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Their inedibility, which they owe to a more or less coriaceous epidermis and an armature of strong sharp spines (Fig. 6).
Their brilliant colors--glistening black and white, yellow, fiery gold, metallic silver, rose-color, blue, orange and blood-red.
Their habit of hanging always exposed in the centre of the web.
[Ill.u.s.tration: FIG. 6.--GASTERACANTHA CREPIDOPHORA (from Cambridge).]
In an interesting discussion of the protective value of color and marking in insects, Poulton says that "the smaller convergent groups of nauseous insects often present us with ideally perfect types of warning patterns and colors--simple, crude, strongly contrasted--everything subordinated to the paramount necessity of becoming conspicuous," the memory of enemies being thus strongly appealed to.
This proposition is well ill.u.s.trated by the Gasteracanthidae. Among larvae the warning colors are almost invariably black and white, or black (or some very dark color), in contrast with yellow, orange and red. These are the colors that also constantly recur among the Gasteracanthidae.
Cases that may be more justly considered exceptions to the rule that these hard, uneatable spiders are conspicuous are such species as Acrosoma rugosa (Fig. 7). One of this species was sent me by Mrs.
Treat last summer. It lived for several weeks in my window, making no regular web, but hanging among a few irregular strands. It ate nothing, although provided with insects, but drank greedily of water.
It might seem that its black and white coloring would make it conspicuous, but in connection with its irregular shape and its way of hanging motionless in the web it had the opposite effect.
We have no reason to suppose that the cla.s.s represented in rugosa is like that touched upon by Poulton, in which very protectively colored larvae suddenly a.s.sume a terrifying aspect on the near approach of an enemy; still they do enjoy a kind of double protection.
[Ill.u.s.tration: FIG. 7.--ACROSOMA RUGOSA. Left-hand figure female, right-hand figure male (from Emerton).]
They are inconspicuous, and thus likely to escape attack, but in case they are attacked they have still the advantage of being quickly rejected. This experience cannot be as fatal to them as to the soft and thin skinned larvae. Their hard covering and projecting spines would protect them to such an extent as to give them a fair chance of surviving.
In one respect the inconspicuous Gasteracanthidae have a decided advantage over their bright-colored relatives. The birds, indeed, avoid the conspicuous ones, but their brilliancy serves to attract another enemy against which spines are no protection--the hunter wasp, which, as we have seen in the work of Bates, sometimes provisions its nest wholly with spiders of this family. Mr. Smith gives like testimony, saying:
"Spines on the abdomen of certain spiders would serve as a protection against vertebrate enemies, though they do not protect against the hunter wasps, which frequently provision their nests with these species." He adds, however, that most of the spiny spiders are common, and that their colors make them conspicuous; just as b.u.t.terflies that are protected by an odor are common and bright-colored....
MIMICRY.--Mimicry, or the imitation of animal forms, while it is a form of indirect protection, differs in no essential respect from the imitation of vegetable and inorganic things. As Bates has said, the object of mimetic tendencies is disguise, and they will work in any direction that answers this purpose.
In nearly all respects spiders come under the three laws given by Wallace, as governing the development of mimetic resemblances in several large cla.s.ses. These laws are as follows:
1. In an overwhelming majority of cases of mimicry, the animals (or the groups) which resemble each other inhabit the same country, the same district, and in most cases are to be found together on the very same spot.
2. These resemblances are not indiscriminate, but are limited to certain groups, which, in every case, are abundant in species and individuals, and can often be ascertained to have some special protection.
3. The species which resemble or "mimic" these dominant groups, are comparatively less abundant in individuals, and are often very rare.
The second and third of these laws are confirmed by what we know of mimetic resemblances among spiders. They mimic ants much oftener than other creatures, and ants are very abundant, are specially protected, and are much more numerous than the mimetic spiders. To the first law, also, they conform to a great extent, since everything tends to show that in tropical America and in Africa the ant and the spider, the one mimicked and the other mimicking, are always found together.
So far as I can discover, however, the ant-like spiders of North America are not found in company with any species of ant which they resemble. This may be because they do not mimic any particular species, but only the general ant-like form; or, considering that the genera which contain their nearest relatives are much more abundant in Central and South America, it may be that these forms were originally tropical, mimicking some tropical species of ants, and that after the Glacial Epoch they migrated northward, leaving the ants behind them.
However this may be, their peculiar form has served them well, since they have maintained themselves as fairly abundant species with a lower fecundity than is found in any other group of spiders.
The cases in which one species mimics another may be divided, according to the kind of benefit derived, into four cla.s.ses: Cla.s.s 1.
As a rule, where we find one species mimicking another, the mimicked species possesses some special means of defence against the enemies of both. This defence may consist of a disagreeable taste or odor, as in the Heliconidae, which are mimicked by other b.u.t.terflies; of some special weapon of offence, as where wasps and bees are mimicked by flies and moths, or poisonous vipers by harmless caterpillars; or of a hard sh.e.l.l, as where the coriaceous beetles are mimicked by those that are soft-bodied.
Instances of this rule are exceedingly numerous; indeed, Wallace says that specially protected forms are always mimicked; still we have nothing mimicking our Gasteracanthidae.
Cla.s.s 2. The mimetic may prey upon the mimicked species, its disguise enabling it to gain a near approach to its victims; as the mantis, mentioned by Bates as exactly resembling the white ants upon which it feeds; and the flies which mimic bees, upon which they are parasitic, and are thus able to enter the nests of the bees and lay eggs on the larvae.
Cla.s.s 3. The mimetic species may, by its imitation, be protected from the attacks of the creature it mimics, as is the case with the crickets and gra.s.shoppers which mimic their deadly foe, the hunter wasp.
Cla.s.s 4. The mimetic species may prey upon some creature which is found commonly with, and is not eaten by, the mimicked species.
No two of these cla.s.ses are mutually destructive so that in any case of mimicry a double advantage may be gained.
Let us see which of these advantages has directed the development of mimetic tendencies among spiders.
While among beetles and b.u.t.terflies we most commonly find mimicry of one species by another within the same order, we have no instance of a spider mimicking another spider. This may be accounted for by the fact that the specially protected spiders depend for their safety upon the possession of hard plates and spinous processes, and although the hardened epidermis might be imitated (we know that hard-sh.e.l.led beetles are mimicked by others that are soft), spines could scarcely be imitated by a soft-bodied creature with sufficient accuracy to insure disguise.
While spiders most commonly mimic ants, we hear also of their imitating beetles, snail-sh.e.l.ls, ichneumons and horseflies. There is also a curious Madagascar species which looks exactly like a little scorpion, the resemblance being heightened by its habit of curving its flexible tail up over its back when irritated.
Those that resemble beetles comprise nearly all the species of the genera Coccorchestes and Homalattus. These are small spiders with short, convex bodies. The abdomen fits closely over the cephalothorax, and the epidermis, which has usually a metallic l.u.s.tre, is sometimes coriaceous. Striking examples are found in H. coccinelloides, which bears a strong resemblance to beetles of the family Coccinelloidae, and in C. cupreus, in which certain marks on the abdomen imitate the elytra of beetles.
The following account of a spider which mimics a snail-sh.e.l.l is given by Mr. G. F. Atkinson;--
"An undescribed species of _Cyrtarachne_ mimics a snail-sh.e.l.l, the inhabitant of which, during the summer and fall, is very abundant on the leaves of plants in this place. In the species of Cyrtarachne the abdomen partly covers the cephalothorax, is very broad at the base, in this species broader than the length of the spider, and rounds off at the apex. When it rests upon the under side of a leaf with its legs retracted it strongly resembles one of these snail-sh.e.l.ls by the color and shape of its abdomen. The two specimens which I collected deceived me at first, but a few threads of silk led me to make the examination.
The spider seemed so confident of its protection that it would not move when I jarred the plant, striking it several hard blows. I pulled the spider forcibly from the leaf, and it did not exhibit any signs of movement until transferred to the cyanide bottle." ...
Trimen gives an account of the imitation, by spiders, of horseflies, a case falling into Cla.s.s 2, as follows:--
"Hunting spiders are in some cases very like their prey, as may everywhere be noticed in the case of the species of Salticus which catch horseflies on sunny walls and fences. The likeness is not in itself more than a general one of size, form and coloring; but its effect is greatly aided by the actions of the spider, which walks hurriedly for short distances, stopping abruptly, and rapidly moving its falces, in evident mimicry of the well-known movements so characteristic of flies."
Instances of spiders mimicking ants are very numerous, and in many cases the resemblance is so close as to, at first sight, deceive a trained naturalist. This resemblance is brought about by the spider's body being elongated and strongly constricted, so that it appears to be composed of three segments instead of two, by the color, by the way in which the spider moves about, zig-zagging from side to side like an ant, and by its habit of holding up one pair of its legs and moving them in such a way that they look exactly like the antennae of an ant.
Ants may be regarded as specially protected, by their sharp, acid flavor, and in some species by the possession of stings or of h.o.r.n.y processes.
On the ground that there are birds which do eat ants, and eat them greedily, it has been thought by some naturalists that they cannot be considered specially protected creatures, and that, as spiders can therefore derive no protection from mimicking them, all cases of such mimicry depend upon the spider's increased ability to capture the ants as prey, but I am convinced that this is too hasty a conclusion. It is unquestionably true that some birds feed almost exclusively upon ants, but these are the exceptions. It is a common thing to find that specially protected groups, which are safe from the attacks of most creatures, have their special enemies. Thus, even the nauseous Heliconidae are preyed upon by certain spiders and wasps; and bees, in spite of their stings, are preferred to other insects by the bee-eaters. Moreover, the ant-devouring birds are found largely among the wood-p.e.c.k.e.rs, which eat the ants that run on the trunks of trees, and are therefore not a source of danger to the ant-like spiders, the American species of which, so far as I can learn, live entirely upon the ground.
In the United States comparatively small numbers of either ants or spiders are eaten by birds, but in tropical America there are enormous numbers of humming-birds feeding almost exclusively upon spiders, and there the protective advantage of looking like ants must be of great importance to the smaller species.
Belt considers that the advantages gained by ant-mimicking Central American spiders lies entirely on the side of protection. In relation to this subject he says: "Ant-like spiders have been noticed throughout tropical America and also in Africa. The use that the deceptive resemblance is to them has been explained to be the facility it affords them for approaching ants on which they prey. I am convinced that this explanation is incorrect, so far as the Central American species are concerned. Ants, and especially the stinging species are, so far as my experience goes, not preyed upon by any other insects. No disguise need be adopted to approach them, as they are so bold that they are more likely to attack a spider than a spider them. Neither have they wings to escape by flying, and generally go in large bodies easily found and approached. The use is, I doubt not, the protection the disguise affords against small insectivorous birds. I have found the crops of some humming-birds full of small, soft-bodied spiders, and many other birds feed on them. Stinging-ants, like bees and wasps, are closely resembled by a host of other insects; indeed, whenever I found any insect provided with special means of defence I looked for imitative forms, and was never disappointed in finding them."
The ant-like species are probably protected by their appearance from the attacks of many of the larger spiders. We have kept great numbers of Attidae in captivity, and, although they devoured flies, gnats, larvae, and other spiders, they would never touch ants. Among spiders, however, as among birds, we find that certain groups subsist almost entirely upon ants.
The cla.s.s of spiders whose mimicry protects them from their enemies, whether they are birds or other spiders, probably includes at least two of our own ant-like species, Synageles picata and Synemosyna formica, which, in confinement, are always hungry for gnats, but will not touch ants, even of small size.
The existence of a cla.s.s of spiders which mimic the particular species of ants upon which they prey is not to be questioned, but it is doubtful whether the benefit to the spider is increased facility in capturing the ant, or whether it is merely protective. It may be that the spider, by virtue of its resemblance to the ant, not only gets an abundant supply of food, but also escapes being eaten itself, and thus enjoys a double advantage. Both Bates and Wallace take the ground that the advantage derived by the spider consists in greater ease in the capture of prey, but both of these writers refer to spiders only incidentally to ill.u.s.trate a general proposition, without special consideration of their peculiar conditions.
Mr. Herbert Smith, who has paid a good deal of attention to this subject, is inclined to believe that the mimicry in question is entirely protective. He writes as follows:--
"In the United States there are a few rare spiders that mimic ants.
Here at Taperinha we find a good score of species of these spiders aping the various kinds of ants very closely; even the odd, spiny wood-ant, _cryptocerus_, furnishes a pattern, and there are spiders that mimic the wingless ichneumons. We find, after a while, that the spiders prey upon ants just as our spiders catch flies; indeed, this fact has already been noted by other observers. But we go a step beyond the books when we discover not only that the spiders eat the ants, but that they eat the particular ants which they mimic. At all events, we verify this fact in a great number of cases, and we never find the spiders eating any but the mimicked species.... I do not like to hazard a theory on this case of mimicry. It is difficult to suppose that the quick-witted ants would be deceived even by so close a resemblance; and, in any case, it would seem that the spiders do not require such a disguise in order to capture slow-moving ants. Most birds will not eat ants; it seems likely, therefore, that this is simply another example of protection; the spider deceives its enemies, not its prey; it mimics the particular species that it feeds on, because it is seen in that company when it is hunting, and among a host of similar forms is likely to pa.s.s unnoticed."
At first sight, and especially in view of the fact that such cases are not uncommon among insects, it would be naturally supposed that the object of the mimicry was to enable the spider to approach its victim without exciting suspicion; and it is difficult to account, on any other supposition, for the very close resemblance between certain species of spiders and the particular species of ants which they prey upon. It seems as though the highest point of _protective_ benefit would have been reached long before the resemblance of the spider to the ant had become so close as it really is. On the other hand, it is difficult to believe that ants are deceived, even by those spiders which mimic them most closely, when we remember that their perceptions are so keen that they discriminate not only between ants of their own and different species, but even between ants of their own species living in two different communities.
The mimicry of ichneumon flies by spiders was noted some years ago by Mr. Herbert Smith. This case comes under Cla.s.s 3, in which one species mimics another which preys upon it. Great destruction is caused by ichneumons which lay their eggs on the bodies of the live spiders, and the disguise probably protects the spider by leading the fly to mistake it for one of its own species.
We have no proof that spiders ever mimic ants as a method of escaping from them, but it is possible that this sometimes happens. We know that some ants prey upon them. The foraging ants of South America destroy spiders as well as many kinds of insects, and Wallace mentions a small, wood-boring ant which fills its nest with small spiders.
If the spiders that feed upon ants deceive them by their mimicry those which are preyed upon by ants would gain an advantage by a similar disguise. I once placed a little ant-like spider of the genus Herpyllus in a bottle with three ants no larger than itself, which I had caught with it in the sweep-net. In a very few minutes the ants had killed and begun to devour the spider. It may be that the resemblance was sufficiently close to deceive them in the open, but failed when spider and ants were confined together in close quarters.