This work records an investigation of the anatomy of the cranial blood supply in various species of mammal. The findings were based on a study of radiographs of specimens injected with a radioopaque mass and of casts of the vessels made by injection of neoprenc latex, and on dissections of injected preparations. A well-developed carotid rete was found to be present in the cat, sheep, goat, ox and pig, and a rudimentary form of this structure was found in the dog. In the cat the rete is situated extracranially, but in the other species it lies within the cranium in the cavernous sinus. There is no carotid rete in the rabbit or the rat. The carotid rete consists of a compact network of intertwined, freely anastomosing arteries, and is related to the branches of the trigeminal nerve. When the rete is situated intracranially there is a variable degree of communication across the mid-line with the rete of the opposite side. Whether situated intracranially or extracranially the rete lies within a venous lake. The presence of a well-developed carotid rete was associated with the non-persistence of, or a degenerating internal carotid artery. A thin fibrous cord was the only remnant of this artery found in the cat. In the goat, sheep and pig the internal carotid artery was absent proximal to the rete, but the large trunk which connected the rete with the circle of Willis was identified as representing the still persisting distal segment of this artery. In the ox (a young animal) a similar trunk connecting the rete with the circle of Willis formed the continuation of a still persisting but relatively narrow proximal segment of the internal carotid artery. The main vessels which may supply the carotid rete are the internal maxillary artery (usually via the ramus anastomoticus and the arteria anastomotica), the ascending pharyngeal artery, and the occipital artery. In the cat, sheep, goat and ox the chief vessel of supply is the internal maxillary artery, but in the pig it is the ascending pharyngeal artery. In the ox a substantial contribution is provided by the occipital artery. Both the ramus anastomoticus and the arteria anastomotica form connexions between the external and internal carotid systems and they join one another within the cavernous sinus. It is at his site that the intracranial carotid rete is developed, but the extracranial carotid rete, seen in the cat, is situated around the internal maxillary artery at the site of origin of the arteria anastomotica of the artiodactyls. The variations in the situation of the carotid rete in different species is along the line of the arteria anastomotica. It is suggested that the arteria anastomotica and the ramus anastomoticus respectively are homologous with the recurrent meningeal and the middle meningeal arteries of the rabbit and man. The arteries which supply the orbital and the ethmoidal regions are described and the homologies of the external ophthalmic artery are discussed. The great variability in the supply of these territories which was seen in the cat is thought to be associated with the presence of an extracranial carotid rete. In spite of variations in different species of animal a basic pattern can be discerned in the major arteries supplying the head. It is suggested that this basic pattern is related to a primitive stapedial artery, and that the variations seen are due to modifications of the branches of this earlier vessel. The circle of Willis was found to derive its blood supply from one or more of five sources: the internal carotid artery, the external carotid arterial system, the ascending pharyngeal artery, the vertebral artery (via the basilar artery) and the occipital artery. In the absence of an internal carotid artery the greatest contribution of blood passes to the circle of Willis through the carotid rete. An occipito-vertebral anastomosis seems to be of some importance in supplying the circle of Willis in the cat, pig, dog and rabbit. In the sheep, goat and ox the direction of the flow of blood in the basilar artery would appear to be away from and not towards the circle of Willis. In the pig the two anterior cerebral arteries anastomose in the mid-line and continue forward as a single vessel. The peculiar structure of the vessels which compose the carotid rete suggests that this compact network has a haemodynamic significance, and since the rete lies in the pathway of the major artery or arteries which supply the brain, its existence and possible influence should be borne in mind when problems of the cerebral circulation are considered in species in which this structure is present.