Helonias bullata Linnaeus (1753: 342)

Tanaka, Noriyuki, 2019, Taxonomy, evolution and phylogeography of the genus Helonias (Melanthiaceae) revisited, Phytotaxa 390 (1), pp. 448-450 : 448-450

1. Helonias bullata Linnaeus (1753: 342) , p.p., excl. Ephemerum phalangoides virginianum flosculis arbuteis bullatis aureis in spicam dispositis in Plukenet (1692: t. 174, f. 5; 1696: 135) and in Morison (1699: 606, sect. 15, t. 2, f. 1).

( Figs. 1A , 4 , 5 ).

Type (lectotype designated by J.L. Reveal in Jarvis et al. 1993: 53):― U.S.A. Habitat in Pensylvaniae paludosis, Pehr Kalm s.n. (LINN No. 471.1*!) . Isolectotype: BM n.v.

≡ Helonias latifolia Michaux (1803: 212) .

Veratrum racemo simplicissimo , corollis patentibus, staminibus longioribus Miller (1760: 181, t. 272).

Veratrum scapo fistuloso squamoso, spica stricta Ehret in Trew (1771: 41, t. 77, p.p., excl. A–C).

Colloquial name:―Swamp pink.

Japanese name: Amerika (America)-shôjôbakama (nov.).

Description:―Rhizome vertical or ascending, cylindrical, closely annulate with numerous scars, to 6 cm or longer, to 3 cm in diam. Roots filiform, some contractile, to 2 mm in diam., pale dull orangish (or pinkish) brown (or pale brick colored) when dry. Leaves slightly hysteranthous, main fresh rosette leaves 6–8(–13), usually persistent for 1 year, spatulate or oblanceolate, to 32 cm long, to 4 cm wide, tapering to cuneate base, margins entire, apex acute or acuminate, apiculus 0.5–0.8 mm long. Flowering stem 22–40 cm long at anthesis, to 72 cm long in fruit; peduncle terete, 20–37 cm long at anthesis, elongating to 62 cm long in fruit; scale-like leaves on peduncle 9–11 (excl. basal ones), becoming shorter and laxer upward, ovate, oblong-ovate, lanceolate or narrowly deltoid, to 3.8 cm long, margins submembranous, apex acute or acuminate; inflorescence racemose, ovoid, narrowly ovoid or ellipsoid, 3–5 cm long, 2–2.5 cm in diam., rachis 2–4 cm long at anthesis, elongating to 12.5 cm in fruit; pedicels terete, 6-ribbed, pale pink, 2.5–6 mm long at anthesis, 5–10 mm long in fruit. Flowers 30–72 per inflorescence, spirally arranged, crateriform or funnelform, 7–10 mm across, facing upward (distal flowers), spreading horizontally (middle flowers) or downward (proximal flowers) at mid anthesis, fragrant, nearly adichogamous. Tepals ascending, pink, darker distally, slightly imbricate proximally, narrowly oblong or narrowly oblanceolate, 5–7 mm long, 1.5–2.1 mm wide, apex obtuse or (sub)acute, veins 3 or 5, adaxial surface slightly canaliculate toward base; basal nectary almost flat, abaxial basal portion slightly inflated. Stamens 6; filaments ascending, pale pink, filiform-subulate, slightly complanate, 4.5–8 mm long, gradually widening toward base, apex subacute; inner filaments adnate to base of ovary; anthers extrorse (antrorse apically), adaxially basifixed, ovoid-ellipsoid, base sagittate, unilocular, abaxial side longitudinally septate (furrowed) between thecae, adaxial side not septate nor furrowed, 1.5–2 mm long, whitish with blue tint, dark blue along suture of sacs, exserted beyond tepals. Pistil 1; ovary ovoid or ellipsoid, obtusely trigonous, apex emarginate, 2–3 mm long, 2–3 mm in diam., dark (greenish) purplish-pink; styles 3, often shortly connate basally into a single column to 0.6 mm long, free part of style ventrally stigmatic, papillulate, linear, 1.5–2(–2.5) mm long, recurved, white; ovules 6–18 per locule, biseriate on central axile placentae. Capsules antrorse, obpyramidal (obcordate laterally), 5–7 mm long, 10–12 mm broad, apex emarginate-sunken, trilobed; lobes ovoid-conic, rounded at apex, ascending. Seeds ascending within capsular cavity, with white testa, subfusiform, 4.5–5.5 mm long, 0.7–1 mm wide, slightly flattened, often somewhat falcate, subcaudate proximally, caudate distally; body of seed located on one side in middle within testa, fusiform, 1.1–1.8 mm long, 0.4–0.7 mm wide, brown.

Additional specimens examined:―U.S.A. Delaware: New Castle Co., Belltown Run, NW of Glasgow Station , 21 April 1938, fl., R.R. Tatnall 3662 (GH); In paludosis civitatis, fl., Canby 15276 (BM-001118053); Farnhurst?, 13 May 1896, fr., A. Commons (GH); Swamps near Wilmington, Wm. M. Canby s.n. (K) . New Jersey: Burlington Co., Buddtown, 30 April 1932, fl., H.M. Rusk & H.K. Svenson 6126 (P-02101625*); Camden Co. , cultivated from roots taken at Atco, May 1871, fl., J. H. Redfield s.n. (TI); swamps, near Atco, 3 May 1872, fl., J. H. Redfield s.n. (TI, 3 sheets); Middlesex Co ., New Brunswick, 8 May 1932, M.A. Chrysler s.n. (P-02101626*); Morris Co., Budd’s Lake, 7 May 1905, fl. & fr., K. K. MacKenzie 1209 (E-00115524); Budd’s Lake , 13 May 1906, 11 June 1905, fl. & fr., W.W. Eggleston s.n. (K, P-01879677*); Salem Co. , Whiglane, 14 April 1935, fl., B. Long 45636 (GH); Newfield, May 1900, fl., H.E. Stone s.n. (BM-001118054) . North Carolina: Transylvania Co., Pisgah National Forest , 7 May 2007, G. Kokubugata 6414 (TNS) .? Pennsylvania: Near Philadelphia , fl., Torrey s.n. (K) . Virginia: Henrico Co., Whitecoak Swamp, west of Elko Station , 21 September 1938, fr., Fernald & Long 9295 (GH). Cultivated material: Botanic Gard. of Harvard Univ. , 1879, fr., A.Gray (K); Hort . Edinburgh ( Bot. Gard. Edinburgh), 14 May 1895, fl. (E-00115523) .

Distribution:― U.S.A. Southern uplands of the Blue Ridge Mountains and on coastal plain northward; Delaware, Georgia, Maryland, New Jersey, New York, North Carolina, South Carolinas, Pennsylvania (not reported since the late 19th century), Virginia ( Gleason 1952; Utech 1978, 2002). The population in New York is considered extirpated ( USFWS 1988, 1991) ( Figs. 6 , 31-1 ).

Habitat:―Freshwater wetlands including spring seepages, swamps, bogs, meadows, and margins of small streams at elevations 0–1100 m ( USFWS 1988, 1991, Utech 2002, Chafin 2007).

Conservation status:― Helonias bullata is threatened throughout the range ( USFWS 1988, 1991). Threats to the species are outlined by Chafin (2007) as follows: Conversion of habitat to farmland and pasture, pollution runoff and sedimentation into wetlands, ditching and draining of wetlands, poaching, and encroachment by shrubs and trees. Conservation measures and recovery plans for threatened or endangered populations of this species are detailed in USFWS (1991). The populations vary extensively in size, stability, and quality of habitat conditions ( USFWS 1991). While H. bullata is threatened, endangered or extirpated in several sites, it is locally abundant in New Jersey ( Sutter 1984). It is noteworthy that a population in North Carolina contains over 100,000 rosettes, and the Blue Ridge populations in Virginia has a total of 15,000 plants ( USFWS 1991). Having such large populations, H. bullata as a species is not assigned to any of the threatened categories CR, EN or VU, when assessed by the criterion D or D 1 in the IUCN Red List Categories and Criteria (2001).

Flowering:―April–May.

Ripening:―June.

Remarks:―Although Helonias bullata was reported by Linnaeus as growing in a swamp in Pennsylvania, Gates (1918) stated that “the plant was originally discovered by Kalm near Philadelphia, probably on April 26, 1749, at Pennsneck, N[ew]. J[ersey]. It was formerly supposed to have occurred in eastern Pennsylvania, but this record was probably a mistake.” The persistence of its occurrence in that state has been doubted at least since the late 19th century (e.g. Britton & Brown 1896, 1913; Gates 1918; Utech 1978, 2002). I examined Torrey’s specimen, recorded as from ‘near Philadelphia’ (K, originally in Herbarium Hookerianum 1867). The specimen is cited under Pennsylvania in the list below, but it is uncertain whether it was actually collected in Pennsylvania or in nearby New Jersey. Gray (1848, 1868) included Pennsylvania within the range of Helonias . A detailed account of the distribution and habitat of H. bullata is provided in USFWS (1991).

Figures

FIGURE 1. Buds and their sprouting in Helonias. A. H. bullata (material Bu-1; photo on 5 April 2013). B. H. orientalis (O-3; 15 November 2014). C, D. H. thibetica (T-1; C on 19 February 2014, D on 2 March 2014). Flower buds or fresh flowering stem designated by F, leaf buds or sprouting leaves by L, rosulate leaves developed in previous years by R. In Fig. C, two of leaves developed in the preceding year marked with single white circle (dot); two of leaves developed the year before last marked with paired white circles (dots). Photo A by Jim Fowler at site of “Bu-1” in South Carolina; B–D by the author at Hachioji, Tokyo.

FIGURE 4. Helonias bullata (material Bu-1; photos A–E on 28 April 2014, F on 13 June 2002). A. Habit and boggy habitat. B. Young opening flowers with indehiscent anthers. C, D. Young flowers visited by a Bombus species sucking nectar.E. Flowers visited by a halictid bee. F. Fully dehisced, ascending capsules. All photos taken by Jim Fowler.

FIGURE 5. Helonias bullata (A–E; material Bu-2). A. Young flower with indehiscent anthers. B. Anthers seen from above. Three outer anthers dehisced earlier than three inner ones of which one marked with IS. C. Proximal interior of partial flower, showing nectar (n) secreted at adaxial bases of tepals. D. Pistil showing basal connation (*) among three styles. E. Proximal portion of flower in vertical section. IF; inner filament. OF; outer filament. F. Seeds (Bot. Gard. Harvard Univ., year 1879, K). Photos A, C, E on 21 May 1996, B on 22 May 1996, D on 28 May 1996, all taken at Hachioji, Tokyo.

FIGURE 6. Geographic range (dotted areas) of Helonias bullata in eastern North America (Data based principally on Utech 1978).

FIGURE 31. Geographic ranges of two sections Helonias and Heloniopsis. Ancestral plants of Helonias are presumed to have been in northern high latitudes approximate area of which is encircled with broken line.They are supposed to have been separated into two groups, and eventually one group reached eastern N. America and the other eastern Asia as consequence of their southward migrations (A). ① Sect. Helonias. ②–④ Sect. Heloniopsis. ② Subsect. Ypsilandra. ③ Subsect. Heloniopsis ser. Umbellatae. ④ Subsect. Heloniopsis ser. Heloniopsis. For historical aspect of range extension, see also Figs. 32, 33 and text (Part III).