Asplenium × gravesii, commonly known as Graves' spleenwort, is a rare, sterile, hybrid fern, named for Edward Willis Graves (1882–1936). It is formed by the crossing of Bradley's spleenwort (A. bradleyi) with lobed spleenwort (A. pinnatifidum). It is only found where its parent species are both present; in practice, this proves to be a few scattered sites in the Appalachian Mountains, Shawnee Hills, and Ozarks, reaching perhaps its greatest local abundance around Natural Bridge State Resort Park. Like its parents, it prefers to grow in acid soil in the crevices of sandstone cliffs.
Description
Asplenium × gravesii is a small fern, whose fronds grow in loosely bundled tufts. Its stem below the leaf blade is a shiny purplish-brown, while the green, narrowly triangular blades are cut into pinnae near the base, which diminish into lobes in the upper part. The fronds are monomorphic, showing little or no difference between sterile and fertile fronds.
The fronds of A. × gravesii, which are long, are closely spaced along an upward-curving, densely scale-covered rhizome in length. The scales are long and narrow, about long and 0.5 millimeters wide at the base, coming to a pointed tip. They are dark reddish-brown in color and strongly clathrate (bearing a lattice-like pattern). The stipe (the stalk of the leaf, below the blade) is typically long, shiny and purplish-brown, sometimes becoming green near the base of the leaf blade. It is rounded below and flat or grooved above.
The overall shape of the blade is narrowly triangular, from long and wide. The leaf tissue is of a medium texture (neither delicate nor leathery), with clavate (club-shaped) hairs on the underside becoming gland-tipped, narrow scales on the veins. The lower part of the blade is cut into pinnae, diminishing to lobes in the upper part. The basal pinnae are triangular in shape, roughly equilateral, and nearly heart-shaped at their base; they are borne on short stalks. They are shallowly rounded or toothed around the edges. In larger specimens, the basal pinnae bear a pair of rounded lobes at their base. Successive pinnae above the base are narrower and less deeply cut, gradually diminishing into fused lobes or overlapping. The lobes continue to the pointed tip of the blade. The rachis (central axis of the blade) is typically green and flat, with narrow wings, and a groove on the upper side near the base of the blade, where it becomes the stipe. Larger individuals may have a narrower, glossy-brown rachis. A few scales are present on stipe and rachis, becoming narrower and more twisted as they ascend from the rhizome.
The abundant sori are dark brown and variable in shape, fusing with one another as they grow. The sori are covered with firm, white indusia. In wild specimens, they are found beneath the costa (midrib of the pinnae). As a sterile tetraploid hybrid, the spores are seen to be misshapen and abortive under microscopic examination. The species has a chromosome number of 144 (2 × 72) in the sporophyte.
Asplenium × gravesii can potentially be confused with its parental species or with a number of the other Asplenium hybrids in the Appalachian Asplenium complex. Smaller specimens are most similar to A. pinnatifidum, but can be distinguished from that species by their brown stipes (and often rachides), a smaller number of fronds in each tuft, and by the tip of their leaves (which is pointed, but not drawn out at length as in A. pinnatifidum). A. × gravesii also has a slightly finer leaf texture, slightly sharper teeth on its leaves, and darker brown sori than A. pinnatifidum. When compared to A. bradleyi, larger A. × gravesii individuals are most similar, but the blades are not so deeply cut (A. bradleyi being wholly pinnate) nor the pinnae toothed around the edges, the rachis shows at least some traces of winging, and more of the rachis tends to be green. The leaf texture is somewhat more delicate than the leathery blades of A. bradleyi.