Skip to main content

Full text of "Phytoneuron"

See other formats




Bruce A. Sorrie 

North Carolina Natural Heritage Program and 
University of North Carolina Herbarium 

North Carolina Botanical Garden 

Chapel Hill, North Carolina 27599-3280 


Shrub species of Hypericum with needle-like leaves continue to present identification 
challenges, despite stable taxonomy since the 1960s. Here I provide an improved key, identification 
notes, habitats, and range maps for nine species of the North American coastal plain. 

KEY WORDS: Hypericum. Hypericaceae, identification, distribution, habitat. 

The "needle-leaved" Hypericum shrub species have been problematic for 
Many of the species treated in this paper were not recognized or had been synonymized under H. 
fasciculatum Lam. or H. galioides Lam. until W.P. Adams" groundbreaking publication (Adams 
1962). Prior to then, there was abundant misinformation regarding species limits, morphology, 
habitat, and distribution. Adams borrowed specimens from 36 herbaria, so there are a large number 
of correctly identified specimens that serve as a reference base. Nonetheless, botanists continue to 
have difficulty with the group, in part due to the large percent of incorrectly determined specimens in 
herbaria. Adams' summary of southeastern USA Hypericum (1973) and Robson's treatment of 
section Myriandra (1996) are derivative from the 1962 work and incorporate updated information. 
Godfrey and Wooten (1981) and Godfrey (1988) made significant improvements to species 
descriptions and habitat statements; Godfrey (1988) illustrated eight of the nine species. The most 
recent taxonomic treatment is that of Crockett (2000), which recognizes the same species as Adams 


The current study w r as conducted over a period of twenty five years while gathering specimen 
information on endemic species of the North American coastal plain. Specimens annotated by W.P. 
Adams served as a baseline for my research, augmented by several iterations of newly constructed 
keys and my own annotations. I examined specimens at DUKE, FLAS, FSU, GA, GH, MISSA, 
NCSC, NCU, TEX-LL, USCH, USF, and VDB. Leaf measurements were made of the longest leaves 
on individual plants: normally these occur at branch nodes. Only mature plants were used in this 
study; sprouts and seedlings produce elliptical, blunt-tipped leaves strikingly divergent from adult 

Leaf morphology 

The linear leaves of this group of species are needle-like; that is, they are flattened in cross- 
section similar to a conifer needle, with the adaxial surface plane. The leaf margins are parallel until 
the abruptly pointed tip. These species have margins that are termed "revolute" by authors; however, 
they are revolute to an extreme degree, unlike the normal meaning of the term as "rolled under" or 
"curved towards the midrib." In this group, margins are abruptly bent under and form an angle with 
the adaxial surface such that the margins nearly or actually touch the midrib (see illustrations in 

: Needle-leaved Hyperi 

Godfrey 1988). Normally, only the midrib and the margins (each with several round punctations) are 
visible on the abaxial side, with none of the true abaxial leaf surface visible. Moreover', the margins 
are fused to the abaxial surface, making it virtually impossible to "unroll" the margins. In H. 
fasciculatum and H. tenuifolium Pursh (and to a lesser extent in other species) the midrib is raised 
such that the narrow space between the leaf margins and the midrib forms two elongate grooves. 
However, these grooves are often very shallow — a situation which has caused some confusion when 
botanists try to understand what authors mean by "groove" when there appears to be none. In other 
species, such as H. nitidum Lam., H. brachyphyllum (Spach) Steudel, and H. exile P. Adams, the 
midrib is slightly raised or not at all, such that the two-grooved aspect is lost; there appears to be a 
single broad groove or trough with the midrib at the bottom. Note: on any given specimen there may 
be a few leaves that show a narrow strip of abaxial surface on either side of the midrib; it is not 
known whether this is caused by pressing/drying or whether it occurs naturally in life. When 
attempting to identify specimens of needle-leaved Hypericum, it is imperative to examine a number of 
leaves before deciding on morphological characters. 

One apparently unrelated species, Hypericum gaiioides, is included in this paper because its 
normally flat (or slightly revolute) leaves often become strongly revolute in pressing, such that they 
take on the appearance of needle-leaved species. Slender-leaved populations of the widespread H. 
gaiioides have been the cause of many mis identifications and misattributions of range within the 
needle-leaved complex (e.g., Brown & Gandhi 1989). However, in every slender-leaved //. gaiioides 
specimen examined, many leaves clearly showed the abaxial surface. In addition, leaves of H 
gaiioides are oblanceolate or oblinear, contrary to the parallel sides of true members of the needle- 
leaved complex. Finally, the adaxial surface of the leaves of H gaiioides is convex, whereas it is 
plane in H brachyphyllum, H. fasciculatum, and other potentially confounding species. 

Plant height 

Needle-leaved hypericums conveniently fail into two groups: short species normally less than 
0.5 meters tall (//. lloydii (Svenson) P. Adams, H. tenuifolium) and tall species normally 1-2+ meters 
tall (H chapmanii P. Adams, H. fasciculatum, H. gaiioides, H. lissophloeus P. Adams, H. nitidum). 
Two species are intermediate in height: H. brachyphyllum ranges from 0.5 to 1.0 m tall, with 
occasional individuals to 1.5 m (fide Adams 1962 and Godfrey & Wooten 1981); H. exile ranges 
from about 0.4 to 1 m tall, or a bit more. 

Key to the species 

In the key I offer more than one character in each couplet, in part because the variability of 
each species is such that a single character may not distinguish one species from another and in part 

to facilitate identification whether vegetative, fruiting, or f!ow ? ering. In the key "nodes" refers to 
flowers in axillary nodes as well as the terminal cyme. 

1. Longest leaves 5-16 mm. 

2. Adaxial leaf surface convex, merging gradually with revolute margin; leaf shape oblanceolate or 
linear-oblanceolate Hypericum gaiioides 

2. Adaxial leaf surface plane, abruptly angled to revolute portion; leaf shape linear. 

3. Capsules 6-9 mm long; longest leaves 5-10(-ll) mm; corollas 13-15 mm diameter; plants 

of spodosol flatwoods and interdunes Hypericum tenuifolium 

3. Capsules 3^1.5 mm long; longest leaves 7-16 mm; corollas 13-17 mm diameter; plants of 
alfisols and ultisols of wet pine savannas, flatwoods, seq>age bogs Hypericum brachyphyllum 

Some: Needle-leaved Hypericum in the se USA 3 

1. Longest leaves 13-30 mm. 

4. Plant a shrublet or low shrub <4 dm tall, more or less decumbent, forming dense patches; 
inflorescence elongate (flowers at up to 5 nodes); corollas 10-12 mm diameter; dry to mesic soils 
of lower piedmont and inner coastal plain of se VA-NC-SC-GA-c AT,; disjunct to rock outcrops of 

s GA Hypericum lloydii 

4. Plant an erect shrub 0.5-4 m tall, with single main stem branched above; inflorescence elongate 
(3-7 nodes) or short (1-3 nodes in H. fasciculatum and H. chapmanii): corollas 13-26 mm 
diameter; wet soils of coastal plain. 

5. Abaxial surface of most leaves easily seen on both sides of midrib, veins usually obvious on 
undersurface, leaves oblanceolate to oblinear, (1.0— )1.5— 5(— 7) mm wide; inflorescence elongate 

(3-7 nodes) Hypericum galioides 

5. Abaxial surface usually not seen except for midrib (leaf margins nearly touch midrib along its 
whole length), if undersurface visible then no veins visible, leaves linear, needle-like, parallel- 
sided, 0.5-1.5 mm wide; inflorescence elongate or short. 

6. Plant short, <1 m tall; stem <1 cm wide at base; plant unbranched or few-branched, wand- 
like with narrow crown; restricted to FL panhandle Hypericum exile 

6. Plant tall, normally >0.8 m; stem 1-several cm wide at base; crown broader with many 
ascending to spreading branches. 

7. Young branches, leaves, and sepals strongly glaucous; bark of upper stem and branches 
silvery gray and smooth; mature plant 2-4 m tall with ascending branches imparting tree-like 
or vase- like aspect; restricted to shores of sinkhole ponds in Bay and Washington cos., FL 
Hypericum lissophloeus 

7, Young branches, leaves, and sqials not glaucous; bark of upper stem and branches not 
silvery gray and smooth (except some H. chapmanii); mature plants various!}' shaped. 

8. Inflorescence elongate (3-7 nodes); stem bark tight, thin, not exfoliating or exfoliating in 
narrow strips, not revealing buff or pale cinnamon color; if leaf undersurface is exposed it 
is distinctly paler man upper surface; usually associated with flowing water (blackwater 
streams and impoundments) Hypericum nitidum 

8. Inflorescence short (1-3 nodes). Stem bark corky-thickened to spongy, exfoliating in 
broad strips or sheets revealing buff or pale cinnamon color; if leaf undersurface is exposed 
it is about the same color as upper surface; usually associated with static water (Carolina 
bays, impoundments, beaver ponds, borrow pits, flatwoods depressions, cypress-gum 

9. Mature plant 2-3 (^\) m tall; branches ascending and imparting a tree-like or vase-like 
aspect (younger plants may be bushy); youngest intemodes terete; restricted to flatwoods 

depressions and cypress-gum ponds of FL panhandle Hypericum chapmanii 

9. Mature plant 0.8-1.5 m tall; branches spreading and imparting a bushy or gumdrop 
aspect; youngest internodes with distinct winged ridge on either side; Carolina bays, 
impoundments, beaver ponds, borrow pits, widespread Hypericum fasciculatum 

1. Hypericum brachyphyllum (Spach) Steud. is intermediate in stature, but its leaves average 
shorter than all species excqjt H. tenuifolium. North Carolina plants are shorter in height than 
elsewhere, normally less than 5 dm. Flowers are produced at 3-5 nodes. Unusually tall individuals 

Some: Needle-leaved Hypericum in the se USA 4 

may be told from H. fasciculatwn and H. nitidum by characters in the key, plus its later flowering 

period (July -August vs. late May-early July). 

Recently Hypericum brachyphyllum has been reported from western Louisiana (Robson 
1996: Vernon Par.: Anacoco, Demaree 50849 (BM). There are specimens at NCU, USF, TEX-LL, 
and VDB that appear to be this species, collected from more than a dozen sites in eastern Texas and 
western Louisiana. However, all of these specimens prove to be H. galioides, including a duplicate of 
Demaree 50849 at NCU. These specimens have unusually slender leaves, but the abaxial side of 
some to many leaves show exposed leaf surface rather than merely a groove on either side of the 
midrib. See H. galioides text for additional identification criteria. H. bi cl v, ' ylh m inh ibifs id 
pine flatwoods and pitcher plant seepage bogs. Range map 1. 

Adams (1962) thought mat Hypericum limosum Griseb. of western Cuba might be 
synonymous with H. brachyphyllum, but opted to wait for better specimens; Robson (1996) treated H 
limosum as a good species. I have not seen enough specimens to make an informed decision. 

2. Hypericum chapmanii P. Adams and H lissophloeus are by far the tallest of the group, reaching 
3-4 meters. Both usually look like small slender scruffy trees, although some H chapmanii can be as 
short as H fasciculatum and have a bushy-branched aspect. Hypericum chapmanii and H. 
fasciculatwn have flowers in a. terminal cyme plus l-2(-3) axillary nodes, unlike the rest of the 
group, which have flowers in 3-7 nodes. From H. fasciculatum. H. chapmanii can be distinguished 
by the greater development of thick corky bark, which has large, vertical, pale lactifers that stand out 
in contrast to the bark color. From H. lissophloeus, H. chapmanii can be distinguished by green, non- 
glaucous leaves (but beware that some H. lissophloeus leaves are merely glaucescent), smaller 
flowers (15-16 mm diameter vs. 20-22 mm), and lack of metallic silver-gray color of upper stems 
and branches (occasional plants of H. chapmanii show some of this color). 

It inhabit- cypress-gum ponds, small lakes, natural depressions, and borrow pits in eleven 
counties in the Florida panhandle. Range map 2. 

3. Hypericum exile P. Adams has an odd looking gestalt: a wand-like aspect with a few short 
branches, or unbranched. The sepals and capsules are long (6-7 mm and about 7 mm, respectively, 
according to Godfrey and Wooten 1981) in contrast with the much shorter sepals and capsules of H. 
fasciculatum and H. nitidum. The leaf midrib is pale green or pale greenish tan, unlike the darker 
color in H. fasciculatum and H. nitidum. This pale color hardly contrasts with the color of the abaxial 

Hypericum exile is restricted to five counties in the Florida panhandle, where it inhabits 
periodically wet flatwoods and savannas. Range map 3. 

Robson (1996) treated Hypericum exile as H. nitidum subsp. exile (P. Adams) N. Robson. He 
also attributes this taxon to western Cuba, citing several specimens. He states that it is more variable 
in Cuba than in Florida, but gives no data to support his decision to consider Cuban material identical 
with Florida material. Adams (1962) considered all plants of the H fasciculatum complex occurring 
in Cuba to be "closest to H. nitidum" and places them in H. nitidum without additional comment. 
Adams (1973) repeated this course of action. 1 have not examined Cuban specimens. 

4. Hypericum fasciculatum Lam., H. nitidum, and H. galioides have the same gestalt: a single main 
trunk with many branches forming a roughly rounded crown, long leaves, and wetland habitats. The 
trunk and oldest branches of H. fasciculatum have thickened bark with a spongy or corky texture: the 
outer layers peel off in thin sheets to reveal a pale cinnamon, or pinkish-tan color, in marked contrast 

Some: Needle-leaved Hypericum in the se USA 5 

to the bark of H. nitidwn and H. galioides, which is thin and dark and which peels off in small narrow 
strips or not at all. Although all species of this group of Hypericum have fascicled leaves, in. H. 
fasciculatum they are generally more numerous and densely packed than in other species; this 
comparative character can he used with caution in separating vegetative H. fasciculatum from H. 
nitidum. Another gesta It character is the more or less cut-off leaves of H. fasciculatum, as if someone 
clipped the fascicles with scissors. The clipped fascicles often give a "neat" appearance to the 
branches, and expose a good bit of the twig between nodes. Some specimens show this character 
well; others do not, so caution is advised. Populations in the Sandhills of the Carolinas have shorter 
leaves on average than elsewhere; otherwise I find no differences among populations rangewide. A 
crucial character of//, fasciculatum is the short inflorescence with flowers at only 1-3 nodes, in 
marked contrast to all other needle-leaved species except H. chapmanii. This inflorescence character 
becomes critical when other characters of a plant are equivocal. 

Pre- Adams determinations of specimens of this and other members of the needle-leaved 
group were often "Hypericum fasciculatum"' without recognizing the diversity actually represented. 
Thus situation contributed to significant errors in range statements and morphology. 

Hypericum fasciculatum inhabits static wetlands, such as Carolina bays, cypress-gum ponds, 
natural lakes, impoundments, depressions in flatwoods, borrow pits, and roadside ditches; this is in 
contrast with H. nitidum which see. Occasional populations occur in wet streamheads, cypress 
stringers, and apparently along blackwater streams (Godfrey & Wooten 1981). Records from western 
Louisiana and eastern Texas are all mis identifications involving H. galioides or H. br achy phy Hum. 
Range map 4. 

5. Hypericum galioides Lam., although not a true member of the needle-leaved group, often 
produces narrow-leaved forms; in fact, populations west of the Mississippi River almost always 
produce slender leaves. Leaves of these plants become strongly revolute in drying and many 
specimens were originally determined as H. fasciculatum or H. nitidum. However, H. galioides can 
be told by these features: leaves oblanceolate or oblinear (vs. parallel sides), most or all leaves (vs. 
few or none) showing exposed abaxial surface, abaxial surface usually with obvious venation (vs. 
none), and elongate inflorescence with flowers at 3-7 nodes (vs. 1-3 in H. fasciculatum). Slender- 
leaved plants of H. galioides are vexingly similar to H. brachyphyllum, which has similar-sized 
leaves, corollas, and fruits. From H. brachyphyllum, H. galioides differs in its linear-oblanceolate 
leaf shape (slightly broadened distally, vs. linear and with completely parallel sides), in its convex 
adaxial leaf surface (vs. plane), and in in the exposure of abaxial leaf surface in at least several leaves 
(vs. few or none). Due to its variable size, wide distribution, and broad range of habitats, familiarity 
with/7, galioides is fundamental to understanding the true needle-leaved species. 

Habitats include ponded depressions, cypress-gum ponds, beaver ponds, impoundments, 
floodplain swamps, wet savannas, flatwoods, and ditches. Range map 5. 

6. Hypericum lissophloeus P. Adams is not likely to be confused with any other species, due to its 
very tall stature (2-A meters), narrow crown, and tree-like aspect. Hypericum chapmanii occasionally 
produces similar-looking plants, but H lissophloeus differs in its pronounced metallic silver-gray 
color to upper trunk and limbs, glaucous or glaucescent leaves, and large corollas (at 20-22 mm the 
largest in the group). 

Hypericum lissophloeus inhabits sinkhole ponds in deep sand deposits of Bay and 
Washington Counties, Florida. Range map 6. 

Some: Needle-leaved Hypericum in the se USA 6 

7. Hypericum lloydii (Svenson) W.P. Adams and H. tenuifolium are the only species in the needle- 
leaved group that normally inhabit dry soils. Moreover, they are the smallest in stature, never 
exceeding 0.5 m tall. Hypericum lloydii is a compact, bushy-branched shrublet. Among the short 
species of the group (H. brachyphyllum, H. lloydii, H. tenuifolium) H. lloydii has the smallest corollas 
(10-12 mm diameter) and longest leaves (13-25 mm). 

Hypericum lloydii inhabits diy to mesic roadsides, powerlines, semi-shaded rocky or sandy 
slopes, and openings in oak-hickory-pine woodlands. Its distribution is unique within the needle- 
leaved group in that it occupies a narrow band of the lower piedmont and inner coastal plain 
(Sandhills region). Disjunct populations occur on outcrops of Altamaha Grit sandstone in Turner and 
Coffee counties in southern Georgia. While numerous in the Carolinas, H lloydii is apparently rare 
elesewhere. Range map 7. 

8. Hypericum nitidum Lam. resembles H fasciculatum, with which it has been much confounded, 
but differs in a number of features. The leaf fascicles never look clipped like those of//, fasciculatum 
and are more widely distributed and expose less of the twig between nodes, thus giving H. nitidum 's 
branches an unkempt appearance. If on a few leaves the abaxial surface may be seen, the surface is 
much paler tlian the inrolled margins, in contrast to H, fasciculatum. Two excellent characters are the 
number of flowering nodes (3-7 for H. nitidum, 1-3 for H. fasciculatum) and dark, thin (not corky) 
bark that does not flake off in large sheets or strips to reveal a pale cinnamon color as in H. 

Hypericum nitidum normally inhabits the margins of blackwater rivers, streams, and 
flatwoods drainageways, in contrast with the usually static waters inhabited by H. fasciculatum. On 
the East Gulf Coastal Plain, H. nitidum is often a dominant where blackwater streams meet estuaries 
(i.e., fresh-tidal zones), in the company of Sarracenia leucophylla, Eriocaulon decangulare, 
M acr anther a flammea. and other seq^age bog plants. Locally, H. nitidum inhabits borrow pits and 
roadside ditches. 

Reports of Hypericum nitidum from Brunswick County, North Carolina (Adams 1962; 
Godfrey & Wooten 1981) are not supported by vouchers that I have seen and likely pertain to 
specimens of H brachyphyllum. All "nitidum" specimens examined from Louisiana, Mississippi, 
and from west of Mobile Bay in Alabama prove to be mis identified; thus, H. nitidum does not range 
west of the Alabama-Tens aw River estuary. I have seen one specimen from central Florida: "Salt 
Springs, Ocala NF", 3 Jun 1929, Ashe s.n. (NCU), and S. Crockett (pers. eomm.) has seen a specimen 
from adjacent Lake County. Range map 8. 

Hypericum nitidum was attributed to Cuba and Belize by Robson (1996), as H nitidum subsp. 
cubense (Turcz.) N. Robson. One of the specimens he cited is Howard 5201 (many herbaria). Two 
duplicates are at NCU. The general aspect of the branches is like nitidum— -fascicles not dense, 
leaves without the "clipped" look of//, fasciculatum, unkempt appearance, longest leaves up to 16 
mm — but other aspects contradict identity with H. nitidum (and with H. exile): (1) The abaxial leaf 
surface is barely or not paler than inrolled margins, contra H nitidum and H exile. (2) The 
inflorescence consists of 1-3 nodes, contra //. nitidum and //. exile (both with 3-7 nodes). (3) Plants 
are extremely short for //. nitidum: Howard's labels state "low woody herb seldom to 1. 1/2 feet" 
This is short even for //. exile. (4) The habitat is dry; Howard's labels state "dry open grassy 
meadow," in marked contrast with H nitidum and H exile. Basal on this admittedly small sample, I 
am of the opinion that tax on cubense does not belong with H. nitidum. 

9. Hypericum tenuifolium Pursh (= H reductum P. Adams) has the shortest leaves of any of the 
needle-leaved group. Godfrey and Wooten (1981) stated that leaves do not exceed 5 mm, but plants 

. long. A key character is the long c; 

in the Carolinas routinely have leaves 4-1 mi 
much longer than other short statured species. 

In North and South Carolina Hypericum tenuifolium inhabits moist to dry sandy flatwoods, 
sandrims of Carolina bays, and ecotones of depression ponds; southward it also inhabits maritime 
interdune swales, pine-scrub oak sandhills on inland or "fossil" dunes, and pond ecotones. 

i distribution between Pasco and Franklin 
ly this gap reflects a lack of dune and sand 

On the Gulf Coast of Florida there i; 
Counties, with the exception of Dixie County, 
ridge habitats. Range map 9. 


Adams, [W.]P. 1962. Studies in the Guttiferae. 1. A synopsis of Hypericum section Myriandra. 

Contr. Gray Herb. 189: 1-51. 
Adams, [W.]P. 1973. Clusiaceae of the southeastern United States. J. Elisha Mitchell Sci. Soc. 89: 

Brown, L.E. and K.N. Gandhi. 1989. Notes on the flora of Texas. Phytologia 67: 394-399. 
Crockett, S.L. 2000. Taxonomy and medicinal chamistry of the Hypericum fasciculatum 

alliance (Clusiaceae: Hypericum section Myriandra). Unpublished Masters thesis, 

Univ. of Georgia, Athens. 
Godfrey, R.K. 1988. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia 

and Alabama. Univ. Georgia Press. Alliens. 
Godfrey, R.K. and J.W. Wooten 1981. Aquatic and Wetland Plants of Southeastern United 

States. Univ. Georgia Press, Athens. 
Robson, N.K.B. 1996. Studies in the genus Hypericum L. (Guttiferae). 6. Sections 20. Myriandra to 

28. Elodes. Bull. Nat. Hist. Mus. London (Bot.) 26: 75-21 7. 

Figure 1. Distribution of Hypericum brachyphyllum. 

Some; Needle-leaved Hypericum in the se USA 8 

H/fCyT^- ^tv^iirYtnL^4- 



t-Tl./' '- V" '-'. ; ^\ :; '" 




rWTvJSw-H " •■■'(- 

>nM Vvrii ""'' ^^^^BP^ ~ ■ 

■■■..>.;"■.■, Brc 



Figure 2. Distribution of Hypericum chapmanii. 

Figure 3. Distribution of Hypericum exile. 

Some: Needle-leaved Hypericum in She se USA 9 

Figure 5. Distribution of Hypericum galioides. 

Some: Needle-leavedi^vper/'cumin the se USA. 10 

Figure 7. Distribution of Hypericum lloydii. 

Sorrie: Needle-leaved Hypericum in the se USA 1 1 

Figure 9. Distribution of Hypericum tenuifolium. 

: Needle-leaved Hyperi 


Selected Specimen records 

1. Hypericum brachyphyllum. Alabama. Baldwin Co.: sand dunes. Ft. Morgan peninsula. 28 Jul 
1954, Demaree 59903 (NCU); Conecuh Co.: hillside seepage bog 3^1 mi W of Castleberry, 11 Sep 
1989, Orzell & Bridges 11 731 (NCU, TEX); Geneva Co.: pond cypress depression by County Road 
4, 12 Sep 1995, Sorrie 8589 (AUA, NCU). Florida. Collier Co.: grassy prairie surrounded by pond 
cypress swamp. 20 Sep 1965, Ward 5233 (NCU); Citrus Co.: hardwood swamp, Chassahowitzka. 7 
Oct 1972, Genella & Fleming 1601 (NCU); Duval Co.: near Jacksonville, 3 Jul 1891, Sudworth 888 
(NCU); Wakulla Co.: wetland limestone savanna E of St. Marks Refuge, 14 Sep 1989, Orzell & 
Bridges 12099 (TEX). Georgia. Calhoun Co.: edge of low pine woods, 26 May 1961, Adams 793 
(USF); Pierce Co.: borrow pit 0.2 mi N of Alapaha River, 9 Jun 1967, Bozeman 9404 (NCU); Taylor 
Co.: Little Whitewater Creek by Georgia 137, 12 Aug 2002, Krai 93044 (TEX). Louisiana. St. 
Tammany Par.: roadside ditch S of Talisheek, 20 Oct 1990. Brawn 15023 (TEX); Tangipahoa Par.: 
near Hammond, 15 Jun 1929, Ashe s.n. (TEX). Mississippi. Hancock Co.: acid savanna 1 mi N of 
Necaise, 6 Jul 1967, Jones 14122 (USF); Jackson Co.: pine meadows about halfway between 
Pascagoula and Fontainebleu, 22 Aug 1962, Harper 4532 (NCU); Wayne Co.: 5 mi N of State Line, 
low wet grassy flat in open pineland, 18 Jul 1956, Ray, Jr. 7046 (USF). North Carolina Brunswick 
Co.: Camp Branch Savanna, 23 Sep 1994, LeBlond4115 (NCU); Columbus Co.: Schulkens Savanna 
on S.R. 1928, 6 Aug 1994, LeBlond 3999 (NCU); Onslow Co.: Haws Run Mitigation Site, 14 Jun 
2003, LeBlond 5 771 (NCU); Pender Co. : Shaken Creek Savanna, 9 Oct 2002, LeBlond 5 736a (NCU). 

2. Hypericum chapmanii. Florida. Franklin Co.: shallow ponds near Apalachicola, 1893, Chapman 
5735a (GH, NCU); Holmes Co.: pond margin near Bonifay, 23 May 1930, Blanton 6560 (GH); 
Liberty Co.: Taxodium ascendens stringer in wet savanna, 3 mi sw of Kern, Apalachicola National 
Forest, 1989, Orzell & Bridges 12061 (GH. NCU): Walton Co.: Grayton Beach, boggy area by stream 
running into Deer Lake, 31 Jul 1983, Godfrey 80824 (FSU). 

3. Hypericum exile. Florida. Gulf Co.: 2.5 mi SE of Port St. Joe, pine flatwoods, 18 Jun 1958, 

Adams 456 (DUKE, FSU, GH); Liberty Co.: sandy-peaty shallow ditch by forest road 123C, 
Apalachicola National Forest, 18 Aug 1989. Godfrey (GH). Cuba. Prov. Pinar del Rio, Britton & 
Crowell 9618 (F, GH, K, MO, NY). 

4. Hypericum fasciculatum. Alabama Covington Co.: Conecuh National Forest, Forest Service 
road 96, seepage bog, 12 Jul 1995, LeBlond 4319 (NCU); Geneva Co.: seepage bog in longleaf pine 
sandhills W of Geneva, 25 Jul 1968, Krai 31994 (NCU); Mobile Co.: pine barren flats, hwy 163 
between Mobile and Dauphin Island, 3 Jun 1967, belong 3210.1 (NCU). Florida Alachua Co.: 1 
mile E of hwy 24 on 232, 8 May 1965, Wiggins 2001 4 (NCU); Hernando Co. : wet prairie, section 1 1 
& 12, 19 Sep 1959, Cooley 7040 (NCU); Polk Co.: Tiger Creek Preserve, ephemeral pond edges, 28 
January 2008, Corogin TC624 (TEX); Wakulla Co.: wet thickets. January 1929, Ashe s.n. (TEX). 
Georgia. Chariton Co.: powerline right-of-way, east side of Foikston and just north of GA 40, 25 
May 1999, Nelson 20452 (GA, USCH); Mcintosh Co.: wet depression along GA 99 S of Eulonia, 20 
Oct 1987, Angerman s.n. (NCU); Worth Co.: Cypress-Ilex-Hypericum pond on Ga. 32, 15 Jun 1967, 
Bozeman 998'' (NCU). Louisiana St. Tammany Par.: wet pine savanna. 11 November 1990, 
Urbatsch 6929 (NCU); Washington Par.: wet hardwood forest, 1 Oct 1982, Taylor 2227 (USF). 
Mississippi. Forrest Co.: Shelby State Park, 7 Jun 1964, Jones, Jr. 1864 (NCU); Harrison Co.: De 
Soto National Forest, Forest Sen-ice road 426, seepage bog, 15 Aug 1996, Sorrie 9010 (NCU); 
Jackson Co.: Orange Grove, 2 May 1954, Demaree 35057 (GH). North Carolina. Cumberland Co.: 
wet sandy soil. 15 mi S of Fayetteville, 11 Feb 1940, Totten s.n. (NCU): Hoke/Moore Co.: Fort 
Bragg, boggy margin of Johnsons Mllpond, 9 Oct 1991. Sorrie 5967 (NCU). South Carolina. 
Allendale Co.: swamp 1.2 mi SSW of Barton, 30 Jun 1956, Bell 4001 (NCU); Berkeley Co.: Francis 

Some: Needle-leaved Hypericum in the se USA 13 

Marion National Forest, limesink depression SE of route 654, Jun 1997, McMillan 2636 (NCU); 
Orangeburg Co.: Branchville Bay e of US 21, 1 Sep 1994, Nifong 541 (NCU). 

5. Hypericum galioides. Alabama, Baldwin Co.: low woods bordering Tensaw River, 26 Oct 1967, 
belong 386S (NCU); Clarke Co. : low roadside ca. 4 mi N of Choctaw Bluff, 4 Oct 1966, Clark 9078 
(NCU; Washington Co.: stream margin, Bassetts Creek near hwy 43, 3 Oct 1966, Clark 871 6 (NCU). 
Florida. Columbia Co.: along Fla. 2, halfway between GA state line and Baker County line, roadside 
ditch in weedy, recently logged pine flatwoods, 11 Jul 1984, Hansen 10147 (USCH); Levy Co.: 
highwater mark at edge of floodpiain forest, Suwanee River, 22 November 1974, Godfrey 74133 
(NCU); Okaloosa Co.: wet roadside with Macranthera flammea, S side hwy 90, 1.8 mi E of 85, 20 
Aug 1971, Musselman 4375 (NCU). Georgia. Brooks Co.: margins of slough on Withlacoochee 
River, 14 Jul 1965, Faircloth 2195 (NCU); Chatham Co.: mixed woodland on hwy 21, S of St. 
Augine Creek, 14 Jul 1966, Bozeman 6149 (NCU); Tattnall Co.: cleared floodpiain of Altamaha 
River, NE of Lane's Bridge, hwy 169, 12 Jul 1966, Bozeman 5923 (NCU). Louisiana. Allen Par.: 
pine flatwood along hwy 26, 5 mi WNW of Oberlin, 25 Jul 1975, A lien 6681 (NCU); Calcasieu Par. : 
moist pine woods, 6 Jul 1950, Webster & Wilbur 321 7 (GA NCU. TEX); Sabine Co.: pine woods on 
Peason Ridge Military Reservation, 4 Oct 1980, Thomas 73842 (TEX): Tangipahoa Par.: wet 
roadside ditch along LA 1067, 3 mi SW of Robert, 12 Jun 1978, Allen 8171 (NCU). Mississippi. 
George Co.: swamp forest along Escatawpa River, hwy 612, 24 Jul 1969, Belong 5209 (NCU); 
Hancock Co.: Mississippi Test Facility (NASA), along canal, 7 Aug 1970, Rogers 4001-A (NCU); 
Lawrence Co. : 3 mi S W of Silver Creek, secondary deciduous wood bordering small stream, foot of 
pine coverered slope, 24 Jun 1957, Ray, Jr. 8283 (USF). North Carolina. Columbus Co.: low 
cypress savanna SE of Old Dock, Beonard 1754 (NCU, TEX); Hoke Co.: Redwing Pond, boggy 
shrub margin, 21 Jul 2002, Sorrie 10963 (NCU); Pender Co.: swamp forest on Black River, 3 mi W 
of Montague, 26 Jul 1953, Radford 7387 (NCU). South Carolina. Chesterfield Co.: open, savanna- 
like hillside bog adjacent to Oxpen Lake in Carolina Sandhills NWR 5 Jul 1985, Rayner 2324 
(USCH); Dorchester Co.: Givhans Ferry State Park, bottomland below marl bluffs, banks of Edisto 
River, 14 Jun 1988, Hill 19588 (USCH, USF); Jasper Co. : flatwoods just north of S-94, ca. 1/3 mile E 
of 1-95; NE of Hardeeville, 15 Jul 1984, Aulbach-Smith 3136 (USCH). Texas. Chambers Co.: 
freshwater marsh, 1.1 mi S of FM 1985, Anahuac NWR, 14 Jun 2000, Carr 19031 (TEX); Hardin 
Co.: cut-over longleaf pine, 5 Sep 1924, Tharp 3152 (TEX); Montgomery Co.: Lake Houston State 
Park, utility easement on alluvial terraces, 21 Oct 2003, Sanders 6261 (TEX); Newton Co.: junction 
of Big Cow Creek and FM 1416, 24 Sep 2000, Holmes & Singhurst 1 1059 (TEX). 

6. Hypericum lissophloeus. Florida. Bay Co.: Lake Merial, 15 mi N of Panama City, 1966, Ward 
5958 (NCU); Washington Co.: in sand along margin and in shallow water of Parish Pond, 8 Jun 1990, 

Anderson 12882 (FSU). 

7. Hypericum lloydii. Alabama. Randolph Co.: 2.8 mi SE of Rock Mills, 1.6 mi E of Bacon Level, 
large relatively undisturbed granite flatrock above Wehadkee Creek and W of quarry, 31 Aug 1985, 
Allison 2518 (GA); Tallapoosa Co.: Harper 3691 (GH, PH, US). Georgia. Coffee Co.: sandstone 
outcrop on Rocky Creek about 3 mi S of Ocmulgee, 20 Jun 1966, Bozeman 4591 (NCU); Richmond 
Co.: Auga, 29 Mav 1918. Harbison 14393 (NCU); Turner Co.: Altamaha Grit outcrops, 4 mi N of 
route 112, 25 Jun 1969, Faircloth 5855 (NCU). North Carolina. Pitt Co.: no data, 9 Jul 1956, 
Boyette s. n. (NCU): Scotland Co. : margin of sand road through moist pine flatwoods, Sandhills Game 
Land, 25 Jul 1998, Sorrie 9875 (NCU); Union Co.: roadside poweriine by Austin Road, 12 Jul 2010, 
Sorrie 12623 (NCU). South Carolina. Aiken Co.: Graniteville, 1898, Eggert s.n. (MO, NY, US); 
Chesterfield Co.: Sugar Loaf Mountain, 14 Jun 1918, Coker s.n. (NCU); Greenwood Co.: roadside 
bank on rte. 25, 5.3 mi S of Cuffytown Creek, 24 Jun 1987. Hill 18124 (NCU). Virginia. 
Mecklenburg Co.: roadside of route 58, 7 Jul 1967, Seaman 7460(NCU). 

Some: Needle-leaved Hypericum in the se USA 14 

8. Hypericum nitidum. Alabama. Baldwin Co.: Jack 3004 (GH). Florida. Gulf Co.: flatwoods 
between Panama City and Wewahitchka, 4 May 1926, Small 12823 (TEX); Marion Co.: Salt Springs, 
Ocala National Forest, 3 Jun 1929, Ashe s.n. (NCU); Santa Rosa Co.: swamp along U.S. 90, 
Blackwater River e of Milton, 22 November 1966, Bozeman 8761 (NCU). Georgia. Berrien Co.: 
creek swamp on U.S. 82, E of Fjiigma, Bozeman 9932 (NCU); Brooks Co.: road excavation area, 2.8 
mi E of Barney, 21 November 1959, Adams 376 (USF); Dooly Co.: swale in longleaf pineland, Krai 
51607 (TEX); Tattnall Co.: pocosin 1.9 mi NW of Reidsville, 11 Jun 1961, Ahles 54258 (NCU). 
South Carolina. Darlington Co.: edge of Black Creek near Darlington Country Club, 7 Jul 1940, 
Smith 1122 (USCH); Lexington Co.: Black Creek w of Pelion, Rayner 2569a (USCH); Richland: Fort 
Jackson, sand and much of small islands in Colonel's Creek, 16 Jun 1994, Nelson 15611 (USCH, 

9. Hypericum tenuifolium. Alabama. Baldwin Co.: hwy 182 near Romar Beach, 1 Jun 1977, 
Davenport 96 (USF). Florida. Lake Co. : dry sand of scrub, Ocala National Forest service road 71,2 
Aug 1962, Ward 3048 (NCU); Manatee Co.: 1845, Rugel s.n. (NCU); Martin Co.: pineland, Jonathan 
Dickinson State Park, 21 May 1977. Correll 48587 (NCU); Volusia Co. : scrub near dry pond, road to 
Benson Spring, 14 April 1953, Prichard 583 (NCU). Georgia. Bryan Co.: fossil dunes, Canoochee 
River, Adams 541 (GH, USF); Clinch Co.: low exposed area along rte. 84, 6 mi SW of Homerville, 
30 Jun 1998, McNeilus 98-476 (TEX); Irwin Co.: pine savanna on U.S. 319, 0.7 mile E of Alapaha 
River, 13 Jun 1967, Bozeman 9608 (NCU). North Carolina. Carteret Co.: recently burned longleaf 
pine stand about 13 mi W of Morehead City, 9 Jul 1963, Wilbur 6922 (DUKE, USF); Cumberland 
Co.: flat pine woods 5.7 mi N of Bladen County line on N.C. 53, 27 Jun 1953, Ahles 29873 (NCU, 
USF); New Hanover Co. : Carolina Beach, dry sterile white sand ridge among Firms palustris and 
Quercus catesbaei, Godfrey, PI. Exs. Gr. 1260 (GH, NCU, USF). South Carolina. Beaufort Co.: 
Bluffton, 1874. Mellichamp s.n. (GH); Horry Co.: savanna, Conway, 1 Sep 1940, Schallert s.n. (GH); 
Jasper Co.: savannah 2.3 mi SW of Rklgeland on U.S. 17. 26 Jun 1956, Ahles 15529 (NCU).