Research Article |
Corresponding author: Kenneth R. Wood ( kwood@ntbg.org ) Academic editor: Pedro Acevedo-Rodríguez
© 2016 Kenneth R. Wood, Marc S. Appelhans, 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:
Wood KR, Appelhans MS, Wagner WL (2016) Melicope oppenheimeri, section Pelea (Rutaceae), a new species from West Maui, Hawaiian Islands: with notes on its ecology, conservation, and phylogenetic placement. PhytoKeys 69: 51-64. https://doi.org/10.3897/phytokeys.69.8844
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Melicope oppenheimeri K.R. Wood, Appelhans & W.L. Wagner (section Pelea (A. Gray) Hook. f., Rutaceae), a rare endemic tree from West Maui, Hawaiian Islands, is described and illustrated with notes on its ecology, conservation, and phylogenetic placement. The new species differs from Hawaiian congeners by its carpels basally connate 1/5, narrowed into a strongly reflexed beak 10–15 mm long. It also differs in a combination of leaves with 7–10 pair of secondary veins; cymes to 3 cm long; peduncles 5–6.5 mm long; flowers perfect; capsules 4–9 × 40–52 mm; and a densely appressed short-sericeous ovary. Melicope oppenheimeri is known only from an isolated cliff-base plateau in upper Waihe‘e Valley, West Maui. Its discovery brings the number of recognized Melicope J.R. Forst. & G. Forst. species in the Hawaiian Islands to 49. A table is included indicating the conservation status of Hawaiian Melicope and Platydesma H. Mann., which is nested within Melicope sect. Pelea.
Melicope oppenheimeri falls into the IUCN Critically Endangered (CR) Red List category.
Rutaceae , Melicope , M. section Pelea, new species, Hawaiian Islands, West Maui, Critically Endangered
Melicope J.R. Forst. & G. Forst. is the largest genus of the Citrus family (Rutaceae) and consists of ca. 235 species of shrubs and trees (
All measurements and descriptions were taken from dried herbarium specimens or from notes made in the field and are presented in the descriptions as follows: length × width, followed by units of measurement (mm or cm).
Differs from Hawaiian congeners by its combination of leaves having 7–10 pair of secondary veins; cymes to 3 cm long; peduncles 5–6.5 mm long; flowers perfect; carpels basally connate 1/5, narrowed into a strongly reflexed beak 10–15 mm long, capsules 4–9 × 40–52 mm; and ovary appressed densely short-sericeous.
United States of America. Hawaiian Islands, West Maui: Wailuku District, Waihe‘e Valley, Metrosideros polymorpha var. glaberrima-Cheirodendron trigynum wet forest, 20°54.15'N; 156°33.95'W, 770 m elev., 12 Sep 2006, Hank Oppenheimer & Jill Miller H90609 (holotype:
Trees 3–4 m tall, bark medium brown, young branchlets light brown, glabrate, 3–6 mm wide in third internode, terminal branchlets yellowish brown tomentose with a waxy scurf. Leaves opposite, unifoliolate, coriaceous, the blade obovate to broadly elliptic, occasionally orbicular, 5–17.5 × 3.5–8.5 cm, the margin entire, the base rounded to obtuse, the apex rounded, obtuse to acute, or emarginate, secondary veins usually 7–10 pairs, connected by a moderately arched vein 2–11 mm from margin with higher order venation reticulate, both surfaces glabrous, occasionally glabrate along midrib of abaxial surface, young leaves glabrate to sparsely puberulent on lower surface; petiole shallowly canaliculate, 10–30 × 1–3 mm at middle, glabrate. Flowers: perfect, 3–5 in axillary cymes up to 3 cm long, peduncles 5–6.5 mm long, sparsely short-puberulent, pedicles 5–8 mm, short-puberulent, bracteoles 1.5–3 mm long, sepals broadly ovate, tip short acuminate, externally sparsely short-puberulent, glabrous within, 3 × 2.5–2.8 mm, connate basally 1/4 to 1/3 of length; petals tinged purple, narrowly ovate, lanceolate, 6–8 × 2.5–3 mm, glabrous internally and externally, tips recurved, nectary disk with sparsely scattered hairs; ovary densely appressed short-sericeous; style ca. 1.7 mm long, with finely appressed hairs; stigma capitate, four lobed, glabrous; stamens 8, filaments glabrous, the antesepalous ones 5–7 mm long, antepetalous ones 4–5 mm long, all with pollen. Capsules purple tinged when fresh, 4–9 × 40–52 mm, carpels connate basally for ca. 1/5 their length, narrowed into a strongly reflexed beak 10–15 mm long, exocarp glabrate with few hairs widely spaced over surface, endocarp sparsely to evenly puberulent. Seeds 2 per carpel, ovoid, 5–9 mm long.
To date, Melicope oppenheimeri has been observed with flower buds in January and August, with flowers at anthesis during September, and with fruit during January, February, May, August, September, and November.
We are pleased to name this new species in honor of Hank Oppenheimer, botanist with the Maui Nui Plant Extinction Prevention Program, who collected the type specimen and has made many valuable contributions to the understanding and conservation of the Hawaiian flora.
United States. Hawaiian Islands, West Maui: Wailuku District, Waihe‘e Valley, south side, below and north of Kahoolewa Ridge, 20°54.15'N; 156°33.95'W, 770 m elev., 8 Aug 1998, Wood & Perlman 7408 (
The following couplets can be inserted into the existing key to Hawaiian Melicope (treated as Pelea) by Stone, Wagner, and Herbst (in
19(18) | Exocarp sparsely to densely puberulent or tomentose, at least toward base or along suture | 20 |
19 | Exocarp glabrous or glabrate, sometimes with a few hairs widely spaced over surface (49). | 49 |
49(19) | Endocarp densely and uniformly short-villous; K | M. cruciata |
49 | Endocarp glabrous or sparsely puberulent, especially along suture | 50 |
50(49) | Leaves ternate; O | M. lydgatei |
50 | Leaves opposite. | 51 |
51(50) | Most petioles 0–10 mm long | 52 |
51 | Most petioles over 10 mm long | 58 |
58(51) | Ovary sparsely to densely puberulent or tomentulose, but exocarp glabrate or nearly so in fruit | 59 |
58 | Ovary and exocarp glabrous | 60 |
59(58) | Flowers perfect, carpels narrowed into a reflexed beak 10–15 mm long; |
M. oppenheimeri |
59 | Flowers unisexual, carpels straight or somewhat reflexed, apex not beaked | 59a |
59a(59) | Capsules (16–)25–40(–50) mm wide, carpels connate 1/4–1/3 their length; pedicels 5–20 mm long; L, |
M. volcanica |
59a | Capsules 11–20 mm wide, carpels connate (1/3–)1/2 their length or more; pedicels 2–5 mm long; K, O | M. wawraeana |
Although seven trees of Melicope oppenheimeri have been documented since its discovery in 1998, only three trees are still surviving in the upper headwaters of Waihe‘e Valley, West Maui (Figures
Melicope oppenheimeri K.R. Wood, Appelhans & W.L. Wagner. A Flowering branch, Oppenheimer & Hansen H20505 (
The plant communities of upper Waihe‘e Valley are varied and merge together in and around the 0.25 km2 cliff-base plateau region where trees of Melicope oppenheimeri occur. The plateau itself represents a relictual low statured Metrosideros Banks ex Gaertn. (Myrtaceae)-Cheirodendron Nutt. ex Seem. (Araliaceae) lowland wet forest community. Other relic native components of the plateau include ferns such as Asplenium lobulatum Mett. (Aspleniaceae), Cibotium glaucum (Sm.) Hook. & Arn. (Cibotiaceae), Ctenitis latifrons (Brack.) Copel. (Dryopteridaceae), Diplazium sandwichianum (C. Presl) Diels (Athyriaceae), two endemic genera of ferns, namely Adenophorus Gaudich. (Polypodiaceae) and Sadleria Kaulf. (Blechnaceae), along with herbs, shrubs, and small trees including Antidesma platyphylla H. Mann (Phyllanthaceae), Broussaisia arguta Gaudich. (Hydrangeaceae), Coprosma pubens A. Gray (Rubiaceae), Dubautia plantaginea Gaudich. (Asteraceae), Peperomia hirtipetiola C. DC. (Piperaceae), and various species of Clermontia Gaudich., Cyanea Gaudich. (both Campanulaceae), and Cyrtandra J.R. Forst. & G. Forst. (Gesneriaceae). Steeper ridges and slopes that edge the plateau are dominated by matting ferns of Dicranopteris Bernh.(Gleicheniaceae) (
Unfortunately, past habitat degradation by introduced pigs (Sus scrofa L.) has altered the plant composition of the plateau, which is now being dominated by a succession of invasive weeds, which threaten Melicope oppenheimeri, especially Ageratina adenophora (Spreng.) R.M. King & H. Rob. (Asteraceae), Buddleja asiatica Lour. (Scrophulariaceae), Clidemia hirta (L.) D. Don (Melastomataceae), Erigeron karvinskianus DC. (Asteraceae), and Tibouchina herbacea (DC.) Cogn. (Melastomataceae) (
Towering over the plateau and most outstanding, are vertical caldera-like basalt cliffs dominated by a native sedge and grass association called Machaerina Vahl (Cyperaceae)-Deschampsia P. Beauv. (Poaceae) wet cliff community. The cliffs are seeping with springs and waterfalls and strata of past volcanic flows are clearly evident. Additional components of this Waihe‘e cliff community include Sadleria pallidaHook. & Arn., Pipturus albidus (Hook. & Arn.) A. Gray (Urticaceae), Eragrostis grandis Hillebr., Isachne distichophylla Munro ex Hillebr. (both Poaceae), and Dubautia scabra (DC.) D.D. Keck.
Superb examples of Metrosideros-Cheirodendron montane wet forest dominate above these cliffs in association with windswept wet shrublands and occasional bog communities in and around the locality known as Kaho‘olewa Ridge (
The forests spreading below the plateau are composed of Metrosideros mixed lowland riparian vegetation, which are closed to open in canopy and dissected by deeply carved streams with steep banks 10–30 m in height. The native flora of this lower riparian community has similarities to the cliff-base plateau but with a greater diversity, including Ilex anomala Hook. & Arn. (Aquifoliaceae), Perrottetia sandwicensis A. Gray (Dipentodontaceae), Polyscias oahuensis (A. Gray) Lowry & G.M. Plunkett (Araliaceae), and Pritchardia forbesiana Rock (Arecaceae), along with species of Kadua Cham. & Schltdl. (Rubiaceae), Myrsine L. (Primulaceae), and Psychotria L. (Rubiaceae) (
Melicope oppenheimeri, like all Hawaiian Melicope, falls into section Pelea, which has a distribution ranging from Taiwan, the Philippines, and Borneo eastward to the Hawaiian and Marquesas Islands, and south to New Caledonia. Only Melicope triphylla (Lam.) Merr. is distributed in the Southeast Asian areas mentioned above, and the remainder of sect. Pelea is restricted to New Guinea and Pacific archipelagos (
Although beaked fruit have evolved in a few species of Melicope belonging to sections Melicope and Vitiflorae (
Morphologically, Melicope oppenheimeri resembles the Moloka‘i endemic species M. reflexa, with both having reflexed carpels that are coherent at base. Significant differences between M. oppenheimeri and M. reflexa include capsules connate 1/5, 4–9 × 40–52 mm (vs. capsules connate 1/4, 10–17 × 20–33 mm); endocarp sparsely to evenly puberulent (vs. endocarp glabrous); ovary appressed densely short-sericeous (vs. ovary glabrous); pedicles 5–8 mm (vs. pedicles15–20 mm); and flowers perfect, 3–5, on robust peduncles (vs. flowers unisexual, 1–3, on delicate peduncles).
Two other morphologically similar Hawaiian Melicope, also with carpels coherent at base include M. molokaiense (Hillebr.) T.G. Hartley & B.C. Stone and M. volcanica (A. Gray) T.G. Hartley & B.C. Stone. Melicope molokaiense, which is known from Molokai and Maui, differs from M. oppenheimeri in having capsules connate 1/4, 10–17 × 21–39 mm; endocarp glabrous; ovary glabrous; and flowers unisexual. Melicope volcanica, which is known from Lana‘i, Moloka‘i, East Maui, and the big island of Hawai‘i, similarly differs with capsules connate 1/3–1/4, and flowers unisexual, yet also differs with leaves having 10–20 pair of secondary veins; cymes ca. 6 cm long; and peduncles 7–38 mm long. Melicope oppenheimeri has leaves with 7–10 pair of secondary veins; cymes ca. 3 cm long; and peduncles 5–6.5 mm long.
Melicope species are generally either exclusively hermaphroditic or dioecious, yet there are a few recorded exceptions (
Plant and animal endemics from isolated oceanic islands are often endangered or critically endangered (
Checklist of endemic Hawaiian Melicope and Platydesma with conservation status and island distribution. (Status Symbols: C=candidate for federal listing; E=federally listed as endangered; EX=possibly extinct; PEPP=Plant Extinction Prevention Program (50 or less individuals known in wild); SOC =species of concern. Island Distribution: K=Kaua‘i; O=O‘ahu; Mo=Moloka‘i; L=Lana‘i; M=Maui; H=Big Island of Hawai‘i; Note: Platydesma had been shown to be nested in Melicope [
Taxon | Status | Distribution |
---|---|---|
Melicope adscendens (H.St.John & E.P.Hume) T.G.Hartley & B.C.Stone | E, PEPP | M |
Melicope anisata (H.Mann) T.G.Hartley & B.C.Stone | K | |
Melicope balloui (Rock) T.G.Hartley & B.C.Stone | E, EX | M |
Melicope barbigera A.Gray | K | |
Melicope christophersenii (H.St.John) T.G.Hartley & B.C.Stone | E | O |
Melicope cinerea A.Gray | SOC | O |
Melicope clusiifolia (A.Gray) T.G.Hartley & B.C.Stone | K, O, Mo, L, M, H | |
Melicope cruciata (A.Heller) T.G.Hartley & B.C.Stone | SOC | K |
Melicope degeneri (B.C.Stone) T.G.Hartley & B.C.Stone | E, PEPP | K |
Melicope elliptica (A.Gray) T.G.Hartley & B.C.Stone | O, Mo, M | |
Melicope feddei (H.Lév.) T.G.Hartley & B.C.Stone | K | |
Melicope haleakalae (B.C.Stone) T.G.Hartley & B.C.Stone | SOC | M |
Melicope haupuensis (H.St.John) T.G.Hartley & B.C.Stone | E, PEPP | K |
Melicope hawaiensis (Wawra) T.G.Hartley & B.C.Stone | SOC | Mo, L, M, H |
Melicope hiiakae (B.C.Stone) T.G.Hartley & B.C.Stone | E | O |
Melicope hosakae (H.St.John) W.L.Wagner & R.K.Shannon | O | |
Melicope kaalaensis (H.St.John) T.G.Hartley & B.C.Stone | O | |
Melicope kavaiensis (H.Mann) T.G.Hartley & B.C.Stone | K | |
Melicope knudsenii (Hillebr.) T.G.Hartley & B.C.Stone | E, PEPP | K, M |
Melicope lydgatei (Hillebr.) T.G.Hartley & B.C.Stone | E, PEPP | O |
Melicope macropus (Hillebr.) T.G.Hartley & B.C.Stone | EX, SOC | K |
Melicope makahae (B.C.Stone) T.G.Hartley & B.C.Stone | E | O |
Melicope molokaiensis (Hillebr.) T.G.Hartley & B.C.Stone | Mo, M | |
Melicope mucronulata (H.St.John) T.G.Hartley & B.C.Stone | E, PEPP | Mo, M |
Melicope munroi (H.St.John) T.G.Hartley & B.C.Stone | E | Mo, L |
Melicope nealae (B.C.Stone) T.G.Hartley & B.C.Stone | EX, SOC | K |
Melicope oahuensis (H.Lév.) T.G.Hartley & B.C.Stone | O | |
Melicope obovata (H.St.John) T.G.Hartley & B.C.Stone | EX, SOC | M |
Melicope oppenheimeri K.R.Wood, Appelhans & W.L.Wagner | PEPP | M |
Melicope orbicularis (Hillebr.) T.G.Hartley & B.C.Stone | M | |
Melicope ovalis (H.St.John) T.G.Hartley & B.C.Stone | E | M |
Melicope ovata (H.St.John & E.P.Hume) T.G.Hartley & B.C.Stone | K, O | |
Melicope pallida (Hillebr.) T.G.Hartley & B.C.Stone | E | K, O |
Melicope paniculata (H.St.John) T.G.Hartley & B.C.Stone | E | K |
Melicope peduncularis (H.Lév.) T.G.Hartley & B.C.Stone | K, O, Mo, M | |
Melicope pseudoanisata (Rock) T.G.Hartley & B.C.Stone | M, H | |
Melicope puberula (H.St.John) T.G.Hartley & B.C.Stone | E | K |
Melicope quadrangularis (H.St.John & E.P.Hume) T.G.Hartley & B.C.Stone | E, PEPP | K |
Melicope radiata (H.St.John) T.G.Hartley & B.C.Stone | H | |
Melicope reflexa (H.St.John) T.G.Hartley & B.C.Stone | E, PEPP | Mo |
Melicope rotundifolia (A.Gray) T.G.Hartley & B.C.Stone | O | |
Melicope saint-johnii (E.P.Hume) T.G.Hartley & B.C.Stone | E | O |
Melicope sandwicensis (Hook. & Arn.) T.G.Hartley & B.C.Stone | SOC | O |
Melicope sessilis (H.Lév.) T.G.Hartley & B.C.Stone | Mo, M | |
Melicope volcanica (A.Gray) T.G.Hartley & B.C.Stone | Mo, L, M, H | |
Melicope waialealae (Wawra) T.G.Hartley & B.C.Stone | K | |
Melicope wailauensis (H.St.John) T.G.Hartley & B.C.Stone | EX, SOC | Mo |
Melicope wawraeana (Rock) T.G.Hartley & B.C.Stone | K, O | |
Melicope zahlbruckneri (Rock) T.G.Hartley & B.C.Stone | E, PEPP | H |
Platydesma cornuta var. cornuta | E | O |
Platydesma cornuta var. decurrens B.C.Stone | E | O |
Platydesma remyi (Sherff) O.Deg., I.Deg, Sherff & B.C.Stone | C, PEPP | H |
Platydesma rostrata Hillebr. | E | K |
Platydesma spathulata (A.Gray) B.C.Stone | K, O, M, H |
IUCN Red List Category. When evaluated using the World Conservation Union (IUCN) criteria for endangerment (IUCN 2001), Melicope oppenheimeri falls into the Critically Endangered (CR) category, which designates this species as facing a very high risk of extinction in the wild. Our formal evaluation can be summarized by the following IUCN hierarchical alphanumeric numbering system of criteria and subcriteria: CR B1ab(i,ii,iii,v)+2ab(i,ii,iii,v); C2a(ii); D; which reflects a severely limited Extent of Occurrence (EOO) and Area of Occupancy (AOO) of less than 1 km2 and a wild population of only three individuals. It should be noted that seed collections of M. oppenheimeri have been made by Maui PEPP staff during routine monitoring and there is currently a single cultivated individual being grown at the Olinda Rare Plant Facility on East Maui.
We thank Keahi Bustamente, Greg Hansen, Jill Miller, Hank Oppenheimer, and Steve Perlman for their assistance in forest research. We acknowledge the National Tropical Botanical Garden, the U.S. Fish and Wildlife Service, The Hawai‘i State Department of Land and Natural Resources, and the Plant Extinction Prevention Program of Hawai‘i for their continued focus on conservation. We appreciate the efforts of the Hawai‘i State Division of Forestry and Wildlife and their Olinda Rare Plant Facility on East Maui, especially for the cultivation of Melicope oppenheimeri. Much gratitude to Alice Tangerini for the superb illustration, Timothy Flynn for assistance at the