Collins New Naturalist Library. M. Brian V.

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is known so far it appears that they can detect other animals from a distance of a few centimetres. Thus wood ants are known to be able to see movements 10 cm away. In soil spaces and perhaps in foliage ants may be able to detect the vibrations made by small animals moving through the substratum and they certainly have an acute sense of smell. Once they have received a distance signal they approach slowly, alert for others, probably mainly smells, before actually attacking. As they get nearer to the potential prey they orientate with their head forward, jaws open and antennae retracted and make further exploratory movements. Clearly, they must not throw themselves into the jaws of an enemy and a secure grip on prey is an obvious advantage. Finally they pounce and snap. If a hold is obtained, usually on a leg or antenna, they quickly bring round their gaster and inject poison with their sting if they have one, otherwise they spray penetrating or adhesive toxins. Not many ants are really alone when hunting, there are others nearby usually that help by flushing out prey or gathering round to help pin one down; they are more dog-like than cat-like on the whole.

      Many small invertebrates are easily caught and immobilized; the small larvae of flies and moths and beetles, for example. Many others, particularly adults, have escape mechanisms. Thus, springtails (Collembola), which are very much valued as food by Myrmica and Lasius, can usually escape instantly by jumping but are easily caught while they are changing their skins, stuck in a water droplet or in some way damaged. The larvae of many sawflies and moths can flick their bodies smartly, others are so hairy that ants find them difficult to get hold of and many caterpillars can simply slip away on a thread of silk; some protect themselves with a case of vegetable material. To escape, other insects kick or produce repellents or sticky exudates from special skin glands, such as the cornicles of aphids. Those with a hard, shiny cuticle, like beetles, may be difficult to grip and impossible to sting. The great variety of these defensive mechanisms make it highly probable that many prey animals are only caught when they are incapacitated, perhaps through age, perhaps by mechanical damage (such as being trodden on), by wing failure, by the necessity to moult or even by transient low temperatures or weak light (some insects can only fly in sunlight). Bugs, flies and spiders comprised 80–90% of the number of prey caught by species of Myrmica. Small spiders that live on and near the soil surface are a major constituent of this food (11–38% in different years). In Polish grassland from 1700 to 4000 spiders were caught in a year in one square metre. In May, June and July, when feeding peaks, between sixteen and seventy-four spiders may be taken each day from one square metre.

      Undoubtedly, an important element in the diet of most ants is other ants. Each summer they eat large numbers of sexuals, both of their own and other species, especially those that are unlucky enough to descend after the mating flight on to already occupied territory. Also, in spring when food is scarce and catastrophic fires destroy the vegetation, a wider search for food often leads to fighting between workers of different colonies and species in which a lot die. The corpses are taken back to the nest and sucked dry: an economical way of adjusting the population to a sharp drop in food supply.

      Information is very much needed about the food of different colonies and species at different times of year. Some ecologists have taken samples of foragers on their return to the nest and identified what they were carrying. One has recently invented an ingenious trap which has been used to study the food of the wood ant, a species with two advantages: well-defined tracks above ground and large workers. A whole nest was surrounded by a barrier soaked in repellent oil and workers were induced to pass over this on specially constructed wooden bridges. One was used for incoming and the other for outgoing traffic. This was possible because although ants are prepared to drop a few centimetres from the end of a bridge on to the soil below they cannot reach up to return the same way. So, it was only necessary to place strips with the drop outside the barrier to take outgoing traffic and the drop inside the barrier for ingoing traffic. Those returning dropped into a box from which they could only escape by small holes little bigger than their bodies and they left behind anything they had been carrying in their jaws. Perhaps some day, traps on a similar principle will be devised for ants which forage underground, but this is likely to be much more difficult.

      Wood ants eat many invertebrates which they catch both in the trees and on the ground. A large number of these are flies (including midges and crane flies (Tipula spp.)) and many kinds of aphids; also, in season, winged ants, particularly of the genera Lasius and Myrmica. A lot of these insects are forest pests and the establishment of wood ant populations has become, at least in Europe, an important part of woodland management. In years when defoliating insects are very abundant and trees are stripped of their leaves in summer, conspicuous green islands are left around areas where ants nest. These defoliators include the larvae of various moths and sawflies that feed on oaks, pine, spruce and larch. Pupae and adults are also eaten. There are indications that wood ants will attack moving things in preference to quiet, still ones; yet it was found by trapping that they collected a lot of prey at night. There are of course a great many flying insects that settle in the foliage of trees and bushes during darkness; these could perhaps be easily located by smell. A wood ants’ nest was surrounded by guttering into which they threw their refuse. This not only consisted of empty cocoon fragments as expected but of many other insects and other odd items that were collected but not eaten; evidently they take in a great many more things than they use.

      Food collection by Formica aquilonia has been studied in detail in an old Caledonian forest. Some five or six trackways leave each nest and go to trees on which prey are caught and aphids are tended for honeydew. After leaving the nest, ants pass round the perimeter and then leave on any one of the tracks. There is just a slight tendency for individuals to use the same track out as they use in. This is oddly at variance with other results which have shown taat Formica rufa and its allies come near to partitioning their foraging grounds between groups of workers that are fairly fixed in individual composition. Different species, different types of food collection or just different times of year may explain this apparent contradiction. Many of the foragers of Formica aquilonia leave the trackways to forage in the neighbouring herbs and observations show that this ‘leakage’ occurs at a constant rate and that the search for prey is quite random until very high prey densities are encountered. Then a recruitment mechanism increases the number of ants entering the area. Temperature affects the rate of flow of traffic on the trackways; a rise from 8 to 18% increases it 5 times. This is probably due to a greater availability of prey at high temperatures, as well as to a greater number of foragers joining in.

      Rate of traffic flow is also very much affected by obstacles. If the leaves and twigs are swept from a track the walking rate rises as much as 10%; at 20° C it is normally about a metre a minute. In sections where the tracks pass between rocks traffic density is often so high that collisions are frequent. This causes some delay as the ants stop to examine each other with their antennae. There seems to be very little organization of the flow near the nest: a slight tendency exists for incoming workers to move on the outside and outgoing ones in the centre of the track. In the July of the study there were about seventy thousand foragers active; one in five brought an insect back and it was estimated that about a hundred thousand insects were collected each day.

      All this information is not very well received by entomologists primarily interested in the insects which are destroyed. They claim, not without reason, that wood ants impoverish the fauna, but the interrelationships between insects are so complicated and numerous that it is more likely that they merely prevent any one type from predominating and thus preserve a richer mixture at a lower density. As has been pointed out there is plenty of evidence that they concentrate on prey that is momentarily superabundant. Birds of course do this too, and as there are many which live on insects in forests they might be expected to compete with ants. Curiously, the evidence is to the contrary and it is suggested that ants dislodge many insects whilst hunting which they lose and these are caught by birds. This sort of situation is well known in the Tropics where some birds subsist largely by collecting the prey which escapes from the devastating columns of army ants.

      The food of Lasius flavus was a mystery until quite recently but it is now known that they eat a great many soil animals, including soft-bodied mites, beetle larvae

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