taxonID	type	description	language	source
DB357E4FAB0FFF87FF1DB6EBCDD5FE43.taxon	description	Coeliades keithloa (Wallengren): Acridocarpus natalitius Coeliades forestan (Stoll): Combretum apiculatum, C. bracteosum, Solanum auriculatum, Millettia sutherlandi Coeliades pisistratus (Fabricius): Acridocarpus pruriens Similarly, two small papers with biology information were missed. Sevastopulo (1964) reports that Coeliades sejuncta (Mabille and Vuillot) and C. anchises (Gerstaecker) are attracted to light by night (see also under Baorini below), and Van Someren (1955) reports C. forestan hilltopping, ‘ basking in the sun or chasing any intruder out of its particular territory’.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB0CFF82FF1DB098CD42FD4E.taxon	description	The caterpillar pupated in a semi-transparent plastic container, and R. de Jong took a sequence of images from the day of pupation until the day before emergence. On the day of pupation (Figure 3.1) the caterpillar colours were retained by the pupa on the abdomen, although the thorax and head were red like the head of the caterpillar. The ventral ground colour is pale orange and the black markings are fixed on the day after pupation (Figure 3.2). The ground colour becomes slightly paler over the next 14 days (Figure 3.3 – 5). The day before emergence the head, thorax and wings of the pupa are white, but this appears to be due to a white waxy efflorescence on the surface of the pupa (it is abraded between the wings). There is no sign of the brightly coloured body of the adult (Figure 4) as the pupa cuticle is opaque. This sequence of changes is not unusual amongst Hesperiidae, although the brightly coloured caterpillar markings of the newly formed pupa are especially striking. For those species with a translucent pupa cuticle the changes may be more obvious, especially as the adult colouring becomes visible. Illustrating or describing a pupa based on one occasion can be misleading, when pupae of different ages are compared. We have been aware of this during the course of this sequence of papers, and so have not normally illustrated newly formed pupae or pupae about to emerge, and given the dates of pupation and emergence when recorded.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB0BFF80FF1DB2D7CFFFF9A6.taxon	description	The penultimate instar caterpillar is black with white and yellow markings (Figure 5). Head black, matt, rugose. Pronotum black with a white transverse band, interrupted dorsally. Body black with narrow white subdorsal line T 2 – A 8; stronger yellow dorsolateral line, interrupted T 2 – A 1, thicker and continuous A 2 – A 8; large yellow spot laterally in anterior half of A 3 and A 5, similar but smaller spot on A 1; a short yellow bar extending from dorsolateral line on posterior margin of A 8; 3 – 4 transverse rows of pale dots in posterior half of A 1 – A 6; yellow and white lateral line, thick and yellow T 1 – T 2, thin and white T 3, white with a yellow section in middle of segment, A 1 – A 8; ventrolateral white line. The final instar is a more heavily marked version of the penultimate instar with a red-brown head (Figure 6). Head red-brown, matt, rugose; diffuse dark line each side of epicranial suture, extending indistinctly adjacent to adfrontal suture. Pronotum black, with anterior and posterior margin narrowly white; T 1 laterally yellow in line with lateral yellow and white line of body. Body dark, but not black; scattered short, pale, inconspicuous setae; subdorsal white line T 2 – A 8; thick, yellow dorsolateral line T 2 – A 8, pale yellow towards posterior margin of each segment, pale yellow A 8, diverging towards posterior margin; a short white bar extends laterally from dorsolateral line, near posterior margin of T 2 – A 2 and on posterior margin A 8; three lateral rows of white dots adjacent to the posterior margin of A 1 – A 6, the most anterior of which on T 3 – A 2 are in line with the white bar just mentioned; conspicuous yellow spots in anterior half of A 1, A 3 – A 5 and A 7, those of A 3 and A 5 being significantly larger; a thick yellow and pale yellow lateral line; a thick white ventrolateral line, the area between this and the lateral line speckled white; anal plate unmarked; legs brown; prolegs white-brown; spiracles brown, in lateral line. The pupa is light brown, almost completely covered with a uniform layer of white wax, interrupted with scattered bare dots on abdomen; short, blunt, black, upturned, frontal spike; prominently protruding black T 1 spiracles, and a dark dot slightly dorsal to these; other spiracles black, conspicuous; inconspicuous, erect, pale setae dorsally on thorax. The pupa is attached at the cremaster and supported by a Y-shaped silk girdle. The posterior margin of A 4 and anterior margin of A 5 are curiously striated dorsally; we have not noticed such a pattern before and have no evidence to suggest what function it may have. These caterpillar descriptions and figures of C. ramanatek ramanatek are very different to those in Cock (2010 b) based on blown caterpillars of C. ramanatek comorana Evans in the BMNH. So much so, that we suggest either these two subspecies are two different species with very different caterpillars, or there is additional cryptic diversity within the species, or the material in the BMNH is misidentified. If caterpillars of C. ramanatek comorana can be found and documented, this should help to clarify the position. SCC and Ivan Bampton found this subspecies on a Trema sp. at LaGrille, Grande Comoro in 1991 but it was not documented. Alain Gauthier (pers. comm. 2015) has searched on T. orientalis in the Comoros Islands and found an empty pupa, almost certainly of C. ramanatek comorana. The caterpillars of C. ramanatek ramanatek do not show any close affinities with any other species of Coeliades, suggesting that it is has no close relatives, rather than that it is the Madagascan replacement for C. libeon (Druce) below, as suggested by Chiba (2009).	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB09FF8EFF1DB6BCCE2BFCBD.taxon	description	In the final instar (Figure 8) the head is black, with an irregular pale band across the dorsal part of the face, breaking up into spots ventrally, extending to the posterior margin laterally and along the epicranial suture to the posterior margin. Pronotum black, with a broad whitish anterior margin. Body pale with variegated dark and yellow markings; black dorsal line, narrowly interrupted between segments A 1 – A 8; immediately adjacent a pale subdorsal line and a variegated darker area lateral to this; dorsolaterally a broad yellow line, more or less continuous in the upper half, but in the lower half, interrupted by a large black rectangular marking in the anterior third of each segment, separated into an anterior black rectangle, a white bar and then a dark bar posterior to this, and three irregular dark vertical lines in the posterior two-thirds; these dark markings are heavier on T 2 – T 3, almost continuous on T 2; there is a broad, black bar across the anterior margin of the anal plate, which is otherwise pale; laterally the body is pale, darkly diffused on T 1 – T 3; a white ventrolateral line; spiracles dark, positioned in lower part of white lateral area; legs black; prolegs pale. Cock (2010 b) reported the caterpillar descriptions of Fontaine (1988) in Zaire and R. Paré from Zimbabwe (in Pringle et al. 1994, Henning et al. 1997), expressing concern at the apparent differences between the two. Now in light of Figure 8, it can be seen that Paré’s ‘ lemon yellow with black chequering’ provides a reasonably succinct description, while features reported by Fontaine are also recognisable: a thin black longitudinal dorsal line edged on each side by a yellowish line; on each segment, a lateral pair of square blackish spots; thin, yellow, longitudinal spiracle line, bordered on the dorsal side with a wider, dark band; head and legs scaly black; prolegs yellowish. However, Fontaine’s reference to the upper surface covered with a very fine brownish down, and the dorsal region varies from brown-red to brown-grey are not clear. Nevertheless, we conclude that both published descriptions apply reasonably well to what we report here. The image of the pupa (Figure 9) is less satisfactory. It is pale with dark speckling which is quite heavy in some areas; the short, blunt frontal spike, T 1 spiracles and posterior margin of T 3 are dark; there are paler dorsal and dorsolateral lines on the abdomen. It is not clear to what extent the pupa is covered with a layer of white wax. Fontaine (1988) reported the pupa as having black veins on the wings, but these are not visible in Figure 9. In Cock (2010 b) it was noted that apart from C. libeon, all species of Coeliades and Pyrriades for which food plants have been reported include at least one record from a species of Malpighiaceae. This record of C. libeon from Flabellaria is therefore significant. However, the generalisation is still not complete as C. ramanatek is now reported from Cannabaceae only (above).	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB07FF8EFF1DB504C976F8F5.taxon	description	The final instar caterpillar of T. i. insularis (Figure 11) is very similar to that of T. flesus (Fabricius) (Henning et al. 1997, p. 65; Cock & Congdon 2011 a, Figure 21). The head of the Madagascan T. i. insularis is slightly darker in colour and the apices more acutely angled, while the body appears more intensely coloured. However, without more information on individual variation, it would be premature to assume this was significant. The pupa is very similar to that of T. flesus (Henning et al. 1997, p. 65; Cock & Congdon (2011 a, Figure 22), and immediately recognisable as a Tagiades sp. The pupa of T. i. insularis is darker brown, the shape of the brown wing markings differ, the distal half of the antenna is shining white in T. flesus, and light brown in T. i. insularis, and the shining white area on the ventral abdomen is much more extensive in T. flesus. However, as for the caterpillar, without examining more material, it cannot be suggested that these differences are significant.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB07FF8FFF1DB79ECD95FF1E.taxon	description	The emerged pupa is similar to that of T. i. insularis (Figure 12), but the white markings on the wings are less developed in this individual.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB05FF8AFF1DB754C918FE3E.taxon	description	Figure 13.3 is an enlargement of the freshly cut second shelter, which contained a second instar caterpillar. The edges of the cut are still green, as is the chewed material round the edge of the cut. Note that the caterpillar did not eat the chewed material. During shelter construction, the caterpillar is exposed and vulnerable to predators and weather; hence speed of shelter construction is the priority, and eating and digesting the material cut to make the shelter would have delayed completion. At this stage the flap has not been secured to the leaf surface, but already a tuck can be seen near the top of the flap in the figure. To make this, the caterpillar eats from the leaf upper surface (of the flap within the shelter) to weaken the underside of the roof so that it is easy to pull together with silk. It does not actually cut through the flap. Figure 13.4 is of the same shelter a day later. The flap has been secured to the leaf surface, and the lid on each side of the notch pulled together with silk (Figure 13.5) to form the entrance, and making the underside of the lid concave to form the shelter. The surface of the cut and the chewed material are now dry and brown, as is the flap itself. The final shelter shown in situ in Figure 14 was also documented on the same occasion. Figure 14.1 shows the shelter in details and Figure 14.2 shows the whole leaf. A more or less bell shaped flap was cut from the leaf margin, across one major vein and hinged on a second main vein. The bell-shaped flap was then swivelled underneath the distal part of the leaf held with silk. Silk threads seem to have been used to make both surfaces arched, creating a pocket between them. There is a deep notch in the flap cut across the base of cut main vein, and at the internal end of this, silk has been used to pull the sides together, contributing to the arching of the flap.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB02FF8BFF1DB097CE94FD48.taxon	description	Libert (2014) illustrates four caterpillars and three pupae of the galenus clade from TCEC’s work in Tanzania (Table 1). We have associated these with their corresponding caterpillars, pupae and, where possible, adults (Figures 15 – 17), and clarified some identifications.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB02FF96FF1DB5F3C957FB6A.taxon	description	One source of guidance in similar situations has been to compare their barcodes (e. g. Janzen et al. 2009, 2011). Libert (2014) states that barcodes are of little use to interpret the galenus clade. Nevertheless we compared his barcodes in BOLD (http: // www. boldsystems. org /) for these two species from Kakamega Forest: MLIB- 0033, MLIB- 0034, MLIB- 0044, and MLIB- 0046 for A. opalinus and MLIB- 0039 and MLIB- 0043 for A. kakamegae. We found that the two species in Kakamega Forest have identical or almost identical barcodes, which strongly suggests that they are an interbreeding population. Furthermore, specimens of A. rwandae Libert from Rwanda (MLIB- 0045 and MLIB- 0407) also have barcodes almost identical to those of A. opalinus and A. kakamegae from Kakamega Forest. Barcode Index Numbers (BINs) have been introduced to provide a permanent numbering system for barcode clusters which in a high percentage of cases correspond to known taxonomic species and can also help flag species complexes or clusters needing taxonomic research (Ratnasingham & Hebert 2013, Miller et al. 2016). They provide a useful surrogate for species in ecological diversity estimates but there are many known cases of BIN sharing between apparently good morphological species and different BINs representing deep splits where no morphological differences have been found (e. g. Hausmann et al. 2013, Zahiri et al. 2017). These specimens from Kakamega Forest and Rwanda have a single BIN: BOLD: ACE 5474, and appear to represent one species based purely on their barcodes. However, barcoded individuals of all three species from other regions have barcodes that match other species, often those co-occurring in those locations. Accordingly, it might not be wise to assume that A. opalinus from east of the Rift Valley (type locality Kikuyu), for which no barcodes are available, has identical barcodes to the material barcoded from west of the Rift Valley. Libert (2014) was aware of these types of problems with the barcodes of the galenus clade and considered many uninterpretable when presenting his conclusions based on wing markings and male genitalia. The galenus glade will require further work to clarify what is going on in terms of population dynamics and speciation to reconcile with the barcode patterns. In the meantime, the situation in Kakamega Forest could be made clearer for A. opalinus and A. kakamegae, by (1) the documentation of mating pairs, (2) rearing broods from individual females to see if they breed true or can produce both species, and (3) crossing the two species and documenting the progeny.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB1FFF97FF1DB4FBC8A8FEAE.taxon	materials_examined	As we anticipated (Cock & Congdon 2011 b), A. biseriata (Butler) is a valid species rather than a subspecies of A. galenus (Fabricius). The population from eastern Kenya which we treated as C. galenus biseriata (Cock & Congdon 2011 b, pp. 28 – 32, Figures 27 – 30) is now treated as A. biseriata maculata (Hampson), which was described from the Sabaki River, eastern Kenya (Hampson 1891), and occurs from southern Kenya to Zimbabwe (Libert 2014). Celaenorrhinus handmani Collins & Congdon which we treated as a valid species (Cock & Congdon 2011 b, pp. 32 – 34, Figures 31 – 33) is considered a synonym of A. biseriata maculata (Libert 2014). We collected both populations from Hypoestes forskaolii (Acanthaceae) and saw no diagnostic differences between the early stages. However, when the barcodes for this material are examined, it is not so clear-cut, as individuals named as A. biseriata maculata can be found within several different barcode clusters, including BINs BOLD: ABY 8703, BOLD: AAQ 3589, BOLD: ACE 5675, BOLD: ACE 3207 and BOLD: ABY 8700 (BINs are hereafter referred to by the last seven characters), mostly in isolation, but in one case alongside individuals of other species with identical barcodes. We note in particular that specimens from Mufindi, Tanzania, the type locality of C. handmani are placed in a separate BIN (ACE 3207) together with two specimens from Ethiopia which appear to be a different species with the same barcode. The bulk of Tanzanian specimens form a separate compact cluster (ABY 8703 from Kimboza, Kihansi, Rondo Plateau, Rubeho, Rondo and the Shimba Hills, Kenya). TCEC’s observations on A. handmani and A. biseriatus maculatus indicate allopatric populations at different elevations. Apallaga biseriatus maculatus occurs at lower elevations and seems to occur no higher than at 1800 m in the Chyulu Hills (Van Someren 1939). Apallaga handmani is a moderate to high elevation species, so their ranges do not overlap. Apallaga handmani was thought to occur as low as 1450 m on Mt. Mabu in Mozambique (at that latitude, the elevation is equivalent to 1800 m or more at the equator), but this population comes out as a different BIN (ACE 5675) so there may be additional allopatric diversity here. Again, more work is needed to understand this situation and reconcile taxonomy with barcode patterns (cf. A. opalinus and A. kakamegae above). However, we think the barcode difference justifies reinstating C. handmani stat. rev. as a good species, although not all the type series of A. handmani are likely to represent this species rather than previously unrecognised, very similar but genetically distinct, allopatric species, such as the population from Mozambique mentioned above.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
DB357E4FAB1EFF97FF1DB1B7C892FB54.taxon	description	Cock & Congdon (2012) overlooked the observation by Sevastopulo (1964) that B. borbonica can be attracted to light by night. In our experience, crepuscular species may be attracted to light at dusk or dawn. In addition, many species of Hesperiidae (and other butterfly families) are rarely attracted to lights by night, but we attribute this to individuals being disturbed and taking flight at night rather than true nocturnal behaviour.	en	Collins, Steve C. (2017): Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 12. New information and corrections. Zootaxa 4312 (3): 471-496, DOI: 10.11646/zootaxa.4312.3.4
