The pollination process of the extremely long‐spurred orchids Angraecum sesquipedale and A. sororium is described and documented here for the first time. The pollinaria and viscidia load was examined in moths captured in central and south Madagascar. Visits to orchids by hawkmoths were rarely observed in the field and were therefore systematically recorded in large flight tents using a night‐vision video technique and flashlight photography. Angraecum sesquipedale in Fort Dauphin is pollinated by Xanthopan morgani praedicta and A. sororium on Mt. Angavokely by Coelonia solani. By combining a deep nectar spur of extraordinary length with a protruding labellum functioning as a landing platform, these orchids overcome the moth's stereotypic swing‐hovering flight thus enabling full insertion of the long tongue. Angraecum compactum in Forêt d'Ambohitantely is pollinated by both the shorter and longer‐tongued forms of Panogena lingens which never swing‐hover but is also exploited by X. morgani and C. solani with wastage of pollinaria. The duration of tongue insertion, nectar exploitation and tongue withdrawal were analyzed: legitimate and illegitimate visitors differ in their time budget and approach to the flower. Nectar volume, nectar level and sugar concentration of A. sesquipedale and A. sororium were compared with the nectar requirements of the pollinating hawkmoths. The evolution of very long spurs in these orchids is likely to have involved a series of pollinator shifts. The orchids adapted to different hawkmoth species with increasingly long tongues which primarily evolved to avoid predator attacks during visits to less specialized flowers. This “pollinator shift” model modifies the classical “coevolutionary race” model. The relevance of the taxon Angraecum bosseri Senghas is questioned.

SUMMARY Heartbeat activity in tethered adult drosophilids was recorded using a linear optosensor chip and an IR-light beam. Recording from two to five sensor elements within 250 μm along the anterior heart, it was possible to analyze periodic reversals. In intact Drosophila melanogaster and D. hydei, longer anterograde pulse periods with lower pulse rates generally alternated with shorter retrograde pulse periods having higher pulse rates. These differences are dependent on heart anatomy: a newly discovered first pair of ostia is connected to bilateral thoraco-abdominal hemolymph channels. These channels are part of a venous space separated from the abdominal hemocoel by a septum, consisting of a metanotal ridge and the pericardial diaphragm lined by a special form of fat body. The channels are sealed, and their lumen is possibly controlled by the metathoracic tergo-pleural muscle. During retrograde pulses, the heart chamber works like a suction pump,aspiring hemolymph through the first ostia from the venous channels and discharging it through a newly described caudal opening. During forward beating, the anterior chamber receives hemolymph via all inflow ostia from the entire heart and drives it like a pressure pump through the narrow aorta. Also, during forward pulses, a lateral circulation occurs in the thorax as a result of the venous supply. Inhibition of abdominal mobility leads to an irregular heart rate, with pulse-wise alternating heartbeat reversals. The possible involvement of slow abdominal movements in heartbeat periodicity is discussed. The heartbeat periods are superimposed with intermittent bouts of abdominal pumping movements.

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In this review we provide a detailed description of Darwin's prediction of the coevolution of a long-spurred orchid, Angraecum sesquipedale, and a long-tongued moth, his correspondence on the subject, the history of the moth and the subsequent literature. On seeing the long spur of A. sesquipedale, Darwin predicted that its pollinator would be a moth with a long proboscis. For more than a century following Darwin's prediction this was assumed to be the case. The pollinator was taken to be Xanthopan morganii praedicta, despite the fact that it had not been observed to visit A. sesquipedale flowers. Direct observations, infra-red cinematography and photographs published between 1993 and 1997 and a video made in 2004, all of which show X. morganii praedicta visiting A. sesquipedale flowers and removing pollinia, proved that Darwin's prediction was accurate. Recent research suggests that selection pressure exerted by predators on the pollinators, resulted in the evolution of extreme tongue lengths and a special hovering flight.

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