic montane subspecies Charaxes acuminatus shimbanus. Larger moths that appear to be endemic to coastal eastern Africa are presented and others have been preliminary classified as rare for coastal Kenya including species first recorded from Kenya. Two biogeographical groups of coastal forests were found among ca. 30% of the Kenyan butterfly fauna and the authors believe that a further sub-division of the “Usambara-Kwale local centre of endemism” is possible between coastal forests further inland (e.g. Kaya Muhaka) and those close to the shoreline of the Indian Ocean (e.g. Kaya Kinondo, Kaya Diani). INTRODUCTION The coastal forests of eastern Africa are one of the top ten priority ecosystems for biodiversity conservation on the African continent (Burgess, 2000), covering an area of ca. 3170 km2 from southern Somalia to northern Mozambique including small amounts of forest in south-eastern Malawi and eastern Zimbabwe (Burgess et al., 2000a). The coastal 122 I. Lehmann & E. Kioko forests 1 of Tanzania and Kenya are recognised as an area of global importance and have been earmarked as one of the 25 world’s hotspots of biodiversity due to the concentration of many narrowly endemic plant and animal species in exceptionally small areas (Myers et al., 2000). Estimates of closed-canopy Kenyan forests vary, depending on the definition. According to FAO’s estimate (1999) natural forest, plantations and bamboo occupy only 2.3% or 12 920 km2 of Kenya’s total available land. Due to a new global definition for “forest” FAO’s estimate (2001) is now 30.0% or 170 959.5 km2 and includes a large area of drier forests (mainly woodlands) in the Kenyan lowlands (Holmgren, FAO, pers. comm., 2004). Of it, the currently known 103 coastal forests cover an estimated total area of 787 km2 (Younge et al., 2002). Most of the coastal forests are smaller than 5 km2; the largest are Arabuko-Sokoke (370 km2) and the Shimba Hills forests (ca. 90 km2) (Wass, 1995). The heavily fragmented Kenyan coastal forests are very diverse (e.g. Robertson & Luke, 1993; Mlingwa et al., 2000) and are an important sub-centre of endemism within the “Swahilian regional centre of endemism”. Due to its high number of endemic plant species (>1000), Clarke (1998) classified the latter and modified the former “Zanzibar-Inhambane regional mosaic” as defined by White (1976, 1983). Subsequently he enlarged, divided and renamed it. The term “Swahilian region sensu lato” encompasses both this regional centre of endemism and the adjacent “Swahilian/Maputaland regional transition zone” to the south. Because of the particular geographic concentrations of vascular plant species endemism along the southern Kenyan coast between Mombasa and the Tanzanian border, Clarke et al. (2000) defined this area as the “Kwale local centre of endemism sensu White (1993)”. Burgess (2000) included this area in the “Usambara - Kwale local centre of endemism”. He defined a local centre of endemism as an area with >100 endemic species (not only plants). Many forest patches north of the Pangani River (Tanzania) once enclosed “Kaya or Kaya-like clearings” (Hawthorne, 1993). Based on the authors’ assessment of the data in Robertson & Luke (1993), 70 Kayas and Sacred Groves exist in Kenya, including 46 with an estimated total size of 31 km2. They are protected by tradition of the nine Mijikenda ethnic groups and up to date 39 have also been gazetted by the legal system as National Monuments (NM) since 1992. They are managed by the National Museums of Kenya (NMK) and their Coastal Forest Conservation Unit (CFCU) in collaboration with the Committee of Elders and the district administration. 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). There is debate about Spear’s static picture what Mijikenda society was, about his opinion that all the Kayas were established after the turn of the 17th century and that total village communities of the Mijikenda onced lived within the Kaya before they left all their Kayas between the 1830’s and 1870’s. Most probably the Kaya should be seen rather in the context of the power of old men with which it is clearly linked (Walsh, 1987; Willis, 1993). Knowledge and care of the Kayas are still in the hands of old men now. 1 The term “forest” will be applied here according to the new definition of the FAO (2001) for a stand of trees with a canopy cover of more than 10% that varies in height from 5 m or more. For the Kenyan coast the term “coastal forest” will be used as defined by Hawthorne (1993) which is all forest on “ ... the land over the sedimentary (and intrusive volcanic) rocks of the coastal plains and plateau, to the east of the exposed basement complex land.” Three kaya forests on the south coast of Kenya 123 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 ca. 56% are restricted to forests (de Jong & Congdon, 1993). Congdon & Collins (1998) mentioned “approximately 895” butterfly species for Kenya and listed 25 endemic species. The total number of forest-associated butterfly species found in the coastal forests of eastern Africa may reach ca. 400 including 75 species strictly restricted to these forests (excluding lowland forests of Malawi, eastern Zimbabwe and northeast South Africa). The highest numbers of butterfly species known from single coastal lowland forest sites in Kenya and Tanzania below 300 m above sea level include Arabuko-Sokoke with 320 species and three Tanzanian sites with 241 species from the Pugu Hills (10 km2), 127 species from Dendene forest at Kisiju (2 km2) and 129 species from Zaraninge forest at Kiono (20 km2) (Kielland & Cordeiro, 2000; forest size from Burgess & Clarke, 2000, in appendix 1). The total number of Macroheterocera occurring in the Afrotropical region and in Kenya respectively is unknown including information about the habitat for the majority of species. Hence, the aim of the authors was to undertake a first long-term study on the Lepidoptera fauna and its habitats in a sample of small Kenyan coastal forests. They also would like to show that this sample of tiny coastal forests should receive more attention because of its high conservation value. STUDY AREAS Location and size Kaya Muhaka (KM), also called Kaya Kambe or Kaya Mwadabara, is situated ca. 15 km southeast of the Shimba Hills (SH), close to Muhaka village, 32 km south of Mombasa. The forest is ca. 5.5 km inland from the Indian Ocean at 45 m (UTM 37MEF5623) and is one of the largest Kaya forests in Kwale District 2 with ca. 150 ha. The study area is in the more intact northeastern and central part representing ca. 30 ha. Kaya Kinondo (KK, plate 1e), also called Kaya Ngalaani, is situated ca. 7 km southeast of KM. It lies ca. 5 km north of Chale Point, ca. 100 m inland from the Indian Ocean at 5– 10 m (UTM 37MEF6015) and has a forest size of ca. 30 ha. The study area of ca. 7 ha lies in the central and southern part. Kaya Diani (KD) is situated ca. 7 km northeast of KM. It lies ca. 700 m north of the road from Diani Beach to Ukunda, ca. 500 m inland from the Indian Ocean at 15 m (UTM 37MEF6526) and has a forest size of ca. 20 ha. The study area of ca. 5 ha lies in the central and southeastern part. Recent human disturbance All three Kayas are isolated by „Zanzibar-Inhambane secondary grassland and wooded grassland“ and/or “Zanzibar-Inhambane evergreen and semi-evergreen bushland and thicket” sensu White (1983) and poorly utilised crop land. Fires were seen twice in the grassland near the western forest edge of KM in February 1994 and 2003. 2 The forest size of all three Kayas was estimated by Robertson & Luke (1993); all are gazetted as NM since 1992. 124 I. Lehmann & E. Kioko Kaya Muhaka lost at least five trees of 20–30 cm diameter at breast height (dbh), which were poached from the study area in 2001 (Scorodophloeus fischeri, Hymenaea verrucosa). Pole cutting did increase since 2001 and collecting of fuelwood is common. Kaya Kinondo was the second most important “main Kaya” after Kaya Kwale for the Digo people, who protect the forest up to date. Between 1994–2003 there was no illegal logging, pole cutting or collecting of fuelwood. The Digo use the Kaya again since 1999 for ceremonies near a glade where several graves are located, honoured as a sacred place (Mzee Mnyenze, pers. comm., 2000). It is probably the same place which was used for a large meeting in 1983, mentioned by Robertson (1984). She also mentioned stands of “the superb timber tree Calophyllum inophyllum” which were found by the authors still intact in 1994. The authors conclude that no tree poaching occurred in the forest interior for at least 20 years. Since 2001 the „Kaya Kinondo Pilot Ecotourism Project“ was implemented by the NMK, World Wide Fund for Nature, the Kaya Kinondo community and the private sector. The construction of a thatched hut resulted in some clearance of natural regeneration and treelets e.g. Antiaris toxicaria, Mallotus oppositifolius on the transect figured by Lehmann & Kioko (2000). Pole cutting is very common in KD and two trees of Lecaniodiscus fraxinifolius with >15 cm dbh were cut in the sample area in January 2003. Dead standing stems or dead lying wood is almost absent due to extensive collecting of fuelwood. Temperature, rainfall and relative air humidity Michieka et al. (1978) mentioned 26.1°C for coastal southeast Kenya. The air temperatures vary during the hotter dry season (December–March) and cooler transitional season (July–mid September). Rainfall figures appear to be different between KM and KK/KD according to Jätzold & Schmidt (1983): KM has an average annual rainfall of 1129 mm (23 years of records) with 132 mm during December–March (February is the driest month), 568 mm during the long rains April–June, 172 mm in July/August and 257 mm during the short rains from September–November. Rainfall data for KK/KD appear to be uncertain, showing a 60% reliability of rainfall in 6 out of 10 years of 600–700 mm from March–September and less than 100 mm in November and 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 air humidity is also high throughout the dry season. White (1983) concluded that no month is absolutely dry. Physiography and soils The three Kayas are located on the coastal plain. This is of Pleistocene age, flat to gently undulating, not more than 50 m 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 Kinondo and KD are 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 ferralitic Cambisols, lithic phase). 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) (cf. Michieka et al., 1978). Three kaya forests on the south coast of Kenya 125 MATERIAL AND METHODS Duration of research Fieldwork was done daily for seven hours in the dry season as well as in the rainy season over a total period of 32 weeks in KM/KK (1994, 1996–1998, 2001–2003) and 14 weeks in KD during the dry season (2001–2003); including February 1994 (two weeks); January/ February 1996 (five weeks); June/July 1997 (five weeks); January/February 1998 (five weeks); January/February 2001 (five weeks); June 2001 (one week); February/March 2002 (five weeks) and January/February 2003 (four weeks). Sampling sites for description of forest structure, dominance and floristic composition In each study area, permanent marked “sample sites” were established in 1994 (in KD in 2001). Each sample site is 25 x 25 m or 625 m2 (0.0625 ha) in size. Sample sites were chosen subjectively (e.g. high forest, gaps). Ten sample sites or 6250 m2 (0.625 ha) were established on ca. 30 ha in KM, five sample sites or 0.3125 ha on ca. 7 ha in KK and five sample sites on ca. 5 ha in KD. The sum of all sample sites “per Kaya” (e.g. 0.625 ha) is defined as “sample area”. The term “stratum” will be applied here for a layer or set of tree crowns and shrubs between certain 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). Trees of strata A and B will be termed “overstorey trees”, of the C stratum “understorey trees”. At least one transect of 450 m2 each (75 x 6 m) was marked out in the central part of each Kaya to construct a profile diagram. All plant specimens >5 m tall and rooting on the sample site were determined, numbered and figured by drawing in their position of stem and approximate shape of crown. Specimens of ≥ 3 cm dbh were measured in regard to total height, lower limit and width of crown, dbh and diameter at base of trunk (dab). Plants <3 cm dbh were measured on three sample sites per Kaya (including one gap) and on all transects. Heights >2.5 m were measured with a Suunto clinometer. The dominant species was determined for each sample site and for each stratum. The “most dominant species” was determined for the sample area. Dominance for trees is usually defined as basal area (Mueller-Dombois & Ellenberg, 1974). The basal area is π/4 (d2) and was computed for each specimen ≥3 cm dbh including dead standing stems. Dead trees as well as dead wood on the ground (length ≥0.5 m and midpoint (mp) of length at least 5 cm in diameter) were measured. Their volume was computed (π/4 x mp2 x l). Lepidoptera Macroheterocera (termed here “larger moths”) and butterflies were recorded in the study areas by netting, light-trapping and bait-trapping. The latter was not possible in KD due to vervet monkeys (Chlorocebus aethiops (L.)), which ate all fruits from the traps. A portable ultra-violet light-trap with a 15 W lamp was used for light-trapping at the forest edge and a mercury-vapour light-trap with a 125 W lamp in the forest interior. Light-trapping in KD was not undertaken. Collected specimens were killed with potassium cyanide, pinned, prepared, labelled and assimilated into the collection of the NMK. Species identification Plants were identified in the field, difficult specimens were collected and later identified by Quentin Luke. Literature used for identification and nomenclature was Haines & Lye (1983), Johns (1991), Flora of Tropical East Africa and Beentje (1994). 126 I. Lehmann & E. Kioko Literature used for identification and nomenclature of butterflies was Henning (1989), Kielland (1990), Ackery et al. (1995), Larsen (1996) and Congdon & Collins (1998). The identification of larger moths was done at The Natural History Museum (BMNH, London). The collection at the NMK (Nairobi) was used for comparison. Nomenclature follows the BMNH, Carcasson (1976), Fletcher & Nye (1982), Poole (1989), Goodger & Watson (1995), Häuser & Boppré (1997), Scoble (1999) and Vári et al. (2002). RESULTS Classification of forest types Kaya Muhaka is a “Mixed dry forest” following Clarke & Robertson (2000) but will be classified here as a “Wetter mixed semi-deciduous forest” because: firstly, White (1983) defined a “Dry forest” as a forest that experiences a dry season lasting several months during which atmospheric humidity is low. White’s definition was accepted by Clarke & Robertson (2000) but this is problematic since e.g. Schmidt (1991) recorded 40% relative air humidity in the driest month in the Shimba Hills forests, near KM. Hence, most probably the relative air humidity is also high in KM during the driest month. Secondly, KM is locally dominated by caesalpiniaceous trees such as Cynometra and Scorodophloeus in strata A and B, which is similar to West and Central African forests described as “Wet evergreen forests” by Hamilton (1989). Thirdly, Synsepalum brevipes, the most dominant species of stratum C, is linked to wetter forests of the Guineo-Congolian region after Hawthorne (1984, 1993). Fourthly, two genera and at least three species of Lepidoptera are mainly linked to wetter forests of West and Central Africa: Euptera (a rain forest genus), Euriphene (a genus centred on the rainforest zone), Anthene liodes (often common in the main equatorial rainforest zone, Larsen, 1996) and the larger moths Pseudogiria polita, Rougeotiana xanthoperas (own studies). Kaya Kinondo will be classified as a “Groundwater forest on coral rag” because: firstly, the Moraceae family, common in groundwater forest (Clarke & Robertson, 2000), is dominant in strata A and B with large trees of Antiaris toxicaria and Milicia excelsa. Secondly, Sorindeia madagascariensis, a species which indicates a good supply with groundwater (e.g. Andrews et al., 1975), is most dominant in stratum B and locally dominant in stratum C. Thirdly, Ficus species are locally common in strata B and C such as Ficus scassellatii, a species of riverine or groundwater forest (Beentje, 1994). Fourthly, the paucity of the legumes (Caesalpiniaceae, Mimosaceae, Papilionaceae) is remarkable (Lehmann & Kioko, 2000) and is according to Lowe & Clarke (2000) typical of undisturbed forests in groundwater sites (which supports the authors opinion that KK is largely undisturbed, cf. forest structural types). Noteworthy are patches of the “Mixed scrub forest” (Clarke & Robertson, 2000) where Combretum schumannii is dominant and where two trees of the rare Vitex zanzibarensis occur. Kaya Diani will be classified as a “Maritime scrub forest” (Clarke & Robertson, 2000) since its canopy has typically little vertical stratification (stratum A largely absent, B patchy, C has largest crowns, only D is more or less continuous), is 4–10 m tall, with few emergents up to 23 m and many species that are recorded as dominant in this vegetation type (listed by Burgess & Clarke, 2000, in appendix 2) were found to be dominant or most dominant in the sample area e.g. Adansonia digitata, Grewia plagiophylla, Haplocoelum inoploeum. The authors observed that the driest month is most severe in KD where many shrubs and treelets Three kaya forests on the south coast of Kenya 127 appear quite limp and fully green leaves hang down as if wilted, which is noteworthy because mean annual temperature, rainfall and soils are similar in KK. Forest structural types and assessment of logging Three forest structural types have been identified based on four structural parameters (table 1): 1. Very high total mean dbh, high mean tree height and basal area, low stem density (KM). 2. Very high total basal area, high stem density, intermediate mean dbh/tree height (KK). 3. Low total mean tree height, basal area and stem density, intermediate mean dbh (KD). Table 1. Comparison of strata A-D for trees of ≥ 10 cm dbh; KM (1), KK (2), KD (3). Strata Kaya A 1 2 3 1 2 3 1 2 3 1 2 3 1 2 3 B C D Total Mean dbh (m) 0.633 0.783 0.320 0.469 0.365 0.624 0.226 0.207 0.257 0.110 0.103 0.125 0.426 0.305 0.329 Mean tree height (m) 27.9 27.9 22.3 17.8 17.5 17.1 9.1 9.6 8.4 3.4 3.5 3.6 17.6 13.6 10.0 Living individuals ( ≥ 10 cm dbh.ha-1) 61 32 3 54 118 51 67 214 119 5 6 32 187 370 205 Basal area (m2.ha-1 ) 21.9 18.2 0.3 10.4 14.2 18.6 3.5 9.2 8.2 0.04 0.05 0.4 35.8 41.6 27.5 A comparison with seven vegetation structural groupings presented by Lowe & Clarke (2000) for Tanzanian coastal forests was done to assess the degree of logging; KM was found to be closest to group 7, KK to group 4 and KD to group 5: 1. Kaya Muhaka has very tall and thick trees. Two parameters are similar to more or less undisturbed coastal dry and moist forests but all have at least 250 stems.ha-1 and 39 m2.ha1 . 2. Kaya Kinondo has tall and thick trees. Two parameters are similar to coastal moist and dry forests including one disturbed groundwater forest but all have a total basal area of <32 m2.ha-1 (the groundwater forest has only 21.3 m2.ha-1) and at least 440 stems.ha-1. 3. Kaya Diani has short and thick trees. Two parameters are similar to undisturbed coastal dry forests, partly on coral rag too, but all have at least 340 stems.ha-1 and a higher total basal area in those forests where Adansonia digitata also occurs, including one where the latter is dominant having 43.7 m2.ha-1. From the comparison it becomes clear that logging reduces the basal area and stem density. The authors conclude that KK is largely untouched because of its very high total basal area and high mean stem density for a forest on an extremely rocky site (cf. figure 1). The two other Kayas are largely moderately logged (cf. figure 2 for KD). Kaya Muhaka has a few patches in the central part, which appear to be only lightly logged having at least 16 stems of ≥10 cm dbh on 625 m2 (cf. profile diagram of KM in Lehmann & Kioko, 2000). 128 I. Lehmann & E. Kioko These conclusions are consistent with figures for stem density and basal area in Kibale forest (Uganda) represented and discussed by Struhsaker (1997). Key: 4 Sorindeia madagascariensis; 10 Drypetes natalensis; 11 Lecaniodiscus fraxinifolius; 20 Antiaris toxicaria; 35 Asteranthe asterias; 41 Uvariodendron kirkii; 45 Diphasia sp. A of FTEA; 48 Diospyros ferrea; 49 Mallotus oppositifolius; 78 Garcinia livingstonei; 90 Rinorea ilicifolia; 96 Cola minor; 107 Encephalartos hildebrandtii; 110 Hunteria zeylanica; 112 Lasiodiscus pervillei; 120 Blighia unijugata; 148 Lannea welwitschii; 173 Salacia elegans; 176 Coffea pseudozanguebariae; Dt.=Standing dead stem; Dw.=Dead wood; U.=Unidentified. Figure 1. Profile diagram and crown projections of a groundwater forest on coral rag (KK, 2003). The strip of forest is 75 m long and 6 m wide, showing five discontinuous strata, stratum E is almost absent. The openness of all strata is due to the cropping out of coral (densely stippled). It is likely that the open forest on pure coral (left and centre) is very close to primary vegetation since the scarcity of strong light-demanding pioneer species in all strata, the occurrence of one of the tallest specimens in KK of the slow growing Encephalartos hildebrandtii (total height 4.3 m, trunk height 2.1 m), the absence of cut stumps as well as many pieces of dead wood on the ground partly >3 m in length and covered with mosses, a rare feature in all three Kayas, are outstanding. The crown projections below show also large tree species which are growing on pure coral (e.g. Diphasia sp. A, Diospyros ferrea). Dead standing trees, annual natural tree mortality and dead wood on ground Dead standing trees can be seen everywhere in undisturbed forests and can supply together with dead wood on the ground a considerable part of diet e.g. for caterpillars of some larger moth species. In the sample areas two of four dead standing stems of ≥10 cm dbh fell down in KM but were replaced by three new ones in 1999; two of three stems in KK fell down but were replaced by two new ones in 1999/2001 (records 1994–2003). The total number of dead standing stems per ha increased in KM from 6 to 8 after 1996 but remained constant with 10 stems in KK. In KD, they are generally rare which is a characteristic structural feature. None was recorded on 0.3125 ha in 2001–2003. Table 2 shows the number of dead standing stems and their percentage of the total basal area over a Three kaya forests on the south coast of Kenya 129 period of five years with the highest value in KK in 2003. The rather low mean height suggests several windbreaks of trees of strata A and B. Key: 6 Fernandoa magnifica; 11 Lecaniodiscus fraxinifolius; 35 Asteranthe asterias; 43 Adansonia digitata; 46 Trichilia emetica; 49 Mallotus oppositifolius; 110 Hunteria zeylanica; 111 Ludia mauritiana; 113 Erythroxylum emarginatum; 122 Suregada zanzibariensis; 129 Maytenus undata; 133 Monanthotaxis fornicata; 148 Lannea welwitschii; 149 Haplocoelum inoploeum; 152 Balanites maughamii; 154 Vepris eugeniifolia; 157 Millettia usaramensis; 166 Allophylus rubifolius; 169 Ochna thomasiana; 172 Uvaria acuminata; 174 Flacourtia indica; 176 Coffea pseudozanguebariae; 189 Cordia monoica; 190 ?Salacia stuhlmanniana; 195 Cussonia zimmermannii; 201 Acacia adenocalyx; 202 Grewia vaughanii; 203 ?Deinbollia borbonica; 204 Cissus sp. ; U. =Unidentified. Figure 2. Profile diagram and crown projections of a maritime scrub forest on coral rag (KD, 2003). The strip of forest shows four strata (A is absent, B is represented by a single specimen of Adansonia digitata). Strata C and D are more or less continuous; stratum E is locally absent. Several signs of disturbance are visible like the occurrence of species which are more common on the coastal plain outside of forests (e.g. Adansonia digitata, Millettia usaramensis), species in strata C/D which are common in forest edges or depleted coastal forests (e.g. Cussonia zimmermannii, Trichilia emetica) and denser vegetation including cut stumps (left). Dead wood does not occur. The patch right is the habitat of Deudorix diocles, an uncommon butterfly in KD. The crown projections below show a clumped dispersion of trees in the centre including several specimens of Trichilia emetica. Table 2. Basal area of dead standing stems of ≥10 cm dbh on 0.625 ha in KM (1) and on 0.3125 ha in KK (2) in January 1998 (a) and January 2003 (b). Kaya 1 (a) 1 (b) 2 (a) 2 (b) Number of dead standing stems 4 5 3 3 Mean height (m) 5.3 4.8 11.6 8.0 Mean dbh (cm) 51.0 43.6 32.7 61.8 Basal area (m2) 0.93 0.93 0.25 1.0 Percent of total basal area 3.9 3.9 1.9 7.2 130 I. Lehmann & E. Kioko The average annual mortality in the sample areas for trees of ≥10 cm dbh was 0.6% in KM, 0.7% in KK over ten years and 0% in KD during 2001–2003 (sample size: 117 trees in KM; 116 in KK; 64 in KD). This is below the value of Gentry & Terborgh (1990) who found an average annual tree mortality rate of 1–2% for a wide range of tropical forests. Although the sample size is a small fraction of the whole tree population it shows that the low stem densities in KM/KD are not due to a natural tree mortality. Noteworthy is that in KM large branches often break off adult Julbernardia magnistipulata (dominant in strata AD) but that a dead tree of this species was not seen since 1994 and that the only baobab in KK (dbh 1.34 m in 1998; 1.40 m in 2002) was older than 50 years (Mzee Mnyenze, pers. comm., 2000) but less than 100 years 3 before it fell in 2003 (table 3). Table 3. Natural annual mortality for trees of ≥10 cm dbh in the sample areas of KM (1) and KK (2) over a period of ten years (1994–2003). Tree species Kaya 1 Total 2 Antiaris toxicaria ? Antiaris toxicaria Lecaniodiscus fraxinifolius Scorodophloeus fischeri Sorindeia madagascariensis Adansonia digitata (baobab) Diphasia sp. A of FTEA Drypetes natalensis Stadmania oppositifolia Total Number of stems 1 1 1 4 Percent of total number per year 7 2 0.6 1 1 3 1 8 Notes windbreak 1994 windthrow 2003 windthrow 1997 standing dead stems of 1994/1999 standing dead stems of 1996/1999 windthrow 2003 windthrow 2000 windthrow 1997 windbreak 2001 0.7 Firewood collecting was found to be absent in KK where the total volume of dead wood per hectare lying on the ground is twice as much as in KM. A significant decrease in the volume of dead wood lying on the ground over a period of five years (1998–2003) was found in KM. This decrease is certainly due to more firewood collecting since 2001, the year when also tree poaching increased. In KK, many of the pieces of 1998 were still on the sample sites in 2003. In contrast, KD has almost no dead wood on the ground. Only 17 pieces or 0.86 m3 were recorded on 0.3125 ha in 2003, but none which was several years old. The authors assume that one cubic metre of dry dead wood weighs about 600 kg. An estimate from the coastal lowland forests of East Africa is that one person needs between 332–572 kg of firewood per year where dead wood makes up the bulk of the firewood gathered (Cambridge-Tanzania Rain Forest Project 1994 cited by Burgess et al., 2000b). This is between 0.5 and 1 m3. If one further assumes that a family consists of five members, they will need between 2,5 and 5 m3 of firewood per year. Hence, one hectare of forest in the study area of KM had firewood for 8–16 families in 1998 but for only 2–4 families five years 3 The age was estimated according to Wickens (1982) who stated that the trunk of the baobab attaining 4.5 m in diameter after 100 years and that the branches of young trees 30-40 years old are more erect; both was not observed in the specimen of KK. Three kaya forests on the south coast of Kenya 131 later. Kaya Kinondo might have supplied 2–4 families in 1998 but 4–7 in 2003, when KD had firewood to supply just one family (table 4). Table 4. Volume of dead wood on the ground of 0.625 ha in KM (1) and 0.3125 ha in KK (2) in January 1998 (a) and January 2003 (b). Kaya 1 (a) 1 (b) Pieces of dead wood 109 88 Volume (m3) 26.5 5.9 Volume (m3 .ha -1) 42.4 9.4 2 (a) 2 (b) 67 64 3.4 5.7 11.0 18.2 Floristic diversity Three hundred and thirty one plant species in 77 families have been recorded so far in KM 4 . This represents ca. 7.4% of all species estimated for the Zanzibar-Inhambane regional mosaic/Swahilian region sensu lato or ca. 4.7% of all species known for Kenya 5 . In the sample area, 40 families and 95 species were recorded in strata A-E; in KK, 64 families and 192 species have been found, of which 49 in 31 families in the sample area; in KD, 60 families and 183 species were recorded including 79 in 38 families in the sample area (in appendix 1). In all three sample areas (1.25 ha) 174 species have been found in strata A-E of it 67 only in KM, 51 only in KD and 20 only in KK; 13 occur in all Kayas, seven are shared between KD/KM, eight between KD/KK and eight between KM/KK. In KD, 26 families occur in strata A-D on 0.3125 ha representing 45 species of ≥3 cm dbh This is the highest plant family diversity compared to KM (24 families on 0.625 ha) and KK (16 families on 0.3125 ha) (Lehmann & Kioko, 2000). An important aspect in the family composition for plants ≥3 cm dbh is, that the Caesalpiniaceae comprise eight species in the sample area of KM in strata A-D but none in KK and only one (Tamarindus indica) in KD. The majority of families (62%) and 40% of species of the whole KD were recorded on 0.1875 ha (strata A-E) including the highest number among the Rubiaceae (nine species). In both other Kayas, 50% of all families but only 22% of all species were represented on 0.1825 ha including the highest numbers amongst the Euphorbiaceae (KM, nine species) and Annonaceae/Sapindaceae (KK, five species each). But the species diversity is decreasing when a larger area is considered. Clarke et al. (2000) mention that between 300 and 800 vascular plant species are present per coastal forest. This might imply that KK/KD have depauperate floras as both have less than 200 species. However, this lower diversity seems to be natural for KK, which was found to be largely undisturbed. Kaya Diani is most diverse in strata C/D but the plant species composition of stratum D seems to be locally a consequence of pole cutting. Table 5 (sample size: 251 individuals in 45 species) represents ten species with the highest relative densities in the sample area. It shows that they comprise 56% of all individuals while the remaining 35 species occur at low 4 The total number of plant species and families per Kaya is based on the CFCU database of February 2000 and the additional records made by the authors until January 2003. 5 Clarke et al. (2000) mention at least 4500 plant species for the Zanzibar-Inhambane regional mosaic/Swahilian region sensu lato. Bennun & Njoroge (1999) mentioned ca. 7000 plant species for Kenya. 132 I. Lehmann & E. Kioko densities of more or less 1%. Trichilia emetica and Fernandoa magnifica together comprise 10% indicating some disturbance. Lecaniodiscus fraxinifolius, Markhamia zanzibarica and Ludia mauritiana are common in strata B and/or C but are absent to rare in stratum D were poles occur if the authors estimate that they are ca. 2.5 m long with a dbh of 3.5 cm. In KM/KK the ten species with highest relative densities comprise none which is typical for disturbed forest habitats and all are well distributed in stratum D except for Antiaris toxicaria showing largely an apparent lack of regeneration in KK (Lehmann & Kioko, 1998). Table 5. Ten species ≥3 cm dbh with the highest relative density on 0.3125 ha in KD. Species Stratum Lecaniodiscus fraxinifolius Synadenium pereskiifolium Vepris eugeniifolia Asteranthe asterias Trichilia emetica Haplocoelum inoploeum Fernandoa magnifica Markhamia zanzibarica Grewia plagiophylla Ludia mauritiana B, C C C, D C, D C, D C, D C, D C, D C, D C, D Percent of total number of Accumulative percent of plant individuals (N) N 7.2 7.2 6.8 14.0 10.8 24.8 6.4 31.2 5.6 36.8 5.2 42.0 4.4 46.4 4.0 50.4 2.8 53.2 2.8 56.0 Endemic plant genera and endemic plant species Clarke et al. (2000) list 33 endemic genera for the coastal forest belt of the ZanzibarInhambane regional mosaic/Swahilian region sensu lato. Based on this list eight of these occur in KM (Lettowianthus*, Ophrypetalum*, Zamioculcas*, Asteranthe, Grandidiera*, Mkilua*, Pseudobersama*, Schlechterina); the latter five in strata C-E in the sample area (table 6). Genera marked ‘*’are forest-dependent; tree genera are underlined. As endemic plant genera are thought to represent ancient phylogenetic lineages and endemic tree genera are considered to represent more ancient endemics compared to shrubs and herbs (Clarke et al., 2000) the high number of forest dependent endemics including three tree genera suggests that KM is an older forest site when compared to KK/KD where Ophrypetalum, Asteranthe, Schlechterina and Zamioculcas have been found in KK (the latter three in the sample area) and Ophrypetalum, Asteranthe and Schlechterina in KD (the latter two in the sample area). Table 6. Number of endemic plant genera in strata A-D of ≥3 cm dbh in the sample areas of KM (1), KK (2) and KD (3). Endemic genera Kaya 1 2 3 Total Overstorey trees 0 0 0 0 Understorey trees 3 1 1 3 Shrubs treelets 4 1 1 4 Lianes Total 0 0 0 0 4 1 1 4 Clarke et al. (2000) identified 1356 endemic plant species for the Swahilian regional centre of endemism sensu lato. Of these, 786 occur only in coastal forests. When divided by the 3170 km2 of forest area this gives 1 endemic per 4 km2 forest. We identified 99 Three kaya forests on the south coast of Kenya 133 endemic species occurring in KM (on 1.5 km2 of forest) 54 in KK and 50 in KD. Sixtysix endemics were found in the three sample areas but only five or 7.5% occur in all three Kayas. Almost 50% have a narrow species range along the Kenya or Kenya/Tanzania coast and each Kaya has different endemics, which are restricted to less than five localities. The sample area of KM has 39 endemics (41% of plant species). Nineteen endemics or 49% are restricted to coastal forests; 22 to the Kenya or Kenya/Tanzania coast including five which are rare in Kenya. Gigasiphon macrosiphon and Keetia lukei are restricted to less than five localities and the latter species is endemic to the Kenya coast. Kaya Kinondo has 18 endemics in the sample area (37% of plant species). Five endemics or 28% are forest dependent; 12 are restricted to the Kenya or Kenya/Tanzania coast; four are rare in Kenya. Diphasia sp. A of FTEA and Ziziphus robertsoniana are endemic to the Kenya coast and are restricted to less than three localities. The sample area of KD has 28 endemics (35% of plant species). Only four endemics are forest dependent. Hence, KD has the least number of forest dependent endemic genera and species. Eleven endemics are restricted to the Kenya or Kenya/Tanzania coast; two are rare in Kenya. Synsepalum subverticillata and Mitriostigma greenwayi are Kenyan endemics and the latter is restricted to less than five localities. Pole cutting of endemic plant species Endemics of ≥3 cm dbh account for at least 35% (KM), 32% (KK) and 38% (KD) of the of 62, 25 and 45 species (including unidentified species) recorded in strata A-D per sample area. Table 7 presents the total number of endemics having a size which makes them especially vulnerable to pole cutting or illegal logging.There are 22 endemic species (56% of endemics) in KM; eight in KK (44%) but 17 in KD (61%). Table 7. Number of endemic plant species in strata A-D of ≥ 3 cm dbh found in the sample areas of KM (1), KK (2) and KD (3). Category Kaya Coastal forest 1 2 3 1–3 Total Overstorey trees 6 2 3 11 Endemic species Understorey trees 13 5 13 27 Shrubs treelets 19 5 15 33 Lianes Total 0 0 0 0 22 8 17 35 From the conservation point of view it is also noteworthy that stratum E has between 25 and 50% of the total number of endemics, which were not recorded in other strata. Since stratum E is well developed in small natural gaps it shows how important they are for those endemics. Plant species distribution and dominance Clarke et al. (2000) found that 33% of the coastal forest flora is endemic/near endemic while 49.3% is widespread. Only 3.4% are species that occur in both the Swahilian and GuineoCongolian regions. Unfortunately, a list was not presented for those species. Hence, the authors follow Hawthorne (1993) who summarized eight chorological categories for plant species of the East African coastal forests and assigned species to each category. Table 8 shows that widespread species comprise less than 12% per Kaya and species which were 134 I. Lehmann & E. Kioko considered as Guineo-Congolian by Hawthorne (1993) comprise ca. 17% in KM, 7% in KK and 5% in KD. Table 8. 140 plant species found in KM (1), KK (2) and KD (3) on 0.1875 ha and assigned to chorological categories after Hawthorne (1993). Category Coastal Kaya 1 2 3 Species per stratum A B 2 1 1 2 0 1 C 4 4 10 D 9 2 14 E 18 11 23 Total 22 12 26 Eastern 1 2 3 1 0 0 3 3 1 5 2 4 4 1 5 8 5 5 7 5 6 Oceanic 1 2 3 0 0 0 0 0 1 0 0 3 0 1 5 3 3 4 3 3 7 Guineo-Congolian 1 2 3 3 1 0 2 0 1 3 0 1 4 1 2 10 3 3 13 3 4 Zambesian 1 2 3 0 0 1 0 0 1 0 0 0 0 0 1 1 1 0 1 1 2 Wide African 1 2 3 0 1 0 0 0 1 0 0 3 2 1 5 6 5 5 7 5 8 Unknown 1 2 3 1 2 3 0 0 0 6 3 1 0 2 3 6 7 9 5 3 5 17 9 26 4 2 12 23 8 44 24 11 19 70 39 59 22 13 21 75 42 74 Total The sample area of KM has 16 dominant species; KK/KD have 12 species each; KM/KD showing the least similarity (table 9). According to table 9 only KM has a high representation of dominant species, which are either endemics (50%) or linked to the Guineo-Congolian region. The latter include the genera Scorodophloeus and Mildbraedia. Both are unique to the Guineo-Congolian region and coastal forest belt (Clarke et al., 2000). This indicates that KM must be a relict of the once extensive “Pan-African lowland forest” (cf. Lovett, 1993). Additionally, the dominance of a number of caesalpiniaceous tree species in KM is similar to forests of West and Central Africa described as “Wet evergreen forests” by Hamilton (1989). As mentioned before, trees represent more ancient endemics than shrubs and hence, KM is an older forest site than KK/KD since both have less dominant endemic tree species. Three kaya forests on the south coast of Kenya 135 Table 9. Dominant species of sample sites and most dominant species with basal area (m2) and its percentage of total basal area of the sample area in KM, KK (1998) and KD (2003). Endemics are underlined. Forest Strata Dominant species KM A Julbernardia magnistipulata, Scorodophloeus fischeri, Parkia filicoidea, Antiaris toxicaria B Cynometra suaheliensis, C. webberi; Julbernardia magnistipulata, Scorodophloeus fischeri, Sorindeia madagascariensis, Synsepalum brevipes C Craibia brevicaudata; Drypetes natalensis, D. reticulata, Fernandoa magnifica, Julbernardia magnistipulata, Scorodophloeus fischeri, Sorin-deia madagascariensis, Synsepalum brevipes D Acalypha neptunica var. neptunica, Cynometra suaheliensis, Julbernardia magnistipulata, Mildbraedia carpinifolia var. carpinifolia, Mkilua fragrans, Streblus usambarensis KK A Antiaris toxicaria, Adansonia digitata, Trichilia emetica B Diospyros ferrea, Diphasia sp. A, Sorindeia madagascariensis C Diphasia sp. A, Drypetes natalensis, Sorindeia madagascariensis, Terminalia catappa D Grewia plagiophylla, Polysphaeria parvifolia, Rinorea elliptica, Uvariodendron kirkii KD A Diospyros ?squarrosa B Adansonia digitata, Balanites maughamii, Lannea welwitschii, Lecaniodiscus fraxinifolius C Balanites maughamii, Lecaniodiscus fraxinifolius, Tamarindus indica, Trichilia emetica, Sterculia appendiculata D Asteranthe asterias, Balanites maughamii, Grewia plagiophylla, Haplocoelum inoploeum, Vepris eugeniifolia Most dominant species Scorodophloeus fischeri 3.3 m2 or 24.1% Scorodophloeus fischeri 1.76 m2 or 27.2% Synsepalum brevipes 0.59 m2 or 22.1% Acalypha neptunica var. neptunica 2 0.056 m or 16.7% Antiaris toxicaria 2.16 m2 or 37.9% Sorindeia madagascariensis 3.27 2 m or 73.5% Drypetes natalensis 0.65 m2 or 22.2% Rinorea elliptica 0.009 m2 or 15% 0.08 m2or 100% 6 2 Adansonia digitata 1.95 m or 33.6% Tamarindus indica 0.73 m2 or 26.5% Haplocoelum inoploeum 0.052 m2 or 18% Butterfly diversity So far 127 species of butterflies have been recorded in the study area in Kaya Muhaka over a period of ten years (1994–2003). This is ca. 14% of the currently known Kenyan butterfly fauna. Ninety-three species were recorded on only 0.625 ha. A comparison to the Tanzanian forest sites mentioned in the introduction shows that KM is very rich in butterfly species, probably one of a few smaller coastal forests in Kwale District with such a high diversity. Only 56 species have been recorded in the study area of KK over ten years although extensive field work was undertaken. Thirty-seven species were found on 0.3125 ha. Kaya Diani has 77 species which have been recorded in the study area during 2001–2003 including 6 This specimen of Diospyros was the only one in the sample area in stratum A. Generally, the tallest trees are Ficus species and Sterculia appendiculata. The latter is probably most dominant in stratum A in the study area. 136 I. Lehmann & E. Kioko 49 found on 0.3125 ha. In all three study areas (ca. 42 ha) five families and 156 species occur (17% of the Kenyan fauna): 53 species only in KM, 11 only in KK, 14 only in KD; 26 in all Kayas; 33 are shared by KM/KD, 15 by KM/KK, four by KK/KD (appendix 2). Of interest is, that the Pieridae comprise only five (mainly widespread) species in KK. As the Pieridae are more closely linked to open formations than other butterfly families, their weakly representation perhaps indicates in general a more closed, less disturbed coastal forest. Table 10. Total number of butterfly species per family in the three study areas. Family Papilionidae Pieridae Lycaenidae Nymphalidae Hesperiidae Total Butterfly species and percentage of total number KM % KK % KD 9 19 24 60 15 127 7 15 19 47 12 100 7 5 9 28 7 56 12.5 9 16 50 12.5 100 10 18 15 30 4 77 % 13 23 20 39 5 100 Total Number 11 23 33 67 22 156 Seasonal or local butterfly species About 23% of species of KM/KK (listed by Lehmann & Kioko, 2003) were found in 1994– 1999 but not between 2000–2003. Fifteen species of KM and 11 of KK were first recorded after the year 2000 such as the large Pseudacraea boisduvalii trimeni in KK in 2003. This shows that even for small forest patches, many years of field work are needed to get a more or less complete species list. One reason is that several species are seasonal (e.g. Libythea labdaca laius, Sallya amulia rosa with no dry season record) or are rare and restricted to a few patches of ca. 1000 m2 of closed forest (e.g. Baliochila minima, Deudorix dinomenes, Euxanthe t. tiberius, Hypolimnas usambara, Salamis cacta amaniensis). The latter are especially sensitive to habitat disturbance e.g. Salamis cacta amaniensis disappeared from two sites in KM after 1999 probably due to logging/pole cutting since both sites became more open and dry. This may result in extinction of any species with a small population size. Rare butterfly species Ten rare species have been found in KM, four in KK and none in KD. The rarity status follows Larsen (1996) and Burgess & Clarke (2000, in appendix 9); underlined species were represented with more than three specimens per year mentioned. KM: Teriomima micra (1996–2003), Baliochila minima (1996–1998), Deudorix dinomenes (1996), Charaxes acuminatus shimbanus (1996, 2001–2003), Charaxes pythodoris nesaea (1997– 1998), Euxanthe t. tiberius (1997–1998), Pseudacraea eurytus conradti (1994–2003), Hypolimnas usambara (1994, 1996–1998), Acraea aubyni (1996–2003), A. zonata (1996–1998). KK: Charaxes acuminatus shimbanus (1996), Sallya amulia rosa (1997; migrant species with a few records from southeast Kenya and hence, the authors treat it here as rare for Kenya), Hypolimnas usambara (1998/99), Borbo f. ferruginea (2003). The records of the rare montane Kenyan endemic subspecies Charaxes acuminatus shimbanus are the first ones near sea-level (Lehmann & Kioko, 2000; Kroon, 2001). Endemic butterfly genera and species The genera Euthecta and Eresinopsides are endemic to the coastal forests of eastern Africa as well as the near-endemic genus Teriomima (Kielland & Cordeiro, 2000). The latter occurs in Three kaya forests on the south coast of Kenya 137 KM and KD. Burgess & Clarke (2000, in appendix 9) presented the 134 butterfly species or subspecies, which are mostly confined to coastal forests or semi-evergreen vegetation close to the coast of eastern Africa. Based on their list, 25 endemic species or 19% were recorded: 23 in KM, 7 in KK, 9 in KD. Eleven species occur only in KM; Borbo f. ferruginea in KK and Graphium philonoe in KD only. Six endemics penetrate also into non-forest vegetation. The genus Baliochila,which is very closely related to the near-endemic genus Teriomima, is only well represented in KM with two forest-dependent species and specimens of the Kenyan endemic Charaxes acuminatus shimbanus have been found locally three times. Both other Kayas share a high number of endemics (87%) with KM. In contrast, only 7.5% of endemic plant species occur in all three Kayas. Butterfly species distribution If the butterfly species of the three Kayas and of SH (based on the authors assessment of the data published by Sevastopulo (1973, 1974a,b; see here in appendix 2) are assigned to biogeographical elements after Larsen (1996), two biogeographical groupings were found among 266 species or almost 30% of the Kenyan butterfly fauna (table 11): Table 11. Biogeographical elements after Larsen (1996) found in four coastal forests. Element Number of butterfly species per area SH % KM % KK % KD % Total % General distributions All forest zones Main forest zone Equatorial forest zone Coastal forest zone Transitional species All Africa open formations Sudanian open formations Somali open formations Zambesian open formations Montane distributions Special habitats Unknown 29 36 1 2 56 13 36 12 15 0.4 0.8 23 5 15 20 21 1 0 37 7 16 16 17 0.8 0 29 5 13 14 8 0 0 17 3 2 25 14 0 0 30.4 5.4 3.6 19 12 0 0 17 5 7 25 15.5 0 0 22 6.5 9 31 37 2 2 60 13 38 11.6 14 0.8 0.8 23 5 14 3 1 0 0 0 0 0 0 3 1 7 3 1 0.8 0 0 1 1.3 7 2.6 43 17 12 9 5 9 9 11.7 46 17 1 2 17 0.4 0.8 7 1 1 10 0.8 0.8 8 1 1 5 1.8 1.8 9 0 1 6 0 1.3 7.7 1 2 24 0.4 0.8 9 Total 246 100 127 100 56 100 77 100 266 100 1. Coastal forests with a high number of species with “General distributions” (>20%) and of the “Coastal forest zone” comprising together about 50% or more of the total number of butterfly species per forest (KK/KD). 2. Coastal forests above and well below 300 m where at least one species with “Montane distributions” and a high number of species from the “Coastal forest zone”, “All forest zones” and with at least one genus and/or species from the “Main forest zone” and/or “Equatorial forest zone” (genus and/or species centred on West and Central African 138 I. Lehmann & E. Kioko rainforests) occur. They comprise for about 40% or more. Species with “General distributions” have less than 20% (SH/KM). Both, SH and KM include a higher number of forest-dependent species e.g. the genera Euriphene (Euryphura) and Euptera which are centred on the rainforest zone and Charaxes acuminatus shimbanus representing the “Montane distributions”. Because of these striking similarities it seems likely that KM was once connected to the forests of SH. Larger moth diversity and composition Further research is needed on the following results because the authors assume that their species lists are still incomplete (especially for KD), that additional species might be defined as rare, forest-dependent and endemic, and that the taxomomic status of others is unclear at the moment. In all three study areas, a total number of 16 families and 273 larger moth species have been recorded so far. Sixteen families and 225 species in KM; 11 families and 98 species in KK; 8 families and 36 species in KD. Kaya Kinondo has a remarkable low diversity of larger moths and all light-trapping sessions since 1994 were disappointing with sometimes less than ten species per session and together with extensive field work during daytime only 40 additional species were found after 2000. Noteworthy is also that the superfamily Cossoidea (families Cossidae, Metarbelidae, Limacodidae) represents only one species in KK, none in KD but 11 in KM where the family Metarbelidae is diverse including a Salagena ?sp. nov. and Teragra simillima, which are locally common. This difference in the family composition to both other Kayas is of interest because it might reflect the higher rainfall in KM or the wetter forest type respectively (table 12 and in appendix 3). Rare, forest-dependent and endemic larger moth species The results presented here are based on Fletcher (1974), Sevastopulo (1979, 1982), Krüger (2001), as well as on material seen in the BMNH (London), NMK (Nairobi), the Museum für Naturkunde (Berlin) and own records. The authors treat a species as “rare” for coastal Kenya if: 1. A species was first recorded from Kenya (underlined) or coastal Kenya and less than three specimens were seen in 1994–2003 (species marked ‘*’ were listed by Lehmann & Kioko, 2000); including for KM: Euproctis rufopunctata*, Lymantria leucerythra* (plate 1a) (Lymantriidae); Grammodes congesta*, Metaleptina nigribasis*, Oglasodes nyasica, (plate 1d) (Noctuidae); Chiasmia orientalis and for KK: Racotis breijeri* (Geometridae). 2. A species is currently known from five localities or less along the Kenya coast and less than three specimens were seen during 1994–2003; including for KM: Antharmostes p. papilio, Mixocera viridans (Geometridae); Conigephyra pallidula, Dasychira callipepla (Lymantriidae); Secusio drucei (Arctiidae); Dysgonia conjunctura, Heliophisma xanthoptera, Marcipa insulata, Rougeotiana xanthoperas (plate 1b), Ulotrichopus primulinus and for KK: Ozarba perplexa (Noctuidae). 3. A species is known from five localities or less along the Kenya coast but 3–10 specimens were seen in 1994–2003; including for KM: Stracena bananae (Lymantriidae), Rhanidophora cinctigutta; for KK: Oglasa nana and for KK/KD Thyas arcifera (Noctuidae). Three kaya forests on the south coast of Kenya 139 Based on own observations several species appear to be forest-dependent and do not penetrate into open vegetation. The authors compared these species with labels of museum specimens in collections mentioned above. If literature notes and/or 90% of labels per species indicated again a forest site then it will be treated here as forest-dependent; species marked ‘*’ occur also in forests of West and/or Central Africa but, based on Vári et al. (2002), do not occur in southern Africa; rare species underlined: Salagena ?sp. nov. (Metarbelidae); Scotinocerides conspersa (Limacodidae); Chiasmia feraliata, Chiasmia orientalis, Plegapteryx anomalus, Zamarada euphrosyne*, Zamarada keraia, Zamarada rufilinearia (Geometridae), Acropteris erycinaria* (Uraniidae), Mimopacha tripunctata* (Lasiocampidae), Eurystaura griseitincta (Notodontidae), Dasychira barbara, Stracena bananae (Lymantriidae), Secusio drucei* (Arctiidae), Aburina sobrina*, Anomis punctulata*, Caryonopera breviramia*, Ercheia subsignata, Giria pectinicornis*, Marcipa insulata*, Metaleptina nigribasis*, Ophiusa gonoptera, Pseudogiria polita*, Rougeotiana xanthoperas*, Ugia amaponda, Thyas arcifera (Noctuidae). This list indicates that at least 11% of larger moths in KM and 6% in KK are forest-dependent including at least 5% and 2% respectively which are also distributed in forests of West and/or Central Africa but do not occur in southern Africa. Table 12. Larger moth families and their number of species in three Kaya forests. Larger moth species per family and area KM Family Cossidae Metarbelidae Limacodidae Geometridae Uraniidae Lasiocampidae Eupterotidae Saturniidae Sphingidae Notodontidae Thyretidae Lymantriidae Arctiidae Herminiidae Aganaidae Noctuidae Total KK % KD % % Total Number % 2 7 2 48 3 3 1 2 10 4 2 13 11 3 4 110 0.8 3.1 0.8 21.3 1.3 1.3 0.4 0.8 4.4 1.7 0.8 6 5 1.3 1.7 49 0 0 1 22 2 0 0 0 2 1 3 3 8 1 3 52 0 0 1 22 2 0 0 0 2 1 3 3 8 1 3 52 0 0 0 7 1 0 0 1 2 0 0 1 4 0 2 18 0 0 0 19.5 2.8 0 0 2.8 5.5 0 0 2.8 11.1 0 5.5 50 2 7 3 59 3 3 1 2 11 4 3 15 17 3 4 136 0.7 2.5 1 22 1 1 0.4 0.7 4.0 1.5 1 5.5 6.2 1 1.5 50 225 100 98 100 36 100 273 100 According to the material seen in the museums mentioned above, literature notes and own observations, five species from KM, including one (Paraptychodes tenuis, plate 1c) occurring also in KK/KD, were up to date only found in coastal eastern Africa, mainly in Kenya and/or Kenya/Tanzania. These larger moth species will be treated here as endemic to the vegetation of the coast of eastern Africa and are mentioned in appendix 3. 140 I. Lehmann & E. Kioko DISCUSSION Logistic/historical factors for levels of biodiversity The three Kayas are continental habitat islands with high levels of biodiversity (using species richness and the degree of endemism as a measurement). Particularly rich in species is KM (table 13) due to three main factors. Firstly, the generally higher number of species in KM reflects its larger area. This follows Forman et al. (1976) who stated that island size is generally an important predictor of species numbers. Secondly, KM is an older forest, a relict of the once extensive “pan-African lowland forest”, and it has also been isolated longer. This becomes clear from the high number of forest-dependent endemics as well as plant and Lepidoptera genera and/or species linked to the forests of the Guineo-Congolian region. The number of such genera and/or species is decreasing in younger forest sites (e.g., KK/KD) where a higher number of species with “General distributions” occur due to colonisations of dispersive species with high levels of reproduction and a short generation time. Thirdly, KM receives a higher average annual rainfall which enables drought intolerant plant and Lepidoptera species to persist. Wieringa & Poorter (2004) described an increasing plant species richness along the rainfall gradient with an optimum around 2500 mm in West African forests and studies of Janzen & Schoener (1968) in forests of Costa Rica made clear that insect abundance and diversity is increasing with moisture levels. Kielland & Cordeiro (2000) pointed out that the moist coastal climate stretches much further inland in Tanzania when compared to Kenya and species of the coastal butterfly fauna similarly range further inland. Kaya Muhaka has 127 endemic plant and Lepidoptera species while the other two Kayas have well below 70 endemics each. According to the definition presented by Burgess (2000) that areas with more than 100 endemic species (not only plants) are considered as local centres of endemism, the authors believe that a further sub-division of the “Usambara-Kwale local centre of endemism” is possible may be between coastal forests further inland (e.g. KM) and those close to the shoreline of the Indian Ocean (e.g. KK/KD). The latter should be less diverse in endemic species since in general, endemic species are an indication of a longer period of divergence in isolation (MacArthur & Wilson, 1967). Such a period was certainly longer for older forest sites further inland. Table 13. Plant and Lepidoptera species diversity of three Kaya forests in Kwale District. ! =Coastal endemics; GC = Species linked to the Guineo-Congolian region. Note that the number of species is preliminary since only the total number of plants and the endemic plant species were found in the whole Kaya but all Lepidoptera in the smaller study area and rare/GC plant species in the smaller sample area. Group Plants Butterflies Larger moths Total KM Total 331 127 225 683 Preliminary number of species per Kaya KK ! Rare GC Total ! Rare GC 99 5 13 192 54 4 1 23 10 1 56 7 4 0 5 18 12 98 1 4 2 127 33 26 346 62 12 3 KD Total 183 77 36 296 ! Rare 50 2 9 0 1 1 60 3 GC 3 0 0 3 Habitat availability Although the flora in KK is diverse, Lepidoptera do not respond to it. The groundwater forest of KK has a low Lepidoptera diversity including only one genus (Euptera) which Three kaya forests on the south coast of Kenya 141 is centred on the rainforest zone; KD has no such genus. In contrast, KM receives ca. 150 mm more rainfall annually than KK/KD. This rather small additional amount of rainfall seems to be enough for two Lepidoptera genera (Euriphene, Euptera) to persist, which are centred in the rainforest zone. A higher number of Lepidoptera species in KM are also linked to forests of West and Central Africa, probably favouring wetter forest habitats. Such species appear to be sensitive to small scale habitat changes e.g. Euriphene achlys disappeared locally in KM from several sites which became drier due to tree poaching. Disturbance Disturbance occurred in KK where ceremonies were or are still carried out. Since 1994, only two more or less open patches each ca. 700 m2 in size are still visible in the study area through a clumped dispersion of the strong light demanders Antiaris toxicaria-Milicia excelsa-Terminalia catappa-Trichilia emetica in strata A and B near a sacred place (figure 4 in Lehmann & Kioko, 2000) and near a small pond with large Terminalia catappa trees. Since no other signs of disturbance were found, the authors do not believe that an entire group of Digo people once lived in the forest interior of KK (cf. Spear, 1978). The opinion of the authors is supported by Willis (1993) who stated that Spear’s presentation of Mijikenda residence in the Kayas is a problematic one. Both other Kayas are moderately disturbed probably rather due to tree poaching than past residences within the forest. But it is encouraging to see that KM still supports a very rich endemic flora and Lepidoptera fauna when compared to KK. Kaya Kinondo indicates that a coastal forest which is largely undisturbed is not automatically rich in Lepidoptera species. These results confirm Hawthorne (1993) who stated, that disturbance is not a main cause of variation in any community, having a subordinate role. Conservation The setting of priority sites for conservation of coastal forests was suggested by Rodgers & Burgess (2000) by using species richness or the degree of endemism or the magnitude of threat as a measurement. The authors recommend focusing firstly on “priority areas” defined as all habitats (not only forest) of any large potential species source (e.g. SH to protect also non-forest-dependent Kenyan endemics like the butterfly Acraea matuapa) and nearby forests (e.g. KM) to raise each island’s immigration from a potential species source because extinctions are more likely on smaller islands and on islands more distant from the source. Secondly, on the “diversity of forest types” along the Kenya coast including patches of rare types which might have rather low levels of species richness and/or endemism (e.g. KK). Thirdly, on “any single-site or near single-site endemic” because more than one Kenyan endemic plant species including at least one different species which is restricted to less than five localities have been found per sample area. The latter are near to single-site endemics (e.g. Keetia lukei in KM, Diphasia sp. A in KK, Mitriostigma greenwayi in KD). Levels of such near single-site endemism are lower among Lepidoptera (Charaxes acuminatus shimbanus, probably Zamarada keraia in KM only). The sacred Kaya forests are part of a cultural ethnosystem that has up to now preserved rare species and endemics, which have disappeared elsewhere. It shows that traditional management under the leadership of the Committee of Elders is able to protect and conserve cultural as well as natural values in an symbiotic relationship. The control of the local community upon KK is an outstanding example for sustainable management of an 142 I. Lehmann & E. Kioko coastal forest, partly achieved through the development and implementation of community centred conservation, education and ecotourism projects. It becomes clear how effective participation of local communities could be elsewhere in helping to protect the originality and biodiversity of all Kaya forests, which are an important part of East Africa’s cultural landscapes. 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Unpublished report. Three kaya forests on the south coast of Kenya 147 Appendix 1. Species list of plants recorded in the sample areas of KM , KK and KD. Nomenclature follows mainly Beentje (1994) and CFCU database (February, 2000). Families or species marked ‘*’ are new records (= not included in 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 (K or K/T = Kenya or Kenya/Tanzania only); ! (s) = Coastal endemic (southwards); !! = restricted within coast; !!! = very restricted within coast; ? = Unknown. Species marked ‘! *’ are Coastal endemics after Burgess & Clarke (2000, in appendix 3) but were not treated as such by Hawthorne (1993). Species marked ‘F’ are forest-dependent Coastal endemics. Family KAYA MUHAKA Acanthaceae Anacardiaceae Annonaceae Apocynaceae Asclepiadaceae Bignoniaceae Buxaceae Caesalpiniaceae Celastraceae Combretaceae Connaraceae Dichapetalaceae Ebenaceae Euphorbiaceae Flacourtiaceae Guttiferae Loganiaceae Malvaceae Plant Species and Subspecies Stratum Category Monothecium aristatum T.Anderson Whitfeldia elongata (Beauv.) C.B.Clarke Sorindeia madagascariensis DC. Asteranthe asterias (S.Moore) Engl.& Diels ssp. asterias Mkilua fragrans Verdc. Uvaria lucida Benth. ssp. lucida Uvariodendron kirkii Verdc. Xylopia parviflora (A.Rich.) Benth. Ancylobotrys petersiana (Kl.) Pierre Landolphia watsoniana Romburg Saba comorensis (Bojer) Pichon Tylophora apiculata K.Schum. Fernandoa magnifica Seem. Buxus obtusifolia (Mildbr.) Hutch. Cynometra suaheliensis (Taub.) Baker f. Cynometra webberi Baker f. Dialium holtzii Harms Erythrophleum suaveolens (Guill. & Perr.) Brenan Gigasiphon macrosiphon (Harms) Brenan Hymenaea verrucosa Gaertn. Julbernardia magnistipulata (Harms) Troupin Scorodophloeus fischeri (Taub.) J.Léonard Salacia madagascariensis (Lam.) DC. Combretum illairii Engl. Agelaea pentagyna (Lam.) Baill. Ellipanthus madagascariensis (G.Schellenb.) Keraudr. Dichapetalum ruhlandii Engl. Tapura fischeri Engl. Diospyros abyssinica (Hiern) F.White ssp. abyssinica Diospyros greenwayi F.White Diospyros kabuyeana F.White ? Acalypha fruticosa Forssk. * Acalypha neptunica Muell.Arg. var. neptunica Alchornea laxiflora (Benth.) Pax & K.Hoffm. Antidesma venosum Tul. Drypetes natalensis (Harv.) Hutch. var. leiogyna Brenan Drypetes reticulata Pax Mallotus oppositifolius (Geiseler) Muell.Arg. Mildbraedia carpinifolia (Pax) Hutch. var. carpinifolia ? Oldfieldia somalensis (Chiov.) Milne-Redh. * Suregada zanzibariensis Baill. Tragia furialis Bojer Dovyalis macrocalyx (Oliv.) Warb. Grandidiera boivinii Jaub. Garcinia livingstonei T.Anderson Garcinia volkensii Engl. Strychnos panganensis Gilg Gossypioides kirkii (Mast.) J.B.Hutch. E E A, B, C, D, E C, D, E C, D, E E D, E E E E C, D, E E C, D, E E A, B, C, D B, C, D C, D, E A, E A, E A, E A, B, C, D, E A, B, C, D, E E D, E E C, E D, E E D, E E E D C, D, E D, E D, E B, C, D, E C, D, E D D E E E E D, E D E E E ? GC + E (z) !; K/T !!; F; K/T ? !; K/T A ? ! *;F; K/T GC + ! *;F; K/T !* !; F; K/T !; K/T !; F; K/T ! GC !!; F; K/T O !; K/T !; F OM ! ? !; F; K/T E GC A ! *; F !; F; K/T ? GC A A E ! (s) GC + !; F ! O ? ? !; F A ? ! ! 148 Family KAYA MUHAKA Melastomataceae Meliaceae Mimosaceae Moraceae Papilionaceae Passifloraceae Rhamnaceae Rubiaceae Sapindaceae Sapotaceae Sterculiaceae Thymeleaceae Tiliaceae Ulmaceae* Verbenaceae Violaceae Vitaceae Commelinaceae Cyperaceae Dracaenaceae Gramineae Davalliaceae Total: 40 KAYA KINONDO Zamiaceae Anacardiaceae Annonaceae I. Lehmann & E. Kioko Plant Species and Subspecies Stratum Category Warneckea sansibarica (Taub.) var. sansibaricum Pseudobersama mossambicensis (Sim) Verdc. Trichilia emetica Vahl Newtonia paucijuga (Harms) Brenan Parkia filicoidea Oliv. Antiaris toxicaria Lesch. Dorstenia kameruniana Engl. Ficus sp. Streblus usambarensis (Engl.) C.C.Berg Trilepisium madagascariensis DC. Craibia brevicaudata (Vatke) Dunn ssp. brevicaudata Schlechterina mitostemmatoides Harms ? Lasiodiscus mildbraedii Engl. ssp. ferrugineus (Verdc.) Faden* Chazaliella abrupta (Hiern) Petit & Verdc. var. abrupta Coffea sessiliflora Bridson Keetia lukei Bridson Keetia zanzibarica (Klotzsch) Bridson ssp. zanzibarica Polysphaeria parvifolia Hiern Psychotria capensis (Ecklon) Vatke ssp. riparia Psychotria lauracea (K.Schum.) Petit Vangueria randii S.Moore ssp. acuminata Verdc. Blighia unijugata Baker Chytranthus obliquinervis Engl. Lecaniodiscus fraxinifolius Baker ssp. vaughanii (Dunkley) Friis Lepisanthes senegalensis (Poir.) Leenh. Majidea zanguebarica Oliv. Pancovia golungensis (Hiern) Exell & Mendonca Paullinia pinnata L. Synsepalum brevipes (Baker) Pennington ? Synsepalum msolo (Engl.) Pennington * Synsepalum subverticillata (E.A.Bruce) Pennington Cola minor Brenan Nesogordonia holtzii (Engl.) Capuron * Synaptolepis kirkii Oliv. Grewia plagiophylla K.Schum. Celtis mildbraedii Engl. * Clerodendrum incisum Klotzsch Lantana camara L. * Rinorea ilicifolia (Oliv.) Kuntze var. ilicifolia Rinorea squamosa (Tul.) Baill. ssp. kaessneri (Engl.) Grey-Wilson Cissus sciaphila Gilg Cissus sylvicola Masinde & L.E.Newton Commelina sp. Kyllinga cartilaginea K.Schum. Dracaena mannii Baker (= usambarensis Engl.) Panicum sp. Davallia denticulata (Burm.f.) Kuhn var. denticulata 95 (9 unidentified species omitted) E C, D, E D A, B A, E A, B D D D B C, D, E E D E E D E C, D, E E E D, E D C, D, E B, C, D, E E A, E C, D, E E B, C, D, E D D, E D, E E E C, D, E C E D, E E D E E E E C E E ? !; F A !; F; K/T GC GC GC ? ? GC ? ! !; F; K GC !; F; K/T !!! *; F; K ? E (n) Z ? ? GC + !; F; K/T E (v) A ? (!) A ? GC GC !!; K !; K ! !; F ! *; K/T GC !* ? GC OM !* ! *; K/T ? ! *; K/T GC ? ? Encephalartos hildebrandtii A.Braun & C.D.Bouché var. hildebrandtii Sorindeia madagascariensis DC. Asteranthe asterias (S.Moore) Engl. & Diels ssp. asterias Monanthotaxis fornicata (Baill.) Verdc. Ophrypetalum odoratum Diels ? Polyalthia stuhlmannii (Engl.) Verdc. * Uvariodendron kirkii Verdc. E ! B, C, E C, D, E E E E D, E E (z) !; K/T ! !; F; K/T !; K/T !; K/T Three kaya forests on the south coast of Kenya Family KAYA KINONDO Apocynaceae Bombacaceae* Celastraceae Combretaceae Connaraceae Ebenaceae Erythroxylaceae Euphorbiaceae Flacourtiaceae Guttiferae Meliaceae Montiniaceae Moraceae Ochnaceae Oleaceae Passifloraceae Rhamnaceae Rubiaceae Rutaceae Sapindaceae Sterculiaceae Thymeleaceae Tiliaceae Verbenaceae Violaceae Vitaceae Araceae Gramineae Total: 31 KAYA DIANI Acanthaceae Amaranthaceae* Anacardiaceae Annonaceae Apocynaceae Balanitaceae Family 149 Plant Species and Subspecies Stratum Category Hunteria zeylanica (Retz.) Gardn. Saba comorensis (Bojer) Pichon Adansonia digitata L. * Salacia elegans Oliv. Terminalia catappa L. Agelaea pentagyna (Lam.) Baill. Diospyros ferrea (Willd.) Bakh. Erythroxylum emarginatum Thonn. Drypetes natalensis (Harv.) Hutch. var. leiogyna Brenan Drypetes reticulata Pax Drypetes usambarica (Pax) Hutch. var. mrimae Mallotus oppositifolius (Geiseler) Muell.Arg. Ludia mauritiana J.F.Gmel. Calophyllum inophyllum L. Garcinia livingstonei T.Anderson Trichilia emetica Vahl Grevea eggelingii Milne-Redh. var. keniensis Verdc. (= Grevea madagascariensis Baill. ssp. keniensis Verdc.) Antiaris toxicaria Lesch. Ficus sp. Ochna thomasiana Engl. & Gilg Chionanthus battiscombei (Hutch.) Stearn Adenia gummifera (Harv.) Harms var. gummifera Lasiodiscus pervillei Baill. ssp. pervillei Ziziphus robertsoniana Beentje Polysphaeria parvifolia Hiern ? Tricalysia pallens Hiern * Diphasia sp. A of FTEA Toddalia asiatica (L.) Lam. Chytranthus obliquinervis Engl. Lecaniodiscus fraxinifolius Baker ssp. vaughanii (Dunkley) Friis Lepisanthes senegalensis (Poir.) Leenh. Majidea zanguebarica Oliv. Pancovia golungensis (Hiern) Exell & Mendonca Stadmania oppositifolia Poir. ssp. oppositifolia ? Cola minor Brenan* Synaptolepis kirkii Oliv. Grewia plagiophylla K.Schum. Premna hildebrandtii Gürke Rinorea elliptica (Oliv.) Kuntze Cissus sylvicola Masinde & L.E.Newton Zamioculcas zamiifolia (Lodd.) Engl. * Oplismenus burmannii (Retz) P.Beauv. 49 (6 unidentified species omitted) E D, E A E A, C, E E B, C, E E B, C, D, E B, C C C, D, E E E E A, E E O GC + ? ? ? ? ? A E ! (s) !! GC + O ? A A !! A, E B, C E E C, D, E D, E E D, E E A, B, C, D, E E C, E E E E D, E A, B, E D, E E D, E E D, E B, C E E GC ? !; K/T ? E (z) ? !! *; F; K E (n) ? !!!; F; K ? !; F; K/T E (v) A ? (!) A E (z) !; K !; F ! *; K/T ! *; K/T O ! *; K/T Z (!*) ? E E C, D, E B, D, E C, D, E C, D, E D, E D, E E C, D, E E C, D, E C, D, E B, C, D, E Stratum ! A E (z) GC !; K/T ! ! *; F !; K/T A O GC E (z) !* ! *; K/T Category Ecbolium amplexicaule S.Moore Scadoxus multiflorus (Martyn) Raf. * Sorindeia madagascariensis DC. Lannea welwitschii (Hiern) Engl. var. ciliolata Engl. Asteranthe asterias (S.Moore) Engl. & Diels ssp. asterias Monanthotaxis fornicata (Baill.) Verdc. Uvaria acuminata Oliv. Uvariodendron kirkii Verdc. Xylopia parviflora (A.Rich.) Benth. Hunteria zeylanica (Retz.) Gardn. Pleiocarpa pycnantha (K.Schum) Stapf * Schizozygia coffaeoides Baill. Tabernaemontana elegans Stapf Balanites maughamii Sprague ssp. acuta Sands Plant Species and Subspecies 150 KAYA DIANI Bignoniaceae Bombacaceae Boraginaceae Burseraceae Caesalpiniaceae Celastraceae Combretaceae Connaraceae Dichapetalaceae Ebenaceae* Erythroxylaceae Euphorbiaceae Flacourtiaceae Labiatae Meliaceae Moraceae Ochnaceae Papilionaceae Passifloraceae Rhamnaceae Rubiaceae Rutaceae Sapindaceae Sapotaceae Sterculiaceae Thymeleaceae Tiliaceae I. Lehmann & E. Kioko Fernandoa magnifica Seem. Markhamia zanzibarica (DC.) Engl. Adansonia digitata L. Bourreria nemoralis (Gürke) Thulin Cordia goetzei Gürke* Cordia monoica Roxb. Commiphora edulis (Kl.) Engl. ssp. boiviniana Tamarindus indica L. Salacia elegans Oliv. Salacia stuhlmanniana Loes. Combretum pentagonum Laws. Agelaea pentagyna (Lam.) Baill. Dichapetalum madagascariense Poir. Diospyros squarrosa Klotzsch* Erythroxylum emarginatum Thonn. Alchornea laxiflora (Benth.) Pax & K. Hoffm. Drypetes reticulata Pax Mallotus oppositifolius (Geiseler) Muell.Arg.* Suregada zanzibariensis Baill. Synadenium pereskiifolium (Baill.) Guill. Flacourtia indica (Burm.f.) Merrill * Ludia mauritiana J.F.Gmel. Oncoba spinosa Forssk. Hoslundia opposita Vahl Azadirachta indica A.Juss. Trichilia emetica Vahl Turraea wakefieldii Oliv. Ficus lingua De Wild. & T.Durand ? Ficus sansibarica Warb. Ochna thomasiana Engl. & Gilg Dalbergia vacciniifolia Vatke Millettia usaramensis Taub. ssp. usaramensis Adenia gummifera (Harv.) Harms var. gummifera Schlechterina mitostemmatoides Harms Ziziphus mucronata Willd. ssp. mucronata Coffea pseudozanguebariae Bridson Mitriostigma greenwayi Bridson Polysphaeria parvifolia Hiern Psydrax sp. Rytigynia sp. * Tarenna nigrescens (Hook.f.) Hiern * Tarenna supra-axillaris (Hemsl.) Bremek. ssp. supra-axillaris Tricalysia ovalifolia Hiern ? Tricalysia sp. Vepris eugeniifolia (Engl.) Verdoorn Zanthoxylum holtzianum (Engl.) P.G.Waterman Allophylus rubifolius (A.Rich.) Engl. Haplocoelum inoploeum Radlk. Lecaniodiscus fraxinifolius Baker ssp. vaughanii (Dunkley) Friis Lepisanthes senegalensis (Poir.) Leenh. Majidea zanguebarica Oliv. Manilkara sulcata (Engl.) Dubard Sideroxylon inerme L. ssp. diospyroides (Baker) J.H.Hemsl. Synsepalum subverticillata (E.A.Bruce) Pennington Sterculia appendiculata K.Schum. Cola sp. Synaptolepis kirkii Oliv. Carpodiptera africana Mast. Grewia ectasicarpa S.Moore * Grewia plagiophylla K.Schum. C, D, E C, D, E B D, E D E D C E D, E C, D E E A, D C, D, E C, D, E C, D, E C, D D, E C, D, E E C, D, E E D D, E C, D, E E B B D, E E C C, D, E E D D, E E D, E E E D E D, E D C, D, E C C, D, E C, D, E B, C, D, E D E B, C, D, E B, D D B, C, E D, E D, E C, D, E E C, D !* ? ? !* ? ? ? ? ? !* E (z) ? GC Z A A ! (s) GC + O !; K/T ? O ? ? ? A !; F A Z !; K/T ! *; K/T ? (!) E (z) ! A !; K/T !!!; F; K E (n) ? ? O O ? (O) ? ? ! ? ! E (v) A ? (!) ! O !!; K ! ? !; F ! ! *; K/T ! *; K/T Three kaya forests on the south coast of Kenya Family KAYA DIANI Ulmaceae Verbenaceae* Violaceae Vitaceae Asparagaceae Total: 38 151 Plant Species and Subspecies Stratum Category Celtis philippensis Blanco Lantana camara L.* Rinorea arborea (Thouars) Baill. Cissus phymatocarpa Masinde & L.E.Newton Asparagus setaceus (Kunth) Jessop 79 (16 unidentified species omitted) C, E D C, D, E E E ? ? OM !* ? 152 I. Lehmann & E. Kioko Appendix 2. Species list of butterflies recorded in SH by Sevastopulo (1973, 1974a, b) and in our study areas of KM, KK and KD. Nomenclature follows Larsen (1996), Ackery et al. (1995) and Vári et al. (2002). Rare species in bold. Categories 1-8 after Larsen (1996): 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 endemics after Congdon & Collins (1998). Species marked ‘2g*’ were treated as Coastal endemics by Burgess & Clarke (2000, in appendix 9). Species marked ‘F’ are forest-dependent Coastal endemics. Family Butterfly Species and Subspecies Superfamily Papilionoidea Papilionidae Papilio dardanus Brown Papilio constanus Ward Papilio nireus lyaeus Doubleday Papilio ophidicephalus Oberthür Papilio demodocus Esper Graphium angolanus (Goeze) Graphium philonoe (Ward) Graphium leonidas Fabricius F Graphium kirbyi (Hewitson) Graphium colonna (Ward) Graphium polistratus (Grose-Smith) Graphium policenes (Cramer) Graphium antheus (Cramer) Graphium porthaon (Hewitson) Subtotal Pieridae Catopsilia florella (Fabricius) Eurema hecabe solifera (Butler) Eurema floricola orientis (Butler) Eurema brigitta brigitta (Stoll) Pinacopteryx eriphia melanarge (Butler) Nepheronia argia (Fabricius) Nepheronia thalassina (de Boisduval) Nepheronia buquetii buquetii (de Boisduval) Eronia cleodora Hübner Eronia leda (de Boisduval) Colotis protomedia (Klug) Colotis ione (Godart) Colotis regina (Trimen) Colotis hetaera (Gerstaecker) Colotis danae eupompe (Klug) Colotis aurora (Cramer) Colotis auxo (Lucas) Colotis antevippe (de Boisduval) Colotis euippe omphale (Godart) Colotis daira (Klug) Colotis amata (Fabricius) Colotis evagore (Klug) Colotis eris (Klug) Belenois aurota aurota (Fabricius) Belenois creona (Cramer) Belenois gidica (Godart) Belenois thysa (Hopffer) Dixeia orbona vidua (Butler) Dixeia charina (de Boisduval) Dixeia spilleri (Spiller) Appias epaphia orbona (de Boisduval) Appias lasti lasti (Grose-Smith) Appias sabina phoebe (Butler) Leptosia alcesta inalcesta Bernardi Mylothris agathina (Cramer) Mylothris rueppelli rhodesiana Riley SH KM KK KD Category X X X X X X X X X X X X X X 14 X X 0 X X X X X X X X X X X X X X X X X X X X X X X X 0 X X X X X X X X X 0 X X X X 0 0 X X 0 X X 0 9 X X X X 0 X X X X 0 0 X 0 0 X 0 0 0 X 0 0 0 0 X X 0 X X 0 0 X X 0 X X 0 X X X X 0 0 0 0 X 0 0 X X 0 7 X 0 0 X 0 0 0 0 0 0 0 X 0 0 X 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 2a 3 2a; 3 2g 1 4a (?); 2g* 3 2g; 2g* 2g; 2g* 2g; 2g* 2a; 3 2a; 3 2g; 3 X X X X X 0 X 0 X X 0 X X 0 10 X 0 0 X 0 X X 0 X 0 0 X 0 0 X 0 0 X X 0 X 0 0 0 X 0 X 0 X 0 X X X X X 0 1 1 ? 1 4a 2a 2a; 3 4a 4a 4c 4c ? ? 4d 4a 4b 4d 4a ? 4a ? 4a 4a 4a 4a 4a 4d 4a 4d 3; 4d 3 2g 2a 2a 1 3; 4d Three kaya forests on the south coast of Kenya Family Pieridae Lycaenidae F F F F F F F Butterfly Species and Subspecies Pontia glauconome Klug Subtotal Alaena picata Sharpe Pentila tropicalis (de Boisduval) Pentila rogersi rogersi (Druce) Ornipholidotos peucetia peuceda (Grose-Smith) Teriomima subpunctata Kirby Teriomima micra (Grose-Smith) Baliochila hildegarda (Kirby) Baliochila minima (Hawker-Smith) Baliochila latimarginata (Hawker-Smith) Deloneura ochrascens ochrascens Neave Aslauga purpurascens Holland Spalgis lemolea Druce Lachnocnema bibulus (Fabricius) Cigaritis nyassae (Butler) Cigaritis victoriae (Butler) Cigaritis apelles (Oberthür) Cigaritis homeyeri (Dewitz) Chloroselas pseudozeritis tytleri Riley Axiocerses harpax ugandana Clench Axiocerses amanga (Westwood) Axiocerses punicea (Grose-Smith) Iolaus diametra (Karsch) Iolaus silanus silanus Grose-Smith Iolaus mermis (Druce) Iolaus pallene (Wallengren) Iolaus lalos lalos (Druce) Iolaus crawshayi littoralis (Stempffer & Bennett) Hemiolaus caeculus littoralis (Stempffer) Hypolycaena philippus philippus (Fabricius) Hypolycaena buxtoni rogersi Bethune-Baker Leptomyrina hirundo (Wallengren) Deudorix caerulea obscurata Trimen Deudorix antalus (Hopffer) Deudorix diocles Hewitson Deudorix dinochares Grose-Smith Deudorix dinomenes Grose-Smith Deudorix dariaves Hewitson Deudorix lorisona (Hewitson) Anthene lemnos loa (Strand) Anthene lasti (Grose-Smith & Kirby) Anthene liodes (Hewitson) Anthene lunulata (Trimen) Anthene amarah amarah (Guérin-Méneville) Anthene kersteni (Gerstaecker) Cupidopsis jobates jobates (Hopffer) Cupidopsis cissus (Godart) Pseudonacaduba sichela sichela (Wallengren) Lampides boeticus (Linnaeus) Cacyreus lingeus (Stoll) Leptotes pirithous (Linnaeus) Zizeeria knysna (Trimen) Zizula hylax (Fabricius) Actizera lucida (Trimen) Azanus jesous (Guérin-Méneville) Azanus mirza (Plötz) Eicochrysops hippocrates (Fabricius) Euchrysops malathana (de Boisduval) Euchrysops osiris (Hopffer) Euchrysops barkeri (Trimen) 153 SH X 35 X X X X X X X X 0 X X 0 X X X X X X X X X X X 0 X X 0 X X X X X X X X 0 X X 0 X 0 X X X X X X X X X X X X X X X X 0 X KM 0 19 0 X 0 0 X X 0 X X 0 0 X 0 0 X 0 0 0 0 0 X 0 X 0 0 0 0 0 X X 0 X 0 X 0 X 0 X 0 X X 0 0 X 0 0 0 0 0 X 0 X 0 0 0 X X X X KK 0 5 0 X 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X X 0 X 0 0 0 0 0 0 0 0 0 0 X 0 X 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 X 0 0 0 KD 0 18 0 X 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 X 0 X 0 0 0 X X X 0 X 0 X 0 0 0 0 X 0 0 0 0 0 X 0 X 0 0 0 X 0 0 0 Category ? 4d 2g 2g; 2g* 2g; 3 2g; 2g* 2g; 2g* 2g; 3 2g; 2g* 2g; 2g* 4d 4d 2a ? 4d 4d 4d 4d 4d 4b 4a 2g; 2g* 4d 2g; 2g* 2g 4d 2g; 4d 4d 2g 4a 4d 4d 4a 1; 4a 2g 4d ? 2g 4a ? 2g; 2g* 2b 4a 4a 2g 4a 8 4a 1 1 1 1 1 ? 4a 4a 8 4a 4a 4d 154 I. Lehmann & E. Kioko Family Lycaenidae Nymphalidae F F F F F F F Butterfly Species and Subspecies Lepidochrysops peculiaris peculiaris (Rogenhofer) Chilades trochylus (Freyer) Subtotal Libythea labdaca laius Trimen Danaus chrysippus (Linnaeus) Tirumala petiverana (Doubleday) Amauris niavius dominicanus Trimen Amauris ochlea ochlea (de Boisduval) Melanitis leda (Linnaeus) Gnophodes betsimena diversa (Butler) Bicyclus ena (Hewitson) Bicyclus campina ocelligera (Strand) Bicyclus anynana anynana (Butler) Bicyclus safitza safitza (Westwood) Henotesia perspicua (Trimen) Ypthima asterope asterope (Klug) Physcaeneura leda (Gerstaecker) Charaxes varanes vologeses (Mabille) Charaxes acuminatus shimbanus van Someren Charaxes candiope candiope (Godart) Charaxes protoclea azota (Hewitson) Charaxes lasti lasti Grose-Smith Charaxes jasius saturnus Butler Charaxes castor flavifasciatus Butler Charaxes brutus alcyone Stoneham Charaxes bohemani Felder & Felder Charaxes violetta maritima van Someren Charaxes cithaeron kennethi Poulton Charaxes pythodoris nesaea Grose-Smith Charaxes etesipe tavetensis Rothschild Charaxes jahlusa kenyensis Joicey & Talbot Charaxes ethalion littoralis van Someren Charaxes viola picta van Someren & Jackson Charaxes contrarius van Someren Charaxes guderiana rabaiensis Poulton Charaxes pleione oriens Plantrou Charaxes zoolina zoolina (Westwood) Euxanthe wakefieldi (Ward) Euxanthe tiberius tiberius Grose-Smith Euriphene achlys (Hopffer) Bebearia chriemhilda (Staudinger) Bebearia orientis orientis (Karsch) Euphaedra neophron littoralis Talbot Euphaedra orientalis Rothschild Hamanumida daedalus (Fabricius) Aterica galene theophane Hopffer Harma theobene blassi (Weymer) Cymothoe coranus Grose-Smith Euptera pluto kinugnana (Grose-Smith) Pseudathyma lucretioides lucretioides Carpenter & Jackson Pseudacraea eurytus conradti Oberthür Pseudacraea lucretia expansa (Butler) Pseudacraea boisduvalii trimeni Butler Neptis saclava marpessa Hopffer Neptis kiriakoffi Overlaet Neptis alta Overlaet Neptis rogersi Eltringham Neptis trigonophora trigonophora Butler Neptis goochi Trimen Cyrestis camillus sublineata Lathy SH X X 53 X X X X X X X 0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X KM 0 0 24 X X X X X X 0 0 X X X X X X X X 0 0 X 0 0 X 0 X X X 0 0 0 0 X 0 0 X X X X 0 X X 0 0 X 0 X X 0 KK 0 0 9 0 X X X X X 0 X X X X 0 0 0 0 X X 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 X X 0 0 0 0 X X 0 KD X 0 15 0 X X X X 0 0 0 0 0 X X X X X 0 X 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 X X 0 0 0 0 X 0 0 X 0 0 0 0 Category 4d 4a X X X X X X X X X X X X X 0 0 0 0 0 X 0 0 X X 0 0 0 0 0 0 0 0 X 0 X 0 0 0 0 0 0 2a 2a ? 1 2e ? 2g; 2g* 2g 2g 2a 2a 1 1 1 2g 1; 3 2a 4d 4d 4d 1; 3 4d 4a 2g; 2g* 1 5; 2g* 1 2a; 3 2g; 2g* ? 1 3 4d 2g; 2g* 4d (2a); 2g* 2a; 3 4d 4d 4b 2g; 2g* 4d 2a 4d 2g 2g; 2g* 2g 2g; 2g* 3 2g; 2g* 2g; 2g* 4a (2a); 2g* 2a 2g 2g; 2g* 2g; 2g* Three kaya forests on the south coast of Kenya Family Nymphalidae F F F Butterfly Species and Subspecies Sallya garega (Karsch) Sallya amulia rosa (Hewitson) Byblia ilithyia (Drury) Byblia anvatara acheloia (Wallengren) Neptidopsis fulgurata platyptera Rothschild & Jordan Eurytela dryope angulata Aurivillius Hypolimnas misippus (Linnaeus) Hypolimnas deceptor deceptor (Trimen) Hypolimnas anthedon wahlbergi (Wallengren) Hypolimnas usambara (Ward) Salamis parhassus (Drury) Salamis anacardii (Linnaeus) Salamis cacta amaniensis Vosseler Junonia orithya madagascariensis Guenée Junonia oenone oenone (Linnaeus) Junonia hierta cebrene Trimen Junonia natalica natalica (Felder & Felder) Junonia terea elgiva Hewitson Junonia antilope (Feisthamel) Catacroptera cloanthe cloanthe (Stoll) Cynthia cardui (Linnaeus) Lachnoptera ayresii Trimen Phalanta phalantha aethiopica (Rothschild & Jordan) Phalanta eurytis eurytis (Doubleday) Acraea aubyni Eltringham Acraea encedon encedon (Linnaeus) Acraea esebria esebria Hewitson Acraea eponina (Cramer) Acraea petraea de Boisduval Acraea egina areca Mabille Acraea braesia Godman Acraea equatorialis anaemia Eltringham Acraea oncaea Hopffer Acraea pudorella pudorella Aurivillius Acraea natalica de Boisduval Acraea zonata Hewitson Acraea rabbaiae mombasae Grose-Smith Acraea satis Ward Acraea zetes acara Hewitson Acraea chilo chilo Godman Acraea anemosa Hewitson Acraea boopis ama Pierre Acraea quirina rosa Eltringham Acraea cuva cuva Grose-Smith Acraea insignis insignis Distant Acraea neobule neobule Doubleday Acraea matuapa Grose-Smith Acraea adrasta adrasta Weymer Acraea aganice Hewitson Acraea epaea epitellus Staudinger Pardopsis punctatissima (de Boisduval) Subtotal Superfamily Hesperioidea Hesperiidae Coeliades libeon (Druce) Coeliades anchises anchises (Gerstaecker) Coeliades forestan forestan (Stoll) Coeliades pisistratus (Fabricius) Coeliades sejuncta (Mabille & Vuillot) Celaenorrhinus galenus (Fabricius) Tagiades flesus (Fabricius) 155 SH X 0 X X X KM 0 0 0 X 0 KK 0 X 0 0 0 KD 0 0 0 X 0 Category ? ? 4a 4a 2g; 2g* X X X X X X X X X X X X X X X X X 0 X X X X X X X X X X X X X X X X X X X X X X X X X X X X 105 X X X 0 X X X X 0 X 0 X X 0 0 0 X X 0 X 0 X X 0 X X 0 0 0 X X X X 0 0 0 X X 0 0 0 X 0 X 0 X 60 X X 0 0 X X 0 0 0 X 0 0 0 0 0 0 0 X X 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 28 X X 0 0 0 0 X 0 0 X 0 X X 0 0 0 0 X 0 0 0 0 X 0 0 0 0 0 0 X X X 0 0 0 X 0 0 0 0 0 0 0 0 0 0 30 1 1; 4a 2g 2a 2g; 2g* 2a 2a; 3 ? 4a 1 4a 3; 4d 2a; 3 4d 4a 4a 2g 1 2a 2g; 2g* 1; 4a 3; 4d 1 2g 4a 4c 4c 4d 4d 3; 4d (4d); 2g* 2g 2g 3; 4a 4c 3; 4d 2g 2a 2g; 2g* 4d 1; 4a 2g; 2g* 2g; 2g* 3; 4d (2a) 2g* ? X X X X X X X 0 X X 0 0 X X 0 0 0 0 0 0 0 0 0 0 0 X 0 0 2a 4d 1 1 2g 2a 2a 156 I. Lehmann & E. Kioko Family Hesperiidae F Butterfly Species and Subspecies Eagris sabadius ochreana Lathy Eagris nottoana nottoana (Wallengren) Sarangesa motozi (Wallengren) Sarangesa maculata (Mabille) Caprona pillaana Wallengren Netrobalane canopus (Trimen) Abantis paradisea (Butler) Spialia kituina (Karsch) Spialia spio (Linnaeus) Spialia diomus diomus (Hopffer) Spialia confusa obscura Evans Spialia dromus (Plötz) Spialia zebra bifida (Higgins) Gomalia elma elma (Trimen) Astictopterus stellata stellata (Mabille) Ampittia capenas capenas (Hewitson) Gorgyra subflavidus Holland Gorgyra diva Evans Gorgyra johnstoni (Butler) Pardaleodes incerta (Snellen) Teniorhinus herilus (Hopffer) Acada biseriata (Mabille) Parosmodes morantii morantii (Trimen) Paracleros biguttulus (Mabille) Acleros ploetzi Mabille Acleros mackenii (Trimen) Semalea arela (Mabille) Andronymus neander neander (Plötz) Andronymus caesar philander (Hopffer) Zophopetes nobilior (Holland) Artitropa reducta Aurivillius Artitropa erinnys radiata Riley Fresna nyassae (Hewitson) Pelopidas thrax inconspicua (Bertoloni) Borbo fatuellus fatuellus (Hopffer) Borbo lugens (Hopffer) Borbo detecta (Trimen) Borbo ferruginea ferruginea (Aurivillius) Borbo borbonica borbonica (de Boisduval) Subtotal TOTAL SH X 0 X X X X X X 0 X X X X X X X X X X X 0 X 0 X X X X X X 0 X X X 0 X X 0 X X 39 246 KM 0 0 0 0 0 0 0 0 X 0 0 X 0 0 X 0 0 0 0 X X 0 X 0 0 0 0 0 X X 0 0 0 X 0 X X 0 0 15 127 KK 0 X X 0 X 0 0 0 0 0 0 0 0 X 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 0 X 0 7 56 KD 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 X 0 0 0 4 77 Category 4d 2g 4d 2e 4d 4d 4d 4c; 9 ? 4a (4d); 2g* 4a 4c 1 2g 4d 2g; 2g* (?); 2g* 2g 2a 2g 4d ? 2a 2a 2a 2a ? 2a ? ? 4d 2b 1 2a 4d 4d 2g; 2g* 4a Three kaya forests on the south coast of Kenya 157 APPENDIX 3. Species list of larger moths recorded in the study areas of KM, KK and KD. Nomenclature follows those of the collection in the BMNH (London) as well as Carcasson (1976), Fletcher & Nye (1982), Poole (1989), Goodger & Watson (1995), Häuser & Boppré (1997), Scoble (1999) and Vári et al. (2002). Rare species of the Kenya coast in bold; species marked ‘F’ are forest-dependent Coastal endemics (after literature notes, museum material, own records 1994-2003). Family Superfamily Cossoidea Cossidae Metarbelidae Limacodidae F Moth Species KM KK KD Azygophleps sp. Brachylia terebroides Felder Metarbela ?dialeuca Hampson Metarbela haberlandorum Lehmann Metarbela latifasciata Gaede Salagena ? irrorata Le Cerf Salagena tessellata Distant Salagena ? sp. nov. Teragra simillima Hampson Ectropa sp. nov. Micraphe lateritia Karsch Scotinocerides conspersa (Kirby) (Coastal endemic) Subtotal X X X X X X X X X 0 X X 11 0 0 0 0 0 0 0 0 0 X 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 X X X 0 0 X 0 X X X X X X X X X 0 X X 0 X X X X X 0 X X X 0 X X X X X X 0 X 0 X 0 0 0 0 X 0 X 0 0 X 0 0 X X X X 0 X 0 X 0 0 0 X X X 0 0 X X X 0 X 0 0 0 X 0 X 0 0 0 0 X 0 0 0 0 0 0 0 0 X 0 X 0 X 0 0 0 0 0 0 X 0 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 Superfamily Geometroidea Geometridae Acanthovalva ?inconspicuaria (Hübner) Allochrostes biornata Prout Antharmostes papilio papilio Prout Cartaletis libyssa (Hopffer) Chiasmia feraliata (Guenée) Chiasmia orientalis Krüger (first Kenyan record) Chiasmia sororcula (Warren) Chrysocraspeda leighata leighata Warren Cleora munda (Warren) Cleora rothkirchi rothkirchi (Strand) Colocleora ?comoraria Oberthür Colocleora divisaria divisaria (Walker) Comibaena esmeralda (Warren) Comibaena rufitornus Prout Conolophia conscitaria conscitaria (Walker) Erastria albosignata albosignata Walker Erastria leucicolor leucicolor (Butler) Ereunetea reussi reussi Gaede Eucrostes disparata (Walker) Geolyces ? sp. nov. Gyalomia elatina Prout Heterostegane ? sp. nov. Isoplenia trisinuata Warren Isturgia catalaunaria (Guenée) Isturgia supergressa (Prout) Metallochlora grisea Prout Miantochora sp. nov. Mixocera viridans Prout Omizodes rubrifasciata (Butler) Ozola ? sp. nov. Paraptychodes tenuis Butler (Coastal endemic) Phaiogramma ? faustinata (Milliére) Pingasa rhadamaria alterata (Walker) Pingasa ? sp. nov. Pitthea trifasciata Dewitz Plegapteryx anomalus Herrich-Schäffer Problepsis digammata Kirby Racotis apodosima Prout Racotis breijeri (Prout) (first Kenyan record) Racotis squalida squalida (Butler) 158 I. Lehmann & E. Kioko Family Geometridae F Superfamily Uranioidea Uraniidae Moth Species Scopula addictaria (Walker) Scopula atricapilla atricapilla Prout Scopula cassioides Prout Scopula donovani (Distant) Scopula lactaria (Walker) Scopula minorata minorata (de Boisduval) Scopula ? ossicolor (Warren) Scopula ? paradelpharia Prout Traminda neptunaria (Guenée) Traminda obversata obversata (Walker) Traminda vividaria (Walker) Xenochroma ? palimpais Prout Zamarada ? crystallophana Mabille Zamarada ? cucharita Fletcher Zamarada euphrosyne Oberthür Zamarada keraia Fletcher (Coastal endemic; K only?) Zamarada ? plana denticincta Hampson Zamarada rufilinearia Swinhoe Zamarada sp. nov. Subtotal KM X X X 0 X X X X X 0 X X X X X X X X X 48 KK 0 0 0 X X 0 0 0 X X 0 0 0 0 0 0 0 0 0 22 KD 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 7 Acropteris erycinaria (Guenée) Epiplema barbara Warren Leucoplema dohertyi (Warren) Subtotal X X X 3 X 0 X 2 0 0 X 1 X X X X X X X X X 0 X X X X X X X 16 0 0 0 0 0 0 0 0 0 X 0 0 0 0 0 0 X 2 0 0 0 0 X 0 0 X 0 0 0 0 0 0 0 0 X 3 X X X X X X 0 X X X X X 0 0 X X 0 0 0 X X X 0 0 0 0 0 X X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 Superfamily Bombycoidea Lasiocampidae Beralade continua Aurivillius Mallocampa ? leighi Aurivillius Mimopacha tripunctata (Aurivillius) Eupterotidae Vianga sp. nov. Saturniidae Decachorda aspersa Bouvier (? ssp. ) Imbrasia zambesina (Walker) Sphingidae Acherontia atropos (Linnaeus) Andriasa contraria Walker Centroctena imitans (Butler) Euchloron megaera (Linnaeus) Neopolyptychus compar septentrionalis Carcasson Nephele bipartita Butler Nephele funebris (Fabricius) Nephele peneus (Cramer) Praedora marshalli tropicalis Rothschild & Jordan Pseudoclanis postica postica (Walker) Temnora marginata (Walker) Subtotal Superfamily Noctuoidea Notodontidae Eurystaura griseitincta Hampson Odontoperas ? sp. nov. Paracleapa psecas Druce ? Peratodonta extensa Gaede Thyretidae Automolis helga (Kiriakoff) Automolis rufescens (Walker) Thyretidae Automolis ? sp. nov. Lymantriidae Conigephyra pallidula (Hering) (Coastal endemic) Cropera testacea Walker Dasychira barbara Hübner Dasychira callipepla Collenette (Coastal endemic) Euproctis aethiopica Snellen Euproctis pygmaea Walker Euproctis ? rubricosta Fawcett Euproctis rufopunctata (Walker) Three kaya forests on the south coast of Kenya Family Lymantriidae Arctiidae Arctiidae Herminiidae Aganaidae Noctuidae Moth Species Hemerophanes xanthopa Collenette Knappetra fasciata (Distant) Laelia extorta (Distant) Leucoma parva (Plötz) Lymantria leucerythra Collenette (first Kenyan record) Marblepsis macrocera Sharpe Stracena bananae (Butler) Amata phoenicia (Hampson) Amerila phaedra Weymer Amerila vitrea Plötz Amphicallia bellatrix (Dalman) Argina amanda (de Boisduval) Argina astrea (Drury) Asura anticraspeda Hampson Eilema distigmata Hampson Eyralpenus melanocera (Hampson) Nyctemera restrictum (Butler) Secusio drucei Rothschild Secusio strigata Walker Spilosoma lineatum Walker Spilosoma lutescens Walker Spilosoma ? sp. nov. Utetheisa pulchella (Linnaeus) Xanthetis ichorina (Butler) Alelimma pallicostalis Hampson Progonia ? aenicta Fletcher Simplicia extinctalis (Zeller) Asota speciosa (Drury) Digama africana Swinhoe Soloe tripunctata Druce Soloe sp. nov. Aburina poliophaea Hampson Aburina sobrina Möschler Acantholipes trimeni Felder & Rogenhofer Achaea catella Guenée Achaea dasybasis Hampson Achaea faber Holland Achaea lienardi (de Boisduval) Achaea mercatoria (Fabricius) Acontia nitidula (Fabricius) Amyna axis Guenée Amyna punctum (Fabricius) Androlymnia torsivena (Hampson) Anoba atriplaga (Walker) Anoba hamifera (Hampson) Anoba ? sp. nov. Anomis flava (Fabricius) Anomis leona (Schaus & Clements) Anomis punctulata (Holland) Anomis sabulifera (Guenée) Anomis simulatrix (Walker) Anticarsia irrorata (Fabricius) Attonda adspersa (Felder & Rogenhofer) Axiopoeniella sp. nov. ? Baniana sp. nov. Blenina quadripuncta Hampson Bryophilopsis tarachoides Mabille Calesia zambesita Walker Caligatus angasii Wing Caryonopera breviramia Hampson Cerynea thermesialis (Walker) 159 KM X X X X X X X X X X 0 0 0 X X X X X X 0 0 X 0 X X X X X X X X 0 X X 0 X 0 X 0 X 0 X X X X X X X X X X X X X X X 0 0 X X X KK 0 0 X 0 0 0 0 0 X 0 X X X X 0 0 0 0 X X X 0 0 0 X 0 0 X X X 0 0 0 X X 0 X X X 0 X X 0 0 0 X X X X X X 0 X 0 0 X X 0 0 0 X KD 0 0 0 0 0 0 0 0 X 0 0 X 0 0 0 0 0 0 X 0 0 0 X 0 0 0 0 X X 0 0 X 0 0 0 X 0 X 0 0 0 X 0 0 0 0 0 0 0 X 0 0 X 0 0 X 0 X 0 0 0 160 Family Noctuidae I. Lehmann & E. Kioko Moth Species Chasmina tibialis (Fabricius) Corgatha ozolica Hampson Cyligramma limacina (Guérin-Méneville) Diparopsis castanea Hampson Dysgonia conjunctura (Walker) Dysgonia torrida (Guenée) Egnasia vicaria (Walker) Egybolis vaillantina (Stoll) Enispa flavitincta Hampson Entomogramma pardus Guenée Ercheia subsignata (Walker) Erebus walkeri (Butler) Ericeia congregata (Walker) Eublemma ? aurantiaca Hampson Eublemma baccalix (Swinhoe) Eublemma ? sp. nov. Eudocima divitiosa (Walker) Eudocima materna (Linnaeus) Eutelia amatrix Walker Eutelia musicalis Berio Facidia vacillans (Walker) Facidina semifimbria (Walker) Fodina embolophora Hampson Geniascota patagiata Hampson Gesonia obeditalis Walker Giria pectinicornis (Bethune-Baker) Gracilodes caffra Guenée Gracilodes nysa Guenée Grammodes congesta Berio (first Kenyan record) Grammodes stolida (Fabricius) Heliophisma xanthoptera (Hampson) Hypena laceratalis Walker Hypena masurialis Guenée Hypena varialis Walker Lacera alope (Cramer) Lamprolopha melanephra Hampson Lophocrama phoenicochlora Hampson Lophoptera litigiosa (de Boisduval) Lophotavia globulipes (Walker) Marathyssa cuneata (Saalmüller) Marcipa insulata (Walker) Marcipa mediana Hampson Marcipa pyramidalis (Hampson) Maxera marchalii (de Boisduval) Mazuca strigicincta Walker Metaleptina nigribasis Holland (first Kenyan record) Mocis conveniens (Walker) Mocis mayeri (de Boisduval) ? Nagia microsema Hampson Nola chionea Hampson Odontestis fuscicona (Hampson) Oglasa nana (Walker) Oglasodes nyasica Hampson (first Kenyan record) Ogovia sp. Oligia ? sp. nov. Oligia ? sp. nov. Ophiusa gonoptera Hampson Ophiusa sp. nov. Oraesia emarginata (Fabricius) Ozarba abscissa (Walker) Ozarba accincta (Distant) KM X X X X X 0 X X X X X X X 0 X X X X X X X 0 X X X X X X X 0 X X X X 0 X X X X X X X X X X X X X 0 X X 0 X X X 0 X X X X 0 KK 0 0 X 0 0 X 0 X 0 0 X X X X X 0 0 0 0 0 X X 0 0 0 0 X 0 0 0 0 0 0 0 X 0 0 0 X 0 0 0 X 0 0 0 X 0 X 0 0 X 0 0 0 X X 0 0 0 X KD 0 0 X 0 0 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 X 0 0 X 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Three kaya forests on the south coast of Kenya Family Noctuidae Moth Species Ozarba domina (Holland) Ozarba heliastis (Hampson) Ozarba perplexa Saalmüller Pantydia sp. Paralephana argyresthia Hampson Parasada sp. Phytometra ? sp. nov. Plecoptera aspila (Hampson) Plecoptera flavilinea Hampson Plecoptera melanoscia Hampson Plecoptera rufirena (Hampson) Plecoptera tripalis (Wallengren) Plusiodonta commoda Walker Pseudogiria polita Berio Pteronycta fasciata Fawcett Radara subcupralis (Walker) Rhanidophora agrippa Druce Rhanidophora cinctigutta (Walker) Rhesala moestalis (Walker) Rhesala sp. Rhynchina ?leucodonta Hampson Rhynchina revolutalis (Zeller) Rhynchina sp. Rougeotiana xanthoperas (Hampson) ? Saaluncifera uncinata (Saalmüller) Selepa ? transvalica Hampson Serrodes partita (Fabricius) Sphingomorpha chlorea (Cramer) Spodoptera cilium Guenée Spodoptera exempta (Walker) Spodoptera exigua (Hübner) Spodoptera littoralis (de Boisduval) Stictoptera confluens (Walker) Tatorinia fumipennis (Felder & Rogenhofer) Taveta eucosmia Hampson Tavia instruens Walker Tavia nycterina (de Boisduval) Tephrialia vausema Hampson Thyas arcifera (Hampson) Thyatirina achatina (Weymer) Trichopalpina ? sp. nov. Trigonodes hyppasia (Cramer) Ugia amaponda (Felder & Rogenhofer) Ugia sp. nov. Ulotrichopus primulinus (Hampson) Subtotal TOTAL 161 KM X X 0 X X X X X 0 X 0 X X X X X X X 0 X X X 0 X X X X 0 X X X X X X X 0 X X 0 X X 0 X 0 X 147 225 KK 0 0 X 0 0 0 0 X X 0 X 0 0 0 0 0 0 0 X 0 0 0 X 0 0 0 0 X 0 X 0 X 0 0 0 X X 0 X 0 0 X X X 0 71 98 KD 0 0 0 0 0 0 0 0 0 0 0 X 0 0 0 X 0 0 X 0 0 0 0 0 0 0 0 0 0 X 0 X 0 0 0 0 0 0 X 0 0 0 0 0 0 25 36 162 I. Lehmann & E. Kioko Plate 1. A. Lymantria leucerythra (ex KM, 17 January 1998) is presented as a new species for Kenya. Few other records suggest that this species occurs also in southeast Uganda, eastern Tanzania and southeast Malawi but it is currently not known from southern Africa. B. Rougeotiana xanthoperas (ex KM, 12 July 1997) is classified here as a forest species centred on the Guineo-Congolian region. Several museum specimens suggest that the species is linked to wetter forests. It is currently not known from southern Africa. Only one other Kenyan record was found which is a specimen in the NMK’s collection (ex Shimba Hills). C. Paraptychodes tenuis (ex KD, 21 March 2002) is classified here as a coastal endemic species. The genus has probably a disjunct distribution between eastern Africa and the Guineo-Congolian region and has in southern Africa so far only been recorded recently from southern Mozambique. D. Oglasodes nyasica (ex KM, 29 January 1996, no other record) is presented as a new species for Kenya and is classified here as a potential coastal endemic species which is probably rare. Only two specimens were seen so far, both are in the BMNH, collected at 700 m on the foothills of Mount Mulanje in 1913, southeast Malawi. Of interest is, that Mount Mulanje has patches of wetter evergreen and semi-evergreen lowland forest at this altitude and that the area belongs to the western limit of the eastern African Coastal Forests (Swahilian/ Maputaland regional transition zone). The species is currently known only from these two localities within the Coastal Forest belt. The genus occurs also in West Africa but is not known from southern Africa. E. A forest stand in the eastern part of KK having a mixed species composition in strata B-E in the background. In the foreground is, in contrast, a single dominant forest patch of the tree Sorindeia madagascariensis. A characteristic feature is the absence of stratum D. Stratum E is represented mainly by the grass Oplismenus burmannii. Local dominance of only one tree species in natural forest most probably indicates an undisturbed forest stand. Photographs by Ingo Lehmann Three kaya forests on the south coast of Kenya E View publication stats 163