CHAPTER IV. New-World Mimics

NEW-WORLD MIMICS

Of all the continents South America affords the greatest wealth of butterfly life, and it is in the tropical part of this region that many of the most beautiful and striking cases of mimicry are to be found. Viewed as a whole the butterfly population presents several features which serve to mark it off from that of the other two great tropical areas. In the first place the proportion of gaily coloured forms is higher. Bright red, yellow or fulvous brown contrasted with some deep shade approaching black form the dominant notes. Sombre coloured species are relatively scarcer than in the Oriental and African regions. In the second place when looking over collections from this part of the world one cannot help being struck by the frequency with which similar colour combinations occur over and over again in different as well as in the same groups. Now it is a simple scheme of black with an oblique scarlet band upon the fore wings--now an arrangement with alternating stripes of bright brown and black relieved with patches of clear yellow--now again a scheme of pure transparency and black. {38} Gay and pleasing as are the designs turned out the palette is a small one and invention is circumscribed. Under such conditions it might well be supposed that instances of close resemblance between different species would be numerous, and this in effect is what we find.

As in Asia with its Euploeines and Danaines, and in Africa with its Danaines and Acraeines, so in S. America are the fashions set by two dominant groups of models. These are the Heliconinae and the Ithomiinae, both peculiar to this region and both characterised, like the Old-world Danaids, by slow flight and great tenacity of life. Both live on poisonous plants--the Heliconines on Passifloras and the Ithomiines on Solanaceae. In both groups, but more especially in the Ithomiinae, the species are numerous, and the number of individuals in a species often beyond computation. From the point of view of mimicry these two groups have so much in common that they may conveniently be considered together.

It was from among the Ithomiines, as already pointed out, that the models came for the Pierine mimics of the genus Dismorphia upon which Bates founded the theory of mimicry. Though the Pierine mimics are the most striking the Heliconines and Ithomiines are mimicked by members of other groups. A few Papilios (Pl. X, fig. 8), certain Nymphalines such as Protogonius (Pl. X, fig. 9), Eresia, Phyciodes and Colaenis (Pl. XI, fig. 4), together with various day-flying moths, more particularly of the genera {39} Castnia and Pericopis, are among the well-known mimics of this group of models. The models themselves are very variable in appearance. In one locality the predominant pattern is black with a warm red-brown diagonal bar occupying rather more than a third of the fore wing (Pl. XV, fig. 5), in another it consists of parallel bands of black and fulvous brown with clear yellow patches at the tips of the fore wings (cf. Pl. X, fig. 7), while in yet another locality it is different again. Different localities often have their own peculiar pattern and this affects the various mimics as well as the Ithomiine and Heliconine models.

These groups of different species, some belonging to palatable and some to unpalatable groups, all exhibiting a close resemblance in colour and pattern, are far more strikingly developed in S. America than in either Asia or Africa, and it is not uncommon for eight or ten species to enter into such an association. A group of this sort which possesses unusual interest is the so-called "Transparency Group" from certain parts of the Amazon region. It was originally described by Bates with seven species belonging to six different genera. To-day it is said that no less than 28 species of this peculiar facies are known, though some are excessively rare. The majority are Ithomiines, but two species of the Danaine genus Ituna, the Pierine Dismorphia orise (Pl. XII, fig. 2), the Swallow-tail Papilio hahneli, and several species of diurnal moths belonging to different families (cf. Pl. XII, fig. 4) also enter into the combination. {40} In connection with it there is a feature of peculiar interest in that the transparent effect is not always produced in the same way. In the Ithomiines such as Thyridia, where there are normally two kinds of scales, the wider ones for the most part lose their pigment, become much reduced in size and take on the shape of a stumpy V (Pl. XIV, fig. 3). Also they stand out for the most part more or less at right angles to the wing, and the neck by which they are joined to the wing membrane is very short. The longer and narrow form of scales also tend to lose their pigment and become reduced to fine hairs. In Dismorphia the scales, which are of one sort, are also reduced in size though apparently not in number. Like the wider scales of the Thyridia they tend sometimes to project at right angles to the wing membrane, though not to the same extent as in the Ithomiine: possibly because the neck of the scale is not so short. As in Thyridia these reduced scales lose their pigment except in the transition region round the borders of the transparent patches. In Ituna there is a difference. The scales are not reduced to the same extent in point of size. Their necks are longer as in normal scales and they lie flat on the wing membrane. The majority of the scales, as in the preceding cases, lose their pigment, but mixed up with them is a certain proportion, about one-quarter, {41} in which the pigment is retained. In Castnia and in Anthomysa the scales on the transparent parts which are without pigment are also somewhat reduced in size, being stumpier than the normal ones. At the same time they tend to stand out at right angles to the wing membrane. The neck here again is shorter in the transparent than in the pigmented scales. A good deal of stress has been laid upon this case by some supporters of the theory of mimicry, since it is supposed to shew that a similar effect can be brought about in a variety of ways; consequently the existence of this assembly of similar transparent forms belonging to various families cannot be put down as due to the effect of similar conditions, but must be regarded as having arisen in each instance in a different manner through the independent action of natural selection. It is doubtful, however, whether such a conclusion necessarily follows from the facts. In all of the cases the process would appear to be similar: loss of pigment, reduction in the size of the scales, and eventually a tendency for the scales to stand at right angles to the wing--this last part of the process apparently depending upon the reduction of the neck of the scale. It has been said that greater transparency is brought about by the scales standing out at right angles in this way, but as the scales {42} themselves are already transparent there would appear to be no reason why this should be so. Of course the process has not proceeded in all of the forms to the same extent. There is least change in Ituna where the scales are not much reduced in size and where a fair proportion are still pigmented. There is probably most in an Ithomiine such as Thyridia, where the scales are not only small and entirely without pigment, but also are for the most part neckless so that they stand out at right angles to the wing. Having regard to the fact that several widely separate genera with different types of scaling formed the starting points, the final results do not seem to preclude the supposition that the transparency has arisen through a similar process in all of them.

It is somewhat remarkable that no Satyrine exhibits mimicry in S. America, in spite of the fact that transparency of the wings, as in so many of the butterflies of this region, is quite common in the group. On the other hand the relatively large number of more or less mimetic Pierines is a striking feature of S. America. For the most part they belong to the genera Dismorphia and Perrhybris, and resemble the yellow, black, and brown Heliconines and Ithomiines, though some of the former genus are mimics of the small transparent Ithomiines. Some of the species of Pereute with their dark ground colour and the bright red bar across the fore wing (Pl. XI, fig. 6) resemble Heliconius melpomene, as also does Papilio euterpinus. But some of the most interesting Pierine {43} mimics are several forms belonging to the genus Archonias (Pl. XI, fig. 10) which exhibit the simple and striking arrangement of black, red and white so characteristic of the Swallow-tail Poison-eaters of S. America. They form one of the rare instances of a Pharmacophagus Papilio being mimicked by a butterfly which does not belong to the Swallow-tail group.

As everywhere in the tropics the Papilios of S. America supply a goodly proportion of the mimicry cases. A few, such as P. zagreus (Pl. X, fig. 8), enter into the black-brown and yellow Ithomiine-Heliconine combination; P. euterpinus resembles Heliconius melpomene (Pl. XI, fig. 5); P. pausanias is like Heliconius sulphurea (Pl. XI, figs. 1 and 2). But this practically exhausts the list of Papilios which mimic Heliconines and Ithomiines. The great majority of mimicking Swallow-tails in S. America find their models among the Poison-eaters of their own family, offering in this respect a contrast to those of Asia where the majority of models are among the Danaines and Euploeines, and of Africa where they are exclusively Acraeines or Danaines.

The Poison-eaters of S. America fall into two well-marked groups which we may call the red-spotted and the dark green groups respectively. The red spotted group form a remarkably compact and uniform assemblage. The general ground colour is a deep black-brown (Pl. XI, figs. 8 and 9), the hind wings are almost invariably marked with red near the centre or towards the outer margin, and the fore wing may {44} or may not bear a patch which is generally whitish in the female, though often of a brilliant blue or green in the male. This simple colour scheme with variations runs throughout about three-quarters (some 40 species) of the Poison-eaters. The same general colour scheme is also found in about two dozen species of the unprotected Swallow-tails. As the total number of the unprotected species is placed by Seitz at less than 100 this means that fully one-quarter of them fall into the general colour scheme adopted by the majority of the Poison-eaters. In many cases the resemblance between mimic and model is so close as to have deceived the most expert entomologists before the structural differences between the groups had been appreciated (cf.