Giant cladoxylopsid trees resolve the enigma of the Earth’s earliest forest stumps at Gilboa

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Abstract
The evolution of trees of modern size growing together in forests fundamentally changed terrestrial ecosystems. The oldest trees are often thought to be of latest Devonian age (about 380-360 Myr old) as indicated by the widespread occurrence of Archaeopteris (Progymnospermopsida). Late Middle Devonian fossil tree stumps, rooted and still in life position, discovered in the 1870s from Gilboa, New York, and later named Eospermatopteris, are widely cited as evidence of the Earth's 'oldest forest'. However, their affinities and significance have proved to be elusive because the aerial portion of the plant has been unknown until now. Here we report spectacular specimens from Schoharie County, New York, showing an intact crown belonging to the cladoxylopsid Wattieza (Pseudosporochnales) and its attachment to Eospermatopteris trunk and base. This evidence allows the reconstruction of a tall (at least 8 m), tree-fern-like plant with a trunk bearing large branches in longitudinal ranks. The branches were probably abscised as frond-like modules. Lower portions of the trunk show longitudinal carbonaceous strands typical of Eospermatopteris, and a flat bottom with many small anchoring roots. These specimens provide new insight into Earth's earliest trees and forest ecosystems. The tree-fern-like morphology described here is the oldest example so far of an evolutionarily recurrent arborescent body plan within vascular plants. Given their modular construction, these plants probably produced abundant litter, indicating the potential for significant terrestrial carbon accumulation and a detritus-based arthropod fauna by the Middle Devonian period.