Morphology of the prothoracic discs and associated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae)
Introduction
Acanthocnemus nigricans is a small dark beetle having a body length of only 3–6 mm. The species nigricans is the only recent species within the genus Acanthocnemus and is distributed all over Australia. Since the middle of the 19th century A. nigricans has also been found on other continents like Asia, Africa and Europe (Lawrence and Britton, 1994). Most probably, the reason for this recent expansion all over the world is the commercial export of Australian wood or cereals infested with A. nigricans. Therefore, the species can be also regarded as a cosmopolitan to warmer parts of the Old World (Champion, 1922). In contrast to its inconspicuous appearance, A. nigricans shows a remarkable behaviour as beetles of both sexes are attracted by forest fires. The beetles invade a freshly burnt area immediately and approach areas where glowing remnants of trees or hot ashes are still present (Champion, 1922, Schmitz and Schmitz, 2002). The beetles often land very close to spots with still high surface temperatures. After a short period of quickly running around, the beetles hide in small openings in the ash or under the bark of burnt trees. The reason for this so-called pyrophilous behaviour is unknown but it can be speculated that the described ‘hot spots’ serve as meeting places for the sexes. Most probably, the females deposit their eggs into the ash or under the bark of burnt trees. Currently, the larval food is unknown. Because it is reasonable to suppose that larvae were exported out of Australia inside wood (see above), it is very probable that the cambium layer of freshly burnt trees represents the primary source of food for the first instars.
In two pyrophilous genera of buprestid beetles, Melanophila and Merimna, unique extraantennal sensory organs have been found which are used in fire detection. Beetles of the genus Melanophila have two pairs of metathoracic infrared (IR) organs which contain about 70 arch-shaped photomechanic IR sensilla (Evans, 1966a, Evans, 1966b, Vondran et al., 1995, Schmitz et al., 1997). The Australian ‘fire-beetle’ Merimna atrata has one to three pairs of abdominal IR receptors (Schmitz et al., 2001, Mainz et al., 2004) which are innervated by thermoreceptive multipolar neurones (Schmitz and Trenner, 2003). Therefore, the receptors function as microbolometers. However, the prothoracic sensory organs in A. nigricans are located on a different part of the body and, moreover, are fundamentally different from the IR receptors of the pyrophilous buprestid beetles. A. nigricans possesses two prothoracic sensory organs which are located directly anterior to the coxae of the prothoracic legs. Each of the organs consists of a tiny cuticular disc (diameter 110–185 μm), which is held by a lateral cuticular stalk above an air-filled cavity. First morphological investigations have already shown that the disc is filled with the somata of many sensory cells, which contain large numbers of mitochondria. By these findings, a sensory function of the disc was made very probable (Schmitz et al., 2002). Additionally, on the outer surface of the disc, small pegs were discovered which represent the cuticular apparatus of sensilla. In the present paper, we describe the morphology of the cuticular sensilla in detail. The possible role of the prothoracic sensory organ in fire detection is discussed.
Section snippets
Animals
Adult beetles were collected in 2002 and 2003 on freshly burnt areas in Western Australia. Animals were kept for several weeks in plastic boxes and fed with raisins, peanuts and walnuts; drinking water was given ad libitum.
Scanning electron microscopy
Twelve beetles were air-dried and glued on holders by Leit-Tabs (Plano, diameter 12 mm) with either the dorsal or the lateral side. Sensory discs from seven beetles were isolated and cleaned by sonication in a mixture of chloroform/ethanol (2:1) for 2 min. Discs were air-dried
The prothoracic sensory organs
One pair of unusual organs is located on the prothorax in both sexes. The two organs are situated directly anterior to the coxae of the prothoracic legs (Fig. 1(A) and (B)). The most striking component of an organ is a more or less round cuticular disc, which is somewhat sunken into the surface of the prothorax. At its posterior edge, the disc is held by a stalk which originates at the dorso-anterior border of the coxal cavity (Fig. 2(A) and (B)). When analysing the composition of the cuticle
Classification and proposed function of the sensilla
It is generally accepted that the bauplan of the cuticular apparatus and its interaction with the sensory dendrite is highly modality-specific in an insect sensillum (Steinbrecht et al., 1989). Therefore, the detailed study of the morphology of a sensillum gives valuable insight into its possible function, e.g. chemoreception, mechanoreception or thermo-/hygroreception. When analysing the bauplan of the small sensilla found on the sensory disc (in the following called ‘disc sensilla’) it turned
Acknowledgements
Mike Cantelo and Brian Inglis from CALMfire, Perth, WA, enabled us to visit bush fires for the collection of beetles and supported us with all necessary equipment. Karl-Heinz Apel and Tobias Mainz helped us to collect the beetles. The Department of Environment and Heritage (Canberra) and the Wildlife Branch of CALM (Perth) issued the collecting and export permits. We are grateful to the NEES Institute for Biodiversity of Plants at the University of Bonn for the use of the SEMs. Mareike Rüdinger
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