1.1. The earliest beetles and their systematic treatment

1.2. Reflectance Transformation Imaging

2.1. Material

2.2. Imaging

2.3. RTI dome

2.4. RTI processing

2.5. Image processing

2.6. Graphic design

2.7. Data availability

Coleopsis

Coleopsis archaica

4.1 New interpretation of morphological features

4.2. Systematic interpretation

4.3. Coleopteran autapomorphic character states documented in the fossil (features shared with other extinct and extant beetles)

4.4. Plesiomorphic character states of the fossil (features not shared with extant beetles)

4.5. Possible affinities within Coleoptera

4.6. RTI technique