The Painted Lady
Vanessa cardui (Lepidoptera: Nymphalidae) Linnaeus, 1758
Vanessa cardui perching on butterfly bush flowers Buddleja. Image by Thom Mitchell (Permission pending)
Acknowledgment: Data were provided by the Butterfly and Moth Information Network and the many participants who contribute to its Butterflies and Moths of North America project.
Vanessa cardui, also known as the painted lady butterfly, boasts the world's farthest known butterfly migratory route, beating the well-known North American migratory Monarch butterfly Danaus plexippus! As one of the most cosmopolitan insect species in the world, the painted lady is found in all continents except Antarctica. Some of the reasons for its widespread distribution include a wide variety of plants it feeds and lays eggs on, the ability to migrate and avoid winter, and continuously reproducing. This almost global distribution means that its pretty wings can be observed around the world and easy to rear for scientific studies. Of course, they are also not classified as endangered anywhere.
Recently, efforts have been devoted to understanding the exact migration routes V. cardui take. If you want to learn and contribute to knowing where they travel to and breed, you can become a Citizen Scientist by contributing to this global project on V. cardui. But first, just scroll down to find out more about this extravagant Painted Lady butterfly!
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I. Biology
Morphology
Like all other insects, adult butterflies have three body parts - the head, thorax and abdomen, as well as three pairs of legs attached to the thorax. Butterflies are known for their two pairs of charismatic wings, often dotted and striped in different colours.
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Dorsal surface | Ventral surface |
Images by Didier Descouens, licensed under CC BY-SA 4.0
With wing spans ranging from 5.1 to 7.3 cm, the painted lady butterfly exhibits different colour patterns on the dorsal (upperside) and ventral (underside) wing surfaces, contributing to its conspicuity.
The dorsal surface consists of white and black pattern elements like spots and stripes, against an orange background. The ventral surface, on the other hand, had a greater variety of colours including blue, yellow, brown and grey on top of the three colours observed on the dorsal surface. Of particular interests are the eyespots, which are made up of concentric rings of different colours, and found on the ventral hindwing of these butterflies.
Wings of these lepidopteran insects are widely diverse in colours and patterns. So what are these wings made of? Studies have found that butterfly wings are made up of scale cells, and each wing scale (approximately 0.8 to 40 mm2) only one colour. This means that a single eyespot is dotted by a variety of scale cells of different individual colours. It is a wonder how dead scale cells give rise to the vibrant array of colours on the wings!
Life cycle
Found in temperate and subtropical regions, the painted lady butterfly exhibits seasonal migration and breed throughout the year. The female lays eggs during its migration, contributing to its almost global distribution. It does not have subspecies, possibly a result of migratory behaviour that prevents the diversification of local populations1 .
Like other holometabolous organisms, the painted lady butterfly undergoes metamorphosis as part of its life cycle that ranges from 1 month in subtropical areas to 2 months in the temperature region2 . They can be found in open or disturbed urban areas. Eggs are laid singly on plant leaves and develop over the course of 5 days, before the caterpillar hatches. Five larval stages, also known as instars, span over 25 days where the caterpillar feeds continuously and grows extensively, followed by a pupation stage when metamorphosis takes place. The adult butterfly emerges from the pupa after a week and finds a mate before settling on a leaf to lay eggs and start the cycle again.
A single V. cardui egg. Image by Harald Süpfle, licensed under CC BY-SA 3.0
Eggs are laid singly on upperside leaves of foodplants. The adult female butterfly first finds a host plant where its caterpillars would have abundant food, then oviposits about 500 eggs.
After hatching, the caterpillar crawls to the underside of the leaf and start its voracious feeding life. Born with six eyes, it then undergoes a total of four moults, where it sheds its skin to grow larger.
Vanessa cardui larva hiding under a silk tent made on a thistle leaf. Image by Michael J. Plagens (permission pending).
The caterpillar builds a silk tent across one or more leaves that it feeds on and excrement accumulates, making it easy to spot. It leaves the tent when it is ready to pupate.
Vanessa cardui undergoing pupation (left) then metamorphosis (right). Video adapted from Birdy Official (permission pending).
After emerging from its chrysalis, the adult butterfly rests and unfolds its wings slowly to dry. A few hours later, it will be able to find food and mates to produce the next generation of painted ladies.
Diet
More than 100 host plant species of the painted lady butterfly has been recorded around the world. Among the host plants, V. cardui prefers thistles (Asteraceae), hollyhock and mallow (Malvaceae), and various legumes (Fabaceae)3 .
Tricoloured buddleia. Image by © YouGarden 2018
While V. cardui is a polyphagous species, some of its favourite food as a larva is thistle like Buddleia, which can be an invasive weed, and common mallow Malva sylvestris (Malvaceae). Unlike the adults with a proboscis mouthpart, larvae have chewing mouthparts. These mouthparts allow them to munch on an immense volume of leaves that support their size increase4 .
The painted lady butterfly prefers open fields and urban areas, like flower meadows and rural hills. In Singapore, they have been observed a few times, at one of our urban hill parks, at an open wasteland in the north and one at Pasir Panjang Nursery5 .
Multi-generational migration
Migration
The most amount of attention diverted to V. cardui is its world's longest return migratory journey. In particular, the most well documented trip was by Stefanescu et al. (2013), who reported an exceptional large number of V. cardui migrating across Europe in 2009. Vanessa cardui returned to the Europe from its overwintering range in Africa and the trip was observed to climb to high altitudes about 1000 metres above sea level.
Distribution
Described as cosmopolitan, the extant Vanessa cardui has one of the world's largest distribution, found in almost all continents including Europe, Africa, Asia, North and South America, except Antarctica6 .
Proposed global distribution of Vanessa cardui. Map by Discover Life.
Exhibiting multi-generational migratory behaviour, the painted lady butterfly stops in Singapore occasionally during winter migrations from East Asia, with sightings since 20077 . Migratory behaviour that moves them from temperate regions, where they spend spring and summer, to subtropical regions in autumn and winter ensures constant access to resources and avoid parasitism8 . Travelling over thousands of kilometres, these butterflies have been found in Svalbard, Norway, vastly distant from breeding grounds. The abundance and wide range habitats of V. cardui allows for the non-listing as an endangered species.
The interesting thing about migration in V. cardui is the span of the migration cycle over multiple generations. Unlike vertebrates including large mammals, which have generation times longer than the migration cycle, migratory insects with short lifespans either travel a shorter distance, or make stops to reproduce during migration9 . Well studied migration routes include between North Africa and Europe, and between North and South America. In both cases, these butterflies make a round trip and these annual circuits span a minimum of 10 generations10 . Other than the Americas, Europe, Africa and Asia, the painted lady is not found in areas east of the Wallace line including south Australia, as well as Antarctica.
Locally, there has only been sporadic sightings of V. cardui in Singapore between 1990 and 201711 . Hopefully, we get to see more of this elegant Painted Lady in years to come.
Heat regulation
As with other invertebrates, V. cardui is ectothermic. This means that it is unable to regulate its body temperature and is heavily dependent on the environment. Correspondingly, it usually avoids the cold due to its inability to overwinter as well, by migrating to other regions where the temperature is warmer. The painted lady butterfly takes advantage of the sun by basking in the heat.
V. cardui basking on a hot sunny afternoon. Image by © 2018 North American Butterfly Association.
Basking is the act of using specific wing postures and orientation according to solar and wind direction. When it gets too hot, the butterflies simply fold their wings back to prevent overheating.
Apart from warming up, V. cardui also have to face ephemeral periods of cold in their breeding regions in the Northern Hemisphere, before they make their long migration. Shivering behaviour in butterflies involves the contraction of flight muscles against each other to produce heat12 . This behaviour is observed in insects generally, but is not well studied. A further investigation of this shivering behaviour could elucidate the ability of such tiny insects to traverse across long distances and high elevations.
Did you know?
Different colours observed on butterfly wing patterns could be a result of either pigment colours (like green chlorophyll) or structural colours (like blue iris in the human eye)!
The great migration
Migration is a common tactic used by many living organisms to move away from predators, unfavourable climate or lack of food. However, organisms, especially small insects, face problems of drifting away from historical routes due to the wind or water currents13 . The painted lady butterfly overcomes this by solar navigation14 .
II. Species diagnosis
As of today, there are a total of 22 species in the genus Vanessa.
Vanessa species with dorsal (left) and ventral (right) wings shown15
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The most closely related species to V. cardui is Australian painted lady, V, kershawi.
Enlarged images of dorsal and ventral wing surfaces of V. cardui and V. kershawi. Adapted from Abassi & Marcus (2015).
In Singapore, V. cardui is distinct from other butterflies in Singapore so any luck with finding a butterfly with a white stripe with splodges of orange and black should make it easy to identify as V. cardui.
Vanessa kershawi, the Australian painted lady. Image by Fir0002/Flagstaffotos, under CC BY-NC.
Note the huge similarity in wing colour patterns between the two Vanessa species. Despite the glaring similarities of the two species in behaviour and morphology, the most prominent difference is phenotypic plasticity in V. kershawi. Phenotypic plasticity is exhibited by different appearance or behaviour of organisms when they are exposed to different environments like varying temperatures. In V. cardui, stress or low temperatures can induce darker colour wing patterns like those in V. kershawi16 . Talk about becoming darker under stress!
III. Uses in scientific studies
Vanessa cardui is widely used for scientific studies due to its abundance and ease of rearing. The studies range from examining nanostructures of wings to biological studies of environmental effects on organisms.
Material Science
Following the studies of shells, bones and honeycombs, butterfly wings have become the next natural architecture to be studied17 . As mentioned previously, butterfly wings are made up of many scales, each contributing a single colour. These scales have been found to be made of chitin, which is the second most abundant organic compound after cellulose, and chitin is also a component of crab shells and squid pens (a hardened internal body part)18 . The dazzling structures of butterfly wings make them candidates for replicating these biomimetic structures for solar cells and advanced sensors. However, these nature-engineered structures are difficult for mass production at the moment, and the ongoing evolutionary process means that these structures are continuously altered but it is difficult to replicate natural modifications19 . These studies are pioneering the field and future research could resolve these issues for the replication of naturally occurring structures.
Phenotypic plasticity
Vanessa cardui was first found to be temperature-sensitive in a study by Nijhout (1985), where cold-shocks administered to led to individuals developing aberrant colour patterns20 .
This observation was then further explored by Yamanaka et al. (2009), where pupae of V. cardui were found to exhibit phenotypic plasticity. Different shades of wings appeared under varying temperature conditions, as observed from the image below.
An increase of 14°C led to a shrink in the black central spot (indicated by the white triangles) in the eyespots on the ventral hindwings of V, cardui. Image © 2016 Connahs et al.21
This led on to further investigation of plasticity in nymphalid butterflies, where Junonia coenia and Bicyclus anynana have also been observed to exhibit varying eyespot sizes with different temperatures. Preliminary comparative work has identified plasticity in nymphalid butterflies to have multiple, independent origins22 .
Spot the differences
How many different points can you pick out from the two species of butterflies?
So how do we differentiate the two species?
If it is too difficult to take note of the details of the differences between the two species, just remember that V. cardui is everywhere except the Southern region of Australia. The next time you see a butterfly with these colourful wings on your trip to Tasmania, you know it is V. kershawi!
IV. Taxonomy and systematics
The Nymphalidae family is the family with the largest number of butterfly species.
Vanessa cardui belongs to a subgenus known as Cynthia, commonly known as painted ladies, together with V. kershawi, V. virginiensis and V. annabella. The species concepts for Lepidoptera appears to be significantly skewed towards the Biological Species Concept23 , where reproductive isolation can be easily tested by interbreeding studies and the past two decades of phylogenetic analyses have incorporated the time of divergence of clades in Lepidoptera.
Nomenclature
Scientific name: Vanessa cardui
Homotypic synonym: Cynthia cardui (Linnaeus 1758)
Common name: Painted lady butterfly, Thistle butterfly, Cosmopolitan
The genus Vanessa means butterfly in Greek. The origin of cardui is not known.
Original description
Original description of Papilio (Nymphalis) cardui by Linnaeus (1758). Image by Biodiversity Heritage Library24
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First described as Papilio (Nymphalis) cardui by Carl Linnaeus in 1758 25
, Papilio cardui was revised by William D. Field (1971) to the genus Cynthia, along with other Vanessa species26
. Following the extensive revision by Field, Wahlberg et al. (2005) resynonymised the genera Bassaria and Cynthia with Vanessa based on maximum parsimony analysis of molecular and morphological characters, leading to the current-day binomial name of Vanessa cardui27
, Despite the changes at the genus level, the species level taxonomy was stable.
Type information
The specimen described by Linnaeus in 1758 and designated in 176428
did not survive and another specimen was by Field (1971) and chosen as lectotype. The following is the type locality as described.
Type Locality: “Europae, Africae”; defined as “Sweden” by W. D. Field (1971), Smithsonian Contrib. Zool. (84): 43, based on Linnaeus’ reference to Fauna Svecica.
Today, the surviving specimen designated by Honey and Scoble (2001)29 is named a lectotype as well.
Images of lectotype in LINN 0299 Papilio cardui (Ins Linnaeus).Images by The Linnean Collections, UK (permission obtained).
Taxonavigation
Table showing classification of V. cardui 30 ,
| Kingdom | Animalia |
| Subkingdom | Bilateria |
| Infrakingdom | Protostomia |
| Superphylum | Ecdysozoa |
| Phylum | Arthropoda |
| Subphylum | Hexapoda |
| Class | Insecta |
| Subclass | Pterygota – winged insects |
| Infraclass | Neoptera – modern, wing-folding insects |
| Superorder | Holometabola |
| Order | Lepidoptera – butterflies, moths |
| Superfamily | Papilionoidea Latreille, 1802 – butterflies, papillons |
| Family | Nymphalidae Rafinesque, 1815 – Brushfooted Butterflies |
| Subfamily | Nymphalinae Rafinesque, 1815 – True Brushfoots |
| Tribe | Nymphalini Rafinesque, 1815 |
| Genus | Vanessa (Fabricius, 1807) |
| Species | Vanessa cardui (Linnaeus, 1758) – Painted Lady |
Phylogeny
Butterflies are one of the most well studied group of organisms, in particular the ecology and evolutionary processes of the largest family of butterflies, Nymphalidae. The following will highlight some of the attempts at resolving phylogenetic relationships within the Nymphalinae subfamily that V. cardui is grouped in.
The latest phylogenetic analysis of the subfamily Nymphalinae was conducted by Su et al. (2017).
Phylogenetic relationships obtained based on maximum likelihood and Bayesian analysis of specimens within the Nymphalinae subfamily. Position of V. cardui is demarcated by the red arrow31 .
A total of 353 specimens of 269 described species covering 55 of all 56 nymphaline genera was used to reconstruct the specimen-level tree. Sequences of three genes, EF1a, COI and wingless were concatenated and aligned using MUSCLE in MEGA5 software and checked manually. Time priors for Bayesian analysis was based on the fossils Vanessa amerindica and Prodryas persephone, from about 35 Ma, representing the divergence of Vanessa from its sister genus Hypanartia. In addition, secondary time priors using two host plant dates, Acanthaceae were used to estimate divergence time. For phylogenetic reconstruction, Bayesian, maximum likelihood (ML) and maximum parsimony (MP) were used. ML was conducted using raxmlGUI1.5b1 and bootstrap values were obtained using 1,000 pseudoreplicates. As for MP, tree-bisection reconnection (TBR) branch-swapping technique, together with heuristic search over 500 replicates gave bootstrap support values. The three analyses were mostly concordant with each other, especially well-establishing the relationship between sister genera Hypanartia and Vanessa.
This analysis built upon data analyses from the previous decades for the phylogenetic reconstruction of the butterfly subfamily Nymphalinae. Within the Nymphalidae family, the first support for the tribe Nymphalini as a monophyletic group was obtained using data from 161 species from 19 genera, DNA sequence data from three genes. namely the mitochondrial gene COI (Cytochrome Oxidase subunit I), and nuclear genes EF-1α (Elongation Factor 1 alpha) and wingless was obtained and combined, with gaps treated as fifth character states32 . Eight trees were obtained, from which the most parsimonious tree was produced using Partitioned Bremer Support analysis, which was preferred over bootstrap values for the variability between 1 and infinity in the former. A dispersal-vicariance analysis showed that dispersal gave rise to the divergence of Nymphalidae across three biotic regions. The study then hypothesised that the ancestral species of Vanessa had a relatively widespread distribution and speciation resulted during periods of isolation due to climatic or other factors.
Then in 2011, the phylogeny for the genus Vanessa was constructed by Wahlberg & Rubinoff33
, using one mitochondrial and eight nuclear protein coding genes. Bayesian topology was obtained with estimated divergence time from BEAST v1.5.2 analysis and the most parsimonious reconstruction was provided for the evolution of vagility in Vanessa species. A fossil Vanessa is known from the late Eocene, from ca 34 Ma34
, and has been used previously as a minimum age constraint for the split between the lineages leading to extant Vanessa and Hypanartia. Corresponding conclusion in this study used biogeographic data to reveal rapid expansions and inconsistent speciation within the genus Vanessa, where some lineages are constrained in speciation like V. cardui, but other taxa diverging into multiple species in different locality over time.
Comprehensive, dated, higher level phylogeny of butterflies inferred from genomic data, the history of phylogenetic analyses in the subfamily Nymphalinae exemplifies the increasing amount of resolution within larger clades. The most recent phylogenetic reconstruction by Su et al. (2017) conducted a concurrent analysis using DNA sequences, host plant divergence time and main evolutionary events after the Cretaceous–Paleogene extinction event 66 Ma. These analyses provided a well-encompassing estimates of possible basis for butterfly species divergence and global colonisation, namely warmer climates and angiosperm recovery.The original hypotheses of the time of divergence of Nymphalinae at 66 Ma (Wahlberg et al., 2005) was still supported in the most recent analysis (Su et al., 2017). Additionally, Su et al. (2017) showed that Vanessa diverged ca. 28.9 Ma and that Nymphalini colonised the New World in the Eocene from the Old World. This depicts the benefits of incorporating more data that is relevant for understanding the diversity of butterfly species observed today. Nonetheless, some of the groups remain unresolved, but the phylogenetic relationship of the vagrant V. cardui is well resolved over the past decades of phylogenetic analyses.
References
| Ref | Notes |
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| 1 | Ecuador, G. I. (1992). World distribution of the Vanessa cardui group (Nymphalidae). Journal of the Lepidopterists’ Society, 46(3), 235-238. |
| 2 | Hammad, S. M., & Raafat, A. M. (1972). biology of the painted lady butterfly, Vanessa (Pyrameis) cardui L. (Lepidoptera: Nymphalidae). Bulletin. |
| 3 | Janz, N. (2005). The relationship between habitat selection and preference for adult and larval food resources in the polyphagous butterfly Vanessa cardui (Lepidoptera: Nymphalidae). Journal of Insect Behavior, 18(6), 767-780. |
| 4 | Kristensen, N. P. (1998). Handbook of Zoology, Volume IV Arthropoda: Insecta. Part 35. Lepidoptera, Moths and Butterflies. Volume 2: Morphology, Physiology and Development. |
| 5 | "A Christmas Exclusive - Singapore's own Painted Lady," by Khew S.K. Butterflies of Singapore, 22 December 2007. URL: http://butterflycircle.blogspot.com/2007/12/christmas-exclusive-singapores-own.html (accessed on 4 Nov 2018). |
| 6 | Braby, M. F. (2000). Butterflies of Australia: their identification, biology and distribution. CSIRO publishing. |
| 7 | Jain, A., Khoon, K. S., Gan, C. W., & Webb, E. L. (2018). Butterfly extirpations, discoveries and rediscoveries in Singapore over 28 years. Raffles Bulletin of Zoology, 66, 217-257. |
| 8 | Stefanescu, C., Askew, R. R., Corbera, J., & Shaw, M. R. (2012). Parasitism and migration in southern Palaearctic populations of the painted lady butterfly, Vanessa cardui (Lepidoptera: Nymphalidae). European Journal of Entomology, 109(1). |
| 9 | Stefanescu, C., Páramo, F., Åkesson, S., Alarcón, M., Ávila, A., Brereton, T., ... & Hill, J. K. (2013). Multi‐generational long‐distance migration of insects: studying the painted lady butterfly in the Western Palaearctic. Ecography, 36(4), 474-486. |
| 10 | Suchan, T., Talavera, G., Sáez, L., Ronikier, M., & Vila, R. (2018). Pollen metabarcoding as a tool for tracking long‐distance insect migrations. Molecular ecology resources. |
| 11 | Jain, A., Khoon, K. S., Gan, C. W., & Webb, E. L. (2018). Butterfly extirpations, discoveries and rediscoveries in Singapore over 28 years. RAFFLES BULLETIN OF ZOOLOGY, 66, 217-257. |
| 12 | Ducatez, S., & Baguette, M. (2016). Inter‐individual variation in shivering behaviour in the migratory painted lady Vanessa cardui. Ecological entomology, 41(2), 131-137. |
| 13 | Chapman, J. W., Reynolds, D. R., & Wilson, K. (2015). Long‐range seasonal migration in insects: mechanisms, evolutionary drivers and ecological consequences. Ecology letters, 18(3), 287-302. |
| 14 | Nesbit, R. L., Hill, J. K., Woiwod, I. P., Sivell, D., Bensusan, K. J., & Chapman, J. W. (2009). Seasonally adaptive migratory headings mediated by a sun compass in the painted lady butterfly, Vanessa cardui. Animal Behaviour, 78(5), 1119-1125. |
| 15 | Abbasi, R., & Marcus, J. M. (2015). Color pattern evolution in Vanessa butterflies (Nymphalidae: Nymphalini): non‐eyespot characters. Evolution & development, 17(1), 63-81. |
| 16 | Otaki, J. M. (2007). Stress-induced color-pattern modifications and evolution of the Painted Lady butterflies Vanessa cardui and Vanessa kershawi. Zoological science, 24(8), 811-819. |
| 17 | Zhang, D., Zhang, W., Gu, J., Fan, T., Liu, Q., Su, H., & Zhu, S. (2015). Inspiration from butterfly and moth wing scales: Characterization, modeling, and fabrication. Progress in Materials Science, 68, 67-96. |
| 18 | Schiffman, J. D., & Schauer, C. L. (2009). Solid state characterization of α-chitin from Vanessa cardui Linnaeus wings. Materials Science and Engineering: C, 29(4), 1370-1374. |
| 19 | Gu, J., Zhang, W., Su, H., Fan, T., Zhu, S., Liu, Q., & Zhang, D. (2015). Morphology genetic materials templated from natural species. Advanced Materials, 27(3), 464-478. |
| 20 | Nijhout, H. F. (1984). Colour pattern modification by coldshock in Lepidoptera. Development, 81(1), 287-305. |
| 21 | Connahs, H., Rhen, T., & Simmons, R. B. (2016). Physiological perturbation reveals modularity of eyespot development in the painted lady butterfly, Vanessa cardui. PloS one, 11(8), e0161745. |
| 22 | Monteiro, A., Tong, X., Bear, A., Liew, S. F., Bhardwaj, S., Wasik, B. R., ... & Cao, H. (2015). Differential expression of ecdysone receptor leads to variation in phenotypic plasticity across serial homologs. PLoS genetics, 11(9), e1005529. |
| 23 | Wheeler, Q., & Meier, R. (Eds.). (2000). Species concepts and phylogenetic theory: a debate. Columbia University Press. |
| 24 | Caroli Linnaei...Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Image obtained from Biodiversity Heritage Library (BHL). Retrieved on 2 Nov 2018 from https://www.biodiversitylibrary.org/item/10277#page/497/mode/1up. |
| 25 | Linnæus, C. 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. - pp. 475, 1-824. Holmiæ. (Salvius). |
| 26 | Field, W. D. (1971). Butterflies of the genus Vanessa and of the resurrected genera Bassaris and Cynthia (Lepidoptera: Nymphalidae). |
| 27 | Wahlberg, N., Braby, M. F., Brower, A. V., de Jong, R., Lee, M. M., Nylin, S., ... & Zakharov, E. (2005). Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers. Proceedings of the Royal Society of London B: Biological Sciences, 272(1572), 1577-1586. |
| 28 | Linnaeus, C. V. von-1761. Fauna Svecica. Stockholmiae, editio altera:(48), 1-578. |
| 29 | Honey, M. R., & Scoble, M. J. (2001). Linnaeus's butterflies (Lepidoptera: Papilionoidea and Hesperioidea). Zoological Journal of the Linnean Society, 132(3), 277-399. |
| 30 |
Integrated Taxonomic Information System on-line database, URL: https://www.itis.gov/ (accessed on 2 Dec 2018). Global Biodiversity Information Facility. URL: https://www.gbif.org/ (accessed on 2 Dec 2018). |
| 31 | Su, C., Shi, Q., Sun, X., Ma, J., Li, C., Hao, J., & Yang, Q. (2017). Dated phylogeny and dispersal history of the butterfly subfamily Nymphalinae (Lepidoptera: Nymphalidae). Scientific reports, 7(1), 8799. |
| 32 | Wahlberg, N., Brower, A. V., & NYLIN, S. (2005). Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society, 86(2), 227-251. |
| 33 | Wahlberg, N., & Rubinoff, D. (2011). Vagility across Vanessa (Lepidoptera: Nymphalidae): mobility in butterfly species does not inhibit the formation and persistence of isolated sister taxa. Systematic Entomology, 36(2), 362-370 |
| 34 | Miller, J. Y., & Brown, F. M. (1989). A new Oligocene fossil butterfly, Vanessa amerindica (Lepidoptera: Nymphalidae), from the Florissant formation, Colorado. Bulletin of the Allyn Museum (USA). |
This page is authored by Loh Ling Sheng
Last curated on 2 Dec 2018
