L.H. Bailey Hortorium, Plant Biology Section, School of Integrative Plant Science, Cornell University, Ithaca, NY, 14853-4301, USA.
Department of Entomology, American Museum of Natural History, Central Park West at 79th New York, NY, 10024, USA.
Am J Bot. 2018 Aug;105(8):1424-1435. doi: 10.1002/ajb2.1103. Epub 2018 Jun 14.
An inflorescence with three pistillate flowers in amber from the early Upper Cretaceous (Turonian, ~90-94 million years ago) of central New Jersey represents the oldest known flowers with features present in an early stem complex of the Fagales. The inflorescence has characteristics of Nothofagaceae, but also has strikingly distinct characters that suggest it is intermediate between Nothofagus and other Fagales. This intermediacy is consistent with its northern hemisphere distribution.
We investigated this new fossil by comparing it with extant and fossil members of the Fagales using light microscopy and nano-computed tomography. In addition, for exploring its relationships, we mapped the morphological characters onto a widely accepted molecular-based tree of modern basal Fagales using standard methods of character optimization.
The phylogenetic position of the fossil inflorescence can be unequivocally determined by the presence of unique features, singly and in combination, that are found only in "basal" members of Fagales. The fossil adds critical information on the features of the early stem Fagales, evolution of the cupule in Nothofagaceae and Fagaceae, and a reasonable biogeographic hypothesis for the differentiation of southern (e.g., Nothofagaceae) and northern hemisphere Fagales.
This new fossil provides insight into the early evolution of Fagales and suggests that early stem Fagales that had not yet differentiated into modern families were present in the Late Cretaceous of North America. Based on available evidence, the fossil is best interpreted as an early stem member of the Fagales, with features that suggest a transition from a more generalized Nothofagus-like fagalean ancestor with some unique presumably plesiomorphic features. The presence of an enlarged perianth and flexuous styles also suggests the possibility of insect pollination, which has been lost in all Fagales with the exception of some members of subfamily Castaneoideae in Fagaceae sensu stricto (which otherwise are very different from this fossil). The poorly developed, bract-like cupule valves of the fossil can be interpreted as primitive (i.e., incipient) or as reduced from more developed cupules that are found in most modern Fagaceae and Nothofagaceae.
来自新泽西州中部早白垩世(Turonian,约 90-94 百万年前)的一块带有三朵雌花的花朵琥珀代表了已知最古老的具有山毛榉目早期茎复杂特征的花朵。该花序具有山矾科的特征,但也具有明显不同的特征,表明它在山矾属和其他山毛榉目之间处于中间位置。这种中间状态与它在北半球的分布一致。
我们通过与山毛榉目现存和化石成员进行比较,使用光学显微镜和纳米计算机断层扫描来研究这个新化石。此外,为了探索其关系,我们使用标准的特征优化方法,将形态特征映射到广泛接受的现代山毛榉目分子基础树上。
化石花序的系统发育位置可以通过存在独特的特征来明确确定,这些特征单独存在或组合存在,仅存在于“基部”山毛榉目中。该化石增加了有关早期山毛榉目茎特征、山矾科和山毛榉科的杯状结构进化以及南半球(例如山矾科)和北半球山毛榉目分化的合理生物地理假说的关键信息。
这个新化石提供了对山毛榉目早期进化的深入了解,并表明尚未分化为现代科的早期山毛榉目茎在北美晚白垩世就已经存在。基于现有证据,该化石最好解释为山毛榉目的早期茎成员,具有从更具普遍性的山矾属山毛榉目祖先向具有一些独特的可能祖征特征过渡的特征。增大的花被和弯曲的花柱的存在也表明了昆虫授粉的可能性,而除了山毛榉科狭义的山毛榉亚科中的一些成员外(它们在其他方面与这个化石非常不同),所有山毛榉目都失去了这种授粉方式。化石中发育不良、类似苞片的杯状结构可以解释为原始的(即初生的),或者是由更发达的杯状结构简化而来的,这些杯状结构存在于大多数现代山毛榉科和山矾科中。