Lyssa+zampa

=Tropical Swallowtail Moth= Every few years in Singapore, swarms of huge moths appear downtown in places like the National Library; the species responsible is //Lyssa zampa//, known as the tropical swallowtail moth or Laos brown butterfly. Because of its flashy size and behaviour, it often makes local news headlines across Southeast Asia. Despite its prevalence in urban areas, its biology is not studied in great detail.

**//Behaviour//**
//Lyssa zampa// is attracted to the bright lights of urban areas, including Singapore. It is most famous for its "swarming" behaviour; this may not be true swarming but merely mass emergence. //Lyssa zampa// is present during other times of the year, between April and August, but only occasionally congregate in large numbers. Various outlets (BBC, Straits Times, Stomp, etc.) have reported on swarm sightings in the city. The reason for these sudden large numbers is not well understood. It is thought that high population numbers may occur in years with favourable weather conditions or low predation levels. Mass emergence has occurred in the past following dry spells, supporting the weather based hypothesis. In this model, a dry spell is followed by a mass flowering and subsequent foliage growth. The additional foliage is able to support more larvae, causing a mass emergence and swarming. In Singapore, mass recent mass swarms have occurred in 2005, 2010, and 2014.

media type="youtube" key="9i3OQdbo_eE" width="560" height="315" The Star news report of //L. zampa// in Malaysia

The behaviour within Uraniidae, the family containing //L. zampa//, varies alongside morphology. When alighting in the wild, the more conspicuous species generally alight on top of a leaf, then reverse their position so that the head points downward. This tactic confuses predators by drawing attention to the hind wing tails, a non-vital part of the animal. Other species of //Lyssa//, which often have cryptic brown coloration with disruptive white bands, rest in dark places or under leaves where they are completely out of sight. //Lyssa zampa// itself exhibits the former habit of turning upside down after alighting in conspicuous locations. The caterpillars leave a trail of silk behind them as they forage. When threatened, they use this thread to drop from the host plant and avoid predation.



**//Morphology//**
//Lyssa zampa// is a large moth commonly known as the tropical swallowtail moth or the Laos brown butterfly. The wingspan of //L. zampa// is 10-16 cm ; //Lyssa zampa// has a light stripe in the middle of both dorsal wings; there is a lighter region immediately distal to this stripe. On the underside there is a more or less even dispersal of dark striation distal to the white band. The hind wings of //L. zampa// have two tails, the longer one being more central. The ventral side has a smooth brown region distal to the light stripe, and rippled striation proximal to the stripe. Females have a slightly larger wingspan and body, typical in many moths and butterflies. Females also have a slightly lighter colour.



Like all moths and butterflies, //L. zampa// goes through a complete metamorphic life cycle, changing through egg, larva, and pupa to adult. The majority of a moth's life span is in the larval stage, where it does its eating and growing. The adult moth phase typically lasts a few days and is largely for reproduction and dispersal. The larvae are pale yellow-green with red-brown head and legs. Most segments have pale brown markings that are emphasised by dramatic skin wrinkling. The spiracles are white and ringed with black, so they stand out clearly. Young larvae have more extensive black markings and darker green coloration. Pupation occurs in leaf litter; the pupa is dark brown in colour. This is a widespread method of pupation that provides protection and camouflage.

Lyssa zampa is in the subfamily Uraniinae, which contains about 50 described species in seven genera and contains two common trends in morphology. Three of these genera are largely diurnal and brightly coloured. Their bright bands of color (typically iridescent green or blue) is warning coloration communicating their toxicity. Of these, the Madagascan sunset moth is famous for its bright colours. The remaining four, including Lyssa, have duller coloration and are largely nocturnal. The hindwing tails and general shape of Uraniidae bears strong resemblance to papilionid butterflies. For this reason, Linnaeus at one point described L. zampa in the family Papilionidae. Among all the genera, Lyssa is notable in containing the largest species on average.



**//Range//**
//Lyssa zampa// can be found from North East Himalaya to Southern China and the Malay Peninsula. The majority of sightings occur in cities like Hong Kong and Singapore, where //L. zampa// is attracted to light sources. The colour of //L. zampa// varies slightly with geography and can be used as a locality indicator.



Within Singapore, there are also a great deal of sightings. Sightings are concentrated in the downtown core and other more densely populated areas, but occur across the country. media type="custom" key="28849584"

Local editor of Habitatnews, N. Sivasothi, curates records of //L. zampa// sightings using an [|online form]. This form has been collecting submissions since 2010 after a mass emergence.

**//Host Plant Interactions//**
Most larval feeding in Uraniidae occurs on plants in the family Euphorbiaceae. The tree //Endospermum diadenum// is recorded as a food plant for three of the primarily nocturnal genera in Uraniidae (//Lyssa, Urapteroides//, and //Cyphura//); this tree is native to Singapore but vulnerable. The feeding habits of //L. zampa// have not been explored in-depth, but the feeding habits of other uraniids have. In //Chrysiridia rhipheus,// caterpillars eat exclusively leaf tissue for the first 3-4 days of life then move on to flowers, fruits, young stems, and other portions of the plant. In some adults of Uraniidae, nectar feeding occurs largely in white flowers; this indicates a visual aspect to food selection.



The tree is used for low grade timber, as a shade plant, and in alley cropping with rubber trees. The seeds of //E. diadenum// are difficult to obtain because of heavy damage by caterpillars; trees are often grown from wild saplings because of this.

The following is a table of hostplants for //L. zampa// in various areas. //L. zampa// has been captivity reared in Malaysia on //Eudospermum malaccense.

// Lyssa macleayi //sequesters alkaloidal glycosidase inhibitors (AGIs) from the hostplant //Endospermum medullosum;// it’s very likely that //L. zampa// does so as well. These compounds block digestive glycosidase enzymes, but uraniid species are resistant to this effect. Interestingly, these same compounds are being investigated in their ability to prevent the progression of an HIV infection by halting antigenic diversity.

**//Other Species Interactions//**
The predators of //L.// // zampa // in Singapore include various birds, which consume the larvae. The moths are also host to various parasites; in Vietnam, //Forcipomyia// biting midges have been observed parasitising //L. zampa//. The midges have a special spine and a comb on each of their tarsal claws, which allow them to cling to the moth scales in a uniform fashion with heads pointed to wing base.



//Taxonomy//
The genus //Lyssa// was described in 1823 by Hübner. //Lyssa// contains 28 species, which live in Southeast Asia and the Pacific. The name was introduced as a general name for two species, one of which was described by Linnaeus. The //Nyctalemon// genus was described by Dalman in 1825 and approached in 1953 by C. O. van Regteren Altena. This paper demonstrated that several genus names, including //Papilio// and //Urania// had been previously used to describe a cluster of species including //L. zampa//; several of these names were already in use for species from different genera and could not be considered. At the time, //Nyctalemon// was the synonym most predominantly used that was not already in use for other genera. This paper recognised that the original //Lyssa// should be used based on the rule of priority, but wanted to discontinue its use in favour of //Nyctalemon.// Ultimately, //Nyctalemon// was suppressed as it is a junior synonym.
 * Kingdom || Aimalia ||
 * Phylum || Arthropoda ||
 * Class || Insecta ||
 * Order || Lepidoptera ||
 * Superfamily || Geometroidea ||
 * Family || Uraniidae ||
 * Genus || //Lyssa// ||
 * Species || //zampa// ||

//Lyssa zampa// was originally described by Arthur G. Butler in 1869 under the synonym //Nyctalemon zampa//; he very briefly described it as “a local representative of //N. hector//”(20). Its type specimen is kept in the British Museum. The name was later changed to //Lyssa zampa// with the revision of the //Nyctalemon// genus. The description was published along with descriptions of several other species in “Descriptions of species of Lepidoptera, confounded with others described by Linnaeus and Fabricius” in Entomology Monthly Issue 5, page 273. //Lyssa zampa// has several synonyms, all in the //Nyctalemon// genus.

The family Uranidae is placed in the superfamile Geometriodea. The following are two phylogenetic trees. The first is of the families and subfamilies of geometroid moths based on morphological analysis. The second is a more detailed tree of Uraniinae based on DNA bar codes ; the tree focusses on //Urania// but includes //L. zampa.//

Phylogenetic Trees Placing Uraniinae and //L. zampa//