Tropical Swallowtail Moth

Lyssa zampa dorsal resting.jpg
Image by Bernard Dupont, under the CC BY-SA 2.0 License.
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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[2] . 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[3] . 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[4] . In Singapore, mass recent mass swarms have occurred in 2005, 2010, and 2014[5] .

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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[7] . 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[8] .

Lyssa other moth.jpg
Urania leilus. Image by D. Gordon and E. Robertson under CC BY-SA 3.0 License.
[9]

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
[10] ; 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[11] . Females also have a slightly lighter colour.

Lyssa zampa life stages.png
L. zampa larvae, pupa, adult, and copulation. Images by Kelvin Lim, permission pending.
[12]

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[13] . Pupation occurs in leaf litter; the pupa is dark brown in colour. This is a widespread method of pupation that provides protection and camouflage[14] .


Lyssa Uraniidae Scientific Illustration.jpg
Urania sloanus by William Swainson, under public domain.
[15] Lyssa zampa is in the subfamily Uraniinae, which contains about 50 described species in seven genera and contains two common trends in morphology[16] . 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[17] . 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[18] . Among all the genera, Lyssa is notable in containing the largest species on average.



Lyssa sunset moth resting.jpg
Chrysiridia rhipheus, the Madagascan sunset moth. Image by Bernard Dupont under CC BY-SA 2.0 License.
[19]

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Range

Lyssa zampa can be found from North East Himalaya to Southern China and the Malay Peninsula[21] . 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[22] .

Lyssa zampa map.png
Map of 111 sightings of L. zampa from 2000 to 2010. Image under CC-BY-SA 3.0 License.
[23]

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.
[24]

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[25] .


Host Plant Interactions

Most larval feeding in Uraniidae occurs on plants in the family Euphorbiaceae[26] . 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[27] . 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[28] . In some adults of Uraniidae, nectar feeding occurs largely in white flowers; this indicates a visual aspect to food selection.

Lyssa zampa host plant.jpg
Endopermum diadenum, image by Cerlin Ng under CC BY-NC-SA 2.0 License

















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The tree is used for low grade timber, as a shade plant, and in alley cropping with rubber trees[30] . 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[31] .

The following is a table of hostplants for L. zampa in various areas. L. zampa has been captivity reared in Malaysia on Eudospermum malaccense[32] .
Lyssa zampa Host plants.png


Lyssa macleayi sequesters alkaloidal glycosidase inhibitors (AGIs) from the hostplant Endospermum medullosum; it’s very likely that L. zampa// does so as well[33] . These compounds block digestive glycosidase enzymes, but uraniid species are resistant to this effect[34] . Interestingly, these same compounds are being investigated in their ability to prevent the progression of an HIV infection by halting antigenic diversity[35] .


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[36] . 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.

Lyssa zampa with midge.png
L. zampa with midges.
Lyssa zampa midge closeup.png
Closeup of midge on wing surface. Images by Andrei Sourakov, permission to use granted.













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Taxonomy

Kingdom
Aimalia
Phylum
Arthropoda
Class
Insecta
Order
Lepidoptera
Superfamily
Geometroidea
Family
Uraniidae
Genus
Lyssa
Species
zampa
The genus Lyssa was described in 1823 by Hübner. Lyssa contains 28 species[38] , 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[39] . 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[40] . 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.

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[41] . 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[42] . Lyssa zampa has several synonyms, all in the Nyctalemon genus[43] .

The family Uranidae is placed in the superfamile Geometriodea [44] . The following are two phylogenetic trees. The first is of the families and subfamilies of geometroid moths based on morphological analysis[45] . The second is a more detailed tree of Uraniinae based on DNA bar codes[46] ; the tree focusses on Urania but includes L. zampa.

Phylogenetic Trees Placing Uraniinae and L. zampa[47] [48]
Lyssa zampa phylogeny.pngLyssa zampa specific phylogeny.png
  1. ^ “Tropical Swallowtail Moth (Lyssa zampa)” by Bernard Dupont. Flickr, 20 Feb 2007. URL: https://www.flickr.com/photos/berniedup/6654060775/ (accessed on 15 Nov, 2016).
  2. ^
    “Lyssa zampa”. Revolvy. URL: https://www.revolvy.com/main/index.php?s=Lyssa%20zampa&uid=1575 (accessed on 15 Nov, 2016).
  3. ^ “Giant Moths Have Descended on Malaysia, And No One Knows Why” by Mary Beth Griggs. Smartnews, 16 Jun 2014. URL: http://www.smithsonianmag.com/smart-news/giant-moths-are-flying-around-malaysia-180951746/?utm_source=facebook.com&no-ist (accessed on 15 Nov, 2016).
  4. ^ “Citizen science contributions to tracking the seasonal abundance of the tropical swallowtail moth (Lyssa zampa) (Butler, 1869)” by Lynn NG, Anuj Jain, and N. Sivasothi. Biodiversity of Singapore Symposium IV, 2015. URL: https://biodiversitysg4.files.wordpress.com/2015/07/iii-04-lynn-ng-lyssa-zampa.pdf (accessed on 15 Nov, 2016).
  5. ^ “Citizen science contributions to tracking the seasonal abundance of the tropical swallowtail moth (Lyssa zampa) (Butler, 1869)” by Lynn NG, Anuj Jain, and N. Sivasothi. Biodiversity of Singapore Symposium IV, 2015. URL: https://biodiversitysg4.files.wordpress.com/2015/07/iii-04-lynn-ng-lyssa-zampa.pdf (accessed on 15 Nov, 2016).
  6. ^
    “Massive Moths Invade Malaysia” by GeoBeats News YouTube Channel. YouTube, 12 Jun, 2014. URL: https://www.youtube.com/watch?v=9i3OQdbo_eE&feature=youtu.be (accessed on 17 Nov, 2016).
  7. ^
    Lees, D. C., 1991. Foodplants of the Uraniinae (Uraniinae) and their Systematic, Evolutionary and Ecological Significance. Journal of the Lepidopterists' Society, vol. 45: 296-347
  8. ^
    “Lyssa zampa Butler” by Jeremy Daniel Holloway. The Moths of Borneo, 1998. URL: http://www.mothsofborneo.com/part-8/uraniinae/uraniinae_1_1.php (accessed on 15 Nov, 2016).
  9. ^ “Green-banded Urania (Urania leilus)” by D. Gordon and E. Robertson. Wikimedia Commons, 15 Nov 2011. URL: https://en.wikipedia.org/wiki/Urania_leilus#/media/File:Green-banded_Urania,_Tambopata_Park,_Peru.jpg (accessed on 15 Nov, 2016).
  10. ^
    "Lyssa zampa”. Revolvy. URL: https://www.revolvy.com/main/index.php?s=Lyssa%20zampa&uid=1575 (accessed on 15 Nov, 2016).
  11. ^ “Citizen science contributions to tracking the seasonal abundance of the tropical swallowtail moth (Lyssa zampa) (Butler, 1869)” by Lynn NG, Anuj Jain, and N. Sivasothi. Biodiversity of Singapore Symposium IV, 2015. URL: https://biodiversitysg4.files.wordpress.com/2015/07/iii-04-lynn-ng-lyssa-zampa.pdf (accessed on 15 Nov, 2016).
  12. ^
    “Lyssa zampa Metamorphosis” by Kelvin Lim. News from Lee Kong Chian Natural History Museum, 6 Jun 2014. URL: https://lkcnhm.net/2014/06/06/swallowtail-moth-lyssa-zampa-in-singapore-biodivesity-records/ (accessed on 15 Nov, 2016).
  13. ^
    “Lyssa zampa Butler” by Jeremy Daniel Holloway. The Moths of Borneo, 1998. URL: http://www.mothsofborneo.com/part-8/uraniinae/uraniinae_1_1.php (accessed on 15 Nov, 2016).
  14. ^ “Lyssa zampa Butler” by Jeremy Daniel Holloway. The Moths of Borneo, 1998. URL: http://www.mothsofborneo.com/part-8/uraniinae/uraniinae_1_1.php (accessed on 15 Nov, 2016).
  15. ^
    “Plate from Zoological illustrations, Volume 3, 2nd series: Accepted as Urania sloanus (extinct)” by William Swainson. Wikimedia Commons, 1 Jan 1829. URL: https://en.wikipedia.org/wiki/Urania_sloanus#/media/File:Zoological_Illustrations_Volume_III_Series_2_129.jpg (accessed on 15 Nov, 2016).
  16. ^ Lees, D. C., 1991. Foodplants of the Uraniinae (Uraniinae) and their Systematic, Evolutionary and Ecological Significance. Journal of the Lepidopterists' Society, vol. 45: 296-347.
  17. ^ Lees, D. C., 1991. Foodplants of the Uraniinae (Uraniinae) and their Systematic, Evolutionary and Ecological Significance. Journal of the Lepidopterists' Society, vol. 45: 296-347.
  18. ^ Altena, C. O., 1953. “A Revision of the Genus Nyctalemon Dalman (Lepidoptera, Uraniidae) with Notes on the Biology, Distribution, and Evolution of its Species”. Zoologische Verhandelingen, 19: 1-58.
  19. ^ “Madagascan Sunset Moth (Chrysiridia rhipheus)” by Bernard Dupont. Wikimedia Commons, 20 Sept 2004. URL: https://commons.wikimedia.org/wiki/File:Madagascan_Sunset_Moth_(Chrysiridia_rhipheus)_(7621728054).jpg (accessed on 15 Nov, 2016).
  20. ^
    “Plate from Zoological illustrations, Volume 3, 2nd series: Accepted as Urania sloanus (extinct)” by William Swainson. Wikimedia Commons, 1 Jan 1829. URL: https://en.wikipedia.org/wiki/Urania_sloanus#/media/File:Zoological_Illustrations_Volume_III_Series_2_129.jpg (accessed on 15 Nov, 2016).
  21. ^
    “Lyssa zampa Butler オオツバメガ Cat.2940”. JPMoth. URL: http://www.jpmoth.org/~dmoth/Digital_Moths_of_Asia/85_GEOMETROIDEA/03_URANIIDAE/02_Uraninae/01_Lyssa/Lyssa_zampa/Lyssa_zampa.htm (accessed on 15 Nov, 2016).
  22. ^ Altena, C. O., 1953. “A Revision of the Genus Nyctalemon Dalman (Lepidoptera, Uraniidae) with Notes on the Biology, Distribution, and Evolution of its Species”. Zoologische Verhandelingen, 19: 1-58.
  23. ^
    “Lyssa zampa Butler, 1869”. GBIF Backbone Taxonomy. URL: http://www.gbif.org/species/1953196 (accessed on 15 Nov, 2016).
  24. ^
    “2014 Singapore Swallowtail moth (Lyssa zampa) sighting map” by Butterflysm Facebook group. GoogleMaps, 2014. URL: https://www.google.com/maps/d/u/0/viewer?mid=1otLh1L0zXb3sAVt1jxPaRN3iLmY&hl=en_US&ll=2.5197237603477096%2C105.91671645641327&z=6 (accessed on 17 Nov, 2016).
  25. ^
    “Submit your record of Lyssa zampa, the tropical swallowtail moth, in Singapore” by N. Sivasothi. Habitat News, 22 May 2014. URL: http://habitatnews.nus.edu.sg//index.php?phrase=lyssa+zampa&submit=Search+Habitatnews (accessed on 15 Nov, 2016).
  26. ^
    “Lyssa zampa Butler” by Jeremy Daniel Holloway. The Moths of Borneo, 1998. URL: http://www.mothsofborneo.com/part-8/uraniinae/uraniinae_1_1.php (accessed on 15 Nov, 2016).
  27. ^ “Endospermum diadenum” by Ken Fern. Useful Tropical Plants, 16 Aug 2016. URL: http://tropical.theferns.info/viewtropical.php?id=Endospermum+diadenum (accessed on 15 Nov, 2016).
  28. ^ Lees, D. C., 1991. Foodplants of the Uraniinae (Uraniinae) and their Systematic, Evolutionary and Ecological Significance. Journal of the Lepidopterists' Society, vol. 45: 296-347.
  29. ^
    “Endospermum diadenum” by Cerlin Ng. Flickr, 16 Mar 2014. URL: https://www.flickr.com/photos/89906643@N06/13288944785/ (accessed on 15 Nov, 2016).
  30. ^
    “Endospermum diadenum” by Ken Fern. Useful Tropical Plants, 16 Aug 2016. URL: http://tropical.theferns.info/viewtropical.php?id=Endospermum+diadenum (accessed on 15 Nov, 2016).
  31. ^ “Endospermum diadenum” by Ken Fern. Useful Tropical Plants, 16 Aug 2016. URL: http://tropical.theferns.info/viewtropical.php?id=Endospermum+diadenum (accessed on 15 Nov, 2016).
  32. ^ //
    “A Database of the World's Lepidopteran Hostplants”, G. S. Robinson. HOSTS, 2010. URL: http://www.nhm.ac.uk/our-science/data/hostplants/search/list.dsml?searchPageURL=browse.dsml&Family=Uraniidae&Genus=Lyssa&Species=zampa&Country=&sort=Family (accessed on 15 Nov, 2016).
  33. ^
    Sourakov, A., 2013. “The Tropical Swallowtail Moth, Lyssa zampa (Uraniidae): Another Victim of Lymph-Thirsty Parasites in Vietnam”. News of the Lepidopterists’ Society, 55 (3): 106-107.
  34. ^ Kite, G. C., 1997. “Alkaloidal Glycosidase Inhibitors and Digestive Glycosidase Inhibition in Specialist and Generalist Herbivores of Omphalea diandra”. Journal of Chemical Ecology, 23 (1): 199-135.
  35. ^ Dealler, S., 1994. “”Alkaloidal glycosidase inhibitors (AGIs) as the cause of sporadic scrapie, and the potential treatment of both transmissible spongiform encephalopathies (TSEs) and human immunodeficiency virus (HIV) infection”. Med Hypotheses, 42 (2): 69-75.
  36. ^
    Sourakov, A., 2013. “The Tropical Swallowtail Moth, Lyssa zampa (Uraniidae): Another Victim of Lymph-Thirsty Parasites in Vietnam”. News of the Lepidopterists’ Society, 55 (3): 106-107.
  37. ^
    Sourakov, A., 2013. “The Tropical Swallowtail Moth, Lyssa zampa (Uraniidae): Another Victim of Lymph-Thirsty Parasites in Vietnam”. News of the Lepidopterists’ Society, 55 (3): 106-107.
  38. ^
    Kite, G. C., 1997. “Alkaloidal Glycosidase Inhibitors and Digestive Glycosidase Inhibition in Specialist and Generalist Herbivores of Omphalea diandra”. Journal of Chemical Ecology, 23 (1): 199-135.
  39. ^ Altena, C. O., 1953. “A Revision of the Genus Nyctalemon Dalman (Lepidoptera, Uraniidae) with Notes on the Biology, Distribution, and Evolution of its Species”. Zoologische Verhandelingen, 19: 1-58.
  40. ^ Altena, C. O., 1953. “A Revision of the Genus Nyctalemon Dalman (Lepidoptera, Uraniidae) with Notes on the Biology, Distribution, and Evolution of its Species”. Zoologische Verhandelingen, 19: 1-58.
  41. ^
    Altena, C. O., 1953. “A Revision of the Genus Nyctalemon Dalman (Lepidoptera, Uraniidae) with Notes on the Biology, Distribution, and Evolution of its Species”. Zoologische Verhandelingen, 19: 1-58.
  42. ^ Butler, A. G., 1869. “Descriptions of species of Lepidoptera, confounded with others described by Linnæus and Fabricius”. The Entomologist's Monthly Magazine, 5:270-273.
    tree
  43. ^ “Lyssa zampa”. Revolvy. URL: https://www.revolvy.com/main/index.php?s=Lyssa%20zampa&uid=1575 (accessed on 15 Nov, 2016).
  44. ^
    Beljaev, E. A., 2009. “Phylogenetic relationships of the geometroid lepidopterans
    (Lepidoptera: Cimeliidae, Epicopeiidae, Sematuridae, Drepanidae, Uraniidae, Geometridae)”. Spixiana 32 (1): 134-136.
  45. ^ Beljaev, E. A., 2009. “Phylogenetic relationships of the geometroid lepidopterans
    (Lepidoptera: Cimeliidae, Epicopeiidae, Sematuridae, Drepanidae, Uraniidae, Geometridae)”. Spixiana 32 (1): 134-136.
  46. ^ Nazari, V., 2016. “Century-Old DNA Barcodes Reveal Phylogenetic Placement of the Extinct Jamaican Sunset Moth, Urania sloanus Cramer (Lepidoptera: Uraniidae)”. PLoS ONE, 11 (10).
  47. ^
    Beljaev, E. A., 2009. “Phylogenetic relationships of the geometroid lepidopterans
    (Lepidoptera: Cimeliidae, Epicopeiidae, Sematuridae, Drepanidae, Uraniidae, Geometridae)”. Spixiana 32 (1): 134-136.
  48. ^ Nazari, V., 2016. “Century-Old DNA Barcodes Reveal Phylogenetic Placement of the Extinct Jamaican Sunset Moth, Urania sloanus Cramer (Lepidoptera: Uraniidae)”. PLoS ONE, 11 (10).