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A reassessment of collections of Marquesan Apocynaceae assigned to the genera Neisosperma Raf., Ochrosia Juss., and Rauvolfia L. revealed that two nomenclatural changes are necessary: 1) transfer of Neisosperma brownii Fosberg & Sachet to the genus Ochrosia, as Ochrosia brownii (Fosberg & Sachet) Lorence & Butaud, comb. nov., and 2) transfer of Ochrosia nukuhivensis Fosberg & Sachet to Rauvolfia as Rauvolfia nukuhivensis (Fosberg & Sachet) Lorence & Butaud, comb. nov. As a result, two species each of Ochrosia and Rauvolfia are recognized from the Marquesas Islands, all endemic. Recent field work has yielded important new data on their distribution, habitat, and conservation status. It is recommended that all four species should be added the IUCN Red List at the Critically Endangered (CR) category.
Apocynaceae, French Polynesia, Marquesas Islands, Neisosperma, Ochrosia, Rauvolfia
In his Flora of Southeastern Polynesia, Forrest
During the preparation of an account of Apocynaceae
for the Vascular Flora of the Marquesas Islands and Flore de la
Polynésie française, additional field work by J-FB and study of
additional herbarium specimens have greatly increased our knowledge of
the Marquesan taxa and their correct identity. For example, collections
from field work associated with the project resulted in the discovery
of a new species of Lepinia, a genus previously not even known from the Marquesas (
Traditionally Neisosperma [incorrectly spelled Neiosperma by Fosberg & Sachet (1972)] was separated from Ochrosia
based on two morphological features: the gynoecium possesses a nectary
comprised of two thin lobes or scales present in the former but lacking
in the latter; secondly, the fruit endocarp is fibrous and muricate
with no air cavity in the former and fibrous but relatively
smooth-surfaced with two air cavities in the latter (
Based on morphological and molecular evidence, recent classifications of Apocynaceae tend to recognize only a single genus Ochrosia (
All measurements given herein are taken from dried
herbarium specimens, although certain features such as shapes were
supplemented with information from field notes, alcohol-preserved
specimens, and digital photos. Measurements are presented in the
descriptions as follows: length × width (or length × width × thickness
in the case of fruits and seeds), followed by units of measurement (mm
or cm). All specimens cited in this paper have been seen by the authors.
Specimens from the following herbaria were studied: AD, BISH, K, MO,
MU, PAP, PTBG, and US. The area of occupancy (distribution) for
these species was calculated using herbarium collection data and field
observations, and the conservation status is proposed following the
IUCN Red List Category criteria (IUCN
1a | Calyx lobes c. 2 mm long; corolla lobes 6-7 mm long; mericarps ellipsoid 37-53 × 27-37 mm; Nuku Hiva | Ochrosia brownii |
1b | Calyx lobes c. 2.5 mm long; corolla lobes 9-10 mm long; mericarps ovoid-ellipsoid 61-78 × 32-44 mm; Fatu Hiva | Ochrosia fatuhivensis |
urn:lsid:ipni.org:names:77112735-1
http://species-id.net/wiki/Ochrosia_brownii
Figs 1A, BMarquesas Islands: Nuku Hiva: without precise locality, 15 July 1921, F. B. H. Brown 541 (Holotype BISH-500905!).
Ochrosia parviflora sensu F. Br., non (G. Forst.) Henslow
Tree to 13 m tall, trunk to 24 cm diam., branchlets glabrous, leafy twigs 4−4.5 mm in diam, terete, drying wrinkled, older leafless twigs 6−7 mm in diam., latex white. Leaves opposite on smaller branchlets, ternate on larger branchlets, petiolate, leaf axils with dark brown, linear-digitate colleters 1−1.5 mm long secreting pale yellow resin; blades obovate-elliptic, 9.4−16.5 × 3.2−6.4 cm, base narrowly cuneate, attenuate, apex shortly acuminate, glabrous, discolorous, drying brown above, yellowish-brown below, when fresh green to yellow green above, pale green below, both surfaces glossy, secondary veins 15−20 on each side, secondary and tertiary veins visible above, prominulous beneath, margins conspicuously and tightly revolute; petioles 17−33 mm long, 1.7−2 mm in diam. Inflorescence terminal, 7−12 cm long, trichotomous, branching to the third degree, with 2 primary branches at apex of peduncle, each with 12−18 flowers, glabrous, axes with small, scale-like bracts; flowers on pedicels 1.5−4 mm long, 5-merous; calyx lobes obtuse to rounded, 2 × 2.5 mm; corolla in bud to 10 mm long; corolla at anthesis white, fragrant, corolla tube 6−7 × 3 mm, corolla lobes 5, contorted to the right, 6−7 × 2−25 mm, rounded at apex; ovary conical, 0.7−0.8 mm long, bicarpellate, style 0.8 mm long, stigmatic head ovoid, 0.7-0.8 mm long with tuft of hairs at apex and green ring (but no collar) at base, nectary 0.3 mm, 2-lobed, the lobes alternating with and partly covering the carpels. Infructescence with peduncle c. 17 cm long; fruits composed of two fleshy mericarps, when fresh orange at maturity, ellipsoid, 37−53 × 27−37 × 25−36 mm, mesocarp orange, 5 mm thick; endocarp externally fibrous, internally woody, 31−48 × 23−33 × 19−27 mm, the fibers to 1.5 mm in diam. Seed 2 per fruit, ellipsoid, 25 × 23−26 × c. 4 mm.
Nuku Hiva, Marquesas Islands, originally
known only from the type collected in the vicinity of the Toovii plateau
at about 900 m (
This species occurs in evergreen wet forest between 730 and 900 m elevation, with species of Hibiscus, Ixora, Metrosideros, Pandanus, Phyllanthus, Premna, Wikstroemia, Xylosma, and ferns including Asplenium australasicum (J. Sm.) Hook., Histiopteris incisa (Thunb.) J. Sm., and Microsorum grossum (Langsd. & Fisch.) Brownlie.
When evaluated using the IUCN criteria for endangerment (IUCN
Marquesas Islands: Nuku Hiva: Matahamo, Vaioa, fin de la piste forestière, 758 m, 2 Jan. 2003, Butaud 8 (PAP); Matahamo, Vaioa, 758 m, 9 Feb. 2003, Butaud 10 (PAP); Vaipupui, amont de la piste forestière menant à la dernière parcelle de pins, 737 m, 27 Jan. 2010, Butaud & Benne 2586 (PAP).
Since 2006, a conservation plan proposed by the Environment Direction of French Polynesia has been implemented, with conservation ex situ (seed collection and conservatory plantings) and in situ (fencing enclosure of the population and enrichment planting of seedlings inside).
A, B Ochrosia brownii. A inflorescence and open flower and buds showing right-contorted corolla lobes (Vaipupui, Nuku Hiva, Feb. 2003, Butaud unvouchered) B branchlet with apocarpous fruit composed of paired mericarps (Vaipupui, Nuku Hiva, Butaud 8); C, D Ochrosia fatuhivensis. C inflorescence with open flower and buds, Hanativa (Fatu Hiva, Nov. 2009, Butaud 2458) D ripe mericarps, seed showing fibers and germinating seed (Hanativa, Fatu Hiva, Nov. 2009, Butaud 2458).
http://species-id.net/wiki/Ochrosia_fatuhivensis
Figs 1C, DType. Marquesas Islands: Fatu Hiva: Oia [Ouia], c. 800 m, 19 Jan. 1922, F. B. H. Brown 886 (Holotype BISH-500706!).
Tree 10−14 m tall, trunk 15−20 cm in diam., branchlets glabrous, leafy twigs 2.5−7 mm in diam., bark wrinkled when dry, latex white. Leaves opposite on smaller branchlets, apparently ternate on larger branchlets; blades elliptic, 5.8−20.2 × 2.4−10.1 cm, base narrowly cuneate, apex shortly acuminate, glabrous, discolorous, when fresh dark green above, pale green below, drying brown, margins slightly revolute; secondary veins 9−20 on each side, prominulous on both surfaces, intersecondaries and tertiaries prominulous on both surfaces; petioles 14−36 mm long. Inflorescence terminal, tri- or quadrichotomous, branching twice, cymose-corymbiform, 52−88 mm long, axes and flowers glabrous, sessile, with 3 primary branches (6−)20−64 × c. 2 mm, subquadrangular, bracts ovate-triangular, 1−1.2 × 1−1.2 mm, ultimate axes with flowers crowded in cymules; flowers with pedicels 2−5 mm long, each with a single bract, calyx cup 2.5 mm long, calyx lobes 5, triangular-ovate, 2.5 × 3 mm, apex obtuse; corollas in bud to 10 mm long; corolla at anthesis white, fragrant, corolla tube 7−8 × 3 mm, corolla lobes 5, contorted to the right, 9−10 × 2−2.5 mm; ovary and nectary not seen. Infructescence with peduncle c. 11.5 cm long. Fruits apocarpous, composed of 2 fleshy mericarps, when fresh orange at maturity, ovoid-ellipsoid, 61−78 × 32−44 × 30−40 mm, mesocarp c. 7 mm thick; endocarp 54−72 × 27−40 × 23−34 mm, fibrous without, woody within, single-seeded. Seed 2 per fruit, ellipsoid, 29−37 × 18−19 × 3−4 mm.
Marquesas Islands, Fatu Hiva where originally known only from the type collected in Ouia Valley at 800 m.
This species was recently rediscovered in 2009 by Jean-François Butaud and Ravahere Taputuarai following the directions of Joseph Mititai. This population in Hanativa valley, a remote hanging valley north of Ouia, consists of 13 adult plants and numerous juveniles and seedlings at 480 m elevation. It occurs in mesic to wet forest with species of Aleurites, Allophylus, Artocarpus, Boehmeria, Cerbera, Inocarpus, Macropiper, Metrosideros, Musa troglodytarum L., Pandanus, Phyllanthus, Pipturus argenteus (G. Forst.) Wedd., Wikstroemia, the introduced Coffea arabica L. and Dioscorea spp., and the ferns Nephrolepis biserrata (Sw.) Schott, Pteris comans G. Forst., and Tectaria jardinii (Mett. ex Kuhn) E. D. Br.
Ochrosia fatuhivensis is currently known only from a single population. When evaluated using the IUCN criteria for endangerment it falls into the Critically Endangered (CR) category, which designates species facing the highest risk of extinction in the wild. IUCN Red List Category: Critically Endangered (CR) B1a, b; B2a, B2b (i–iii): B1, extent of occurrence estimated to be less than 100 km2, and B1a, known to exist at only a single location; B1b (i-iii), continuing projected decline in (i) extent of occurrence, (ii) area of occupancy and (iii), area, extent and quality of habitat; B2, area of occupancy estimated to be less than 10 km2, and B2a, a single population known. B2b (i–iii), habitat continuing decline inferred. The suitable habitat for Ochrosia fatuhivensis on Fatu Hiva (c. 85 km2) is indicated as an endangered environment, threatened by human activity (deforestation and fire), feral animals, and invasive plants, reducing the extent of the forest.
Marquesas Islands: Fatu Hiva: Hanativa, Affluent Sud de la vallée, vallon perché, 481 m, 7 Novembre 2009, Butaud, Taputuarai & Mititai 2458 (PAP).
Fosberg & Sachet (1972) suggested Ochrosia fatuhivensis may be related to Ochrosia compta
K. Schum. of Hawaii, but no molecular-phylogenetic studies have been
carried out involving these two species. The local name is ho’ei in Fatu
Hiva, similar to the name holei used for species of Ochrosia in Hawaii (
A single species of Rauvolfia, Rauvolfia sachetiae Fosberg was described from Hiva Oa, Marquesas based on a single collection (
A recent Nuku Hiva collection, Perlman 10014 distributed under the name Rauvolfia sachetiae, compares very well with Brown 432, the type of Ochrosia nukuhivensis, and clearly represents the same species. Furthermore, Perlman 10014 has subglobose or globose, syncarpous fruits characteristic of many Pacific species of Rauvolfia
in which normally each of the two carpels contains two ovules, but
frequently one aborts so that at maturity the fruit is usually 2-seeded,
but sometimes one carpel does not fully develop, resulting in a
1-seeded syncarpous drupe. In Ochrosia the fruits consist of a pair of well separated, distinct or bally basally connate oblong mericarps (
1a | Leaves narrowly ovate to narrowly elliptic or oblong, L:W ratio 2.9:1 to 3:1, secondary veins 21-25 on each side; calyx total width distally 3.2-4 mm; calyx lobes 1.5-2 × 1.5-2 mm | Rauvolfia nukuhivensis |
1b | Leaves ovate to elliptic, L:W ratio 1.7:1 to 2:1, secondary veins 18-19(-21) on each side; calyx total width distally, 2-2.5 mm; calyx lobes 1-1.2 × 1-1.2 mm | Rauvolfia sachetiae |
urn:lsid:ipni.org:names:77112735-1
http://species-id.net/wiki/Rauvolfia_nukuhivensis
Figs 2A, B, CMarquesas Islands: Nuku Hiva: without precise locality, ca. 1000 m, 20 June 1921, F. B. H. Brown 432(Holotype BISH-500695!).
Tree 8−15 m tall, trunk 40−50 cm in diam., bark furrowed, pale orange to beige in color, leafy branchlets 3−3.5 mm in diam., glabrous, bark wrinkled, brown, leafless branchlets 6 mm in diam., gray-brown, latex white. Leaves ternate, leaf axils and adjacent petiole bases with brown, glandular digitate colleters 1−2 mm long; blades narrowly ovate to narrowly elliptic or oblong-elliptic, 6−17.6 × 1.5−6.2 cm, base narrowly cuneate to cuneate, apex acute, chartaceous, glabrous, when fresh pale green, discolorous, margin slightly revolute, secondary veins 21−25 on each side, intersecondary and tertiary veins prominulous on both surfaces, petioles 12−36 × 1.2−1.5 mm, adaxially flattened. Inflorescences terminal, displaced by terminal shoot growth, 5−7 × 7−10 cm, including the verticellate peduncles 1.7−5.4 × 2 mm, branchlets slightly zigzag, glabrous, branching three to four times, ultimate branches trichotomous, bracteoles 1.5 mm long, triangular, acute; calyx total width distally 3.2−4 mm, calyx lobes 5, broadly triangular-ovate, 1.5-2 ×1.5-2 mm, obtuse at apex; corolla cream color when fresh, tube 8−13 mm long, lobes 2−3.5 × 2−2.5 mm, slightly contorted to the left, anthers 1.2-1.3 mm long, narrowly triangular-subulate, sagittate at base, ovary conical, 1.5 mm long, nectary disc annular, 0.4-0.5 mm long, style 5 mm long, stigma head cylindric, 0.4 mm long, with membranous collar at base. Fruit subglobose when mature, 12−15 mm in diam., fleshy, turning black at maturity, drupaceous with 1−3 seeds. Seeds compressed, obliquely oblong-ovoid, with one straight and one curved edge, base sub-truncate, apex diagonal from straight edge upward to a blunt point, sides coarsely and shallowly rugose.
Marquesas Islands, known only from Nuku Hiva between 198 and 627 m elevation.
Rauvolfia nukuhivensisis very rare and localized with fewer than 50 living trees currently known in valley or plateau dry forest with species of Cerbera, Ficus, Maytenus, Phyllanthus, Sapindus, and Xylosma. Currently known in the Terre-Deserte region in the valleys of Motuee, Hakaavao, Haahopu, Haatuatua, Hakaoa, Tapueahu and Uea, and on the plateaus of Tohuahee, Vaiteheii, Maauu, and Putata. As many dead trees as living trees are known. Formerly known from the north coast of Nuku Hiva, between the Haataivea peninsula and behind Aakapa village.
The suitable habitat for Rauvolfia nukuhivensis on Nuku Hiva (c.
340 km2) is indicated as an endangered environment, threatened by
human activity (deforestation, exploitation for leaves and bark),
feral animals, and invasive plants, reducing the extent of the forest.
Following the criteria and categories of IUCN
Marquesas Islands: Nuku Hiva: Tapueahu (labels erroneously says Matatekouaehi) Valley, 650 ft. (198 m), 1 July 1988, Perlman 10014 (AD, BISH, K, MO, MU, P, PAP, PTBG, US); Terre Déserte, branche gauche de la moyenne vallée de la Baie Marquisienne, face au Keiaki, 340 m, 30 July 1987, Florence 8434 (BISH, PAP, US); broad interfluve above Uea Valley, near Baie Marquisienne, 20 Apr. 1964, Decker 2046 (US); Ivipuovoteahi, NO de Keiaki, 400 m, 8°55’ S, 140°12’ W, February 1987, Jourdan 128 (PAP); Terre Deserte, Pipiheihe, 400 m, 8 April 1989, Toutain 4275 (PAP); Plateau de Maauu, au Sud du deuxième vallon de la forêt naturelle, 456 m, 9 April 2004, Butaud 418 (PAP).
Collectors’ notes indicate the leaves and bark are used for traditional medicine, another factor likely contributing to the decline of this species. Since 2006, a conservation plan lead by the Environment Direction of French Polynesia was implemented, with conservation ex situ (seed collection and conservation plantings) and in situ (enclosure of several populations). Local name: tueiao
A, B, C, Rauvolfia nukuhivensis A branchlet with leaves and inflorescence with open flowers, and buds with left-contort corolla lobes (Terre-Deserte, Nuku Hiva, May 2007, Butaud (unvouchered) B infructescence with syncarpous green fruits (Maauu, Nuku Hiva, April 2004, Butaud 418) C inflorescence with buds and open flower (Keiaki, Nuku Hiva, 27 Dec. 2002, J.-F. Butaud unvouchered); D Rauvolfia sachetiae, portion of isotype specimen showing leaves and inflorescence (Hiva Oa, above Taaoa, 15 Jan. 1975, Sachet 2115, Isotype BISH-578658).
http://species-id.net/wiki/Rauvolfia_sachetiae
Fig. 2DMarquesas Islands: Hiva Oa: dry crest above Taaoa, SW of village, 350 m, M.-H. Sachet 2115 (Holotype US!; Isotype BISH-598409!).
Tree 6−10 m tall, trunk 20−25 cm in
diam., bark furrowed, branches more or less horizontal, twigs and
leaves in whorls of 3−4, entire plant glabrous, latex abundant;
branchlets 4−6 mm in diam., proximal internodes to 9 cm long, distal
ones 0.5−1 cm long toward end of season’s growth, terminal buds with
abundant clear, brown resin. Leaves elliptic to broadly
elliptic, 5.5−22 × 2.5−6.8 cm apex acute to obtuse, base acute to
occasionally obtuse, slightly decurrent on petiole, glabrous,
chartaceous, light to bright yellowish green when dried, secondary
veins 12−15 on a side, brochidodromous, forming a wide angle with
costa, secondary and tertiary veins prominulous on both surfaces;
petioles 15−35 × 1−2 mm in diam. with 20−30 brown digitate, glandular
colleters 0.5−1.2 mm long in each leaf axil. Inflorescences
terminal, pseudoaxillary by displacement, compound cymes 3−6 × 8−12.5
cm including the corollas, peduncles verticellate, 1.5−3 cm long, 3−6
times ramified, branches open, widely divergent, ultimate branches
trichotomous with central branch usually represented by a single
pedicellate flower, most flowers early caducous from all but the most
distal branches, each lateral branch subtended by a scale-like,
ovate-triangular bract 1−1.5 × 1−1.5 mm, a bract pair subtending each
lateral bud, axils of bracts glandular as those of leaves, pedicel of
central flower 2−2.5 mm long, lateral buds of a triad subsessile,
globose, tending to develop into a new triad with only the central
flower of a triad developing to anthesis; calyx turbinate-campanulate,
lobed almost to base, lobes 1.5−2 × 1.5−1.7 mm, imbricate, obtuse,
margins thin; corolla yellow in bud, at anthesis cream colored, 12−14
mm long including limb, tube 9.5−14 × 1−1.5 mm in diam. medially,
slightly dilated at apex and base, lobes slightly left-contorted,
ovate, 2.5−3.5 × 2 mm, obtuse or rounded at apex, anthers narrowly
ovate, somewhat bluntly acuminate, sagittate at base, subsessile,
1.5 mm long, inserted 1.5 mm below throat, a very few hairs around and
below insertion on inside surface of corolla tube; ovary surrounded by
cup-like nectary disk with minutely crenulate margin, carpels oblong,
free distally, united in lower half, style glabrous, 7−9 mm long,
style head thickened, cylindrical (not seen: floral description from
Marquesas Islands, Hiva Oa, previously known only from a single tree at the type locality above Taaoa. In 2011 one dead trunk probably representing this species was found in a small gulch above Tanaeka valley, at around 430 m elevation (Butaud, pers. obs.).
Dry secondary forest or shrubland with Casuarina.
Despite careful searches this species has not been found since the last collection was made at the type locality in 1977. In 2010 the area was observed to be overgrazed by goats and invaded by Syzygium cumini.(L.) Skeels (Butaud, pers. obs.). Rauvolfia sachetiaefalls into the Critically Endangered (CR) category, which designates species facing the highest risk of extinction in the wild. IUCN Red List Category: Critically Endangered (CR) B1a, b; B2a, B2b (i–iii): B1, extent of occurrence estimated to be less than 100 km2, and B1a, known to exist at only a single location; B1b (i-iii), continuing projected decline in (i) extent of occurrence, (ii) area of occupancy, and (iii) area, extent and quality of habitat; B2, area of occupancy estimated to be less than 10 km2, and B2a, a single population known. B2b (i–iii), continuing habitat decline inferred. The suitable habitat for Rauvolfia sachetiae on Hiva Oa (c. 315 km2) is indicated as an endangered environment, threatened by human activity (deforestation and fire), feral animals, and invasive plants, reducing the extent of the forest.
Marquesas Islands: Hiva Oa: Ridge above Taaoa, SW of village, 350 m, 20 Nov 1974, Sachet & Decker 1885 (BISH, CBG, CHR, L, MO, NSW, P, PTBG, US), Taaoa, sur la presqu’île, versant SW, 350 m, 8 March 1977, Schäfer & Oliver 5293 (BISH, CBG, CHR, MO, NSW, PTBG, US).
We thank the French Polynesian Direction de l’Environnement for funding the surveys in the Marquesas Islands and also the conservation plan for two of the species. We are grateful to the Musée de Tahiti et des Iles (PAP), Bernice P. Bishop Museum (BISH), and Smithsonian Institution (US) for access to herbarium specimens. Napua Harbottle (BISH) kindly provided digital images of the types and Mel Gabel composed the color plates. Special thanks are extended to Joseph Mititai, Roberto Maraetaata and Ravahere Taputuarai on Fatu Hiva for assistance in locating the single population of Ochrosia fatuhivensis, and to Frédéric Benne and Frédéric Jacq on Nuku Hiva for work on Rauvolfia nukuhivensis and Ochrosia nukuhivensis. We thank Mary Endress and Warren Wagner for helpful discussions regarding Apocynaceae systematics and comments on the manuscript, and Mary Endress and David Middleton for reviewing the manuscript.