Blue Moon

Male blue moon butterfly Hypolimnas bolina, photographed by Comacontrol.

My native country of New Zealand is not home to a large diversity of butterflies. Only a couple of dozen or so species are known from the entire country. It would not be unreasonable for a keen butterfly spotter to attempt to track down them all. But one particular species of butterfly generally included in New Zealand lists would a touch of luck: the aptly named blue moon Hypolimnas bolina.

This is because the blue moon is not a regular resident of New Zealand (I've personally never spotted one). It is native to a wide region stretching from Madagascar and India to Japan and northern Australasia where it is usually referred to by the more prosaic name of common or greater eggfly. The only examples found in New Zealand are vagrants who lost their way on southwards migrations. Nevertheless, such vagrants are regular enough for its local appellation to be thought worth coining. Not only does it reflect their rarity, it also describes the appearance of the male, with the wings bearing blue-ringed white spots on a black background.

Two females of Hypolimnas bolina. On the left, a mimetic individual, copyright Greg Hume; on the right, a non-mimetic individual, copyright W. A. Djatmiko.

The appearance of the female is a bit harder to explain because it can vary between individuals. Females of the eggfly genus Hypolimnas are commonly mimics of other, poisonous butterflies of the subfamily Danainae, to which eggflies are only distantly related (both groups belong to the family Nymphalidae but eggflies are placed in the subfamily Nymphalinae). For instance, the diadem or danaid eggfly H. misippus of Africa and Asia (and also introduced into parts of the Americas adjoining the Caribbean) is a mimic of the plain tiger Danaus chrysippus. The chosen model of H. bolina in the western part of its range is the common crow Euploea core and in the region of India almost all females are a remarkably good copy of that species (above left). But as one moves east, one starts seeing females of H. bolina that are not mimics like the individual shown above right; by the time one reaches Australia these make up the greater part of the population. Mimetic females may also vary to resemble different Euploea species, depending on which model is locally present.

Female danaid eggfly Hypolimnas misippus, copyright Raju Kasambe. Males of this species are similar to those of H. bolina but lack the blue rings around the white spots on the wings.
There are about two dozen species of Hypolimnas eggflies found in various parts of the Old World tropics. Hypolimnas misippus is also found in parts of the Americas around the Caribbean where its presence is usually explained as the result of an early introduction (possibly, and somewhat poignantly, in connection to the slave trade). Their vernacular name is probably derived from the unique behaviour (for butterflies) of a number of species whose females stand guard over their eggs, beating their wings over them to protect them from predators until hatching. About two-thirds of Hypolimnas species are mimics. In some of these species, both sexes are mimetic; others resemble H. bolina and H. misippus in that only the females are mimics (Vane-Wright et al. 1977). One might be tempted to ask why this variation exists. One point to be considered is that there are limits on when mimicry is likely to be effective. The mimic needs to be much less abundant than its model, otherwise potential predators may not learn to associate the distinctive coloration with the toxic original. Swinhoe (1896) noted that males of Hypolimnas misippus were very active, aggressively defending their territories from other butterflies, and suggested that this agility might provide males with alternative defences to mimicry. The more sedentary females (especially when egg-guarding) might be expected to benefit more from the passive protection mimicry provides, but mimesis might be expected to disappear in areas where their model is less abundant.

REFERENCES

Swinhoe, C. 1896. On mimicry in butterflies of the genus Hypolimnas. Journal of the Linnean Society, Zoology 25: 339–348.

Vane-Wright, R. I., P. R. Ackery & R. L. Smiles. 1977. The polymorphism, mimicry, and host plant relationships of Hypolimnas butterflies. Zoological Journal of the Linnean Society 9: 285–297.

Large Yellow Underwings

Above is an example (copyright Richard) of the large yellow underwing Noctua pronuba, the most widespread species of its genus. Noctua pronuba is a relatively large moth, with a wingspan of up to 60 mm. The forewings are fairly dull and dark in colour, but the hindwings are a bright yellow-orange (hence the vernacular name) with a black border. It can usually be found out and about in mid summer to early autumn. Its larvae are one of the types of caterpillar known as 'cutworms', which live buried in the soil during the day and emerge at night to feed. They get their vernacular name because their soil-dwelling habits mean that they tend to feed on plants from the base, often toppling small plants and seedlings like a lumberjack taking down a tree. The larvae of the large yellow underwing are not overly discerning in their food preferences; though they most often feed on grasses, they will quite happily dine on other herbaceous flowering plants such as legumes.

Larva of Noctua pronuba, copyright Nigel Richards.

The large yellow underwing is native to a wide part of the Palaearctic region, i.e. Europe and northern Asia. In 1979, it was also found introudced to Nova Scotia in North America. Since then, it has spread rapidly and can now be found over much of temperate North America, reaching British Columbia in the west and Louisiana in the south (Copley & Cannings 2005). The large yellow underwing is a strong flier and is known to undertake significant migrations in its native range. Females may also lay large numbers of eggs at a single time on the underside of leaves or on non-host plant substrates, where they can easily be carried between locales by human transportation. For the most part, a significant impact of the large yellow underwing on native or horticultural production in North America has not been recognised, though a number of authors have suggested that this species' generalist diet may lead to any such impact going unnoticed.

The Azorean Noctua atlantica, copyright Jens Jacobasch.

Historically, a large number of species have been included at one time or another in the genus Noctua, but a revision of this and closely related genera by Beck et al. (1993) cut it down on the basis of larval and male genital morphology to just two species. The only other species retained in Noctua sensu stricto by Beck et al. was N. atlantica, a species endemic to the Azores islands west of Portugal. Noctua atlantica is somewhat smaller than N. pronuba with duller hindwing coloration. It is restricted in its home range to the laurisilva, a particular subtropical forest type dominated by laurels and other evergreen, broad-leaved trees. Noctua pronuba is also found on the archipelago but inhabits are wider range of habitats. Genetic comparisons between the two species suggest that their populations diverged about five million years ago, a time frame that is not inconsistent with the origin of the Azores archipelago about four million years ago (Montiel et al. 2008). Perhaps an early population of N. pronuba became isolated on the Azores long enough to evolve into a distinct species, followed by a later re-colonisation of N. pronuba from the mainland. This would be similar to patterns seen in my home country of New Zealand, where repeated immigrations from Australia have lead to species pairs such as the endemic takahe Porphyrio hochstetteri and the more widespread pukeko P. melanotus, or the endemic (now extinct) Eyles' harrier Circus eylesi and the modern swamp harrier C. approximans.

REFERENCES

Beck, H., L. Kobes & M. Aloha. 1993. Die generische Aufgliederung von Noctua Linnaeus, 1758 (Lepldoptera, Noctuidae, Noctuinae). Atalanta 24 (1–2): 207–264.

Copley, C. R., & R. A. Cannings. 2005. Notes on the status of the Eurasian moths Noctua pronuba and Noctua comes (Lepidoptera: Noctuidae) on Vancouver Island, British Columbia. J. Entomol. Soc. Brit. Columbia 102: 83–84.

Montiel, R., V. Vieira, T. Martins, N. Simões & M. L. Oliveira 2008. The speciation of Noctua atlantica (Lepidoptera, Noctuidae) occurred in the Azores as supported by a molecular clock based on mitochondrial COI sequences. Arquipélago 25: 43–48.