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2013 SOUTHEASTERN NATURALIST 12(4):790–808
Coefficient of Conservatism Rankings for the Flora of
Georgia: Wetland Indicator Species
Wendy B. Zomlefer1,*, Linda G. Chafin2, J. Richard Carter3, and David E. Giannasi1
Abstract - Wetland habitats currently cover about one-fifth of Georgia and have been
reduced in acreage by as much as twenty-five percent over the past two centuries due to
anthropogenic activities. Accurate identification and careful study of these areas are crucial
for their preservation and for compliance with federal and stat e environmental regulations.
Several vegetation-based biological assessment methodologies have been developed to define
wetlands and to assess their quality. One major wetland delineation system, mandated
by federal law, incorporates the National Wetland Plant List (NWPL), a classification system
ranking plant species in five indicator categories according to fidelity and preference
for wetlands or uplands. These rankings were recently updated via a comprehensive and
collaborative nationwide effort involving four government agencies and teams of wetland
specialists. Another expert-based indicator system, coefficients of conservatism, is the foundation
of the floristic quality index, a metric widely used in the United States for assessing
ecological condition of wetlands (as well as other plant communities). The coefficients
are based on breadth of habitat preference(s) and tolerance to disturbance, with exotic and
ruderal species receiving the lowest scores and ecologically conservative species assigned
the highest scores. A team of four botanists, proficient with the flora of Georgia, convened
to assign coefficient of conservatism rankings to the 2262 NWPL species for the state. The
resulting web-accessible database, which includes information such as regional wetland
rankings and conservation status, is described here.
Introduction
Georgia is a keystone for understanding the complex floristic diversity of
the southeastern United States; with over 3800 species (W.B. Zomlefer, D.E.
Giannasi, and J.R.Carter, unpubl. data), the state has been ranked seventh nationally
for vascular plant diversity (Stein 2002), second to Florida (≈4200 spp.;
Wunderlin and Hansen 2008) among the eastern states. Underlying this diversity
are fourteen major river systems (e.g., Altamaha, Chattahoochee, and Savannah),
extending from the upland physiographic regions to the outer Coastal
Plain, representing ancient migration corridors and Pleistocene refugia responsible
for complex floristic patterns with admixtures of diverse geographical origin
(Wharton 1978). Depending on the classification system, the state includes portions
of as many as six basic physiographic zones and ecoregions (Fig. 1A; e.g.,
US EPA 2012a, Wharton 1978). These areas have substantial floristic overlap
with both Atlantic and Gulf Coastal Plains and with subtropical Floridian and
Interior Continental systems.
1Department of Plant Biology, University of Georgia, Athens, GA 30602-7271. 2State Botanical
Garden of Georgia, University of Georgia, Athens, GA 30605-1674. 3Department
of Biology, Valdosta State University, Valdosta, GA 31698-0015. *Corresponding author
- wendyz@plantbio.uga.edu.
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Wetlands, in particular, contribute to the high level of biodiversity in Georgia.
As defined here, the hydrology of these habitats ranges from permanent inundation
(shallow water) to periodic soil saturation at or near the soil surface (seasonally waterlogged;
Tiner 2012). Approximately 20 percent (3,116,000 ha [7,700,000 acres])
of the state’s area is classified as wetland, including a wide variety of estuarine and
palustrine habitats. Estuarine wetlands comprise approximately 148,500 ha (367,000
acres), and the remaining 2,954,000 ha (7,300,000 acres) are forested, scrub-shrub,
and emergent freshwater wetlands (GA-DNR 2010a). A recent classification of
Georgia’s wetlands by physiographic province (Edwards et al. 2013) recognizes 14
general wetland types based on hydrologic origins (Table 1; Chafin 2011).
One of the most significant wetlands in Georgia is the extensive swamp and
bottomland system associated with the Altamaha River, the largest drainage basin
in the state (3,678,000 ha; 9,088,000 acres) and the third-largest freshwater
contributor to the Atlantic Ocean (GA-DNR 2004). Two other outstanding types
of Georgia wetlands defy easy classification and contribute disproportionally to
Georgia’s biodiversity: vernal pools (Piedmont granite outcrops) and the Okefenokee
Swamp (southeast Georgia Atlantic Coastal Plain). Piedmont granite
outcrop pools were created by weathering of certain granite types and provide
habitat for three federally listed plant species (Amphianthus pusillus Torr. [Pool
Sprite], Isoetes melanospora Engelm. [Black-spored Quillwort], I. tegetiformans
Rury [Mat-forming Quillwort]) and the state-listed Isoetes melanopoda J. Gay &
Durieu (Blackfoot Quillwort) (GA-DNR 2010b, 2011). The Okefenokee Swamp,
one of the largest freshwater wetland systems in the world (177,300 ha; 438,000
Figure 1. Ecological regions of Georgia. A. Environmental Protection Agency Level III
Ecoregions (US EPA 2012a). SA = Southwestern Appalachians. B. Wetland regions designated
by the US Army Corps of Engineers for the National Wetland Plant List (Lichvar
and Minkin 2008; USACE 2010, 2012a). Both maps modified by W.B. Zomlefer from the
referenced sources.
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2013 Southeastern Naturalist Vol. 12, No. 4
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acres), is a complex of wetland types fed mainly (80 percent) by rainwater and
has been designated a “wetland of international importance” by the United Nations
(Ramsar Convention on Wetlands 2012).
An estimated 20 to 25 percent of Georgia’s original wetland acreage has been
lost in the 230 years since colonial settlement (GA-DNR 2010a), and development
and encroaching urbanization continue to impact remaining wetlands. Accurate
identification and careful study of these areas are crucial for preserving them and
for compliance with federal and state environmental laws. Vegetation, an accessible
and conspicuous component of wetlands, can serve as a sensitive measure of
anthropogenic effects. Plant species characteristically have differential tolerances to
environmental change, and they have been utilized extensively as indicators to define
wetlands and in assessments of wetland condition and quality (Fennessy et al. 2002).
An example of a plant-based indicator scheme is the recently revised National
Wetland Plant List (NWPL; Lichvar 2012, Lichvar and Kartesz 2012), a ranking
system applied to 7828 species in the United States, which serves as a standard
research reference for wetland identification and delineation. The five wetland
indicator categories (Table 2) are based on qualitative ecological descriptions
of each species’ fidelity to and preference for wetlands or uplands (Lichvar et
al. 2012, Lichvar and Minkin 2008), depending upon its occurrence within 10
broad geographic regions (Lichvar 2012). Georgia lies within two of these regions
(Fig. 1B)—the Atlantic and Gulf Coastal Plain region (USACE 2010) and Eastern
Table 1. Summary of wetland types for the six Level III Ecoregions of Georgia (US EPA 2012a)
depicted in Figure 1A. Data from Chafin (2011) and Edwards et al. (2013). BR = Blue Ridge, P =
Piedmont, RV = Ridge and Valley, SA = Southwestern Appalachians (Cumberland Plateau), SCP =
Southern Coastal Plain (outer Coastal Plain and barrier islands), SP = Southeastern Plains (inner
Coastal Plain).
Ecoregion
Wetland type BR P RV SA SCP SP
Isolated freshwater wetlands
Depression marshes × ×
Depression swamps × × ×
Sag ponds, sinkholes × ×
Seepage forests, meadows, bogs, fens × × × × × ×
Spray cliffs ×
Wet savannahs, flatwoods × × × ×
Riverine wetlands
Beaver ponds × × × ×
Bottomland hardwood forests × × × ×
Floodplain swamps × × × × × ×
Tidally influenced wetlands
Interdunal wetlands ×
Salt marshes, brackish marshes ×
Sand bars, intertidal flats ×
Tidal freshwater marshes ×
Tidal swamps ×
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Table 2. Wetland-indicator status rankings and indicator-rank tallies and example species for Georgia. For more detailed descriptions of these categories
including plant characteristics and exemplar species, see Lichvar and Minkin (2008), Lichvar et al. (2012), and Lockwood (2012). For authorities and
common names of species given in the Table, see Lichvar and Kartesz (2012). Hydrophyte = plants that grow in water or on a substrate that is saturated
at a frequency and duration during the growing period sufficient to affect plant occurrence (Tiner 2012); wetland = a habitat with hydrologic conditions
ranging from permanent inundation (shallow water habitats) to periodic soil saturation at or near the soil surface (seasonally waterlogged habitats; Tiner
2012). AGCP = Atlantic and Gulf Coastal Plain region (USACE 2010), EMP = Eastern Mountains and Piedmont region (USACE 2012a), NWPL = National
Wetland Plant List (Lichvar 2012). For Georgia, the AGCP comprises 2241 species and the EMP comprises 2042 species. Percentages given in brackets
for number of species is for percentage of total in that region .
Number of Georgia
NWPL species
Short version
Ecological description of indicator descriptor AGCP EMP
Indicator status [code] (Lichvar and Minkin 2008) (Lichvar et al. 2012) [%] [%] Example species in Georgia
Obligate [OBL] Almost always a hydrophyte, Almost always occur 634 540 Alnus maritima, Lemna
rarely in uplands in wetlands [28.3] [26.4] minor, Nuphar lutea,
Pontederia cordata,
Taxodium distichum
Facultative Wetland [FACW] Usually a hydrophyte but Usually occur in wetlands, 564 466 Agalinis purpurea,
occasionally found in uplands but may occur in non-wetlands [25.2] [22.8] Cornus asperifolia, Quercus
laurifolia, Rhynchospora
plumosa, Viburnum obovatum
Facultative [FAC] Commonly occurs as either Occur in wetlands or 484 409 Agrostis scabra, Carpinus
a hydrophyte or non-hydrophyte non-wetlands [21.6] [20.0] caroliniana, Staphylea
trifolia, Ulmus rubra, Zizia
aurea
Facultative Upland [FACU] Occasionally a hydrophyte but Usually occur in non-wetlands, 501 538 Ambrosia artemisiifolia,
usually occurs in uplands but may occur in wetlands [22.4] [26.3] Carya ovata, Oenothera
biennis, Ostrya virginiana,
Prunus serotina
Upland [UPL] Rarely a hydrophyte, almost Almost never occur in wetlands 58 89 Acer spicatum [AGCP only],
always in uplands [2.6] [4.4] Actaea pachypoda [EMP],
Aristida longespica [EMP],
Rhus copallinum [AGCP],
Tillandsia usneoides [EMP]
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Mountains and Piedmont region (USACE 2012a)—and has a total of 2262 NWPL
species for the state (USACE 2012b).
Updating the NWPL was a well-established collaborative effort involving four
federal agencies: the US Army Corps of Engineers (USACE), the US Environmental
Protection Agency, the US Fish and Wildlife Service, and the USDA Natural
Resources Conservation Service. The wetland indicator rankings were assigned by
extensive review and consensus of national and regional panels, professional scientists,
and other technical experts, and are based on regional botanical/ecological
expertise and field observations, reappraisal of previous indicator status designations
(i.e., Reed 1988), literature review, and herbarium specimen data (details in
Lichvar 2012, Lichvar and Gillrich 2011, Lichvar and Minkin 2008). The official
list has a specific legal role in defining the hydrophytic vegetation parameter for
wetland delineation protocols required under Section 404 of the Clean Water Act
and the Swampbuster provision of the 2008 Farm Bill (Lockwood 2012). More
generally, the indicator rankings are also used to support wetland restoration and
research and as indicators to measure various environmental conditions (e.g., for
the National Wetland Inventory Program [USFWS 2013a]).
The coefficient of conservatism (C, C of C, or CoC) is another indicator system
with rankings assigned to plant species by regional experts and is also often used
in assessing wetlands, as well as other plant communities (Table 3). Each rank,
an integral value from 0 to 10, reflects a particular species’ fidelity to habitat and
tolerance of disturbance within a specified geographical region. The least conservative
species (exotic and ruderal; ranked 0–3) occur in a wide range of habitats and
thrive with disturbance, while the most conservative species (9 and 10) are habitat
specialists restricted to a narrow range of ecological conditions and are intolerant
of disturbance (Table 3; e.g., Swink and Wilhelm 1979, 1994; Taft et al. 1997).
Each coefficient, thus, represents an ordinal weighting factor reflecting the degree
of that particular species’ habitat fidelity in relation to other species of that region
(Andreas et al. 2004).
Coefficient values have been incorporated into community-based site assessment
methods for wetland biological integrity—in particular, the floristic quality
assessment index (FQI or FQAI), a robust metric of species richness used in evaluation
of natural areas and verification of mitigation and restoration efforts (Fennessy
et al. 2002, Lopez and Fennessy 2002). Assessment methodologies based on FQI
have been developed by regulatory agencies of several states (e.g., Bernthal et al.
2003, Herman et al. 2006, Taft et al. 1997) to monitor wetland conditions in compliance
with Section 305(b) of the Clean Water Act, including the current nationwide
effort, the National Wetland Condition Assessment (US EPA 2012b).
The FQI is most commonly calculated by multiplying the average C (“Mean C”)
by the square root of the total number of species in an inventory unit (Swink and
Wilhelm 1979, 1994; Taft et al. 1997), although there are modifications of this basic
equation, including the inclusion or exclusion of non-native species (see Andreas
et al. 2004, Bourdaghs et al. 2006). The Mean C relates directly to aggregate conservatism,
while FQI values are sensitive to factors that increase species richness.
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Table 3. Summary of coefficient of conservatism (C of C) ranking definitions used for the Georgia NWPL database, modified from Andreas et al. (2004).
NWPL = National Wetland Plant List (Lichvar and Kartesz 2012). For authorities and common names of species given in the Table, see Lichvar and Kartesz
(2012). Percentages given in brackets for number of species is for perc entage of total assigned C values.
Number of NWPL
C of C Georgia species
ranking Definition Examples of NWPL species in Georgia [%]
0 Invasive non-native species that displace native species, to th e Alternanthera philoxeroides, Ligustrum sinense, 36 [1.6]
extent of altering ecosystem function or community structure an d Murdannia keisak, Panicum repens, Pueraria montana
diminishing native species composition
1 Relatively benign non-native species Amaranthus spinosus, Morus alba, Murdannia nudiflora, 294 [13.0]
Persicaria hydropiper, Ranunculus sardous
2 Native invasives (native weedy species) very tolerant of Campsis radicans, Mikania scandens, Persicaria 126 [5.6]
anthropogenic disturbance and exhibiting a broad ecological hydropiperoides, Pinus taeda, Typha latifolia
amplitude
3 Native species with an intermediate range of ecological toleran ces; Acer rubrum, Andropogon glomeratus, Juncus effusus, 240 [10.6]
typifying a stable phase of some native community, but thriving Morella cerifera, Smilax glauca
under substantial natural or anthropogenic disturbance
4 Native species with an intermediate range of ecological toleran ces; Amorpha herbacea, Boehmeria cylindrica, Cornus 427 [19.0]
typifying a stable phase of some native community, but persisting amomum, Mimulus ringens, Vitis aestivalis
under moderate disturbance
5 Native species with an intermediate range of ecological toleran ces; Asimina parviflora, Chasmanthium laxum, Crataegus 448 [19.9]
typifying a stable phase of some native community, but persisting viridis, Quercus alba, Vitis vulpina
under limited disturbance
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Table 3, continued.
Number of NWPL
C of C Georgia species
ranking Definition Examples of NWPL species in Georgia [%]
6 Native species with a more or less narrow range of ecological Carphephorus corymbosus, Decumaria barbara, 352 [15.6]
tolerances, typifying a stable or near climax community (includ ing Pinguicula caerulea, Sapindus saponaria, Sarracenia
fire-dependent and other disclimax communities); tolerating limi ted minor
disturbance
7 Native species with a somewhat narrow range ecological Aesculus pavia, Calopogon tuberosus, Lindera benzoin, 203 [9.0]
tolerance, typifying a stable or near climax community; not Polygonatum biflorum, Ulmus americana
tolerating disturbance
8 Native species with a narrow range of ecological tolerance; Cypripedium parviflorum, Juglans cinerea, Parnassia 71 [3.1]
typifying a stable or near climax community; not tolerating grandiflora, Sabal minor, Salix floridana
disturbance; showing a moderate degree of habitat fidelity
9 Native species with a narrow range of ecological tolerance; Alnus maritima, Calopogon multiflorus, Helonias bullata, 35 [1.5]
exhibiting a high degree of fidelity to a narrow range of habita t Mitella diphylla, Sedum pulchellum
requirements; often listed as threatened or rare (but rarity no t due
to its distribution at the limit of geographical range)
10 Native species with a very narrow range of ecological tolerance ; Diamorpha smallii, Hymenocallis coronaria, Isoetes 20 [0.9]
exhibiting a high degree of fidelity to a narrow range of habita t melanospora, Trichomanes boschianum, Uniola paniculata
requirements; often listed as threatened or rare (but rarity no t due
to its distribution at the limit of geographical range)
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In general, a higher Mean C and FQI for a site indicate greater floristic quality and
biological integrity and a lower level of disturbance impacts. Thus, coefficient of
conservatism and floristic quality indices are relative values that provide baseline
reference data for tracking a single wetland over time and for comparing different
sites. Advantages and limitations (e.g., plant community effects) in their application
are summarized in Andreas et al. (2004), Bernthal et al. (2003), Bourdaghs et
al. (2006), Ervin et al. (2006), Lopez and Fennessy (2002), and Taft et al. (1997).
The National Park Service (NPS) is incorporating C and FQI indices as part
of a long-term ecological inventory and monitoring program initiated in 1999 to
establish baseline data on park ecosystems for resource management decisions (see
Fancy et al. 2009). This program included comprehensive floristic surveys of the
20 parks comprising the Southeast Coast Network (SECN), covering ≈74,460 ha
(184,000 acres) in Alabama, Florida, Georgia, North Carolina, and South Carolina
(DeVivo et al. 2008). Every SECN park includes considerable wetlands; six
parks are classified as inland riverine, and 14 as coastal (DeVivo et al. 2011). The
vouchered inventories established a species list for each park (e.g., Zomlefer and
Giannasi 2005; Zomlefer et al. 2004, 2012).
When the NPS contracted coauthors W.B Zomlefer, L.G. Chafin, and D.E. Giannasi
to generate coefficient of conservatism rankings for the ≈3000 SECN park
species, we discovered that Georgia lacked a C of C treatment, although indices
were available for some surrounding states in the Southeast (e.g., Mississippi [Herman
et al. 2006]; Florida [Mortellaro et al. 2012]). Based on our C-assignment
database for the NPS (W.B. Zomlefer, L.G. Chafin, and D.E. Giannasi, unpubl.
data), we subsequently assembled a coefficient list for Georgia. To create this new
database, we reassessed our SECN coefficient assignments with focus on the state,
rather than the five-state region, and limited coverage to wetland species, as defined
by the new NWPL.
Methods
We compiled a database for the 2262 NWPL species for Georgia (USACE
2012b), based on previous coefficient of conservatism assignments generated for
the NPS (W.B Zomlefer, L.G. Chafin, and D.E. Giannasi, unpubl. data), and including
newly generated C rankings for the state. The database, in Excel format,
comprises 10 columns (A–J) of information, detailed below.
Column A. Family
Family circumscriptions follow FNA (1993) for gymnosperms, Smith et al.
(2006) for lycophytes and monilophytes (“ferns and allies”), and APG III (2009)
for angiosperms.
Column B. Scientific name
These are the official names (required for reporting federal wetland delineations)
taken directly from the National Wetland Plant List (Lichvar and Kartesz 2012) that
adopted the nomenclature of the Biota of North America Program (Kartesz 2009).
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Column C. Authority
The authorities are from Tropicos® (http://www.tropicos.org/) and The International
Plant Names Index (IPNI; http://www.ipni.org/), using standard author
abbreviations established by IPNI.
Column D. Nativity
The native status follows the Flora of the Southern and Mid-Atlantic States
(Weakley 2012), which incorporates the most current information on non-native
taxa (whether naturalized, persistent, waif, etc.) targeting the primary southeastern
flora area. Weakley also indicates and/or discusses species of questionable nativity.
The designation in the database reflects the probable status for Georgia. An asterisk
indicates some uncertainty about nativity in the state (as noted by Weakley 2012),
briefly summarized in “Column J. Nativity and other notes”.
Column E. Coefficient of conservatism rankings
The coefficient rankings were assigned by a team of the following four professional
plant taxonomists, representing many years of regional field experience:
(1) J. Richard Carter (Curator, Valdosta State University Herbarium) is an expert
on sedges (e.g., Bryson and Carter 2008, Carter and Bryson 2000) and floristics in
Georgia, including rare plant surveys (e.g., Carter et al. 2009); (2) Linda G. Chafin
(Conservation Botanist, State Botanical Garden, University of Georgia) has particular
expertise in floristics and rare and invasive plants of Florida (Chafin 2000)
and Georgia (Chafin 2007); (3) David E. Giannasi (Emeritus Director, University
of Georgia Herbarium) taught wetland plant courses for over 20 years and has been
involved with plant surveys throughout the state (e.g., Zomlefer et al. 2010, 2013);
and (4) Wendy B. Zomlefer (Curator, University of Georgia Herbarium) studies
petaloid monocots (e.g., Zomlefer et al. 2006) and has collected extensively in
Florida (e.g., Zomlefer 1994, Zomlefer et al. 2007) and Georgia (e.g., Zomlefer et
al. 2008, 2012). Chafin, Giannasi, and Zomlefer compiled the original coefficient
list, and Carter reviewed the previous assignments for the NPS park taxa and also
independently ranked over 600 species (Cyperaceae, Poaceae, and several other
crucial taxa such as Juncus and Xyris).
The eleven rankings (Table 3) generally follow the recommendations detailed in
Andreas et al. (2004) with the exception of the assignment options for non-native
species. As advocated by Michael Byrne, Terrestrial Ecologist for the NPS SECN
(Cumberland Island National Seashore, St. Marys, GA, pers. comm.), exotics are
here treated more precisely—in two ranks (0 for invasive; 1 for relatively [and
currently] benign) —rather than as in one group (all ranked 0). We devised a dichotomous
key as a guide for assigning coefficient rankings (Fig . 2).
The coefficient assignments were based on the authors’ collective field experience,
supplemented with specimen label data from the extensive herbarium
collections at the University of Georgia (264,000 specimens) and Valdosta State University
(65,000), and regional manuals such as Cronquist (1980), Godfrey (1988),
Godfrey and Wooten (1979, 1981), Isely (1990), Radford et al. (1968), Snyder and
Bruce (1986), Weakley (2012), Wunderlin and Hansen (2000, 2011), and appropriate
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volumes of the Flora of North America (FNA 1993+). The team also considered various
state and federal indices of conservation or invasive status (see “Columns F–G.
NWPL rating” and “Column H. State and federal rankings” below).
We tested our assignment criteria by carefully deliberating coefficients for a
sample set of 50 species and comparing our results. Coauthors Chafin, Giannasi,
and Zomlefer then convened in a series of 14 face-to-face meetings (27 Feb.–23
Nov. 2009; 29 Oct. 2012–31 Jan. 2013) and conference calls (later joined by coauthor
Carter) to reevaluate our previous rankings for 1499 species extracted from the
NPS SECN list (W.B. Zomlefer, unpubl. data) and to generate new C of Cs for the
Dichotomous Key for Coefficient of Conservatism Rankings
(a) Non-native species..................................................................................................(b)
(b) invasive ………….……………………….............……..……………...…...rank 0
(b) relatively benign ………….……..……………….……..….............……….rank 1
(a) Native species……………………………………………….….............………...….(c)
(c) opportunistic, broad range of ecological tolerance, more or less restricted
to areas subject to human disturbance….……………...............……….…..rank 2
(c) non-opportunistic, intermediate to narrow range of ecological tolerance...........(d)
(d) intermediate range of ecological tolerance, typifies a stabl e phase of
some native community, thrives and/or persists under natural or
human disturbance…………………....…………........…...……...........…....(e)
(e) persists and/or thrives under natural or human disturbance..............rank 3
(e) persists but does not thrive under limited natural or human
disturbance…………………………...............………….................…….(f)
(f) persists with some disturbance………………..…...............….rank 4
(f) persists with a little disturbance………………….............…... rank 5
(d) narrow range of ecological tolerance, typifies a stable or n ear
climax community (including fire-dependent disclimax communities ),
tolerates little to no disturbance (unless surrogate for fire or other
natural disturbance)…………………..…..…..............................................(g)
(g) moderate fidelity to a narrow habitat requirement, may or ma y
not tolerate limited disturbance………………..…….…….…..........…..(h)
(h) more or less narrow range of ecological tolerance, tolerate s
limited disturbance...................................................................rank 6
(h) narrower range of ecological tolerance, does not tolerate
disturbance......................................................................................(i)
(i) somewhat narrow range of ecological tolerance.............. ....rank 7
(i) narrow range of ecological tolerance……..……................. rank 8
(g) high fidelity to a narrow range of habitat requirement, does not
tolerate disturbance……………………… ………………..........…….(j)
(j) narrow range of ecological tolerance, relatively high fideli ty
to a narrow range of habitat requirement………................…. rank 9
(j) very narrow range of ecological tolerance, very high fidelit y
to a very narrow range habitat requirement…………............rank 10
Figure 2. Dichotomous key for assigning coefficient of conservat ism rankings.
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balance of 763 species to complete the Georgia NWPL. Throughout the process, we
periodically confirmed consistent application of the concept of conservatism—for
example, that a rare plant did not always receive a high coefficient ranking, and a
widespread species was not always necessarily assigned a low ranking (see “Results”).
Consensus on conflicting coefficients was achieved through discussion of
the particular species and consultation of appropriate referenc e data.
Columns F–G. NWPL rating
The database lists the NWPL wetland indicator rankings for 2241species of the
Atlantic and Gulf Coastal Plain region (AGCP; USACE 2010) and 2042 species
for the Eastern Mountains and Piedmont region (EMP; USACE 2012a), with an
overlap of 2021 species (89.3%) in common for these two regions in Georgia. The
rating for a particular species may vary depending upon the reg ion.
Column H. State and federal rankings
Government agency rankings in this column include Georgia non-native invasive
categories (GA-EPPC 2006) and state, federal, and global conservation
status/rank (GA-DNR 2010b, 2011; USFWS 2013b) for listed species. These
rankings are defined and referenced under the “Legend for Wetland Plant List”
tab of the database.
Column I. Synonyms
When differing from the NWPL, the scientific name (synonym) used in Weakley
(2012) is provided for database users since this flora is the primary source for
identifying plants in Georgia. Additional commonly used synonyms are included
in Weakley (2012), and more comprehensive listings are available at the PLANTS
Database (http://plants.usda.gov) and the Integrated Taxonomic Information System
(http://www.itis.gov/).
Column J. Nativity and other notes
The annotations in this column briefly address concerns about nativity (see “D.
Nativity”, above) and on the dubious occurrence of a particular species in Georgia.
Noted here also are pertinent intraspecific issues, including species with subspecies
or varieties assigned state or federal rarity rankings and/or species with both native
and non-native infraspecific elements.
Results
For the complete database including new coefficient of conservatism rankings
for the 2262 species of the two NWPL wetland regions of Georgia, see
Supplemental File 1, available online at https://www.eaglehill.us/SENAonline/
suppl-files/s12-4-S1195-Zomlefer-s1, and, for BioOne subscribers, at http://dx.doi.
org/10.1656/S1195.s1. Coefficient values were assigned to all species supported
by vouchers at the University of Georgia and Valdosta State University herbaria. A
few species (such as Proserpinaca intermedia Mack. [Intermediate Mermaidweed])
lacked vouchers for the state but were indicated in historical or Georgia Department
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2013 Southeastern Naturalist Vol. 12, No. 4
of Natural Resource records as occurring in Georgia. These were ranked with
reference to literature and specimen-label data from adjacent states. However, 10
species (e.g., Thuja occidentalis L. [Northern White Cedar] and Carex vexans F.J.
Herm. [Florida Hammock Sedge]) included in the NWPL very likely never have
occurred in Georgia and remain unranked.
Table 3 and Figure 3 summarize the number of species in each coefficient category
for the remaining 2252 species. Three hundred and thirty species (≈14.6%) on the
list are non-native (C = 0 or 1): 36 are ranked as invasive exotics (C = 0), and 294, as
relatively benign (C = 1). The frequency distribution of coefficients of conservatism
ranks for native species (C = 2–10; 1922 spp.; Fig. 3, light gray bars) is somewhat
skewed towards the less conservative end of the spectrum due to the highest number
of species ranked at C = 4 (427 spp.), 5 (448 spp.), and 6 (352 spp.). The most-generalist
natives (native invasives; C = 2; 126 spp.) and the most-conservative species
(C = 8, 9, 10; collectively 126 spp.), comprise the lowest number of species.
Discussion
Since the NWPL specifically targets wetland species in the state, we initially
expected somewhat higher-end fidelity reflected in the C values. However,
rankings 4–6 comprised ≈63.4 percent of the native species on the list (Fig. 3).
Figure 3. Bar graph depicting distribution of coefficient of conservatism designations for
the 2252 Georgia National Wetland Plant List (NWPL) species (see Table 3; USACE 2012b)
ranked for this project.
W.B. Zomlefer, L.G. Chafin, J.R. Carter, and D.E. Giannasi
2013 Southeastern Naturalist Vol. 12, No. 4
802
This result reflects that wetland-adapted species may be likely to survive—despite
disturbance—as long as sufficient water is available. Further, wetland
species are generally tolerant of a wide range of soil moisture levels, and some
wetland habitats, such as alluvial floodplains and exposed bars and banks along
streams and coasts, are naturally maintained by disturbance. Many wetland
species are heliophytes, adapted to exposed sunny areas with reduced competition
from taller shading trees and shrubs in these communities dependent on
disturbance (Bryson and Carter 2008, Carter 2005). Various wetland species,
such as sedges, have intrinsic characteristics (e.g., rapid growth, vegetative
proliferation, extended seed dormancy) that promote population expansion after
disturbance and may have originally evolved as colonizers of disturbed habitats
(Baker 1965, 1974).
As reported for other large-scale regional C-assignment efforts (e.g., Bried et
al. 2012), members of our team usually were most confident ranking the least (C =
2 or 3) and most (C = 8–10) conservative native species. For example, several ubiquitous
natives with low coefficients, such as Ambrosia artemisiifolia L. (Common
Ragweed) (C = 2 in our scale), likely would receive similar C value assignments
throughout their geographical range. Other species, such as Saxifraga micranthidifolia
(Haw.) Steud. (Branch-lettuce) (C = 9), are so restricted in their ecological
requirements, they were obvious candidates for the other extreme of the coefficient
scale. The most difficult rankings involved assignment of species to coefficient 6 vs.
7, particularly when floras and herbarium specimen labels provided limited habitat
data. Tree species (e.g., Quercus alba L. [White Oak], Fagus grandifolia Ehrh.
[American Beech]), which may be long-lived components of climax forests, were
also sometimes difficult to assign with certainty due to lack of information about reestablishment
of populations after disturbance.
Various indices for rare or invasive species were helpful guides but not necessarily
definitive for assigning C values. Many species of conservation concern are
indeed highly conservative (C = 9 or 10) and restricted to specific remnant natural
communities. An example is Sedum pusillum Michx. (Granite Stonecrop) (stateranked
threatened; C = 10), confined to granite outcrops, a rare habitat (Chafin
2007). However, since rarity is not always due to limited ecological tolerance (Taft
et al. 1997), conservation status did not always correlate with high conservatism
rankings. For example, some rare species, such as Fimbristylis perpusilla R.M.
Harper ex Small & Britton (Harper’s Fimbry; state-ranked endangered, C = 4),
can thrive in very disturbed areas, while others, such as Pinguicula primulifolia
Wood & Godfr. (Clearwater Butterwort) (threatened; C = 6), typically occur in
pristine habitats but exhibit some tolerance of disturbance (Godfrey and Wooten
1981). Furthermore, several highly conservative species are not rare in Georgia,
such as Uniola paniculata L. (Sea Oats) (not state listed; C = 10), a perennial grass
restricted to beach dunes (not an uncommon habitat).
The coefficient key (Fig. 2; see “Methods, Column E. Coefficient of conservatism
rankings”) provided an effective approach for ranking species, allowed
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W.B. Zomlefer, L.G. Chafin, J.R. Carter, and D.E. Giannasi
2013 Southeastern Naturalist Vol. 12, No. 4
modification as the team gained experience, and serves as a model for other state
floras. Overall, team members exhibited some bias due to differing perspectives,
from one career focused on rare species vs. others emphasizing broad ranging
floristic surveys and/or collecting in disturbed areas for instructional purposes.
Generally, one botanist consistently assigned lower or higher coefficients compared
to another. An informal comparison of Chafin vs. Zomlefer and Carter vs. Zomlefer
of the assignments for ≈300 non-graminoid species (using our key) revealed
approximately one-third of species with the same rank, and another one-third,
within one integer. Most discrepancies were resolved by direct field experience
and personal observation confirming a particular species’ sensitivity to disturbance.
Despite differences in opinion for ranking species, team effort is valuable, especially
when members have wide-ranging field experiences with the flora.
These coefficient assignments represent the first such effort for Georgia and
will facilitate floristic quality assessment in the state. The National Wetland
Plant List for Georgia, which includes at least 60 percent of the species in the
state, serves here as a foundation for developing C coverage for the entire flora.
The values were applied provisionally in a study of the Atlantic Coastal Plain
limestone forest association (Lynch 2012) and were provided to S. Fennessy
(Kenyon College, Gambier, OH, pers. comm.) for data analysis as part of the
National Wetland Condition Assessment project (US EPA 2012b) and to K. Gianopulos
(North Carolina Department of Environment and Natural Resources,
Raleigh, NC, pers. comm.) as reference data for the Southeast Wetlands Workgroup
(SEWWG 2013). We encourage further input and refinement of these
C-assignments from experts as we continue forward with our goal of ranking
the entire flora of Georgia—including intraspecific taxa (varieties, subspecies)—
based on a vouchered checklist compiled from an ongoing collaborative
herbarium digitization endeavor supported by the National Science Foundation
(Wichmann et al. 2012, Zomlefer and Carter 2012).
Acknowledgments
We thank Richard Reaves for alerting us to the revised National Wetland Plant List
(then in progress) and for providing appropriate Web links, Walter S. Judd for confirming
family circumscriptions in the database for conformance to APG III, Brenda
L. Wichmann for supplying information on updated NatureServe conservation-status
designations, Sharon Carter for assistance checking specimen label data at the Valdosta
State University Herbarium, and Harald Scherm for helping with the graph. Siobhan
Fennessey and two anonymous reviewers generously provided constructive criticisms of
the manuscript. We also appreciate the support of National Park Service personnel: Joe
DeVivo, Network Coordinator for the Southeast Coast Network (SECN), and Michael
W. Byrne, Terrestrial Ecologist, for the initial phase of this project that focused on coefficients
of conservatism for SECN park plants. Portions of this work were funded by
National Park Service contract agreement J2114-08-0021 (W.B. Zomlefer, PI) and the
National Science Foundation, DBI 1054366 (J.R. Carter, PI) and DBI 1054329 (W.B.
Zomlefer, PI, and D.E. Giannasi, coPI).
W.B. Zomlefer, L.G. Chafin, J.R. Carter, and D.E. Giannasi
2013 Southeastern Naturalist Vol. 12, No. 4
804
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