The evolution of the dorsal pallium in the telencephalon of amniotes: cladistic analysis and a new hypothesis.

The large body of evidence that supports the hypothesis that the dorsal cortex and dorsal ventricular ridge of non-mammalian (non-synapsid) amniotes form the dorsal pallium and are homologous as a set of specified populations of cells to respective sets of cells in mammalian isocortex is reviewed. Several recently taken positions that oppose this hypothesis are examined and found to lack a solid foundation. A cladistic analysis of multiple features of the dorsal pallium in amniotes was carried out in order to obtain a morphotype for the common ancestral stock of all living amniotes, i.e., a captorhinomorph amniote. A previous cladistic analysis of the dorsal thalamus (Butler, A.B., The evolution of the dorsal thalamus of jawed vertebrates, including mammals: cladistic analysis and a new hypothesis, Brain Res. Rev., 19 (1994) 29-65; this issue, previous article) found that two fundamental divisions of the dorsal thalamus can be recognized--termed the lemnothalamus in reference to predominant lemniscal sensory input and the collothalamus in reference to predominant input from the midbrain roof. These two divisions are both elaborated in amniotes in that their volume is increased and their nuclei are laterally migrated in comparison with anamniotes. The present cladistic analysis found that two corresponding, fundamental divisions of the dorsal pallium were present in captorhinomorph amniotes and were expanded relative to their condition in anamniotes. Both the lemnothalamic medial pallial division and the collothalamic lateral pallial division were subsequently further markedly expanded in the synapsid line leading to mammals, along with correlated expansions of the lemnothalamus and collothalamus. Only the collothalamic lateral pallial division--along with the collothalamus--was subsequently further markedly expanded in the non-synapsid amniote line that gave rise to diapsid reptiles, birds and turtles. In the synapsid line leading to mammals, an increase in the degree of radial organization of both divisions of the dorsal pallium also occurred, resulting in an 'outside-in' migration pattern during development. The lemnothalamic medial division of the dorsal pallium has two parts. The medial part forms the subicular, cingulate, prefrontal, sensorimotor, and related cortices in mammals and the medial part of the dorsal cortex in non-synapsid amniotes. The lateral part forms striate cortex in mammals and the lateral part of dorsal cortex (or pallial thickening or visual Wulst) in non-synapsid amniotes. Specific fields within the collothalamic lateral division of the dorsal pallium form the extrastriate, auditory, secondary somatosensory, and related cortices in mammals and the visual, auditory, somatosensory, and related areas of the dorsal ventricular ridge in non-synapsid amniotes.