taxonID	type	description	language	source
6F3587EC322A1B4CAB9F5836FC6AE61A.taxon	description	This is a Madagascan endemic genus of about 15 described species, including the synonymised genus Miraja (Lees et al. 2003); considering material collected to date, the radiation is probably at least twice this size (D. C. Lees, pers. comm. 2013). In a future paper, we will report that P. albiplaga (Oberthür) and P. ismael (Oberthür) are bamboo feeders. Here we present limited information on a species reared from palm.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC322A1B4CAB9F5B10FAACE71F.taxon	description	An ABRI collector reared this species from an unidentified palm at Marhamiiza, near Andasibe, 25 Dec 2002. The emerged pupa in ABRI is brown; flattened frontally; T 1 spiracle slightly raised; proboscis 3 mm beyond wing cases; shelter lightly covered with white wax. There is no associated final instar caterpillar head capsule.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC322A1B4CAB9F5D2FFB02E38E.taxon	description	Mabille (1877) described Hesperia amygdalis in a footnote to his paper on the Hesperiidae of Congo, based on two specimens from Nosy Bé (as Nossi-Bé) and Madagascar. Adults have been collected at light more recently at Nosy Bé (specimens in ABRI). Published reports are restricted to the north and east of Madagascar. Food plants. There have been reports of this species feeding on coconut, Cocos nucifera (Viette 1956, Mariau 2001), but the life history has not previously been documented. ABRI collectors have found caterpillars on C. nucifera on Île Ste Marie, off the east coast of northern Madagascar (Figure 1 – 2), as well as a caterpillar and pupa on the indigenous golden palm, Dypsis lutescens, at Antsiranana (Diego-Suarez) and Montagne des Français respectively, in the far north, and emerged pupae (1 ♂, 2 ♀) on a Raphia sp. palm at Ranomafana, in the south-east of Madagascar. Life history. The ova are laid in longitudinal rows on a palm leaflet, touching each other. They are 1.9 mm in diameter (9 eggs = 17 mm), dome-shaped with about 54 fine ribs. The eclosing caterpillar eats most but not all the shell apart from the base. The caterpillars (Figure 2.1 – 2) are unusual in that they are gregarious in the early instars (Figure 2.2) and do not make leaf shelters. In the final instar (Figure 2.1) the caterpillars disperse, and make individual rolled leaflet shelters, in which they pupate (Figure 1.1). The pupa (Figure 2.3) is 29 – 32 mm long, brown, smooth, without projections, the proboscis barely projecting beyond the wing cases. The inside of the shelter and the pupa are both lightly covered with white waxy powder. Natural enemies. In one row of nine eggs, four had emergence holes of an egg parasitoid; in another of seven eggs all emerged normally. Pupae were parasitized by a large, solitary yellow-brown ichneumonid parasitoid; it emerged by pushing off the head of the pupa along suture lines, rather than cutting a hole in the cuticle. Other fieldcollected pupae had round parasitoid emergence holes compatible with a Brachymeria sp.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC322A1B4CAB9F5A1FFDD0E000.taxon	description	The genus Ploetzia was established by Saalmüller (1884) for the single species P. amygdalis (Mabille). Amongst African Hesperiidae, this is one of very few species with gregarious caterpillars (Pteroteinon laufella, below, is another), supporting its unique position.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC322B1B4DAB9F5CCFFD4BE370.taxon	description	This small genus has been considered to comprise five species of large skippers (Ackery et al. 1995). Lindsey (1925) made Z. dysmephila (Trimen) the type species of the genus. All food plant records from this genus are from palms. In the French West African literature on palm pests, early records of Z. dysmephila appear to have been misidentifications for Z. cerymica (Hewitson), rather than Z. quaternata (Mabille), which is closest in appearance (see below). Zophopetes cerymica replaces Z. dysmephila in more recent publications on palm pests (Mariau et al. 1981, Mariau 2000, 2001), and the descriptions of the early stages (e. g. Mariau & Morin 1974) match Z. cerymica rather than Z. dysmephila. Thus, the records of Z. dysmephila from oil palm (Elaeis guineensis) at Anguédedou, Côte d’Ivoire (Genty 1968), and as pests of oil palm and coconut in West Africa (Mariau & Morin 1974) are considered to be of Z. cerymica. However, the record of Z. dysmephila from Phoenix reclinata at Lamto (Vuattoux 1999), is referable to Z. quaternata (Larsen 2005).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	This sexually dimorphic species was described as Pamphila dysmephila by Trimen (1868) from South Africa (type locality ‘ Bashee River, Kaffraria’, i. e. Mbhashe River, Eastern Cape Province) based on a female. The male was subsequently described as Hesperia schulzi Plötz (1882) from Angola and as Hesperia mucorea Karsch (1892) from Cameroon. Evans (1937) treated these two names as synonyms of Z. dysmephila, and this view has not been questioned. In preparing this paper, it became apparent that there is geographical variation in the caterpillars and male genitalia, suggesting either a complex species or a species complex. The first illustration of the male genitalia is that of Murray (1959) based on a specimen from Port Elizabeth (Eastern Cape Province, and hence representative of the type locality), which shows more or less symmetrical tapered, pointed valves. Examination of the BMNH collection suggests that W. H. Evans examined the male genitalia of just one specimen, which was from ‘ Zululand’ (largely overlapping with KwaZulu-Natal Province, north of the type locality). Although Evans (1937) shows only one valve in his diagram, his preparation shows that the valves are asymmetrical, the left valve resembling that shown in Evans’ figure, but the right more like those shown by Murray (1959). Larsen (2005, Figure 7.21) illustrates the right valve of a Nairobi specimen, which has a strongly upturned apical portion, differing from the earlier illustrations.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	materials_examined	In preparing this paper, MJWC compared the dissected male genitalia of single Kenyan specimens from Kakamega Forest (western Kenya, west of the Rift Valley), Nairobi (central Highlands, east of the Rift Valley) and Bamburi at the coast. The Nairobi specimen was clearly very different from the other two, resembling Larsen’s (2005) figure, with the apical portion of the valves turned upwards and inwards. Brushing the scales off the ventral surface of male genitalia, particularly if the genitalia are extruded when preparing fresh specimens, will reveal this character in many pinned specimens and showed that all six other MJWC specimens from Nairobi have valves of this type, as does one from Nyeri, Central Highlands, Kenya, in the Hope Entomological Collections, Oxford University Museum (HEC). This form is referred to as ‘ Nairobi typical’ below. The specimens from Kakamega Forest and Bamburi are similar to each other, but still show potentially significant differences, e. g. the right valve of the Bamburi specimen is more elongate with a distinctly rounded apex, compared to the pointed apex of the specimen from Kakamega Forest. A male from Eldoret, western Kenya, in HEC probably matches those from Kakamega Forest — certainly it is not of the Nairobi form. A male from Eritrea in the BMNH has the male genitalia displayed in situ and appears to match the Bamburi male. However, T. B. Larsen (pers. comm. 2013) has dissected males from various localities and finds substantial variation and no clear pattern. Thus, apart from the series from Nairobi, no material in series has been examined from any localities, and although the male genitalia of the Nairobi form seem constant, this may not be the case elsewhere. As yet no female genitalia have been documented. The taxonomy of this ‘ species’ needs further research. Here, we provide information on the early stages as we and others have observed them in different parts of the range of Z. dysmephila. The main population stretches from the coastal areas of South Africa, as far west as Western Cape Province (Henning et al. 1997) to most of eastern Africa, north to Kenya and Uganda, but not much west of the Albertine Rift, although MJWC found early stages at Butare, Rwanda (90 / 202). It seems commonest in South Africa, but as indicated below, its apparent general scarcity may be more apparent than real. The type localities of the synonyms schulzi from Angola and mucorea from Cameroon lie outside this area, which as T. B. Larsen (pers. comm. 2013) points out, is puzzling. There are three or four specimens of typical dysmephila from Bom Jesus, Angola which might well have been imported with palms from Mozambique (T. B. Larsen pers. comm. 2013). Indeed, the movement of ornamental Phoenix spp. is not rare and populations of Z. dysmephila may have been redistributed with them, both within and occasionally between countries.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	In Kenya and Tanzania this species is probably to be found wherever its main food plants, Phoenix spp. palms, grow (or are grown as ornamentals). It is particularly common in Nairobi where many P. reclinata have been planted in gardens and public places, and many of these will have at least one leaf shelter, often several.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	The margin of the apical part of the male forewing upper surface is white, an unusual feature in Hesperiidae, but one shared with Carystoides a Neotropical genus where again only males show it (Evans 1955, Cock 2005), and some species of the Oriental genus Erionota, e. g. E. hiraca (Moore) (Swinhoe 1912 – 1913, Evans 1949 as E. acroleucus (Wood-Mason & De Nicéville )) and E. surprisa De Jong & Treadaway (1993). The white band of the antenna club is also conspicuous and reminiscent of the white antennae of Chondrolepis spp., which are pure white above in the male and have the club partly or completely white above in the female (Evans 1937, De Jong 1986). All four genera fly at dusk and this colouring may well be a signalling device, for mate location and acceptance, as suggested by De Jong (1986) for Chondrolepis. Food plants. This is one of the first skipper life histories to have been documented from Africa. Trimen (1889) reports J. H. Bowker’s 1881 observations of the caterpillar, pupa and food plant from South Africa, based on A. D. Millar’s discoveries at Durban (= D’Urban). The food plant reported is the indigenous Phoenix reclinata grown as a garden plant. It is not clear whether subsequent records of P. reclinata from other countries represent local observations, independently reporting this food plant, or simply repeat the earlier information from South Africa. However, locally attributable records from Uganda (Le Pelley 1959), Kenya (MJWC, SCC), Tanzania (TCEC) and Zambia (TCEC) suggest that P. reclinata is a normal food plant throughout this skipper’s range, and this probably represents the original food plant. Other species of Phoenix are also reported as food plants. The introduced ornamental P. canariensis is a suitable food plant, and Jannone (1948) reported Z. dysmephila as a pest on this palm in Eritrea, where he considered it to be the preferred food plant (la specie … che viene particolarmente presa di mira). An introduced population has also been reported on this palm in Cape Town (Geertsema 1985) although a subsequent report by Claassens & Dickson (1986) referred to date palms or common date palms –– by implication P. dactylifera, although this was not made explicit. MJWC found caterpillars on a palm provisionally identified as P. canariensis at Diani Beach, Kenya (95 / 102). Dickson & Kroon (1978) report that C. G. Clark ‘ frequently obtained larvae from the leaves of the common date palm, P. dactylifera ’ at Port Elizabeth, South Africa, whereas Pringle et al. (1994) refer to the common date palm, without a scientific name, and then list the food plants as P. ‘ reclinata (wild date palm) ’ and P. ‘ dactylifera (date palm) ’. Jannone (1948) reported that in Eritrea, in the proximity of infested P. canariensis, the palms Chamaerops humilis (a Mediterranean species) and P. dactylifera are also frequently attacked (riscontrandosi frequentemente nelle prossimità di Phoenix canariensis più o meno infestate dalla specie in questione, esemplari inquinati e danneggiati di Chamaerops humilis e Phoenix dactylifera). In a similar observation, A. J. M. Claassens found caterpillars in Cape Town ‘ on P. canariensis and another palm, tentatively identified as Chrysalidocarpus (Areca) lutescens [i. e. Dypsis lutescens], the butterfly palm, standing between specimens of P. canariensis ’ (Claassens & Dickson 1986). MJWC recently found empty shelters (Figure 4.1) and feeding on P. reclinata (Figure 4.2), P. sylvestris (an Indian species, Figure 4.3) and Chamaerops humilis (Figure 4.4) planted on the campus of the World Agroforestry Centre (ICRAF), Gigiri, Nairobi (15 Nov 2013, MJWC 13 / 03 – 04), which can be attributed to Z. dysmephila, as no other palm-feeding skippers occur in the Nairobi area. In addition to Phoenix spp., Van Someren (1974) lists Raphia spp. and Borassus spp. (Arecaceae) as food plants for East Africa. Probably the latter two can be referred to R. farinifera (raffia palm) and B. aethiopum (borassus or African fan palm), the only indigenous members of these genera in Kenya (Beentje 1994). Sevastopulo (1975) lists Raphia, but did not rear this species himself (Sevastopulo unpublished). These food plants have been repeated by subsequent authors. However, in Tanzania TCEC has confirmed R. farinifera to be a food plant, but not B. aethiopum. Although TCEC has collected caterpillars from B. aethiopum, he has never found any Hesperiidae on it. Accordingly we consider the record from Borassus likely to be an error. As discussed under Zophopetes above and Z. cerymica below, early records from French West Africa of Z. dysmephila as a pest of oil palm and coconut (Genty 1968, Mariau & Morin 1974) are misidentifications for Z. cerymica, which have been repeated in the economic literature (e. g. Lever 1969, Howard & Abad 2001). Larsen (1991) includes Cocos (i. e. C. nucifera, the only species of the genus) as a food plant of Z. dysmephila. We have not found Z. dysmephila feeding on coconut in Kenya or elsewhere. To summarise, the original food plant is most probably the indigenous P. reclinata. Raphia farinifera is used as a food plant, but we do not know if this behaviour is widespread or common. The introduced P. canariensis is an acceptable food plant, but it is not clear how regularly the introduced P. dactylifera and P. sylvestris are used when available. Carter’s (1992) observation that the caterpillar ‘ feeds on date palm (Phoenix dactylifera) and other palms’ is therefore slightly misleading. The records on Chamaerops humilis and D. lutescens may represent spillover effects to less-preferred or marginal hosts from populations on Phoenix spp. Records from Cocos nucifera and Borassus sp (p). are not accepted at this time. The suitability of date palm, P. dactylifera, should be further evaluated, since if this important crop is readily accepted, Z. dysmephila would be a significant quarantine pest for the date-growing areas of the world. Life history. Not only was Z. dysmephila one of the earliest African skippers to have its food plant documented, it was also one of the first to have its life history documented and illustrated. Trimen (1889) included a brief description of the caterpillar, pupa and shelters. A few years later, Kelley (1912) described and illustrated the caterpillar and pupa on P. reclinata at Pietermaritzburg (= Maritzburg), KwaZulu-Natal Province, South Africa. Jannone’s (1948) account of this species as a pest of P. canariensis in Eritrea includes good drawings of all stages, shelters and damage. These two accounts are synthesised briefly by Lepesme (1947). G. C. Clark (in Dickson & Kroon 1978) provided his usual excellent detailed paintings of the complete life history. Here we document the history from Kenya, supported by observations from Rwanda, Tanzania and Zambia.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	Ovum. MJWC found the ovum of the Nairobi population is dark red when newly laid and turns white; it is hemispherical 1.7 mm wide (range 1.61 – 1.76, n = 7) x 1.1 mm high (range 1.06 – 1.18, n = 4), with 27 – 29 fine ribs which peter out close to the flattened micropyle. One ovum from Kakamega Forest, Kenya, measured 1.57 x 0.98 mm wide x high. Ova found at Butare, Rwanda measured 1.6 mm wide (range 1.49 – 1.65, n = 21) x 1.0 mm high (range 0.98 – 1.06, n = 5), with 34 – 38 (n = 3) fine ribs. In Eritrea, Jannone (1948) reports the ova to be 1.4 – 1.7 x 1.0 mm wide x high. Ova are laid on palm leaflets, usually singly, on the upper surface near the apex, or in the upper surface hollow at the base of the leaflet. MJWC found that normally ova are laid singly or in pairs, although once a group of four ova was found at Nairobi (90 / 24) and a group of 24 at Butare, Rwanda (90 / 202). In Eritrea, Jannone (1948) found that ova are normally laid singly or in groups of up to six, but occasionally groups of up to 40 may be laid; this could be a reflection of high skipper densities. Dead ova are quite commonly found, and many of these have emergence holes of egg parasitoids (below). Ova took 20 – 30 days to develop at 2300 m in Eritrea (Jannone 1948). Leaf shelters, Nairobi typical. The newly hatched caterpillar eats the shell of the ovum, apart from the base, and then pulls together the edges of a leaflet towards the apex to form a small channel in which it hides (Figure 5.2). It seems likely that there is considerable predation of first instar caterpillars by spiders and wasps due to the inadequate protection provided by this shelter; certainly hatched ova with no associated shelter, and empty first shelters with no nearby subsequent shelters are quite common. The caterpillar feeds basally or distally to the shelter before making a new shelter lower down the leaf, which it is then able to close to provide much better protection (Figure 5.4). Mature caterpillars will shelter between two leaflets held by silk, and pupate between several leaflets of a young leaf drawn together. Caterpillar. Clark (in Dickson & Kroon 1978) indicates rearing through six instars, but in Nairobi MJWC documented only five instars (90 / 24). Below, we segregate the available information by locality.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	Kelley’s account includes black and white photographs of the caterpillar and pupa from KwaZulu-Natal Province, South Africa. These are not very clear, but the caterpillar can be seen to have a pale head, with the epicranial suture dark and black lines adjacent to the epicranial suture from the vertex, and laterally from vertex to stemmata. There are pale subdorsal lines on the body. We note that the lines adjacent to the epicranial suture extend to the vertex, and seem more or less straight, with little obvious divergence at the adfrontals, although the head is positioned so that this is not clear. Thus, the photo resembles Clark’s (in Dickson & Kroon 1978) figure of the penultimate instar 5, rather than the final instar. Clark (in Dickson & Kroon 1978) provides a full series of paintings of the life history at Port Elizabeth, Eastern Cape Province, South Africa. The first instar is pale with a dark head. The second instar is pale green and the dark head has a trace of a pale area lateral to the adfrontals. Instar 3 is green with a dark head, with the adfrontals and a streak lateral to them pale. Instar 4 has a dark green dorsal line and the head is now pale, with dark sutures, a dark line adjacent to the epicranial suture and a strong lateral line from the vertex to stemmata, which may extend to the posterior margin. Instar 5 is similar, but the body now has pale subdorsal lines. In the final instar 6 the head and body are paler; the lines adjacent to the epicranial suture start well below the vertex, and diverge sharply along the adfrontal sutures, ending in a slight swelling; the lateral line from vertex to stemmata is broad and appears to extend to the posterior margin; the sutures are dark, and in one of two illustrations, the upper part of the adfrontals is dark.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Caterpillar, Eritrea. Jannone (1948) has provided a description and drawings of the caterpillar of Z. dysmephila in Eritrea (which may be indigenous or introduced). His figure 3.2 shows a mature caterpillar in dorsal view with a dark dorsal line but no clear subdorsal lines. The head has black lines adjacent to the epicranial and adfrontal sutures very similar to those illustrated by Henning et al. (1997) and we report below from Zambia. Jannone adds that there is a black band on the posterior margin of the head (una stretta fascia nera in corrispondenza del foro occipitale), and the dorsal part of the adfrontals is black in some specimens (La zona triangolare compresa tra i due rami anteriori della sutura metopica, in qualche esemplare può essere anche nera). The caterpillar develops in 30 – 55 days at 2300 m in Eritrea depending on the season (Jannone 1948).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	Caterpillar, Nairobi typical. MJWC reared this form several times, and found it to be variable in the head markings, and in the timing of their development in instars 2 – 4. This is described in some detail to set the observations from other locations in context. Instar 1 (87 / 6) has a black head, shiny, slightly rugose, 1.0 mm wide (range 0.98 – 1.10, n = 6) x 1.1 mm wide (range 1.06 – 1.18, n = 6); narrow, black pronotum; body dull shiny green; A 8 slightly and A 9 quite strongly tinted red; long erect white setae on anal plate. Figure 6 shows a first instar caterpillar about to moult. Instar 2 (87 / 6) similar; head dark, 1.3 mm wide (range 1.25 – 1.37, n = 4) x 1.4 mm high (range 1.37 – 1.49, n = 4); body more yellow-green, anal colouring reduced to yellow tint on A 9, and white setae on anal plate not obvious. From the second instar, caterpillars may begin to show characteristics of the head markings of the final instar, although the markings are darker and more diffuse. However, in some individuals the head remains mostly black until the fourth (penultimate) instar. Thus, although the head of caterpillars of instar 2 are normally dark, one of two caterpillars collected at Thika (87 / 6) had the ground colour pale brown; epicranial and clypeal sutures dark; weak diffuse dark line parallel to epicranial suture from well below vertex to and merges into a darker ventral area; diffuse dark band from vertex to stemmata; posterior margin narrowly black. Instar 3 head 1.75 (range 1.65 – 1.84, n = 13) x 2.05 mm high (range 1.88 – 2.16, n = 13). Of the 13 individuals examined, two had a dark head, without distinguishable markings, while at the other extreme, one individual had the head pale brown; epicranial and clypeal sutures black; a diffuse brown line parallel and close to epicranial suture, from well below the vertex to the top of the adfrontals; vertex to apices diffusely brown; posterior margin narrowly dark. There was continuous variation between these extremes, with most caterpillars having the line adjacent to the epicranial suture longer and darker, extending half way down the level of the frons, and a dark band from vertex to apices and stemmata. Instar 4 head 2.4 mm wide (range 2.24 – 2.71, n = 11) x 2.8 mm high (range 2.71 – 2.94, n = 11); variable markings. At one extreme, one of 11 caterpillars had the ground colour brown (as opposed to light or pale brown); epicranial, adfrontal and clypeal sutures blackish; dark line adjacent to epicranial suture fused with the dark epicranial suture in lower part, widening and becoming more diffuse to the level of the bottom of the frons; adfrontals pale brown; a wide, dark lateral band from vertex, reaching the posterior margin laterally; posterior margin black. At the other extreme, some caterpillars had markings similar to those of the final instar: ground colour light brown; epicranial and clypeal sutures black; adfrontal suture very slightly darkened in dorsal half; black line adjacent to epicranial suture runs from well below the vertex to the top of the adfrontals; no trace of dark spot on edge of epicranium adjacent to adfrontals; slightly dark, diffuse, lateral band from vertex to apex to half way down side of head; posterior margin narrowly dark. More commonly the dorsal parts of the adfrontals and frons may be dark; there may be a spot on the epicranium, adjacent to the adfrontals, level with half way down the frons; the lateral band may continue to the stemmata. The head of the final, fifth instar caterpillar measures 3.3 x 4.2 mm wide x high, and the head markings are variable although less so that in the previous two instars. The following description of the final instar (87 / 2) is based on one of the first skipper caterpillars that MJWC reared in Nairobi, and represents the typical form –– i. e. the form most frequently found around Nairobi, similar to, but not matching those documented from South Africa (above). Length 48 mm when collected, six days before it stopped feeding prior to pupation; width 6 mm; height 4 mm. Other individuals of at least 50 mm have been measured (89 / 35). Head shiny and slightly crinkly; ground colour whitish brown; ventral part of face and adfrontals slightly paler; clypeus pale grey-brown; epicranial suture slightly darker; narrow black line from just below vertex, near and parallel to epicranial suture, diverging slightly and thickening at the level of the top of the adfrontals where it stops; half way from the end of this line to the mouthparts a black spot on the edge of the epicranium, adjacent to adfrontals; stemmata black. The head is held with the mouthparts directed anteriorly (semiprognathous), and the dorsal part resting on top of T 1, which has a narrow black pronotum. Body dull green, T 2 yellowish green; dark green transparent dorsal line about 1 mm wide T 3 – A 7; separated by a narrow line of ground colour, a slightly diffuse dorsolateral line T 2 – A 8; anal plate semicircular, nearly flat, somewhat shiny; spiracle T 1 brown, other spiracles pale brown; all legs concolorous. As can be seen in Figure 7, the intensity of the line adjacent to the epicranial suture is variable, as is the size and intensity of the lower spot adjacent to the adfrontals, but none have been found where the line and spot are joined; similarly, the ground colour varies in intensity, there may be a trace of a lateral band, and the posterior margin may be dark. Prior to pupation, the bodies become more opaque and yellowish and the dorsal line is more pronounced. The wax glands developed ventrally on A 1 – A 8, covering the whole ventral surface apart from a narrow ventral line, the prolegs, and the posterior half of A 7 and A 8 (MJWC 89 / 105). In other caterpillars (MJWC 90 / 69, 95 / 102) the wax glands extended to whole ventral area A 1 – A 8 apart from the prolegs. The young caterpillars feed only from the edge of the leaflet, but mature caterpillars are able to eat through the mid-rib as well, and leave the tip of the leaflet truncated, coming to a point at or near the mid-rib. This is quite characteristic of palm feeding skippers (Figures 4.1, 5.5), and MJWC has noted it in several other genera around the tropics. Total caterpillar development takes about 48 – 57 days in Nairobi. Instars durations of 7, 7, 7, and 7 – 8 were noted for the first four instars.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Caterpillar, Nairobi atypical. All the caterpillars reared from Nairobi fitted a common pattern, treated above (Figure 7) and referred to as ‘ Nairobi typical’ except for one caterpillar (89 / 35; Figure 8) which was sufficiently different to suggest it was some other species, but which produced an adult male indistinguishable from others reared at Nairobi. It was collected as a penultimate instar premoult caterpillar, Kyuna Estate, Nairobi, 12 May 1989. The penultimate instar resembled a very heavily marked fourth instar of the typical form: 2.4 x 2.8 mm wide x high; ground colour light brown; epicranial suture and clypeal sutures heavily black; heavy dark line adjacent to epicranial suture and contiguous with marking over epicranial suture except portion nearest vertex, widening ventrally to the top of the adfrontals and continuing to level of bottom of the frons; apical portion of adfrontals black; frons brown; dark lateral band from vertex to stemmata, widening towards stemmata; posterior margin narrowly dark. The final instar caterpillar measured 30 mm when newly moulted and grew to 50 mm shortly before it pupated. Head similar in size (c. 3.0 x 4.0 mm wide x high), shape and orientation to Z. dysmephila Nairobi typical, but with strikingly different black and white markings; a broad white stripe, narrowed at each end, running down each side of the head; from the vertex, a short, narrow white line runs adjacent to the epicranial suture, about ¼ the distance to the adfrontals; adfrontals, except at apex and labrum white; adjacent to the dark posterior margin, there is a ventrolateral white area. By the time the caterpillar is full-grown, the white markings turn very pale brown, in this regard, resembling Nairobi typical Z. dysmephila. T 1 white, with a narrow black pronotum which extends laterally to just short of the orange-brown T 1 spiracle. Body translucent yellowish green; a 0.5 mm wide subdorsal white stripe, yellowish T 2 – A 2, and containing the yellow gonads at A 5; spiracles conspicuous orangebrown; all legs concolorous. The pupa was not distinguished from that of Z. dysmephila Nairobi typical. In colouring this caterpillar is closest to the caterpillars found in Kakamega Forest, Kenya (below), sharing the pale ground colour of collection 91 / 23 (male reared), but with markings very similar to those of Z. quaternata from Gambia (Figure 16). The genitalia seem to be the same as those of other Nairobi specimens, so the caterpillar may represent an extreme variation or an aberration of Nairobi typical. Caterpillar, Kakamega Forest, Kenya. MJWC has reared a male and two females of what appeared to be Z. dysmephila from Kakamega Forest, western Kenya, the adults being indistinguishable in appearance from those reared in Nairobi. However, reviewing reared material for this paper in discussion with T. B. Larsen, MJWC realised that the single male reared from Kakamega (91 / 23) is not the same as that reared from Nairobi, based on clear differences in the male genitalia, which TBL already suspected from other parts of Africa. One individual (90 / 76 A) was reared from an ovum and had 5 instars, matching what was found at Nairobi, and so treated as typical here.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	Male 91 / 23 was collected 31 Mar 1991 as an instar 3 caterpillar on the same P. reclinata palm in northern Kakamega Forest as earlier collections of Z. dysmephila and of Caenides dacela (Hewitson) below. A detailed description was not prepared (at that time it was thought to be within the range of variation of Z. dysmephila), but fortunately the last two instars were photographed (Figure 9). The head of the instar n- 2 caterpillar measured 1.6 x 1.8 mm wide x high; brown with the epicranial suture darker. The head of the fourth instar caterpillar measured 2.2 x 2.7 mm wide x high; ground colour dark brown; a diffusely edged brown patch from near vertex to in front of stemmata; labrum brown. The head of the final instar caterpillar measured 3.1 x 3.8 mm wide x high; colour almost white, with black markings: from posterior margin of vertex to stemmata, parallel to the epicranial suture, a spot on epicranium adjacent to adfrontals at level of mid frons, epicranial, adfrontal and clypeal sutures. Both instars have T 1 white with a narrow black pronotum on the posterior margin. The spiracles are strikingly orange-brown. Female 90 / 76 A, collected 20 Jul 1990 as an ovum on the same P. reclinata was more carefully documented. The ovum was laid between the apical and adjacent leaflets, right at the base. It was 1.6 x 1.0 mm diameter x height, with very fine ribs, the top flattened and dark. There were five instars, the head capsules for instars 2 – 5 measuring 1.2 x 1.4 mm, 1.7 x 1.9 mm, 2.3 x 2.8 mm, and 3.2 x 4.0 mm respectively (that for instar 1 was lost). Instar 1 had a black head, yellow-green body with anal segments tinted orange. Instar 2 was similar although the head was very dark brown; it measured 8 mm when newly moulted. Instar 3 (Figure 10.1) measured 12 mm when newly moulted; head dark brown; epicranial and clypeal sutures black; the middle of each epicranium slightly paler; black pronotum now evident; body translucent light green, white subdorsal line. Instar 4 (Figure 10.2 – 4) measured 25 mm newly moulted; head brown; epicranial and clypeal sutures very dark; heavy dark line adjacent to epicranial suture, widening and becoming more diffuse adjacent to adfrontals, extending to level of near bottom of frons; upper part of adfrontals dark; a broad, dark lateral band from vertex to stemmata, wider ventrally; narrow band of ground colour before dark posterior margin. T 1 with narrow black pronotum; body translucent dark green with white subdorsal line. This fourth instar individual resembles that of the male treated above, except the pale markings on the head are stronger. The final, fifth instar was not described or photographed; it resembles that described above for the male (Figure 9.2). Under Nairobi conditions, this individual took 9, 7, 7, 11, 19 and 23 days for the five instars and pupa. There are differences between these early stages and those documented above as Nairobi typical, even given the level of variation observed in Nairobi typical and the small number of observations from Kakamega. Thus, in the final instar, from Kakamega Forest, the paler ground colour of the head, lateral band from vertex to stemmata, and more conspicuous orange-brown spiracles may be significant.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Caterpillar, Kenya Coast. MJWC collected Z. dysmephila at the Kenya Coast (Bamburi, Malindi) on P. reclinata, and it probably occurs throughout the developed part of the coast where ornamental P. reclinata are planted. A female and crippled male were reared from one collection made at Bamburi (90 / 58). Unfortunately, at the time they were thought to be the same as the caterpillars from Nairobi and so were neither photographed nor described. The following is based on an examination of the associated head capsules of the reared male and compared with all material collected at the coast. The penultimate instar head measured 1.9 x 2.1 mm wide x high (n = 3), rather smaller than in the same instar for Nairobi typical (2.4 x 2.8 mm). However the final instar was almost the same: 3.2 x 4.1 mm (n = 4) compared to 3.3 x 4.2 mm. Penultimate instar (90 / 58 B): head ground colour pale brown; epicranial and clypeal sutures dark; slightly darker brown line adjacent to epicranial suture as far as adfrontals; no spot against adfrontals; laterally and around stemmata concolorous; posterior margin narrowly dark; the reared female (90 / 58 A) was similar and another individual from Kikambala, Kilifi District (87 / 39), differed only in that the posterior margin was not dark. Final instar (90 / 58 B): head ground colour pale brown; epicranial and clypeal sutures dark; dark line adjacent to epicranial suture to level with the top of the frons; dark spot on epicranium against adfrontal at about the middle of the adfrontal; patch around stemmata dark; posterior margin narrowly dark; the reared female (90 / 58 A) was similar; in two individuals from Kikambala (87 / 39) the posterior margin was narrowly dark, and one of the two had a diffuse brown area laterally on the head. Thus, the caterpillars from the coast were at the pale extreme of variation observed in Nairobi typical Z. dysmephila. Caterpillar, Rwanda. A cast final instar caterpillar skin, separate head capsules from the two previous instars and ova were collected at Butare, Rwanda on an ornamental Phoenix sp. The head of the final instar caterpillar differed from others shown here; it measured 3.0 x 4.2 mm wide x high and was light brown with blackish markings: a line from vertex laterally to stemmata, widening from about half way and dividing into two basally to cover the anterior and posterior stemmata; epicranial suture dark; a strong line from just below vertex (where narrowest), parallel to epicranial suture and on epicranium adjacent on adfrontals (widest at top of adfrontals); upper half frons dark; adfrontals and posterior margin of head not dark; pronotum narrow, black; a light covering of white waxy powder; no associated pupa.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	A single penultimate instar head capsule measured 2.6 x 3.1 mm wide x high; light brown with heavy dark markings: posterior margin narrowly dark; epicranial and clypeal sutures dark, but adfrontal sutures not; dark line from vertex laterally to half way down head; black line parallel to epicranial suture from a little below vertex (where narrowest) to level with top of adfrontals; dark line through anterior stemmata. Three n- 2 instar head capsules measured 1.8 x 2.1 mm wide x high and were variable in markings. One was brown with extensive dark markings: posterior margin broadly dark to edge of face; epicranial and clypeal sutures dark; a diffuse dark area parallel to epicranial suture and on epicranium adjacent to adfrontals, from below vertex to level of mid frons, then running laterally to stemmata where it joins the dark posterior border; frons darker brown than brown of face. The other two were dark, with slightly paler areas on the upper epicranium and from the lower half of the adfrontals to the ventral stemmata, similar to the third instar caterpillar from western Kenya shown in Figure 10.1.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Caterpillar, Mafwemiru, Rubeho Mountains, Tanzania. A 2013 collection by TCEC on Phoenix reclinata in the Rubeho Mountains produced Z. dysmephila caterpillars with a pale brown head, no lateral line, a strong black line parallel to the dark epicranial suture, a spot around the stemmata, and individuals with and without a spot adjacent to the lower adfrontals (Figure 11). These caterpillars resemble those documented above from Nairobi, but not elsewhere.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	description	Caterpillar, Mufindi, Southern Highlands of Tanzania. A caterpillar from Mufindi (Figure 12) has the pale brown head of Nairobi (Figure 7) and southern African populations. The heavy dark line adjacent to the epicranial suture is separate from a lower black spot adjacent to the adfrontal suture, resembling individuals from Kenya (all three populations, Figures 7, 9, 10), central Tanzania (Figure 12) and north-east Zambia (Figure 13), but not elsewhere. There is a heavy black band from the dorsal posterior margin to the stemmata, widening basally, resembling western Kenya specimens (Figure 9, 9), and that documented from Zambia below, but not those from Central Tanzania (Figure 11), Nairobi (Figure 7) or the Kenya Coast. Caterpillar, north-east Zambia. Caterpillars recently collected by TCEC in north-east Zambia have the head pale brown, strongly marked with black, similar to the most heavily marked Nairobi examples (Figure 7.2), with a dark lateral line, although not as strong as that of the individual from Mufindi, Tanzania; the body had yellow subdorsal lines rather than white (Figure 13). It appears to be almost identical otherwise to the caterpillar from Mufindi, Tanzania. Pupa. The pupa (based on Nairobi typical material, Figure 14) is 28 mm long; head, appendages and dorsal areas dark brown; abdomen pale brown ventrally; becoming darker during development; surface shiny under a light covering of white waxy powder; rounded, with the frons and eyes slightly protuberant; thorax and eyes (except vertical bar down middle) with light covering of reddish brown, erect setae; the proboscis does not protrude beyond the wing cases. The inside of the shelter is heavily covered with white waxy powder, but the pupa itself only lightly so. The pupae from other populations did not appear to differ. The pupal stage lasts about 18 days in Nairobi (range 17 – 21, n = 3), compared to 18 – 25 days at 2300 m in Eritrea (Jannone 1948).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Natural enemies. South Africa. Dickson & Kroon (1978) record? cynipid and chalcid egg parasitoids, Pimpla sp. (ichneumonid third instar caterpillar parasitoid), tachinid larval-pupal parasitoids Nemorilla cruciata (Wiedemann), Erycia sp. and Theocarcelia incedens (Rondani) (? synonym of T. acutangula (Macquart )), and chalcid pupal parasitoids. They also note that insectivorous birds take a heavy toll of caterpillars.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	materials_examined	Nairobi, Kenya. The ova are parasitized by gregarious trichogrammatid egg parasitoids; for example one produced 6 tiny, brown adult wasps 18 days after it was collected (Nairobi, 90 / 20). On two occasions, MJWC found dead caterpillars in their shelters in Nairobi, one associated with four pupae (90 / 122) which produced one male and three females of an? Elasmus sp. (Eulophidae) and another (fourth instar, 90 / 96) with nine naked black pupae of a euplectrine, which emerged 3 – 4 days later, yielding one male and seven female adults (and one escape).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Kakamega, Kenya. Twice MJWC found large caterpillars in North Kakamega Forest parasitized by a small gregarious tachinid fly. A fourth instar caterpillar produced 3 puparia which failed to emerge, and a fifth instar caterpillar produced 13 small puparia, but only one crippled adult fly emerged. It appears to be the same species, Ceranthia sp., which has also been reared from Hesperiinae Pelopidas mathias (Fabricius) (Cock & Congdon 2012) and Chondrolepis leggei (Heron) (MJWC unpublished) in western Kenya.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	materials_examined	Kenya Coast. At the Kenya coast the ova are attacked by a dark egg parasitoid, larger than those mentioned above. An ovum collected on a small, but partially defoliated Phoenix reclinata near Malindi, 22 Sep 1989 (Figure 3.1) produced a single such parasitoid on 5 Oct (MJWC 89 / 63).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32281B5BAB9F59EDFE74E183.taxon	discussion	Rwanda. The material collected at Rwanda (90 / 202) revealed a high level of egg parasitism. In the batch of 24 ova, three were completely missing having probably become detached from the substrate, three had eclosed normally, 14 had parasitoid emergence holes and 4 were dead, perhaps due to parasitism. Two individually laid ova produced two different species of solitary egg parasitoid 21 and 22 days after collection. One small leaf shelter typical of Z. dysmephila contained three black euplectrine pupae, one of which emerged seven days later. Discussion. We have shown geographical and local variation in the caterpillars of Z. dysmephila, some of which correlates with unanticipated variation in male genitalia, where this has been investigated. More rearing is needed to assess the extent of variation at most sites, as it is not clear whether this variation reflects a variable species throughout its range, or is a function of geography. The material from Nairobi suggests a discrete, constant form of the male genitalia, but there is not enough material in series from other areas to assess the observed differences reported here. Zophopetes dysmephila is a complex species, or a species-complex, and more rearing, male and female genitalia examination and barcoding will be needed to work out the full picture. This needs to be followed up with further study.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC323D1B5EAB9F5C9BFF68E107.taxon	description	Mabille (1876) described Pamphila quaternata, which closely resembles Z. dysmephila (above), from Senegal. Evans (1937) treated Z. quaternata as a valid species, but suggested it might be a subspecies of Z. dysmephila, and subsequently he did make it a subspecies (Evans 1947). Lindsey & Miller (1965), Gillies (1982), Ackery et al. (1995) followed Evans in this regard, for example Gillies considered that of 16 specimens in his collection from Gambia, 14 resembled quaternata and two dysmephila. However, Larsen (2005) recognized them as two separate species with consistent differences in the male genitalia, and this is the situation today. Adult Z. quaternata are readily recognised by the underside having a dense irroration of tiny blackish flecks, often with a slight reddish tone (the red tone not being evident in Figure 15) (T. B. Larsen pers. comm. 2013). Zophopetes quaternata is now known from Gambia and Senegal, east to Ghana (Larsen 2005), with additional records from Nigeria (Figure 15) and Cameroon, and a cluster in DR Congo (Virunga), south-west Uganda and north-west Tanzania (T. B. Larsen pers. comm. 2012). Records of Z. dysmephila from West Africa are misidentifications of Z. quaternata (Larsen 2005), as Z. dysmephila is at most found as far west as southern Cameroon. Adult behaviour. We have no observations, but in Gambia, Gillies (1982) reported Z. quaternata ‘ flying at great speed round Phoenix palms at dusk’. Food plants. The record of Z. dysmephila from Phoenix reclinata at Lamto (Vuattoux 1999) is referable to Z. quaternata (Larsen 2005). SCC has reared Z. quaternata from P. reclinata in Gambia (reported below), and MJWC has reared what appears to be Z. quaternata from P. dactylifera at Ibadan, Nigeria (94 / 107, 94 / 109).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC323D1B5EAB9F5C9BFF68E107.taxon	discussion	Life history, Gambia. As the type locality for Z. quaternata is Senegal, the following observations from Gambia by SCC must represent the true Z. quaternata. In October 2010, SCC searched for skipper early stages on palms in the vicinity of Banjul, Gambia, where the naturally growing P. reclinata palms are under threat as the old swampy areas are being drained. He found two caterpillars of Z. quaternata on P. reclinata in a forest remnant amongst the deepest undergrowth, and reared one through. The caterpillar (Figure 16.1 – 2) is similar to those of the Z. dysmephila group, but the head has markings which are different, although more material is needed to assess variation. SCC’s photographs and the cast skin in ABRI show the head to be light brown, with dark brown-black markings on the posterior part of the head extending to the posterior margin, the epicranial suture, a heavy black line each side of the epicranial suture, which meet just above the adfrontals, and then run down the epicranium adjacent to the adfrontals to level with the clypeus, and the whole of the frons, but not the adfrontals. The pupa (Figure 16.3) resembles those of the Z. dysmephila group.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC323D1B5EAB9F5C9BFF68E107.taxon	materials_examined	Life history, Nigeria. MJWC reared this species at Ibadan, Nigeria, where they were causing significant damage to small Phoenix palms planted as ornamentals in Mar 1994 (94 / 107, 94 / 109).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC323D1B5EAB9F5C9BFF68E107.taxon	description	Caterpillars of 6 and 8 mm (instar n- 3): head 1.2 x 1.4 mm wide x high (n = 2); light brown; epicranial suture dark; dark black pronotum; body translucent green. A 13 mm caterpillar (instar n- 2) was similar: head 2.0 x 2.2 mm wide x high; broadly and diffusely dark in a band from vertex to stemmata; epicranial suture and upper part of clypeal suture dark; slightly separated from epicranial suture, and adjacent to adfrontal sutures very diffuse, slightly darker band from below vertex to level with mid frons, with inner margin more sharply defined. Five penultimate instar caterpillars of 22 – 28 mm were collected, and a description was prepared of one of 23 mm. Head nearly oval, but widest below centre, slightly indent at vertex, 2.4 x 2.9 mm wide x high (n = 6); semiprognathous; ground colour light brown with dark brown markings: dorsal half of posterior margin, a detached diffuse line from bottom end to stemmata, stemmata, epicranial suture, upper part of adfrontals, a heavy stripe each side of epicranial suture, a diffuse mark on epicranium adjacent to adfrontals level with the top of frons, which is also dark; mouthparts brown. Narrow black pronotum. Body dull yellow-green; dorsal line dull green; pale subdorsal line (within ground colour) T 2 – A 8; anal plate with brown tint; spiracles brown; legs concolorous. The intensity and extent of the markings on the face varied between individuals (Figure 17). A narrow black line across the front of the anal plate (Figure 17.2) in one individual was thought to be an artefact where a piece of the moulted skin of the previous instar with the pronotum had stuck to the body of the moulted caterpillar. The final instar caterpillar was not described or photographed at the time, and the following is based on four caterpillars preserved in ethanol. Head 3.3 x 4.2 mm wide x high (n = 4); rugose; ground colour light brown, as penultimate instar; diffuse dark brown band from vertex to stemmata, the upper third against posterior margin, the lower part swinging forwards to the stemmata, the intensity varying between individuals; epicranial suture dark, but not strongly so; upper part of adfrontal and clypeal sutures dark in some individuals; black line from below vertex, close to epicranial suture to level with top of adfrontals, a black spot on the epicranium adjacent to adfrontal, at level of mid frons, more or less in line with the black line (similar to Figure 17.1 but more strongly marked). In another individual the black line continues to join the spot against the adfrontal, while two were intermediate with the black line partially or almost separated from the black spot; two individuals had a dark line down the centre of the frons. Narrow black pronotum, divided at the dorsal line. The pupa (Figure 18) is similar to those of the Z. dysmephila group: 24 mm; smooth in outline with no projections; head and thorax light chestnut brown; appendages and abdomen light brown; spiracle T 1 dark brown. Groups of scattered brown, forward-directed setae on thorax and eyes. The inside of the shelter and the pupa lightly dusted with white waxy powder.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC323D1B5EAB9F5C9BFF68E107.taxon	discussion	Discussion. The caterpillar of Z. quaternata from Nigeria is of the same general form as that of Z. quaternata from Gambia, and in common with that population, the posterior part of the head is dark. However, the markings on the face, especially the black frons of the Gambia population differ from the reduced markings found in Nigeria. The possibility that Z. quaternata occurs in geographically separated subspecies should be considered. Furthermore, the caterpillars of Z. quaternata from Nigeria (Figure 17) seem closer to those of Nairobi typical Z. dysmephila (Figure 7) than to Gambian Z. quaternata (Figure 16). Given the variation in caterpillars and male genitalia discovered in Z. dysmephila above, it seems clear that further investigation should be directed at both species.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B5EAB9F5C27FAD5E328.taxon	description	Evans (1937) described this species from Côte d’Ivoire, commenting that it is ‘ generally similar to [Z.] dysmephila ’. It is known from Ghana to Kinshasa, DR Congo, but is rare (Larsen 2005, T. B. Larsen pers. comm. 2014). SCC found caterpillars of what he suspected was Z. ganda on Phoenix at 1500 m on the Obudu Plateau, East Nigeria, 7 – 11 Apr 2007. The three pupae reared were all parasitized by a larval-pupal tachinid fly. The pupae are preserved in ABRI and two have cast final instar caterpillar skins. The head capsule is light brown, with the posterior margin and epicranial suture narrowly dark; a black line from apex, close to epicranial suture to just below apex of adfrontals; adjacent to the adfrontals about half way down, a black spot, slightly wider than the line above; an inverted V at the apex of the frons; a dark line through anterior stemmata; narrow black pronotum. These markings are within the range observed within Z. dysmephila and Z. quaternata. However, the caterpillars were subjectively smaller than those of Z. dysmephila and Z. quaternata, and no other members of this species group have been reported from eastern Nigeria, so we treat this as a provisional association, needing confirmation.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	description	Hewitson (1867 – 1871) described Hesperia cerymica from Akwa Akpa (= Old Calabar, south-eastern Nigeria). Evans (1937) treated cerymica as two subspecies: the nominate cerymica in West Africa, from Senegal as far east as Cameroon, and what he referred to as nobilior Holland, the eastern subspecies, described from Gabon (Lambarene, French Congo), but for which Evans only listed specimens from Uganda, Kenya and Tanzania, although it also occurs in DR Congo (Kielland 1990, Ackery et al. 1995) and north-west Zambia (Heath et al. 2002). Lindsey & Miller (1965) treated cerymica and nobilior as full species, based on differences in the male genitalia, and this has been followed by subsequent authors, although Larsen (1991) suggested they may be subspecies, when documenting the population from the Kenya Coast as Z. nobilior. T. B. Larsen (pers. comm. 2013) has now established that the taxon referred to as nobilior is not that species, which is currently only known from two females from Gabon, and near Mbandaka in western DR Congo that are much smaller than Z. cerymica and with a rather different hind wing underside. Although Lindsey & Miller (1965) considered that the male genitalia of Z. cerymica from West Africa and East Africa (their Z. nobilior) represent different species, T. B. Larsen (pers. comm. 2013) notes that the material treated as nobilior by Evans falls within the range of variation in colouring and male genitalia seen in Z. cerymica, so that for the moment at least, it would be more appropriate to treat this as a single variable species without subspecies. We have seen caterpillars from Togo, Côte d’Ivoire, Kenya (coast), Tanzania and Zambia, which we illustrate separately here. There are no significant differences between the early stages of these populations, supporting the view that Z. cerymica is a somewhat variable species found from Senegal, east to Kenya and south to Zambia. However, given the variation that we have reported in caterpillars and male genitalia indicating complex species or species-complexes in the other Zophopetes spp. above, this conclusion perhaps should be considered provisional pending more detailed studies.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	discussion	In Kenya this species is restricted to the coast, where coconut is a regular food plant. Although not as common as those of Gretna carmen capra Evans on coconut, caterpillars can be found without too much difficulty. In Tanzania, it is found in forests on the eastern shore of Lake Tanganyika, and in eastern Tanzania it is known from Dar es Salaam, the Pugu Hills, and the Lukosi River near Iringa (Kielland 1990). It is not confined to forest, for example, on the Lukosi River as it flows through dry thorn scrub and rocky Acacia woodland for the last 20 – 30 km to its junction with the Great Ruaha, it breeds on Raphia palms growing on the banks and islands among the channels of the fast flowing river bordered by very degraded remnants of riverine forest. In Zambia it is restricted to the north-west (Heath et al. 2002).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	discussion	Food plants. As noted above, early reports of Z. dysmephila as a palm pest in the French West African literature were misidentifications for Z. cerymica (rather than Z. quaternata, which Z. dysmephila resembles). The associated descriptions and figures of the early stages (e. g. Genty 1968, Mariau & Morin 1974) match Z. cerymica rather than Z. dysmephila, and Z. cerymica replaces Z. dysmephila in more recent publications (Mariau et al. 1981, Mariau 2000, 2001). Thus, Z. cerymica (as Z. dysmephila) is recorded from oil palm (E. guineensis) at Anguédedou, Côte d’Ivoire (Genty 1968). Mariau & Morin (1974) give a general account of Z. cerymica (as Z. dysmephila) and Pteroteinon laufella (see below) as pests of oil palm and coconut in West Africa. The pinned adult butterfly they illustrate as Z. dysmephila appears to be a female Z. cerymica, and the authors refer to the head of the caterpillar as uniformly golden yellow (uniformément jaune d'or), which is diagnostic for Z. cerymica (Figure 20 – 24) compared to Z. dysmephila (Figures 7 – 13). We note that although the paper title specifies hesperiid defoliator pests of oil palm and coconut, the text only refers to oil palm. Vuattoux (1999) only reared this species from E. guineensis in Côte d’Ivoire. In Ghana, T. B. Larsen (pers. comm. 2013) found it most frequently along beaches with many coconut palms (especially between Cape Coast and Takoradi). Early food plant records from East Africa were under the name Z. cerymica, anticipating the present arrangement. Thus Van Someren (1974) gives Raphia spp. and Cocos nucifera (as Cocoa nucifer) as food plants of Z. cerymica in his list of food plants of East African butterflies. Sevastopulo (1975) repeats these records, and later adds Phoenix (Sevastopulo 1981). Treating this species as Z. nobilior, Kielland (1990) lists the food plants as Phoenix, Borassus and Raphia palms, but doesn't mention coconut. Larsen (1991) adds coconut, based on MJWC’s rearing, and repeats these food plants for Z. cerymica (Larsen 2005). Ackery et al. (1995) list the food plants of Z. cerymica as Borassus, Phoenix and Raphia. Heath et al. (2002) give the food plant in north-west Zambia as Raphia sp. In Kenya, MJWC found this species on Cocos nucifera only, whereas TCEC found caterpillars on E. guineensis in Tanzania, and on Raphia farinifera in Tanzania and north-west Zambia, although some were transferred from R. farinifera to C. nucifera in order to rear them through. Both authors have tried independently to transfer caterpillars to feed on leaves of P. reclinata, and failed. MJWC tried with one caterpillar that only fed a little and then died (89 / 18 B). We doubt that Phoenix spp. are normal food plants of Z. cerymica. We also have reservations about the repeated records from Borassus spp., which TCEC has searched without finding any Hesperiidae early stages. MJWC has also reared Z. cerymica from C. nucifera at Lomé, Togo (90 / 213), and found a caterpillar on an unidentified planted palm in Côte d’Ivoire (88 / 214). Leaf shelters. On R. farinifera at Lukosi River, Tanzania, TCEC noted that the young caterpillars roll the tip of a leaf into a tube and eat their way back down the leaf as they grow. When the midrib becomes too tough, the shelter is abandoned, after the caterpillars have eaten down to it from one or both sides.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	description	The medium grown caterpillars (penultimate or previous instar) feed from the apex of a leaflet, where they make a shelter by rolling one or both edges of the leaflet over upwards. Rolling the flap upwards is the line of least resistance, as the young leaflets are naturally V-shaped, and if they rolled downwards caterpillars would have to deal with the sharp thorns on the underside of the midrib of the leaflet. Fully grown caterpillars make a tubular shelter, usually at the apex of the partly consumed leaflet, by rolling both edges of the leaflet under. Feeding usually leaves the distal end of the leaflet truncate, but pointed at the midrib.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	discussion	Ovum. The ova are laid on the leaflets of C. nucifera; they are similar to those of Z. dysmephila but larger. Mariau & Morin (1974) and Mariau et al. (1981) refer to the ova of Z. cerymica and Pteroteinon laufella as 1.5 mm in diameter and laid in variable numbers (en nombre variable) on the leaflet under surface. Ova from the Kenya coast measured 1.8 x 1.2 mm wide x high (n = 2). Ova hatch in 5 – 6 days (Genty 1968). We have found ova of Z. cerymica only singly, so suggest that the references to ‘ variable numbers’ should be considered to apply to P. laufella only, until new observations suggest otherwise.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	description	Caterpillar. Brief descriptions of material from the Kenya coast were prepared based on collection 87 / 22 (Diani Beach, 29 Sep 1987), which was not successfully reared, but considered to match later collections that were. When collected in the penultimate instar, the caterpillar measured 24 mm; head light brown with diffuse central marking on face; narrow dark pronotum; body creamy green with a yellow dorsolateral stripe; anal plate speckled yellow. This brief description is compatible with Figure 20.1, where the detail of the marking on the central face can be seen, the pronotum is white on the anterior margin, and the spiracles are inconspicuous. Four head capsules measured 3.0 x 3.5 mm wide x high (range 2.9 – 3.1, 3.3 – 3.7). The mature caterpillar (Figure 20) is one of the largest of Kenyan skipper caterpillars, measuring up to 46 mm and taking about 14 days to complete development. The head capsule measures 4.0 x 5.2 mm wide x high (range 3.8 – 4.3, 4.6 – 5.6, n = 6). Caterpillar 87 / 22 moulted to the final instar three days after collection, and on the following day (4 Oct) it measured 28 mm (Figure 20.4). Head 3.9 x 4.8 mm wide x high; shiny, rugose; plain yellow brown except for a black posterolateral band from just before vertex, where it is narrow with a brown posterior margin, widening to the bottom where it runs from the posterior margin to the stemmata. Shiny black pronotum to just above spiracles. Body pale green; T 2 – A 9 a dorsolateral yellow-white line, about 1 mm wide; spiracles light brown; anal plate almost chordate; flanged posteriorly; a network of yellow markings anteriorly. Subsequent collections have shown that the black lateral marking of the head may be reduced to a basal bar from near the posterior margin to the stemmata (Figure 21.3) or just a small black spot covering the stemmata (Figure 20.2). When it has finished feeding, the caterpillar turns dirty yellow, losing the dorsolateral lines (Figure 20.6) The final instar caterpillar recorded in Tanzania is similar (Figure 21). The head marking of the caterpillar shown (Figure 21.3) is an extensive black marking from the vertex laterally, and anteriorly to the stemmata, as heavy as the heaviest documented from Kenya (Figure 20.4). Instar 2 (Figure 21.1 – 2) has the dorsal two-thirds of the head dark, almost black, and the ventral one-third brown; body pale green with pale subdorsal lines. The caterpillars from Zambia are also similar. The early instars were not reported from Kenya (above), but instar 1 from Zambia (Figure 22.1) can be seen to have a brown head, with a dark brown line adjacent to and close to the epicranial suture; darker on the frons; narrow black pronotum and a plain yellow-green body. In the penultimate instar (Figure 22.2 – 3) the yellow-brown head has a W-shaped mark on the face, the outer arms parallel to the epicranial suture, and a diffuse dark patch over the dorsal part of adfrontals and adjacent epicranium. The final instar shown in Figure 22.4 – 6 appears to be mature and starting to turn yellow. It is similar to that from Kenya; in the example photographed there is a minimal black marking around stemmata (Figure 22.4), comparable to one example from Kenya (Figure 20.2). No detailed description of the caterpillar was prepared for the caterpillars from West Africa, but they were documented with photographs (Figures 23 – 25). The n- 2 instar head (Togo 90 / 213) measures 2.1 x 2.6 mm wide x high (n = 3); pale brown; bottom of epicranial suture and clypeal sutures dark; short diffuse, dark line parallel to epicranial suture from just above top of adfrontals to level with top of frons; posterior margin dark. The penultimate instar measures 3.1 x 3.7 mm wide x high (n = 4); face of the penultimate instar from Togo (Figure 23) has a variable diffuse W-shaped mark across the top of the adfrontals and adjacent epicranium; stemmata black, sometimes with a black patch around them; posterior margin dark in some individuals. This instar is intermediate to that shown from Kenya (Figure 20.1) and that shown from Zambia (Figure 22.3). In the final instar from Togo (Figure 24), the head is 4.2 x 5.1 mm wide x high (n = 3, 6); plain orange brown, apart from a black patch around the stemmata (Figure 24.2), matching Figure 20.2 from Kenya; T 1 broadly white, with a narrow black pronotum; body white, more or less opaque, with broad yellow dorsolateral stripes with diffuse edges; T 1 – 3 and A 9 and anal plate with a strong yellow tint; spiracles pale brown, contrasting with white body; legs concolorous. The caterpillar takes about a month to develop (Genty 1968). The body of the caterpillar from Côte d’Ivoire (Figure 25) is a brighter white than those from Togo (Figure 24), but otherwise there is no great difference. It too has a small black spot over the stemmata like Figure 20.2 from Kenya.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32381B57AB9F5E17FC2FE4F5.taxon	discussion	Pupa. Pupae were not described, but are similar in all populations; the interior of the pupal shelter and the pupa are both covered with white waxy powder (Figure 26). They are superficially similar to those of Z. dysmephila, but larger (25 – 34 mm), and the proboscis sheath extends 7 mm beyond the wing cases (Figure 26.4). The pupa is held by a girdle consisting of 10 – 14 strands of thick silk; erect setae dorsally on the thorax and forward directed setae dorsally on the posterior part of A 1 (which grip the silk girdle); erect setae dorsally on A 2 – A 4; a protuberance frontally, a pair of protuberances just above this and one at the base of the antenna. One character noted on West African and Zambian material is a tuft of backwardly angled stiff brown setae dorsolaterally on a thickened area at the base of the cremaster. Kenyan material is similarly thickened but lack the tuft of setae (Figure 27). We do not know the function or significance of this character. However, there are rare observations of more robust structures in this situation in other species of Hesperiinae, which suggest a possible function. Cock (2010 b) noted a robust dorsal plate on the dorsal posterior end of the pupa of the Neotropical Anthoptini species Corticea corticea (Plötz), with two robust posterolateral spikes on each side and suggested that since pupation is in a tightly rolled grass leaf shelter, this posterior plate may be used to prevent access by parasitoids or small predators which may enter the pupal shelter at the posterior end; the heavily chitinized plate would be difficult for small predators to attack, and the posterolateral spikes at the side could be used to crush an intruder against the side of the shelter. The pupa of the Asian palm-feeder Gangara thyrsis (Fabricius) (Hesperiinae, incertae sedis) has a very strong posteriorly directed spike at this point (Figure 28.2) which may also have a defensive function. The pupal stage lasts 21 – 23 days under Nairobi conditions, although in Côte d’Ivoire, Genty (1968) recorded the duration as about a month. Natural enemies. Mariau & Morin (1974) did not distinguish between Pteroteinon laufella and Z. cerymica, when they treated the natural enemies of these two skippers in West Africa. However, more recently, Mariau (2001) treats all the same natural enemies under P. laufella and mentions none for Z. cerymica, so we follow this arrangement here. The only exception to this is the record of an unnamed entomopathogenic fungus, which is illustrated on a pupa of Z. cerymica (Mariau & Morin 1974, Figure 9). The single caterpillar which MJWC found at Adiopodoumé, Côte d’Ivoire, had several macrotype tachinid ova on its body (Figure 25), but it died at pupation. MJWC reared several parasitoids from material collected on C. nucifera at the Kenya coast. Ova are parasitized; two adults of an unidentified orange species emerged from an ovum collected at Diani Beach (29. iii. 1989; 89 / 23). The corpse of a fourth instar caterpillar was found in its shelter surrounded by empty euplectrine pupae (Diani Beach, 22. viii. 1995; 98 / 110), but living material of this parasitoid has not been obtained. A field-collected pupa from Diani Beach (26. iii. 1989; 89 / 18 A; Figure 26.4) was parasitized by a gregarious Brachymeria sp. (Chalcididae), which appears to be the same species reared from Gretna carmen capra (below); 37 adult wasps emerged from the one pupa, 20 days after collection. A three-quarter circle of diameter 2.2 mm was cut in the sub-dorsal abdomen, through which the adult wasps emerged. Although there was some white waxy powder on the inside of the pupal shelter, there was very little on the parasitized pupa itself (Figure 26.4). An empty field-collected pupa collected on the same occasion (89 / 18 C) had a 3.0 mm diameter emergence hole probably made by a Brachymeria sp. Economic damage. Mariau & Morin (1974) state that Z. cerymica (as Z. dysmephila) is commoner on oil palm than Pteroteinon laufella, but do not separate the two species in their assessment of the damage. Genty (1968) describes an outbreak in Côte d’Ivoire where up to three-quarters of the foliage was eaten. Outbreaks can occur on oil palms of all ages, including newly planted estates. Defoliation at this stage can be severe (Genty 1968) and set back crop development. Attacks on older palms are less common, the damage usually less serious and normally localised on the edge of plantations. An economic threshold of 20 hesperiid caterpillars per frond has been proposed for mature palms, based on a sample of two fronds per ha at the point of heaviest attack (Mariau et al. 1981). There have been no reports of Z. cerymica occurring in high enough numbers to cause damage to palms in East Africa, or elsewhere in its range, perhaps because of the action of natural enemies keeping populations in check. Nevertheless, Z. cerymica should be considered a quarantine pest for areas outside Africa where palms, especially coconut and oil palm are grown. Discussion. The caterpillars and pupae of Z. cerymica appear almost identical throughout its range. The variation in the extent of the black ventrolateral marking seen on the head of the final instar caterpillar from different collections in Kenya encompasses all forms seen elsewhere. The presence of the tuft of setae dorsolaterally at the base of the cremaster of West African pupae but not on those from East Africa is the only consistent difference we have noticed between the two areas. Nevertheless, at this stage, treating Z. cerymica as a single slightly variable species throughout its range seems reasonable.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32311B57AB9F5AB3FC34E19C.taxon	description	This is a genus of six Afrotropical skippers (Evans 1937, Ackery et al. 1995). Gretna as currently constituted includes some species that feed on palms and at least one that feeds on Marantaceae, an unusual combination in one genus. Below we treat Gretna carmen capra, G. waga (Plötz) and G. balenge balenge (Holland), all of which feed on palms. Vuattoux (1999) reared G. cylinda (Hewitson) from three species of Marantaceae only. MJWC found a Gretna sp. caterpillar on a Marantaceae in Uganda, which based on the known distribution of Gretna spp. is probably G. cylinda, but could be G. zaremba jacksoni Evans. This species will be treated in a later part of this series, dealing with Marantaceae-feeders. Based on the early stages of these three species and G. cylinda, there are three distinct groups of species: (1) Gretna carmen and G. waga, (2) G. balenge, and (3) G. cylinda. As we show below, the first two groups have substantial differences, but also some key similarities. Once we have treated G. cylinda along with other Marantaceae-feeders, we will further discuss the integrity of the genus.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32311B6EAB9F5C9EFED3E5F8.taxon	description	The nominate subspecies, carmen Evans, was described from Cameroon and recorded from Zaire (Evans 1937), but is now known from Côte d’Ivoire, east to the western parts of Kenya, Tanzania and Zambia (Larsen 2005). Subspecies capra was described from a single male from Rabai-Mombasa (Evans 1937). Since then it has been found to be widespread on the Kenya coast (Larsen 1991), and is reported from inland in Tanzania intermittently from the foothills of the Eastern Arc forests to the east coast of Lake Tanganyika, western Tanzania (Kielland 1990, Ackery et al. 1995, TCEC unpublished) and Zambia (Heath et al. 2002). It is not clear from the published literature whether the records of both subspecies in western Tanzania and Zambia reflect overlapping populations, or that both subspecies names have been applied to the same populations. In any case, T. B. Larsen (pers. comm. 2013) has suggested that it may be more appropriate to treat G. carmen as a single species without subspecies. He found no genitalia differences between specimens from Cameroun, eastern DR Congo, Burundi, Zambia, and the Kenya Coast. We note that material from eastern Africa (i. e. Kenya, Tanzania and Zambia) shows quite a lot of variation, but that where a reasonable series is available from one place, e. g. Shiwa Ng’andu (northern Zambia) or the Kenya coast, the underside markings are fairly constant for that locality. In our treatment below, we describe the Kenyan life history in some detail and provide additional information from Tanzania and Zambia. Adult behaviour. The adults rest with their wings closed (Figure 28). They fly low and quickly, and are difficult to follow by eye because of their nondescript grey-brown colouring which blends in so well. Occasionally they will settle on coconut palm trunks, and can even be caught by hand with a stealthy approach. A female was observed showing ovipositing behaviour around a Roystonea sp. palm at Diani Beach at about 18.00 h on 25 Aug 1995 [MJWC], and TCEC has observed adults flying at dusk rarely. Food plants. Sevastopulo (1975), Kielland (1990), Larsen (1991) and Heath et al. (2002) list Borassus, Phoenix and Raphia spp. palms as food plants. Sevastopulo may not have reared this species himself (Sevastopulo unpublished), so his records may have been derived from others. If these observations were from the Kenya coast, as seems most likely, the palms in question are most probably B. aethiopum, P. reclinata and R. farinifera (see under Zophopetes dysmephila above). However, MJWC’s experience has been that this subspecies normally feeds on Cocos nucifera at the Kenya coast (Larsen 1991). MJWC also found caterpillars on an ornamental palm he identified as Roystonea sp. (Diani Beach, 22 Aug 1995, 95 / 109; Figure 29), but noting that in captivity the caterpillars were feeding much more slowly than another collection on C. nucifera, changed the rearing to C. nucifera. At inland sites in Tanzania, TCEC found this species on R. farinifera and Elaeis guineensis (oil palm). At Shiwa Ng’andu, Zambia, Ivan Bampton and TCEC found early stages on R. farinifera. We consider, therefore, that food plant records from Borassus and Phoenix should be treated as unconfirmed and unlikely to be normal food plants. Larsen (2005) and Vande weghe (2010) treat G. carmen in West Africa and Gabon respectively (with no subspecies) but the food plants that they list are those recorded from eastern Africa; Borassus, Cocos, Phoenix, and Raphia. Thus we have no food plant records that are associated with the ssp. carmen (or the western population of G. carmen, if it were treated as a monotypic). Ovum. The ovum is slightly more pointed than hemispherical, with a dark spot on the top. Leaf shelters. I n Kenya, the fourth and fifth instar caterpillars rest in a rolled C. nucifera leaflet, with the edges held together by strands of silk. Pupation is in a similar shelter. TCEC noted in Tanzania that the caterpillars cut and roll the edges of palm leaves to make a shelter from which to feed. They sometimes use the dead, curled ends of leaflets, and are then very hard to find. Caterpillar. Based on a limited number of observations of the early instars, there seems to be six instars in Kenya. The first instar is 4 mm long, white with a black head, 0.8 x 0.6 mm wide x high (n = 1); it eats the shell of the ovum and three days after hatching moults without further feeding. The second instar is 4.5 mm when newly moulted; it has a light brown head, 1.1 x 1.2 mm wide x high (n = 2), dark centrally and ventrally, and covered with long dark setae, mostly about 0.3 mm long, but up to 0.8 mm; as it grows, white wax powder develops laterally on the face; the body is light green with a row of eight quadrate, white dorsolateral spots on A 2 – A 10; covered with long pale setae. In the third instar (Figure 30.1) the head is 1.7 x 1.8 mm wide x high (n = 1), pale brown, broadly dark along epicranial suture, on adfrontals and frons; T 1 matt dull green; body matt dark green dorsally, sharply defined by a broad subdorsal-lateral line of white waxy powder, diffusely defined ventrolaterally; the white waxy line becomes more pronounced during the course of the instar; head and body covered with long pale erect setae. The head measures 2.4 x 3.0 mm wide x high (n = 1) in the fourth instar (Figure 30.2); light brown, adfrontals and frons darker; long pale setae up to 1.5 mm long, darker in ventral area; white waxy powder spreads over much of the head as the caterpillar grows. The only fifth (penultimate) instar caterpillar for which a description was prepared had head markings rather like those shown in Figure 33.2 from Tanzania; 2.9 x 3.5 mm wide x high (n = 1); ground colour pale brown; broad dark line with diffuse edges along epicranial suture from vertex to adfrontals, continuing much narrower along dorsal half of adfrontal suture; a contiguous diffuse dark spot on the epicranium adjacent to the top of the adfrontals; dark spot covering dorsal third of frons and adjacent adfrontals; setae minimal on dorsal half of head, but long on ventral half, around 1.2 mm, and up to 1.8 mm long; a heavy vertical stripe of white waxy powder on epicranium near adfrontals, running from level with top of adfrontals to bottom of frons, interrupted at the level of the bottom of the dark spot in the frons. The head of the final instar of this individual was plain pale brown. The head of the sixth and final instar caterpillar measures 3.7 x 4.5 mm wide x high (range 3.4 – 3.8 x 4.4 – 4.7, n = 6). In some cases there are black markings on the light brown head (Figure 31.1), but the head is usually plain light brown (Figure 31.2 – 3). The following description of the final instar is based on individual 88 / 77 D, collected as a penultimate instar at Bamburi Beach, Kenya, moulted to final instar 18 Sep when it was described. Length 33 mm; Head oval slightly wider basally; semiprognathous so that dorsal part of head covers T 1; light brown with dark line along epicranial suture; dark spot at top of frons in centre of face; stemmata dark; between mouthparts and stemmata a weakly differentiated yellowish bar; head surface matt, very slightly rugose, covered with long, fine white setae, especially ventrolaterally. Body ground colour pale greenish white; narrow pale dorsal line bordered by darker subdorsal line and pale yellow dorsolateral line; body with long, 2 mm, fine white setae, especially dense on A 9; T 1, legs and spiracles concolorous. By 28 Sep, the sides of the head and body, including the pale yellow dorsolateral line were covered with white waxy powder (cf. Figure 32), and the caterpillar pupated 2 Oct. The prepupa becomes covered with wax before pupation. One caterpillar was recorded to take 18 days for the fifth instar. The caterpillars TCEC documented from Tanzania are rather different. An early instar caterpillar from Tukuyu, south-western Tanzania has the head dark with a pale brown anterolateral stripe, unlike any observed in Kenya (compare Figures 33.1 and 30.1). The final instar caterpillar from Kigoma Town, western Tanzania (Figure 33.3) lacks the black line along the adfrontal suture seen in the caterpillars from Tukuyu, south-western Tanzania (Figure 33.2,4 – 5), but both are much more heavily marked than those shown above from coastal Kenya (Figure 31). Furthermore, Figure 33.2 from Kigoma Town indicates yellow interrupted dorsal and dorsolateral lines on the anterior body, whereas one of those from Tukuyu has a solid yellow dorsal line on the anterior abdomen segments (Figure 33.2), while the other has five or six brownish dorsal marks on the abdomen (Figure 33.4 – 5). More observations are needed to assess the amount of variation in caterpillar markings at each locality.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32311B6EAB9F5C9EFED3E5F8.taxon	materials_examined	Pupa. The pupa (Figure 34) is formed in a single rolled leaflet, and supported by a band of several strands of silk around the thorax. The pupa and the inside of the shelter are covered with white waxy powder. Individual 88 / 77 C pupated 8 Sep 1988 and was described 18 Sep; 23 mm; smoothly contoured, no projections; short fine pale setae dorsally; longer (> 1 mm) on head and anterior and posterior third of the eyes; ground colour under translucent cuticle light green; as the pupa develops it turns paler, then brownish, especially the thorax and appendages and the eyes become red; T 1 with dark dorsal line; T 2 a thick dorsal dash near posterior margin; T 3 a faint short dorsal mark; A 4 – A 6 two pairs of transverse dashes each side of slightly darker dorsal line; similar but faint, diffuse marks on A 3, A 7; spiracle T 1 inconspicuous, a short brown vertical line at base of forewing; other spiracles light brown, inconspicuous. The black markings are variable (cf. Figure 34). The pupal stage lasts 17 days (range 14 – 20).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32311B6EAB9F5C9EFED3E5F8.taxon	discussion	MJWC reared a female of a solitary species of Brachymeria (Chalcididae) from a field collected pupa at Bamburi Beach, Kenya (6 Sep 1988, 88 / 77 B). This individual survived for a month fed on honey, but would not parasitize the pupae of Helicoverpa armigera (Hübner) (Noctuidae) which were exposed to it. A different, gregarious Brachymeria sp. (apparently the same species as reared from Z. cerymica above), was also reared from field-collected pupa from Diani Beach: eight adults emerged from one approximately 2.5 mm diameter hole cut through the wing sheaths of one pupa (26 Mar 1989, 89 / 19 C), while 15 adults emerged from three holes (two of 1.4 mm and one of 2.6 mm) cut in the thorax of another (23 Aug 1995, 95 / 107 D). Similar material from coconut, Watamu, coastal Kenya (Oct 1993, SCC) is preserved in ABRI. Empty field-collected pupae with similar emergence holes of diameter about 2.5 mm were found on more than one occasion at the Kenya coast, which could have been due to either of these Brachymeria spp. Finally, from material collected on Raphia farinifera at Shiwa Ng’andu (northern Zambia), TCEC reared a large yellow and brown ichneumonid larval pupal parasitoid, similar to that reared from Ploetzia amygdalis (above); it too emerged by pushing the head off the host pupa. Discussion. As noted in the introduction to this species, at present the scattered records from Kenya, Tanzania and Zambia suggest isolated populations. From what we know now of the palm food plants (above), it seems quite likely that further collecting will show that these populations are connected, at least along rivers. We have noted geographical variation in adult phenotype between these populations, and we document some differences between caterpillars from different areas, in particular between the Kenya coast population and those of Tanzania and Zambia. However, we have noted that there is some variation in body and head markings amongst the final instar material from the Kenya coast (Figure 31), but we don’t have this information for Tanzanian and Zambian populations, so cannot assess to what extent the differences in markings may overlap when variation is taken into consideration. As yet, we have no observations from the western part of the range of C. carmen. Thus, our observations do not resolve the question of whether this is a single variable species, or whether there are distinctive geographically separated populations, which may or may not merit subspecies status. More detailed observations and documented individual rearing from several areas should help clarify things, and the use of molecular methods should also help.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32081B6FAB9F5F34FBDEE72E.taxon	description	The associated final instar head capsules are more or less covered with white waxy powder; the body is covered with long white setae; the ventral half of the head has downward directed pale setae, but the dorsal half is more or less clear. The head is pale brown; the one from Côte d’Ivoire has the epicranial suture black and the adfrontal suture dark; there is an irregular black mark at the apex of the frons, but no markings on the epicranium. Any markings on the Ghana specimen are obscured by the heavy white waxy powder.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32091B63AB9F5A0FFC6AE54D.taxon	description	There are two subspecies recognised for Gretna balenge: the nominate subspecies which Holland (1891) described from Benita, Gabon, occurs from Nigeria to Uganda, western Tanzania, much of DR Congo, and north-western Zambia, while ssp. zowa Lindsey & Miller was described from Liberia and is found from Sierra Leone to the Volta area (Lindsey & Miller 1965, Larsen 2005). In Tanzania, ssp. balenge is now known from Bukoba, Kigoma and Mpanda. Adult behaviour. Carcasson (1981) notes that it flies at both dawn and dusk. Larsen (2005) reports ABRI collectors catching this species coming to ‘ various kinds of foul bait’ near collecting camps in Cameroon and Central African Republic. In TCEC’s experience this is not a true forest species, but occurs wherever palms grow, including open situations on river banks, etc. Food plants. TCEC has reared this species from Raphia farinifera in Tanzania, and Eremospatha sp. rattan palms in north-western Zambia. The latter is the climbing palm food plant in Zambia referred to by Larsen (2005) and Vande weghe (2010). Motshagen (2013) has reared it from the ornamental Madagascan palm Dypsis lutescens in his garden at Port Harcourt, Nigeria (Figure 37.3). Ovum. The ovum (Figure 38) is very large, even for a palm-feeding skipper. It is rounded, wider than high, the widest part being at slightly less than half the height. The surface is covered with faint, irregular polygons dorsally, merging into irregular faint striations laterally.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32091B63AB9F5A0FFC6AE54D.taxon	description	Caterpillar. The newly hatched caterpillar (Figure 39.1) eats the whole of the egg shell and then spins a shelter in which it moults without feeding. Figure 39 shows a caterpillar trying to make a shelter between the bottom and side of the plastic rearing container (behind which a leaf has been placed). Normally, this behaviour would pull closed a leaf shelter made from a single leaflet, but because the container is rigid, the caterpillar remained in view where it could be photographed. It does not start to eat the palm leaf until it has changed into the second instar (Figure 39.2). Second and subsequent instar caterpillars have scattered long setae; the body and setae are covered with white waxy powder, giving the impression of some fungus-infested body (Figures 39.2, 40.2 – 3). Much of the white wax was removed to reveal the underlying pattern of the final instar caterpillar in Figure 40.3; white with an interrupted dark dorsal line and brown head. Pupa. The pupa is very unusual in having a pair of large, elaborate processes frontally on the head, reminiscent of the head of an ant lion (Figure 41). The upper part is in two halves, each arising just above the eye, and extending dorsally well above the head, and spreading laterally and anteriorly in a broad but narrowing arc, the points ending up directed towards each other, separated by about the width of the lower part, but well above and anterior to it; this lower part projects forward to a pair of points separated by a broad, smooth notch; just ventrally and laterally to the base of this lower part is a small tuft of erect setae (Figure 41.6). There is a similar tuft on the anterior margin of the eye, centred on a small projection (Figure 41.4 – 5). When disturbed, the pupa vibrates violently and rattles quite noisily against the sides of the shelter, behaviour probably to deter predators or parasitoids. We think the rattle is made by the frontal processes striking the sides of the shelter, not the thoracic segments, and the vibration is a result of flexing the abdominal segments, but we do not have specific observations on this.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32091B63AB9F5A0FFC6AE54D.taxon	discussion	Discussion. This skipper and its life history are remarkable in several ways. It is amongst the largest of African skippers, ‘ a monster of a butterfly, possibly the most muscular of all African butterflies’ (Larsen 2005). The caterpillar, covered with white waxy powder, forming projections based on the long setae, is unique amongst African skippers known to us. The complex frontal projections of the pupa are unparalleled in our experience, while the pupa’s ability to vigorously vibrate in its shelter has not been noted in other African skippers. The nearest parallel that we are aware of is the Asian species Gangara thyrsis (Hesperiinae, incertae sedis), which is a large, polyphagous palm feeder, described in detail by Bell (1925). It too has a caterpillar covered with white waxy powder, which forms projections (Figure 42.1). The mature caterpillar makes an unusual pupal chamber by rolling one or two palm leaflets in a spiral, forming a rigid tube (Figure 42.3), and within this, when disturbed, the pupa (Figure 42.2) vibrates rapidly and violently, drumming against the sides of the shelter, which is certainly a deterrent to an inexperienced predator (or indeed entomologist in MJWC’s personal experience), and may also be a behaviour to crush any intruding predator or parasitoid insect (note the heavily chitinized spurs laterally before the cremaster, discussed under Zophopetes cerymica above).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32051B63AB9F58F3FB76E63D.taxon	description	Watson (1893) fixed P. laufella as the type species for this genus, when he established the genus Pteroteinon. There are ten species of this Afrotropical genus, which is restricted to West and central Africa (Larsen 2005, Vande weghe 2009). All reported food plants are palms, and P. laufella is a recognised minor pest in West Africa.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32051B65AB9F5BFEFDB7E271.taxon	description	This species forms a part of a small group of closely related species, the others being P. ceucaenira (Druce) and the recently described P. concaenira Belcastro & Larsen (Larsen 2005) and P. komo Vande weghe from Gabon (Vande weghe 2009). The first three species have clear genitalia differences (T. B. Larsen pers. comm. 2013) but the genitalia of P. komo have not been documented yet. Pteroteinon caenira (Hewitson) (Figure 43) is found from Sierra Leone, though West Africa to Central African Republic, DR Congo, western Uganda and northern Zambia (Larsen 2005). Previous reports of P. ceucaenira in Tanzania (e. g. Congdon & Collins 1998) are incorrect, the specimens actually being a new subspecies of concaenira or caenira, or even a distinct species (T. B. Larsen pers. comm. 2014). Food plants. Vuattoux (1999) reports rearing this species from introduced palms only in Côte d’Ivoire (Phoenix dactylifera and Washingtonia filifera) but as he notes, it can be expected to have indigenous palms as food plants as well. MJWC (in Larsen 2005) found early stages of this species on an ornamental palm at Ibadan, Nigeria (94 / 101); at the time, this was thought to be an ‘ Areca sp. ’, but most likely was Dypsis lutescens. Larsen (2005) notes that these records may refer to any of the three species of this small complex of Pteroteinon spp. MJWC found one final instar caterpillar (94 / 101 A), a pupa which died formed up (94 / 101 B, Figure 45.3), and two dead pupae (94 / 101 C, D), one of which was missing the head probably due to a predator, but had fully formed up showing the distinctive male forewing markings of P. caenira (94 / 101 D, Figure 45.4). The other pupa (94 / 101 B, Figure 45.3) showed markings compatible with female P. caenira; MJWC associated all four because of the similarities of the pupae and because they were collected on the same palm at the same place at the same time. Nevertheless, the association of the caterpillar although most likely, is not definite. Leaf shelters. The final instar caterpillar (94 / 101 A) was in a silk lined shelter made from a single leaflet; a 20 mm wide notch was cut to the midrib basal to the shelter; the shelter was 60 mm long, formed by rolling the leaflet downwards; feeding was distal to the shelter and the leaflet apex had been consumed. The live pupa (94 / 101 B) was in a shelter formed from two leaflets, the edges held together and strengthened with short bands of silk along the join, which were duplicated along the midrib (probably to roll the leaflet into shape); it was lined with silk and white waxy powder. Caterpillar. The final instar caterpillar (Figure 44) measured 30 mm; head 3.2 x 4.3 mm wide x high (n = 2); oval, indented at vertex; shiny, rugose; uniform light chestnut brown, except for dark stemmata. T 1 pale anteriorly, dark posteriorly; pronotum concolorous in posterior half. Body dull whitish green; dorsal line dark green, bordered by yellow-white subdorsal line; spiracles and T 1 legs dark; T 2 – 3 legs and prolegs concolorous; gonads not evident. The caterpillar was noted to flick its frass at least 25 cm horizontally. The caterpillar of P. concaenira (Figure 48) is very similar; there are no clear-cut differences, so the possibility of the two species occurring together in MJWC’s Nigeria collection 94 / 101 cannot be ruled out. Pupa. Collection 94 / 101 B was a 25 mm pupa (Figure 45.1 – 3); elongate, 4 mm wide at front of thorax; short, blunt, dark, bifurcate frontal projection, 1.1 mm long, 1.9 mm between the divergent tips, a few pale brown setae forward directed on anterior surface; proboscis sheath projects to cremaster tip; head and thorax brown; abdomen yellow brown with slightly darker dorsal line, paler ventrally; spiracles slightly darker than surrounding ground colour. The adult formed up in this pupa, but failed to emerge: spots were visible in the forewing in cell, spaces 2, 3 and 6 – 8, i. e. compatible with female P. caenira. Pupa 94 / 101 D was already dead, with the head missing (? eaten), but the forewing markings (Figure 45.4) clearly indicating its identity as a male P. caenira. Natural enemies. Pupa 94 / 101 C was empty and contained a single light grey-brown cocoon in which was a fully formed wasp. The front part of the pupa was missing (similar to 94 / 101 D, Figure 45.4) and fitted directly onto the front of the pupa was a mud-dauber wasp cell, which contained an adult wasp and cocoon remains similar to the last. This was opportunistic use of the empty pupa as a suitable niche for a mud cell, but could easily have been misinterpreted as a parasitism event.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC32031B67AB9F5FAAFAEDE360.taxon	description	This species was described from Ghana (Belcastro & Larsen 1996) and is found from there, Côte d’Ivoire, and Nigeria to western Uganda, north-western Tanzania and Zambia (Larsen 2005). Pteroteinon concaenira is distinguished from P. ceucaenira and P. caenira by the presence of a white forewing band in the female, the other species having white spots. The male of nominate P. concaenira has a narrower forewing band than P. caenira. The Zambian material (Figure 46) appears to represent an undescribed subspecies, having a much wider male forewing band than other populations. In Tanzania it is found in Kikuru and Munene forests, Kagera Region. Food plants. TCEC found this species in western Zambia feeding on Raphia farinifera (Heath et al. 2002), and anticipates that it feeds on the same plant in Tanzania. Doubtless, this is the source of the record from Raphia in Vande weghe (2010). Life history. An ovum (Figure 47), caterpillars (Figure 48) and an emerged pupa (Figure 49) are shown. The caterpillars pull the sides of the end of a leaflet into a tube, in which they rest, eating the leaflet lamina basal to this. This leads to a much longer and more untidy tube than is made by other palm-feeders such as Zophopetes spp.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321E1B78AB9F59EDFCCDE2D6.taxon	description	Hewitson (1867 – 1871) described Hesperia laufella from Akwa Akpa (= Old Calabar, south-eastern Nigeria), and it is known from Sierra Leone, east to Cameroon, and south to northern Angola and adjacent DR Congo (Larsen 2005). Larsen (2005) had no records from Benin, but many have since been collected there by A. Coache (T. B. Larsen pers. comm. 2013) and MJWC has a male from Cotonou (29 May 1989). There is a limited literature on it as a pest of oil palm in West Africa. Food plants. As early as 1919 and 1929, this species was already appearing in German texts as a pest of oil palm in Cameroon (Lepesme 1947). As discussed in Cock (2010 a) early reports of Pyrrhochalcia iphis (Drury) and Coeliades bixae (Linnaeus) (Hesperiidae, Coeliadinae) feeding on palms are likely to be misidentifications for Pteroteinon laufella. Thereafter, Pteroteinon laufella reappears in the economic literature for French West Africa. As discussed above under Zophopetes cerymica, Mariau & Morin (1974) report on two hesperiid defoliators of oil palm and coconut: Zophopetes cerymica (as Z. dysmephila) and P. laufella, but do not separate the two in their account of the damage, economic thresholds, and natural enemies. Mariau et al. (1981) and Mariau (2000, 2001) summarise information on P. laufella as a pest of oil palm and coconut. In Côte d’Ivoire, Vuattoux (1999) reared this species primarily from oil palm (Presque tous les adultes obtenus ont été élevés sur Elaeis guineensis), and only two specimens from coconut; these two food plants are repeated in Vande weghe (2010). Life history. The published life history information (Mariau & Morin 1974, Mariau et al. 1981, Herder et al. 1994, Mariau 2000, 2001) lacks detail, but does include some photographs. The ova are laid in batches; Mariau et al. (1981) show a batch of 17 ova, which have a dark dorsal circle around the micropyle.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321E1B78AB9F59EDFCCDE2D6.taxon	discussion	The caterpillar is illustrated in black and white in Mariau & Morin (1974), Mariau et al. (1981) and Herder et al. (1994), and in colour in Mariau (2000), which is reproduced here as Figure 50.3. The head is black with a broad brown stripe from vertex to clypeus, narrow at the vertex and widening evenly to include the adfrontals and part of the adjacent epicranium ventrally; conversely, Mariau & Morin (1974) describe the head of the caterpillar as golden brown (like Z. cerymica) with broad brown patches on the sides (avec de larges taches brunes sur les côtés). T 1 white, black pronotum in the posterior half; spiracle conspicuous, orange-brown; legs black and white. Body greenish white; spiracles pale brown; legs brown; prolegs not visible. The pupa is dark in colour, with a distinctive bifurcate frontal projection (Mariau & Morin 1974, Mariau et al. 1981), similar to that of P. caenira (Figure 45.1 – 2). Natural enemies. Mariau & Morin (1974) do not distinguish between P. laufella and Z. cerymica, when they discuss the natural enemies of these two skippers. However, more recently, Mariau (2001) treats all the same natural enemies under P. laufella and mentions none for Z. cerymica, so we follow this arrangement here. Although these two palm-feeding skippers may well have natural enemies in common, this should not be assumed. The ova are parasitized by a Tetrastichus sp. and the pupae are host to tachinids, an ichneumonid and two chalcids: Brachymeria sp. and Stomatocerus sp. As discussed in Cock (2010 a), the true host of the egg parasitoid Telenomus thoas Nixon (Scelionidae), described from ova of Pyrrhochalcia iphis, is probably Pteroteinon laufella.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321E1B78AB9F59EDFCCDE2D6.taxon	discussion	Economic damage. See under Z. cerymica, regarding the comments in Mariau & Morin (1974), Mariau et al. (1981) and Mariau (2000), which treat P. laufella and Z. cerymica together. Herder et al. (1994) refer to regular outbreaks of P. laufella on oil palm in southern Côte d’Ivoire.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321F1B79AB9F5D97FE44E3BE.taxon	description	Evans (1937) introduced this genus, which he considered closely related to Pteroteinon and Caenides Holland, for nine species which previously were included in Caenides. Although he distinguished them by their ‘ antennae which are white or ochreous above’, he also noted that the male genitalia indicate a cohesive group. Lindsey & Miller (1965) concluded that the antennae and genitalia do not separate the two genera, and Leona and Caenides are congeneric, Caenides being the senior name. They recognised two groups within the combined genus: the dacela group with a hair tuft UPH between M 3 and Cu 2 (Evans refers to ‘ overlying the bases of veins 2, 3 and 4 ’) and the leonora group without, which align partially, but not completely, with Evans’ Caenides and Leona respectively. Carcasson (1981) and Ackery et al. (1995) follow Lindsey & Miller (1965), but Larsen (2005) reverted to Evans’ (1937) arrangement, and we follow this here as the most recent comprehensive treatment. It is likely that an analysis of the early stages and food plants will throw light on this question, but as yet, those of relatively few species are known, and the records for both genera include palms and softer-leaved monocotyledons such as Zingiberaceae. Leona halma Evans is recorded from Zingiberaceae: Amomum (Kielland 1990) and Aframomum (Heath et al. 2002, Vande weghe 2010), although it is not impossible that one of these is an error for the other, most probably the former for the latter. Heath et al. (2002) suggest that the food plant of L. leonora dux Evans is possibly also Zingiberaceae. Leona lissa Evans has been reared from Dracaena sp. (Asparagaceae) (SCC in Larsen 2005, Vande weghe 2010), but the genitalia are so different that it is probably not congeneric with the larger Leona spp. (T. B. Larsen pers. comm. 2014).	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321D1B7BAB9F5BADFC5AE122.taxon	description	Caenides, as currently defined, contains species which feed as caterpillars on Arecaceae, Zingiberaceae, Costaceae and Colchicaceae (some lilies). The palm-feeder C. dacela is treated here, whereas C. dacena (Hewitson) has been reared from Costus, Costaceae (Vuattoux 1999, Vande weghe 2010, MJWC unpublished observations), and will be treated in a subsequent part of this series. Furthermore, we note, and will document in this future part, that the early stages of these two species are sufficiently different to indicate that they are not congeneric. T. B. Larsen (pers. comm. 2014) advises that he expects to move C. dacena to a genus of its own, close to Hypoleucis and Semalea, based on the structure of the male genitalia. Other workers provide additional insight: C. kangvensis Holland has been reared from the lily Gloriosa superba (Colchicaceae, formerly Liliaceae) and Thalia welwitschii (Marantaceae), but the early stages were not documented (Vuattoux 1999, repeated in Vande weghe 2010). Larsen (2005) expresses surprise at the record from a lily (Liliaceae), being aware of no other records from the family, however, MJWC found a skipper caterpillar (89 / 202) on what he thought was a Liliaceae in Côte d’Ivoire, but failed to rear it through or preserve a sample of the food plant. Caenides soritia (Hewitson) has been reported from an undetermined Zingiberaceae (Sevastopulo 1975, unpublished) and reared once from Elaeis guineensis in Côte d’Ivoire (Vuattoux 1999); these contrasting food plants which are repeated in Vande weghe (2010), need confirmation.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
6F3587EC321D1B7DAB9F5CF8FC38E2AF.taxon	description	Food plants. Le Pelley (1959), Sevastopulo (1975) and Kielland (1990) give P. dactylifera as the food plant, the first reference referring to Uganda. Heath et al. (2002) refer to a specimen reared by TCEC and I. Bampton, and give the food plants as P. dactylifera and Raphia spp. (repeated in Vande weghe 2010), but since TCEC did not rear C. dacela from these palms in his collaboration with Bampton, we discount these food plant records. In Kakamega Forest, Kenya, MJWC collected this species from P. reclinata (not P. dactylifera as stated in Larsen 1991). TCEC reared it from the same food plant in western Tanzania (forests of Mpanda, Kigoma and Kagera), but from a rattan palm, Eremospatha sp. in northern Zambia. Leaf shelters. The first instar shelter (Kenya) is made either by cutting two notches from the edge of a leaflet and folding the resultant flap over upwards, or by cutting a notch from one side just before the apex of a leaflet, and pulling together the distal portion. The shelter of the young caterpillar (Zambia) is in the tip of a tatty old leaflet, and the caterpillar is coloured to match (Figure 54.1). The shelter has been opened, and the caterpillar is in the process of pulling it shut again. From a human perspective, this is not the sort of place one would look to find a caterpillar, so may be a good survival strategy for the caterpillar. The fifth instar caterpillar forms a shelter between two leaflets. Ovum. Two ova collected on Phoenix in association with early stages of C. dacela (Alupe, near Busta, western Kenya, 18 Jun 2006, SCC) are thought to be C. dacela as they are not Zophopetes dysmephila, the species commonly found on Phoenix spp. These eggs are 1.5 mm diameter, dome-shaped, with 31 weak ribs. Caterpillar. The following notes are based on collection 90 / 76 on P. reclinata, north Kakamega Forest. There are five instars, the head capsules measuring 0.8 x 0.8, 1.2 x 1.2, 1.7 x 1.7, 2.4 x 2.7 and 3.7 x 4.0 mm wide x high, respectively. The first instar has a uniformly dark head. The second instar caterpillars have a light brown head with the posterior margin dark; body green. The third instar is shown when newly moulted in Figure 53.1: 8 mm; head light brown, reticulate pattern faintly apparent on front of epicranial, posterior margin dark; body dark brown dorsally, light brown ventrally. The fourth instar, also newly moulted, is shown in Figure 53.2: head similar to previous instar, but reticulate pattern slightly more evident; faint trace of lateral band. The mature fifth instar is shown in Figure 53.3 – 4: 32 mm; head with broad brown stripe down centre; light brown band down anterolateral margin; light brown posteriorly; dark posterior margin. Body dull grey; dorsal line dark; subdorsal and dorsolateral lines dull yellow; paler ventrolaterally and ventrally; anal plate paler on posterior margin; all legs concolorous; spiracles light brown, inconspicuous. The wax glands develop as a continuous ventral mass from A 1 to A 8. Two caterpillars collected in the first instar moulted after 6 days, and the following instars lasted 6, 8, 10 and 17 – 21 days respectively. The caterpillars which TCEC documented from Tanzania and Zambia (Figure 54) are compatible with those found in Kenya by MJWC (Figure 53). Pupa. The pupa is formed in a shelter of one or two leaflets, and held with a single strand silk girdle (Figure 55); rather uniformly brown, no setae evident. The inside of the shelter and the cast final instar skin are well covered with white waxy powder, but the pupa only lightly so. The pupal stage lasts 20 – 24 days under Nairobi conditions. Natural enemies. One of two ova collected by SCC at Alupe, western Kenya, 18 Jun 2006, had an egg parasitoid (probably Trichogrammatidae) exit hole of 0.35 mm diameter.	en	Cock, Matthew J. W., Congdon, Colin E., Collins, Steve C. (2014): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa 3831 (1): 1-61, DOI: http://dx.doi.org/10.11646/zootaxa.3831.1.1
