Key to Hexastylisin North Carolina
Key adapted from Blomquist (1957 [Brittonia]), Gaddy (1987 [Castanea]), Whittemore and Gaddy (1997 [FNA]), Weakley (2008), and Gaddy (2011 [Phytoneuron]). Photos by Krings, unless otherwise indicated. Line drawings from Britton & Brown (1913; An illustrated flora of the northern United States, Canada and the British Possessions). Maps courtesy of USDA PLANTS and the North Carolina Natural Heritage Program.
1. Style extension distinctly bifid to stigma; leaves triangular to ovate-sagittate or subhastate...H. arifolia
![Hexastylis](images/keys/Hexastylis_arifolia_complete.jpg)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings6_sml.jpg)
Leaf of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings1_sml.jpg)
Leaves of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings7_sml.jpg)
Flower of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings2_sml.jpg)
Flower of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings3_sml.jpg)
Flower of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings5_sml.jpg)
Flower detail of Hexastylis arifolia (Photo: Krings)
![Click to enlarge Hexastylis arifolia](http://herbarium.ncsu.edu/rare/images/Hexastylis_arifolia_Krings4_sml.jpg)
Flower detail of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
(A-B, var. arifolia: common, mesic to dry deciduous forests, Mt, Pd, CP;
C, var. ruthii: uncommon, ultramafic outrop barrens, upland forests, Mt)
1’. Style extension notched or divided at the apex, but not bifid to the stigma; leaves cordate, subreniform,
orbicular, or triangular-cordate...2.
2. Leaf margins minutely toothed...H. rosei
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6750_Krings_habitat_sml.jpg)
Habit of Hexastylis rosei (Photo: Krings)
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6693_Krings_lf_sml.jpg)
Leaf of Hexastylis rosei (Photo: Krings). Note minute teeth along leaf margins, as well as the impressed midvein.
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6695_Krings_fl2_sml.jpg)
Leaf of Hexastylis rosei (Photo: Krings). Note minute teeth along leaf margins, as well as the impressed midvein. In Galax urceolata (inset), the midvein is distinctly raised.
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6696_Krings_fl3_sml.jpg)
Leaf of Hexastylis rosei (Photo: Krings).
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6707_Krings_fl_sml.jpg)
Flower of Hexastylis rosei (Photo: Krings).
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6712_Krings_fls_sml.jpg)
Flowers of Hexastylis rosei (Photo: Krings).
![Click to enlarge Hexastylis rosei](http://herbarium.ncsu.edu/rare/images/Hexastylis_rosei_IMG_6703_Krings_fl4_sml.jpg)
Flower of Hexastylis rosei (Photo: Krings).
Hexastylis rosei
(rare; acidic cove forests; Mt)
2'. Leaf margins entire...3.
3. Inner surface of calyx lobes pilose with whitish hairs...H. lewisii
![Hexastylis lewisii](images/keys/Hexastylis_lewisii_complete.jpg)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings1_sml.jpg)
Habit of Hexastylis lewisii. Note diffuse, non-clumping, arrangement along elongate rhizome (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings2_sml.jpg)
Habit of Hexastylis lewisii (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings3_sml.jpg)
Leaf of Hexastylis lewisii (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings4_sml.jpg)
Leaf of Hexastylis lewisii (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings5_sml.jpg)
Calyx of Hexastylis lewisii with free lobes removed; showing pilose trichomes in throat (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings6_sml.jpg)
Pilose pubescence of inner calyx of Hexastylis lewisii (Photo: Krings)
![Click to enlarge Hexastylis lewisii](http://herbarium.ncsu.edu/rare/images/hexastylis_lewisii_krings7_sml.jpg)
Androecium and gynoecium of Hexastylis lewisii (Photo: Krings)
Hexastylis lewisii
(rare; pocosin ecotones and upland forests; Pd, CP)
3’. Inner surface of calyx lobes puberulent...4.
4. Calyx tube broadly urceolate-campanulate or rhombic-ovate...5.
5. Calyx tube urceolate-campanulate, lobes 10–22 mm wide at base...H. shuttleworthii var. shuttleworthii
![Hexastylis shuttleworthii](images/keys/Hexastylis_shuttleworthii_complete.jpg)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_habit_sml.JPG)
Habit of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_lvs1_sml.JPG)
Leaf of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_fl1_sml.JPG)
Leaf of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_fl2_sml.JPG)
Calyx tube of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_interior1_sml.JPG)
Interior of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_interior2_sml.JPG)
Interior calyx surface of Hexastylis shuttleworthii (Photo: Krings)
![Click to enlarge Hexastylis shuttleworthii](http://herbarium.ncsu.edu/rare/images/Hexastylis_shuttleworthii_interior3_sml.JPG)
Interior of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii var. shuttleworthii
(uncommon; acidic deciduous forests, frequently along creeks; Mt)
5’. Calyx tube rhombic-ovate (broadest near the middle), lobes 3–8 mm wide at base...6.
6. Internal calyx tube reticulations absent or poorly developed, ridges < 1.0 mm if present; ovary ca. 1/3 inferior
...H. contracta
![Hexastylis contracta](images/keys/Hexastylis_contracta_complete.jpg)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings1_sml.jpg)
Leaf of Hexastylis contracta (Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings2_sml.jpg)
Hexastylis contracta (Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings3_sml.jpg)
Calyx lobes of Hexastylis contracta (Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings4_sml.jpg)
Internal throat hairs of Hexastylis contracta (calyx lobes removed; Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings5_sml.jpg)
Androecium and gynoecium of Hexastylis contracta. Note also the very low ridges on the internal tube walls (Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings6_sml.jpg)
Low internal ridges at base of tube of Hexastylis contracta (Photo: Krings)
![Click to enlarge Hexastylis contracta](http://herbarium.ncsu.edu/rare/images/hexastylis_contracta_krings7_sml.jpg)
1/3 inferior ovary of Hexastylis contracta (Photo: Krings)
Hexastylis contracta
(rare [State E, S1 G3]; acidic deciduous forests; Mt)
6’. Internal calyx tube reticulations well developed, ridges 1.5–2 mm tall; ovary superior...H. rhombiformis
![Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/Hexastylis_rhombiformis_complete2.jpg)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings1_sml.jpg)
Leaf of Hexastylis rhombiformis (Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings2_sml.jpg)
Calyx tube of Hexastylis rhombiformis (Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings3_sml.jpg)
Variation in calyx tube of Hexastylis rhombiformis (Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings4_sml.jpg)
Variation in calyx tube of Hexastylis rhombiformis (Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings5_sml.jpg)
Calyx lobes of Hexastylis rhombiformis (Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings6_sml.jpg)
Tall internal ridges of calyx tube of Hexastylis rhombiformis (upper portion of tube removed; Photo: Krings)
![Click to enlarge Hexastylis rhombiformis](http://herbarium.ncsu.edu/rare/images/hexastylis_rhombiformis_krings7_sml.jpg)
Androecium and gynoecium, as well as calyx wall, of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
(rare [SR-L, S3 G3]; acidic deciduous forests, frequently along creeks; Mt)
4’. Calyx tube cylindrical to narrowly cylindro-urceolate...7.
7. Calyx tube cylindrical to narrowly cylindro-urceolate, lobes 2–4 mm long, erect to slightly spreading...H. virginica
![Hexastylis virginica](images/keys/Hexastylis_virginica_complete.jpg)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings1_sml.jpg)
Leaf of Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings2_sml.jpg)
Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings3_sml.jpg)
Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings4_sml.jpg)
Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings5_sml.jpg)
Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings6_sml.jpg)
Hexastylis virginica (Photo: Krings)
![Click to enlarge Hexastylis virginica](http://herbarium.ncsu.edu/rare/images/hexastylis_virginica_krings7_sml.jpg)
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
(common; upland forests; Mt, Pd, CP)
7’. Calyx tube cylindrical, lobes 4–15 mm long, moderately spreading to reflexed...8.
8. Calyx tube usually wider than long, orifice wider than the lobe lengths (lobes 9-16 x 8-16 mm);
ovary ca. 1/3 inferior...9.
9. Calyx tube 12–25 mm long, leaves always densely variegated...H. minor
![Hexastylis minor](images/keys/Hexastylis_minor_complete.jpg)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_habitat_krings_sml.jpg)
Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings2_sml.jpg)
Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings3_sml.jpg)
Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings4_sml.jpg)
Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings7_sml.jpg)
Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings5_sml.jpg)
Trichomes on calyx lobe of Hexastylis minor (Photo: Krings)
![Click to enlarge Hexastylis minor](http://herbarium.ncsu.edu/rare/images/hexastylis_minor_krings6_sml.jpg)
Hexastylis minor (Photo: Krings)
Hexastylis minor
(common; dry to moist forests; Pd, CP)
9'. Calyx tube 8–18 mm long, leaves sparsely variegated to not variegated...Hexastylis sorriei
![Hexastylis sorriei](images/keys/Hexastylis_sorriei_complete.jpg)
Habitat of Hexastylis sorriei: Seep in sandhill Pinus palustris savanna (Photo: Krings)
Leaf of Hexastylis sorriei (Photo: Krings)
Leaf of Hexastylis sorriei (Photo: Krings)
Flower of Hexastylis sorriei (Photo: Krings)
Flower of Hexastylis sorriei (Photo: Krings)
Flower of Hexastylis sorriei (Photo: Krings)
Flower of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei
(infrequent; pocosin ecotones, margins of sandhill seeps; Pd/CP)
8’. Calyx tube at least as long as wide or longer, orifice narrower than the lobe lengths; ovary superior or
half-inferior...10.
10. Calyx tube 7–14 mm wide, lobes 6–17 mm wide; ovary superior; leaves typically not variegated
...H. heterophylla
![Hexastylis heterophylla](images/keys/Hexastylis_heterophylla_complete.jpg)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings1_sml.jpg)
Leaf of Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings2_sml.jpg)
Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings3_sml.jpg)
Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings4_sml.jpg)
Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings6_sml.jpg)
Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings5_sml.jpg)
Hexastylis heterophylla (Photo: Krings)
![Click to enlarge Hexastylis heterophylla](http://herbarium.ncsu.edu/rare/images/hexastylis_heterophylla_krings7_sml.jpg)
Androecium and gynoecium of Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
(common; dry to mesic forests; Mt, Pd)
10’. Calyx tube 4–7 mm wide, lobes 4–7 mm wide; ovary half-inferior; leaves freq. variegated...H. naniflora
![Hexastylis naniflora](images/keys/Hexastylis_naniflora_complete.jpg)
Habitat of Hexastylis naniflora: Moist Piedmont slope (Photo: Krings)
Habitat of Hexastylis naniflora: Moist Piedmont slope (Photo: Krings)
Leaves of Hexastylis naniflora (Photo: Krings)
Flower of Hexastylis naniflora (Photo: Krings)
Flower of Hexastylis naniflora (Photo: Krings)
Flower of Hexastylis naniflora (Photo: Krings)
Flower of Hexastylis naniflora (Photo: Krings)
Hexastylis naniflora
(rare [Fed T, S3 G]; acidic deciduous forests; Pd; late winter to spring)
**Errata in: Gaddy (
2011: A new species of
Hexastylis (Aristolochiaceae) from the Sandhills of North and South Carolina. Phytoneuron 2011-47: 1–5):
Under
Specimens Cited, the first specimen cited from Cumberland Co., NC, is actually
Hexastylis arifolia and should be deleted from the list of paratypes. Radford's collection number of the Richmond Co. specimen is
11336, not
11356.
Federally listed taxon—Hexastylis naniflora (Fed T, State T | S3 G3 |
NHP |
USFWS| Five-year review)
Habitat. "Acidic sandy loam soils along bluffs and nearby slopes, hillsides and ravines, in boggy areas adjacent to creekheads and streams. Soil type is the most important habitat requirement (Pacolet, Madison, or Musella types)" (USFWS)
Status. NHP recognizes seventy-six populations extant in North Carolina, of which forty-one are judged of good viability (as of Sep 2010). In the five-year review (2011; see above), the USFWS recognized 108 populations. This species may be a candidate for delisting in the future.
Threats. Residential/industrial development, conversion of woodlands to pasture, reservoir construction, herbicides.
![](images/lecture%20images/Hexastylis_naniflora_distr_SEUS.jpg) |
![Habitat, Hexastylis naniflora](images/lecture images/Hexastylis_naniflora_habitat_Hickory_Krings.jpg)
|
Distribution (courtesy SE Flora Atlas) |
Habitat (acidic, deciduous woods) |
![](images/lecture%20images/Hexastylis_naniflora_fls_Blomquist1957.jpg)
|
![Leaf](images/lecture images/Hexastylis_naniflora_lf_Hickory_Krings4.jpg)
|
Floral tube; detail of androecium and gynoecium |
Leaf (note variegation) |
![Leaf](images/lecture images/Hexastylis_naniflora_lf_Hickory_Krings3.jpg)
|
![Leaf](images/lecture images/Hexastylis_naniflora_lf_Hickory_Krings.jpg)
|
Leaves (note variegation) |
Leaf (note variegation) |
![Leaf](images/lecture images/Hexastylis_naniflora_lf_Hickory_Krings2.jpg)
|
![Flower](images/lecture images/Hexastylis_naniflora_fl_Hickory_Krings.jpg)
|
Leaf (note variagation) |
Flower |
![Flower](images/lecture images/Hexastylis_naniflora_fl_Hickory_Krings2.jpg)
|
![Flower](images/lecture images/Hexastylis_naniflora_fl_Hickory_Krings3.jpg)
|
Flower (lateral view) |
Flower |
![](images/lecture%20images/Hexastylis_naniflora_pollen_Padgett2004.jpg)
|
![](images/lecture%20images/Hexastylis_heterophylla_pollen_Padgett2004.jpg)
|
Hexastylis naniflora pollen (no "bumps") [Padget
2004] |
Hexastylis heterophylla pollen (note surface bumps)
[Padget 2004] |
![](images/lecture images/Hexastylis_minor_pollen_Padgett2004.jpg)
|
|
Hexastylis minor pollen (not surface bumps) [Padget
2004] |
|
Similar taxa:
Leaves of Asarum canadense may at first glance be taken for Hexastylis. However, note the distinct differences in flower structure in the images below. In contrast to Hexastylis, the flowers of Asarum canadense are conspicuously pubescent and the perianth lobes are not completely fused into a tube.
Asarum can be distinguished from
Hexastylis by the following key:
1. Sepals distinct (although touching valvately and forming a
well-defined false tube); anthers each with prominent terminal appendages;
styles connate in a column...Asarum
1'. Sepals connate for most of their length; anthers lacking
terminal appendages; styles distinct (except sometimes at extreme base)...Hexastylis
![](images/lecture images/Asarum_Hexastylis_FNA.gif)
Illustration from Flora of North America (1997)
Leaf of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Flower of Asarum canadense (Photo: Krings)
Kelly (1998 [Amer. J. Bot.]) has argued for submerging
Hexastylis into Asarum, based on molecular evidence. He proposed this
action as a "conservative" approach, arguing that the alternative of recognizing
two clades would require over fifty new combinations. However, Kelly did
not consider an additional alternative of recognizing more than two clades,
which would require negligible new combinations and be consistent with the
morphological/molecular distinctness he himself noted.
![](images/lecture images/Asarum_Hexastylis_phylogeny_Kelly1998.gif)
Fig. 4. The strict consensus tree of the 288 equally most
parsimonious trees from the simultaneous analysis of ITS and morphological data.
Taxon areas are indicated to the right. The 9-bp deletion is mapped on the tree.
Sepal morphology is mapped for species of Asarum s.s.; the characters
shown were not included in the morphological analysis because of continuous
variation. Mapping indicates that the extremes of variation have been
independently derived in North America and in Asia (Kelly 1998 [Amer. J. Bot.]).