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Kadua lichtlei Lorence & W. L. Wagner, sp. nov., a new species from Ua Huka, Marquesas Islands, is described and illustrated. This new species differs from the three other Marquesan Kadua species by its broadly elliptic to broadly ovate or subcircular leaf blades as well as much smaller and more numerous (80-300) flowers and smaller capsules. Known from a single population of about 30 individuals, we conclude this new species should be added to the IUCN Red List as critically endangered (CR). A key, amended descriptions, conservation status, and specimen citations for the three previously described Marquesan species, Kadua lucei, K. nukuhivensis, and K. tahuatensis are also provided.
Conservation, French Polynesia, Kadua, Marquesas Islands, Rubiaceae
Until recently the Marquesas Islands were relatively
poorly explored botanically. Prior to initiation of the Flore de la
Polynésie française project under the auspices of Jacques Florence at
IRD (formerly ORSTOM) no species of Rubiaceae belonging to tribe Spermacoceae were known from these islands (
Generic delimitations in Hedyotis L. and related genera in Hedyotidinae have not been fully resolved, although recent morphological and molecular studies suggest that Hedyotis
s. str. is an Old World genus ranging from southeastern Asia to the
Caroline Islands of Micronesia and with two widespread species ranging
into western Polynesia (
Based on their salverform, fleshy corollas with
appendaged lobes and non-diplophragmous capsules (i.e., that do not
separate into paired cocci after dehiscing), the three Marquesan
species formerly placed in thegenus Hedyotis (Florence & Lorence 2000) have been transferred to the genus Kadua by
Results of an unpublished molecular analysis (
All measurements given herein are taken from dried
herbarium specimens, although certain features such as shapes were
supplemented with information from alcohol-preserved flowers and fruits,
field notes, and digital photos. Measurements are presented in the
descriptions as follows: length × width, 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, BR, K, MO, NY, P, PAP, PTBG, and US. The area of
occupancy (distribution) for this species was calculated using herbarium
collection data and field observations, and the conservation status is
proposed following the IUCN Red List Category criteria (
1a | Inflorescences with 80–300 flowers; corolla tube 1.8–2.2 mm long; capsules 4–5 mm long, 3.5–4 mm wide; seeds 0.5–0.6 mm long, 0.35–0.4 mm wide, irregularly ovoid to ellipsoid | Kadua lichtlei |
1b | Inflorescences with 12–30 flowers; corolla tube 13–28 mm long; capsules 7–22 mm long, 6–12 mm wide; seeds 0.9-1.3 mm long, irregularly trigonous or angulate (unknown inKadua lucei) | 2 |
2a | Inflorescences 3–6 cm long; corolla tube 13–16 mm long; Tahuata | Kadua tahuatensis |
2b | Inflorescences 8–13 cm long; corolla tube 22–28 mm long; Nuku Hiva; Fatu Hiva | 3 |
3a | Corolla lobes 8–10 mm long; capsules 7–8 mm long, 6 mm wide; Fatu Hiva | Kadua lucei |
3b | Corolla lobes 10–15 mm long; capsules 15–22 mm long, 7–12 mm wide; Nuku Hiva | Kadua nukuhivensis |
urn:lsid:ipni.org:names:77112739-1
http://species-id.net/wiki/Kadua_lichtlei
Figs. 1, 2A, B, 3Differt a congeneribus Marquesanis laminis late ovatis vel late ellipticis vel rotundis (3–) 5–17.5 × (1.8–) 3.5–11.5 cm, inflorescentia 6.5–14 × 9–12 cm, floribus in inflorescentibus 80–300, parvis, hypanthio 1–1.8 mm longo, corollae tubo 1.8–2.2 mm longo, corollae lobis 1.5–2 mm longis, et capsulis minoribus 4–5 × 3.5–4 mm.
Marquesas Islands: Ua Huka: Hane/Hokatu cliff zone, 520m elevation, 14 December 2003, K.R Wood and J.-Y. Meyer 10554 (holotype PTBG-44091!; Isotypes AD!, BISH!, BR!, K!, MO!, NY!, P!, PAP!, US!).
Shrub or small tree reaching 4 m tall, glabrous except for inflorescence, moderately branched, branches diffuse or often decumbent, twigs 3–3.5 mm in diam., internodes compressed, bark smooth to striate, dark brown. Leavesopposite, those of a pair at a node equal or sometimes unequal, blade (3–) 5–17.5 × (1.8–) 3.5–11.5 cm, broadlyovate, broadly elliptic, broadly obovate-elliptic or subcircular, chartaceous, glabrous, when fresh glossy dark green above, light green beneath, costa greenish white, margins entire, base acute to obtuse or rounded, shortly decurrent, apex obtuse or rounded, tip abruptly short acuminate, 0.5–1.5 cm , secondary veins 6–9(–11) per side, festooned brochidodromous, venation prominulous and conspicuous to 3° adaxially and to 4° abaxially; petioles (0.5–)1–2 cm long, winged distally; stipules interpetiolar (occasionally also intrapetiolar), fused with adaxial petiole bases, the body forming a short, broadly triangular sheath 1.5–3 × 3–7 mm, apiculate or with a short lateral ridge, glabrous, persistent. Inflorescences terminal, 6.5–14 × 9–12 cm (including the corollas), cymose-paniculate or -corymbiform, trichotomous, 80–300-flowered, sessile or sometimes with peduncle 3–4 cm long, the basal primary branches subtended by a pair of short petiolate, ovate foliaceous bracts 2–3.5 cm long, branching to the 4° (–5°), axes and pedicels minutely papillose-puberulent, subtended by brown triangular acuminate bracts 0.5–2 × 0.5–1 mm. Flowers hermaphroditic, apparently monomorphic and protandrous, borne in dichasial cymules on ultimate branches, central flower often sessile, lateral ones on pedicels to 4 mm long, hypanthium green when fresh, 1–1.8 × 1.2–1.6 mm, broadly obovoid or obconical, tuberculate, calyx tube 0.2–0.4 mm long, glabrous externally and internally, calyx lobes 4 (–5), 0.2–0.5 mm long, triangular, glabrous;corolla in bud 4-angular, apex obtuse, slightly or not depressed, at anthesis shortly hypocrateriform, when fresh with white lobes and green tube, tube 1.8–2.2 × 1–1.3 mm medially, externally and internally glabrous, lobes 4, triangular-ovate, 1.5–2 × 1.3–1.6 mm, apex with a small incurved appendage, externally papillose, internally rugulose; anthers 4, 0.7–1.4 mm long, ellipsoid, apiculate, bilobed basally, sessile, attached below top of tube, tips exserted; style 1.5 mm long, stigma lobes 0.5 mm long, included in tube. Capsules 2/3 inferior, broadly obovoid to subglobose, 4–5 × 3.5–4 mm in diam., apex (beak portion above the calyx) rounded to conical, glabrous, dark brown when fresh, vascular bundles becoming visible with age, loculicidal, apex splitting into 4 segments. Seeds c. 200, dull brown, 0.5–0.6 × 0.35–0.4 mm, irregularly ovoid to ellipsoid, laterally compressed, laterally cuneate with a marginal punctiform hilum, the testa irregularly reticulate with areoles enclosing granulate-verrucose mounds.
This new species is known only from Ua Huka, Marquesas Islands, where it is apparently restricted to the Hokatu cliff zone above Hane village.
Rare and localized, this new species occurs in mixed wet shrubland and herbland growing on basalt cliffs and rock outcrops above wet forest of Hibiscus tiliaceus L., Pandanus tectorius Parkinson, and Freycinetia impavida (Gaudich. ex Hombr.) B.C. Stone. Other associates include species of Bidens, Boehmeria, Maytenus, Peperomia, Alyxia, Morinda, Phyllanthus, Miscanthus, Macropiper, Xylosma, and diverse pteridophytes. It was collected in flower in December and June (in bud), and in fruit in June and December (old fruit with a few seeds).
We are pleased to name this new species in honor of Mr. Léon Lichtle, Mayor of Ua Huka, for his generous help and logistic support when we conducted field work on the island and also in recognition of his strong support for conserving the island's native flora and fauna.
The area of occupancy (distribution) for the
species was calculated using herbarium collection data and field
observations, and the conservation status is proposed following the
IUCN Red List Category criteria (
Marquesas Islands: Ua Huka: Hane valley, ridge and cliffs above tiki marae, back of valley on west side, 8°54.48'S, 138 °3.54'W, 518 m, 12 Jun 2004, Perlman et al. 19007 (BISH, P, PAP, PTBG, US); Hane/Hokatu cliff zone, 520m, 11 December 2003, Wood & Meyer 10514 (BISH, P, PAP, PTBG, US), 550m, 14 December 2003, Wood & Meyer 10544 (PTBG, US), 10547 (PTBG, US), 12 June 2004, 8°54.72'S, 139 °31.60'W, 520 m, Wood et al. 10737 (PTBG), 19738 (PTBG, US).
Hawaiian Islands: Kaua`i: Koloa District, National Tropical Botanical Garden, horticulture center nursery, 5 October 2005, Lorence 9476 (PTBG, US).
Within the Marquesan clade Kadua lichtlei
differs markedly from the other three species by a number of
characters, notably its broadly elliptic to broadly ovate or
subcircular leaf blades, much smaller and more numerous flowers (80-300
per inflorescence), and smaller capsules. In addition, seeds of Kadua nukuhivensis and Kadua tahuatensis (those of Kadua lucei
were not available) are more sharply angulate with well defined
marginal ridges. Seed morphology has proven useful in the infrageneric
classification of Kadua (
Kadua lichtlei Lorence & WL Wagner A habit, fruiting branch B inflorescence C node showing stipule and petiole bases D, E flowers in bud and at anthesis F mature capsule, dehisced G seeds, lateral and dorsal (center) views. A, F, G. based on the type collection Wood & Meyer 10554; B, C, D, E based on Lorence 9476.
Seeds of three Marquesan Kadua species, whole seed and surface detail A, B Kadua lichtlei, Wood 10514 (PTBG) C, D Kadua nukuhivensis C Perlman 15029 (PTBG) D Perlman 15054 (PTBG) E, F Kadua tahuatensis Perlman 16020 (PTBG).
http://species-id.net/wiki/Kadua_lucei
Marquesas Islands: Fatu Hiva: au pied de la crête à Touaouoho, env. 1000 m, 3 Aôut1999, J.-P. Luce s.n. (Holotype P!)
Glabrous shrubs 1.5–2 m tall, leafy twigs 4–6 mm in diam., internodes compressed, bark brown, smooth to striate or rugulose. Leaves of a pair equal to subequal, blade elliptic to obovate-elliptic, (5.3–)10–14.5 × (3.3–)7–10 cm, discolorous, chartaceous to subcoriaceous, base acute to obtuse or rounded, apex acute to obtuse or rounded, often abruptly short acuminate, margin entire, weakly revolute; secondary veins 8–10 pairs, loosely brochidodromous, secondary veins prominulous on both surfaces, tertiary veins slightly prominulous below; petioles purple when fresh, 2–4 × 2–5 mm, stout, adaxially flattened and slightly winged; stipules inter- and intrapetiolar, apex shortly cuspidate, sheath semicircular, 3 × 4–7 mm, fused with the adaxial petiole bases. Inflorescences(only seen with old flowers and young fruits) terminal, corymbiform cymose, 8–9 × 6 cm, peduncle 35 mm long, 2.5–3 mm in diameter, compressed, primary axis 2.5–3 cm × 2–2.5 mm with two pair of secondary branches, the lower pair often branching once again, the upper pair unbranched, the ultimate branches each bearing 2–3 flowers, lower bracts foliaceous, suborbicular, 15 mm in diam. Flowers glabrous, about 30 per inflorescence, bracts broadly triangular, scarious, 1 × 1.5 mm, pedicels stout, 2–3 × 0.5–1 mm; hypanthium obconical, 2–3 mm long, calyx limb cupuliform, 2–3 mm long, calyx lobes 4, broadly triangular, 0.5–0.7 × 1.5 mm; corolla fleshy, white when fresh, buds not seen, at anthesis salverform, tube purple tinged when fresh, 22–26 mm long, 1.7–2 mm wide medially, lobes 4, linear to oblong, recurved, 8–10 × 1.5–2 mm, apex with a hook-like appendage 0.5–1 mm long; flowers possibly dimorphic; stamens included, anthers linear, 3–3.5 × 0.3–0.4 mm, connective attached 2 mm from apex of tube; style included, 20 mm long, two stigmatic lobes linear, 2.5 mm long. Fruits on stout pedicels 5–7 mm long, capsules turbinate to obovoid-turbinate, old capsules 7–8 × 6 mm, consisting of network of persistent vascular bundles enclosing endocarp, apex (beak portion above the calyx) 1–1.5 mm long. Seeds not seen.
Marquesas Islands, Fatu Hiva where known only from a single small population on the summit ridge between Tekou and Touaouoho peaks.
This new species was collected at 915–1000 m elevation on a steep, precipitous ridge crest in wet shrubland with species of Alsophila, Freycinetia, and Histiopteris.
The specific epithet honors its discoverer and first collector, Mr. Jean-Pierre Luce, an amateur naturalist, in recognition of his efforts to explore the most rugged mountainous zones of the Marquesas and thus increase our knowledge of their flora and vegetation.
The suitable habitat for Kadua lucei
on Fatu Hiva (ca. 85 km2) is indicated as an endangered environment,
threatened by feral animals and invasive plants, reducing the extent of
the forest. Estimated population size is about 3–4 individuals.
Following the criteria and categories of
Marquesas Islands: Fatu Hiva: sous la crête entre Tekou et Touaouoho, 915 m, 15 février 2000 (st), J.-Y. Meyer & J.-P. Luce 835 (PAP, PTBG).
This species is apparently related to Kadua tahuatensis from which it differs by its larger inflorescence reaching 8–9 cm long and 6 cm wide, more numerous flowers, about 30 per inflorescence, and larger white corollas with a longer purple-tinged tube 22–26 mm long. Although mature fruits of Kadua lucei are not known, old fruits are smaller than those of Kadua nukuhivensis and Kadua tahuatensis.
http://species-id.net/wiki/Kadua_nukuhivensis
Marquesas Islands: Nuku Hiva: route Toovii–Terre Déserte, 5 km après le col, 8°52'S, 140°10'W, 1020 m, 5 juin 1984, J. Florence 6914 (Holotype P!; Isotypes BISH!, P!, PAP!, PTBG!, US!).
Glabrous shrubs or small trees 2.5–6 m tall, trunk to 20 cm in diam., leafy twigs cylindrical, 7–9 mm in diam., internodes compressed, bark grayish- to blackish-brown. Leaves of a pair equal to sometimes unequal, blade obovate to obovate elliptic or oblong, 8–18 × 6–10.5 cm, chartaceous to subcoriaceous, drying brown or blackish-brown, base cuneate, decurrent, apex obtuse to rounded, secondary veins 7–9 pairs, loosely brochidodromous, prominulous above, tertiary veins slightly prominulous, higher order venation obscure, midrib sulcate above, rounded below, margin entire, plane to slightly revolute; petioles 8–13 × 2–3 mm in diam., sulcate; stipules inter- and intrapetiolar, sheath truncate, cupuliform, 2.5–3 × 5–6 mm, caducous but becoming thickened and horseshoe-like on adaxial surface of petioles. Inflorescences terminal and rarely also axillary in upper leaf axils, cymose, corymbiform, 7–13 cm long including corollas, on a stout, compressed peduncle 3–6 cm long or sessile with a pair of basal branches, often subtended by a pair of reduced subsessile leaves, bracts foliaceous, 1.5–3 cm long, caducous. Flowers 12–30, possibly dimorphic, fragrant when fresh, subtended by caducous triangular bracts c. 1 mm long, hypanthium obconical, 3 mm long, calyx limb cupuliform, 2 mm long, calyx lobes broadly triangular, 0.3–0.5 mm long; corolla fleshy, white or pinkish white when fresh, apex flat to slightly depressed in bud, at anthesis salverform, tube 26–28 × 2 mm in diam. distally, lobes 4, linear-oblong, recurved, 10–15 × 3–5 mm, apex with a recurved hook-like appendage; stamens with anther tips exserted for 1.5 mm, mucronulate, linear, 3–4 × 1 mm, attached 1.5–2 mm below apex of tube; style 17–19 mm long, included, stigmatic lobes 2, adnate, 3 mm long. Capsules on stout pedicels to 6 mm long, turbinate to obpyriform, 15–25 × 7–14 mm, strongly compressed, bisulcate, strongly 8-ribbed, 2/3 inferior, apex (beak portion above the calyx) obtuse, 5–6 mm long, smooth, dehiscence loculicidal then septicidal, old capsules disintegrating leaving network of vascular bundles enclosing the persistent endocarp. Seeds 1–1.3 × 0.5–0.9 mm, irregularly angulate to broadly ellipsoid, trigonous, margins thin, testa dull, dark brown, surface finely papillose.
Marquesas Islands, Nuku Hiva, where known only from the island's central mountain crest, on the leeward side of the Terre Déserte in the upper Tapuaehu Valley, between 1000 and 1065 m elevation.
This species occurs in wet forest with species of Hernandia, Ilex, Metrosideros, and Weinmannia in the canopy and the understory with species of Cyrtandra, Melicope, and Psychotria. Numerous pteridophytes occur terrestrially and as epiphytes. Kadua nukuhivensis also occurs in shrubland on ridge crests with species of Alsophila, Bidens, Dicranopteris, Elaphoglossum, Freycinetia, Pennisetum, and Styphelia.
The specific epithet refers to the only known island of occurrence for this species.
The suitable habitat for Kadua nukuhivensis on Nuku Hiva (c.
340 km2) is indicated as an endangered environment, threatened by
human activity (deforestation and fire), feral animals, and invasive
plants, thus reducing the extent of the forest. Based on the IUCN
criteria and categories this species is assigned a preliminary Red List
status of Endangered (EN): B1, B2b (i-iii): B1 extent of
occurrence <5, 000 km2; B2: total area of occupancy less than 500 km2
(c. 75 km2); B2b (i-iii), habitat continuing decline inferred in (i)
extent of occurrence, (ii) areas of occupancy, and area, (iii) extent
and/or quality of habitat. This status is a revision from VU originally
suggested by
Marquesas Islands: Nuku Hiva: Route Toovii—Terre Déserte, haute Tapuaehu, 8°52'S, 140°11'W, 1020 m, 14 fév 1986, Florence 7545 (BISH, P, PAP, US); off the old Airport road west of the summit crest of Peak #1227 m, drainages of Tapueahu Valley, 0.75 miles south of Airport Road, bottom of valley, 3340 ft., 24 sep 1995, Perlman15054 (AD, BISH, MO, P, PAP, PTBG (2), US); Ooumu area (sic), top of Tapueahu off new Hwy, 8°51'53"S, 140°10'63"W, 3500 ft, 23 jun 1997, Wood, Meyer & Luce 6337 (BISH, P, PAP, PTBG, US).
Kadua nukuhivensis resembles Kadua tahuatensis,
and molecular evidence places these two as sister species in the same
clade within the larger clade of Hawaiian and French Polynesian species (
Kadua lichtlei, flowering plant growing the National Tropical Botanical Garden nursery, 5 Oct. 2005, Lorence 9476 (PTBG).
http://species-id.net/wiki/Kadua_tahuatensis
Fig. 2E, F.Marquesas Islands: Tahuata: ridge between Amatea and Haaoiputeomo, south-facing slope, 2580 ft. [780 m] elevation, 19 july 1997, S. P. Perlman 16020 (Holotype: PTBG-30160!; Isotypes: BISH!, MO!, P!, PAP!, US!).
Glabrous shrubs to 2 m tall, leafy twigs 4–6 mm in diam., internodes strongly compressed, bark pale brown, smooth to striate. Leaves of a pair equal to subequal, blade elliptic to obovate-elliptic, 4.2–15 × 2–8 cm, discolorus, chartaceous to subcoriaceous, base acute to cuneate or narrowly cuneate, apex obtuse to rounded, tip sometimes abruptly short acuminate, secondary veins (5-) 6–9 pairs, weakly brochidodromous, secondary and tertiary veins prominulous on both surfaces, higher order venation obscure, margin thickened, plane; petiole stout, 2–5 × 2–3 mm, adaxially sulcate; stipules inter- and intrapetiolar, sheath cupuliform, truncate, 3 × 5–6 mm, persistent, fused with adaxial petiole surfaces and becoming thickened and horseshoe-like. Inflorescences terminal, thyrsiform cymose, 23–25-flowered, 5–6 × 4.5–6 cm (including corollas), on peduncle to 2 cm long, 1.5–2 mm in diam., flattened, primary axis 15 × 1.5–2 mm with 2 pairs of lateral branches, the basal one often branching once, lower bracts foliaceous, ovate, 1.2–2 × 1–1.5 cm, upper branch pair unbranched, ultimate branches ending in 2–3 flowers. Flowers glabrous, on stout pedicels 2–3 × 0.8–1.4, compressed, bracts scarious, ovate-trangular, 1 × 1 mm, hypanthium obconical, 3–4 × 1.5–2 mm, calyx limb campanulate, 2–3 × 4–5 mm, lobes ovate-triangular, 1.5-2 × 2-2.5 mm; corolla fleshy, pale green when fresh, lobes with dark purple margins, in bud fusiform with non-depressed apex, at anthesis salverform, tube 13–16 × 1.5–2 mm in diam. medially, lobes 4, linear-oblong, recurved, 8–10 × 1.5–2 mm, apex with a hooked appendage 1 mm long; flowers possibly dimorphic, stamens exserted for 1.5–2 mm, linear, 3-3.5 × 0.5–0.6 mm, attached 1–1.5 mm below apex of tube, apex slightly mucronulate; style included, 11–12 mm long including 2 coalescent stigmatic lobes 2.5 mm long. Fruits on stout pedicels 3–8 mm long; capsule tubinate to obovoid-turbinate, 12–20 × 6–8 mm, sub-quadrangular, 2/3 inferior, apex with short beak portion 5–6 mm long above the calyx, dehiscence at first loculicidal then septicidal, old capsules disintegrating into network of vascular bundles enclosing persistent, bisulcate endocarp. Seeds irregularly trigonous or angulate, 0.9–1.2 mm long and wide, margins with narrow wing 0.1-0.3 mm wide, testa finely reticulate.
The specific epithet refers to the only known island of occurrence for this species.
Marquesas Islands, Tahuata, known only from the island's summit crest and high southeastern slopes between 780 and 835 m elevation.
Occurs in wet montane forest and shrubland with species of Alsophila, Cheirodendron, Crossostylis, Hibiscus, Metrosideros, Reynoldsia, Weinmannia, and pteridophytes including Asplenium, Blechnum, Elaphoglossum, Lycopodiella and Nephrolepis,
The suitable habitat for Kadua tahuatensis on Tahuata (c.
61 km2) is indicated as an endangered environment, threatened by feral
animals and invasive plants, reducing the extent of the forest. This
species is extremely rare, with only five plants known from two
localities. Following the criteria and categories of
Marquesas Islands: Tahuata: ridge between Amatea and Haaoiputeomo, southeast-facing slopes over Hanatetena village, 2740 ft. (835 m) elevation, 11 July 1997, Perlman 15954 (P, PAP, PTBG, US).
Morphologically Kadua tahuatensis closely resembles Kadua nukuhivensis, and molecular evidence places these two as sister species in the same clade as Kadua rapensis F. Br. and Kadua romanzoffiensis Cham. & Schltdl. within the larger clade of Hawaiian and French Polynesian species (
This study was greatly facilitated by useful discussions, field observations, photos, and specimens provided by Dr. Jean-Yves Meyer (Délégation à la Recherche, Polynésie française), Ken Wood and Steve Perlman (National Tropical Botanical Garden), and Jacques Florence (IRD, Paris). Important observations and collections resulted from fieldwork carried out during trips in 2003 and 2004 as part of the “Vascular Flora of the Marquesas Islands” project, which was supported by a generous project donation. We are grateful to the Délégation à la Recherche (Papeete, Tahiti), the Musée de Tahiti et des Iles, Mr. Léon Lichtle and the Service du Développement Rural for logistic support on Ua Huka, 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 illustration was skillfully drawn by Alice Tangerini (US). We thank Marjorie Knowles and Harold Robinson of the Smithsonian Institution for preparing the SEM seed images. We are grateful to Harold Robinson and Charlotte Taylor for reviewing the manuscript. The contribution by WLW to this study was partially supported by the Smithsonian Research Opportunities Fund and while appointed as McBryde Chair at the National Tropical Botanical Garden.