Carex and Associates of Three Sisters Springs 2015

A new report: Carex and Inventory of Three Sisters Springs Wildlife Refuge 2015  is available from the author at  The previous 2014 report follows this 2025 brief update.

3 sister springs view n. June

Photo by Joyce Kleen USF&W Crystal River October 5, 2015.Three Sisters Springs Wildlife Refuge (TTSWR) was entered by tour buses previously from Kings Bay Road from the north entry until this year when a new entrance and roadway were constructed across Cutler Blvd, now named Three Sisters Springs Trail on the east side during a bulldozing and  dredging for a tidal water treatment pond (Edmunds 2012). Visitors still can enter via the tour buses or ride the new Crystal River trolleys with several pick up stops in Crystal River. No car parking is allowed except for staff and researchers with USF&W Special Use Permits SUP. A gated guard station at the entry monitors visitors and researchers. Three Sisters Spring was opened to the public when manatee viewing season began on Nov. 15, 2015. Manatees were reported mating in the lagoon on Dec. 24, 2015. High tides were record 6′ Jan.17, 2016.









The serpentine basin for a new water treatment pond was dredged in 2015 on the east portion of TTSWR and some of the spoils were placed in the southeast end  an artificial lake dredged in the 1970’s. Formerly named Lake Lynda, it is now Lake Crystal. As a seed bank, that area is very interesting and has more than seeds, as the Jack-in-the-pulpit corms were in the forest understory. One plant had an albino spathe. Another was 3 feet tall and would have a corm larger than an baking potato tuber.

Rows of planted pine seedlings were removed by mowing and tilling, but existing trees were left,  mainly red cedar, Juniperus virginiana and tall cabbage palms, Sabal palmetto. Once a pristine gulf hydric forest into the 1960’s, the repeated attempts at development since has altered its natural return to hydric forest. Natural areas reproduce their former ecosystem from roots and seeds.

No plant inventory existed at Three Sisters Springs. The USFWS manage the Refuge and granted me permits to collect. I began a  working list of other plants from my 4 years of Carex field search that listed associates. Also, the plant inventory from Crystal River Parks and Preserves (Morin 2014) and Waccasassa Bay State Preserve (Abbott & Judd 2000) were consulted to form a check list to help document the TTSWR  species as they were identified. The WBSP inventory was updated to the newest nomenclature following the forthcoming edition of Gleason and Cronquist.

Most of the native plants used in and around the water treatment pond were not previously on the property. Three Sisters Treatment Wetland Conceptual Design (W471) was prepared for Southwest Florida Water Management District, March 27, 2012  by Wetland Solutions, Inc., Jones Edmunds.  The final 2015 report was sent to USFWS Jan 11, 2016. One error was found in  species list table and that was the pestiferous Brazilian Pepper Schinus terebinthifolia whose family name is Anacardiaceae. It also was misspelled …folius.

One person reported that error, Jim Solomon at Mobot. Thanks, Jim.

Other errors reported by Chris Davis, in Malvaceae, I misspelled genera Kosteletzkya,

and Abutilon. Thank you Chris. Do email Linda at if you spot an error.

Carex and associates of Three Sisters Springs Wildlife Refuge

2014 Based on the Report by Linda Curtis, botanist, for USF&W.

In comparison of 2014 & 2015 aerial images, note the continuous hydric forest fringe along the tidal creek along Cutler Spur Boulevards. About 100 feet of forest and understory were removed to put in new entry.

Three Sisters Springs

Photo by Joyce Kleen-USFWS

(USFWS) is part of the Florida Gulf Coastal lowland, Crystal River, Kings Bay, Citrus County, Florida. Above image faces West with Cutler Spur Boulevard on the lower portion of the aerial photograph.

Above image: 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. Lake Lynda in center is a dredged barrow.The forested fringe along Cutler Spur Blvd  has a varied understory of herbaceous plants and shrubs that included 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 University of Florida-Tampa (USF), University of Florida-Gainesville (FLAS), and Florida State University-Tallahassee (FSU).

Carex and associates of Three Sisters Springs, Crystal River, Florida by Linda Curtis 2014

Report filed Dec 31, 2014 by Linda W. Curtis, USFWS contacts were Joyce Kleen,, and Ivan Vincente contact

Abstract: 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 2014. 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 was 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 N28 53.519 and longitude W82 35.469. The parking lot for tour bus drop-off is near the boardwalk. The southern boundary canal is 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 60o 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 (Fig. 2, map 1974). A pond barrow pit was dug for limestone, later filled in. The eight acre pit of dredged limestone, named Lake Lynda, is 40 ft deep and has native Typha domingensis, (southern cattail) and Cladium jamaicense, (sawgrass), along its shore and invasive  submersed Hydrilla verticillata (waterthyme) (Fig. 3. Map 1984).

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 3). 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 (Fig. 3, Map 1984).

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).


Three Sisters Spring did not have a site-specific plant inventory except for Carex (Table 1), therefore 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) was previously misidentified as C. godfreyi, (Godfrey’s sedge) and was collected in April 2008 from Churchhouse Hammock, and annotated by Curator Bruce Hansen (USF).

C. paeninsulae was collected in the Archaeological 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 and 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. 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  2012-14 Linda Curtis, Collector

Scientific name                             Common name                         Habitat
C. chapmanii Steud.                        Chapman’s Sedge            Wet  calcareous hammocks

C. gholsonii, Naczi & Cochrane     Gholson’s  Sedge             Wet clearings   

C. godfreyi Naczi                               Godfrey’s Sedge              Wet  calcareous hammocks

C. fissa Mack. var aristata Herm  Hammock Sedge              Wet  clearings 
C. leptalea Wahlenb                         Bristly Stalked Sedge      Swamps, wet hammocks

C. longii Mack                                     Long’s Sedge                     Clearings, roadsides

C. styloflexa Buckley                         Bent Sedge                        Moist or wet hammocks
C. vexans R. J. Herm                         Florida Hammock Sedge     Wet hammock

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. Most Carex cannot be identified solely by leaves and the specimens must have mature seed heads and identified by a key in  Guide to the Vascular Plants of Florida (2011) or Flora of North Am. vol 23: Cyperaceae (2002).


Carex of Crystal River Preserve State Parks 2008-14 Linda Curtis

Scientific name                              Common Name                            Dates collected

C. chapmanii Steud.                     Chapman’s Sedge                          4-20-08, 3-16-09

C. dasycarpa Muhl.                       Sandywoods Sedge         3-20-08,  4-4-10, 4-21-14

C. floridana Schwein.                   Blackedge Sedge                           4-20-08, 3-16-09

C. gholsonii Naczi & Cochrane    Gholson’s Sedge                                         3-16-09

C. godfreyi Naczi                             Godfrey’s Sedge                                        4-26-09

C. leptalea Wahlenb.                       Bristly stalked Sedge                               3-26-08

C. longii Mack.                                Long’s Sedge                                                3-26-09

C. lupuliformis Sartw. ex Dewey  False Hop Sedge                                           4-4-08

C. paeninsulae Naczi                       Peninsula Sedge                         4-24-08, 4-19-14

C. stipata Muhl.                               Awlfruit Sedge                                           4-21-08
C. vexans R. J. Herm.                     Florida Hammock Sedge                           4-24-09

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 2014 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 plus shrub 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 USDA 1984 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 planted 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. Vegetation skirts around trees in mowed areas preserves wildflowers.

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 3. Restoration trees and shrubs planted 2014 at Three Sisters Springs by Citrus Co. by members of Citrus 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.

Special thanks to Richard Wunderlin, Dick Hansen, Kent Perkins, Austin Mast

Figure 2. Below: Three Sisters Spring surrounded by Gulf Hammock Hydric Forest 1944, Highway 19 us far right, then Cutler Spur.

figure 1 Figure 3. Below: Canal and pond dredged in 1970’s preparing for development.


figure 2


Figure 4 below: Soils Map 1984 with dredged Lake Lynda.

image 5 figure 3


image 6 figure 3


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