Sandstone rockhouses of the eastern United States, with particular reference to the ecology and evolution of the endemic plant taxa

Rockhouses are semicircular recesses extending far back under cliff overhangs that are large enough to provide shelter for humans. The largest sandstone rockhouses in the eastern United States are at the heads of gorges, and they are in stream valleys cut during the Pleistocene; most are formed in Mississippian and Pennsylvanian-age rocks. Compared to the surrounding environment, the interior of rockhouses is shaded, is warmer during winter and cooler during summer, and has lower evaporation rates and higher humidities. Water enters rockhouses primarily by groundwater seepage and by dripping from the ceiling. Soil consists mostly of sand with low pH, but high levels of some nutrients are associated with saltpeter earth and with ecofactual and artifactual remains left by human occupants during prehistoric time. Most plant taxa in sandstone rockhouses in eastern United States are native C3 phanerophytes or hemicryptophytes, and similarities in species composition among rockhouses are low. Eleven plant taxa belonging to eight families of flowering plants and ferns are endemic or nearly endemic to sandstone rockhouses in eastern United States. Three endemics are restricted to the gorges of a single river, and only one taxon ranges far north of the Wisconsinan Glacial Boundary. The endemic ferns are Tertiary relicts derived from tropical taxa. The majority of endemic flowering plants are derived from temperate taxa that grow in habitats in the vicinity of rockhouses; their relative age ranges from Late Tertiary to the Recent. All the endemic taxa are perennial; two ferns occur as independent gametophytes. The endemic taxa of rockhouses are threatened primarily by disturbances associated with recreation.