decompose wood also comes from the Late Devonian. The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by rain. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper waters, increasing the extinction rate among deep-water animals. Land invertebrates Animals had to change their feeding and excretory systems, and most land animals developed internal fertilization of their eggs. The difference in refractive index between water and air required changes in their eyes. On the other hand, in some ways movement and breathing became easier, and the better transmission of high-frequency sounds in air encouraged the development of hearing. The oldest known air-breathing animal is Pneumodesmus, an archipolypodan millipede from the Middle Silurian, about . Its air-breathing, terrestrial nature is evidenced by the presence of spiracles, the openings to tracheal systems. However, some earlier trace fossils from the Cambrian-Ordovician boundary about are interpreted as the tracks of large amphibious arthropods on coastal sand dunes, and may have been made by euthycarcinoids, which are thought to be evolutionary "aunts" of myriapods. Other trace fossils from the Late Ordovician a little over probably represent land invertebrates, and there is clear evidence of numerous arthropods on coasts and alluvial plains shortly before the Silurian-Devonian boundary, about , including signs that some arthropods ate plants. Arthropods were well pre-adapted to colonise land, because their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. The fossil record of other major invertebrate groups on land is poor: none at all for non-parasitic flatworms, nematodes or nemerteans; some parasitic nematodes have been fossilized in amber; annelid worm fossils are known from the Carboniferous, but they may still have been aquatic animals; the earliest fossils of gastropods on land date from the Late Carboniferous, and this group may have had to wait until leaf litter became abundant enough to provide the moist conditions they need. The earliest confirmed fossils of flying insects date from the Late Carboniferous, but it is thought that insects developed the ability to fly in the Early Carboniferous or even Late Devonian. This gave them a wider range of ecological niches for feeding and breeding, and a means of escape from predators and from unfavorable changes in the environment. About 99% of modern insect species fly or are descendants of flying species.