Article: Bennettitalean leaf cuticle fragments (here Anomozamites and Pterophyllum) can be used interchangeably in stomatal frequency-based palaeo-CO2

Bennettites are an abundant and frequently well-preserved component of many Mesozoic fossil floras, often playing an important ecological role in flood plain vegetation communities. During a recent study focusing on stomatal indices of Triassic–Jurassic fossil plants, it became evident that the leaf fragments of two bennettite genera Anomozamites Schimper (1870) emend. Harris (1969) and Pterophyllum Brongniart (1825) display a significant overlap of leaf shape as well as cuticular characters. Owing to the preference of recognition of single taxa (ideally species) for the stomatal method, we use a database of 70 leaf fragments of Anomozamites and Pterophyllum compressions from five isotaphonomic Late Triassic sedimentary beds of Astartekløft in East Greenland to test whether leaf and cuticle fragments of the two genera can be separated using a range of quantitative and qualitative morphological and statistical analyses. None of the observed characters – including stomatal frequencies – could be applied to separate the fragments of the two genera into well-defined groups. Our results therefore indicate that fragmented material and dispersed cuticles cannot be utilized to distinguish between Anomozamites or Pterophyllum at the genus level, but that instead these cuticle fragments may be used interchangeably as stomatal proxies. Classification of fossil leaves into either of these genera is thus only possible given adequate preservation of macro-morphology and is not possible based solely on cuticle morphology. We suggest that this large inter- and intra-generic morphological variation in both leaf and cuticle traits within Anomozamites and Pterophyllum may be related to the bennettites’ role as understory plants, experiencing a range of micro-environmental conditions, perhaps depending mainly on sun exposure. Based on the results obtained in this study, we conclude that Anomozamites and Pterophyllum cuticle fragments can be employed interchangeably in palaeo [CO2] reconstructions based on the stomatal method, thus potentially annexing a plethora of bennettitalean fossil plant material as CO2 proxies, including dispersed cuticles.