Hesperiinae

Cock, Matthew J. W., Congdon, T. Colin E. & Collins, Steve C., 2016, Observations on the biology of Afrotropical Hesperiidae (Lepidoptera). Part 9. Hesperiinae incertae sedis: Zingiberales feeders, genera of unknown biology and an overview of the Hesperiinae incertae sedis, Zootaxa 4066 (3), pp. 201-247 : 238-244

Discussion, Hesperiinae incertae sedis

This paper completes our treatment of Hesperiinae incertae sedis. We have treated this assembly of genera in five parts: dicotyledon feeders ( Cock & Congdon 2013), palm feeders ( Cock et al. 2013), grass and bamboo feeders (Cock & Congdon 2014), Dracaena feeders (Cock et al. 2015) and Zingiberales feeders (this paper). The 37 genera treated and their food plant families are summarised in Table 2 , leaving only 11 genera for which nothing is known of the food plants or life history.

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1 Families in bold were documented in our work, others are from the literature.

2 + from literature or museum specimens; ++ our observations; +++ photograph.

3 As noted above, T.B. Larsen (pers. comm. 2015) intended to create a new genus for this species. Evans (1937) placed it in Xanthodisca in Ceratrichia group, but its affinities are not clear.

Ova. A priori , it seems likely that differences between ova are likely to be more conservative, than any observed amongst caterpillars or pupae, which are likely to show adaptations to avoid predation and parasitism and related to their food plants and leaf shelters (e.g. Greeney et al. 2012). However, this is not necessarily true as mortality of ova has been shown to be important in pest Lepidoptera such as the polyphagous noctuid Helicoverpa armigera (Hübner) , where egg predators play an important role in Kenya (Van den Berg et al. 1999). MJWC noted that significant numbers of Artitropa erinnyis (Trimen) ova disappeared from a Dracaena steudneri plant in his garden in Nairobi (Cock et al. 2015). Hence, adaptations to maximise adhesion of ova to different types of plant surfaces, or minimise the ability of predators such as ants to remove ova are certainly possible. We have images for only 14 genera ( Figure 41 ), but they do seem to indicate some groups within the incertae sedis. The genera Parosmodes , Semalea and Platylesches are united by have having wall-like ribs; further, the ova of Parosmodes and Semalea are somewhat flattened compared to the relatively round ova of Platylesches , and the dorsal surface of the Parosmodes ova is rugose. Hence, there are significant differences between these three genera as well as a striking similarity. There is a group of genera with rather similar ova: dome shaped, with fine ribs, including Erionota , Zophopetes , Pteroteinon , Leona , Artitropa and the dicotyledon feeding genera Acleros and Andronymus . The similarity of the Asian Erionota and the African Zophopetes and similar genera is striking. Gretna balenge differs in that the ribs are very fine and the egg shape is bun-shaped (an oblate spheroid). The two grass feeding genera, Astictopterus and Ceratrichia are united by smooth dome-shaped ova, without apparent ribs. The bunshaped ovum of Gorgyra spp. differs from others, although this could be linked to its placement between leaves.

Caterpillars. The caterpillars of Hesperiinae incertae sedis do not divide as easily into groups ( Figure 42 ). In general, the dicotyledon feeders have stouter caterpillars, and the grass feeders tend to have more slender caterpillars. The species currently placed in Gretna can be grouped together because of the well-developed setae and white waxy powder on the head and body. Apart from this no clear groupings can be recognised, although Semalea and Caenides dacena have relatively narrow heads, Parosmodes and Melphinyet have distinct scale-like setae on the head (the similarities of the leaf shelters of these two genera have already been noted in Cock & Congdon (2013)), Acleros , Andronymus and tube shelter building Platylesches have strongly contrasting pale markings on the head, and so on.

Pupae. Representative pupae of the Hesperiinae incertae sedis are shown in Figure 43 . None are of the Baorini type, which is perhaps slightly surprising since other tribes of Hesperiinae do include genera with pupae of this type ( Cock & Congdon 2012). Possible groupings based on the pupae are not that obvious. Gorgyra and Pteroteinon spp. have a bifurcate frontal projection, Leona has a blunt frontal projection, Chondrolepis has a short down-turned frontal projection, Semalea and to a lesser degree Xanthodisca have a small frontal protuberance and the other genera have none; this does not seem to be a helpful character. We have noted above that the three species of Gretna which are doubtfully congeneric have quite similar caterpillars, but the pupae are very different, implying they belong in three different genera, none of which show much affinity to any other genera. The frontal projection of the pupa of G. balenge is unique, and the tufted pupa of G. cylinda is also very distinctive, but this may reflect that it is the only Afrotropical species that we have documented to form its pupa on the food plant leaf without the construction of any shelter. As previously noted the pupae and pupal shelters of Parosmodes and Melphinyet show affinities ( Cock & Congdon 2013), and the pupae of at least some Platylesches spp. are generally similar. The pupae of Acleros and Andronymus show affinities in shape and colouring as well as pupal shelter construction. Above, we have noted that the pupae of Caenides (both the palm-feeding C. dacela and the Costaceae-feeding C. dacena ), Semalea , Xanthodiscus , and Hypoleucis are united by the broad C-shaped rim to the T1 spiracle. The pupae of Gorgyra and Osmodes do not show any obvious affinities to any other genera.

The pupae of Zophopetes , Moltena , Monza and Perrotia are similar in shape and colour, while those of Gamia , Artitropa , and Erionota are similar but paler. Chondrolepis , Pteroteinon and Leona are also quite similar to this group, but have a frontal projection and the cremaster of Chondrolepis is less developed and may not be functional, although this may be a reflection of a tightly rolled pupal chamber obviating the need for pupal attachment. The dicotyledon-feeding Meza spp. may also relate to this group, but their exposed pupation site is perhaps linked to the unusual dorsal markings. The simple pupa of Acada , which could easily be mistaken for that of a member of the Noctuoidea, might also belong with this group. The pupae of Pardaleodes and Ankola are of similar shape and both are very flimsy (Cock & Congdon 2014), but their shape suggests they may have affinities with this group. There may also be affinities with Kedestes .

In summary, based on the pupae alone, we have nine possible groups:

1 Parosmodes, Melphinyet and perhaps Platylesches

2 Acleros and Andronymus

3 Caenides dacela , Semalea , Xanthodiscus , and Hypoleucis 4 Zophopetes , Moltena , Monza , Perrotia , Gamia , Artitropa , and Erionota with possible subgroups or related groups (a) Pteroteinon and Leona , (b) Chondrolepis , (c) dicotyledon-feeding Meza , (d) Acada , (d) Pardaleodes and Ankola and (e) Kedestes

5 Gorgyra

6 Osmodes

7 Gretna carmen 8 Gretna balenge 9 Gretna cylinda

These are not incompatible with the tentative groupings suggested for the limited observations of ova. The weak groupings suggested for caterpillars are also mostly compatible except that the similarities of the caterpillars of the three species of Gretna suggest that they are more closely related than do the very dissimilar pupae.

We look forward to seeing if and how these groups may align with future development of the classification of Hesperiinae incertae sedis based on the T.B. Larsen’s work on male genitalia and adult characters (mostly unpublished at present) and future expansion of the recent molecular and adult character analysis ( Warren et al. 2009). In doing so, we recognise that any classification based on the early stages (and hitherto we are only considering a morphological classification rather than a phylogeny) will not necessarily align with classifications or phylogenies based on the adults, but analyses based on both combined are likely to yield a more robust phylogeny ( Meier & Lim 2009). For example, Penz et al. (2013) showed that a phylogeny based on early stages of Brassolini (Nymphalidae) was less well resolved than one based on adult characters, but that the phylogeny based on the combined dataset was similar, but not identical to that based on adult characters only. On the other hand, caterpillars and pupae living in an exposed way (as Brassolini do) are likely to be subjected to stronger and perhaps more convergent natural selection, compared to those of Hesperiidae , which mostly live in shelters. We have already commented on the similarity of the early stages of diverse Hesperiinae that feed partially exposed, particularly on grasses and other monocotyledons. The caterpillars mostly have green bodies with longitudinal lines, and the pupae have similar colouring and a pointed frontal spike, which we have referred to as Baorini type pupae (Cock & Congdon 2014). At the time we hesitated to suggest whether this was a conservative character or the result of convergent evolution to address a common problem; today we are more inclined to the later explanation. We have noted another example of convergent evolution that will obscure relationships between genera in this series of papers: the development of ‘counterfeit predator eyes’ on the heads of caterpillars and pupae which live in shelters and should gain some advantage by frightening small vertebrate predators ( Janzen et al. 2010). Thus natural selection will obscure phylogeny where selection acts to converge on particular models for life style reasons.

Figures

FIGURE 41. Representative examples of the ova of the genera of Afrotropical Hesperiinae incertae sedis. The species are Parosmodes morantii, Platylesches moritili, Semalea pulvina, Acleros mackenii, Erionota torus, Zophopetes dysmephila, Pteroteinon concaenira, Leona maracanda, Artitropa sp. SCC 01, Andronymus caesar, Gretna balenge, Astictopterus stellata, Ceratrichia semlikensis, Gorgyra bibulus. See the relevant part of this series for details, except the image of E. torus was cropped from a photograph by P. Manoj (©, see acknowledgements).

FIGURE 42. Representative examples of the caterpillars of the genera of Afrotropical Hesperiinae incertae sedis. The species are Gorgyra mocquerysii, Acada biseriata, Parosmodes morantii, Acleros mackenii, Platylesches moritili, Andronymus caesar, Melphinyet tarace, Platylesches galesa, Platylesches robustus, Ploetzia amygdalis, Zophopetes dysmephila, Gretna carmen, Gretna cylinda, Gretna balenge (© Dirk Motshagen), Pteroteinon concaenira, Leona maracanda, Caenides dacela, Astictopterus stellata, Prosopalpus styla, Kedestes callicles, Ceratrichia semlikensis, Pardaleodes bule, Ankola fan, Perrotia albiplaga, Chondrolepis leggei, Monza alberti, Gamia shelleyi, Artitropa usambarae, Semalea arela, Hypoleucis ophiusa, Caenides dacena, Moltena fiara. See the relevant part of this series for details.

FIGURE 43. Representative examples of the pupae of the genera of Afrotropical Hesperiinae incertae sedis. The species are Gorgyra mocquerysii, Acada biseriata, Parosmodes morantii, Melphinyet flavina, Meza larea (specimen in The Natural History Museum, London), Acleros mackenii, Andronymus caesar, Platylesches moritili, Zophopetes quaternata, Gretna carmen, Gretna balenge, Pteroteinon caenira, Leona maracanda, Caenides dacela, Kedestes wallengrenii, Pardaleodes tibullus, Ankola fan, Perrotia albiplaga, Chondrolepis leggei, Monza cretacea, Gamia buchholzi, Artitropa usambarae, Semalea arela, Hypoleucis ophiusa, Xanthodisca vibius, Caenides dacena, Osmodes adon, Gretna cylinda, Moltena fiara. See the relevant part of this series for details.

Tables

TABLE 2. Summary of the genera of Hesperiinae incertae sedis and their food plant families.

Genus (species)	Evans’ Genera Group	Food plant families1	Stages documented2 Ovum Caterpillar Pupa
Acleros	Acleros	Fabaceae , Anacardiaceae , Malpighiaceae	+++ +++ +++
Andronymus	Acleros	Fabaceae , Sapindaceae	+++ +++ +++
Hypoleucis	Acleros	Costaceae	+++ +++
Meza (sensu lato)	Acleros	Fabaceae , Linaceae , Dichapetalaceae	+ +++
Meza (sensu stricto)	Acleros	Poaceae
Paronymus	Acleros	Dicotyledon	++
Gorgyra	Ampittia	Connaraceae Dichapetalaceae	+++ +++ +++
Kedestes	Ampittia	Poaceae	+++ +++
Prosopalpus	Ampittia	Poaceae	+++
Astictopterus	Astictopterus	Poaceae	+++ +++
Lepella	Astictopterus	Poaceae
Tsitana	Astictopterus	Poaceae	+ + +