Figure eight moth Ophiusa parcemacula, copyright Heath Hunter.

Belongs within: Erebinae.

Piercing fruit and piercing souls
Published 2 February 2021

The moths of the superfamily Noctuoidea are one of the most diverse subsections of the Lepidoptera, with probably somewhere between fifty and seventy thousand species known to date (Zahiri et al. 2012; as with other massively diverse clades, the lack of proper checklists and revisions makes the question of species number surprisingly difficult to answer). For many people, the classic image of a ‘moth’ will evoke a noctuoid: broad-winged, often nocturnal, often predominantly brown or grey in colour. Obviously, a group this size is going to have a complex taxonomy, and one of the significant subgroups of the noctuoids is the tribe Ophiusini.

Variable drab moth Ophiusa mejanesi, copyright Bernard Dupont.

Historically, the classification of noctuoids has been something of a mess. One researcher commented in 1975 that “It is exceptional to find any two authors who use the same combination of subfamily names within the Noctuidae” and Zahiri et al. admitted in 2012 that the validity of this statement still stood. Until recently, the majority of noctuoids were dumped in a broad family Noctuidae but recent studies (particularly influenced by molecular data) have lead to a significant rearrangement. As a result, the Ophiusini went from being usually placed in the family Noctuidae, subfamily Catocalinae, to the family Erebidae, subfamily Erebinae. A number of genera previously included in the Ophiusini were also transferred elsewhere; most notably, these included all New World representatives so the Ophiusini are now regarded as an exclusively Old World group.

Thyas juno, copyright Alexey Yakovlev.

The Ophiusini are mostly robust-bodied moths with wings of a fairly uniform background colour marked with simple, linear lines on the forewings. The males lack well-developed coremata (eversible structures used for dispersing pheromones) on the genital valves. The caterpillars are elongate semi-loopers with the front two pairs of abdominal prolegs much reduced compared to the rear two pairs. Larvae have been recorded from a wide range of host plant families but the most commonly exploited hosts are members of the Combretaceae and Myrtaceae (Holloway 2005). The pupa lacks the waxy bloom found in many other erebines.

Caterpillar of guava moth Ophiusa disjungens, copyright Robert Whyte.

Many members of the Ophiusini also have a modified apex to the adult proboscis bearing strong, enlarged spines and reversed, erectile hooks (Zahiri et al. 2012). This formidable apparatus is used to pierce the skins of fruits, allowing the moth to feed on their juice. As well as damage caused by browsing caterpillars, ophiusins may therefore also be of concern to horticulture due to damage from this fruit-piercing behaviour. As well as the damage caused by the moth itself, the resulting holes may allow the fruit to be attacked by disease or other insects not capable of breaching the rind themselves. The modified proboscis may also function in what is somewhat daintily referred to as lachrymal feeding: the process of applying the proboscis to the eyes of mammals (more rarely birds) and feeding on secreted fluids. Yes, these are moths that can potentially destroy an orchardist’s crop… and then proceed to drink his tears.

Ophiusini corrections
Published 8 December 2021

Earlier this year, I presented the above section on the noctuoid moth tribe Ophiusini. As it turns out, that post includes some notable errors. One of the main sources I used, Zahiri et al. (2012), stated that Ophiusini “have a strongly modified apex to the proboscis, with strong and enlarged spines and erectile, reversed hooks that are used in fruit-piercing or lachrymal-feeding behaviour“. As reviewed by Zilli (2021), such hooks on the proboscis are unique to a separate subgroup of the family Erebidae, the Calpinae. Ophiusini have thin, nail-like spines on the proboscis but no erectile hooks. They are still fruit-piercers but no ophiusins have been observed to date engaging in lachrymal feeding.

Artena dotata, copyright Shipher Wu.

Zilli (2021) had further comments on the historically fraught concept of Ophiusini. As noted in my earlier post, ‘Ophiusini’ has historically been recognised as a cosmopolitan group of moths but molecular studies have lead to its restriction to the Old World, North American exemplars being transferred to the related tribe Poaphilini. However, though the two groups are each supported as monophyletic by molecular data, they are not well defined morphologically. Characters previously thought distinct to one or the other do not always hold true. Ophiusini have been described as having reduced coremata but some ophiusins have coremata larger than those of some poaphilins. Ophiusins have been supposed to lack the waxy bloom on the pupa found in other noctuoids but some species do indeed have such a bloom. Some have pointed to the use of Euphorbiaceae as host plants by Poaphilini but not Ophiusini, but not all poaphilins feed on Euphorbiaceae and their use of this plant family is generally correlated with species being more generalist feeders overall.

One character that may yet distinguish the two tribes is the location of the androteca, a groove along the top of one of the leg segments in the male that contains a long brush of dense hairs (I’m not sure just what the function of this structure is meant to be but I would suspect something to do with dispersing pheromones). In Ophiusini, this structure is found on the femur of the fore leg. In Poaphilini, it is on the tibia of the mid leg. Nevertheless, Zilli (2021) questions the reliability of this feature: both arrangments are found in other tribes and neither alone is diagnostic.

Conversely, molecular phylogenies support the two tribes as sister taxa, and they share a number of distinctive features of the terminalia. While he does not formalise the suggestion, Zilli (2021) seems to feel that we might be better served by a return to a broader Ophiusini uniting the two tribes as one. I commented in my previous post that noctuoid classification has been in a continuous flux for as long as it has been a thing. It would be presumptuous to believe that it has finally been settled.

Systematics of Ophiusini
<==Ophiusini [Focillidae, Ophiusidae, Ophiusidi, Ophiusina, Ophiusinae]HM03
|--+--‘Ophiusa’ tirhaca (Cramer 1777)ZK11, Z21
| `--Clytie Hübner 1823ZK11, Z21
| |--C. deviaZK11
| `--C. illunarisSKR96
`--+--Artena Walker 1858ZK11, Z21
| |--A. dotata Fabricius 1794Z21
| |--A. reggiantii Zilli & Lourens 2018Z21
| |--A. rubida (Walker 1863)Z21
| `--A. velutina (Prout 1919)Z21
`--Ophiusa Ochsenheimer 1816ZK11, Z21
|--O. algira [incl. Noctua achatina, N. triangularis]L49
|--O. arctotaeniaWM66
|--O. coronataNC91
|--O. dilecta Walker 1865Z21
|--O. disjungens (Walker 1858)Z21
|--O. geometricaL49
|--O. lunarisL49
|--O. parcemaculaMC13
|--O. pastinumF92
|--O. stolida [incl. O. singularis]L49
|--O. stuposaWM66
`--O. trapezium (Guenée 1852)Z21

Ophiusini incertae sedis:
Ophiodes triphaenoidesWM66
Trigonodes hyppasiaWM66
Bocula S01 [incl. EudraganaJG19]
|--B. erota Swinhoe 1901S01
|--B. limbata (Butler 1889) [=Eudragana limbata]JG19
`--B. punctilineataS01
Motina shumara Swinhoe 1901S01
Tiruvaca Swinhoe 1901S01
`--*T. subcostalis [=Thermesia subcostalis]S01
Platyja minutipuncta Swinhoe 1901S01
Thyas Hübner 1824Z21, WL09
`--T. coronata (Fabricius 1775)Z21
|--S. reducta Mabille 1880Z21
`--S. tumiditermina (Hampson 1910)Z21
Anua Walker 1858 [Anuini]FL05
`--A. coronataC70
Zethes multiplaga Swinhoe 1901FL05, S01
Eubolina diffundensFL05, H01
Bendis Hübner 1823 [Bendidae]FL05
Egnasia costalis (Moore 1883) [=Acharya costalis]S01
|--D. almana Swinhoe 1901S01
|--D. diffusifascia Swinhoe 1901S01
`--D. lichenosaS01
Rhesala Walker 1858 (see below for synonymy)S01
|--R. asphalta Swinhoe 1901S01
|--R. digramma [=Raparna digramma]S01
|--R. iada Swinhoe 1901S01
|--R. imparataS01
|--R. laevis Swinhoe 1901S01
|--R. mansueta [=Daona mansueta, Xanthoptera mansueta; incl. Byturna rufifascia Hampson 1892]S01
|--R. ochreipennisS01
`--R. transversaS01
Focilla Guenée 1852FL05
Lagoptera Guenée 1852 [Lagopteridae]FL05
Phaeocyma Hübner 1818 [Phaeocymini]FL05

Rhesala Walker 1858 [incl. Byturna Moore 1883, Daona Walker 1864, Enea Walker 1864, Hingula Moore 1882, Raparna Moore 1882, Vescisa Walker 1864]S01

*Type species of generic name indicated


[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[F92] Fan Z. 1992. Key to the Common Flies of China 2nd ed. Science Press: Beijing.

[FL05] Fibiger, M., & J. D. Lafontaine. 2005. A review of the higher classification of the Noctuoidea (Lepidoptera) with special reference to the Holarctic fauna. Esperiana 11: 7–92.

[H01] Hampson, G. F. 1901. The Lepidoptera-Phalaenae of the Bahamas. Annals and Magazine of Natural History, series 7, 7: 246–261.

Holloway, J. D. 2005. The moths of Borneo (part 15 & 16): family Noctuidae, subfamily Catocalinae. Malayan Nature Journal 58: 1–529.

[HM03] Holloway, J. D., & S. E. Miller. 2003. The composition, generic placement and host-plant relationships of the joviana-group in the Parallelia generic complex (Lepidoptera: Noctuidae, Catocalinae). Invertebrate Systematics 17: 111–128.

[JG19] James, D. J., P. T. Green, W. F. Humphreys & J. C. Z. Woinarski. 2019. Endemic species of Christmas Island, Indian Ocean. Records of the Western Australian Museum 34 (2): 55–114.

[L49] Lucas, H. 1849. Exploration Scientifique de l’Algérie pendant les années 1840, 1841, 1842 publiée par ordre du gouvernement et avec le concours d’une commission académique. Sciences physiques. Zoologie. II. Histoire naturelle des animaux articulés. Troisième partie. Insectes (suite). Imprimerie Nationale: Paris.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[SKR96] Sobhian, R., L. Knutson & J. S. Rodier. 1996. Biology and host specificity notes on Styphlus penicillus Gyllenhal (Coleoptera: Curculionidae), examined as a biological control agent for Crupina vulgaris in the United States with remarks on its host plant. Proceedings of the Entomological Society of Washington 98 (2): 317–323.

[S01] Swinhoe, C. 1901. New genera and species of eastern and Australian moths (continued). Annals and Magazine of Natural History, series 7, 7: 489–501.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

[WL09] Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook & G. W. Krantz. 2009. Order Trombidiformes. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 233–420. Texas Tech University Press.

Zahiri, R., J. D. Holloway, I. J. Kitching, J. D. Lafontaine, M. Mutanen & N. Wahlberg. 2012. Molecular phylogenetics of Erebidae (Lepidoptera, Noctuoidea). Systematic Entomology 37: 102–124.

[ZK11] Zahiri, R., I. J. Kitching, J. D. Lafontaine, M. Mutanen, L. Kaila, J. D. Holloway & N. Wahlberg. 2011. A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera). Zoologica Scripta 40 (2): 158–173.

[Z21] Zilli, A. 2021. Tabwecala robinsoni gen. nov., sp. nov., from Vanuatu and its systematic postion in the ‘Ophiusini-Poaphilini’ clade (Lepidoptera, Erebidae). Nota Lepidopterologica 44: 193–211.

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