An Introduction to the Longwing (Heliconid) Butterflies By Nigel Venters

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2 By Nigel Venters (This paper was first issued as part of the Painted Lady and Longwing Butterflies workshop.) Introduction As I mentioned in Article (and think is worth repeating here) - it may be considered as strange to combine two very different families of butterflies into a single workshop. The Painted Ladies (Vanessa species from the Nymphalidae family) and the Longwings (Heliconid species from the Heliconiinae family) are far apart scientifically. I am well aware that many taxonomists would groan in disbelief at my combining these two families into the same workshop! However, my aim is to show how to breed butterflies from both these families in captivity in the most efficient way, minimizing the work, equipment and space required. From a butterfly breeder s point of view, these butterflies share many similar characteristics and raising methods! Of course the host plants between these two butterfly families are very different, but they also vary considerably within each family. I will cover these host plants in some depth later in this paper. As I mentioned in Article 5.3.1, Monarch breeders need not worry about the parasite Ophryocystis elektroscirrha (such an awful name and best known as O.e.) as it does not occur in any of these species, being entirely restricted to the Danidae family of butterflies. So relax, if you breed these species you will be totally free from O.e.! The Zebra Longwing (Heliconius charatonia) shown on the left, and the Gulf Fritillary (Agraulis vanillae) shown on the right, are probably the most commonly bred butterflies from the family. In the wild they inhabit different biotypes, but can sometimes be seen flying together at the edges of forests or dense woodland. 1 By Nigel Venters

3 This section of the workshop is structured to first give the student a thorough understanding of the species you are expected to breed. Article covers the Painted Lady/Red Admiral Butterflies (the Vanessas) and this article presents the Longwing (Heliconid) species. Article will go into depth, teaching the most efficient methods to successfully breed large numbers of all the Painted Ladies/Red Admirals from around the world, as well as all of the Longwing species (which are restricted to the Americas (North, Central and South America). Detailed techniques are provided to easily breed butterflies from both these families by using optimal conditions and the best host plants. I. Distinguishing Between the Different Longwing Species Many species of Longwing butterflies use Mullerian mimicry to avoid being eaten by predators. Mullerian mimicry (named after its discoverer, Frans Muller) is when a number of different distasteful species gain protection by all looking similar to each other. The Longwings above are all different species! The Longwing Species (Heliconiinae) - A Brief Overview The Longwing species are extremely variable, and within this family the Longwings can be broken into two main groups - from a butterfly breeder s point of view! The Longwings Group One These are the faster flying, and mostly orange coloured Longwings. Most of this group generally have wider wings than those from the Longwings Group Two. An essential difference is that none of the Group One Longwings can process pollen to feed on. This makes them a shorter-lived butterfly than those in the Group Two Longwings that can live as adults for up to nine months in optimum conditions. 2 By Nigel Venters

4 The lifespan also affects the egg laying sequence and timing. The Group One Longwings live shorter lives and the total egg laying period is much quicker within this group, than in the Group Two Longwings. This is discussed in some detail in the next article. One characteristic that all Longwing butterflies share is that in captivity (and to some extent, in the wild) they tend to lay many eggs off their host plant. This can be a problem for the breeder and solutions to this challenge are discussed in the next article in this workshop. Some of the Group One Longwings are strong migrants, and can be found well away from their usual haunts. A good example is the Gulf Fritillary (Agraulis vanillae), which occurs along forest edges but is also quite at home in much drier conditions in open country. Although they are quite powerful fliers, they do not have the same vigour and strength as the Painted Ladies and Red Admirals. All of the Longwings (Heliconids) feed on host plants from the Passion Vine plant family (Passiflora). However, all of the Longwings in Group One are generalist feeders, feeding on many different host plants in the Passion Vine family. (Host plants discussed in detail later). Interestingly, the Passion Vine plants which are almost always climbers (a few exceptions of course) produce toxins in their leaves. In a similar way to the well-known Monarch butterfly, this deters predators from eating the Longwings. It is also strange that while all the leaves and stems of these plants are toxic, the fruits are not! Many fruits from this family are commercially produced for the food market, and perhaps the best known is Passiflora edulis The Passion Fruit. It is worth pointing out that any plant with a scientific name that is followed with a specific name of Edulis means that it is edible! The Longwings Group Two The second group within this Longwing butterfly family, is the true Longwings. They really do have longer and narrower wings than those species in Group One. They are also essentially forest/woodland dwellers, and flutterers. They always appear to be weakly flying through the forest edges and open areas within the forest where (for example) a tree has fallen down to expose a sunny glade, or a cleared path through the forest. Actually, it is surprising how quickly they will turn on a burst of speed to escape from a swinging butterfly net! It is rare to find these species very far away from the forest or forest edges, flying in more open, arid areas. Although Group Two is in the same family of butterflies as Group One, they display some interesting differences in their behaviour, lifespan and egg laying routines. Perhaps the most different aspect between the two groups is that the Longwings in Group Two collect pollen as well as nectar from plants to feed upon. The Group Two butterflies are the only butterflies in the World that can collect and process pollen into a highly nutritious food source! In the wild they are heavily dependent on tropical climbing plants from the cucumber, melon, and pumpkin family, (Cucurbitaceae) 3 By Nigel Venters

5 which are known as Psiguria species. These flowers are rich in pollen and nectar, but it is the pollen they concentrate on! The pollen is collected and held as a small ball of pollen at the base of the tongue, and at its leisure, the butterfly can then release a unique enzyme from its tongue to dissolve the pollen into a very high protein liquid. This liquid is sipped in the usual way back through the tongue and into its digestive system. It is not vital to have Psiguria plants to breed this species of course, and these Longwings can collect pollen from many different sources to ensure they can live a long life as an adult. In this brief overview (expanded in detail later) there is one other aspect that needs to be mentioned, and this is the egg laying by the females in this group. As mentioned above, the females of Group One will freely lay their eggs over a fairly short time span, but the butterflies in Group Two are quite different! The females in this group live long lives, and often lay just a few eggs per day over a long period of time. In captivity, this can be speeded up (covered later). Since the butterflies are so rapid in growth and lifecycle in the larval stage, many butterfly display houses restrict breeding to the Group Two Longwings, which lay eggs slowly over a long period. This prevents a complete decimation of their host plants within their display, by those species that lay their eggs over a short period as seen in Group One. However, regardless of belonging to Group One or Group Two, the adults only pair once. Once paired, the females then produce a scent (pheromone) from their abdomens, which deters and prevents other adventurous males from trying to pair with them. Many of the different species of Group Two Longwings look very similar to each other, and use Mullerian mimicry for general protection from their predators. Mullerian mimicry is basically where a number of different distasteful butterfly species look very similar, and are all unpalatable to predators. The slow unconcerned flight helps their potential predators recognise and avoid them! See the photo on page two. Interestingly, the Longwings in Group One are also distasteful, generally displaying an orange wing colour warning. Their more open biotypes coincide with many other similar but different species of butterflies which also display the same general wing colour, but are in fact quite palatable! Therefore, it is more practical for the Longwing species in Group One to be much faster fliers to avoid mistakes by potential predators. A Brief Overview of Host Plants When is comes to host plants, the Longwings in Group Two are split into two separate groups! Later, I will describe the host plants for each species. However, an easy way to separate these species is by calling them Group Two-A and Group Two-B. Group Two-A consists of generalist feeders that feed on a wide range of different Passion Vine host plants. Group Two-B is a smaller group made up of very specialised feeders that feed only on an extremely limited number of Passion Vine host-plants, refusing all others. (For Central and South American species, there is also the further complication of some 4 By Nigel Venters

6 species females laying eggs singly or in egg batches. workshop articles.) This is covered later in these A General Note I hope this overview of Longwings does not appear too complicated, and that you will not be discouraged from breeding them. They are in fact, all easy species to breed once you have the set up organised correctly. The remainder of this article is written to provide you with an easy identification guide between the different species, some regional distribution information, and host plant preferences, etc. The third article in this workshop will show how to easily breed them with the minimum effort and maximum results! II. The North American Longwing Species Identification Guide If you live in North America, there are only six species you need to distinguish. Four of these species belong to Group One (and two are extremely rare migrants). These are the more open-country species that are not restricted to forest areas. They are also generalist feeders and are not too fussy on which Passion Vine that they feed on. The remaining two of these species belong to Group Two-A. These two species require a forest biotype. One of these, the Crimson Patched Longwing, (Heliconius erato) is restricted to the far south of Texas. Both of the US Longwing species in this group are generalist feeders and feed on a huge range of different Passion Vine plants. There are no species in North America that fall into Group Two-B. This group is restricted to Central and South America, and are forest species with a very restricted range of Passion Vine host plants. The North American Longwings, Group One The Gulf Fritillary (Agraulis vanillae) Gulf Fritillaries. The male (on the left) and female (right) look very similar, but the male is always a brighter orange with less heavy black markings on the wings. The undersides are identical. 5 By Nigel Venters

7 The Gulf Fritillary is probably the best known, most widely distributed, and most often captive bred species of all the Longwing species in the US. This is due to the fact that it occurs quite a good way north in the US, compared to the other Longwings. This species has a huge range throughout South America, Central America, The Caribbean Islands and into the US northwards. In the US it is found from San Francisco south to California and beyond, resident throughout the Southern States, and it is a great migrant that travels north into the Great Basin, The Rockies and Midwest, also the Great Lakes area and mid-atlantic states. The further north this butterfly is found, the more sporadic its occurrence. Depending on the year, it can appear commonly in some years and absent in other years. Of course in the northern part of this range, it is a temporary migrant, and cannot survive the cold winters there. However, having said that, this species is the most cold resistant of all the Longwing species, and, as both an adult and as a larvae, it can tolerate and survive temperatures close to freezing for short periods. The larvae just stop feeding when it is cold, and the adults tuck themselves away into the vegetation waiting for the temperatures to rise. As soon as it warms up, they become active again. The Gulf Fritillary is so distinctive with its bright silver under-wing spots, you d think it would be impossible to confuse it with any other species of Longwing. However, if you live in the south part of Texas, you may also see the much rarer Mexican Silverspot (Dione moneta). The photo below (under the heading Mexican Silverspot) shows how to distinguish between these two species that look almost identical to each other! It may seem strange for two so similar looking butterflies as the Gulf Fritillary and the Mexican Silverspot to be able to inhabit the same biotype, with almost identical requirements for survival. In Argentina where I live, both species are equally common in my garden throughout the year. They are fairly hard to distinguish from each other while flying, but the Mexican Silverspot flies slower than the Gulf Fritillary. They also (generally) choose different Passion Vine host plants to lay their eggs on. The host plants for the Gulf Fritillary are the many different Passion Vine species, and I have not known a single Passion Vine species it would not accept as a larvae. However, the best host plant you can choose is the Blue Passion Vine (Passiflora caerulea). The Blue Passion Vine has several advantages for breeding the Longwing butterflies over the all the other Passion Vine species you may find locally available. The further north you live in the US, it is an especially hardy plant. More detail on this in Article Three. 6 By Nigel Venters

8 The Mexican Silverspot (Dione moneta) The Gulf Fritillary (Agraulis vanillae) is almost identical to The Mexican Silverspot (Dione moneta). There are always more black spots in the Gulf Fritillary s forewing, but the best way to be sure, is to look for silver spots in the forewing (see red arrows in photo). The top photo with silver spots in the forewing is the Gulf Fritillary, and the bottom photo without silver spots is the Mexican Silverspot! Much of the information required to distinguish between this species has been provided in the photo above. You can see from this photo just how similar these two species are and how to be certain of what you have caught. The Mexican Silverspot is less tolerant of cold conditions than the Gulf Fritillary. However, I have found large groups of adults here in Argentina, sitting out the cooler winter conditions. They gather together, similar the Monarchs in their overwintering grounds in Mexico, but in far smaller total numbers, of course. The groups stay together for a few days and then move out to feed on warmer days and continue their lifecycle. 7 By Nigel Venters

9 As previously mentioned, unless you live in the southern part of Texas, you are unlikely to see the Mexican Silverspot butterfly. Even there, it relies on migration from the south, and can be reasonably common in some years and absent in others. They have a very beautiful pupa, which of course has a similar shape to all the other Longwing butterflies. However, the pupa has some bright, metallic silver spots on the sides, not so very different to the silver spots on the adult s hind-wing. The main host plant is Passiflora morifolia (Mulberry Leaved Passion Vine) but it also uses P. andenopoda, P. capsularis, P. edulis, P.mollissima and several other species. The larvae will eat Passiflora caerulea in captivity, but in my experience, the adults do not usually choose this Passion Vine as a natural host plant in the wild. The Julia (Dryas Julia) The Julia (Dryas julia) in flight, may look a little like the Gulf Fritillary and Mexican Silverspot at first glance. However, it has much more elongate wings than either of the other two species and is much more orange with minimal black markings on the forewings. The main difference is the complete absence of silver spots on the underside of the wings. The female is duller than the male. Both sexes are distinctive, bright butterflies that rely on protection from predators by using toxins obtained from the larval host plants. 8 By Nigel Venters

10 A very beautiful sub-species of this butterfly occurs in the Ecuadorian Amazon, where the bright orange colour in the top wings is replaced by the most stunning crimson red, that almost shines as it flies! I am not entirely sure if this is a true sub-species or if this striking colour is climate induced. It may be of interest to breeders of this species to raise the temperature and humidity levels to see if this is the case. Humidity in this case needs to be close to 100%! The Julia is a very common butterfly in southern Florida and southern Texas. However, it is hard to catch with a net as it is a rapid flier! This is also a species with a far less cold tolerance than the Gulf Fritillary, and it does not often migrate or wander north from its favoured habitats. A few vagrants do wander north occasionally, but of course they can never establish themselves in the colder climates they experience. Once you venture south, from the southern US, this butterfly is extremely common everywhere as far south as Argentina. It is common at forest edges, and sunny glades within the forest, but also quite happy in gardens and open countryside within its range. The host plant range for the Julia is huge! It is a generalist Passion Vine feeder and uses a staggering number of different Passion Vine species as it host plants. Again, I have to say, that I have never had this species refuse any Passion Vine I offered it to lay on, and the resulting larvae have always completed their development. The Striped Longwing (Draydula phaetusa) The Striped Longwing (Draydula phaetusa). This is quite a distinctive species and if you do find it, you are unlikely to confuse it with any other Longwing species. I had been uncertain if this butterfly ever reached the southern US, but given its ability to migrate and turn up in places far from its normal range, I considered it likely to do so. Recently, I was interested to learn from Steve O Neil, who lives in southern Texas, that my assumption was correct. Steve had caught and bred this species there from the migrants that wander up from regions south of him. 9 By Nigel Venters

11 You are unlikely to confuse this butterfly with any other Longwing species, after seeing the photos. The female is always a little duller than the male, which can often be a bright orange colour. The wing marking for both sexes are identical, however the male does tend to have a less rounded, Longwing shape rather than the more rounded wings of the female. Although the Striped Longwing frequents forest edges and clearings, it is often seen as a more open-country species. It has much rounder wings than most other Longwings, and the two dark coloured stripes on its hind-wings make this an easy species to identify. The host plant range is again huge, and similar to all of the more open-country Longwing species. As for all this family, the host plants are all Passion Vines (Passiflora species). III. The North American Longwings, Group Two-A We now move on to the Heliconid Longwing group, and as I mentioned earlier in this paper, I have separated this group of Longwings (from a butterfly breeders point of view) from the more open-country, ubiquitous species described above, into a second group that really do rely on a forest or heavy woodland environment to do well. These species do have the occasional wanderer into more open habitat, but will never breed and develop a separate colony in these conditions. They can be most often found as forest edge species, but also wander through closed forest, and frequent paths, clearings, or an area where a large tree has fallen to allow some sunlight into the forest. I will cover their raising requirements in the next article, but mention here that to be successfully bred, they will never do well in hot, dry conditions, and the eggs need to be protected from direct sunshine as they desiccate very quickly in these conditions. The Zebra Longwing, (Heliconius charitonius) The Zebra Long-wing (Heliconius charitonius, but note: some literature uses the scientific name Heliconius charithonia). It is unmistakable, and is quite different to almost all of the other Longwings in this family. It is the most widely distributed Longwing species from Group Two-A, and has a huge range through Central and South America. 10 By Nigel Venters

12 The Zebra Longwing is unlikely to be confused with any other Long-wing species in the US, as it is so distinctive. Further South, in Central and South America, there are other species that do not look so dissimilar, but even here it is quite simple to separate from other Longwing species. It has the most lovely, slow, fluttering flight, and initially seems totally unconcerned when approached. However, don t be fooled, when needed, they can put on an impressive turn of speed to avoid the butterfly net! The Zebra Longwing is a resident from Texas and Florida in the South to South Carolina. In some years in suitable climatic conditions, it wanders north to the Great Basin, Colorado and the Great Plains. It also occurs in the Caribbean, and far to the south of the US. Attached at the end of this article you will find text and photos kindly provided by Dale Clark, President of the Dallas County Lepidopterists Society to help identify the two Zebra Longwing subspecies, Heliconius c. vazquezae (Texas), and Heliconius c. Tuckeri (Florida). Essentially, it is a forest edge and clearings butterfly, and to be sure of finding it, you really need to search these ideal environments. Although a forest species, it does have the ability to migrate or wander to areas far to the north of its expected range. However, it is never truly happy unless it finds a suitable forest/woodland environment. Of course, the winter will kill off all the northern colonies, established temporarily by this wandering activity. The host plant range is again huge, and they are all Passion Vines (Passiflora species) as they are the only family accepted as host plants by the Longwing butterflies. I have never found a Passion Vine species that this species would not accept in captivity, however, I personally use Passiflora caerulea to breed this species. The Crimson Patched Longwing (Heliconius erato) The Crimson Patched Longwing (Heliconius erato) is perhaps the most beautiful of all the North American Longwing species. Unfortunately, unless you live in southern Texas, you are unlikely to ever see it! It becomes very common as you venture south into Central and South America. 11 By Nigel Venters

13 The Crimson Patched Longwing is the last species described here that flies in North America. However, it is really a fully tropical species, and is only found in southern Texas in the US. It is unmistakable you may think - however, and although I can find no reliable US records for the almost identical species, Heliconius melpomene in the US. These two species can be almost impossible to separate. Heliconius melpomene occurs just to the south of the US, and it would not be a great surprise to me that this separate species may also wander into southern Texas. (To separate these two species see the next section on Heliconius melpomene.) The Crimson Patched Longwing is a true forest species, and can be found wandering through closed forest with its slow, fluttering flight pattern. However, it is mostly found on forest edges and sunny paths and glades within the forest, in very similar conditions to the Zebra Longwing described above. Like the Zebra Longwing, this species also has a slow fluttering flight, which can rapidly change when pursued by a collector with a butterfly net! This species is also a generalist when it comes to choosing it host plants for egg laying. Of course, all the host plants are Passion Vines. There is an absolutely huge list of recorded Passion Vine host plants recorded for this species. The list is so long it is hardly worth mentioning here. Basically, if it is a Passiflora species, then the Crimson Patched Longwing will use it as a host plant. Again, I use the Blue Passion Vine (Passiflora caerulea) when I am breeding this species. Host Plants for the Longwings You will have seen from my notes above, that I have continually referred to the Blue Passion Vine (Passiflora caerulea) as the best host plant to use for all the US species. There are a number of reasons why I use this plant. First, all the US Longwing butterflies will use this as a host plant. For some species, it may not be their first option in the wild, but for breeding my Longwings in captivity, it is the best choice for several reasons. By using this host plant, I don t have to keep too many other species of host plants available to use. And in the third article for this workshop (breeding), you will see that there are many other reasons for using it that really benefit the butterfly breeder. Eggs and Egg Laying All of the North American species of Longwings lay their eggs singly. However, once you venture south into Central and South America, this can change and some species lay a huge batch of eggs together, while others will lay only a very few eggs singly each day. I will go into some detail on this, and of course the detailed breeding notes, in Article Three. IV. Central and South America - The Group Two-A Longwings Further south from the US, in Central and South America, there is a minefield of different Longwing species you will need to be able distinguish, to be successful in your breeding. All 12 By Nigel Venters

14 the more open country species described for North America in this article also occur to the south, and it seems unnecessary to repeat this information here. Please just refer back to this article to obtain the information you need for these different Longwing species. As I mentioned, once you leave the US and go south to Central and South America, things become a lot more complicated. This is due to the huge number of different Longwing species that occur there, and the variation in both colour and wing pattern for some species. The only problem from a butterfly breeder s point of view, is to ensure a correct identification of each separate species, so that you will be sure to use the correct host plant. Once you have established this, breeding these species is as simple as all the US species once you have provided the optimum breeding conditions. For the Central and South American Longwings, I have to use scientific names in this article, as local names vary from country to country. A quick search on the Internet using the species name will give you the local common name for all of the species, depending on where you live! Heliconius melpomene I think it best to start this section on Central and South American Longwings with a photo of a single species, Heliconius melpomene, as an example of the incredible wing pattern and colour form variation! In some regions no two butterflies of this species are the same! It is hard to believe that all of these variations are in fact a single butterfly species! This graphic will give you an understanding of the problem I now face with describing the Central and South American Longwing butterflies! However, I will do my best and I think I can separate them all for you! 13 By Nigel Venters

15 Ok! Now to get this into perspective! Heliconius melpomene is is the most difficult of all the Longwings to identify. It has a wide distribution, ranging through Central America, South America and the Caribbean Islands. Heliconius melpomene appears to be heavily involved in a huge Mullerian mimicry ring, so the forms vary, depending on the location. Almost no two specimens look identical. But, regardless of where you see this butterfly, the colour forms tend to be dominated by a selection of wing pattern and colour forms that mimic the commonest other local Longwing, (Heliconid) species. The range of specimens shown in the photo above comes from all over this butterflies distribution range, so don t expect to see these different forms in the same location where you live. This butterfly is a great favourite with the European Butterfly display houses. When released into the exhibit, to the general public and visitors, it looks as though there are many different species of slow-flying and beautiful tropical butterflies present. A Missed Opportunity? It is worth mentioning here that many butterfly display houses import Heliconius melpomene from a single source, (country) from tropical Americas. This seems such a shame, when by locating two different sources of this species, which are widely separated, and then releasing them into the enclosure, such a wonderful display can be created. By releasing two separate races of this butterfly into a butterfly display house, the resulting offspring are truly amazing! The genetic interchange goes wild! And apart from no two butterflies looking alike, the range of colours and wing patterns are truly bizarre! Before I continue with this species description and hostplant requirements, I thought it worthwhile to show you a second photo of Heliconius melpomene, flying in a butterfly display house from the UK. (These are from Butterfly World UK, and many thanks to Gareth Welsh for these photos!) If you look closely at the photos below, you will see that none of them match perfectly any of the H. melpomene shown in the first photo (above). However, I want to draw your attention to the species shown on the far right, which is the Crimson Patched Longwing, (Heliconius erato) described earlier under the US Longwing species section. It looks so different here when shown against H. Melpomone. But, look again at the selection of H. melpomene photos shown above! You will see that the photos shown bottom right and bottom left of H. melpomene are identical to the Crimson Patched Longwing, (Heliconius erato). So the following photo, and description of the differences, will help you be certain of what species you have caught and are breeding. The red spots are entirely restricted to these two species and it will be simple for you to separate them using this guide. 14 By Nigel Venters

16 Heliconius melpomene, a favourite species with butterfly display houses. This species is so variable in wing markings and colouring, throughout its wide range in central and South America and Caribbean Islands. If you look at the photo on the far right, you ll see a specimen of H. erato, a totally different species. I have only included this photo, because H. melpomene, also has a form that is identical to this. Only by looking for some tiny red spots near the main forewing vein (close to the junction with the abdomen) in the forewing underside, can you be sure to separate these two species. H. melpomene has THREE small red spots and H. erato has FOUR small red spots! Additional Notes on Heliconius melpomene Remember there are many other Central and South American Longwing species that look identical to many forms of Heliconius melpomene. However, none of these other species have the red spots described under the photo on the previous page! To see how close some of these different species are, look back at the photo shown on page two of this article. Heliconius melpomene is a forest species, and the females are often seen struggling through the dark enclosed forest looking for host plants to lay their eggs, but most often at the sunny forest edges, and paths, and openings occurring within the forest. It does not stray far into the drier, more open areas that surround the forests. It is a slow flying, fluttering species like so many of the other Longwings in Group Two-A. Heliconius melpomene s host plants are of course Passion Vines (Passiflora species) and it is not surprising that given the huge range of this species, that also a vast range of Passion Vines are used as host plants. Once again, I always use Passiflora caerulea to breed this species. 15 By Nigel Venters

17 V. The Other Central and South American Longwing Species It is probably unnecessary for me to start this section with a separate title, and it is only because I have spent so much time describing the complexities involved in identifying Heliconius melpomene that I thought it was now worthwhile to focus our minds on the other Longwing species! I don t think it is possible within the scope of this article to cover all of the other Longwings in detail, as there are so many different species involved. However, I will now show you how to separate all the other main Longwing species that you are likely to encounter in Central and South America, and supply their host plant information for butterfly breeders. I will continue this next section with general colour and wing patterns of some well-known and widely distributed species, and will then follow up with photos of most of the other known species for you to compare them with. Host plant details so far in this paper have been easy to provide, as all species of Longwings I have described have been generalist feeders. I will of course continue to supply a host plant list for each species, but it does get more complicated when we reach the Longwing species that I consider as falling into Group Two-B, the specialist feeders. Of course, as usual, all these species will only accept Passion Vine (Passiflora) species as host plant for their larvae to feed on. Fortunately, apart from Heliconius melpomene, most of the other Central and South American Longwing butterflies are relatively stable in wing pattern and colouring. This is not to say there is no variation over huge distribution areas, but compared to Heliconius melpomene, these are not so very difficult to separate into their own distinct species. Some of the species described below will have two colour forms, a dominant form, and less common form that may be quite different in colour, but the wing pattern stays relatively stable. Where this occurs I will describe this variation, show photos of similar species, and will give an easy identification guide to distinguish them from each other. Now we move on the individual Longwing species, that you are likely to encounter in this region. 16 By Nigel Venters

18 Heliconius cydno Heliconius cydno is found from Mexico to Ecuador, and has several subspecies. It is quite distinctive in pattern and colour, but can be confused with other species. An explanation of how to separate these species follows. Heliconius cydno is a widely distributed species, ranging from Mexico down to Ecuador and Columbia. It does not seem to be able to survive much further south than this, although the climate seems suitable to my mind. Perhaps it cannot compete with other similar species, that are found in this region. It is part of a complex of blue/black coloured Longwing species, with patches or stripes of either white or yellow in the forewings. This is not a rare species. It inhabits thick forest and even secondary forest growth, but never, in my experience, seems very common. Most specimens I have seen are solitary individuals, struggling through or along forest paths. However, at night they seem to be able to locate others of the same species and roost communally in groups of up to 50 or more. I came across once such gathering in the rain forest in Costa Rica, late in the evening when I was out looking for moths. Communal roosting at night is a common practice amongst the Longwings from this group, and gatherings of the same species at night can be quite large. I once found more than 100 Heliconius erato gathered together early one morning, and as the sun warmed up it was similar to a snow shower as they all took off together. Perhaps the most dramatic way to see this is to disturb the cluster of butterflies by gently tapping the tree or plant that they have roosted on, and an immediate response is assured! One other strange factor that indicates that these butterflies are quite aware of their surroundings, is that they will gather each night in the same place. 17 By Nigel Venters

19 The host plants within the Passion Vine family that this species uses is so vast, that it is considered as a generalist feeder. Again, I have found that the Blue Passion Flower, (Passiflora caerulea) is effective in captive breeding. Heliconius doris Heliconius doris is easily distinguished from H. cydno, as it always has two white or pale yellow bands across the forewings. (Compare with previous photo of H, cydno.) Heliconius doris is found from Mexico to the Amazon basin, just like H. cydno, although the biotype seems suitable it does not occur to the south of this area. Note: Heliconius doris does have a red form, which does not look so very different from H. Melpomene. However, there are no red dots on the underside forewing, near the main vein in H. doris. This is a fairly common species that has different approach to reproduction than most Longwings, as it lay its eggs in huge batches. It has a far more restricted host plant range than many other Longwings, and in the wild it favours some of the more uncommon Passion Vine species, such as Passiflora oerstedii. However, once again, I found no trouble with breeding this species on Passiflora caerulea. 18 By Nigel Venters

20 Now compare H. cydno and H. doris, to the other species you may confuse it with. Heliconius sapho H. sapho is similar to H. cydno, but the forewing always has a white spike in its wing pattern as shown above. The white patch in the forewing of H. cydno always reaches the terminal forewing edge, and is never broken - it is always a white patch. If in doubt, a good way to separate H. cydno and H. sapho when collecting live specimens is to smell the abdomen, as H. sapho (both sexes) always emits a strong, sweet smell, which is absent in H. cydno. Heliconius sapho is found from Mexico to Costa Rica, but has a very restricted range of host plants, feeding on only Passiflora macrophylla, and Passiflora pittieri. The female lays large egg clusters. Heliconius sara H. sara is similar to H. doris, but is always a smaller species. The yellow (or sometimes white) forewing bar that is nearest the body (the base) never reaches the strong forewing vein (the costa) at the top edge of the forewing. Heliconius sara is found from Mexico through to the Amazon basin. It uses a wide range of Passion Vine host plants but perhaps the most often used is Passiflora auriculata. This is one of the few Longwings that will not accept P. caerulea, however, having said that, it will take P. edulis, which is the main species used in the production of Passion Fruit that we all see in the supermarkets. This is an easy way to obtain seeds, and also eat a delicious fruit! The female lays her eggs in clusters. Heliconius wallacei H. wallacei is also similar in colour and marking to H. doris. However H. wallacei is a more southern species, which ranges from the southern Amazon basin down to Bolivia and possibly northern Argentina. It is generally darker, with less of a blue sheen than H. doris, and always has a large, more rounded yellow patch in the forewing. Helliconius wallacei has a rather restricted range of Passiflora host plants and uses P. coccinea, P. glandulosa, P. quadriglandulosa, P. variolata, and P. vitifolia in the wild. The female lays her eggs in clusters. I have not bred this species, but I would certainly try P. caerulea in captivity. 19 By Nigel Venters

21 Heliconius hecale Heliconius hecale is a easy species to identify, and is large! This has to be the largest of all the Longwings in this group, and is much bigger than a Gulf Fritillary (Agraulis vanillae) for example. However, as usual there are some other Longwing species that look relatively similar! H. hecale is a very distinct species. It is larger than most of the other forest Longwings, and occurs over a huge range of Central and South America. If you live south of the US, you are sure to find it there in suitable forest localities, from Mexico down to Peru. This species may also be found in areas adjacent to the forest and is not quite so restricted to closed forest, as are most of the other Longwings in this group. It is not unusual to find it nectaring in open pastures. This butterfly loves to feed and collect pollen at the base of its tongue, from Psiguria and Gurania flowers. Unusually for the Longwings, it is quite aggressive to other butterflies that try to feed from the same flower source, and will actively chase them away! This butterfly has a large number of subspecies over its vast range, and can be quite variable. The photo shows this species from Costa Rica. However, it is important that I show you some of the variations of this species you may encounter. It is also important to mention that the dominant form is shown in the photos above, and the photos below show other forms which are less commonly encountered. The eggs are laid singly, on a wide range of Passion-vines, and as usual the Blue Passionvine (Passiflora caerulea) is a hostplant. 20 By Nigel Venters

22 H. hecale and some colour variations within its huge range south of the US This grapic shows the variation that may occur when you find Heliconius hecale. I kept the images small to show just a pictorial view of the whole variation you may encounter within this species range. Now compare H. hecale with other species you may confuse it with This graphic shows two separate species of Longwings that you may confuse with H. hecale. The photo on the far right is Lycorea cleobaea, which is in the same family as the Monarch (Danaidae) and of course not a Longwing species! Heliconius ismenius This species is always smaller than H. hecale and has a narrower wing shape. It is far more restricted to the forest edges and light gaps caused by fallen trees or paths cut through the forest. It is uncommon to find them wandering far from their favoured environment of primary rainforest. H. ismenius occurs from Mexico south to Ecuador, and feeds on a wide variety of Passion Vines as host plants. In the wild, these are mainly Passiflora alata, P. pedata, P.ambigua and P.platyloba. However, once again, like so many other Longwing species, it feeds on the Blue Passion Vine (Passiflora caerulea) in captivity. Heliconius numata This is an even smaller species than H. ismenius, and is again, a closed primary rainforest species. This species has a much more limited range and can be seen from Columbia southwards to Brazil. H. numata feeds on a wide variety of Passion Vine species throughout its range, and notably one of them is P. edulis, which is the main species used in the 21 By Nigel Venters

23 production of Passion Fruit that we all see in the supermarkets. I have no records of it feeding on Passiflora caerulea, however, I would expect this highly likely as a host plant in captivity. VI. Other Species of Longwings in Central and South America Green is a rare colour for butterflies to have on the upper wings, and here are two examples which are exceptions. On the left is the Longwing Philaethria dido, which is closely mimicked by the Malachite (Siproeta stelenes), which is a Nymphalid. Just look for the tails on S. stelenes to decide which species you have caught! Philaethria dido This species is quite unmistakable amongst the Longwing butterflies, however, see the photo above to make sure you have a correct identification of what you have caught, as it is closely mimicked by the Malachite (S. stelenes). It is a rainforest species that often is found flying high in the canopy and is therefore not often seen. However, the adults do come down to nectar on flowers, and the females fly along the forest edges looking for suitable host plants to lay their eggs. The larvae feed on a great range of host plants from the Passion Vine family, and accept Passiflora caerulea in captivity. It is interesting that P. dido ranges from Mexico to the Amazon basin, yet its mimic, the Malachite (S. stelenes) (which also occupies the same area in Central and South America) is found as far north as southern Texas and Florida, where Philaethria dido is not found. A Final Note About Mimicry, by Other Species - Ithomids, Nymphalids and Additional Butterfly Families I have attempted to provide a detailed account to help you identify the Longwing butterflies in your region, and their associated host plants. However, there are more Longwing species in Central and South America, and this paper covers all the species that you are most likely to find and be interested in breeding. 22 By Nigel Venters

24 For example, there are two more Longwings that look like small Gulf Fritillaries - Dione glycera and Dione juno. Quite frankly, they are nowhere as nice as the species I have illustrated, and as I said, they are smaller species. I have not attempted to delve too deeply into the complex mimicry ring found in Central and South America, where there are a number of species from the Ithomiidae family that may look identical to the Longwings. They are fairly easy to separate from the Longwings as they all have pale yellow or white antennae. The Painted Lady & Longwing Articles Workshop Article This is the first in a series of four articles in this section. It is intended to give an introduction to understanding and identifying the Painted Ladies and Red Admirals from around the world. Article follows with an introduction to understanding and identifying the Longwing butterflies (Heliconid) of the Americas. Workshop Article Article goes into some depth on breeding the butterflies from both these families as they share common breeding requirements. This article covers breeding these butterflies using natural host plants. Workshop Article Article instructs on raising Painted Ladies on artificial diets. The article provides several professional butterfly farmers methods to successfully raise healthy and beautiful butterflies using special diets instead of live host plants. Methods and instructions are provided by Melanie McCarthy (The Monarchy Butterfly Farm), Linda Rogers (Timshell Farm), and the late Dean Kirkwood (Kirkwood Butterfly Company). Many thanks to Dean s wife, Pam Kirkwood, for sharing Dean s raising methods with the butterfly world. Since I raise only on live host plant, I thought it best to have these professionals who have perfected the use of artificial diets, provide this information. I am grateful to these contributors for their parts in this chapter, which rounds out the breeding instruction! Acknowledgments Special thanks to Dale Clark, President of the Dallas County Lepidopterists Society for kindly allowing us to include his text and photos to help in the correct identification of the two Zebra Longwing subspecies, vazquezae (Texas) and tuckeri (Florida). Butterfly farmers in Texas, Florida and the southern states must be able to distinguish these subspecies. Thanks to Linda Rogers and Paul Chesterfield, who are my Butterfly Boutique partners for spending endless hours making sense of what I wrote, proof reading and then turning it into an article that is well presented and readable! 23 By Nigel Venters

25 And always, my thanks to Gracie, my wife, best friend and butterfly buddy in the whole world! Nigel Venters, Argentina, October 2012 The Blue Passion Vine (Passiflora caerulea). This is a great host plant for all these species of Longwing butterflies! Attached are text and photos from Dale Clark For identification of the two Zebra Longwing subspecies. 24 By Nigel Venters

26 Attachment I Identifying the Zebra Longwing Subspecies: Heliconius charithonia vazquezae and Heliconius charithonia tuckeri By Dale Clark, President, Dallas County Lepidopterists Society Zebra Longwing Subspecies Texas and Florida, USA The Zebra Longwing (Heliconius charitonius) occurs in two subspecies within the USA. Heliconius charithonia vazquezae is the Texas subspecies, and Heliconius charithonia tuckeri is the Florida subspecies. Butterfly farmers in Texas, Florida and the southern states must be able to distinguish these subspecies. Heliconius charithonia vazquezae Heliconius charithonia tuckeri Heliconius charithonia vazquezae Ventral hindwing showing the yellow band extending beyond the discal cell. Heliconius charithonia tuckeri Ventral hindwing showing yellow band just touching the lower edge of the discal cell. 25 By Nigel Venters

27 Heliconius charithonia vazquezae Close-up of ventral forewing showing solid yellow band. Heliconius charithonia tuckeri Close-up of ventral forewing showing broken yellow band. Heliconius c. vazquezae, the Texas subspecies: On the hind wing the yellow band is thicker, extending past the discal cell. This is easier to see on the ventral (underside) side. Also on the ventral side, on the forewing, the yellow band is thick and continuous. Heliconius c. tuckeri, the Florida subspecies: On the hind wing, the yellow band is thinner, just reaching the edge of the discal cell. This is easier to see on the ventral (underside) side. Also on the ventral side, on the forewing, the yellow band is thin and usually broken, sometimes being almost nonexistent. This is quite variable. Other traits: Heliconius c. vazquezae: The dark coloring is a more brown appearance; the bands of yellow have a somewhat orange tinge. Heliconius c. tuckeri: The dark coloring is a darker brown/black; the bands of yellow are almost greenish/lime colored. Natural range: Heliconius c. vazquezae: Mexico, Arizona, New Mexico, Texas, northward to Oklahoma, Kansas, rarely further north. Heliconius c. tuckeri: Florida, Georgia, South Carolina. Mississippi and Louisiana?? 26 By Nigel Venters