Papilio+demoleus







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**1.Introduction** The Lime Butterfly**, //Papilio demoleus//** is a commonly widespread swallowtail butterfly found in open areas that spans from South Asia to Australia The subspecies **//Papilio demoleus malayanus//** is found mainly in the region of Southeast Asia and can usually be sighted visiting flowers n the gardens of Singapore residential areas. The name Lime Butterfly is often derived from its foodplant that its caterpillars/larva feeds on //Citrus spp.// plants such as __[|Key Lime]__(//Citrus// //aurantifolia//) .Due to the import of __[|kumquats]__ (//Citrus japonica//) during Chinese New Year season in Singapore, you can see them more often as there are more food for their caterpillars to feed on.

**1.1 Etymology** “Papilio” is derived from the Latin term “papilionem” and the French term “papillon”. Both term mean “butterfly, moth”. There is no clear meaning behind the species demoleus coined by Linnaeus as the name ”demoleus” refers to a greek man who had been slain by Aeneas, a Trojan involved in the war of Troy. “Malayanus”,as the name suggested refers to the Malay peninsula where Wallace establish the subspecies as a local form in Southeast Asia.



1.2 A Pretty Pest
Although the adult lime butterfly may look pretty in our eyes, the caterpillars have a large appetite is considered as minor garden pest in Singapore. For countries with large lime plantation industry, these caterpillars can pose a serious threat in the __[|Old World]__ such as India and Saudi Arabia. They are aggressive pests and spread across foreign countries very quickly. Recent findings have shown that this species has spread to the __[|New World]__ such as Dominican Republic and Puerto Rico, threatening the livelihood of farmers that depends on lime cultivation. Therefore the lime butterflies pose as a serious threat to these countries Guerrero, K. A., Veloz, D., Boyce, S.L., Farrell, B.D. . (2004). First New World documentation of an Old World citrus pest, the Lime Swallowtail //Papilio demoleus//(Lepidoptera:Papilionidae), in the Dominican Republic (Hispaniola). //American Entomologist, 50//(4), 227-229.

In this species page, we will try to understand more about the biology of //Papilio demoleus// in order to show the impact lime butterflies have on humans as an invasive species and an economic pest. Taxonomic information will be provided the help in identification of the butterfly through its __description__, __diagnosis__ and __distribution__ throughout the world. Lastly, __systematic information__ about the species and its related outgroups will be provided show the comparison between //Papilio demoleus// and other similar looking species, of which, some are rare and can only be found in Madagascar. Difficult term definitions can be found in the hyperlink for your convenience

 Think you spotted the lime butterfly ? . Check out our __[|diagnosis]__ to see if you are correct! = 2. Life cycle and behavior = As all butterflies go through __[|holometabolous development]__ or complete metamorphosis, that is there are 4 stages in the life cycle of the lime butterfly. However it is difficult to distinguish lime butterfly from other species when it is larvae or pupa stage. In Singapore, the larval and pupa stages of lime butterfly often resemble that of [|Common Mormon](//Papilio polytes//) and __[|Great Mormon]__ (//Papilio memnon//). Therefore upon spotting of these caterpillars, we can guess they belong to these 3 species. The following description of the life cycle is based on the book “Caterpillars of Singapore’s Butterflies” by Horace Tan and Khew Sin Khoon.

Description
Eggs are laid singly on young leaves or on the underside of the leaves as well as the surface of twigs on lime host plant. The eggs laid are spherical and smooth edged. Due to the tropical climate of Singapore, the time for embryo development is very short, about 2-3 days.

Description
The lime butterfly caterpillar goes through 5 __[|instar]__ stages.During its early stages, the caterpillar is light brown in colour with white markings, closely resembling a bird’s droppings as camouflage. As it grows, it will molt or shed its exoskeleton. Every time it molts, it will grow from the 1st instar to the 2nd instar. From the 1st to the 4th instar, it appearance does not change much other than its size. The white markings in the following 3rd instar specimen (meaning it has shed its skin 2 times) is V-shaped that covered the top and the side of its body. A row of spines can also be observed on the top side of the body. As the caterpillar grows, the size of the white marking will extend and the color will shift from light brown to dark brown or black. Refer to this __[|link]__ for more details

On the 5th instar, the caterpillar changes to yellowish green / green in color with no spines, closely resembling the color of its lime host plant. In the middle or 3rd __thoracic segment__, there are 2 lateral __false eyespots__ with a yellowish brown band spanning across the segment. Base on this specimen, the pillar has brown marking on both sides of its body. It is during this latter stage that the larva can die very easily.

Feeding behavior
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Video 1: Lime catepillar feedin behaviour (Source: Youtube, Video by Kang Rui) Based on personal observations, the caterpillar is not active during most of the days. Instead, it simply lay motionless on the host plants throughout or simply “hiding in plain sight”. Upon feeding, it will move to the edge of the leave and search for the leave edge. The __prolegs__ will grasp onto the the leaf and chew from the edge onwards with the help of it 3 pairs of __thoracic legs__.

Description
The pupae take the colour of either green or brown depending on the surface that it pupate on. Pupating on non-woody or smooth surface usually results in a green pupae while pupating on woody or brown surface gives a brown pupae. . The pupa is supported by the silk girdle across the thorax and attachment on __cremaster__.

Pupation
When the caterpillar is ready to pupate, it will orientate itself to the wall with its head facing upwards and expelling its or waste from its system. A network of __silk pad__ will be spun onto the surface and attach its body to the __silk girdle__ for extra support. Then the body of the caterpillar will shorten in length gradually forming a hunch. After a day, the body will harden into a __[|chrysalis]__ (unlike moth that usually spin a cocoon), forming a harden pupa with a pair of __anterior horns__ and bent away from the support.

Description
(see the __description__ and __diagnosis__ section)

Eclosion
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Video 2: Ecolsion of lime butterfly (Source: Youtube, Video by Kang Rui) After the pupal stage, the butterfly will under go __[|eclosion]__ or emerge from its pupa. Firstly the pupa colour will change to black and yellow, resembling the wings color of the adult butterfly. A slit will be open at the anterior portion of the chrysalis. The butterfly will wiggle from the chrysalis and push itself away from the wall to free its 2 front legs. After achieving that, it will climb out onto the silk pad or branch to dry its wings. It will slowly expand and dry its wings before fluttering off to feed.

Feeding behaviour
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Video 3: Locomotion and feeding behaviour of the adult lime butterfly (Source: Youtube, Video by Kang Rui) Based on personal observation, the butterfly will come to visit the flower at 8-9am in the morning. The latest time a lime butterfly was sighted was at 4pm. Compared to other butterflies in the garden, the lime butterfly fast very fast and actively particularly in the late morning and early afternoon where the temperature is the hottest inthe day. It will dart quickly along the bushes of the flowers and feed on the nectar of any nectar plants such as //__[|Ixora spp]__// and //__[|Bougainvillea spp].__// There is no special preference of flower it feeds on.

= 3. Adaptations = So how does it survive so well out in the wild and urban environment? The lime butterfly is packed with many forms of defense and adaptations to avoid and fend off their prey.

Body camouflage
As mentioned in the life cycle, the caterpillars of the lime butterfly resemble 2 things: bird droppings (__1st – 4th instar__) and green leaves like its host plant (5th instar). Its camouflage will trick potential predator into ignoring them as inanimate objects or hard to see

Fig.3.1 The colouration of the catepillar body resemble inanimate object such as bird's dropping or lime leaf

False eyespot
Eyespots can be found on both the adult butterfly and as well as on the caterpillar in its 5th instar. It can serve as a form of mimicry whereby the eyespot will look the the eyes of another animals, tricking potential predators by drawing their attention away or appear as an dangerous animal.



Fig. 3.2 False eyespots of the lime caterpillar (left) and adult lime butterfly wing (right) (Source: Eunice & Kang Rui)

Osmeterium – a smelly weapon
media type="custom" key="24538590" align="center" Video 4: Eversion of osmeterium as defense by lime caterpillar (Source: Youtube, Video by Kang Rui) __ Osmeterium __ is a defensive gland that produces a pungent smell to ward off predators. When threatened, it will protrude its osmeterium from between the head and __prothorax__ for a short period of time. The form of defense is present in all __instars__ of the lime butterfly caterpillar include other //Papilio// spp. (Swallowtails). The strong odor comprises a mixture of volatile organic acids (isobutyric acid and 2-methylbutyric acid) as __[|pheromones]__.

Large appetite
The caterpillar have large appetites and is recorded that they can eating up 83% of the leaves on young grove trees in Andra Pradesh, India.

So how does it find the correct leaves to feed on? According to Khattar (1978), the caterpillar find its food by detecting the green colour as well as the odour of the //Citrus spp.// host plant called __[|Citral]__.

= 4. General Anatomy =

4.1 Adult wing anatomy
Identification of butterfly species is heavily based on morphology and in particular the wing anatomy. The venation pattern is based on **Comstock-Needham system** where the venation categories are based on insect instead of Lepidoptera i.e butterflies and moths (Venation of the wings, starting from the anterior are as follows :-

- __ Subcosta __ (Sc), - __ Radius __ (R1 - R5), - __ Medius __ (M1- M3), - __ Cubitus __ (Cu), - __ Anal veins __ (1A,2A)



4.4 Pupa anatomy
==  = 5 General Description = So what does the lime butterfly //Papilio demoleus// look like?

Butterflies are generally classified based on their wing patterns which are distinct between species. The description pertains to the species //Papilio demoleus//. Description of the subspecies can be found in the diagnosis section. Butterflycircle.com also provides concise information about //Papilio demoleus malayanus// in Singapore. The current description is based on Bascombe(1999), Braby (2000) and Corbet (1992).

5.1Dorsal (Upper side) Wing
a) Dorsal side of wings are brownish-black in color with a chain of irregular light yellow spots along the __discal__ (red), __submarginal__  (green)  and  __marginal__  (blue) regions    b) __ Hindwings __ has a black __costal__ eyespot near __apex__ that gradually spreads over the blue scales shaped as a crescent. c) In __space 1b__ on the dorsal hind wing, there is a red __tornal__ spot for both sexes**.** Different genders usually have distinctly dissimilar color composition ** . ** See __Differentiation of gender__

5.2 Ventral(Underside) Wing
a) Ventral side of wings are brownish-black (forewing) / yellow (hindwing) in color with a chain of irregular light yellow spots along the __ discal __ (red), __ submarginal __ (green) and __ marginal __ (blue) regions   b) A series of dull-yellow spots are located on the __sub apical__ (below the apex) of the fore wing c) Underside of hindwings has a series of elongated central (__post-discal__) orange spots edged with iridescent blue and black.  d) __ Costal __ spot near apex is orange edged with blue and black.

5.3 Sexual dimorphism
** a) **** Tornal spot pattern **  Males – tornal spot on upper side hind wing is **mostly orange** with little blue scale    Females - tornal spot on upper side hind wing is **partially orange** with more blue scales  ** b) ** ** Ventral hind wing color ** Males – underside of hind wing are **light yellow** in the discal and submarginal region Females – underside of hind wing is **dull yellow i**n the discal and submarginal region

In many specimens the light yellow spots are darken to dull yellow with age

** c) ** ** Shape of terminal abdomen segment **  Male – **sharper** abdomen shape    Female – **round** abdomen shape

Gender differentiation is most reliably determined by the presence of __[|claspers]__, part of the male genitalia.

= 6. Diagnosis =

**// 6.1 Papilio demoleus //**** group (Lime Swallowtails) **
The name lime swallowtail is shared by 5 different species and is often confused with //Papilio demoleus//. Some of which are // Papilio demoleus // group comprises the following 5 species that are considered lime swallowtails and has the same wing pattern of each other 1 See Appendix for visual photo

** 1) //__Papilio demoleus__// Linnaeus,1758 ** ** 2) //__Papilio morondavana__// Grose Smith, 1891 ** ** 3) //__Papilio grosesmithi__// Rothschild,1926 ** ** 4) //__Papilio demodocus__// Esper,1799 ** ** 5) //__Papilio erithonioides__// Grose Smith, 1891 **

** 6.2 Differentiation of //Papilio demoleus// from other Lime Swallowtails **
Based on the dichotomous keys provided by Smith & Vane-Wright(2008).There are 2 essential character states that differentiate Papilio demoleus from other lime swallowtails

**// a) //** **// P.demoleus vs P.morondavana & P.grosesmithi //** **// P. demoleus //** – tail on hindwing vein M3 is rudimentary or very short, less than twice as long as those of neighbouring veins   **// P.morondavana, P.grosesmithi //** – hind wing vein M3 extended to form distinct tail at least 3 times as long as extensions of neighbouring veins

**// b) //** **// P. demoleus vs P. erithonioides & P.demodocus //** **// P.demoleus //** - forewing upperside cell M1 with post-discal mark absent   **// P.erithonioides & P.demodocus //** – forewing upperside cell M1 with post-discal mark present

** c) ** ** External genitalia **

The external male genitalia differs from other members of the genus papilio is that the __[|clasper]__ continues strongly upwards posteriorly to end in aserrate plate beneath and lateral to the __[|pseuduncus]__.

**// d) //** **// Locality //**

**// Papilio demoleus - //** South Asia, East Asia, South-east Asia, Australia and Arabian Penninsular (Green) **// Papilio demodocus – //** Africa and Arabian Peninsular (Red) **// Papilio morondavana- //** Madagascar (Yellow) **// Papilio grosesmithi- //** Madagascar (Yellow) **// Papilio erithonioides- //** Madagascar (Yellow)

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Map 1: Distribution of Lime Swallowtails throughout the World (Yellow: morondavana, grosesmithi, erithonioides, Red: demodocus, Green: demoleus) (Source : Google Maps, Created by Kang Rui) =** 7. Distribution **=

7.1 Distribution in the World (//Papilio demoleus)//
// Papiio demoleus // is distributed across the world stretching from South asia to East Asia. The species then spread down south all the way to Australia. The three most common subspecies are:-

- **// Papilio demoleus demoleus //**, Linnaeus 1758 which dominate the Arabian peninsular, South Asia and East Asia - **// Papilio demoleus malayanus //** Wallace, 1865 which dominate the Southeast Asian region and the Malay Archipelago - **// Papilio sthenelus //**, Macleay, 1826 which dominate the Australian continent.

media type="custom" key="24538880" align="center" Map 2: Distribution of Papilio demoleus subspecies throughout the World (Red : P.d.demoleus, Blue: P.d. malayanus, Green: P.d.sthenelus) // Papilio demoleus malayanus // however have recently been introduced into the New World as specimens collected from Jamaica .Dominican Republic Puerto Rico appear to resemble the subspecies. Methods of introduction are unknown according the articles

7.2 Distribution in Singapore
Generally, the subspecies //Papilio demoleus malayanus// can be found in most open spaces of urban Singapore. The planting of nectar flowers around residential estate help to attract these butterflies to the areas. In old estates such as Hougang areas, many residents would plant lime such as [|Key lime](//Citrus aurantifolia//) at their backyard, providing favourable host plants for the larvae of lime butterflies to feed on. The following map shows some of my personal recording in collecting lime butterfly specimens.

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Map 3: Locations of Lime butterfly spotted in Singapore = 8. Taxonomy =

8.1 Name
**Binomial: **  Species name //Papilio demoleus // Linnaeus 1758

**Synonyms ** //Princeps demoleus // Linnaeus 1758 //Papilio erithonius // Cramer,1779 //Papilio epius // Fabricius,1793 //Papilio demoleinus // Oberthur,1879

**Common names ** <span style="font-family: Arial,sans-serif; font-size: 16pt;"> Common Lime Butterfly <span style="font-family: Arial,sans-serif; font-size: 16pt;"> Lemon Butterfly <span style="font-family: Arial,sans-serif; font-size: 16pt;"> Lime Swallowtail <span style="font-family: Arial,sans-serif; font-size: 16pt;"> Chequered Swallow tail <span style="font-family: Arial,sans-serif; font-size: 16pt;"> Citrus Swallowtail

8.2Taxonavigation
Animalia Arthropoda Insecta Lepidoptera Ditrysia Papilionoidea Papilionidae Papilioninae Papilionini ** // Papilio // ** ** // demoleus // **

8.3 Other lime swallowtails (//Papilio demoleus// group)
1) //Papilio demoleus// Linnaeus,1758 2) //Papilio morondavana// Grose Smith, 1891 3) //Papilio grosesmithi// Rothschild,1926 4) //Papilio demodocus// Esper,1799 5) //Papilio erithonioides// Grose Smith, 1891a

** 8.4 //Papilio demoleus// subspecies **

1) //P.d .malayanus// Wallace, 1865 2) //P.d .demoleus// Linnaeus, 1758 3) //P. d. sthenelus//, Macleay, 1826 4) //P.d. stenelinus// Rothschild, 1895 5) //P.d. novoguineensis// Rothschild,1908 6) //P.d .libanius// Fruhstorfer, 1908 (= //P.d.demoleus// Linnaeus, 1758)

8.5 Types series
<span style="font-family: Arial,sans-serif; font-size: 16pt;">The holotype of the //Papilio demoleus// is found in Royal Belgian Institute of Natural Sciences Collection holotype catalogue no. : 312055

Honey & Scoble (2001) discussed the specimens probably seen by Linnaeus and from which he first described //Papilio demoleus// Linnaeus, 1758. The specimen selected from Guangzhou (Canton), China as the lectotype.

The male lectotype of //Papilio demoleus malayanus// is found in the British Museum of Natural History in London, United Kingdom. The specimen acquired was from habitat in Singapore. The lectotype is a specimen from Wallace’s private collection

= 9. Systematics = Fig. 8.2. One of two equally parsimonious trees (TL 52533, CI50.548, RI50.554) based on combined dataset. Most parsimonious trees were different only in some outgroup rearrangements. Bayesian analysis produced tree with almost identical topology (-lnL5 17024.14) with different position of Papilio anactus shown by a circled star. Histograms above nodes show partitioned Bremer supports, numbers under nodes indicate bootstrap proportions, total Bremer support, and clade posterior probabilities from Bayesian analyse (Diagram from Zakharov et al.,(2004a) Permission not acquired but within the limits of Fair Use) __ Cladistic analysis __ done by Zakharov et al. (2004a) involves the molecular __phylogeny__ and morphological characteristic to test for phylogenic relationship between the species in the //P.demoleus// group and outgroups of the //Papilio// genus. Based on the __cladogram__ derived from the combined data (COI, COII, Wg, EF1a genes and morphology), here is a strong support between the species in the P.demoleus group, showing that //P.demoleus// group is monophyletic.

Shift of basal position from P.demoleus or P.demodocus to P.morondavana. The strongest evidence indicates that a clear sister relationship between //P.demodocus// and //P.erithonioides// is establish with //P.demoleus// likely to be their sister taxon. //P.grosesmithi// is the sister taxon of the 3 species and //P.morondavana// takes the basal position. The data no longer support that //P.demodocus// or //P.demoleus// is the most basal species. Therefore the finding suggest that we should consider conservation projects on //P.morondava// as it holds the oldest lineages in the group. **Despite sharing many similarities with the pest //P.demoleus, P.morondavana// is endemic and should be protected for preservation of the group’s lineage.**

= 10.Human Impacts =

10.1 P.demoleus as a global pest.
With a large dispersal range, high reproductive range and wide range of larval host plants, the spreading of //P.demoleus// population are often fast and rampant. Based from Atluri et al. (2002), the //P.demoleus// is capable of reproducing up to six generation annually in India and has a preference for 19 species of citrus varieties including cultivated species. There is also a positive correlation of //P.demoleus// abundance with seasonal increase in temperature and relative humidity in West Bengal, India. Originally located in the Old World, a recent record of //P.demoleus// species which is likely to be the subspecies from Southeast Asia or //P.d.malayanus// was found in Dominican Republic in 2004, followed by Puerto Rico in 2006

Beginning in the 1970s, invasion of //P.demoleus// have already occurred in the islands of Java, Borneo, Philippines and Sumatra. The invasion was facilitated by deforestation and the increase availability of citrus grove. //P.demoleus// is also known to plague citrus cultivation in Saudi Arabia and Iran. . has also reported that up to 83% defoliation of young grove tress in Andra Pradesh, making //P.demoleus// an economic pest in India as well.

The recent invasion of //P.demoleus// happened to have occurred by a single introduction in 2004 in Dominican Republic as evidence showed that there is a lack of genetic variation among the specimens sampled in Dominican Republic. It is however not known how the species reached the Dominican Republic. Homziak & Homziak (2006) suggested that the invasion of P.demoleus in Puerto Rico in March 2006 is likely originated from the Dominican Republic.

With the presence of favorable climatic conditions, favorable host plant and potential lack of predators, the establishment of //P.demoleus// in Dominican Republic is inevitable



10.2 Economic Importance
The invasion of //P.demoleus// into the New World such as Dominican Republic and Puerto Rico has raised alarm with regards to the potential threat this species may have on the agricultural industry of United States and Hispaniola. Based on estimates from the Caribbean cooperative citrus association (CARICOM), the production of Citrus products amounts to an approximate 1,100,000 in CARICOM countries such as Cuba, Dominican Republic and Haiti, generating an amount of US$61 million annually (Homziak & Homziak,2006) Furthermore, small citrus farms make up between 93 and 98 percent of all CARICOM farms. Therefore the industry is critical to the economic survival of many small farmers .Based on National Agricultural Statistic Service (2004), the value of Florida citrus crop exceed, US 1.5 billion dollars and United States total worth was more than US$2.6 billion dollar National Agricultural Statistical Service (2004). USDA Citrus Fruits Final Estimates, 1997–2002. //United States Department of Agriculture Statistical Bulletins, 997//(4), 1-30. . With the population of //P.demoleus// established in Dominican Republic, it is likely that the population can easily spread through the West Indies and surrounding areas, including the United States where impact on the agricultural industry could be catastrophic.

Biological control
Many literatures have proposed biological control of //P.demoleus// through the use of predators and natural pesticide. Biological predators that could be utilized to control the population - Larva parasitoids - Pupal parasitoids - Adult predators Biopesticides[| Bacillus thuringiensis] and //__[|Beauveria bassiana]__// as well as neem seed kernel extract and [|Azadirachtin] were shown to have effects on // P.demoleus // in India.
 * - Egg parasitoids
 * o //__ [|Ooencyrtus malayensis] __// (Hymenoptera: __[|Encyrtidae]__), Thailand
 * o // [|Tetrastichus] // sp.(Hymenoptera: __[|Eulophidae]__), Thailand
 * o [|Chalcidoid] (Hymenoptera), Jamaica
 * o // [|Erycia nymphalidophaga] // (Diptera: [|Tachinidae]), Thailand
 * o // [|Apanteles papilionis] //, (Hymenoptera: [|Braconidae])India
 * o // [|Bracon hebetor] // (Hymenoptera: [|Braconidae]), India.
 * o // [|Brachymeria]sp. // (Hymenoptera: [|Chalcididae])
 * o // [|Pteromalus puparum] // (Hymenoptera: [|Pteromalidae]).
 * o // [|Cantheconidea furcellata] // (Hemiptera: [|Pentatomidae])
 * o [|Reduviidae]//sp//. (Hemiptera)
 * o Spiders
 * o [|Sphecids] (Hymenoptera)
 * o Chameleons and birds || [[image:pdm_braconid_parasitoid.jpg caption="Bracoid parasitoid feeding off P.demoleus larvae (Photo by J.M. Garg,(2009)from Wikicommons)"]] ||

__[| Neem]__ based insecticides(multineem and neemark) could serve to alter the life-processing behavior that the insect can no longer feed, breed or undergo metamorphosis. The insecticide could induce defecation and sporadic jerking of the body. Heavy pigmentation in the 5th instar larvae, disappearance of red spot and change in wing shape, significant reduction of oocytes (eggs) number are some of the effects on //P.demoleus//

Non-biological control
Regulation of fatty acid has known to influence the growth development of insects. Methat (1970) has demonstrated that the use of fatty acids could serve to inhibit the growth of caterpillar larva. Treatment of lime leaves with oleic, linoleic and erucic acid at concentration higher than 0.25% proved to be toxic to the larva and inhibit pupation.

= 11. Conclusion = In summary, we can see that the appearance of //P.demoleus//, an economic pest, shared many similarities between its sister species in the //Papilio demoleus// group. Some of which are endemic. Thus, this species page hopes that people can identify such difference in the hopes that we eradicate the correct species instead of destroying an endemic one. Who is the true criminal. = 12. Useful links. =  1) [|Butterflycircle.com] - A discussion forum pertaining to butterflies in Singapore

2) [|Singapore butterfly checklist]- Provides an easy checklist of the butterflies in Singapore 3) [|Life History of Lime butterfly] - A blog from butterfycircle that features a great amount of detail on the life stages of the lime butterfly in Singapore 4) [|Samui butterflies] - A aesthetically pleasing photo gallery featuring butterflies of southeast asia and of cours lime butterflies 5) [|Learn about butterflies] - Provides a generally knowledge about butterflies in the world.

= 13. Appendix = Other lime swallowtails

// [|Papilio morondavana] //

 * Description

1) Hindwing vein M3 extended to form **distinct tail** at least 3 times as long as extensions of neighbouring veins. 2) Forewing upperside discal cell with basal pale scales apparently **randomly scattered**, not forming any noticeable pattern. 3) Forewing cells R3 – M1 with submarginal marks **large and clear and sharp edged**, that in R3 forming a bar across the cell 4) Hindwing upperside cells R5- M3 either **shallowly concave** or with **slight indentation** distally

Locality - Madagascar || ||  ||

// [|Papilio grosesmithi] //

 * Description

1) Hindwing vein M3 extended to form **distinct tail** at least 3 times as long as extensions of neighbouring veins. 2) Forewing upperside discal cell with **basal pale scales aggregated into narrow transverse** markings 3) Forewing cells R3-M1 with submarginal marks **small and faint** 4) Hindwing upperside cells R5-M3 with submarginal marks **deeply indented** distally, that in M3 chevron-shaped.

Locality - Madagascar || ||  ||

// [| Papilio erithonioides] //
2) Forewing upperside cell M1 with post-discal mark **present** 3) Basal pale marks in hindwing underside cell R1 and discal cell very **broad**, separated from discal marks only by a narrow dark band. Locality - Madagascar || ||  ||
 * 1) Tail on hindwing vein M3 rudimentary or very **short**, less than twice as long as those on neighbouring veins

// [|Papilio demodocus] //
1) Tail on hindwing vein M3 rudimentary or very **short**, less than twice as long as those on neighbouring veins 2) Forewing upperside cell M1 with post-discal mark **present** 3) Basal pale marks in hindwing underside discal cell and cell R3 **narrow**, separated from discal marks by a dark band much broader than the basal marks Locality  - African continent  - Arabian peninsular ||  ||  ||
 * Description

// [|Papilo demoleus] //
1) Tail on hindwing vein M3 rudimentary or very **short**, less than twice as long as those on neighbouring veins 2) Forewing upperside cell M1 with post-discal mark **absent** Locality - Arabian peninsular - South Asia - East Asia - Southeast Asia - Australian continent || ||  ||
 * Description