Cranial morphology of Javanese Homo erectus: new evidence for continuous evolution, specialization, and terminal extinction.

Our current knowledge of the evolution of Homo during the early to middle Pleistocene is far from complete. This is not only because of the small number of fossil samples available, but also due to the scarcity of standardized datasets which are reliable in terms of landmark identification, interobserver error, and other distorting factors. This study aims to accurately describe the cranial morphological changes of H. erectus in Java using a standardized set of measurements taken by the authors from 18 adult crania from Sangiran, Trinil, Sambungmacan, and Ngandong. The identification of some obscure landmarks was aided by the use of micro-CT imaging. While recent studies tend to emphasize evolutionary conservatism in Javanese H. erectus, our results reinforce the theory that chronologically later groups experienced distinct morphological changes in a number of cranial traits. Some of these changes, particularly those related to brain size expansion, are similar to those observed for the genus Homo as a whole, whereas others are apparently unique specializations restricted to Javanese H. erectus. Such morphological specializations in Java include previously undescribed anteroposterior lengthening of the midcranial base and an anterior shift of the posterior temporal muscle, which might have influenced the morphology of the angular torus and supramastoid sulcus. Analyses of morphological variation indicate that the three crania from Sambungmacan variously fill the morphological gap between the chronologically earlier (Bapang-AG, Bapang Formation above the Grenzbank zone in Sangiran) and later (Ngandong) morphotypes of Java. At least one of the Bapang-AG crania, Sangiran 17, also exhibits a few characteristics which potentially indicate evolution toward the Ngandong condition. These strongly suggest the continuous, gradual morphological evolution of Javanese H. erectus from the Bapang-AG to Ngandong periods. The development of some unique features in later Javanese H. erectus supports the hypothesis that this Javanese lineage went extinct without making significant contributions to the ancestry of modern humans.