Lygodium palmatum subsp. puskariorum Metzgar 2025, subsp. nov.

Lygodium palmatum subsp. puskariorum Metzgar subsp. nov. Figs 1, 2, 3, 5 Type. United States. New York • Saratoga, Northumberland Township, 147 Gurnspring Road, in opening behind houses on east side of road, 26 July 2003, J. Metzgar 44 (holotype: DUKE! 397033). Diagnosis. Lygodium palmatum subsp. puskariorum differs from the similar Lygodium palmatum subsp. palmatum by the absence of white to reddish-orange hairs on the abaxial surface of the sterile leaflets, the sterile leaflet segment petioles, the abaxial surface of the fertile leaflets, and the fertile leaflet segment petioles. Description. Roots with white hairs. Rhizome approximately 1 mm wide and subterranean with dark red hairs. Leaves are evergreen and grow to three meters in length with a thin, twining stipe. Compound pinnae alternately arranged with a pinna stalk borne from the rachis. The pinna stalk branches into two primary pinna branches. Dormant pinna bud that can be glabrous or pubescent found at this branching junction. Each sterile primary pinna branch has a segment petiole terminating in a sterile segment (pinnules). Sterile segments palmately lobed, with 4–6 lobes per segment. Segment petioles and the abaxial surface of sterile segments are glabrous. Fertile segments are dimorphic with respect to the sterile segments. Fertile segments have reduced lamina with sporangia borne on projecting sorophores. The abaxial surface of the fertile segment lamina and the fertile segment petiole lack hairs. Sporangia are indusiate and occur in 4–10 pairs on a sorophore. Spores mature in the fall or early winter. Etymology. The epithet “ puskariorum ” is in honor of both Allan and Donna Puskar (Fig. 1 C), retired school teachers from Wellsboro, PA. Their family name is pronounced “ push-car. ” Allan and Donna taught my high school ecology and biology courses, fueling my love for science and natural history. They have always possessed a deep commitment to their students, with a special compassion for me and many other students who did not quite belong, were down on their luck, or yearned for a larger world. They are also lifelong fern enthusiasts who, one fateful summer in 2001, taught me the basics of fern identification and inadvertently launched my career as a pteridologist. Lygodium palmatum has long been a favorite fern of theirs. Common name. I suggest “ Puskar’s climbing fern ” and “ Puskar’s Hartford fern ” as common names for L. palmatum subsp. puskariorum and suggest “ southern climbing fern ” and “ southern Hartford fern ” as common names for L. palmatum subsp. palmatum. Geographic distribution and habitat. Lygodium palmatum subsp. puskariorum is distributed across the northeastern and mid-Atlantic USA, being found in Connecticut, Maryland, Massachusetts, New Jersey, New York, North Carolina, Pennsylvania, and Rhode Island (Fig. 3). Southern New Jersey, southeastern North Carolina, and central Massachusetts represent areas of secondary contact with L. palmatum subsp. palmatum. The Puskar’s climbing fern grows in moist, acidic soil in forests, bogs, and roadsides. Conservation assessment. Lygodium palmatum sensu lato has a global conservation ranking of G 4 (“ apparently secure ”; Weakley 2025). However, most states in L. palmatum subsp. puskariorum ’ s range already list L. palmatum s. l. as Critically Imperiled (S 1), Imperiled (S 2), or Vulnerable (S 3; NatureServe 2025). Additionally, the taxon is sensitive to development and generally does not survive transplantation or cultivation (Montgomery 1989). I recommend Imperiled status (G 2) for L. palmatum subsp. puskariorum due to its more narrow distribution and scarcity throughout most of its range. Additional specimens examined. United States. Connecticut • Hartford, Plainville, Sep. 1877, J. Bishop s. n. (CM) • Hartford, probably from vicinity of Hartford, Ziegler s. n. (CM) • Hartford, Southington, s. coll. s. n. (CM); no further locality, H. Denslow s. n. (DUKE) • Plainville, Aug 1883, A. Seymour 134 (DUKE); Maryland • Anne Arundel, along Severn River near Benfields, 18 Aug 1966, L. Henry s. n. (CM) • Anne Arundel, North side of Severn Run, 19 June 1981, M. Windham 81-26 (DUKE) • Charles, Sevier River, 18 Aug. 1960, J. Benedict 6432 (NCU, VPI) • Prince Georges, Bladensburg, 26 Mar. 1910, H. Bartlett 1887 (DUKE, NCU) • Prince Georges, Hyattsville, Nov. 1899, C. Pollard, Louis s. n. (NCU) • Prince Georges, Suitland, 9 Sept. 1899, E. Steele (DUKE); Massachusetts • Hampden, Palmer, 25 Aug 1925, G. Bennett G 163 (DUKE) • Hampden, Springfield, 31 Aug. 1934, F. Sargent (DUKE) • Hampden, Wilbraham, 11 Sept. 1935, F. Seymour 4406 (DUKE) • Hampshire, Amherst, 11 July 1980, C. Hellquist 14577 (CM) • Hampshire, Amherst, 28 June 1928, S. Ewer s. n. (NCU) • Hampshire, Amherst, 4 Nov. 1889, G. Nash 99 (CM) • Hampshire, Amherst, S. Estabrook s. n. (NCU) • Hampshire, East Hampton, 29 June 1872, E. Miller s. n. (CM) • Hampshire, North Amherst, 23 Aug. 1895, A. Beals 37 (DUKE) • Hampshire, North Hadley, 29 Nov. 1922, R. Sanborn 2082 (DUKE) • Middlesex, Ayer Junction, July 1883, S. Steele (DUKE) • Middlesex, Shirley, F. Sargent (DUKE) • Middlesex, Still River, Oct. 1910, R. Ware 766 (NCU) • no further locality, Canby (CM) • no further locality, H. Mann s. n. (CM) • no further locality, s. coll. s. n. (NCU) • Plymouth, Lakeville, 12 Sept. 1986, B. Sorrie 3652 (NCU); New Jersey • Burlington, 1 mi NNE of Taunton, 10 Aug. 1926, W. Benner s. n. (CM) • Burlington, Hampton Furnace, 3 July 1922, G. Bassett (DUKE) • Burlington, near New Lisbon, 11 June 1950, D. Krouse s. n. (CM) • Middlesex, Milltown, 25 Dec. 1924, G. Johnson (DUKE) • Middlesex, New Brunswick, 1899, J. Blodgett s. n. (CM) • Monmouth, Shark River, 5 Oct. 1855, F. Hexamer s. n. (CM) • Sussex, Pine Swamp, 1838, Wolle Herbarium s. n. (CM); New York • Onondaga, Cicero Swamp, near Syracuse, 22 Nov. 1966, M. Faust s. n. (CM) • Saratoga, near Gansevoort, 30 Oct. 1938, H. House 26344 (NCU) • Saratoga, Town of Northumberland, Oct. 1938, O. Phelps s. n. (NCU); North Carolina • Bladen, 1 June 1979, R. Rebertus s. n. (NCU) • Cumberland, 1 mile east of Stedman, 26 Dec. 1932, H. Totten s. n. (NCU) • Cumberland, 1 mile southeast of Stedman, 26 May 1961, A. Radford 43676 (NCU) • Cumberland, 7 miles south of Fayetteville, 17 Oct. 1931, M. Smyth s. n. (NCU) • Cumberland, About 1 mile southwest of Stedman, 26 Nov. 1964, H. Totten 1832 (NCU) • Cumberland, Along bank of Rock Fish Creek, 8 Oct. 1939, A. Radford s. n. (NCU) • Cumberland, Along bank of Rockfish Creek, 8 Oct. 1939, L. Stewart s. n. (DUKE) • Cumberland, Along Rockfish Creek, 8 May 1938, D. Correll 9052 (DUKE) • Cumberland, Ardlussa, 1 Oct. 1931, M. Vardell s. n. (DUKE) • Cumberland, Dunns Creek, 16 Oct. 1982, L. Raubeson 399 (NCU) • Cumberland, Fayetteville, 23 Sept. 1934, J. Benedict 3078 (VPI) • Cumberland, North side of Rockfish Creek, 28 Oct. 1969, S. Leonard 2804 (NCU) • Cumberland, Rock Fish Creek, 15 March 1964, T. Creem 316 (NCU) • Cumberland, Vicinity of Fayetteville, 1 May 1937, R. Clark (DUKE) • Onslow, East side of Sandy Run Swamp, 8 Aug. 1997, M. Strong 1571 (NCU) • Sampson, 6 miles south of Clinton, 27 Mar. 1938, L. Anderson 6159 (DUKE); Pennsylvania • Carbon, Mauch Chunk, Oct. 1873, C. Shaefer 11191 (CM) • in Poconos, 17 July 1970, E. Laport s. n. (NCU) • Luzerne, Hells Kitchen, Nescopeck Creek, 19 Sept. 2022, R. Sim s. n. (CM) • Luzerne, Nescopeck State Park, 10 July 2004, J. Metzgar 87 (DUKE) • Luzerne, Ten Mile Run, 1 Oct. 1938, T. Darling (VPI) • Monroe, near Tannersville, July 1874, S. Knipe s. n. (CM) • Monroe, Tunkhannock Township, 6 Sept. 1936, F. Trembley 603 (DUKE) • Monroe, Tunkhannock Township, 9 Sept. 1936, F. Trembley s. n. (NCU) • Monroe, Tunkhannock Twp, 8 July 2015, W. Gleason F 15 GLE 35 - E (CM) • Pike, Pocono Mountain, Turners Swamp, July 1874, S. Knipe s. n. (CM); Rhode Island • Kent, Anthony, 18 Sept. 1886, J. Collins s. n. (NCU) • Kent, Anthony, 26 Sept. 1885, G. Leland s. n. (CM) • Kent, Coventry, 16 Oct. 2020, J. Collins s. n. (NCU). Representative specimens examined of Lygodium palmatum (Bernh.) Sw. subsp. palmatum. United States. Kentucky • Breathitt, Near Morris Fork, S. VanderMeer s. n. (NCU) • Harlan, Above US 421, about 1 mile from Virginia line, 17 Feb. 1965, M. Smyth 508 (VPI) • McCreary, Cumberland Falls State Park, 15 Sept. 1982, F. Utech 82–462 (CM) • Powell, Trail to Rock Bridge Natural Bridge Park, 14 Apr. 1958, C. Gunn K 30 (NCU) • Whitley, Cumberland Falls State Park, 24 Aug. 1961, L. Henry s. n. (CM) • Wolfe, 12 Sept. 1969, P. Higgins s. n. (NCU); Massachusetts • Hampden, Hampden, Oct. 1913, S. Bliss s. n. (NCU); Michigan • Kalamazoo, S of Kalamazoo, 14 Oct. 1974, R. Simpson s. n. (LFCC); New Jersey • Burlington, 1 mi NNE of Taunton, 10 Aug. 1926, W. Benner s. n. (NCU); North Carolina • Alleghany, 9 Sept. 2012, D. Goldman 4220 (NCU) • Ashe, Long Hope Creek Valley, 11 Oct. 1969, J. Moore 2799 (NCU) • Bladen, 1.6 mi ESE of Ammon, 14 July 1999, B. Sorrie 10217 (NCU) • Buncombe, 2 miles southwest of Black Mountains, 29 Aug. 1948, E. Browne s. n. (NCU) • Burke, 3 mi N of Optimist Park, 3 Apr. 1971, R. Harbison Pharr 1013 (NCU) • Cabarrus, Cabarrus, 16 Aug. 1979, D. Wickland 2126 (NCU) • Chatham, In ore mine at Mt. Vernon Springs, 13 May 1953, C. Lindley s. n. (NCU) • Cherokee, Above Granny Squirrel Gap, 17 July 2002, C. McCartney s. n. (NCU) • Cumberland, Rockfish Creek, 7 Aug. 1957, H. Ahles 33420 (NCU) • Davidson, near Silver Hill, 8 May 1988, R. Wilbur 54883 (DUKE) • Durham, NC Gamelands Trail, 30 Aug. 1992, R. Wilbur 61438 (DUKE) • Graham, approx. 3.5 mi I from turnoff to Joyce Kilmer Memorial Forest, 15 Sept. 2000, C. McCartney s. n. (NCU) • Harnett, west of Stewarts Creek, 25 Aug. 2005, B. Sorrie 11702 (NCU) • Henderson, at Pine Knob (or Brickyard) Mine, 10 Sept. 1978, D. Wickland 2452 (NCU) • Hoke, Fort Bragg, 16 Dec. 1992, B. Sorrie 7117 (NCU) • Jackson, headwaters of Tuckaseigee River, 28 June 1970, S. Leonard 3315 (NCU) • Macon, near Snow Hill, March