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Home , Asplenium, Asplenium rhizophyllum, Asplenium scolopendrium, Asplenium trichomanes, Stolon

rhizophyllum L. walking

Photos by Michael R. Penskar State Distribution

Best Survey Period

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Status: State threatened Niagara Escarpment. Elsewhere, this occurs locally on alkaline bedrock outcrops in Dickinson, Global and state rank: G5/S2S3 Schoolcraft, and Houghton counties, with additional : (spleenwort family) local populations found in the Lower Peninsula on South Manitou Island (Leleenau County). It is also Synonyms: Camptosorus rhizophyllus (L.) Link known from a sinkhole in Alpena County, and from an unusual occurrence in Berrien County where a small : This very distinctive species has been but vigorous was discovered on a limestone segregated by several authors and placed in the boulder along a stream. Camptosorus (Morin et al. 1993), a name under which it is known in many manuals and other publications. It Recognition: is an extremely forms part of a complex of Appalachian spleenworts distinctive fern, characterized by its tendency to form researched by Wagner (1954) in a well-known study of dense colonies by reproducing via tip-rooting on hybridization and backcrossing. -covered dolomite boulders and other types of rock outcrops. Individual consist of clumps of Total range: Walking fern occurs in eastern North fronds (leaves) arising from short, scaly . The America, ranging from southern Ontario and Quebec small, 1-3 cm wide fronds, which have net-like (reticu- in Canada south to Georgia, Alabama, and Mississippi, late) veins and may range up to ca. 30 cm in length, occurring west to Wisconsin, Iowa, Kansas, and are stalked, and have slender, long-tapering, lance- Oklahoma. shaped blades with heart-shaped (cordate) to State distribution: Michigan represents somewhat of strongly lobed (auriculate) bases. On the underside a disjunct region of occurrence for this fern, which is of the fronds, the spores are produced in somewhat much more common in the southern portion of its irregular, linear, brown masses of sporangia scat- range. Most of the state’s 27 localities are clustered in tered between the midrib and leaf margin. the eastern Upper Peninsula, primarily in the eastern As indicated by the common name, this species portion of Mackinac County, where there is a concen- “walks” by proliferating through the development of tration of exposed limestone and dolomite boulders long-tapering leaf tips, which elongate and arch — along the spine of the geological feature known as the stolon-like — across moss mats, forming new plants

Michigan Natural Features Inventory P.O. Box 30444 - Lansing, MI 48909-7944 Phone: 517-373-1552 walking fern, Page 2

where the tips contact the surface. Large clonal (green spleenwort) and the federal threatened and state colonies may develop in this manner, sometimes endangered hart’s-tongue fern. Moist moss mats, carpeting substantial portions of large dolomite composed of several different moss species (e.g., boulders or cliff walls. Walking fern may be mistaken Rhodobryum roseum, several Mnium and Brachy- by the inexperienced eye for the very rare hart’s- thecium spp.) appear to be a critical element of this tongue fern, L. Hart’s- species’ microhabitat, as well as the availability of tongue fern is often associated with walking fern, but moist crevices and other concavities that serve as can be distinguished by its much larger, broader fronds colonization niches. that are somewhat shiny, wavy-margined (puckered), and have relatively abrupt, blunt-pointed tips. In Elsewhere within Michigan, walking fern occurs on addition, Hart’s-tongue fern in Michigan occurs similar alkaline rock outcrops, such as in Dickinson primarily as scattered clumps of plants growing in low County, Schoolcraft County and on the limestone face ledges and concavities, and does not form dense of a sinkhole in Alpena County. One of the more colonies on boulders as is so characteristic of walking atypical habitats is on South Manitou Island, where fern. this species occurs on mossy, decaying white cedar logs in an old-growth cedar forest on a sand dune, and Best survey time/phenology: Although this species is associated with green spleenwort. can be recognized throughout the year due to the evergreen nature of its fronds and its very specific Biology: Walking fern occurs primarily on alkaline habitat, it is best inventoried during the heart of the rock substrates, and appears to require moist micro- growing season when new foliage is produced. habitat and moderately to densely shaded conditions. Although this species thrives when growing in moist Habitat: In Michigan this fern occurs primarily in moss mats, it may require exposed, moist rock sub- association with shaded, moist boulders and outcrops strate for colonization. Once established, walking fern of Niagaran limestone is able to proliferate rapidly through tip-rooting, and and dolomite. In the then as mature plants develop, this perennial is able to eastern Upper Penin- produce spores for dispersal to available microhabitat. sula, where most Although this species occurs mostly on bedrock, it is Michigan localities of also known throughout its range to inhabit organic this species are substrates, as long as these are alkaline. Occasionally known, walking fern this species is epiphytic, occurring on trees with usually occurs on circumneutral bark (TNC 1993), and sometimes it can moss-covered boul- be found on and other acidic bedrock ders, outcrop cliff (Morin et al. 1993). faces, and low rock ledges in second- D. J. Evans (1997) found walking fern to occur under growth mesic northern a wider range of canopy cover than the state endan- forest. Typical canopy gered Hart’s tongue fern (A. scolopendrium var. trees include Acer americana) which occupies similar dolomite boulders. saccharum (sugar Her study also showed a high frequency of walking maple), Tilia americana (basswood), Abies balsamea fern on all aspects of the boulders in contrast to the (balsam fir), Fraxinus americana (white ash), and significant preference for cool, dark, north and north- Betula alleghaniensis (yellow birch). Common herba- east aspects exhibited by Hart’s tongue fern. This may ceous species associated with walking fern include reflect the ability of walking fern to “move” across a Polystichum lonchitis (northern holly fern), boulder by -tipping. Walking fern was also found Polypodium virginianum (common polypody), Gera- to occupy a higher vertical position on the rocks, nium robertianum (herb-Robert), Cystopteris fragilis indicating an ability to better withstand periodic (fragile fern), (maidenhair intense sunlight in early spring and late fall during spleenwort), and occasionally such rare plants as the leaf-off periods. state threatened Asplenium trichomanes-ramosum

Michigan Natural Features Inventory P.O. Box 30444 - Lansing, MI 48909-7944 Phone: 517-373-1552 walking fern, Page 3

Conservation/management: The conservation of this Evans, D.J. 1997. Habitat characteristics and Distri- rare fern will depend, over the short term, on protec- bution Patterns of Three Rare Aspleniums in tion of its habitat from major disturbances. Before Northern Mackinac County, Michigan. Master’s long-term management can be undertaken, much more thesis. Michigan Technological University, needs to be learned about this fern’s life history and its Houghton, MI. 112 pp. response to habitat changes, especially overstory Wagner, W.H., Jr. 1989. Kathryn E. Boydston (1897- removal. However, maintenance of at least a partial 1988): Michigan’s fern hybridist and two new canopy is likely crucial to perpetuating the moist moss examples of her work. Mich. Bot. 28:51-57. mats that provide the necessary microhabitat for this Wagner, W.H., Jr. 1954. Reticulate evolution in the species (TNC 1993). Appalacian Aspleniums. Evolution 7:103-118. Comments: This species is frequently cultivated by Wagner, W.H., Jr., R.C. Morin, and C.R. Werth. 1993. horticulturalists, who also artificially hybridize Aspleniaceae Newman – Spleenwort Family. Pages 228- walking fern with a number of other related taxa to 245. In: Flora of North American North of Mexico, Vol. produce unusual ornamentals and study specimens 2. Morin et al. Oxford Press, New York. 474 pp. (Wagner 1989). Research needs: Long-term monitoring to determine population trends, and the effects of such factors as Abstract Citation: light availability and disturbance, would be the most Penskar, M.R. and P.J. Higman. 1997. Special useful investigations. abstract for Asplenium rhizophyllum (walking fern). Michigan Natural Features Inventory, Lansing, MI Related abstracts: Mesic northern forest, Assiniboia 3 pp. sedge, fairy-bells, ginseng, goblin moonwort, Hart’s- tongue fern, northern goshawk, red shouldered hawk, rapids clubtail, woodland vole. Selected references: Cody, W.J. and D.M. Britton. 1989. and fern allies of Canada. Research Branch, Agriculture Canada. Publication 1829/E. 430 pp. Mickel, J.T. 1979. How to know the ferns and fern allies. The Pictured Key Nature Series. Wm. C. Brown Co. Dubuque, Iowa. 229 pp. Morin, N. et al. 1993. Flora of , North of Mexico. Volume 2: Pteridophytes and Gymno- sperms. Oxford Univ. Press. New York, NY. 475 pp. The Nature Conservancy. 1993. The Nature Conser- vancy, Conservation Science Division, in associa- Updated August 2001. tion with the Network of Natural Heritage Pro- grams and Conservation Data Centers. Asplenium Copyright 2004 Michigan State University Board of Trustees. rhizophyllum, Element Stewardwhip Abstract Michigan State University Extension is an affirmative-action, equal- Report. Arlington, VA. opportunity organization. Lellinger, D.B. 1985. A field manual of ferns and fern Funding for abstract provided by Michigan Department of allies of the United States and Canada. Natural Resources-Forest Management Division and Smithsonian Inst. Press. Wash, D.C. 389 pp. Wildlife Division.

Michigan Natural Features Inventory P.O. Box 30444 - Lansing, MI 48909-7944 Phone: 517-373-1552