Carex and Associates of Homosassa Springs and Three Sisters Springs

Linda W. Curtis

Three Sisters Springs Wildlife Refuge (USFWS) is part of the Florida Gulf Coastal lowland in Crystal River and Homosassa Springs Wildlife State Park FDEP is 5 miles south, both in Citrus County, Florida.  Search for Carex species continued in 2022 with 12 species for the site inventory of TSS and 6 found in HSWSP. 

Image of great blue heron drinking in the spring run next to a log with a clump of Carex. Most Carex in Florida bloom March through May, setting seed heads of spikes with perigynia, sacs around seed-like achenes. The Gulf Coast has large tracts of hydric forest, mesic forest. basin swamps and wetlands with many under-reported Carex.

The identification of Carex requires magnification and often lab imaging, so field dats are often listed as just Carex sp. Specimens were imaged in the field with macro lens and with long-range camera recording GPS. Later, the plants are micro-imaged and data sent to Florida Herbaria and Harvard University Herbarium-Cambridge, HUH in 2021..

Scanned images of the specimens and their county distribution maps may be viewed at

Three Sisters Springs

Formerly a dense hydric Gulf forest, the remaining fringes of vegetation around the springs, roadsides and the canal perimeter are regenerating by natural succession back to hydric forest. The understory of herbaceous plants and shrubs includes Carex species discovered in 2012. More Carex were found in 2014 and the listing of their associates initiated a partial on-site plant inventory. Voucher specimens of Carex were sent to herbaria at Universitiy of Florida-Tampa (USF), University of Florida-Gainesville (FLAS), and Florida State University-Tallahassee (FSU). A new entry and trolley road were constructed from Cutler Spur in 2019 and a tidal wetland project excavation to relieve pollution in Kings Bay completed in 2015.

Carex and associates of Three Sisters Springs, this report 2014. Annual reports continued into 2022

Linda W. Curtis, consultant botanist


Seven Carex species grow at the Three Sisters Springs site in the headwaters of Crystal River, Kings Bay, Citrus County, Florida, a USFWS National Wildlife Refuge. Four of the seven Carex species grow in fringes remaining of coastal Gulf hydric hammock forest around the springs pool, roadside and canal borders and three grow in the large central ruderal area of tilled-over planted pines, an area groomed for restoration. An inventory of Carex associates of trees, understory shrubs, and herbaceous ground layer was begun, including the native trees and shrubs donated and planted by the Florida Native Plant Society, Citrus Chapter, in 2013. The existing native vegetation is regenerating, including seedlings of both woody and herbaceous native species but also some invasive and non-native plants.


The purpose of this report is to document Carex species and associated plants at Three Sisters Springs, a 57 acre formerly dense Gulf hydric hammock forest. Three Sisters Springs is one of three springs in Crystal River National Wildlife Refuge and the only one accessible by land (Fig. 1). The locked gate entry road on the northern boundary along King’s Bay Road is latitude N28o 53.519 and longitude W82o 35.469. The parking lot for tour bus drop-off is near the boardwalk at the southern boundary canal at N28 53.290 and W82 35.298.

Cutler Spur Boulevard is the eastern border with a forested tidal creek along the road sidewalk on the southeast end of the navigational canal that feeds high tide water into the roadside ditch. The southern and western borders are canals dug in the 1970s and have re- vegetated limestone banks (Fig. 2)

To the west, Citrus County’s extensive coastal salt marshes are near sea level, whereas the inland Three Sisters Spring gulf hammock is 2–6 feet above the high tide elevation on limestone karst topography. Beaches were not formed as the waves are low energy and do not presently build-up deposits. The only nearby sand beach is at the end of Fort Island Trail and is maintained with imported sand. The Pleistocene coastal swamps along the gulf date from 10,000 to 1.6 million years, in age. The gulf waters rose and fell, alternately depositing and eroding sedimentary soils, and were largely deposited in the recent past when sea levels fell. Gulf waters are now rising due to global warming.

Underground spring water in Three Sisters Spring is moderate at about 72°F and travels 6 miles west to the salt marshes along the gulf where the temperature is usually near 60°F in winter. The Three Sisters Springs inlet was natural, but was altered with canals in the 1970’s. Boulders at the inlet were removed and pipes inserted to restrict boats but allow kayaks and swimmers entry to view the manatees. Tourist visitations by land were limited to Special Use Vendors until November 2014 when one tour company began to manage shuttle-bus tours, restricting tourists to the boardwalk to prevent trampling. A future entry will be a bridge over Cutler Spur Blvd. and that street will be renamed.

The original vegetation of Three Sisters Springs Wildlife Refuge was coastal hydric hammock forest. The site was cleared for development and only forested fringes grew around the springs and site perimeters, with a diverse understory of Carex and other herbs including a 30” tall Arisaema triphyllum, Jack-in-the-pulpit.

Mature non-native shrubs include Schinus terebinthifolia, (Brazilian pepper), Callestemon viminale, (bottlebrush), and Pyracantha coccinea, (firethorn) which escaped from plantings along the west canal boundary.

Native species grow along roadside creeks of both Cutler Spur Blvd. on the east, King’s Bay Drive on the north, and limestone cut banks of the dredged channels on the south and west dug before 1974 (Fig. 2). A pond barrow pit was dug for limestone, later filled in. The eight acre pit of dredged limestone, named Lake Lynda, was 40 ft deep in 1984( Fig 3). Currently  native Typha domingensis, (southern cattail) and Cladium jamaicense, (sawgrass) grow along its shore and invasive aquatic  Hydrilla verticillata (waterthyme) is submersed. .

The central ruderal areas had tilled row plantings of pine seedlings, but were tilled under in 2014 to prepare for a restoration planting of native trees and shrubs that were mostly hand dug into the limestone rock by Citrus County Florida Native Plant Society, (CCFNPS), later in 2014 (Table 4). Sedges Carex fissa, (hammock sedge), C. vexans, (Florida hammock sedge), and C. longii, (Long’s sedge) grow in the tilled ruderal area with other perennials and reseeding annuals like Ruellia caroliniensis, (wild petunia).

Vendor tour buses use a single entrance road from King’s Bay Road around Lake Lynda, a limestone quarry barrow, to the boardwalk where visitors walk around the springs to view manatees, sparing the natural vegetation and restoration plantings from trampling.


Access currently is a single entry road that is gated and locked. Public entry was only on specified days via shuttle buses. For this research, a Special Use Permit #12007-4156 was granted from US Fish and Wildlife Service at their headquarters in Crystal River NWR Complex. Upon each visit, USFWR magnetic signs were attached to my car and labeled yellow vests with SUP (Special User Permit) were worn. Old roads were traveled that were visible on the USDA Soil Survey Map of Citrus County in 1984 (Fig. 3).

Species were photographed in the field with macro-lens. The camera used by photographer James Curtis was Canon EOS5D Mark II that recorded location with GPS Data Logger, a Wireless File Transmitter motorized base, and 2 Wintec G-Rays GPS units mounted on the camera hot shoe. Photos were printed with imbedded GPS data. Although the data is included on labels sent with pressed specimens to herbaria, some curators black out coordinates to prevent poaching of endangered species by collectors.

Carex species of the family Cyperaceae are grass-like and some form leafy tufts to dense clumps in the herb layer of forests and clearings. They were hand-combed to find triangular culms with tubular sheaths and seed heads with sac-like perigynia, unlike the round-stemmed grasses with split sheaths and no sacs. Other species are rhizomatous and have leafy tufts in rows. Variability in clump density of C. styloflexa (bent sedge) and rhizome length of C. chapmannii (Chapman’s sedge) suggests they may be conspecific.

The culms were kept in beakers of rainwater before they were scanned and microimaged. Digital images were sent to Atlas of Florida Vascular Plants at the University of South Florida (USF). Seven Carex species specimens with label data were sent in 2012 -2014 to University of Florida-Tampa (USF), University of Florida Herbarium-Gainesville (FLAS) and Florida State University-Tallahassee (FSU).


Because Three Sisters Spring did not have a site-specific plant inventory except for Carex 2012 (Table 1), other inventories were consulted. The nearest site inventory in Citrus County was Crystal River State Preserve, compiled by Keith Morin, Park Biologist, and included more than 300 species including ten Carex discovered during research 2008- 2012 (Table 3). My research permit #002250812 to collect only Carex was granted by Florida Dept. of Environmental Protection and issued for five sites: Archaeological Museum State Park, Dixie Shores, Yoeman Park, Ecowalk, and Churchhouse Hammock.

The FDEP research permit in 2014 included other Crystal River Parks, Yulee Sugar Mill Runes, and Ellie Schiller Homosassa Springs Wildlife State Park. The lobally imperiled G2 Carex paeninsulae, (peninsula sedge), previously misidentified as C. godfreyi, (Godfrey’s sedge) was collected in April 2008 from Churchhouse Hammock, and annotated by Curator Bruce Hansen (USF).

C. paeninsulae was collected in the Archeological Museum State Park in April 2014. Species that grow in 6-20 planetary locations are G2 and G1 taxa are only in 1-5 populations, as determined by NatureServe, a status system. (

The next nearest inventory in Citrus County was Chassahowitzka National Wildlife Refuge, compiled by Stuart Marcus during 1982-83 which included 120 species listed with C. leptalea the only Carex. Later, C. gholsonii was discovered along the hydric forest along the Chassahowitzka River, west of the county park, in April, 1991. C. paeninsulae was found near Turtle Creek in Waccasassa Bay in Levy County, April, 1996 (Naczi, Bryson, Cochrane 2002).

Waccasassa Bay State Preserve (WBSP) in Levy County had more than 500 plants with five natural communities: tidal marsh, coastal hydric hammock, freshwater pools, basin (cypress) swamp, and mesic to scrubby flatwoods. Its coastal hydric forest hammocks were a mix of smaller communities that constituted, by quantitive analysis, one highly variable community (Abbot and Judd 2002). Nine Carex were listed in the Waccasassa Bay Preserve inventory and eleven species were collected from Crystal River Parks. Seven Carex were collected at Three Sisters Springs. Of the 15 total Carex species, six were in common to Three Sisters Springs and Crystal River Parks, both in Citrus County, and two in common with Waccasassa Bay in Levy County. Six of the nine Carex species at Waccasassa Bay State Preserve grew in shady hydric hammocks (Table 2). Those in brighter clearings and borders were C. fissa var. aristata, hammock sedge), C. godfreyi, (Godfrey’s sedge), and C. vexans, (Florida hammock sedge).

These sites were similar to the ruderal habitat of Three Sisters Springs, with associates of Andropogon sp. (bluestem grasses, (Poaceae), other sedges (Cyperaceae), rushes (Juncaceae), Erigeron vernus, (early fleabane, Asteraceae), Salvia lyrata, (lyreleaf sage, Lamiaceae) and Emilia fosbergii, (Florida tasselflower, Asteraceae).

TABLE 1. Carex of Three Sisters Springs 2012, 2014 collected by Linda Curtis

Species Original Namer Curtis date collected Common name Habitat
C. chapmanii Steud 4-10-12, 3-26-12, 4-3-14 Chapman’s Sedge State endangered Calcareous hydric hammocks
C. gholsonii Naczi & Cochrane 4-5-12, 4-3-14 Gholson’s Meadow Sedge Mesic to wet clearings aside forests, roadsides
C. godfreyi Naczi 4-10-12, 4-3-12 Godfrey’s Sedge Wet to mesic, calcareous hardwood hammocks
C. fissa Mack. var aristata F. J. Herm 4-3-14 Hammock Sedge Wet hammocks, clearings, roadsides
C. leptalea Wahlenb 4-10-12, 4-3-12 Bristly Stalked Sedge Swamps, wet hammocks
C. longii Mack 3-26-12, 4-5-12, 4-10-12 Long’s Sedge Marsh and coastal swales, upland clearings
C. styloflexa Buckley 4-10-12, 3-26-12, 4-3-14 C Bent Sedge Moist or wet hammocks
C. vexans R. J. Herm 4-17-12, 4-10-14 Florida Hammock Sedge Wet hammocks, marshes, roadsides

Fieldwork for Carex in Three Sisters Spring was conducted 2012 –2014 during sedge season in Central Florida beginning late March through June, although a few species re-bloom in autumn. Because most Carex cannot be identified solely by leaves, the specimens must have mature seed heads and identified by a key in either Guide to the Vascular Plants of Florida (2011) or also Flora of North Am. vol 23: Cyperaceae (2002).

TABLE 2. Carex of Waccasassa Bay State Preserve Inventory, Levy County

Species Original namer Common name Habitat
C. blanda Dewey Eastern Woodland Sedge Mesic and wet forests
C. chapmanii Steud Chapman’s Sedge Calcareous hydric hammocks
C. cherokeenis Schwein Cherokee Sedge Wet to calcareous forests
C. fissa var aristata Mack F.J. Herm Hammock Sedge Wet hammocks, clearings, roadsides
C. godfreyi Naczi Godfrey’s Sedge Wet hammocks, clearings
C. hyalinolepsis Steud Shoreline Sedge Marsh and coastal swales
C. lupuliformis Sartw ex Dewey False Hop Sedge Cypress swamps, river shores, wet hammocks
C. paeninsulae* see note Peninsula Sedge Mesic Hardwood Forests
C. vexans R.J.Herm. Florida Hammock Sedge Wet hammocks, clearings

*Naczi, Bryson, Cochrane 2002. Novon 12. no 4. Specimen 8382, Abbott, 4-12-1996

TABLE 3. Carex of the Crystal River Preserve State Parks 2008-2014 Linda Curtis

Species Collector Curtis date collected Common name Habitat
C. chapmanii Steud 4-20-08, 3-16-09 Chapman’s Sedge Calcareous hydric forested hammocks
C. dasycarpa Muhl 3-20-08, 4-24-08, 4-4-10, 4-21-14 Sandywoods Sedge Dry mesic, sandy upland
C. floridana Schwein 4-20-08 and 3-16-09 Florida Sedge Dry mesic, sandy upland
C. gholsonii Naczi & Cochrane 3- 16-09 Gholson’s Meadow Sedge Mesic to wet clearings aside forests, roadsides
C. godfreyi Naczi 4-26-09 Chapman’s Sedge Wet hammocks, clearings
C. leptalea Wahlenb 3-26-08 Bristly Stalked Sedge Swamps, wet hammocks
C. longii Mack 3- 26-09 Long’s Sedge Marsh and coastal swales
C.  lupuliformis Sartw.ex Dewey 4-4 08 False Hop Sedge Cypress swamps, river shores, wet hammocks,
C. paeninsulae Naczi 4-24-08   Peninsula Sedge Mesic hardwood forests, clearings
C. stipata Muhl 4-21-08 Awlfruit Sedge Marshes, wet clearings
C. vexans R.J. Herm 4- 24-09 Florida Hammock Sedge Wet hammocks, clearings

The native Carex associates at Three Sisters Springs include both forest understory and open ruderal species. Trees around the Three Sisters Springs lagoon were similar wetland trees along the roadside and canal-side perimeters. An illustrated page of seven common trees with simple leaves was submitted for the USFW annual report and included Acer rubrum, (red maple), Prunus serotina, (black cherry), Liquidambar styraciflua, (sweetgum), Juniperus silicicola, (red cedar), Ulmus americana, (American elm), Magnolia virginiana, (sweet bay), and Fraxinus caroliniana, (pop ash) (Fig 4).

Also seen were Celtis laevigata, (sugarberry), Psychotria nervosa, (wild coffee), Quercus virginiana, (live oak), Q. laurifolia, (laurel oak), Q. nigra, (water oak), and Tilia americana var. heterophylla, (white basswood).

Gleditsia aquatica, (water locust), a thorny native tree was planted between the plank walk and the springs. Native shrubs include, Itea virginica, (sweetspire), Salix caroliniana, (Carolina willow), Cornus foemina, (swamp dogwood), Baccharis halimifolia, (saltbush), and Erythrina herbacea, (Cherokee bean). The forested perimeters along roadside wet ditches of Cutler Spur Blvd. and King’s Bay Dr. were rich with Carex and Arisaema triphyllum, (Jack-in-the-pulpit), and other understory species such as Sambucus canadensis, (elderberry).

Central ruderal areas had scattered trees, mostly Sabal palmetto, (cabbage palm) and Juniperus virginiana, (red cedar), both evergreen. A potential witness tree, a stout red cedar along the entry road and between Lake Lynda and King’s Bay Road had a 10’ 8” circumference trunk, and its crown was visible on the USDA1984 soils map (Fig. 3). The map revealed a central agricultural landscape with tillage rows and scattered trees. The federally endangered wood storks (Mycteria americana) frequent the site.

During the summer of 2014, many trees along the new plank walk had been de-branched by chain-saw or removed. New railed viewing stations and a manatee rescue gate were added. Some trees that were overhanging the pool on eroded, undercut limestone banks were removed for safety issues, both for visitors and manatees.

The donation of large canopy shade trees will alter the understory species that currently grow in bright sun areas, and shade-tolerant species such as ferns and some Carex will increase. Many of the species planted were already seeding in from surrounding parent trees ensuring hydric forest succession. The native understory plants of ferns, shrubs, and flowering herbs should flourish, unless the site is treated as an arboretum with mowed lawns.

The coastal Gulf hammock forest at Homosassa Springs Wildlife State Park (HSWSP), South of Three Sisters Springs in Citrus County, had similar understory and three specimens collected were C. gholsonii, (Gholson’s sedge) C. godfreyi (Godfrey’s sedge) and C. longii (Long’s Sedge).

Table 4. Restoration trees and shrubs planted 2013 at Three Sisters Springs by Citrus Co. Chapter of Florida Native Plant Society (CCFNPS).


Acer rubrum-red maple
Acer negundo-box elder
Carpinus caroliniana-hornbeam
Carya aquatica-water hickory
Carya glabra-pignut hickory
Celtis laevigata-sugarberry
Fraxinus caroliniana-pop ash
Gleditsia aquatica-water locust
Ilex cassine-dahoon holly
Liquidambar styraciflua-sweet gum
Magnolia grandiflora-southern magnolia
Quercus shumardii-Shumard’s oak
Quercus virginiana-Virginia live oak
Sapindus saponaria-soapberry
Taxodium ascendens-bald cypress
Tilia heterophylla-basswood
Ulmus americana var. floridana– Florida elm

Shrubs and small trees

Euonymous americanus-strawberry bush
Viburnum obovatum-Walter’s Viburnum
Itea virginica-Virginia willow
Cornus foemina-swamp dogwood
C. asperifolia-roughleaf dogwood
Crataegus Marshalii-parsley haw
Chionanthus virginicus-fringe tree
Aesculus pavia-red buckeye
Hamamelis virginiana-witch hazel
Ptelea trifoliata-hop tree
Rhapidophyllum hystrix-needle palm
Sabal minor-blue stem palmetto
Hamelia patens-firebush
Forestiera ligustrina-upland swamp privet
Erythrina herbacea -coralbean
Salix caroliniana-Carolina shrub willow

Selecting the correct trees and shrubs for restoration are facilitated by knowledge of previous inventories for the designated area. Florida has 82 natural communities defined by the Florida Natural Areas Inventory. Google Earth images are also available in Florida Forever Projects.kmz.


Four of the seven Carex species at Three Sisters Springs Wildlife Refuge grow in the shady understory of gulf coast hydric hammock forest remnants around the spring, along the shady roadsides with creeks, wet ditches and on the canal banks. Three Carex species grow in the cut-over disturbed bright sunlight clearings with other ruderal species, introduced and native. Plant inventories from similar gulf hammock forests in Citrus County and one in Levy County were compared and six of fifteen Carex species were in common. The associates of Carex noted in this research will provide a partial inventory that will benefit management and volunteers as an aid for plant identification. The native gulf hammock forest is regenerating and the future understory will support birds, butterflies and other wildlife in one of the most-visited tourist sites in Citrus County.


Abbot, J.R. and W. S. Judd, 2000. Floristic Inventory of the Waccasassa Bay State

Preserve Levy County, Florida. Rhodora, Vol 102, No. 912, pp 439-513 Ball, P. W., and A. A. Reznicek, eds. 2001. Flora of North America, vol. 23.

Cyperaceae. Oxyford University Press, Inc. New York, N.Y.

Curtis, L.W. 2012. Carex of Three Sisters Springs. Special Use Research report for U.S. Fish &Wildlife Service (USFWS), Crystal River and

Environmental Protection Agency 2012 MYWATERS Google Earth.
Federally recommended best practices for stream bank and lakeshore stabilization (FISRWG 1998: USD-FS 2001)

Florida Communities Trust (FCT) 2010. Three Sisters Springs project management plan. Drafted by: The City of Crystal River, The U.S. Fish and Wildlife Service, and the Southwest Florida Water Management District Florida Communities Trust (FCT) Project #08-088-FF8.

Florida Natural Areas Inventory (FNAI) 2010. Guide to the natural communities of Florida: Florida Natural Areas Inventory, Tallahassee, Fl.

U.S. Fish &Wildlife Service, 2013, Coastal Wildlife Conservation Initiative, Nature Coast Region Working Group.

Herrington, S.J. 2012. Three Sisters Springs Focal Area, Conceptual Design for Freshwater Restoration. Draft 1, The Nature Conservancy, Fl Chapter.

Morin, Robin 2014. Crystal River Preserve Plant Inventory. 3266 N. Sailboat Ave., Crystal River, Fl 34428

Naczi, R.F., Bryson, C. T., Cochrane T.S. 2002. Seven New Species and One New Combination in Carex (Cyperaceae) from North America. Novon: 12, No. 4, pp. 508- 532.

NatureServe, 2014. NatureServe Explorer: An online encyclopedia of life.

USDA, 1984 Map # 21, Soil Conservation Service, Soil Survey Citrus Co. USDA

Vince, S. W, S.R. Humphrey, and R. W. Simons, 1989, The Ecology of Hydric Hammocks: A community Profile, U.S. Fish and Wildlife Service, Biological Report 85 (7.26)

Wunderlin, R. P., & B. F. Hansen, 2011, 3rd. ed. Guide to the plants of Florida. University Press of Florida, Gainesville.

Figure 1. Gulf hydric hammock forest along Cutler Spur Blvd., Crystal River, 1944

figure 1

Figure 2. Canals and barrow pit excavated to elevate canal banks. 1974.

 figure 2

Figure 3. Barrow pit filled, Lake Lynda excavated in center. 1984 Soils Map USDA

image 5 figure 3

image 6 figure 3

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