Evolutionary and ecological perspectives of late Paleozoic ferns: Part II. The genus Ankyropteris and the Tedeleaceae
Abstract
The Tedeleaceae are late Paleozoic Filicalean ferns with two known genera, Ankyropteris and Senftenbergia, which partially overlap morphologically and have the same known stratigraphic ranges--Namurian A to Lower Permian in Euramerica. Ankyropteris is anatomically known and Senftenbergia is almost exclusively based on compression specimens. Both genera exhibit abaxially, laminar-borne sori and/or solitary sporangia with some schizaeoid attributes. The sporangia are pyriform to tear-drop shaped, erect to slightly curved with a two to multiseriate apical annulus and mature spores usually of the Raistrickia type. Detailed morphological comparisons of Ankyropteris with Senftenbergia emphasize generic overlaps as well as lack of congruence. Ankyropteris brongniartii, a classic vine in anatomy, defines the stratigraphic range of the genus and provides recurrent ecological observations of habit and associated plants in ecotonal swamp reaches as well as in volcanic environments and on other clastic substrates. Ankyropteris may have originated from the clepsydroid zygopterid clade outside of the swamps. Ankyropteris brongniartii occurred as an epiphytic vine on Psaronius tree ferns as they expanded into coal swamps and often co-occurred with Paralycopodites even earlier in Middle Pennsylvanian. Vine stem anatomy and functional morphology of aphlebiae are emphasized in A. brongniartii with comparisons to Austroclepsis, an early "vining" false tree fern. Anatomical comparisons of Ankyropteris species based on shoots emphasize divergent habits with similar robust sizes; all exhibited aphlebiae. Known only from coal balls, Ankyropteris corrugata (Westphalian A-B) had a rampant habit on exposed peat with a succulent, dichotomous rhizome and erect fronds. The Lower Pennsylvanian specimen of Ankyropteris hendricksii is a compact tree fern with crowded frond bases and intercalated roots. Tyloses commonly occur in A. corrugata and A. brongniartii indicating water-stressed conditions. Apical meristems of shoot and fronds are reported for the first time, suggesting response growth to disturbance and rapid burial.