Research Article |
Corresponding author: Warren L. Wagner ( wagnerw@si.edu ) Academic editor: James Miller
© 2015 David H. Lorence, Warren L. Wagner.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lorence DH, Wagner WL (2015) Heliotropium (Boraginaceae) in the Marquesas Islands (French Polynesia) with description of a new species. PhytoKeys 47: 49-57. https://doi.org/10.3897/phytokeys.47.8767
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During the preparation of the Vascular Flora of the Marquesas Islands a new endemic species of Heliotropium L. (Boraginaceae) has come to light and is described herein: Heliotropium perlmanii Lorence & W. L. Wagner. It is known only from the island of Eiao and appears most closely related to H. marchionicum Decne., also endemic to the Marquesas and known from Nuku Hiva. An amended description of H. marchionicum and key to separate the Marquesan species are given and their differences discussed.
Heliotropium , Boraginaceae , Marquesas Islands, French Polynesia
The Flora of the Marquesas Islands project is a collaborative program primarily between the Smithsonian Institution and the National Tropical Botanical Garden intended to further knowledge of the flora of this remote archipelago. In 1997 the first publications of new species and revisions of genera with at least one endemic species were initiated (
A number of recent studies utilizing both molecular and morphological analyses suggest that the traditional Boraginaceae s.l. should be split into a number of families (see
Heliotropium (Boraginaceae subfam. Heliotropoideae, or Heliotropiaceae of many authors) consists of 280 to 350 species of herbs, shrubs, lianas and small trees from the temperate and warm regions of the world, mostly in arid zones, with the greatest diversity in the New World (
In the Marquesas Islands (SE Polynesia) only a single native species, Heliotropium marchionicum Decne. has been previously recorded (
1 | Stems virgately branched, with two subequal lateral branches developing adjacent to inflorescence; indument of trichomes 0.1–0.2 mm long; leaves opposite, blade with secondary veins obscure; calyx lobes unequal, 1–2 larger, ovate, 1–1.5 × 0.6–1 mm, 3–43 smaller, narrowly ovate to oblong, 1 × 0.3–0.4 mm; corolla 2 mm, tube 1–1.5 mm, lobes 0.5–0.7 × 0.5–0.7 mm; ovary glabrous except for ring of trichomes 0.2–0.3 mm long surrounding base of style; fruits 1 × 1.6 mm | H. perlmanii |
– | Stems sympodially branched, with usually only a single lateral branch developing adjacent to inflorescence; indument of trichomes 0.2–0.5 mm long; leaves subopposite to alternate, blade with secondary veins visible; calyx lobes subequal, 1.7–2.2 × 0.7–1.2 mm; corolla 2.6–3.2 mm, tube 2.0–2.2 mm, lobes 1.2–1.7 × 0.8–1.3 mm mm; ovary densely strigillose; fruits 1.5–2 × 2 mm | H. marchionicum |
Marquesas Islands. Eiao, north side of large valley which is south of Vaittuha Valley, Opituha Valley. Sea cliffs, with Heliotropium, Dodonaea, Cordia lutea. Shrubs 1-2 ft. tall; flower; leaves smaller than Nuku Hiva plants; not silvery, 1050 ft [320 m], 7 Jul 1988, S. Perlman & J. Florence 10052 (Holotype PTBG 009229; Isotypes BISH, F, MO, P, PAP, US). Figure
Heliotropium perlmanii Lorence & WL Wagner A Habit B Upper leaf surface, C Inflorecence D Flower, lateral view E Corolla, face view F Flower, longitudinal section showing stamens and gynoecium G Corolla, sectioned to show stamens and indument, H Fruit and calyx, lateral view I Fruit showing 4 carpels. All figures drawn from Perlman & Florence 10052 (US) and photos from Falaise Est Eiao, 11 March 2007 courtesy of J-F Butaud.
Shrubs 30–60 cm tall, stems decumbent, virgately branched, with two subequal lateral branches developing adjacent to inflorescence; leafy stems 0.8–1.5 mm in diam., terete, brown, moderately shortly strigillose-canescent with white ascendant trichomes 0.1–0.2 mm long; older stems with peeling brown bark. Leaves opposite, blade elliptic to broadly elliptic or obovate-elliptic, 0.8–1.8 × 0.3–1.0 cm, apex obtuse to rounded, or occasionally truncate, usually apiculate, base acutely cuneate, sides slightly attenuate and decurrent, subcoriaceous to coriaceous and brown when dry, bright green when fresh, both surfaces moderately shortly strigillose with appressed trichomes 0.1–0.2 mm, pustular, venation obscure, 1–2 (–3) pairs secondary veins arising near base, petiole 2–4 mm long, 0.4–0.5 mm in diam., shortly strigillose. Inflorescences terminal, scorpioid-cymose, forked 1(–2) times, axes densely shortly strigillose-canescent like the stems, 2–4 cm long, peduncle 0.5–0.8 cm long, primary axes 1.5–3 cm long. Flowers sessile to subsessile, 18–23 per axis, calyx lobes 5, unequal, 1–2 larger, ovate, 1–1.5 × 0.6–1 mm, 3–4 smaller, narrowly ovate to oblong, 1 × 0.3–0.4 mm, densely strigillose toward base, corolla shortly funnelform, 2 mm long, tube 1-1.5 mm long, externally densely villose-strigillose, internally slightly villosulous in throat, lobes 5, subcircular, 0.5–0.7 × 0.5–0.7 mm, margin crisped, stamens 5, attached midway in tube, basifixed, anthers ellipsoid, 0.5–0.6 mm long, apiculate; ovary cylindric-ovoid, glabrous except for ring of trichomes 0.2–0.3 mm long surrounding base of style, style terminal, 0.3 mm long, stigmatic column 0.3–0.4 mm long, cylindrical-conical, apex strigillose, base annular. Fruit broadly ovoid, 1 × 1.6 mm, shortly strigillose, shallowly 4-lobed, dry, splitting into 4 wedge-shaped nutlets. Nutlets 1.3–1.5 × 1–1.1 mm, dorsally strigillose, ventrally glabrous, brown.
Marquesas Islands, known only from three collections made on Eiao.
Grows on windward sea cliffs, with Dodonaea viscosa Jacq., Cordia lutea Lam., and Bidens beckiana (F. Br.) Sherff.
Endangered (EN): B1ab (i, ii, iii) + 2ab (i, ii, iii): B2: total area of occupancy less than 500 km² (ca. 47 km²). B1a, severely fragmented; B1b (1–iii), habitat quality continuing decline inferred. The suitable habitat for Heliotropium perlmanii on Eiao (40 km²) is indicated as an endangered environment, threatened by feral animals and invasive plants, thus reducing the extent of the suitable habitat. Eiao has populations of feral sheep, pigs, cats, and rats (J.-F. Butaud, pers. comm. 2013).
We are pleased to name this new species in recognition of rough-terrain botanist Steven P. Perlman (National Tropical Botanical Garden) in recognition of his contributions to our knowledge of the flora of the Pacific region. Steve collected the type specimen and in his label data noted several differences from H. marchionicum.
Collections of this species were previously distributed as H. marchionicum, which differs by its non-virgate sympodial branching, more densely strigillose indument, larger leaves, and flowers about twice as large with corollas 2.6–3.2 mm long.
Marquesas Islands. Eiao: 20 September 1922, R.H. Beck & W.B. Jones 1537 (A, BISH); NW side of island, Vaituha Bay and summit ridge of island 400 m elevation, 1 August 1977, B.H. Gagné 1295 (BISH).
Iles Marquises [Marquesas Islands], Noukahiva [Nuku Hiva], “toutemanou”, pl. herbacee sur le sommet du montagne, Le Bastard 76 (Holotype P, digital image!). Figure
Heliotropium marchionicum Decne. A Habit B Upper Leaf surface C Inflorescence D Flower, lateral view E Corolla, face view F Flower, longitudinal section showing stamens and gynoecium G Corolla, sectioned to show stamens and indument H Fruit and calyx, lateral view I Fruit showing 4 carpels. Drawn from Perlman 10005 (US) and photos from Nuku Hiva, 24 February 2007 [A], Mercier 1847 (US) and photos from Nuku Hiva, 24 February 2007 courtesy of J-F Butaud [B–I].
Shrubs or suffrutescent perennials 1–2 m tall, stems erect or decumbent, sympodially branched, with usually only a single lateral branch 30–120 cm long developing adjacent to inflorescence, terete, 1.5-3 mm diam., most parts densely silvery white strigillose with ascending white trichomes 0.2–0.5 mm long. Leaves subopposite to alternate, blade elliptic to narrowly elliptic or obovate-elliptic, 1–5 × 0.5–1.5 cm, apex acute, obtuse or rounded, often apiculate, base acute to narrowly cuneate, sometimes attenuate, chartaceous to subcoriaceous, both surfaces strigillose to densely white strigillose with appressed white trichomes 0.2–0.4 mm long, smooth or sometimes pustular, secondary veins 2–3 pairs arising in basal half of lamina; petiole 3–15 mm. Inflorescences terminal and later displaced by growth of one axillary bud, or sometimes leaf-opposed, scorpioid-cymose, densely white strigillose as for stems and leaves, 4–7 cm long, forked once, peduncle 1–2 cm long, primary branches 2.5–7 cm long, each with 17–35 flowers. Flowers sessile or subsessile, calyx lobes 5, densely white strigillose, free to the base, subequal, ovate to lanceolate, 1.7–2.2 × 0.7–1.2 mm, acute to acuminate; corolla shortly funnelform, 2.6–3.2 mm, tube 2.0–2.2 mm, externally strigillose except at base, internally with pubescent lines below the lobes, lobes 5, subcircular, 1.2-1.7 × 0.8–1.3 mm × 0.8 mm, margins crisped, dorsally strigillose medially; stamens 5, attached below middle of tube, basifixed, anthers linear-oblong, 0.6–0.7 mm long, glabrous, not connate; ovary ovoid, densely strigillose, 0.5 mm long, style terminal, 0.3–0.6 mm long, glabrous, stigmatic column 0.4–0.5 mm, cylindrical-conical, papillose, apex strigillose, base annular. Fruit broadly ovoid, 1.5–2.0 × 2.0 mm, shallowly 4-lobed, externally strigillose, dry, splitting into 4 wedge-shaped nutlets. Nutlets 1.4–1.6 × 0.8–1.0 mm, apiculate, dorsally densely strigillose, ventrally glabrous, dark brown.
Marquesas Islands, known only from Nuku Hiva.
This species usually occurs inland on basaltic cliffs and dry ridges, sometimes near waterfalls, in dry land forest with Sapindus saponaria L., Cerbera manghas L., and introduced invasive species including Tecoma stans (L.) Kunth and Leucaena leucocephala (Lam.) De Wit. The label on one collection notes it is a low elevation littoral plant (Brown 542, BISH).
Proposed IUCN Red List Category Endangered (EN): B1ab (i, ii, iii) + 2ab (i, ii, iii): B2: total area of occupancy less than 500 km² (ca. 50 km²). B1a, severely fragmented; B1b (1–iii), habitat quality continuing decline inferred. The suitable habitat for Heliotropium marchionicum on Nuku Huka (ca. 340 km²) is indicated as an endangered environment, threatened by human activity (deforestation), feral animals, and invasive plants, thus reducing the extent of the suitable habitat.
Heliotropium marchionicum is apparently closely related to H. permanii but differs by the characters noted above. A single collection from Taiohae, Nuku Hiva (Florence 8394, BISH, CHR, K, NY, P, US) resembles H. marchionicum superficially but differs in having stems and petioles pilose with hairs to 1 mm long, inflorescence axis pilose, very small flowers (calyx lobes 1–1.1 mm long, corolla 1.1–1.3 mm long) and ribbed fruits 1.1–1.2 × 1.8 mm, covered with bulbous-tuberculate scales, splitting into 4 nutlets. Further collections are needed to determine whether it represents an undescribed taxon or alternatively a naturalized species. It closely resembles H. angiospermum Murr., native to North America and the Caribbean and was identified by M. Strong (US) as this species.
Marquesas Islands. Nuku Hiva: Hakaui, 20 July 1921, F.B.H. Brown 542 (BISH); Moyenne vallée de Hakaui, flanc droit, 125 m, latitude 08°54'S, longitude 140°10'W, 18 May 1984, J. Florence 6695 (BISH, P); Hakaui Valley, 107 m elevation, 26 June 1988, S. P. Perlman 10005 (AD, BISH, F, MO, MU, NY, OS, P, PAP, PTBG, US); Matatekouaehi Valley, about 2 miles in from coast, by 100 ft. waterfall, 1 July 1988, S. P. Perlman 10026 (BISH, PTBG, US); Taiohae, flanc gauche de la baie, S du CJA, 150 m, latitude 08°56’S, longitude 140°05’W, 26 Jul 1987, J. Florence 8394 (BISH, CHR, K, NY, P, US),W shore, 200 m elevation, 20 Oct 1922, W.B. Jones 1591 (BKL); slope on ridge, 18 Oct 1922, E.H. Quayle 1591 (A, BISH); without precise locality, 1841, R. Hinds s.n. (P), Mathias 96 (GH), 1847, M.P. Mercier s.n. (P, US).
We are grateful to the Bishop Museum herbarium (BISH) and MNHN Paris (P) for loan of their collections. Specimens were collected by Steve Perlman and Jacques Florence as part of the Vascular Flora of the Marquesas Islands project, which was supported by a generous private donation to the National Tropical Botanical Garden, and the Flore de la Polynésie française project. We are grateful to the Délégation à la Recherche (Papeete, Tahiti), the Musée de Tahiti et des Iles, and the Service du Développement Rural for logistic support, the Délégation for permission to collect in the Marquesas, and the Délégation and the Haut-Commissariat de la République en Polynésie française for permission to conduct research. The contribution by WLW to this study was partially supported by the Smithsonian Research Opportunities Fund and the National Tropical Botanical Garden’s McBryde Endowment for Hawaiian and Pacific Botany. We thank Alice Tangerini for masterfully preparing the two illustrations. We also appreciate the comments, images, and insights on Marquesas Heliotropium by Jean-François Butaud and Jacques Florence, and the constructive comments provided by two anonymous reviewers.