And thus, summing up the caterpillars, both of b.u.t.terflies and moths, out of eighty-eight spiny and hairy species, only one is green, and even this may not be protectively colored, since it has conspicuous yellow warts. On the other hand, a very great majority of the black and brown caterpillars, as well as those more or less marked with blue and red, are either hairy or spiny, or have some special protection.
Here, then, I think we see reasons, for many at any rate, of the variations of color and markings in caterpillars, which at first sight seem so fantastic and inexplicable. I should, however, produce an impression very different from that which I wish to convey, were I to lead you to suppose that all these varieties have been explained, or are understood. Far from it; they still offer a large field for study; nevertheless, I venture to think the evidence now brought forward, however imperfectly, is at least sufficient to justify the conclusion that there is not a hair or a line, not a spot or a color, for which there is not a reason--which has not a purpose or a meaning in the economy of nature.
[Ill.u.s.tration]
PROTECTIVE RESEMBLANCES IN SPIDERS
BY ELIZABETH G. PECKHAM.[10]
[10] Abbreviated from the occasional Papers of the Natural History Society of Wisconsin, Vol. I., 1889. By permission.
[Ill.u.s.tration]
There are, among spiders, two forms of protective modification: the first, including all cases of protective resemblance to vegetable and inorganic things--that is, all modifications of color or of color and form that tend to make them inconspicuous in their natural relations--I shall call direct protection. The second form, which I shall call indirect protection, includes two cla.s.ses, the spiders which are specially protected themselves and those which mimic other creatures which are specially protected.
Spiders are specially protected when they become inedible through the acquisition of hard plates and sharp spines. The modification of form is frequently accompanied by conspicuous colors, which warn their enemies that they belong to an unpalatable cla.s.s.
The second cla.s.s of indirectly protected spiders--those that mimic specially protected creatures--presents some difficulties, since it is not always easy to determine whether the purpose of mimicry is protection or the capture of prey. The resemblance may, as is frequently the case in direct protection, serve both purposes.
In looking for instances of protective form and color among spiders we encounter one difficulty at the outset. The meaning of a protective peculiarity can be determined only when the animal is seen in its natural home. The number of strangely modified forms depicted in descriptive works on spiders is enormous. Bodies are twisted, elongated, inflated, flattened, truncated, covered with tubercles or spines, enclosed within chitinous plates, colored like bark, like lichens, like flowers of every imaginable hue, like bird droppings, like sand or stones, and in every one of these modifications there is doubtless an adaptation of the spider to its surroundings which, when it is studied out of its natural relations, we can only guess at.
It has been well said that in these protective resemblances those features of the portrait are most attended to by nature which produce the most effective deception when seen in nature; the faithfulness of the resemblance being much less striking when seen in the cabinet....
DIRECT PROTECTION. RESEMBLANCES TO VEGETABLE AND INORGANIC THINGS.--As a general rule the forms and colors of spiders are adapted to render them inconspicuous in their natural homes. Bright colored spiders, ... either keep hidden away or are found upon flowers whose tints harmonize with their own. This rule, while it has numerous exceptions, is borne out by the great majority of cases. A good ill.u.s.tration is found in the genus Uloborus, of which the members bear a deceptive resemblance to small pieces of bark, or to such bits of rubbish as commonly become entangled in old deserted webs. The only species in our neighborhood is Uloborus plumipes, which I have almost invariably found building in dead branches, where its disguise is more effective than it would be among fresh leaves. The spider is always found in the middle of the web, with its legs extended in a line with the body. There has been, in this species, a development along several lines, resulting in a disguise of considerable complexity. Its form and color make it like a sc.r.a.p of bark, its body being truncated and diversified with small humps, while its first legs are very uneven, bearing heavy fringes of hair on the tibia and having the terminal joints slender. Its color is a soft wood-brown or gray, mottled with white. It has the habit of hanging motionless in the web for hours at a time, swaying in the wind like an inanimate object. The strands of its web are rough and inelastic, so that they are frequently broken; this gives it the appearance of one of those dilapidated and deserted webs in which bits of wind-blown rubbish are frequently entangled....
Out of seven examples of the species taken during one summer, five were found in dead tamarack branches, one on a dead bush, and the seventh, an interesting variety, under the eaves of a porch. My eye was caught by what seemed to be a string of eleven coc.o.o.ns (it is not common to see more than four in a web). On attempting to take them down I was surprised to see one of the supposed coc.o.o.ns begin to shake the web violently. Ten were what they seemed to be, but the eleventh was the mother spider, whose color and general appearance was exactly like that of the little cases that she had made for her eggs....
We come now to a large and interesting cla.s.s in genus Epeira. I refer to those species, mostly nocturnal, which are protected during the day, not by hiding in crevices, nor in any way actually getting out of sight, but by the close resemblance which they bear to the bark of the trees to which they cling. This resemblance is brought about in two ways; through their color, which is like that of wood or lichens, and through their tuberculated and rugose forms, which resemble rough bark.
[Ill.u.s.tration: FIG. 1.--CaeROSTRIS MITRALIS (from Vinson).]
[Ill.u.s.tration: FIG. 2.--CaeROSTRIS MITRALIS, in profile (from Vinson).]
One of the most remarkable of these forms is C. mitralis, a Madagascar species, which, looked at in profile, probably resembles a woody knot.
The abdomen is divided into two divergent cones (Fig. 1). The entire upper surface of the body is covered with conical elevations, which render it rough and uneven; the sides of the abdomen are made up of several layers, which form stages, one above another, like the ridges of bark on a woody excrescence. The legs, formed of wide, flattened plates, make the base. The color of the spider is yellowish-gray, varied with white and dark reddish-brown. It has the habit of perching on a branch and clasping it like a bird, so that the elaborate modification of form, which would be useless if the spider hung exposed in the web, is made as effective as possible.
To take an example nearer home, E. infumata is a large, round-bodied spider, with two humps on the abdomen, which Emerton describes from New England as being brown, mottled with white and black; he adds that when it draws in its feet it looks like a lump of dirt. Infumata, in Wisconsin, has always a good deal of bluish-green on the upper surface of the abdomen. This may be a variety which has been so developed as to resemble the lichens which cover the tree to which it clings. It is one of the spiders which bear a good deal of handling without uncurling its legs, or showing any sign of life. Its humpy form and its color give it a very inanimate appearance. It is rather common in our neighborhood and may be caught in the late twilight while building its web, but to search for it in the daytime, even among the trees that it most frequents, is an almost hopeless task. A more grotesque form is E. stellata, in which the abdomen has not two, but twelve or fifteen humps. These are so disposed that the edge of the abdomen, all around, is scalloped. The colors are light and dark brown, modified by gray and white hairs. This spider remains motionless during the daytime, keeping its legs drawn up to its body. It is common on gra.s.s and low bushes. It is not found in Wisconsin, but the description of it suggests a resemblance to a piece of dead leaf.
There are many other spiders in this genus that have humps and are colored in brown, gray or dull yellow, as nordmanii, angulata, solitaria, etc. It is an almost universal habit among the Epeiridae to drop to the ground when threatened, and when a humped gray or brown spider drops to the ground and draws in its legs it is nearly indistinguishable from the lumps of earth, sticks and stones that surround it.
One of the Therididae which has the same protection is Ulesanis americana (Fig. 3). The abdomen, which covers the cephalothorax nearly to the eyes, has a prominent hump in the middle of the back and four or five others behind. Its color is in shades of brown and yellow.
[Ill.u.s.tration: FIG. 3.--ULESANIS AMERICANA (from Emerton).]
a.n.a.logous to the humped Epeiridae is Thomisus foka, of Madagascar, a spider which is regarded with great terror by the natives, as being so poisonous that even its breath is deadly. They say that cattle, when about to lie down, look carefully about to see if one of these spiders is in the neighborhood. This dread is, no doubt, inspired by the strange and uncanny aspect of a perfectly harmless creature. It has a rugose, tuberculated body of trapezoid form, the colors being brown and reddish, while the whole aspect is crab-like. The thick, short legs are reddish, covered with tubercules. The secret of its strange form is made clear when we learn that it resembles in color and general appearance the fruit of Hymenaea verrucosa, a tree common in the forests where this spider is found.
Among the curious forms which must have been developed through advantageous variations but which we are unable to explain, is Eriauchenus workmanni (Fig. 4).
[Ill.u.s.tration: FIG. 4.--ERIAUCHENUS WORKMANNI (from Cambridge).]
Epeira prompta, a variety of parvula, is a common spider in the State of Wisconsin. It is most frequently seen on cedar bushes, where its color harmonizes with that of the foliage and fruit. During the day it usually rests on a branch near its web. The back of the abdomen is of a peculiar bluish-green, exactly like that of the lichens growing on tree trunks. The bluish color is broken by waving black lines which imitate the curling edges of the lichens. The one represented in the plate was found on an old cedar which was covered with lichens. It was kept for two weeks in a gla.s.s-covered box, where it spent most of the time crouching in a corner. It built no web, but spun some irregular lines to run about on. It ate gnats, flies, and once a little jumping spider, S. pulex, which we were keeping in the same box, leaping upon its prey, as noted by Hentz, like an Attus. This seems a curious habit to be acquired by an Epeirid, since spiders, as we have noticed among our captives, are usually dependent for food upon what is caught in their webs. Prompta moves awkwardly, but very rapidly.
Drapetisca socialis, while quite a different looking spider, is protected in the same way--by its resemblance to the bark upon which it lives. Emerton speaks of finding it on the bark of spruce trees, which it "closely resembles in color." Menge says that it is common in Prussia, where it is seen in great numbers on fir trees, whose spotted bark it resembles in color, so that it is not easily seen. We have found them, most commonly, upon birch trees, and in this situation their color adaptation is perfect. Both the spider and the peeling bark of the tree are of a light silvery brown, covered with little blackish marks. On the bark these marks are, of course, irregular, while on the spider they form a pattern made up of straight and curved lines and dots, the legs being silvery, barred with blackish.
Another little Theridion that is found on birch bark has the same colors arranged a little differently. The abdomen above has a large and peculiarly irregular black patch, which shades off into mottled brown and black on the sides and below. The cephalothorax has stripes of brown and black, and the legs are barred with light and dark brown.
Spiders that live upon walls, fences, tree trunks, or on the ground harmonize in color with the surfaces upon which they are found, being usually gray, brown or yellow, mottled with black and white. This proposition is so well established as to need but few ill.u.s.trations.
The Therididae furnish many examples, as T. murarium, a gray spider varied with black and white, said by Emerton to live usually "under stones and fences, where it is well concealed by its color"; and Lophocarenum rostratum, a yellowish-brown spider, found among leaves on the ground. Among the Attidae bright s.e.xual coloring often gains the ascendancy over the protective tints, yet this family gives us good examples in such species as M. familiaris and S. pulex.
To these may be added an as yet undescribed species which we discovered last season in a neighborhood that we had searched thoroughly for eight summers. We found the new spider in great numbers, but could only detect it by a close scrutiny of the rail fences on which it lived, its color being dark gray....
[Ill.u.s.tration: FIG. 5. ORNITHOSCATODES DECIPIENS (from Cambridge).]
The last instance that I shall cite is a predaceous spider which is disguised from both its enemies and its prey by an elaborate combination of form, color, position, and character of web. I refer to Ornithoscatodes decipiens (Fig. 5), first described by Forbes and afterwards by Cambridge, the latter author giving in the same paper descriptions of three other species of the same genus, whose habits have not been noted, but whose protection is evidently of the same order as that of decipiens. I give Forbes"s interesting account of his capture of decipiens, quoting also the remarks by which Cambridge prefaces his description, since his explanation of the gradual development, through Natural Selection, of the spider"s deceptive appearance applies as well to all the cases of protective disguise which have been here enumerated.
The capture is described as follows:--
"On June 25th, 1881, in the forest near the village of Lampar, on the banks of the Moesi river in Sumatra, while my "boys" were procuring for me some botanical specimens from a high tree, I was rather dreamily looking on the shrubs before me, when I became conscious of my eyes resting on a bird-excreta-marked leaf. How strange, I thought, it is that I have never got another specimen of that curious spider I found in Java which simulated a patch just like this! I plucked the leaf by the petiole while so cogitating, and looked at it half listlessly for some moments, mentally remarking how closely that other spider had copied nature, when, to my delighted surprise, I discovered that I had actually secured a second specimen, but the imitation was so exquisite that I really did not perceive how matters stood for some moments. The spider never moved while I was plucking or twirling the leaf, and it was only when I placed the tip of my little finger on it, that I observed that it was a spider, when it, without any displacement of itself, flashed its falces into my flesh.
"The first specimen I got was in W. Java, while hunting one day for Lepidoptera. I observed a specimen of one of the Hesperidae sitting, as is often a custom of theirs, on the excreta of a bird on a leaf; I crept near it, intending to examine what they find in what one is inclined to consider incongruous food for a b.u.t.terfly. I approached nearer and nearer, and at last caught it between my fingers, when I found that it had as I thought become glued by its feet to the ma.s.s; but on pulling gently the spider, to my amazement, disclosed itself by letting go its hold: only then did I discover that I was not looking on a veritable bird"s excreta.... The spider is in general color white, spotted here and there with black; on the underside its rather irregularly shaped and prominent abdomen is almost all white, of a pure chalk white; the angles of the legs are, however, shining jet-black. The spider does not make an ordinary web, but only the thinnest film on the surface of the leaf. The appearance of the excreta rather recently left by a bird on a leaf is well known. There is a pure white deposit in the centre, thinning out round the margin, while in the central ma.s.s are dark portions variously disposed; as the leaf is rarely horizontal, the more liquid portions run for some distance. Now, this spider one might almost imagine to have in its rambles "marked and inwardly discerned" what it had observed, and to have set about practising the "wrinkles" gained; for it first weaves a small, irregular patch of white web on some prominent leaf, then a narrow streak laid down towards its sloping margin ending in a small k.n.o.b; it then takes its place on the centre of the irregular spot on its back, crosses its black-angled legs over its thorax, and waits.
Its pure white abdomen represents the central ma.s.s of the bird"s excreta, the black legs the dark portions of the slime, while the web above described which it has spun represents the more watery marginal part (become dry), even to the run-off portion with the thickened k.n.o.b (which was not accidental, as it occurred in both cases), like the residue which semi-fluid substances ending in a drop leave on evaporation. It keeps itself in position on its back by thrusting under the web below it the spines with which the anterior upper surfaces of the legs are furnished." ...
PROTECTIVE HABITS.--Going along with these forms of protective resemblance, we find certain habits which sometimes serve independently to protect the spider, but oftener are supplemental to color and form. Many species hide in crevices or in leaves which they roll up and bind together at the edges. In the Epeiridae some are like thaddeus, which makes a little tent of silk under a leaf near its web.
The young thaddeus also makes a tent, but spins its little geometrical web on the under side of the leaf, the edges being bent downward. E.
insularis has the more common habit of forming its tent by drawing the edges of two or three leaves together with strands of web; in this it sits all day, but at night descends and occupies the centre of the web during the hours of darkness. I have often found it in this position when hunting nocturnal species by lantern light. It is probable that in tropical countries the monkeys, and perhaps the birds, which devour these large Epeiridae have learned to recognize their webs, which are very large and conspicuous, and to trace them to their hiding places close by; and thus may have arisen the curious habit noticed by Vinson as possessed by E. nocturna and E. Isabella of destroying the web each morning and rebuilding it at night; the spider in this way gaining greater security from diurnal enemies.
Atypus abbotii builds a purse-shaped tube which is found attached to the bark of trees, and which has the external surface dark and covered with sand. The trap-doors which close the nest of some of the Territelariae are wonderful examples of protective industry. They fit with such absolute accuracy into the openings of the nests and are so covered on the upper side with moss, earth, lichens, etc., as to be indistinguishable from the surrounding surface.
The rectilinear lines which are stretched in front of the webs of many Epeirids are useful in taking and sending on to the spider the shock which tells of an approaching enemy. Some spiders, when danger threatens, shake the web so violently as to grow indistinct to the eye, and others, as Pholcus atlanticus, hang by the legs and whirl the body rapidly with the same bewildering result....
A habit common to many spiders, especially among the Epeiridae, is that of dropping to the ground at the approach of danger and resting motionless among the dirt, sticks, leaves, etc., which they resemble in color. The holding of the body in some peculiar position, as in Uloborus, Hyptioides, and the flower-like Thomisidae, is a necessary accompaniment to the color modification.
The coc.o.o.ns of spiders are seldom left exposed and unprotected. We find them in corners and crevices, concealed in rolled up leaves or under bark. Very often the coc.o.o.n itself is covered over with a sheet of web. In some families the mother carries it about with her attached to the underside of the abdomen. In other she carries it in her falces until the young are hatched. The coc.o.o.ns of others, as Uloborus, Argyrodes, etc., while hung out in the web are still concealed by deceptive form and color, or by being covered with rubbish.
Cambridge speaks of A. brunnea, whose coc.o.o.ns "are covered over very soon after they are made and the eggs deposited in them, with a coating of clay, which effectually destroys all their form and beauty.
This coating of clay answers probably two ends: first, the concealment of the coc.o.o.n and its protection from insect enemies; and, secondly, the protection of the eggs from the too powerful rays of the sun, dry clay being (as is well known) one of the best non-conductors of heat."
The peculiar coc.o.o.n of C. bisaccata is described by Emerton as follows: "Only one specimen of this (_bisaccata_) was found on a beech tree at New Haven with two coc.o.o.ns. These were dark brown, as dark as the bark of the tree and as hard. Around the middle of each was a circle of irregular points. One of the coc.o.o.ns was attached by a strong stem to the bark, and the other was attached in a similar way to the first coc.o.o.n. The spider held on to one of the coc.o.o.ns." In this instance the egg has evidently the same protection as that possessed by the gray, bark-haunting spiders, with the added advantage of hardness.
The habit of distributing the eggs through a number of coc.o.o.ns made at intervals of several days, is protective. In this way, although one or two of the coc.o.o.ns may be pierced by the ichneumon, there is a chance that part of the brood may survive.
INDIRECT PROTECTION.--The indirectly protected group includes those spiders which are rendered inedible by the possession of sharp spines and chitinous plates, and also those that mimic other specially protected creatures.
The females of the specially protected group are characterized by the following attributes: