E E  W o r k s h o p: Rare Plants of North Carolina

Hexastylis Raf. (Aristolochiaceae)

Hexastylis is represented by ten species in North Carolina. Only Hexastylis naniflora is listed at the federal level (Fed T, S3 G3). Hexastylis contracta is currently state listed (State E, S1 G3). Hexastylis rhombiformis was state listed as threatened (State T, S2 G2), but was recently reassessed (S3 G3) and downlisted. It continues to be tracked as State SR-L and remains a Federal Species of Concern. No other taxa are listed or proposed for listing.

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
Hexastylis arifolia
Leaf of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
Leaves of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
Flower of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
Flower of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
Flower of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
Flower detail of Hexastylis arifolia (Photo: Krings)
Hexastylis arifolia
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

Hexastylis rosei
Hexastylis rosei
Habit of Hexastylis rosei (Photo: Krings)
Hexastylis rosei
Leaf of Hexastylis rosei (Photo: Krings). Note minute teeth along leaf margins, as well as the impressed midvein.
Hexastylis rosei
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.
Hexastylis rosei
Leaf of Hexastylis rosei (Photo: Krings).
Hexastylis rosei
Flower of Hexastylis rosei (Photo: Krings).
Hexastylis rosei
Flowers of Hexastylis rosei (Photo: Krings).
Hexastylis rosei
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
Hexastylis lewisii
Habit of Hexastylis lewisii. Note diffuse, non-clumping, arrangement along elongate rhizome (Photo: Krings)
Hexastylis lewisii
Habit of Hexastylis lewisii (Photo: Krings)
Hexastylis lewisii
Leaf of Hexastylis lewisii (Photo: Krings)
Hexastylis lewisii
Leaf of Hexastylis lewisii (Photo: Krings)
Hexastylis lewisii
Calyx of Hexastylis lewisii with free lobes removed; showing pilose trichomes in throat (Photo: Krings)
Hexastylis lewisii
Pilose pubescence of inner calyx of Hexastylis lewisii (Photo: Krings)
Hexastylis lewisii
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
Hexastylis shuttleworthii
Habit of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
Leaf of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
Leaf of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
Calyx tube of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
Interior of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
Interior calyx surface of Hexastylis shuttleworthii (Photo: Krings)
Hexastylis shuttleworthii
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
Hexastylis contracta
Leaf of Hexastylis contracta (Photo: Krings)
Hexastylis contracta
Hexastylis contracta (Photo: Krings)
Hexastylis contracta
Calyx lobes of Hexastylis contracta (Photo: Krings)
Hexastylis contracta
Internal throat hairs of Hexastylis contracta (calyx lobes removed; Photo: Krings)
Hexastylis contracta
Androecium and gynoecium of Hexastylis contracta. Note also the very low ridges on the internal tube walls (Photo: Krings)
Hexastylis contracta
Low internal ridges at base of tube of Hexastylis contracta (Photo: Krings)
Hexastylis contracta
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
Hexastylis rhombiformis
Leaf of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
Calyx tube of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
Variation in calyx tube of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
Variation in calyx tube of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
Calyx lobes of Hexastylis rhombiformis (Photo: Krings)
Hexastylis rhombiformis
Tall internal ridges of calyx tube of Hexastylis rhombiformis (upper portion of tube removed; Photo: Krings)
Hexastylis rhombiformis
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
Hexastylis virginica
Leaf of Hexastylis virginica (Photo: Krings)
Hexastylis virginica
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
Hexastylis virginica (Photo: Krings)
Hexastylis virginica
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
Hexastylis minor
Hexastylis minor (Photo: Krings)
Hexastylis minor
Hexastylis minor (Photo: Krings)
Hexastylis minor
Hexastylis minor (Photo: Krings)
Hexastylis minor
Hexastylis minor (Photo: Krings)
Hexastylis minor
Hexastylis minor (Photo: Krings)
Hexastylis minor
Trichomes on calyx lobe of Hexastylis minor (Photo: Krings)
Hexastylis minor
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

Hexastylis sorriei habitat
Habitat of Hexastylis sorriei: Seep in sandhill Pinus palustris savanna (Photo: Krings)
Hexastylis sorriei leaf
Leaf of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei leaf
Leaf of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei flower
Flower of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei flower
Flower of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei flower
Flower of Hexastylis sorriei (Photo: Krings)
Hexastylis sorriei flower
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
Hexastylis heterophylla
Leaf of Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
Hexastylis heterophylla (Photo: Krings)
Hexastylis heterophylla
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

Hexastylis naniflora habitat
Habitat of Hexastylis naniflora: Moist Piedmont slope (Photo: Krings)
Hexastylis naniflora habitat
Habitat of Hexastylis naniflora: Moist Piedmont slope (Photo: Krings)
Hexastylis naniflora habitat
Leaves of Hexastylis naniflora (Photo: Krings)
Hexastylis naniflora flower
Flower of Hexastylis naniflora (Photo: Krings)
Hexastylis naniflora flower
Flower of Hexastylis naniflora (Photo: Krings)
Hexastylis naniflora flower
Flower of Hexastylis naniflora (Photo: Krings)
Hexastylis naniflora flower
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.

Habitat, Hexastylis naniflora

Distribution (courtesy SE Flora Atlas)

Habitat (acidic, deciduous woods)

Leaf

Floral tube; detail of androecium and gynoecium

Leaf (note variegation)

Leaf

Leaf

Leaves (note variegation)

Leaf (note variegation)

Leaf

Flower

Leaf (note variagation)

Flower

Flower

Flower

Flower (lateral view)

Flower

Hexastylis naniflora pollen (no "bumps") [Padget 2004]

Hexastylis heterophylla pollen (note surface bumps) [Padget 2004]

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

Illustration from Flora of North America (1997)

Asarum canadense 
Leaf of Asarum canadense (Photo: Krings)
Asarum canadense 
Flower of Asarum canadense (Photo: Krings)
Asarum canadense 
Flower of Asarum canadense (Photo: Krings)
Asarum canadense 
Flower of Asarum canadense (Photo: Krings)
Asarum canadense 
Flower of Asarum canadense (Photo: Krings)
Asarum canadense 
Flower of Asarum canadense (Photo: Krings)
Asarum canadense 
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.

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.]).