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J. Bot. Res. Inst. Texas 8(2): 419 – 430. 2014
A MORPHOLOGICALLY BASED TAXONOMIC REEVALUATION
OF THE GENUS STIPULICIDA (CARYOPH YLL ACEA E),
WITH COMMENTS ON RANK
Derick B. Poindexter1, Kateland E. Bennett, and Alan S. Weakley
UNC Herbarium (NCU)
North Carolina Botanical Garden
University of North Carolina
Chapel Hill, North Carolina 27599, U.S.A.
and
1Botanical Research Institute of Texas
1700 University Drive
Fort Worth, Texas 76107, U.S.A.
poindexter@unc.edu
abstr act
Stipulicida is a nar rowly endemic , monotypic ge nus as cur rently cir cumscr ibed. Pre vious stud ies have att empted to di agnose add itional t axa
with in the genus, but only one species comprised of t wo varieties i s currently accepted. These two v arieties, St ipulicida seta cea var. setacea
and S. seta cea var. lacerata , demonstrate significant phenotypic divergence and idiosyncratic biogeograph ical patterns. We utilized multi-
variate analyses a nd nonpar ametr ic analyses of varia nce to ascerta in morphological distinctions between these two ta xa and asses s bio-
logica lly appropri ate taxonom ic rank. Ba sed on our morph ometric dat a and biogeogr aphy, we elevate S. setace a var. lacerata to species a nd
provid e a revised ke y to the specie s in the genus.
resum en
Stipulicida es un pequeño género endémico, monotípico tal como se circunscribe nor malmente. Est udios previos habían intentado diag-
nostic ar taxa ad icionales en e ste género, pero sol o se acepta norm almente una e specie con dos va riedade s. Estas dos va riedades , Stipulicida
setacea var. setacea y S. setacea va r. la cerata, muestran una divergencia fenotípica y patrones biogeográficos significativos. Utilizamos un
análisis multivariante y un análisis no-paramétrico de varianza para e stablecer diferencia s morfológicas entre estos dos taxa y valorar el
rango t axonómico apropi ado biológicamente. Ba sados en nuestr os datos morfomét ricos y biogeog rafía, elev amos S. setacea var. lacerata a
espe cie y proporciona mos una clave r evisada de e species de l género.
int roduction
The monotypic genus Stipulicida Michx. (Caryophyllaceae; Paronychioideae; Polycarpeae) is endemic to the
southeastern United States and Cuba (Swanson & Rabeler 2005; Weakley 2012; Kartesz 2013). It consists of
minimalist plants with slender stems, ephemeral basal rosettes, short awl-shaped opposite leaves with incised
stipules, and diffuse, divaricately-branched inflorescences. The flowers are the most conspicuous mor phologi-
cal feature, consisting of 5 dimorphic sepals (often referred to as “inner” and “outer”), 5 white petals, with the
terminal clusters subtended by bracts similar to the leaves. The genus was first described by Andre Michaux
(1803) from collections made “in sabulosis aridis Carolinae” (= sandy, dry soils of the Carolinas) (Fig. 1). Stipu-
licida as a genus has remained unscathed by current phylogenetic and systematic studies of the Caryophylla-
ceae, perh aps only because of its omission from them. In the most recent molecular phylogenetic studie s of the
Caryophyllaceae (e.g., Fior et al. 2006; Harbaugh et al. 2010; Greenberg & Donoghue 2011), Stipulicida has
been one of the few genera omitted. Consequently, exact subfamilial and tribal placement has not been con-
firmed and current circumscription is based solely on morphology. At the time of its description, the genus
was represented by a single species, Stipulicida setacea Michx.
Nash (1895) added another putative species from central peninsular Florida, Stipulicida filiformis Nash,
which allegedly differed from S. se tacea based on a much more slender habit, fewer, sessile flowers, and shorter
bracts. Small (1903, 1913, 1933) maintained this taxon as distinct within his influential manuals and added
420 Journal of the Botanical Research Institute of Texas 8(2)
Fig. 1. One of two presumed isotypes of Stipulicida setacea (Michaux s.n., P, image #P00156962).
Poindexter et al., Reevaluation of Stipulicida (Caryophyllaceae) 421
petal shape (constricted at the middle vs. spathulate) as an additional discriminating feature. Judd (1983)
sought to address the taxonomic status of S. filiformis by conducting extensive fieldwork and observing varia-
tion in situ, in addition to morphometric analyses from herbarium specimens. He focused on two quantitative
features, stem diameter and number of flowers/inflorescence. Judd (1983) concluded that the plants intergrade
broadly with respect to these characters, while also stating that bract size and petal shape failed to provide
separation into two taxa, and consequently recognized S. filiformis as a mere morphological extreme or eco-
typ e of S. setacea, though commenting that its apparent li mitation to the Cent ral Florida Ridge wa s noteworthy
due to the number of taxa that are distributed similarly as endemics of xerophytic sands of this region. Ward
(2001) felt this pattern (in combination with intergrading morphological distinctions) was sufficient to war-
rant maintaining it at varietal rank, Stipulicida setacea va r. filiformis (Nash) D.B. Ward, in order to preserve
some taxonomic distinction.
James (1957) described the only additional member of this genus to date by recognizing a new variety,
Stipulicida setacea var. lacerata C.W. James, restricted to Florida and the Isle of Pines (now the Isla de la Juven-
tud), Cuba. This variety was considered distinguishable from the typical variant by the presence of lacerate
sepals and outer sepals that were mucronate, as well as “essentially allopatric” distributions. James’ (1957) ra-
tionale for infraspecific rank was explicit:
“The collec tion of F.W. Hunnewe ll (no. 8955, Highl ands Co., Fla.), from the e astern lim its of the ran ge of var. lacera ta was t he only one seen
which con sisted of pla nts of both va rieties. The sp ecimen of var. la cera ta in t his collect ion is somewhat at ypical. Th is, in addit ion to the fact
that t here is not a clea r-cut di scontinuit y in the shape of t he sepals, i ndicates t hat there may b e some gene flow b etween thes e taxa.”
During the course of his study, Judd (1983) also noticed the distinctive nature of lacerate-sepaled plants and
felt they were adequately assigned to varietal status due to presence of intermed iate plants from Highlands and
Volusia Counties.
At pre sent, the general consensus (as shown by recent floras covering all or parts of the distribution of the
genus: Wunderlin & Hansen 2011; Swanson & R abeler 2005; Weakley 2012) has been to follow the taxonomic
schema established by Judd (1983) by recognizing a single species within the genus, with two relatively well-
marked varieties, Stipulicida setacea var. setacea (incl. S. filiformis) and S. setacea var. lacerata. The distribution
of the genus as a whole is narrow, restricted to the Atlantic and Gulf Coastal Plain (Fig. 2). Of these two variet-
ies, var. setacea is the most widespread, extending from southeastern Virginia to Florida and west to eastern
Louisi ana. The second var iety, var. lace rata, is much more narrowly distributed in on ly peninsul ar Florida, and
disjunct in Cuba (Isla de la Juventud) (Wunderlin & Hansen 2005; Weakley 2012).
Previous classification attempts have lacked a comprehensive, systematic approach in documenting key
morphological differences between these two entities and have failed to account for what we believe are sig-
nificant characters, in add ition to the seemingly conspicuous sepal mor phology. Consequently, we chose to re-
evaluate the distinctiveness of the two currently recognized varieties using a broader morphometric analysis.
met hods
We examined selected specimens from across the entire distribution of the species (Fig. 2). Physical material
was examined from GH, NCU, and USF. The southernmost locality (Isla de la Juventud, Cuba) was examined
digitally (Britton 14200, NY, image #NY01511627). We also studied the original type description and illustra-
tion of Stipulicida setacea from Michaux (1803), as well as the putative isotypes from the Michaux Herbarium
at the Muséum National d’Histoire Naturelle (Michaux s.n., P, image #P00156961 and #P00156962). In addi-
tion, we examined digital images of types of the other named taxa, including S. filiformis (Na sh 14, GH, image
#GH00038011, MICH, image #MICH1111011, MIN, image #MIN1002897, NY, image #NY00353092, P, image
#P04925565 and #P04925568, US, image #US00103374 and #US00931431) and Stipulicida setacea var. lacerata
(Tracy 6828, GH, image #GH00038012, MSC, image #MSC0092943, NY, image #NY00353093, US, image
#US00103375). We selected 41 representative specimens of Stipulicida setacea (without discrimination be-
tween var. setacea and var. filiformis) and 26 specimens of var. lacerata for morphometric analysis to assess
422 Journal of the Botanical Research Institute of Texas 8(2)
previously described features (see Nash 1895; James 1957; Judd 1983) and to seek additional diagnostic fea-
tures. Specimens measured are denoted within the Appendix (below).
We examined nine continuous characters (Table 1). All characters were selected at random, with the ex-
ception of the length of the longest mucro (LOM), which necessitated a comprehensive scan of each specimen.
In addition, we made general observations regarding the degree of laceration and fimbriate nature of speci-
mens, as well as overall gestalt. All measurements were made with a TDI International, Inc. Micro Ruler.
To determine specimen groups and elucidate the most diagnostic variables, we used principal compo-
nents analysis (PCA). To avoid weighting characters, we examined pairwise correlation coefficients (CC) that
Fig. 2. Distribution of Stipulicida setacea specimens examined for this study. Records that have not been physically examined but are from reliable sources
(Wunderlin & Hansen 2008; Kartesz 2013) are included. State-level occurrence is indicated by shading.
Poindexter et al., Reevaluation of Stipulicida (Caryophyllaceae) 423
Table 1. Morphological characters measured from herbarium specimens with
abbreviations in parentheses. Those characters used in multivariate analyses
are denoted by an asterisk.
1 Length of outer sepal (LOS)
2 Length of inner sepal (LIS)*
3 Width of outer sepal (WOS)*
4 Width of inner sepal (WIS)
5 Stem width below terminal flower cluster (SWF)*
6 Stem width below lowest branching node (SWN)*
7 Length of longest mucro of the outer sepal (LOM)*
8 Difference between inner and outer sepal length (DSL)
9 Difference between inner and outer sepal width (DSW)
may indicate genetically linked characters and ex-
cluded those that exceed a standard threshold of >
0.7. The only exception to this concerned the charac-
ters LOM and WOS, wh ich exhibited a strong correla-
tion, but we believe that this correlation may be artifi-
cial, as it is very unlikely that these two traits would
be due to allomet ric scaling. Thus, we have decided to
retain them both within our analyses. For PCA, we
conducted analyses on a correlation matrix, which
consisted of data that is standardized so that each
variable had a mean of 0 and standard deviation of 1.
We used only the following five continuous charac-
ters in this analysis: LIS, LOM, SWF, SWN, and WOS.
We performed a Bartlett test for homogeneity of group variances on each variable. Variables that demon-
strated variances that were significantly heterogeneous were log-transformed before further analysis. We test-
ed each variable for deviations from normality using the Goodness-of-Fit test. Nearly all variables deviated
significantly from a normal distribution. We chose to maintain a conservative approach using the nonpara-
metric Kruskal-Wallis test and employed the Steel-Dwass method (i.e., a nonparametric version of the Tukey
Honestly Significantly Different test) to further detect the amount of differences between groups. Principal
components and nonparametric analyses were conducted using JMP® version 11 software (SAS Institute,
Cary, NC) on an Apple ® MacBook Pro™ computer.
result s
Principal components analysis yielded strong resolution with Stipulicida setacea var. lacerata and var. setacea
clearly separating within the ordination space (Fig. 3). Principal component 1 was responsible for 57.3% of the
observed variation and principal component 2 was responsible for 16.7%, for a total of 74.0% of the variance
expla ined. An exam ination of specimens likely to be assigned to Stipulicida setacea var. filiformis based on t heir
“wir y” habit and geographic distribution reve aled that they were embedded within t he var. setacea data cluster
(data not shown). Likewise, the Kruskal-Wallis test supported two distinct groups. Both taxa differed from
each other with respect to means of nearly all measured characters except LOS and LSW (Table 2).
discussion
The genus Stipulicida has recently been treated as monospecific and is currently represented by one species,
Stipulicida setacea, and two varieties (Swanson & Rabeler 2005; Weakley 2012): S. setacea va r. setacea and S.
setacea var. lacerata. The paucity of distinctive morphological characters found within the genus have perhaps
led to a general acceptance of this current taxonomic treatment and a lack of interest in further study. This re-
search sought to reevaluate the distinctiveness of these two entities through a more rigorous morphometric
approach that implemented multivariate analyses, in an attempt to address appropriate taxonomic ranks. We
selected several characters that had previously been emphasized by other authors, but also generated new
characters (e.g., outer and inner sepal length differences) that have aided in the further discrimination these
taxa. Despite considerable overlap, many diagnostic trends are apparent. Though both taxa have dimorphic
sepals, those of var. lacerata are distinctly lacerate/fimbriate and usually unequal in length (Fig. 4A– B). Con-
versely, the sepals of var. setacea are mostly entire margined and often even in length (Fig. 4C–D). Stipulicida
setacea var. lacerata exhibits longer and wider in ner sepals on average, and outer sepals that are generally w ider
and with a more pronounced mucronate apex (which is usually obsolete in var. setacea). Variety lacerata also
appears generally stouter than var. setacea in all aspects as evidenced by significantly greater average stem
widths (Table 2).
A combination or suite of characters rather clearly defines these two taxa w ithin ordination space (Fig. 3).
424 Journal of the Botanical Research Institute of Texas 8(2)
Table 2. Means ± 1 SD and ranges for morphological characters measured.
Character abbreviations correspond to those in Table 1. All measurements are
in millimeters. N equals the sample size. Within a row, means with different
superscripts differ significantly (Kruskal-Wallis, P < 0.05).
Character S. setacea S. lacerata
(N=41) (N=26)
1. LOS 1.4 ± 0.2a 1.5 ± 0.29a
(0.9 - 1.8) (1.0 - 2.0)
2. LIS 1.5 ± 0.21a 1.8 ± 0.32b
(1.0 - 1.9) (1.1 - 2.2)
3. WOS 0.4 ± 0.1a 0.8 ± 0.1b
(0.2 - 0.7) (0.6 - 1.1)
4. WIS 0.6 ± 0.1a 1.0 ± 0.2b
(0.4 - 0.9) (0.7 - 1.4)
5. SWF 0.3 ± 0.1a 0.4 ± 0.1b
(0.2 - 0.6) (0.3 - 0.5)
6. SWN 0.5 ± 0.1a 0.8 ± 0.1b
(0.3 - 0.9) (0.5 - 1.1)
7. LOM 0.0 ± 0.04a 0.3 ± 0.1b
(0.0- 0.1) (0.2 - 0.5)
8. DSL 0.1 ± 0.1a 0.3 ± 0.2b
(-0.2 - 0.5) (0.1 - 0.7)
9. DSW 0.2 ± 0.1a 0.2 ± 0.1a
(0.1 - 0.5) (0.0 - 0.4)
As a secondary measure, we selected specimens that
were either originally determined as Stipulicida seta-
cea var. filiformis (or S. filiformis) or were within the
originally described distribution and ecological pref-
erence of this taxon. We mapped these specimens
within our PCA and found them to be centrally nest-
ed within S. setacea var. setacea (data not shown),
which further supports Judd’s (1983) conclusion that
S. filiformis is not distinct from var. setacea. These
data, in conjunction with known biogeographic pat-
terns, suggests that Stipulicida setacea var. lacerata
and S. setacea v ar. setacea a re sufficiently di stinct to be
recognized as discrete entities. However, previous
authors ( James 1957; Judd 1983) have indicated th at a
handful of specimens in areas of sympatry are inter-
mediate. The one specimen (Highlands County, FL,
Hunne well 8955, GH) cited by James (1957) as a mixed
collection was included in our analyses, focusing on
the portion he described as “atypical” Stipulicida
lacerata and it was nested within the var. setacea data
cluster. Though we did not examine two of the addi-
tional specimens mentioned as intermediate by Judd
(1983), we did examine one from Volusia County, FL
(D.S. & H.B. Correll 51934, USF). This specimen ap-
peared to be a mixed collection with one branch that
was unassociated with a parent plant, but mounted on top of another such that it was not easily detectable as
separate. We measured both this stem, which based on our PCA was nested within var. lacerata, and another
full plant on the sheet separately, which was nested within va r. setacea. Since we have not exa mined other puta-
tive intermediates reported, we cannot attest to their identity.
We also recognize that some with in taxon vari ation is apparent and could le ad to perceived intermediacy.
For instance, some trends were observed that may not be readily apparent, e.g., northernmost populations of
var. setacea exhibit acute to slight retuse outer sepal apices, with progressively more retuse to strongly obcor-
date apices as one progresses further south. This pattern is also seen with northern populations possessing
inner and outer calyces that are narrow and subequal in length, while those of southern populations are wider
and often unequal, as well as shorter in comparison. Even the amount of lacerations/fimbriae of var. lacerata
can vary somewhat and an examination of an entire specimen is necessary to evaluate the plasticity in this
qualitative character state. James (1957) also remarked that the specimens from Cuba do not have as strongly
lacerate sepals as those from Florida, though we found this to be within the range of variation of this taxon.
How does this analysis translate to taxonomic decision-making? We find that (despite variation within
each) two morphologically distinctive entities (“lacerata” and “setacea s.s.”) are consistently separable and
identifiable using a suite of morphological characters. The several morphological characters are correlated
with one another, and do not randomly assort. An ordination confirms the more subjective assessment that
two morphologically distinct entities can be recognized. No true intermediates have been seen; and a few “pu-
tative intermediates” cited by previous workers prove not to be, based on careful analysis. Neither are there
situations in which the correlation of the seemingly independent characters breaks down. When mapped,
these t wo morphologically distinct entities have distinctive and different distr ibutions, with a substantial area
of overlap in peninsular and northeastern Florida. In areas of distributional overlap, intermediates are not
seen, alt hough at least one studied collection is mixed. Each of the dist ribution patterns shown by t he two enti-
Poindexter et al., Reevaluation of Stipulicida (Caryophyllaceae) 425
ties is a distr ibution pattern represented by a large number of other angiosperm (and even animal) species: the
“se. Virginia to s. Florida, west to e. Louisiana” distribution of “setacea s.s.” and the “Florida peninsula” distri-
bution (with optional disjunction into the West Indies) of “lacerata”—both commonly repeated distributions
across m any organism groups and therefore likely to show “biogeographic signal” (see Sorr ie & Weakley 20 01,
20 06).
If two taxonomic entities are warranted, what should their taxonomic rank be? Based on the analysis
presented above, we believe that these two entities represent independent evolutionary entities (by biological,
evolutionary, and/or phylogenetic criteria) at this time—species per most modern species concepts. A fre-
quently used, though usually not explicitly stated, assumption suggests that the magnitude of morphological
difference is well-correlated with taxonomic rank—a paradigm that might be characterized as “if you can tell
two entities apart with readily observable, macroscopic characters, they are species, but if the characters re-
quire magnification or are ‘fussy’, they are infrataxa (subspecies or varieties)”. Ward (2012), in a paper propos-
ing the reduction of dozens of Southeastern United States species of vascular plants species to varietal status,
makes some of these points more explicitly, decrying the proliferation of “microspecies”. The following state-
ment regarding Conradina etonia may be regarded as an example: “Details of indumentum, of leaf venation, of
size and pilosity, and of sta men pubescence, while wholly persu asive of the taxon’s genetic separatene ss, do not
rise to the level of difference to be found among related species” (Ward 2012). But modern biological and phy-
logenetic species concepts are not based on the magnitude of morphological variation but on the degree to
which two ent ities are demonstrably separate biological/evolutionary entities—not determinable by the size of
their characters but by inductive reasoning (as in the previous paragraph). The consistency and evolutionary
meaningfulness of the characters is what tells the tale, not their size; small pl ants with small characters are not
less significant evolutionary entities.
Fig. 3. Principal components analysis (PCA) of seven morphological characters (LOC, SWN, SWF, OSL, ISL, OSW, ISW) measured from Stipulicida lacerata
and S. setacea.
426 Journal of the Botanical Research Institute of Texas 8(2)
Gill (2014) has also recently, and significantly, raised the issue of “burden of proof” in modern taxonomic
decision-making. When previous taxonomic studies have proposed a range of conclusions (splitting to lump-
ing), what is the null hypothesi s and on whom does the burden of proof lie; to disprove previous lumping, or to
disprove previous spl itting? Gill (2014) argues that in bi rd studies, allopat ric “populations” have almost a lways
been shown, after det ailed genetic study to wa rrant taxonomic recognit ion at species rank, and that the burden
of proof should now be (based on this preponderant pattern) on those who favor “lumping”. Additionally, this
assumption is more conservationally helpful and parsimonious, in providing a more accurate hypothesis of
significant evolutionary entities requiring conservation attention. In much less well-studied groups (such as
vascular plants of the southeastern United States), patterns of prev ious studies are less clear, but generally also
show that recent rigorous and/or molecular-based studies show a greater general reliability of older morpho-
logical studies (reflected in the treatments of Small, 1903, 1913, and 1913) than the casual lumping seen in
many late 20th century and early 21st century floras (e.g., Radford et al. 1968; Wunderlin & Hansen 2011). As
demonstrated by Weakley (2005), eastern North American floras from the 1960s to 1990s instituted a trend of
lumping, often without additional st udies of the plant s themselves eit her in the field or in herba ria. This ha s led
to a current misimpression that many “splits” (recognized, for instance by J.K. Small in the Southeast and Fer-
nald in t he Northeast) have been disproven, when most ha ve received little or no addit ional study since t he first
half of the 20th century.
For Stipulicida, previous taxonomic studies have concluded that there are one, two, or three entities, and
at various ranks. We concede that we have not presented the ultimate and final analysis, based on an ideal
and complete (as of methodologies available in 2014) synthesis of molecular data, morphological character
analysis, and population biology studies. Such a study would require substantial effort, and with thousands
of such taxonomic problems remaining in the southeastern United States flora and few investigators and even
less funding, it is not our highest priority. We offer this analysis and set of taxonomic conclusions (resulting
in the recognition of two taxa, not one or three) as a current, yet disprovable hypothesis of the taxonomy of
Stipulicida.
We conclude that the best evidence available supports the recognition of two specifically distinct entities
within Stipulicida, and the refore here elevate var. lacerata to t he rank of specie s and provide a revi sed identifica-
tion key to members of the genus.
Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, & Weakley, comb. et stat. nov. Stipulicida seta cea var.
lace rata C.W. James; Rho dora 59:98. 1957. type: UNITED STATES. florida. Pinellas Co.: Dunedin, 14 Apr 190 0, Tracy 6828 (holo-
type: GH!, image; isotypes: MSC!, ima ge, NY!, image, US!, ima ge).
key to s peci es of st ipu lici da
1. Sepal margins lacerate-fimbriate; outer sepals often much shor ter than inner sepals (usually by 0.1–0.5 mm); tips of the
outer sepals with longest mucro (0.1–)0.2–0.4(–0.5) mm; [of FL and Cuba (Isla de la Juventud)] ______________________S. lacerata
1. Sepal margins entire or frayed (not lacerate-fimbriate); outer sepals subequal to slightly shorter or longer than inner
sepals (usually by 0.0–0.2 mm); tips of the outer sepals acute to obtuse or retuse-obcordate; with longest mucro or
apicule 0.0–0.1 mm; [of se. VA south to s. FL, west to e. LA] ____________________________________________________ S. setacea
Additional studies of the interesting, and seemingly isolated, genus Stipulicida are warranted. Population ge-
netic and molecular studies could elucidate additional patterns of variation, and more definitively clarify the
status of the alleged “filiformis” ent ity and the widely disjunct p opulation of S . lace rata in the Isla de la Juventud
(Cuba), as well as providing additional confirmation of our proposed species-level distinction of S. setacea and
S. la cerata. Additionally, the failure to include Stipulicida in recent molecular studies of the family leaves its
phylogenetic affinities and exact taxonomic placement within the Caryophyllaceae uncertain and conjectural.
Poindexter et al., Reevaluation of Stipulicida (Caryophyllaceae) 427
Fig. 4. Mature capsules, calyces and calyx lobes (inner lobes = left, outer lobes = right) of A–B) Stipulicida lacerata (Sarasota County, Florida, Perkins
s.n., NCU) and C–D) S. setacea (Bamberg County, South Carolina, Ahles 25982, NCU). Scale bar = 2 mm.
428 Journal of the Botanical Research Institute of Texas 8(2)
appendi x
List of representative specimens. Those utilized for morphometric analysis are denoted by an asterisk.
Stipulicida lacerata (C.W. James) D.B. Poind., K.E. Bennett, &
Weakley—U.S.A. FLORIDA. Charlotte Co.: NE of Pirate Harbor,
10 May 1990, Orzell 13438 (USF); Prairie/Shell Creek, 11 May 2009,
Franck 1250 (USF*). Collier Co.: Immokalee city limits, 7 Apr 1968,
Lakela 31361 (USF*); vicinity of Lake Trafford, 31 Jul 1968, Lakela
31560 (USF*). De Soto Co.: W side of Peace River, 31 Jul 2008, Franck
771 (USF*). Flagler Co.: NE of US 1, 1 May 2006, Slaughter 14843
(USF*). Glades Co.: Fisheating Creek Wildlife Management Area,
30 May 2010, Franck 2184 (USF*). Hardee Co.: AMAX property, 26
Mar 1979, Arcuri 856 (USF); Goose Pond Rd., 9 Apr 1981, Robinson
129 (USF*). Highlands Co.: S of Old Venus, 1 May 1958, Cooley 6137
(USF*); Archibold Biological Station, 24 Dec 1960, Craighead s.n.
(USF). Hillsborough Co.: Tarpon Springs, 23 Mar 1923, Churchill
s.n. (GH*); Little Manatee River, 21 Feb 1974, Shuey s.n. (USF); Little
Manatee River State Park, 11 Apr 1999, Myers 342 (USF*). Lee Co.:
Myers, 1900, Hitchcock 5 (GH*); Fort Meyers, May 1928, Small s.n.
(NCU*); Koreshan State Park, US 41, 20 Apr 1964, Lakela 26983
(USF); Bonita Springs, 13 Feb 1965, Foster s.n. (USF); Koreshan State
Park, 1 mi S of Estero, 1 Mar 1969, McCart 10,630 (NCU*). Manatee
Co.: Wingate Creek State Park, 14 Mar 1991, Weber WC0046 (USF);
Lake Manatee State Recreation Area, 14 Apr 1992, Weber LM0039
(USF*). Martin Co.: Jonathan Dickinson State Park, 9 Feb 1969,
McCart 10,522 (NCU*); Hobe Mountain, Sep 1987, Woodbury 87
(USF*). Okeechobee Co.: Kissimmee Prairie, 27 Apr 1923, Small
10898 (GH*). Osceola Co.: Kempfer Ranch, 9 Mar 1976, Huck 1728
(NCU*). Pasco Co.: SE of Lacoochee, 29 Jun 1992, Orzell 19761 (USF);
Starkey Wilderness Park, 20 Mar 2004, Ferguson 659 (USF). Pinellas
Co.: NE of Clearwater, 3 Apr 1970, Lelong 5344 (NCU*); Highway
19, 3 Apr 1970, Rogers 2877-B (NCU*); Dunedin Hammock, 1 May
1977, Genelle 2728 (USF*); Magnolia Creek, 24 Mar 1988, Fleming
3981 (USF*). Sarasota Co.: Sarasota, 3 Jan 1943, Perkins s.n. (NCU*);
Oscar Scherer State Park, 27 Aug 1994, Cole OSO257 (USF); Deep
Prairie Creek Preserve, 23 Aug 2007, Franck 332 (USF). St. Johns Co.:
Hastings, 11 Jun 1975, Godfrey 74329 (NCU*). Volusia Co.: Route
415, 26 May 1981, Correll 51934-B (USF*); Tomoka State Park, 15 Mar
2006, Kunzer 1454 (USF*).
Stipulicida setacea Michx.—U.S.A. ALABAMA. Baldwin Co.: Point
Clear, 20 Mar 1883, Mohr s.n. (NCU); Orange Beach, 4 Jul 2012,
Spaulding 13613 (NCU). Henry Co.: NE of Abbeville, 8 May 1989,
Orzell 9538 (NCU). Mobile Co.: Little Dauphin Island, 15 Jul 1965,
Bray 442 (NCU); S of Audubon Bird Sanctuary, 15 Aug 1966, Bray
D995 (NCU); Dauphin Island, 15 May 1972, Lelong 6502 (NCU*).—
FLORIDA. Broward Co.: Deerfield Beach, 17 Jun 1978, Correll 49877
(NCU*); Franklin Co.: near Appalachicola, s.d., Curtiss exsiccata no.
336 (NCU*); S of US Route 98, 22 May 1971, Godfrey 70233 (NCU*).
Gadsden Co.: near Lake Talquin, 29 Apr 1977, Anderson 4430
(NCU*). Highlands Co.: Sebring, 7 Mar 1924, Hunnewell 8955 (GH*);
S of Childs, 25 Apr 1960, Ray 9788 (NCU*); S of Sebring, 25 Apr 1960,
Ray 9731 (NCU*). Hillsborough Co.: W of Plant City, 8 Mar 1963,
Wall 30 (NCU). Indian River Co.: near Sebastian, 3 Apr 1921, Small
9864 (NCU*). Lake Co.: Howey-in-the-Hills, 29 April 1960, Cooley
7352 (NCU*). Martin Co.: Hobe Sound, 22 May 1960, Atwater M-214
(NCU*). Palm Beach Co.: W of Lantana, 5 Aug 1952, Howard 12959
(NCU*); West Palm Beach, 14 Feb 1969, Cassen 461 (NCU*). Seminole
Co.: S of Wagner, 3 May 1960, Cooley 7612 (NCU*). St. Lucie Co.: E
of US 1, 28 Jul 1962, Lakela 25219 (NCU). Volusia Co.: Route 415, 26
May 1981, Correll 51934-A (USF*). Wakulla Co.: Mashes Island, 24
Apr 1955, Godfrey 53207 (NCU*).—GEORGIA. Ben Hill Co.: NNE of
Fitzgerald, 4 May 1968, Faircloth 5189 (NCU*). Candler Co.: 32°16'N,
81°57'W, 3 May 1965, Cook 167 (NCU*); E of the Canoochee River,
13 May 1975, Walker 134 (NCU). Echols Co.: Mayday, 15 May 1969,
Faircloth 5831 (NCU*). Emanuel Co.: N of the Ohoopee River, 18
May 1976, Boufford 18438 (NCU*). Evans Co.: NW of Claxton, 9
May 1977, Oliver 96 (NCU). Hancock Co.: NE of Sparta, 3 Jun 1972,
Williams s.n. (NCU*). Laurens Co.: E of Dublin, 26 Jul 1967, Logue
2113 (NCU). Long Co.: near Beard’s Bluff, 10 Apr 1963, Bozeman 2543
(NCU). Marion Co.: S of Geneva, 17 May 1974, Faircloth 7770 (NCU).
McDuffie Co.: S of Thomson, 29 Jul 1950, Duncan 11558 (NCU); N of
Thomson, 18 May 1952, Duncan 13568 (NCU). McIntosh Co.: NW of
Cox, 4 Jun 1962, Bozeman 710 (NCU). Richmond Co.: near August
Airport, 17 Jun 1950, Radford 5358 (NCU*); E of Briar Creek, 16 May
1958, Duke 614 (NCU). Tattnall Co.: NW of Reidsville, 11 Jun 1961,
Ahles 54183 (NCU*). Taylor Co.: W of Butler, 16 May 1974, Faircloth
7559 (NCU). Warren Co.: NE of Sparta, 3 Jun 1972, Williams s.n.
(NCU).—LOUISIANA. St. Tammany Co.: E of Slidell, 17 May 1983,
Thomas 83658 (NCU*). —MISSISSIPPI. Forrest Co.: Hattiesburg, 29
Jun 1971, Rogers 6585 (NCU*).—NORTH CAROLINA. Anson Co.: SE
of Lilesville, 30 Jun 1956, Radford 13452 (NCU). Bladen Co.: Sandy
places, 10 Jun 1898, Ashe s.n. (NCU); W of White Oak, 22 Jun 1957,
Ahles 29167 (NCU); S of Cumberland County line, 12 May 1971,
Leonard 4784 (NCU). Brunswick Co.: Boiling Springs Lakes Preserve,
6 Jun 2006, Morris 052605-04 (NCU). Carteret Co.: E of Bogue, 21
Jun 1947, Wood 7014 (NCU); N of NC 24, 15 May 1976, Snyder 814
(NCU); Patsy Pond, 22 May 1976, Wilson 1782 (NCU). Craven Co.: W
of Vanceboro, 19 Jul 1958, Radford 37647 (NCU). Cumberland Co.:
S of Fayetteville on NC 53, 11 May 1941, Radford 1063 (NCU); N of
Cedar Creek, 2 Aug 1954, Munson s.n. (NCU); SE of Spring Lake, 4
May 1957, Ahles 24290 (NCU); WSW of Cumberland, 27 Jun 1957,
Ahles 29729 (NCU). Duplin Co.: NE of Magnolia, 27 Apr 1957, Ahles
24047 (NCU*); N of Scotts Store, 15 Jun 1957, Ahles 28407 (NCU*).
Gates Co.: White Oak Pocosin, 30 May 1958, Duke 805 (NCU*);
near Chowan River, 10 Jun 1989, Musselman s.n. (NCU*). Harnett
Co.: SE of Spout Springs, 7 May 1946, Downs 13446 (NCU*); N of
Lillington, 11 May 1946, Radford 3047 (NCU*). Hoke Co.: SW of
Bowmore, 12 May 1957, Ahles 25089 (NCU); Ashley Heights, 26
Jun 1957, Ahles 29353 (NCU). Lee Co.: N of Moore County line,
24 May 1940, Radford 137 (NCU); SE of St. Andrews Church, 7 Jun
1958, Stewart 469 (NCU). Lenoir Co.: W of Deep Run, 22 Jun 1957,
Radford 25701 (NCU). Moore Co.: Weymouth Woods State Park, 13
Jun 1965, Ahles 59622 (NCU); Weymouth Woods-Sandhills Nature
Preserve, 12 May 1973, Carter 556 (NCU); N of Southern Pines, 11
Jun 1978, Reed 106667 (NCU). New Hanover Co.: Wilmington,
May 1867, Canby s.n. (NCU*); NW of Carolina Beach, 12 Jun 1958,
Bell 12691 (NCU); UNC-Wilmington, 2 Jun 1975, Sieren 1330 (NCU).
Pender Co.: SE of Montague, 26 Jul 1953, Radford 7400 (NCU); NE
of Hampstead, 13 Jun 1957, Ahles 28003 (NCU). Richmond Co.: S
of Moore County Line, 4 Jun 1958, Duke 976 (NCU). Robeson Co.:
S of Red Springs, 21 Jun 1957, Ahles 28922 (NCU). Sampson Co.:
SE of Highsmith, 5 May 1957, Ahles 24643 (NCU); W of Clement, 28
Jun 1957, Ahles 30058 (NCU). Scotland Co.: W of Laurinburg, 8 May
1957, Ahles 24817 (NCU); S of Hoke - Scotland County line, 20 Jun
1957, Ahles 28580 (NCU); NW of US 15-501, 18 May 1973, Sharp 1592
(NCU); Green Pond, 2 Nov 1984, Berg 1198 (NCU); Scotland Road
Annual Burn Site, 2 Jun 2007, McCormick s.n. (NCU). Wayne Co.: W
of Seven Springs, 21 Jun 1957, Radford 25480 (NCU). Wilson Co.: NW
of Sims, 28 Jul 1958, Radford 38003 (NCU).—SOUTH CAROLINA.
Darlington Co.: Cannon’s, 8 Jul 1909, Coker s.n. (NCU); Hartsville,
15 May 1910, Coker s.n. (NCU); W of Black Creek, 12 Jul 1920, Norton
s.n. (NCU); W of Hartsville, 22 Apr 1921, Norton s.n. (NCU); Hartsvlle,
10 May 1932, Smith s.n. (NCU); near Black Creek along Highway 34,
23 May 1940, Smith 1664 (NCU); Kalmia Gardens., 27 Sep 1940, Smith
Poindexter et al., Reevaluation of Stipulicida (Caryophyllaceae) 429
1668 (NCU); Witherspoon Island, 3 Jun 1941, Smith 612 (NCU); E side
of Lynches River above Clyde, 18 Jul 1941, Smith 777 (NCU); Coker
College Arboretum, 26 Apr 1944, Matthews s.n. (NCU). Dillon Co.:
NNE of Oak Grove, 18 Apr 1957, Ahles 23281 (NCU); NW of Dillon,
12 Jun 1957, Ahles 27802 (NCU); Edgefield Co.: S of Trenton, 4 Jul
1957, Radford 26445 (NCU). Fairfield Co.: W of Lugoff, 24 Aug 1961,
Radford 44256 (NCU). Florence Co.: NE of Lynches River, 30 Jun
1958, Bell 13449 (NCU). Horry Co.: Myrtle Beach, 23 Jun 1931, Coker
s.n. (NCU). Jasper Co.: Turtle Island, 14 Jun 1997, McMillan 2492
(NCU). Kershaw Co.: N of Bethune, 3 Aug 1958, Duke 1846 (NCU).
Lancaster Co.: SW of Heath Springs, 6 Jun 1957, Ahles 27403 (NCU).
Lee Co.: NE of Bishopville, 6 Jun 1957, Radford 24295 (NCU); NE of
Lucknow, 26 Jul 1957, Radford 17340 (NCU). Lexington Co.: SE of
Lexington, 27 May 1957, Radford 23315 (NCU). Marlboro Co.: Little
Pee Dee River, 10 Jun 1956, Radford 12493 (NCU). Orangeburg Co.:
W of Orangeburg, 18 May 1957, Ahles 25242 (NCU). Richland Co.: E
of Ft. Jackson entrance, 13 May 1958, Duke 529 (NCU). Saluda Co.: S
of Ridge Spring, 26 May 1957, Radford 23128 (NCU). Sumter Co.: N
of Pinewood, 5 Jun 1957, Radford 24011 (NCU). Williamsburg Co.:
N of Mouzon, 12 Jun 1957, Radford 24825 (NCU).—VIRGINIA. Isle
of Wight Co.: S of Lees Mill, 29 May 1966, Svenson 13932 (NCU*).
acknowled gme nts
We would like to thank the curators at GH, MICH, MIN, MSC, NY, P, US, and USF for providing specimen
loans or digital images, and Carol Ann McCormick and Shanna Oberreiter (NCU) for assistance with
databasing and curation of these collections. We are grateful to the Muséum National d’Histoire Naturelle
(MNHN) – Pari s Herbarium (P) for giving u s permission to publish the d igital image of a n isotype of Stipulicida
setacea from the Mich aux Herbarium. We extend our appreci ation to the UNC Chapel H ill Biology Depar tment
and UNC Herbarium (NCU) for providing resources to conduct this study. The quality of this manuscript was
improved by the reviews of Richard Rabeler and Daniel Spaulding. This project was partially funded by the
Charles T. Mohr Internship Fund of the UNC Herbarium / North Carolina Botanical Garden.
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