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Pennaceous feathers. This type of feather is present in most modern birds and has been shown in some species of maniraptoriform dinosaurs. A pennaceous feather has a stalk or quill. Its basal part, called a calamus, is embedded in the skin. The calamus is hollow and has pith formed from the dry remains of the feather pulp, and the calamus opens below by an inferior umbilicus and above by a superior umbilicus. The stalk above the calamus is a solid rachis having an umbilical groove on its underside. Pennaceous feathers have a central shaft (or rachis) with vanes or vaxillum spreading to either side. These vanes are composed of a high number of flattened barbs, that are connected to one another with barbules.
Pennaceous feathers on the forelimb or wing are often attached strongly due to stresses related to flight or other activities. This strong attachment is accomplished by ligaments under the skin, which in some birds and feathered dinosaurs results in raised bumps or marks along the rear forelimb bone (ulna). These bumps, called "quill knobs" (ulnar papillae) are often used as an indirect indication of strongly-attached forelimb feathers in fossil species, and can also indirectly indicate the number of secondary regimes in a given specimen.
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