Savory - Arachnida 1977

Savory - Arachnida 1977 Savory - Arachnida 1977

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88 II. DE ARACHNIDIS capable of transporting for hundreds of kilometres an animal which, without it, would be wafted as many centimetres. In the spring and autumn, seasons in which large numbers of spiders are hatched, gossamer is more than usually prevalent and carries chiefly young ; in the summer the aeronauts include many half-grown and mature examples of the smaller species. The earliest opinion was that one ~pecies, "the gossamer spider", was responsible for all these threads, but this is not so. Emerron in 1918 found 69 American species in the Bristowe in 1930 listed 30 British ones belonging to six families, and Bishop in 1945 identified 25 species in six families from heights between 6·1 and 1,524 m. It is believed today that nearly all the families contain species which undertake aerial travel at some stage in theirlives, though it is clear that this cannot be expected of the largest, which are too heavy to fly. Even so, it cannot be maintained that the "ballooning habit" is common to all, or even to the majority of spiders. Take Emerton's total of69, Bristowe's total of30, and Bishop's total of25; add them together and then, not to seem parsimonious, multiply the sum by ten. This makes a generous allowance for more in different countries, but the product is only 1,240, a very small fraction of the number of species of spiders known to exist. A different method of dispersal is shown by some of the smallest mites that are parasitic on fruit t1 ees. These in the summer, either by grasping the legs of passing insects, or standing erect and leaping vertically if they feel a puff of air. The leap may then help them to be blown some distance. The common British mite Belaustium nemorum is often found holding the legs ofTipulidae; and several ofuropod mites attach themselves by a thread of their own excrement to the bodies of beetles. A more peculiar method of travel is found among false scorpions, and is known as phoresy. False scorpions are essentially lovers of darkness, and yet on occasions they are to be seen clinging to the legs of flies, harvestmen and other creatures, and thus being carried about at no effort to themselves. They grasp the fly's leg in their chelicerae but they are not parasites, for they neither injure nor feed on the fly. 1\\·enty-five or more species belonging to about 20 genera have been found in these circumstances. The habit is not equally prevalent at all times of the year, but during the short time that it appears it is reasonably frequent. Thus Vachon ( 1932) recorded the finding of 78 mens on the legs of 57 harvestmen in one week at the end of August. All belonged to the same species, and all but one were mature females. The highest number found on the legs of any one harvestman was eight. l 0. ECOLOGY: MIGRATION AND DISPERSAL 89 The maturity of the travellers shows that the habit of phoresy has no relation to the dispersal of a brood of youngsters. Vachon found that all his captures were either recently fertilized or alternatively that they had just left the brood-chamber in which they had reared and fed their family. In either case they were in urgent need of food, and he concluded that hunger had made them seize the legs of flies or harvestmen, and that as a result they reached areas where food might be plentiful. An alternative view, which has the support of the authority of Muchmore ( 1971) is that the false scorpion grasps the fly's or harvestman's leg in its search for f(>od, and that the bearer moves on before the passenger lets go. The consequent dispersal is therefore only accidental. A word should be added concerning plain, unspecializcd wanderings. The impression that may be given by the last few paragraphs, or by the casual statement that Arachnida can escape from their kin by merely running avvay and seeking fresh fields, is one that can easily be exaggerated. A reader may think of such an animal as one that has no home, and which, continuously, may find itself many miles from its birthplace. This does not, in happen. Entomologists who have painted coloured spots on ordinary house-flies, so that they can be recognized if they are caught find that flights of a kilometre or so are the exception. Even the wolf spiders that one secs in a meadow arc likely to have been born in it and are as likely to die in it: the individual, in fact, lives normally in a circumscribed area over which it hunts and \\·here it is too busily occupied in daily life to have the opportunity for foreign travel. This is conspicuous when wolf spiders arc kept under observation in the laboratory. In a large round pneumatic trough a solitary spider has an area of about 2,600 cm 2 of sand and gravel, where it energetically hunts and catches any insects that may be offered to it. And somewhere in this field it has found a spot, under a stone or between two stones, which, though to our eyes no different from other spots, has attracted it and has become its "home". Here it usually rests when not on the prowl, and here it returns after its forays, displaying a homing instinct or mnemotaxis. This description of the act of homing should be compared with the account of the function of the lyriform organs, given in Chapter 3. :\figration in such ways as these is so largely dependent on chance that it is not inevitably followed by survival in the new surroundings. The migrant must be adapted to its new circumstances, and the methodical study of the relations between the organism and all the features of its environment has grown into the established science of Ecology. This may be said to present two sides, as autecology and synecology.

90 11. DE ARACHNIDIS Autecology concentrates attention on the lives of the separate species and on their reactions to physical conditions. These include the preYalent temperature and its range of variation, the humidity of the atmosphere, the intensity of the light, the velocity of the winds and the chemical constitution of the soil; all of which are together described as the edaphic factors. Synccology is concerned more with the community or population than with the individual. Thus it takes account of predators and parasites, of food chains, and of changes in the density of population during the year. These arc known collectively as the biotic factors. Clearly it is less elementary than autecology, a characteristic of which is that the effects of physical conditions can often be most precisely determined by experimental study in the laboratory. The synecologist faces a more complex situation, which is most readily approached by a convenient division of the biosphere into three layers. (i) The Gound Layer: the moss and litter layer, up to about 15 cm above the surface. (ii) The Field Layer: the plant and shrub layer, up to about l m above the surface. (iii) The Canopy: the tree layer of trunk and branches, as well as walls and the outsides of buildings. The lowest of these layers is the home of many more individual animals than either of the others, for in there are to be found not only those that are actually underground, but also the vast assembly known collectively as the Cryptozoa. These are the animals that lead lives in conditions that are continuously dark and damp, and in which conditions are less changeable than elsewhere. This constancy is favourable to most Arachnida, and in fact Lawrence ( 1953) estimated that about half the cryptozoa belong to this class. Every field naturalist knows that the sifting of leaf-litter yields spiders, mites, false scorpions and harvestmen, while in the right parts of the world there may also be Palpigradi, Schizomida and Ricinulei. A conspicuous feature of all cryptozoa is the primitive nature of many of their number, relating them to their distant ancestors, who long ago crept from the primaeval ocean. They found in the cryptosphere an environment in which survival was possible for small animals with thin exoskeletons, and tactile and chemotactic sense organs that made blindness of small consequence. Life in the cryptosphere was serene and there was little temptation to leave it and to evolve into more specialized forms. From this arises the statement that whereas the vertebrate palaeontologist looks for fossils the invertebrate palaeontolo- 10. ECOLOGY: MIGRATION AND DISPERSAL 91 gist looks for cryptozoa. The cryptozoic lives of the Palpigradi and Schizomida are examples of this. On the other hand, spiders are among the most highly specialized of the Arachnida, and among the cryptozoic spiders the families Linyphiidae and Erigonidae are prominent. They illustrate the advantages of small size. In addition there are many spiders that may be found in the soil itself. Estimates of their number have ranged from 11,000 to 600,000 spiders per acre. And there are some spiders which, though normally living on the surface, have adopted a habit of burying themselves and thus securing protection. Reiskind ( 1965) has described the steps by which a species of Sicarius digs a hole in sand with its forelegs, enters, and throws sand over itself until it is covered. An environment that may be considered as an extension or modification of the cryptosphere is that of the cave. A cave offers the same conditions of darkness and humidity, and the further a cave may extend from its entrance, the less are these conditions likely to change. The occupants of caves include a proportion of Arachnida, which share with members of other classes the features that may be attributed to the environment. The high relative humidity, which often reaches lOO%, results in a slower metabolic rate. Since oxidation helps to produce melanin, cavedwelling fauna are often pale or colourless. For the same reason, they are often sluggish in movement and less aggressive in behaviour; and this is associated with the reduction in the proportion of chitin in the exoskeleton and also with an increased length of body and legs. The blindness, which is common, is sometimes attributed to the loss of eyes owing to disuse, or alternatively to the fact that organisms, eyeless from some other cause, are at no consequent disadvantage in a cave. Probably the most interesting relation between caves and Arachnida is shown by the Ricinulei. Long regarded as the rarest representatives of their class, these animals have been found by l'vlitchell ( 1970) to be living in their thousands in the caves of Mexico. In the words of one arachnologist, "They were so numerous that it was hard to avoid treading on them". To this there should be added the report by Mitchell ( 1968) of the discovery of three eyeless scorpions, also in Mexican caves. For these, the first blind scorpions recorded, a new genus, Typhlochactas, and a new sub-family were necessary. The species are described as being some 15 mm long, smooth and weakly sclerotized, the exoskeleton so translucent that the alimentary canal could be seen within. The occurrence of such unusual scorpions, at the considerable height of about l ,400 m poses interesting problems in the evolution of the order. The Arachnida that live in the upper, field and canopy layers,

90 11. DE ARACHNIDIS<br />

Autecology concentrates attention on the lives of the separate species<br />

and on their reactions to physical conditions. These include the<br />

preYalent temperature and its range of variation, the humidity of the<br />

atmosphere, the intensity of the light, the velocity of the winds and<br />

the chemical constitution of the soil; all of which are together described<br />

as the edaphic factors. Synccology is concerned more with the community<br />

or population than with the individual. Thus it takes account<br />

of predators and parasites, of food chains, and of changes in the density<br />

of population during the year. These arc known collectively as the<br />

biotic factors. Clearly it is less elementary than autecology, a characteristic<br />

of which is that the effects of physical conditions can often be most<br />

precisely determined by experimental study in the laboratory.<br />

The synecologist faces a more complex situation, which is most<br />

readily approached by a convenient division of the biosphere into three<br />

layers.<br />

(i) The Gound Layer: the moss and litter layer, up to about<br />

15 cm above the surface.<br />

(ii) The Field Layer: the plant and shrub layer, up to about l m<br />

above the surface.<br />

(iii) The Canopy: the tree layer of trunk and branches, as<br />

well as walls and the outsides of buildings.<br />

The lowest of these layers is the home of many more individual<br />

animals than either of the others, for in there are to be found not only<br />

those that are actually underground, but also the vast assembly known<br />

collectively as the Cryptozoa. These are the animals that lead lives in<br />

conditions that are continuously dark and damp, and in which conditions<br />

are less changeable than elsewhere. This constancy is favourable<br />

to most <strong>Arachnida</strong>, and in fact Lawrence ( 1953) estimated that about<br />

half the cryptozoa belong to this class. Every field naturalist knows that<br />

the sifting of leaf-litter yields spiders, mites, false scorpions and harvestmen,<br />

while in the right parts of the world there may also be Palpigradi,<br />

Schizomida and Ricinulei.<br />

A conspicuous feature of all cryptozoa is the primitive nature of many<br />

of their number, relating them to their distant ancestors, who long ago<br />

crept from the primaeval ocean. They found in the cryptosphere an<br />

environment in which survival was possible for small animals with thin<br />

exoskeletons, and tactile and chemotactic sense organs that made<br />

blindness of small consequence. Life in the cryptosphere was serene<br />

and there was little temptation to leave it and to evolve into more<br />

specialized forms. From this arises the statement that whereas the<br />

vertebrate palaeontologist looks for fossils the invertebrate palaeontolo-<br />

10. ECOLOGY: MIGRATION AND DISPERSAL 91<br />

gist looks for cryptozoa. The cryptozoic lives of the Palpigradi and<br />

Schizomida are examples of this.<br />

On the other hand, spiders are among the most highly specialized of<br />

the <strong>Arachnida</strong>, and among the cryptozoic spiders the families Linyphiidae<br />

and Erigonidae are prominent. They illustrate the advantages of<br />

small size. In addition there are many spiders that may be found in<br />

the soil itself. Estimates of their number have ranged from 11,000 to<br />

600,000 spiders per acre. And there are some spiders which, though<br />

normally living on the surface, have adopted a habit of burying themselves<br />

and thus securing protection. Reiskind ( 1965) has described the<br />

steps by which a species of Sicarius digs a hole in sand with its forelegs,<br />

enters, and throws sand over itself until it is covered.<br />

An environment that may be considered as an extension or modification<br />

of the cryptosphere is that of the cave. A cave offers the same<br />

conditions of darkness and humidity, and the further a cave may extend<br />

from its entrance, the less are these conditions likely to change.<br />

The occupants of caves include a proportion of <strong>Arachnida</strong>, which share<br />

with members of other classes the features that may be attributed to the<br />

environment.<br />

The high relative humidity, which often reaches lOO%, results in a<br />

slower metabolic rate. Since oxidation helps to produce melanin, cavedwelling<br />

fauna are often pale or colourless. For the same reason, they<br />

are often sluggish in movement and less aggressive in behaviour; and<br />

this is associated with the reduction in the proportion of chitin in the<br />

exoskeleton and also with an increased length of body and legs. The<br />

blindness, which is common, is sometimes attributed to the loss of eyes<br />

owing to disuse, or alternatively to the fact that organisms, eyeless<br />

from some other cause, are at no consequent disadvantage in a cave.<br />

Probably the most interesting relation between caves and <strong>Arachnida</strong><br />

is shown by the Ricinulei. Long regarded as the rarest representatives<br />

of their class, these animals have been found by l'vlitchell ( 1970) to be<br />

living in their thousands in the caves of Mexico. In the words of one<br />

arachnologist, "They were so numerous that it was hard to avoid treading<br />

on them".<br />

To this there should be added the report by Mitchell ( 1968) of the<br />

discovery of three eyeless scorpions, also in Mexican caves. For these, the<br />

first blind scorpions recorded, a new genus, Typhlochactas, and a new<br />

sub-family were necessary. The species are described as being some<br />

15 mm long, smooth and weakly sclerotized, the exoskeleton so translucent<br />

that the alimentary canal could be seen within. The occurrence<br />

of such unusual scorpions, at the considerable height of about l ,400 m<br />

poses interesting problems in the evolution of the order.<br />

The <strong>Arachnida</strong> that live in the upper, field and canopy layers,

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