ants in frigid or cold temperate regions typically shed their leaves in autumn, whereas in areas with a severe dry season, some plants may shed their leaves until the dry season ends. In either case, the shed leaves may be expected to contribute their retained nutrients to the soil where they fall. In contrast, many other non-seasonal plants, such as palms and conifers, retain their leaves for long periods; Welwitschia retains its two main leaves throughout a lifetime that may exceed a thousand years. The leaf-like organs of bryophytes (e.g., mosses and liverworts), known as phyllids, differ morphologically from the leaves of vascular plants in that they lack vascular tissue, are usually only a single cell thick, and have no cuticle stomata or internal system of intercellular spaces. The leaves of bryophytes are only present on the gametophytes, while in contrast the leaves of vascular plants are only present on the sporophytes, and are associa ted with buds (immature shoot systems in the leaf axils). These can further develop into either vegetative or reproductive structures. Simple, vascularized leaves (microphylls), such as those of the early Devonian lycopsid Baragwanathia, first evolved as enations, extensions of the stem. True leaves or euphylls of larger size and with more complex venation did not become widespread in other groups until the Devonian period, by which time the carbon dioxide concentration in the atmosphere had dropped significantly. This occurred independently in several separate lineages of vascular plants, in progymnos