Pacific Science (1986), vol. 40, nos. 1-4 © 1987 by the University of Press. All rights reserved

A Synopsis of Native Hawaiian ' PORTER P. LOWRY Il2

ABSTRACT: The four genera of Araliaceae native to Hawaii, Cheirodendron, Munroidendron, , and , comprise 13 and 2 subspecies, significantly fewer than have been recognized in previous treatments. Three new combinations are made: C.forbesii (Sherff) Lowry, C. platyphyllum ssp. kauaiense (Krajina) Lowry, and C. trigynum ssp. helleri (Sherff) Lowry. Keys are provided to the genera, species, and subspecies, and a complete syn­ onymy is given for each .

Tms SYNOPSIS is being published as a prelimi­ plasandra oahuensis, and to a lesser degree nary note to a more detailed treatment of Cheirodendron trigynum ssp. trigynum, appear Araliaceae that will appear in the forthcoming to constitute somewhat polymorphic assem­ Manual of the Flowering of Hawai'i blages that are in need ofdetailed, field-based (Wagner et al., in press) . The four native study to determine whether certain popula­ genera of Hawaiian Araliaceae, Cheiroden­ tions might warrant recognition at an infra­ dron, Munroidendron, Reynoldsia, and Tetra­ specific level. plasandra, have received little attention since Detailed descriptions ofthe taxa recognized Sherff (1951, 1952a, 1952b, 1952c, 1952d, here will be provided in the forthcoming 1953, 1954, 1955, 1956, 1962) published an manual, along.with extensive discussions of extensive series of notes and . the genera and species. The two naturalized Sherff applied extremely narrow concepts of species of Araliaceae in Hawaii, helix species and infraspecific taxa, so that variable L. and Schefflera actiophylla (Endl.) Harms, populations were split far more than would will also be treated in the manual. A complete be consistent with contemporary taxonomic listing oftype specimens for all names applied practice. In all, he recogni zed 35 species, 116 to Hawaiian Araliaceae will be published in varieties, and 5 forms , many of which were the Nomenclatural Index ofHawaiian ­ described as new. Sherff did not have the op­ ing Plants (Wagner and Mill, in press). portunity to visit Hawaii, however; and so It is interesting to note that many species of lacked firsthand knowledge of the variation Cheirodendron, Reynoldsia and Tetraplasan­ on which his classification was based. dra are key host plants for mating and larval My recent investigations, based on field ob­ development for a large number of Hawaiian servations and detailed study of herbarium Drosophilidae, including many species of specimens, have led me to the conclusion that Drosophila (Heed 1971; Montgomery 1975). native Hawaiian Araliaceae can best be treated Furthermore, all four genera ofnative Hawai­ as comprising 13 species and 2 subspecies, ian Araliaceae serve as food plants for mem­ necessitating the three new combinations made bers of the endemic insect N esiomiris here. Most of these taxa are well delimited (Heteroptera: Mitidae) and appear to have morphologically, and often geographically, played a critical role in the evolution and spe­ and present no special problems. Only Tetra- ciation of this group (Gagne 1982).

KEY TO THE GENERA 1. palmately compound or trifoliate...... 1. Cheirodendron 1. Leaves pinnately compound.

1 Manuscript accepted November 1985. This work was Division of Biologyand Biomedical Sciencesof Washing­ supported in part by NSF Doct oral Dissertation Im­ ton University. provement Grant BSR83-14691 , an ASPT Herbarium ' Missouri Botanical Garden , PO Box 299, SI. Louis, Travel Award, the Missouri Botanical Garden , and the MO 63166-0299. 79 80 PACIFIC SCIENCE, Volume 40,1986

2. Leaflets membranous, margins lobed or toothed and often undulate; petals and 8-12; plants glabrous throughout or sparsely pubescent on the inflorescence and branch tips 3. Reynoldsia 2. Leaflets coriaceous, margins entire; petals and stamens variable in number; plan ts often pubescent, occasionally glabrous. 3. Inflorescence long, pendant, racemo se, the solitary; carpels 12-15; leaflets densely pubescent with furfuraceous stellate hairs 2. Munroidendron 3. Inflorescence compound-umbellate or racemose-umbellate, the flowers in umbellules; carpels 2- 12 (-16); leaflets glabrous or pubescent with scurfy stellate hairs . · ...... 4. Tetraplasandra

CHEIRODENDRON Cheirodendron Nutt, ex Seem., J. Bot. 5:236.1867.

KEY TO THE SPECIES 1. Styles and carpels 2-4 (- 5); leaflet margins usually toothed, occasionally subentire. 2. Leaflets as wide or wider than long, broadl y to depressed ovate , the margins caudate­ dentate, almost spiny, the teeth more than 1mm long, pointing ou t, with an upcurved tip. 3. Styles and carpels (2- ) 3-4 (-5); leaflets very broadl y ovate to depressed ovate , wider than long, the base truncate or nearly so; 1525-1550 m on Mt. Waialeale, . . ·...... 1. C. domin ii 3. Styles and carpels 2 (very rarely 3); leaflets broadly ovate, about as wide as long, the base obtuse to nearly truncate; 650-1250 m on Kauai 2. C. fauriei 2. Leaflets generally longer than wide , elliptic to broadly ovate, the margins entire or serrate-crenate, the teeth less than 1 mm long, strongly curved up and in toward the margin , , ...... 5. C. trigynum 4. Styles and carpels (2-) 3-4 (-5); , , , , , and Hawaii · -...... 5a. ssp. trigynum 4. Styles and carpels 2 (occasionally 3 in a few flowers); Kauai 5b. ssp. helleri 1. Styles and carpels 5 (rarely 4 in some flowers); leaflet margins entire or remotely serrulate. 5. Leaflets elliptic to obovate, more than twice as long as wide, veins of the upper surface sunken in dried specimens; Kauai ...... 3. C.forbesii 5. Leaflets broadly ovate to depressed ovate, as wide or wider than long, veins of the upper surface raised in dried specimens 4. C. platyphyllum 6. Apex of the leaflets acuminate-mucronate; 5.3-7.5 mm long; Oahu . · 4a. ssp. platyphyllum 6. Apex of the leaflets rounded to acuminate, ra rely mucronate; fruit 3.5-5 mm long; Kauai 4b. ssp. kauaiense

1. C. dominii Krajina, Pre slia 10:94. 1931. 3. C.forbesii (Sherff) Lowry, comb. nov. 2. C.fauriei Hochr., Candollea 2 :487.1925. Basionym: C. kauaiense Krajina var. for­ Syn.: C. fauriei Hochr. var. macdanielsii besii Sherff, Bot. Leafl. 5: 12. Sherff, Bot. Leafl. 5: 13. 1951. 1951. C. trigynum (Gaud.) A. Heller var. Syn.: C. kauaiense Krajina var. keakuense kauaiense H. Lev., Repert. Spec. Krajina, Preslia 10: 98. 1931. Nov. Regni Veg. 10: 154. 1911. 4. C. platyphyllum (H ook. & Arnott) Seem., C. wahiawense Krajina, Preslia 10: J. Bot. 5:236. 1867. 98.1931. 4a. ssp. platyphyllum. C. wahiawense Krajina var. popul­ Syn.: Panax ? platyphyllum Hook. & neum _ Krajina , Presli a 10:98. Arnott, Bot. Capt. Beechey's 1931. Voy ., 84. 1832; Hedera pla- Synopsis of Araliaceae-e-Lownv 81

typhylla (Hook. & Arnott) A. var. ilicoides Sherif, Bot. Gray, U.S. Expl. Exped ., Leafl. 5: 10. 1951. Phan., 720. 1854. C. trigynum (Gaud.) A. Heller 4b. ssp. kauaiense (Krajina) Lowry, comb. var. mauiense H. Lev., Repert. nov. Spec. No v. Regni Veg. 154. Basionym: C. kauaiense Krajina, Pres­ 1911. lia 10 :96. 1931. C. trigynum (Gaud.) A. Heller 5. C. trigynum (Gaud.) A. Heller, Minnesota var. mauiense H. Lev. f. Bot. Studies 1:870. 1897. parvulum O. Degener, I. De­ 5a. ssp. trigynum gener & Sherif, Occas. Papers Syn.: Aralia trigyna Gaud., Bot. Bishop Mus . 22(12) :212. Freycin. Voy., tab . 98. 1826 1962. [text, p. 474. (Jan-Mar) C. trigynum (Gaud.) A. Heller 1830]. var. molokaiense Sherif, Bot. Panax ? gaudichaudii DC., Leafl. 5: 10. 1951. Prodr. 4 :253. (Sep) 1830; C. trigynum (Gaud.) A. Heller Hedera gaudichaudii (DC.) var. molokaiense Sherif f. A. Gray, U.S. Expl. Exped ., angustius Sherif, Bot. Leafl. Phan., 71. 1854; C. gaudi­ 5:1 1.1951. chaudii (DC.) Seem., J.Bot. C. trigynum (Gaud.) A. Heller 5:236. 1867. var. oblongum Sherif, Bot. Panax? ovatum Hook. & Arnott, Leafl. 5: 9. 1951. Bot. Capt. Beechey's Voy., C. trigynum (Gaud.) A. Heller 84. 1832. var. oblongum Sherif f. latius C. trigynum (Gaud.) A. Heller Sherif, Bot. Leafl. 5: 10. 1951. var. acuminatum Skottsb., C. trigynum (Gaud.) A. Heller Meddel. Goteb. Bot. Triidg . var. osteostigma Sherif, Bot. 15:417 .1944. Leafl. 5: 12. 1951. C. trigynum (Gaud.) A. Heller C. trigynum (Gaud.) A. Heller var. confertifolium Sherif, var. rockii Sherif, Bot. Leafl. Bot. Leafl. 5: 2. 1951. 5:8.1951. C. trigynum (Gaud.) A. Heller C. trigynum (Gaud.) A. Heller var, degeneri Sherif, Bot. var. skottsbergii Sherif, Bot. Leafl. 5:4.1951. Leafl. 5: 7. 1951. C. trigynum (Gaud.) A. Heller C. trigynum (Gaud.) A. Heller var . degeneri Sherif f. pau ci­ var. subcordatum Sherif, Bot. florum Sherif, Bot. Leafl. 5: 5. Leafl. 5: 9. 1951. 1951. 5b. ssp. helleri (Sherif) Lowry, comb. nov. C. trigynum (Gaud.) A. Heller Basionym: C. helleri Sherif, Bot. var. fosbergii Sherif, Bot. Leafl. 5: 13. 1951. Leafl. 5:7.1951. Syn.: C. trigynum (Gaud.) A. Heller C. trigynum (Gaud.) A. Heller var. multiflorum Skottsb., var . halawanum Sherif, Bot. Meddel. Goteb. Bot. Triidj. Leafl. 5: 3. 1951. 15:416. 1944;C. helleri Sherif C. trigynum (Gaud.) A. Heller var. multiflorum (Skottsb.) var. hawaiiense H. Lev., Re­ Sherif, Fieldiana, Bot. 29 :40. pert. Spec. Nov. Regni Veg. 1954. 10:154.1911. C. helleri Sherif var. micro­ C. trigynum (Gaud.) A. Heller carpum Sherif, Bot. Leafl. var. hillebrandii Sherif, Bot. 5: 14. 1951. Leafl. 5:5 .1951. C. helleri Sherif var. sodalium C. trigynum (Gaud.) A. Heller Sherif, Bot. Leafl. 6: 6. 1952. 82 PACIFIC SCIENCE, Volume 40, 1986

NOTE: Cheirodendron is a subendemic genus Syn.: R. degeneri Sherif, Bot. Leafl. 6: 15. with a single additional species, C. marquesense 1952. F. Brown, occurring in the Marquesas . R. hillebrandii Sherif, Bot. Leafl. The combination ofopposite, palmately com­ 6: II. 1952. pound leaves and a jointed makes R. hosakana Sherif, Bot. Leafl. 6 : 18. Cheirodendron very distinctive within Ara­ 1952. liaceae. R. huehuensis Sherif, Bot. Leafl. AUTHOR'S NOTE: A new combination for C. 6 : 12.1952. marquesense F. Brown will be published by R. huehuensis Sherif var. brevipes David Frodin. Sherif, Bot. Leafl. 6 : 13. 1952. R. huehuensis Sherif var. intermedia Sherif, Bot. Leafl. 6 : 13. 1952. MUNROIDENDRON R. mauiensis Sherif, Bot. Leafl. 6: 16. Munr oidendron Sherif, Bot. Leafl. 7 :21. 1952. 1952. 1. M. racemosum (c. N. Forbes) Sherif, Bot. R. mauiensis Sherif var. macrocarpa Leafl. 7 :22. 1952. O. Degener & Sherif, in Sherif, Syn.: Tetraplasandra racemosa C. N. Bot. Leafl. 7: 17.1952. Forbes, Occas. Papers Bishop R. oblonga Sherif, Bot. Leafl. 6: 17. Mus. 6(4) : 51. 1917. 1952. M. racemosum (c. N. Forbes) Sherif R. sandwicensis A. Gray var. inter­ var. forbesii Sherif, Bot. Leafl. cedens Sherif, Bot. Leafl. 6 : 10. 7 :23.1952. 1952. M. racemosum (C. N. Forbes) Sherif R. sandwicensis A. Gray var. molo­ var. macdanielsii Sherif, Bot. kaiensis Sherif, Bot. Leafl. 6: 11. Leafl. 7 :24. 1952. 1952. R. venusta Sherif, Bot. Leafl. 6: 13. NOTE: Munroidendron is strictly endemic to 1952. the ofKauai and is now very rare. It is R. venusta Sherif var . lanaiensis closely related to and probably derived from Sherif, Bot. Leafl. 6: 14. 1952. either Tetraplasandra or Reynoldsia, but it is easily distinguished by its remarkable inflores­ NOTE: A genus of seven species with two cence and corolla. These three genera, along each in , the Marquesas Islands, and with (some ten species from East the Society Islands, in addition to Reynoldisa Africa, Madagascar, the and Mas­ sandwicensis, which occurs on the islands of carenes, Malesia, and the Solomon Islands) Oahu, Molokai, Lanai, Maui, and Hawaii. a closely related complex recently dis­ Sherif (1952a ) based his 12 new taxa mainly cussed by Philipson (1970). on characters of leaflet shape, inflorescence size, and pedicel length, which are now known to vary widely within individual populations. REYNOLDSIA Reynoldsia A. Gray, U.S. Expl. Exped. , TETRAPLASANDRA Phan., 723. 1854. I. R. sandwicensis A. Gray, U.S . Expl. Tetraplasandra A. Gray, U.S. Expl. Exped., Exped ., Phan., 724. 1854. Phan. , 727. 1854.

KEYTO THE SPECIES I. Inflorescence racemose-umbellate, the rays with lateral as well as terminal branches; inflo­ rescence and lower surfaces of the leaflets often densely scurfy pubescent.

:..,...... 'i,.-' " Synopsis of Araliaceae-e-Lownv 83

2. Stigmas and carpels 7-13; stamens (2-) 3-4 times as many as the petals; fruit globose, strongly ribbed when dry, the disk flattened to low conical, with only the center raised into a narrow, conical stylopodium, the thus appearing nearly completely inferior; Molokai, Lanai, Maui, and Hawaii 2. T. hawaiensis 2. Stigmas and carpels 2-5 (-7); stamens as many as the petals (rarely 2-3 times as many in T. oahuensis) ; fruit ovoid to cylindrical, without ribs when dry (occasionally ribbed in T. oahuensis) , the disk expanded, conical to ovoid, the ovary thus appearing (!) ~ inferior to completely superior. 3. Ovary disk strongly expanded and hypanthium poorly developed below the ring (i.e., the level of attachment of the petals and stamens), the ovary thus appearing fully superior, even in bud and flower; Koolau Mts., Oahu 1. T. gymnocarpa 3. Ovary disk partially expanded but the hypanthium well developed below the ring, the ovary thus appearing at least! inferior. 4. Ultimate units of the inflorescence racemose; inflorescence and lower surfaces of the leaflets densely scurfy stellate pubescent; Kauai, Oahu (Waianae Mts.), Molokai, Lanai, Maui, and Hawaii 3. T. kavaiensis 4. Ultimate units of the inflorescence umbellate; inflorescence and leaflets glabrous throughout; Kauai, Oahu, Molokai, Lanai, Maui, and Hawaii .. 4. T. oahuensis 1. Inflorescence strictly compound-umbellate, the rays without lateral branches (but often with bracts or bract scars); inflorescence and lower surfaces of the leaflets glabrous. 5. Rays of the inflorescence with one to several bracts or bract scars; stamens 5-20 (-25); fruit ca. 5-12 (-15) mm high when mature; Kauai, Oahu, Molokai, Lanai, Maui, and Hawaii 4. T.oahuensis 5. Rays of the inflorescence without bracts or bract scars (or very rarely with one scar in T. waimeae); stamens 25-50; fruit 20-25 mm high when mature. 6. Inflorescence 2-3 times compound-umbellate; stamens 25-30 (-40); carpels 4-7 (-8); fruit ovoid-cylindrical; Kauai ...... 5. T. waialealae 6. Inflorescence once compound-umbellate; stamens 45-60; carpels (7-) 8-12; fruit globose-urceolate; Kauai 6. T. waimeae

I . T. gymnocarpa (Hillebr.) Sherff, Bot. Leafl. 2. T. hawaiensis A. Gray, U.S . Expl. Exped., 7: 12. 1952. Phan., 728. 1854. Syn.: Pterotropia gymnocarpa Hillebr., Syn.: T. hawaiensis A. Gray var. awinien­ FI. Hawaiian lsI., 150. 1888; sis Sherff, Bot. Leafl. 6: 37. 1952. Heptapleurum gymnocarpum (Hil­ T. hawaiensis A. Gray var. gra­ lebr.) Drake, Ill. FI. Ins. Mar. cilis Sherff, Bot. Leafl. 6: 38. Pacif., 183. 1890; Dipanaxgymno­ 1952. carpa (Hillebr.) A. Heller, Min­ T. hawaiensis A. Gray var.·micro­ nesota Bot. Stud. 1:870. 1897. carpa Sherff, Bot. Leafl. 6: 38. Pterotropia gymnocarpa Hillebr. 1952. var. pupukeensis Degener, FI. 3. T. kavaiensis (H. Mann) Sherff, Bot. Leafl. Hawaiiensis, fam . 281. 1938; T. 7: 10.1952. gymnocarpa (Hillebr.) Sherff var. Syn.: Heptapleurum kavaiense H. Mann, pupukeensis (Degener) Sherff, Proc. Amer. Acad. Arts 7: 168. Bot. Leafl. 8: 12. 1953. 1867;Agalma kavaiense (H. Mann) T. gymnocarpa (Hillebr.) Sherff var. Seem., J. Bot. 6: 103.1868; Ptero­ leptocarpa Sherff, Bot. Leafl. 7: 15. tropia kavaiensis (H. Mann) 1952. Hillebr., FI. Hawaiian lsI., 150. T. gymnocarpa (Hillebr.) Sherff var. 1888; Dipanax kavaiensis (H. megalocarpa Sherff, Bot. Leafl. Mann) A. Heller, Minnesota Bot. 7 : 15.1952. Stud. 1:871. 1897. 84 PACIFIC SCIENCE, Volume 40, 1986

Dipanax mannii Seem., J. Bot. 6: 130. 8 :30. 1894; Triplasandra kaalae 1868. Hillebr. , Fl. Hawaiian lsI., 154. Heptapleurum dipyrenum H. Mann, 1888. Proc. Amer. Acad . Arts 7: 168. T. kaalae (Hillebr.) Harms var. multi­ 1867; Pterotropia dipyrena (H . plex Sherif, Bot. Leafl. 8: 11. 1953. Mann) Hillebr. , FI. Hawaiian lsI., T. kahanana O. Degener & Sherif, in 150. 1888;Dipanax dipyrenum (H. Sherif, Bot. Leafl. 8:6. 1953. Mann) A. Heller, Minnesota Bot. T. kohalae Skottsb., Acta Horti Stud. 1:870. 1897; T. ka vaiensis Goteb. 15:414. 1944. (H. Mann) Sherif var. dipyrena T. lanaiensis Rock, Call. Hawaii (H. Mann) Sherif, Bot. Leafl. PubI. Bull. 1: 12.1911. 7: 11. 1952. T. lihuensis Rock var. graeilipes Pterotropia kaalae Skottsb., Acta Sherif, Bot. Leafl. 8: 9. 1953. Horti Goteb, 10: 151. 1936, non T. lydgatei (Hillebr.) Harms, in Eng!. T. kaalae (Hillebr.) Harms . & Prantl, Nat. Pflanzenfam. III , T. kavaiensis (H. Mann) Sherif var. 8:30. 1894; Triplasandra lydgatei grandis Sherif, Bot. Leafl. 7: 14. Hillebr., FI. Hawaiian lsI., 153. 1952. 1888. T. kavaiensis (H. Mann) Sherif var. T. lydgatei (Hillebr.) Harms var. intereedens Sherif, Bot. Leafl. braehypoda Sherif, Bot. Leafl. 7: 13. 1952. 6:19.1952. T. kavaiensis (H. Mann) Sherif var. T. lydgatei (Hillebr.) Harms var. k oloana Sherif, Bot. Leafl. 7: 12. eoreaeea Sherif, Bot. Leafl. 6:21. 1952. 1952. T. kavaiensis (H. Mann) Sherif var. T. lydgatei (Hillebr.) Harms var. nahikuensis Sherif, Bot. Leafl. forbesii Sherif, Bot. Leafl. 6: 20. 7:13.1952. 1952. T. ka vaiensis (H. Mann) Sherif var. T. lydgatei (Hillebr.) Harms var. oecidua O. Degener & Sherif, in leptoraehis O. Degener & Sherif, Sherif, Bot. Leafl. 8:7.1953. in Sherif, Bot. Leafl. 6:20. 1952. T. kavaiensis (H. Mann) Sherif var. T. meiandra (Hillebr.) Harms, in robustior Sherif, Bot. Leafl. 7: 12. EngI. & Prantl, Nat. Pflanzenfam. 1952. III, 8 :30. 1894; Triplasandra T. mierantha Sherif, Bot. Leafl. 7: 11. meiandra Hillebr., FI. Hawaiian 1952. lsI., 152. 1888. T. turbans Sherif, Bot. Leafl. 7: 16. T. meiandra (HiIlebr.) Harms var. 1952. bisobtusa Sherif, Bot. Leafl. 6:36. 4. T. oahuensis (A. Gray) Harms, in EngI. 1952. & Prantl, Nat. Pflanzenfam . III , 8: 30. T. meiandra (Hillebr.) Harms var. 1894. bryanii Sherif, Bot. Leafl. 8 : 10. Syn.: Gastonia? oahuensis A. Gray, U.S. 1953. ExpI. Exped. , Phan ., 726. 1854; T. meiandra (Hillebr.) Harms var. Triplasandra oahuensis (A. Gray) degeneri Sherif, Bot. Leafl. 6:27. Seem., J. Bot. 6: 139. 1868. 1952. Heptapleurum (?) waimeae Wawra, T. meiandra (Hillebr.) Harms var. Flora 56: 159. 1873, non Tetra­ hillebrandii Sherif, Bot. !. plasandra waimeae Wawra. 6:31.1952. T. bisattenuata Sherif, Bot. Leafl. T. meiandra (Hillebr.) Harms var. 6:26. 1952. hiloensis Sherif, Bot. Leafl. 6:34. T. kaalae (Hillebr.) Harms, in EngI. 1952. & Prantl, Nat. Pflanzenfam. III , T. meiandra (Hillebr.) Harms var. Synopsis of Araliaceae-e-Lownv 85

leptomera Sherff, Bot. Leafl. 6 :30. T. meiandra (Hillebr.) Harms var . 1952. skottsbergii Sherff, Bot. Leafl. T. meiandra (Hillebr.) Harms var. 6: 35. 1952, nom. illeg. longipedunculata Skottsb., Acta T. meiandra (Hillebr.) Harms var. Horti Goteb. 15:412. 1944. tenuistylis Sherff, Bot. Leafl. 6:35. T. meiandra (Hillebr.) Harms var. 1952. makalehana Sherff, Bot. Leafl. T. meiandra (Hillebr.) Harms var . 6: 36. 1952. tenuistyloides O. Degener & Sherff, T. meiandra (Hillebr.) Harms var. in Sherff, Bot. Leafl. 8 : 10. 1953. mauiensis Sherff, Bot. Leafl. 6: 29. T. munroi Sherff, Bot. Leafl. 7 :7. 1952. 1952. T. meiandra (Hillebr.) Harms var. T. oahuensis (A. Gray) Harms var. molokaiensis Skottsb., Acta Horti eradiata Sherff, Bot. Leafl. 8: 5. Goteb. 15:412. 1944. 1953. T. meiandra (Hillebr.) Harms var . T . oahuensis (A. Gray) Harms var. occidentalis Skottsb., Acta Horti fauriei Sherff, Bot. Leafl. 8 :3. Goteb, 15:413. 1944. 1953. T. meiandra (Hillebr.) Harms var . T. oahuensis (A. Gray) Harms var. olowaluana Sherff, Bot. Leafl. hailiensis (0. Degener & Sherff) 6:37.1952. O. Degener & Sherff, in Sherff, T. meiandra (Hillebr.) Harms var. Bot. Leafl. 8: 6. 1953; T. meiandra ovalis Sherff, Bot. Leafl. 6: 31. (Hillebr.) Harms var. hailiensis O. 1952. Degener & Sherff, in Sherff, Bot. T. meiandra (Hillebr.) Harms var. Leafl. 7 : 10. 1952. polyantha Sherff, Bot. Leafl. 6: 28. T. oahuensis (A. Gray) Harms var . 1952. longipes Sherff, Bot. Leafl. 8: 4. T. meiandra (Hillebr.) Harms var. 1952. polystigmata Sherff, Bot. Leafl. T . oahuensis (A. Gray) Harms var . 7:9. 1952. pseudolongipes O. Degener & T. meiandra (Hillebr.) Harms var. Sherff, in Sherff, Bot. Leafl. 8 :4. prolifica Sherff, Bot. Leafl. 6: 32. 1953. 1952. T. oahuensis (A. Gray) Harms var. T. meiandra (Hillebr.) Harms var. pseudorachis Sherff, Bot. Leafl. prolificoides Sherff, Bot. Leafl. 8: 3. 1953. 6: 33. 1952. T. oahuensis (A. Gray) Harms var . T. meiandra (Hillebr.) Harms var. subglobosa O. Degener, I. Degener ramosior Sherff, Bot. Leafl. 6: 36. & Sherff, in Sherff, Occas. Papers 1952. Bishop Mus. 22(12) :211. 1962. T. meiandra (Hillebr.) Harms var. T. oahuensis (A. Gray) Harms var. rhynchocarpa Sherff, Bot. Leafl. venulosior Sherff, Bot. Leafl. 8: 2. 6: 33. 1952. 1953. T. meiandra (Hillebr.) Harms var. T. pupukeensis Skottsb., Acta Horti rhynchocarpoides O. Degener & Goteb, 10: 149. 1936. Sherff, in Sherff, Bot. Leafl. 7: 8. T. pupukeensis Skottsb. var. dicipiens 1952. O. Degener & Sherff, in Sherff, T. meiandra (Hillebr.) Harms var . Bot. Leafl. 8: 8. 1953. rockii Sherff, Bot. Leafl. 6: 33. T. pupukeensis Skottsb. var. megalo­ 1952. poda Sherff, Bot. Leafl. 6 :40. T. meiandra (Hillebr.) Harms var. 1952. simulans Sherff, Bot. Leafl. 6 :35. T. pupukeensis Skottsb. var. megalo­ 1952. poda Sherff f. trigona O. Degener 86 PACIFIC SCIENCE, Volume 40, 1986

& Sherff, in Sherff, Bot. Leafl. suitable for the recognition ofsuch taxa. Also, 7: 7. 1952. the sections of Tetraplasandra recognized by T. pupukeensis Skottsb. var. nitida Sherff were based in large part on O. Degener & Sherff, in Sherff, number and consequently do not appear to Bot. Leafl. 6:39. 1952. represent natural, monophyletic groupings. T. pupukeensis Skottsb. var. venosa O. Degener & Sherff, in Sherff, Bot. Leafl. 6:40.1952. ACKNOWLEDGMENTS T. waianensis O. Degener, Fl. Ha­ waiiensis, fam. 281. 1938. I am grateful to W. L. Wagner and D. R. T. waianensis O. Degener var. pale­ Herbst for numerous stimulating discussions huana Sherff, Bot. Leafl. 8: 11. and many valuable comments and suggestions 1952. and to S. W. Mill for providing extensive bib­ 5. T. waialealae Rock, CoIl. Hawaii Publ. liographic information. I thank P. H. Raven Bull. 1: 10. 1911. for critical comments on the manuscript. J. Syn.: T. lihuensis Sherff, Bot. Leafl. 8: 8. Obata, S. Lucas, R. Hobdy, and H. Le Dantec­ 1953. Lowry generously provided assistance in the T. waialealae Rock var. acrior Sherff, field. I also thank the Pacific Tropical Botani­ Bot. Leafl. 6: 22. 1952. cal Garden for the use ofa vehicle and facili­ T. waialealae Rock var . kohuana ties while on Kauai. The curators of the fol­ O. Degener & Sherff, in Sherff, lowing herbaria provided the loan of speci­ Amer. J. Bot. 43 :476.1956. mens or study facilities for examining material T. waialealae Rock var. pluricostata in situ: A, BH, BISH, BM, BRI, CANB, CAS, CM, G, O. Degener & Sherff, in Sherff, GB, GH, K, MEL, MO, NSW, NY, P, PH, S, UC, US, Amer. J. Bot. 43:476.1956. andw. T. waialealae Rock var. subsessilis Sherff, Bot. Leafl. 6: 23. 1952. T. waialealae Rock var. urceolata LITERATURE CITED Sherff, Bot. Leafl. 6: 23. 1952. T. waialealae Rock var. wahiawensis GAGNE, W. C. 1982. Insular evolution and Sherff, Bot. Leafl. 6:24. 1952. speciation of the genus Nesiomiris in Ha­ 6. T. waimeae Wawra, Flora 56: 158.1873. waii (Heteroptera: Miridae). Entomologia Syn.: T. elstonii Hochr., Candollea 2 :481. Generalis 8:87-88. 1925. HEED, W. B. 1971. Host specificity and T. sherffii O. Degener & 1. Degener, speciation in Hawaiian Drosophila. Taxon Fl. Hawaiiensis, fam. 281. 1962. 20: 115-121. Triplasandra waimeae (Wawra) MONTGOMERY, S. L. 1975. Comparative A. Heller, Minnesota Bot. Stud. breeding site ecology and the adaptive 1:871.1897. radiation of picture-winged Drosophila T. waimeae Wawra var. angustior (Diptera: Drosophilidae) in Hawaii. Proc. Sherff, Bot. Leafl. 6: 26. 1952. Hawaiian Entomol. Soc. 12: 65-103. PHILIPSON, W. R. 1970. A redefinition of NOTE: As defined by Philipson (1970), Tetra­ Gastonia and related genera (Araliaceae). plasandra is endemic to Hawaii; those species Blumea 18:497-505. described from Malesia were referred to Gas­ SHERFF, E. E. 1951. New entities in the genus tonia. The numerous taxa recognized by Cheirodendron Nutt. ex Seem. (fam. Ara­ Sherff (1955) were based primarily on varia­ liaceae) from the Hawaiian Islands. Bot. tions in the number of stamens and carpels Leafl.5 :2-14. and in leaflet size and shape. These same char­ ---. 1952a. Further studies of Hawaiian acters are now known to vary widely within Araliaceae: Additions to Cheirodendron individual populations and, therefore, are not helleri Sherff and a preliminary treatment

• &Zlh 1a·IM4A1&S4!bir!W 1 8 a ¥¥AiI PhiliP ! oJ!4 .J I i P hthA \I !. i#I:@trfflt'iW%'#fI3.ffit'4W !¢£¥ #hliii¥hEe#ii2i.Q¥ Synopsis ofAraliaceae-s-Lownv 87

of the endemic species of Reynoldsia. A. ---. 1955. Revision ofthe Hawaiian mem­ Gray. Bot. Leafl. 6 :6-19. bers of the genus Tetraplasandra A. Gray. - --. 1952b. Additions to our knowledge of Fieldiana, Bot. 29 :49-142. the genus Tetraplasandra A. Gray (fam. --- . 1956. Some recently collected dico­ Araliaceae). Bot. Leafl. 6: 19-41. tyledonous Hawaiian Island and Peruvian - -- . 1952c. Contributions to our knowl­ plants. Am. J. Bot. 43: 475-478. edge of the genera Tetraplasandra A. Gray - --. 1962. Miscellaneous notes on some and Reynoldsia A. Gray (fam. Araliaceae) American and Hawaiian dicotyledons. in the Hawaiian Islands. Bot. Leafl. 7 :7­ Occas. Papers Bishop Mus. 22(12) :207­ 17. 214. - -- . 1952d. Munroidendron, a new genus WAGNER, W. L., D. H. HERBST, and S. H. ofaraliaceous from the island ofKauai. SOHMER. In press. Manual ofthe Flowering Bot. Leafl. 7: 21-24. Plants of Hawaii. Bishop Museum Press, - - -. 1953. Further notes on the genus Honolulu. Tetraplasandra A. Gray (fam. Araliaceae) WAGNER, W. L., and S. W. MILL. In press. in the Hawaiian Islands. Bot. Leafl. 8: 2­ Nomenclatural Index ofHawaiian Flower­ 13. ing Plants. Bishop Museum Press, Hono­ - - -. 1954. Revision of the genus Cheiro­ lulu. dendron Nutt. ex Seem. for the Hawaiian Islands. Fieldiana, Bot. 29: 1-45.

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