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Myriapods (e. g., centipedes and millipedes) display a simple homonomous body plan relative to other arthropods. All members of the class are terrestrial, but they attained terrestriality independently of insects. Myriapoda is the only arthropod class not represented by a sequenced genome. We present an analysis of the genome of the centipede Strigamia maritima. It retains a compact genome that has undergone less gene loss and shuffling than previously sequenced arthropods, and many orthologues of genes conserved from the bilaterian ancestor that have been lost in insects. Our analysis locates many genes in conserved macro-synteny contexts, and many small-scale examples of gene clustering. We describe several examples where S. maritima shows different solutions from insects to similar problems. The insect olfactory receptor gene family is absent from S. maritima, and olfaction in air is likely effected by expansion of other receptor gene families. For some genes S. maritima has evolved paralogues to generate coding sequence diversity, where insects use alternate splicing. This is most striking for the Dscam gene, which in Drosophila generates more than 100,000 alternate splice forms, but in S. maritima is encoded by over 100 paralogues. We see an intriguing linkage between the absence of any known photosensory proteins in a blind organism and the additional absence of canonical circadian clock genes. The phylogenetic position of myriapods allows us to identify where in arthropod phylogeny several particular molecular mechanisms and traits emerged. For example, we conclude that juvenile hormone signalling evolved with the emergence of the exoskeleton in the arthropods and that RR-1 containing cuticle proteins evolved in the lineage leading to Mandibulata. We also identify when various gene expansions and losses occurred. The genome of S. maritima offers us a unique glimpse into the ancestral arthropod genome, while also displaying many adaptations to its specific life history.
The variability of the blazar Markarian 421 in TeV gamma rays over a 14-year time period has been explored with the Whipple 10 m telescope. It is shown that the dynamic range of its flux variations is large and similar to that in X-rays. A correlation between the X-ray and TeV energy bands is observed during some bright flares and when the complete data sets are binned on long timescales. The main database consists of 878.4 h of observation with the Whipple telescope, spread over 783 nights. The peak energy response of the telescope was 400 GeV with 20% uncertainty. This is the largest database of any TeV-emitting active galactic nucleus (AGN) and hence was used to explore the variability profile of Markarian 421. The tithe-averaged flux from Markarian 421 over this period was 0.446 +/- 0.008 Crab flux units. The flux exceeded 10 Crab flux units on three separate occasions. For the 2000-2001 season the average flux reached 1.86 Crab units, while in the 1996-1997 season the average flux was only 0.23 Crab units.