Landmarks are the hallmark of biological shape analysis as discrete anatomical points of correspondence. Various systems have been developed for their classification. In the most widely used system, developed by Bookstein in the 1990's, landmarks are divided into three distinct types based on their anatomical locations and biological significance. As Bookstein and others have argued that different landmark types possess different qualities, e.g., that Type 3 landmarks contain deficient information about shape variation and are less reliably measured, researchers began using landmark types as justification for selecting or avoiding particular landmarks for measurement or analysis. Here, we demonstrate considerable variation in landmark classifications among 17 studies using geometric morphometrics (GM), due to disagreement in the application of both Bookstein's landmark typology and individual landmark definitions. A review of the literature furthermore shows little correlation between landmark type and measurement reproducibility, especially when factors such as differences in measurement tools (calipers, digitizer, or computer software) and data sources (dry crania, 3D models, or 2D images) are considered. Although landmark typology is valuable when teaching biological shape analysis, we find that employing it in research design introduces confusion without providing useful information. Instead, researchers should choose landmark configurations based on their ability to test specific research hypotheses, and research papers should include justifications of landmark choices along with landmark definitions, details on landmark collection methods, and appropriate inter- and intra-observer analyses. Hence, while the landmarks themselves are crucial for GM, we argue that their typology is of little use in applied studies. This article is protected by copyright. All rights reserved.
