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Left: male, sharp tooth above slope at end of each wing case. Right: female, steep slope at end of wing cases densely covered with yellow setae. Left: male, wing cases tapered to rear.

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Right: female, front of head concave. Platypus gracilis - width just under 1 mm, five times longer then wide. Right: female, rounded swelling above slope at end of each wing case. The eggs are cream, shining, 0. They are found at tunnel ends. The larvae pass through five developmental stages. Youngest larvae are flattened and flounder-like, and bear fleshy finger-like projections of the sides and upper surfaces. They are able to move rapidly along the tunnel walls with an undulating motion.

In later stages as the body grows large enough to fill the tunnel it becomes cylindrical and the juvenile projections disappear. Fully grown larvae Fig. The mouthparts are directed downwards, and the jaws are dark brown to black. On each side of the upper surface of the prothorax there is a brown hardened structure which consists of two more or less complete circular ridges linked behind by a transverse ridge, so that the whole resembles a pair of spectacles.

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Such a structure is lacking in earlier stages of larval development. The end of the abdomen may be smooth or may have a minute pointed spine, or a low hardened ridge, or a blunt tooth.

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Top: Platypus apicalis - width just under 2 mm; small spine at end of abdomen. Larvae at all stages move freely throughout the nest. Pupae are found only in pupal chambers, nearly always with heads directed towards the transverse tunnel from which the chambers arise. Life histories and habits Most beetles emerge during the warmer months, though a few may appear on warmer days even in mid-winter. There are at least three kinds of attractants which guide beetles to their hosts. Males of all three species are attracted to dying or freshly felled trees and stumps by volatile substances given off by the stressed tissues.

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These males then emit an attractant of their own probably produced in the hind gut which causes other males and also females to aggregate at the same place. The beetles die before they have penetrated more than a few centimetres. Similar attraction to rapid growth occurs in silver beech, probably accounts for some of the abortive attacks on western balsam poplar and coast redwood, and is very likely of much wider occurrence.

Temporary stress caused by drought or damage to roots may also result in attack on living trees of any species which, if felled or dying, would be utilised by the beetles. Once such stress is relieved, gums or resins soon flood the Platypus tunnels and attractants cease to be emitted.

However, if the tree is susceptible to the introduced pathogenic fungus the sapwood is progressively killed, passage of water from soil to crown is restricted, stress is intensified, further attack occurs, and the tree ultimately dies. Males have been seen to start nests in a red beech log felled 20 years previously, and from which the rotted sapwood had fallen away. Nests containing fully grown larvae have also been found in the stump of a large red beech felled 25 years previously. Selection of a concealed site for the nest entrance is not evident in the other two species.

The males of all species bore radially through the bark and outer sapwood. A circular groove is cut with the tips of the jaws and strands of wood are then seized at one end and torn away from the tunnel face, the strands being passed backwards beneath the beetle and ultimately ejected at the tunnel entrance. The males do not come right to the entrance while frass is being expelled.

Once the male has excavated a sufficient length of tunnel he may be joined by a female. After copulating at the entrance she precedes her mate into the tunnel and takes over the task of extending the nest, the male subsequently ejecting the frass and, at times, excess fungal growth. As the initially radial tunnel continues through the inner sapwood it curves through a right angle and then follows a path close to the boundary between sapwood and heartwood.

When no heartwood is present - e. If either of the pairs should die during the early stages of nest development, the nest fails; if they die at later stages few brood subsequently emerge.

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If the male has been lost, live larvae may fall from the entrance. So, though it is impossible to observe it in detail, parental tending is of great importance to the brood, and social behaviour patterns are well developed. When the nest has been sufficiently developed or perhaps at a given time after copulation , the female lays her first batch of eggs - usually of four to seven - near the end of the tunnel. The presence of the egg batch evidently inhibits further excavations, and a branch in the opposite direction is then started from the curved portion.

This second tangential arm is continued until she lays her second batch of eggs, after which the expulsion of fibrous frass ceases. Development of nests to this stage may take months. In the most recently formed parts of the nest a shining coating soon appears on the tunnel walls.

The yeasts bud profusely on the walls of tunnels and develop strands hyphae which penetrate a short distance into fresh wood which has not been exploited by other fungi. Larvae have been seen to spread some unknown materials on the walls of such parts of the nest with their mouthparts which include rake-like structures, reminiscent of a simple adhesives spreader. Evidently they cultivate this food resource, and do not simply eat whatever happens to appear. In older parts of the nest the initial coating of yeasts is succeeded by a black fungal growth.

While fibrous frass is being ejected much of this growth is abraded, and shortly after adult tunnelling ceases the males scrape excess growth from the walls and eject it as black slimy blobs. Such nest clearings appear periodically from the time adult frass ceases, and are often the last sign of activity after the brood have all emerged.

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It is unknown whether this black fungal growth plays any part in nutrition. In December, January, or February occasionally earlier or later after the cessation of adult frass, a different type of woody frass begins to appear. This is granular, and consists of minute, tightly coiled wood chips cut by the fully grown larvae as they begin to extend the parent tunnels or to make new branches. The ends of the tunnels made by the larvae are concave, whereas those made by the parents are flat. These larval extensions of the nest control the numbers that can be reared, because they extend into previously uninvaded wood suitable for the cultivation of more yeasts, and provide more tunnel ends at which egg batches may be deposited.

As well as extending the nest, mature larvae make groups of short blind tunnels Fig. Groups of pupal chambers usually occur above and below branch tunnels near the junction of the tangential arms and the entrance tunnel. The openings of the upper and lower series are never quite opposite. For larvae which develop from the first egg batches, attainment of the final larval stage is usually followed by a long period of nest extension activity before they pupate; in unfavourable host material, where competing fungi leave no uninvaded wood for yeast cultivation, nest extensions serve no useful purpose and the larvae evidently pupate much sooner.

All three species produce, overall, approximately equal numbers of males and females, though individual nests may produce a marked predominance of either sex. Such a spread of emergence occurs from nests in relatively stable material. Those in hosts which undergo more rapid biodegradation produce few brood after the first flight season.

Though observations were discontinued it seemed clear that some nests could continue to be productive.

During this period the maximum number of brood per pair of beetles was , and the mean for 40 nests was In all three species it is usual to find that twice as many nests produce fewer brood than the mean; a high proportion of the total brood appear from rather few, unusually productive nests. Control No insect parasites or predators are normally found in Platypus nests. Those factors which cause the failure of attempted nests - particularly gum and resin flows when living trees are attacked, and partial drying followed by invasion of the wood by a succession of fungi in dead hosts - are probably the most important in limiting populations.

No control procedures have been attempted in native forests. Traditional selection logging practices, and experimental thinning of pole stands have, indeed, caused populations to reach high levels locally.