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Isoetes hyemalis D.F. Brunton
Winter Quillwort

Isoetes hyemalis by Daniel Brunton. Image subject to copyright.
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Federal Protection: No US federal protection

State Protection: No Georgia state protection

Global Rank: G2G3

State Rank: S1?

Element Locations Tracked in Biotics: Yes

SWAP 2015 Species of Greatest Conservation Need (SGCN): Yes

SWAP 2025 Species of Greatest Conservation Need (SGCN): Yes

2025 SGCN Priority Tier: Highest Conservation Concern

Element Occurrences (EOs) in Georgia: 2

Habitat Summary for element in Georgia: Sandy blackwater creek banks; deciduous swamps


Description

Perennial herb, forming robust tufts of leaves on creek banks. The rootstock (corm) with two lobes, one or more attached to a knotty, horizontal, underground stem (rhizome); new plants develop from these corms. The leaves are 9 - 10.6 inches (23 - 27 cm) long, evergreen, bright green when young, dark brownish-green when older, flared at the base and tapering to a sharply pointed tip, with 4 cross-walls and a single midvein; when growing in flowing water, the leaves are long, wavy, and slightly curled. Spores are produced in the sporangium, a brown-streaked or brown-spotted chamber 0.2 inch (5.3 mm) long in the leaf base, with a transparent membrane (velum) covering 10 - 20% of the chamber opening. Dozens of tiny, grayish-white female spores (megaspores), approximately 0.05 mm across and covered with a dense pattern of bumps (low spines) and very short, loosely connecting ridges, may be seen with 20 - 30x magnification. Olive-gray to tan, dust-sized male spores (microspores) are produced on separate leaves but are indistinguishable without much higher magnification.

Similar Species

Quillworts are distinguished from flowering, wetland plants by their spongy leaves with conspicuous cross-walls and by the presence of sporangia in the flared base of the leaves.

Engelmann’s Quillwort (Isoetes engelmannii) has long leaves (10 - 24 inches, 25 - 60 cm) like winter quillwort, and the velum covers less than 30% of the sporangia, but it has lighter green leaves and smaller megaspores with regularly reticulate ornamentation patterns; Engelmann’s quillwort is the most common quillwort in Georgia but is found mostly in the Piedmont.

Appalachian Quillwort (Isoetes appalachiana), found across the state, has the same size megaspores as winter quillwort but these have an irregularly reticulate ornamentation pattern.

Southern Quillwort (Isoetes flaccida) also occurs in the southwest portion of Georgia’s Coastal Plain in habitats similar to winter quillworts; it also has long leaves (4 - 24 inches, 10 - 60 cm) but its velum completely covers the spore chamber, which is colorless (not streaked with brown), and contains smaller, more plainly ornamented megaspores.

Boom’s Quillwort (Isoetes boomii) and Georgia Quillwort (I. georgiana) both have larger, more coarsely ornamented megaspores and more extensive velum coverage.

Related Rare Species

Twelve quillwort species are state-listed or considered of Special Concern in Georgia. Information about each of these species may be found at these links:

Boom's Quillwort (Isoetes boomii) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=17134

Appalachian Quillwort (Isoetes appalachiana) http://explorer.natureserve.org/servlet/NatureServe?searchName=Isoetes+appalachiana

Glade Quillwort (Isoetes butleri) http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250076861

Florida Quillwort (Isoetes flaccida) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=19963

Georgia Quillwort (Isoetes georgiana) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=16950

Winter Quillwort (Isoetes hyemalis) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=17363

Rush Quillwort (Isoetes junciformis) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=18998

Black-footed Quillwort (Isoetes melanopoda) http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250076862

Black-spored Quillwort (Isoetes melanospora) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=22510

Broxton Rocks Quillwort (Isoetes "snowii," undescribed) https://bioone.org/journals/castanea/volume-83/issue-2/17-122/Interesting-Provocative-and-Enigmatic--Morphological-Observations-on-Southeastern-Quillworts/10.2179/17-122.full

Mat-forming Quillwort (Isoetes tegetiformans) https://www.georgiabiodiversity.org/portal/profile?group=plants&es_id=20422

Mountain Bog Quillwort (Isoetes valida) https://www.jstor.org/stable/1547604?seq=1#metadata_info_tab_contents

Habitat

Densely shaded blackwater creeks with circumneutral to basic water pH, sandy banks along seasonally flowing streams, and flowing water in swamps. Outside of Georgia, Winter Quillwort forms pure stands of several hundred plants in blackwater streams, but also may occur with rushes (Juncus), pondweeds (Potamogeton), and burweed (Sparganium americanum).

Life History

Quillworts are seedless, non-flowering plants that reproduce by spores. Winter quillwort also may rarely reproduce vegetatively by producing plantlets along a horizontal, underground rhizome. Quillwort leaves have hollow chambers at the base where two types of spores are produced: tiny microspores develop sperm-producing structures, and larger (though still minute) megaspores produce eggs. Sperm swim to the eggs in available water and unite to form new plants. Quillworts compete poorly with other aquatic plants and are typically found in relatively sterile sand or silt or in frequently water-worn sites that support few or no other vascular plants.

Survey Recommendations

Surveys in Georgia are best conducted in late spring – early summer when plants are most conspicuous and their leaves have not withered; mature megaspores are best developed in mid – late summer but may be found (from previous year’s growth) in the soil at the base of plants during the following spring.

Range

Coastal Plain and lower Piedmont of Georgia, Florida, Alabama, South Carolina, North Carolina, and Virginia.

Threats

Ditching and draining wetlands, impounding streams, clearcutting in swamps and floodplains. Invasion by exotic pest plants.

SWAP 2025 Threat Matrix

Threat 1 Threat 2 Threat 3
General Threat Transportation & service corridors Biological resource use Human intrusions & disturbance
Specific Threat None None None

Georgia Conservation Status

Isoetes hyemalis is ranked S1? by the Georgia Department of Natural Resources, indicating that the species is probably critically imperiled in the state, but that additional information is needed to make a final ranking. Two populations have been documented in Georgia though neither have been confirmed since 2000; neither population occurs on conservation land.

Conservation Management Recommendations

Protect floodplains and swamps and adjacent areas of local watersheds from damming, clearing, draining, and filling. Eradicate exotic pest plants.


SWAP 2025 Conservation Actions:

  • Action 1: Complete a distributional survey to assess current range, conservation status or to identify best populations
  • Action 2: Protect key populations using land acquisition or easements
  • Action 3: Improve habitat using prescribed fire
  • Action 4: Reassess the conservation status of SGCN before the next revision of Georgia's State Wildlife Action Plan

References

Brunton, D.F. 2015. Key to the quillworts (Isoetes: Isoetaceae) of the southeastern United States. American Fern Journal 105(2): 86-100. https://www.jstor.org/stable/44076059?seq=1#metadata_info_tab_contents

Brunton, D.F., D.M. Britton, and W.C. Taylor. 1994. Isoetes hyemalis, sp. nov. (Isoetaceae): a new quillwort from the southeastern United States. Castanea 59: 12-21. https://www.jstor.org/stable/4033758

GADNR. 2020. Element occurrence records for Isoetes hyemalis. Georgia Department of Natural Resources, Wildlife Resources Division, Social Circle, Georgia.

Musselman, L.J. 2001. Georgia quillworts. Tipularia, Journal of the Georgia Botanical Society 16: 2-19, 40.

NatureServe. 2019. Isoetes hyemalis comprehensive report. NatureServe Explorer. Arlington, Virginia. http://explorer.natureserve.org/servlet/NatureServe?searchName=Isoetes+hyemalis

Weakley, A.S. 2015. Flora of the southern and mid-Atlantic States. University of North Carolina Herbarium, University of North Carolina, Chapel Hill. http://www.herbarium.unc.edu/flora.htm

Authors of Account

Linda G. Chafin and Daniel F. Brunton

Date Compiled or Updated

L. Chafin and D. Brunton, Dec. 2008: original account

K. Owers, Feb. 2010: added pictures

L. Chafin, March 2020: updated original account.

Isoetes hyemalis, illustration by Jean C. Putnam Hancock. Image may be subject to copyright.
Isoetes hyemalis habitat by Daniel Brunton. Image subject to copyright.
Isoetes hyemalis habitat by Daniel Brunton. Image subject to copyright.
Isoetes hyemalis by Alan Cressler. Image may be subject to copyright.