SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 24

Flora of Micronesia, 2: Casuarinaceae, Piperaceae, and Myricaceae

F. Raymond Fosberg and Marie-Hélène Sachet

ABSTRACT

Fosberg, F. Raymond, and Marie-Hélène Sachet. Flora of Micronesia, 2: Casuarinaceae, Piperaceae, Myricaceae. Smithsonian Contributions to Botany, number 24, 28 pages, 1 figure, 1975.—The second installment of the Flora of Micronesia gives systematic treatments, including descriptions, synonymy, pertinent literature references, keys, ethnobotany, citations, geographic records, and specimens examined, of the families Casuarinaceae, Piperaceae, and Myricaceae.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report; Smithsonian Year. SI PRESS NUMBER 5266. SERIES COVER DESIGN: Leaf clearing from the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini.

Library of Congress Cataloging in Publication Data

Fosberg, Francis Raymond, 1908–

Flora of Micronesia.

(Smithsonian contributions to botany, no. 20, 24

Bibliography: v. 1, p. v. 2, p.

Supt. of Docs. no.: SI 1.29:20

Contents: 1. Gymnospermae. 2. Casuarinaceae, Piperaceae, and Myricaceae.

1. Botany—Micronesia. 2. Ethnobotany—Micronesia. I. Sachet, Marie-Hélène, joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to botany, no. 20 [etc.]

QK1.S2747 no. 20, etc. [QK473.M5] 581′.08s [581.9′96′5] 74–11316

Contents

Copyright

Introduction

Casuarinaceae

Casuarina L

Piperaceae

Peperomia Ruiz & Pavón

Piper L.

Synonyms and Excluded or Misapplied Names

Myricaceae

Myrica L.

Literature Cited

Index

Flora of Micronesia,
2: Casuarinaceae, Piperaceae,
and Myricaceae

F. Raymond Fosberg and Marie-Hélène Sachet

F. Raymond Fosberg and Marie-Hélène Sachet, Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

This second installment of the Flora of Micronesia follows essentially the same format as the first installment (Smithsonian Contributions to Botany, number 20). Treated in this installment are the families Casuarinaceae, with one genus and one Micronesian species; Piperaceae, with 2 genera, 15 native and 4 introduced species, including a number of minor taxa; and Myricaceae, with one introduced genus and one species.

More complete bibliographic references may be found in Sachet and Fosberg, Island Bibliographies (1955), and Supplement (1971), with a complete list of serial abbreviations.

For details of the history and circumstances of the present flora, as well as an explanation of its scope and arrangement, reference may be made to the introductory pages of the first installment.

It may be useful to restate here that in citations of specimens the collector and number are italicized if the specimen has been examined by the authors. Herbarium symbols are according to the 5th edition of Index Herbariorum (Lanjouw and Stafleu, 1964), with the additions: “UH” for University of Hawaii, “UG” for University of Guam, and “Fo” for specimens still in possession of the authors. Such symbols are only cited for sheets actually studied, not for sheets that are known to be in other herbaria but not examined in this study.

CASUARINACEAE

Trees with cylindrical, articulate, striated green branchlets which serve as leaves; the true leaves reduced to whorls of minute scales at the nodes; flowers monoecious or dioecious, much reduced, without perianth, arranged in aments, wind pollinated, the staminate aments cylindric, articulate; pistillate aments capitate, developing into woody cone-like structures formed from thickened indurate floral bracts; fruit a samara with a single wing.

A family of a single genus (by some regarded as two), principally Australasian.

Casuarina L.

Casuarina L., Diss. Stickman, 12, 1954 [as Casaarina, sphalm.]; Amoen. Acad., 4:143, 1759.

Characters of the family.

A small genus, principally Australian, with several species in New Caledonia, New Guinea, and Malesia, one widespread in the Indo-Pacific region, including the high islands of Micronesia, and generally introduced in the tropics and subtropics.

There has been much uncertainty about the validity of publication of this generic name by Linnaeus,

as it is not accompanied by a description. However, under the International Code, the reference to an earlier description and plate of the type-species by Rumphius (1743:86, pl. 57) serves to validate it, since it was a monotypic genus as treated by Linnaeus. The same application of Article 42 of the International Code of Botanical Nomenclature makes it necessary to accept the binomial Casuarina litorea in place of the widely familiar C. equisetifolia. This situation will be discussed at length in another paper.

Casuarina litorea L.

Casuarina litorea [Rumphius] L., Diss. Stickman, 12, 1754 [as Casaarina litorea, sphalm.] [type Rumphius, vol. 3, pl. 57].

Casuarina equisetifolia L., Amoen. Acad., 4:143, 1759 [as equisefolia, sphalm.].—Thomas, Jour. Voy. South Seas 168, 1745.—Chamisso, Remarks and Opinions, 77, 1821.—Miquel in de Candolle, Prodr. 16(2) :339, 1868.—Kubary, Ethn. Beitr. Kenntn. Karol. Arch., 101–102, 1895.—Schumann and Lauterbach, Fl. Deutsche Schutzg. Süds., 254, 1901.—Fritz, Mitt. Deutsch. Schutzg., 15:96–118, 1902.—Safford, Contr. U.S. Nat. Herb., 9:220, 1905.—von Prowazek, Deutschen Marianen, 115, 1913.—Merrill, Phil. Jour. Sci. Bot., 9:71, 1914.—Corte [transl.], Guam Recorder, 3:158, 330, 1926 [repr. 1972].—Kanehira, Bot. Mag. Tokyo, 45:274, 1931; Fl. Micr., 80, 1933.—U.S.D.A., Ann. Rept. Guam Agr. Expt. Sta., 1932:17, 1933.—Hosokawa, Kudoa, 2:107–113, 1934.— Kanehira, Enum. Micr. P., 303, 1935.—Okabe, Jour. Anthrop. Soc. Nippon, 56:423, 1941; Nankyo, 2:18, 1943.— Okabe, Nettai Sangyô Kenkyu-sho ihô, 5:2, 1940.—Tuyama, Kagaku Nanyô, 4:16, 1941.—Bryan, Plants of Guam [unpub,. 1946?].—Thompson, Guam and Its People, ed. 3, 200, 1947.—Burcham, Contr. U.S. Nat. Herb., 30:430, 1948.— Guillaumin, Bull. Soc. Bot. France, 99:39, 1952.—Glassman, Bishop Mus. Bull., 209:79, 1952—Glassman, Pac. Sci., 7:295, 1953.—Catala, Atoll Res. Bull., 59:88, 1957.—Stone, Pac. Sci., 13:100, 1959.—Fosberg, Atoll Res. Bull., 67:11, 1959.— Fosberg and Sachet, Atoll Res. Bull., 92:14, 1962.—Whiting, m.s., 1965.—Fosberg and Sachet, Atoll Res. Bull., 123:7, 1969.—Stone, Micronesica, 6:241–242, 1971.—Fosberg, Falanruw, and Sachet, Smithsonian Contr. Bot., 22:20, 1975.

Casuarina mertensiana Ruprecht ex Miquel in de Candolle, Prodr., 16(2) :339, 1868 [type from Guam, Mertens, Chamisso, not seen by us].

Tree, reaching a large size, heart-wood very hard and heavy; branchlets jointed, joints about 1 cm long, about 1 mm or less thick, with 6–8 striae; leaves in whorls of 6 to 8; flowers monoecious to dioecious, staminate in cylindric elongate aments, pistillate in shorter turbinate aments, styles maroon, filiform, fruiting aments cylindric to globose, about 1–1.5 cm thick, up to 2 cm or more long; fruit with a transparent wing.

Indo-Pacific region from East Africa to the Society and Marquesas Islands, north to the Bonins (probably introduced) and Ryukyus. In Micronesia in the Marianas: Maug, Agrigan, Pagan, Alamagan, Anatahan, Saipan, Tinian, Rota, Guam. Carolines: Palau, Mapia, Yap, Namonuito, Truk, Satawan, Ponape, Mokil. Marshalls: Kwajalein, Likiep, Jaluit. Nauru. Gilberts: Butaritari, Tarawa. In the atolls and probably Nauru it is regarded as introduced. What was probably this species was reported from Tinian as “Red-Wood-Tree” or “Iron Wood” by Thomas (1745:168), who visited the island with Anson in 1742.

Found on strands, cliffs, and other open habitats, becoming established as a pioneer on mineral soil, either of limestone or volcanic origin, or even on fresh lava or volcanic ash, apparently able to invade grassland, persisting for a long time in forests but not invading closed woody vegetation. Under a stand of Casuarina, in relatively dry or well-drained areas, few plants of any kind become established, apparently because of the thick accumulation of fallen branchlets or “needles”; but in wetter areas these decompose more rapidly and many plants grow in the resulting humus layers. Characteristic complex-branching nodules on the Casuarina roots are apparently related to the demonstrated nitrogen-fixing ability of the genus. On Guam, when the savannas are protected from fire, they tend to grow up to Casuarina forests.

USES.—The wood is heavy, strong and very hard, but in Guam it is scarcely at all utilized as it is hard to work (Safford, 1905). Used for abortions (Thompson, 1947:200). “Roots are used in a medicinal tea. Roots and trunk (?) ‘without bark’ are used in a douche, reputedly as an abortifacient. Wood of the tree is burned as a mosquito repellent” (Guam: Whiting, MS, 1965). An astringent principle is obtained from the bark and is used for chronic diarrhea (Okabe, 1940:2).

VERNACULAR NAMES.—

ironwood, beefwood, red-wood tree, Polynesian iron-wood, she-oak, Australian pine (English)

gago (Marianas: von Prowazek, 1913)

gago (Pagan: Corte, 1926)

goago (Rota: Kanehira, 1935)

gagu (Rota: Fosberg 24992)

agoho or gago (Guam: Corte, 1926)

gago (Guam: Fosberg 25363, Marche 142, Nelson 133, Stone 4247, Whiting C9; Safford, 1950; Merrill, 1914; Corte, 1926; Bryan, 1946)

gagu (Guam: Stone, 1971)

gao-gao (Guam: Dutton 130)

agas (Palau: Okabe, 1943, n. 18)

agasu (Palau: Okabe, 1941:423)

ngas (Palau: Walleser and Krämer in Tuyama, 1941; Emmons 2)

ngasu (Palau: Kanehira, 1935)

noc (Yap: Wong 465)

lach (Ulithi: Fosberg 46613)

ueuhir (Satawal: Fosberg 46851)

neokuk (Satawan, Ta I.: Anderson 1058)

ueoku (Satawan, Moch I.: Anderson 1000)

ueokuk (Satawan, Ta I.: Anderson 1058)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Gaudichaud (P); planted on Uracas, Asuncion, Guguan, and Medinilla by Fritz, 1902:96–118.

Maug: Seen by Falanruw in 1972.

Agrigan: Fritz, 1902:110; trail around south side of island, on cliffs and bluffs, 10–20 m, Fosberg 31578 (US, BISH, Fo, NY, L).

Pagan: Hosokawa 7949 (US, Fo), 7989 (BISH); Bonham 20 (US, Fo); Kondo in 1949 (BISH); Fritz, 1902:106; isthmus, planted in rows near pier on west side of isthmus, Anderson 580 (US, BISH, Fo, NY, L); Mt. Charlen, on fresh lava flow east of Fresh Water Lake, Fosberg 31411 (US, BISH, Fo, NY, L); Lagona, Villagomez JV–02 (US); Mt. Pagan crater floor, Moore 371 (US).

Alamagan: Around Partido village, on sea bluffs, Fosberg 31646 (US).

Anatahan: Fritz, 1902:98; boulder strewn beach area, sea level, Falanruw 1660 (US); northwest tip of island, 0–10 m, Evans 2447 (US, BISH, Fo, NY).

Saipan: Stephens 51 (Fo), 85 (Fo); Marche 1 (P, Fo); Kanehira 1013 (FU, 2 sheets); southwest corner of island, just north of Agingan Pt., 1–3 m, Fosberg 25257 (US, BISH, Fo, NY, L); Kanehira & Hatusima 4408 (FU); Marpi, Courage 19 (US); near ISCOM Base Command, center of island, 600 ft [180 m], Conover 956 (US, BISH).

Agiguan: Kondo in 1952 (BISH).

Tinian: Thomas, 1745:168; Kondo 33 (BISH); Cameron in 1944 (BISH); Mt. Lasso, 150–175 m, Fosberg 24889 (US, BISH, Fo).

Rota: Grether 4461 (UC); Necker R46 (US); northwest coast of west end of main part of island, between Rota and Tataacho Pt., Fosberg 24992 (BISH); Taipinkoto, Hosokawa 7657 (BISH, Fo); Songsong village and vicinity, 5–10 m, Evans 2031 (US); just behind the west dock, 5–10 m, Evans 1971 (US, BISH, Fo, NY).

Guam: Mertens (P); Marche 142 (P); McGregor 538 (US, BISH, UC); G.E.S. 431 (US, BISH); Chamisso, 1821:77; Agaña, Seale in 1900 (BISH); 4 km east of Agat, Necker 113 (US, BISH); Agfayan Bay, within 100 m of shore, Necker 136 (US, K); 1 mile [1.6 km] south of Barrigada, 400–600 ft [120–180m], G. C. Moore 38 (US, UC); near top of Mt. Tenjo, Rodin 544 (UC); along ocean, south of Ylig River, Rodin 822G (US, UC); Manengon, 150 m, Evans 1789 (US, UH, Fo, K, MO); Manengon, Tarzan Falls, B. C. Stone 4976 (US, UG); Harmon village, Stone 4247 (BISH, UG); Merizo, Whiting C9 (US, Fo); along beach, Fosberg 25363 (US, BISH, Fo, NY, L); Masso, Nelson 133 (BISH); Dadi Beach, Sumay, Bryan 1211 (BISH, K); between Umatac and Cetti Bay 0–10 m, Evans 1555 (US, BISH, Fo, NY, L, P); Yigo, Dutton 130 (US, BISH, Fo).

CAROLINE ISLANDS.—Palau: Ledermann 14052 (K). Kayangl: Salsedo 401 (US). Balbeldaob: Arekalong, Takamatsu 1664 (US, UC, K). Koror: Entomology Lab. Emmons 2 (US); Ngerdis, site of old Japanese plane ramp, Salsedo 67 (US). Ngarakabesang (Arakabesan): Takamatsu 1254 (BISH); west peninsula of island, 10–20 m. Fosberg 32470 (US, BISH, Fo). Ngeanges (Gaingas): in Yoo (Sar) Passage, just west of south point of Urukthapel, 2 m, Fosberg 25838 (US, BISH, Fo, NY, L). Peliliu: Burcham, 1948:430. Angaur: Kanehira, 1931:274 (citing Kanehira 574); Kanehira 1075 (FU); northwest coast, on bluffs above sea, 30 m, Fosberg 25929 (US, BISH).

Mapia: Kubary, 1895:101–102.

Yap: 75 ft [23 m], Wong 465 (US, BISH Fo).

Ulithi Atoll: Asor I., 1–2 m, in village, planted, Fosberg 46435 (US); Falalap I., 1–3 m, Fosberg 46613 (US).

Satawal I.: north side of island, just back of beach, one tree seen, 2 m, Fosberg 46851 (US).

Namonuito: Ulul I., Stone, 1959:100.

Truk: Moen (Harushima): Takamatsu 262 (BISH). Dublon (Natsushima): Takamatsu 4 (BISH). Udot: Hosaka 2739 (US, BISH, Fo).

Satawan: Moch I., Anderson 1000 (US, BISH, Fo, NY, L); Ta I., Anderson 1058 (US, BISH, Fo, NY, L).

Ponape: Colonia, Glassman 2912 (US, BISH).

Mokil: Glassman, 1953:295.

WAKE ISLAND—Seen commonly planted around buildings by Fosberg in 1953, Sachet in 1961, and by Fosberg and Sachet in 1963.

MARSHALL ISLANDS.—Kwajalein: Growing in pot, 1956, seen but not collected by Fosberg; Kwajalein I., St. John 23700 (BISH).

Likiep: Likiep I., Fosberg 36630 (US, BISH, Fo, NY).

Jaluit: Jabor, Lyman 7 (US, BISH, Fo, NY); Fosberg 39469 (US).

NAURU ISLAND.—Several trees introduced from Australia, Burges in 1933 (unpubl.).

GILBERT ISLANDS.—Butaritari (Makin): Catala 65 (P). Tarawa: Seen in garden of Residency at Bairiki by Catala, 1957:88; seen at Bairiki, Betio and Bikenibeu by Sachet in 1967; Bikenibeu I., Herbst & Allerton 2659 (US); Betio Adair 56 (US).

Casuarina litorea var. souderi (Fosberg) Fosberg & Sachet, new combination

Casuarina equisetifolia var. souderi Fosberg, Micronesica, 2:144, 1966.

Branchlets fastigiately arranged, internodes so shortened that the scale-like leaves are imbricate.

Apparently endemic to Guam and very rare there, doubtless a mutation that has arisen from the common form and persists on Guam.

MARIANAS ISLANDS.—Guam: Orote, Naval Station, Souder A–1 (BISH, type, US, Fo).

PIPERACEAE

Shrubs, vines or herbs; leaves simple, entire, alternate, opposite, or whorled; flowers in spikes or racemes, much reduced, without perianth, but each subtended by a peltate bract, or surrounded by 3 bracts, perfect or unisexual; stamens 2 or more, anthers usually 2-celled; ovary 1-celled with 1 basal ovule, stigmas 1 to several, sessile; fruit a drupe or a minute sticky nut.

A pantropical family found in many habitats but common in forest undergrowth, epiphytic, and on rocks. Two very large genera, Piper and Peperomia, are usually recognized, but some botanists divide Piper into several genera. A conservative course is followed here, admitting only the two, keeping Piper intact. Both genera are well-represented in Micronesia.

Key to the Micronesian Genera of Piperaceae

Woody plants, flower with several stamens and stigmas, and 3 bracts, fruit a drupe ........Piper

Herbaceous plants with fleshy stems and (usually) leaves, flower with two stamens, one stigma, and one peltate bract; fruit a sticky nutlet .......................................................Peperomia

Peperomia Ruiz & Pavón

Peperomia Ruiz & Pavón, Prodr. Fl. Peru, 8, 1794.—Kanehira, Enum. Micr. Pl., 303, 1935.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:7–25, 1938; 22:96–97, 1959.—Fosberg, Phytologia, 13:233–234, 1966.

Micropiper Miquel, Comment. Phytogr., 39, 1840.

Herbs, usually fleshy; leaves alternate, opposite, rarely whorled, in Micronesian species with palmate venation, petioles expanded or not at base; spikes terminal, leaf-opposed, or axillary, fleshy, lower leaves on stems often reduced; flowers bisexual, reduced to an ovary and two stamens subtended by a stalked peltate bract; fruit a very sticky minute nut or nut-like drupe (usually described as a berry).

Pantropical, with a number of ill-distinguished species in Micronesia. Common in moist or wet forests, epiphytic or terrestrial, especially on rocks. Several species are grown as ornamental foliage-plants, including, in Micronesia, P. obtusifolia (L.) A. Dietrich, seen as a pot plant in Guam and Likiep (Fosberg 36724 (US)) and P. caperata Yuncker, also said to be grown on Guam. It is probable that the smooth peltate-leafed species P. argyreia Morren (P. sandersii C. de Candolle) is grown as a pot plant in Guam or other islands, but no collections or records are available.

Key to Micronesian Species of Peperomia

1. Leaves cordate or peltate ............................................................2

2. Stem short, leaves rather congested, mostly peltate, bullate, petioles longer than blades .......................................................................................................P. caperata

2. Stem elongate, leaves not congested, petioles shorter than blades, blades thin, never peltate, flat, fruit longitudinally ribbed ..............................P. pellucida

1. Leaves rounded, obtuse or acute at base, fleshy, fruit papillose or verrucose, not ribbed ..................3

3. Stems glabrous ..................................................................4

4. Leaves mostly less than 2 cm long, plant decumbent with erect fertile branches, spikes up to 4 cm long .........................................................P. kraemeri

4. Leaves 2–4 or more cm long, main stems decumbent only toward base, spikes when well developed 3–6 cm long .............................................................5

5. Leaves tending to be broadest at or below the middle, peduncles of spikes mostly less than 1 cm long .........................................................P. mariannensis

5. Leaves tending to be broadest above middle, peduncles mostly 1 cm or more long .........6

6. Leaves leathery, pinnately nerved, spikes terminal, 2–3 mm thick, usually more than 7 cm long ...........................................P. obtusifolia

6. Leaves fleshy, palmately basally nerved, spikes terminal and in upper axils, 2 mm or less thick, usually less than 7 cm long ...............................P. ponapensis

3. Stems more or less hairy ........................................................7

7. Rachis of spike at least sparsely hairy, plant puberulent with appressed or incurved short hairs .......................................................P. palauensis

7. Rachis of spike glabrous, plant variously hairy .....................................8

8. Stems usually rooting at nodes only near base, tending to be erect, spikes mostly exceeding leaves ..........................................................9

9. Plant densely pubescent, leaves all opposite or whorled ............P. leptostachya

9. Plant sparsely puberulent, at least some leaves usually alternate ...........................10

10. Leaves mostly broadest below middle, hairs appressed ......................................................................................P. mariannensis f. saipana

10. Leaves mostly broadest at or above middle, hairs tending to be spreading or merely curved upward .................................................11

11. Leaves ciliolate only near tip, acute or somewhat attenuate at base, stem prominently hirtellous ..........................P. palauensis var. occidentalis

11. Leaves ciliolate all around, obtuse to somewhat rounded at base, stem sparsely hirtellous P. ponapensis var. trukensis

8. Stems prominently rooting at nodes, decumbent, spikes 1–2 cm long ....................12

12. Stems abundantly subappressed hirtellous with short straight hairs, these less than 0.5 mm long, fruit 0.8 mm long .............................................P. breviramula

12. Stems pubescent with curved or long straight hairs, fruit 0.4–0.5 mm long ......13

13. Hairs on stem long and straight, pubescence appearing shaggy, leaves broadly elliptic to orbicular ............................................P. kusaiensis

13. Hairs on stems curved, pubescence crisp, leaves elliptic-lanceolate ..........................................................P. glassmanii

Peperomia breviramula C. de Candolle

Peperomia breviramula C. de Candolle, Bot. Jahrb., 56:503, 1921 [type from Ponape, Ledermann 13739, not seen by us].—Kanehira, Enum. Micr. Pl., 393, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:22–23, 1938; 22:107, 1959.—Hosokawa, Trans. Nat. Hist. Soc. Taiwan, 33:120, 1943.

Herb with decumbent to ascending stems, rooting at nodes, abundantly subappressed hirtellous, hairs straight, less than 0.5 mm long; leaves alternate 1–2 cm long, up to 1.6 cm wide, oval-elliptic to subovate or suborbicular, obtuse at both ends or base somewhat obtusish, trinerved from base, white beneath, abundantly ciliate, moderately hirtellous on both faces when young, becoming glabrate when older, petiole 3–5 mm long, hirtellous; spikes 1–1.8 cm long, pedunculate, terminal or opposite upper leaves, moderately flowered, glabrous; peduncles 5–9 mm long, hirtellous; fruit subglobose, viscid papillose, 0.8 mm long. (Description adapted from Yuncker’s (1938) description and figure.)

Endemic to Ponape, known only from the type and one other collection; growing epiphytically in montane forests.

CAROLINE ISLANDS.—Ponape: Nipit, 350 m, Hatusima 10858 (FU); Paue, Montesanto, de Candolle, 1921:503 (citing Ledermann 13739, type, not seen by us, probably destroyed). Some material previously cited as this species is now regarded as belonging to P. glassmanii.

Peperomia caperata Yuncker

Peperomia caperata Yuncker, Kew Bull., 1957:421, 1958 [type from cultivated material, native home uncertain].

This species was mentioned in a letter from Paul Souder, as growing in Guam, and identified by E. H. Bryan, Jr. We have seen no material. It is a plant with very short stem, erect prominently cordate, usually somewhat peltate, very bullate leaves, which has become common as a house or greenhouse plant in recent years. No circumstances were given as to its occurrence on Guam.

Peperomia glassmanii Yuncker

Peperomia glassmanii Yuncker, Occ. Pap. Bishop Mus., 22:105–107, 1959 [type from Ponape, Glassman 2844].—Glassman, Bishop Mus. Bull., 209:54, 1952 [pro parte; as P. palauensis].

Weak, fleshy, creeping herbs, rooting at lower nodes, stems densely to sparsely puberulent with curved hairs; leaves alternate, well-developed ones elliptic to broadly lanceolate, mostly 0.7–1.5 cm wide, 2–4 cm long, acutish at both ends, 3–5 nerved, nerves from base, outer pair, when 5, weakly developed, blades strongly ciliate, more or less puberulent on both sides, especially on nerves, petiole 3–5 (rarely 10) mm long; spikes pedunculate, terminal and opposite upper leaves, not numerous, 1–1.5 cm long, with peduncles almost as long, peduncle puberulent, rachis glabrous, rather sparsely flowered; fruit about 0.4 mm across, globose ovoid, very few seen.

This species may prove to be too close to P. breviramula when that species is better known.

Endemic to Ponape, where it grows as an epiphyte in montane rain forest and cloud forest at middle to high elevations.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Ponape: Ledermann 13175 (BISH); Koidzumi in January 1915 (TI); Tolomail, Takamatsu 986 (BISH); Mt. Tolomail, Takamatsu 981 (BISH); Mt. Tolotom, 1200 ft [365 m] Glassman 2844 (US, isotype, BISH, type); Mt. Troton, Hosokawa 9534 (BISH, A); Mt. Tolenkiup, 500 ft [150 m], Glassman 2530 (US, BISH); Mt. Kankauzan, Hosokawa 5481 (BISH, A); Mt. Tamatamansakir, 1200 ft [365 m] Glassman 2328 (US, BISH); Niinioanii, Hosokawa 5701 (A); Tolun Nanket, mountain above Nanipil, drainage of Tawenjokola River, Not District, 600 m (above power plant), Fosberg 26423 (US, DPU), Fosberg 26434 (US, BISH, Fo, DPU, NY); 500 m, Fosberg 26456 (US, BISH, DPU); Mt. Nanalaut, 2000 ft [600 m], rain forest, Glassman 2386 (US) (cited by Glassman 1952:54, as P. breviramula); Sankaku-Yama, Kanehira 756 (NY, FU); Nipit, 400 m, Hatusima 10880 (FU); Mt. Seletenreh, U District, Stone 5388 (BISH).

Kanehira 756, Glassman 2328, 2530, 2844 were cited by Glassman (1952) as P. palauensis and are the basis for records of that species from Ponape.

Peperomia kraemeri C. de Candolle

Peperomia kraemeri C. de Candolle, Bot. Jahrb., 56:503, 1921 [type from Palau, Koror, Kraemer, not seen by us].— Kanehira, Enum. Micr. Pl., 304, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:12, 1938; 22:99, 1959.

Plant decumbent, rooting at nodes, almost entirely glabrous; leaves oval-ovate, surfaces glabrous or upper sparsely puberulent near base, margins minutely ciliolate near apex, obtuse at base, obtuse or acutish near apex, petioles 3–6 mm long, glabrous; spikes pedunculate, terminal and opposite uppermost leaves, to 3.3 cm long, glabrous; fruit subglobose.

Very close to P. palauensis.

Endemic to Palau, where it usually occurs on the volcanic islands.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Palau: Babeldaob: Aimion, in monte Ngarua, Tuyama in 1937 (TI); Gatulei-to, Ailai-son, Hosokawa 7305 (A, in part). Koror: de Candolle, 1921:503. Ngarakabesang: Tuyama in 1939 (TI). Urukthapel: Middle of northeast coast of Magaiald (north arm of Urukthapel Island), southwest shore of Malakal Harbor, 1 m, Fosberg 25869 (Fo).

Recorded with doubt from the Marianas by Yuncker, 1938:12–13, but see P. mariannensis for this record.

Peperomia kusaiensis Hosokawa

Peperomia kusaiensis Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:120, 1935 [type from Kusaie, Hosokawa 6278].— Yuncker, Occ. Pap. Bishop Mus., 14:24, 1938; 22:107–108, 1959.

Fleshy decumbent to ascending low herb, rooting at nodes, rather shaggy pubescent with long straightish spreading hairs; leaves alternate, broadly elliptic to broadly oval or orbicular, 1–2 (rarely 2.5) cm long, 1–1.5 cm wide, obtuse to acutish at base and apex, strongly ciliate, sparsely hirsute on both faces, obscurely 3-nerved, petioles 2–5 mm long, pubescent; spikes terminal, 1–2 cm long, sparsely flowered below, on shorter sparsely pilose peduncles; fruits globose, about 0.5 mm diameter.

Very closely related to P. glassmanii and P. breviramula, differing from both in the character of the pubescence, possibly closer to P. glassmanii because of the small fruits. How constant this is in either species is uncertain, as very few fruits are available.

Endemic to Kusaie, where it has only been found growing as an epiphyte on the mossy tree trunks in the cloud forest (Eugenia-Astronia forest).

CAROLINE ISLANDS.—Kusaie: Takamatsu 566 (BISH); Mt. Buache (Matante), Hosokawa 6278 (type in Herb. Taihoku, not seen, A, isotype); Hatusima 11180 (FU); Kanehira 4231 (FU); Fosberg 26604 (US, BISH, Fo, NY, L, DPU)

Peperomia leptostachya Hooker & Arnott

Peperomia leptostachya Hooker & Arnott, Bot. Beechey Voy., 96, 1832 [type from the Sandwich Islands, Lay & Collie, not seen by us.].—Yuncker, Occ. Pap. Bishop Mus., 14:22, 1938; 22:104, 1959.

Erect, fleshy, pubescent herb, leaves opposite or whorled, lower ones tending to be reflexed and caducous in dry seasons and situations, oval, oval-obovate, or rarely broadly ovate, obtuse to acutish at both ends, petioles up to 1 cm long, usually shorter; spikes terminal or in upper axils, pedunculate, slender, elongate, to 6 (or 10) cm or more, peduncles hirtellous, rachis glabrous; fruit globose-subobovoid, 0.9 mm long.

Hawaii and Southeastern Polynesia westward at least to Fiji and the New Hebrides; in Micronesia known only from Palau, where it is uncommon on limestone rocks at low elevations.

CAROLINE ISLANDS.—Palau: Koror: On elevated coral rock, Kanehira & Hatusima 4416 (FU); small unnamed island near Koror, Kanehira 243 (FU, NY).

Peperomia mariannensis C. de Candolle

Peperomia mariannensis C. de Candolle, Prodr. 16(1):442, 1869 [type from “Ins. Mariannis,” Gaudichaud].—Safford, Contr. U.S. Nat. Herb., 9:348–349, 1905.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Bryan, Plants of Guam [unpub., 19465].—Yuncker, Occ. Pap. Bishop Mus., 14:15–17, 1938; 22:99–100, 1959.—Fosberg, Phytologia, 13:239, 1966.—Stone, Micronesica, 6:244–245, 1971.—Fosberg, Falanruw, and Sachet, Smithsonian Contr. Bot., 22: 20, 1975.

Succulent herb, erect to decumbent, or when epiphytic, pendent; leaves alternate or opposite, fully developed ones with blades 2.5–4.5 cm long, 3–5 nerved from base, elliptic to oval-elliptic, acute to obtuse at both ends, strongly punctate beneath, dots dark or obscure, glabrous except for slight ciliation near apex, petiole 2–9 (rarely 12) mm long; spikes pedunculate, terminal, opposite upper leaves, or rarely in upper axils, well-developed ones 4–8 cm long, sparsely flowered near base, basal portions fruiting when apical parts are in anthesis or bud; stigmas pubescent; fruit depressed, globose, papillate.

The type, collected in the Marianas by Gaudichaud, was in the Berlin herbarium and was doubtless lost in the bombing of this herbarium during World War II. A drawing of it in the Geneva Herbarium shows mostly opposite leaves with blades ovate, the larger ones rather acutish, the smaller definitely obtuse. “Coll. Gaudichaud. h. Berol.” A specimen in Paris, “Iles Mariannes,” Gaudichaud 216 might be a duplicate of the type.

Peperomia guamana has always been maintained as a separate species from P. mariannensis, and several other species have been described from the Marianas population of Peperomia. Individuals have also been referred to species from other areas, such as P. membranacea Hooker & Arnott, P. kraemeri C. de Candolle, P. pellucidopunctulata C. de Candolle. There is certainly a considerable range of variation in a number of characters, but there seems to be little correlation between these variations. This has been discussed elsewhere (Fosberg, 1966:239) with the conclusion that only one native species is represented in the Marianas. Its correct name is P. mariannensis C. de Candolle.

The occasional plants of this species with hirtellous stems were placed in a separate variety by Yuncker. Although such plants seem to be more frequent in Saipan, and possibly also occur on Tinian, the glabrous form is also there, indicating that the hirtellous plants probably do not form a separate population, but rather a sporadically occurring form. Since the difference is only slight, it seemed best to reduce var. saipana to the rank of forma.

As Yuncker has pointed out, plants of this species from the Northern Marianas tend to be larger in stature than those from Guam. However, this is only a tendency and is correlated with nothing else. We agree with Yuncker in not regarding it as meriting taxonomic recognition. One of these specimens, Falanruw 1923, from Alamagan, has the more obovate leaves and longer peduncles of P. ponapensis. We prefer, until the Alamagan population can be studied critically in the field, to regard this as an aberrant individual of P. mariannensis.

Distinction from P. kraemeri is a more difficult problem. About the only distinctions seem to be the smaller generally obtuse leaves and shorter spikes of P. kraemeri. However, we have had little chance to study the glabrous Palau material referred here. It seems also very close to P. palauensis, so that reduction might require bringing the latter in, too. For the present these may be regarded as three very closely related species. Plants from Palau referred by Yuncker to P. guamana seem to fit in P. kraemeri with no difficulty and are so disposed of here.

Endemic in the Marianas, where two forms are known.

Key to Forms of Peperomia mariannensis

Stems glabrous .........................................................f. mariannensis

Stems sparsely hirtellous ..............................................f. saipana

Peperomia mariannensis C. de Candolle f. mariannensis

Peperomia mariannensis C. de Candolle in de Candolle, Prodr., 16(1):442, 1869.

Peperomia membranacea sensu Miquel, Syst. Pip., 120, 1843 [non Hooker & Arnott, Bot. Beechey’s Voy. 91, 1841 (1832)].

Peperomia pellucidopunctulata sensu von Prowazek, Deutschen Marianen, 66–115, 1913 [non Elmer, Leafl. Phil Bot., 3:760, 1910].

Peperomia guamana C. de Candolle in Merrill, Phil. Jour. Sci. Bot., 9:72, 1914 [type from Guam, McGregor 629].—Kanehira, Enum. Micr. Pl., 303, 1935.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119–121, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:13, 1938; 22:99–101, 1959—Bryan, Plants of Guam [unpub., 1946?].

Peperomia hoeferi C. de Candolle, Bot. Jahrb., 56:505, 1921 [type from Saipan, Höfer 38, not seen by us.]—Kanehira, Enum. Micr. Pl., 303–304, 1935.

Peperomia mariannensis Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:120–121, 1935 [type from Tinian, Hosokawa 7784, not seen by us] [non C. de Candolle, 1869, based on different type].

Peperomia tinianensis Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:121, 1935 [in syn.].

Peperomia ladronica Hosokawa, Trans. Nat. Hist. Formosa, 25:268, 1935 [nom. nov. for P. mariannensis Hosokawa, non C. de Candolle].

Peperomia kraemeri sensu Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:121, 1935 [non C. de Candolle, 1921].

Stems entirely glabrous.

Plants from the Marianas referred to P. kraemeri C. de Candolle by Hosokawa (1935:121) and Yuncker (1938:12–13) do not differ in any essential respect from P. mariannensis as here circumscribed and are now referred to that species. Likewise, of the Palau specimens annotated in 1955 as P. guamana by Yuncker (1959), the glabrous ones are referred to P. kraemeri and those with appressed hirtellous stems and leaves are P. palauensis C. de Candolle.

USES.—Used against furry tongue; also against fever (Marianas: Fritz acc. von Prowazek, 1913:66).

VERNACULAR NAMES.—

podpod (Marianas: Fritz acc. von Prowazek, 1913:66)

podputput (Marianas: von Prowazek, 1913)

popudpod (Marianas: Fritz acc. von Prowazek, 1913:66)

potpot (Marianas: von Prowazek, 1913)

put pu put (Alamagan, Falanruw 1923)

potpot (Saipan: Kanehira, 1935)

potpotpot (Saipan: Höfer in de Candolle, 1921)

podpod palauan (Guam: Costenoble 1196, Merrill, 1914)

porporput (Guam fide A. Taitano, Evans 1588)

potpupot (Guam: Whiting XI)

pot puput (Guam: Fosberg 35516)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Miquel, 1843:120 (Gaudichaud specimen as P. membranacea Hooker & Arnott); von Prowazek, 1913:115 (as P. pellucidopunctulata C. de Candolle); Yuncker, 1959:99–100 (“collected by Gaudichaud in the Mariana Islands”); s.l. Gaudichaud 216 (P).

Agrigan: Canyon on east side, Fosberg 31621 (US, BISH, Fo, NY, DPU).

Pagan: Hosokawa 7987 (A).

Alamagan: Hosokawa 9908 (BISH, A); Partido village, Fosberg 31678 (US, BISH, Fo, DPU); Anderson 408 (US, BISH, Fo, NY, DPU, L); Fosberg 31674 (US, BISH, Fo, NY, DPU); trail from south-southwest coast, 1025 ft [310 m], Falanruw 1923 (US).

Sarigan (Saligan, Sariguan): Kanehira 2171 (NY, FU); forest above village, 250–300 m, Evans 2381 (US, BISH, Fo, NY); near anchorage, northwest coast, 700 ft [210 m], Falanruw 1797 (US), 325 ft [100 m], Falanruw 1746 (US).

Saipan: Kanehira 986 (NY, FU); s. coll. s.n. (TI); Tuyama in 1937 (TI); “Papeng, Saipan,” Hosokawa 8023 (in part) (BISH); Charlan-Tarhoho, 200 ft [60 m], Hosaka 3008 (US, BISH, Fo, DPU).

Tinian: Hosokawa 7823 (BISH); Tuyama in 1937 (TI); “in a primary forest near Lion-rock,” Hosokawa 7784 (type of P. mariannensis Hosokawa, Herb. Taihoku Imp. Univ., not seen) (Hosokawa, 1935:120–121).

Agiguan: Yuncker, 1959:100; north end, Kondo in 1952 (BISH).

Rota: Necker R71 (US); Yuncker, 1959:100; sabana, 1600 ft [490 m], Stone 5198 (UG); Mt. Kokoa near Tatacho, Hosokawa 7665 (BISH, A); middle-level plateau above Tataacho Point, 150–250 m, Evans 2054 (US, BISH, Fo, K, L, MO).

Guam: Nelson 13 (US); Costenoble 1196 (US); Merrill, 1914:72 (citing R. C. McGregor 629); Ritidian Point, 400 ft [120 m], Hosaka 3093 (US, BISH, Fo, NY, L, DPU), 180 m, Bryan 1174 (BISH), 50–150 m, Bryan 1156 (BISH); Ritidian Point Plateau, Stone 4713 (US, UG); Tarague Beach, 10 m, Anderson 248 (US); Upi (northeast point) 175 m, Bryan 1265 (BISH); Anao, ENE Mt. Santa Rosa, 140 m, Fosberg 35516 (US, BISH, Fo, NY, L, DPU); above Anao Point, Moran 4540 (US, BISH, NY); southwest of Anao Point, 200 m, Fosberg 31939 (US); northeast of College of Guam, Pedrus 70 (BISH); above Pago Bay, north of College of Guam campus, Stone 4136 (US, UG), Stone 4406 (BISH, UG); one mile east of Barrigada, coral shaded by trees, 400–600 ft [120–180 m], Moore 48 (US); north end of Tumon Bay area, 5–100 ft [1.5–30 m], Moore 395 (US); Machanao Distr., 110 m, Bryan 1187 (BISH, FU); Talofofo Point, 90 m, Bryan 1116 (BISH, NY); on rock on north side of Talofofo Valley, ½ mile [0.8 km] from sea, 10 m, Bryan 1044 (US, BISH); near Talofofo River, Rodin 695 (US); Asanite Bay, Talofofo, 25 m, Evans 1881 (US, UH, Fo, K, NY, BRI); Lujuna, just off Route 15, 10–15 m, Evans 1588 (US, UH, Fo, K) (stems microscopically puberulent); along the Piti-Agat road, McGregor 629 (G, type, US, isotype of P. guamana); Mt. Lamlam, 400 m, Anderson 145 (US, BISH, Fo, NY, DPU), near summit, Anderson 334 (US, BISH, Fo, DPU); ridge south of Mt. Lamlam, 385 m, Fosberg 35336 (US); Chalandao Mt., 1 km southeast of Jumujong Mangio Mt., 320 m, Fosberg 35377 (US); Umatac, Whiting XI (US).

Peperomia mariannensis f. saipana (C. de Candolle) Fosberg

Peperomia mariannensis f. saipana (C. de Candolle) Fosberg, Phytologia, 13:240, 1966.—Stone, Micronesica, 6:245, 1971.

Peperomia saipana C. de Candolle in Merrill, Phil. Jour. Sci. Bot., 9:72, 1914 [described again as new by C. de Candolle, Bot. Jahrb., 56:505–506, 1921, same type cited; type from Saipan, Fritz in 1903, not seen by us, presumably destroyed in the bombing of the Berlin Herbarium].—Kanehira, Enum. Micr. Pl., 304, 1935.

Peperomia pacifica Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935 [type from Saipan, Hosokawa 6654, not seen by us] [non Nakai, Bot. Mag. Tokyo, 36:128, 1922].

Peperomia pacificicola Hosokawa, Jour. Jap. Bot., 13(3):63, 1937 [new name for P. pacifica].

Peperomia guamana var. saipana (C. de Candolle) Yuncker, Occ. Pap. Bishop Mus., 14(2):15, 1938, 22:101–102, 1959.

Stems sparsely hirtellous.

USES.—Medicinally against headache and coated tongue (Merrill, C. de Candolle, both quoting Fritz).

VERNACULAR NAMES.—

popudpod (Saipan: Kanehira, 1935)

popudpud (Saipan: Merrill, 1914; C. de Candolle, 1921)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Saipan: Hosokawa 8023 (in part) (A); Kanehira & Hatusima 4319 (FU); Kanehira 3569 (FU); C. de Candolle in Merrill, 1914:72 (as P. saipana, citing Fritz (Herb. Berol.), type, not seen, probably destroyed); Hosokawa, 1935:119, citing Hosokawa 6654 (type of P. pacifica Hosokawa, non Nakai, in Herb. Taihoku Imp. Univ.), 6632, 6663 (none of these seen by us); Yuncker, 1938:15 and 1959:101–102 (as P. guamana var. saipana).

Tinian: Okatani 54 (FU).

The type, Fritz in 1903, was presumably destroyed in the bombing of the Berlin Herbarium. The Arnold sheet of Hosokawa 8023 is somewhat hirtellous and goes here. The sheet in the Bishop Museum is glabrous and is referred to f. mariannensis.

Peperomia obtusifolia (L.) A. Dietrich

Peperomia obtusifolia (L.) A. Dietrich in L., Sp. Pl., ed. 6, 1:154, 1831.

Piper obtusifolium L., Sp. Pl., 30, 1753.

Fleshy glabrous herb with ascending glabrous stems; alternate obovate petiolate leaves, rounded at apex, contracted or cuneate at base, blades to 6 × 4 cm or larger, petiole to 3 cm, spikes terminal, solitary or paired, pedunculate, to 15 cm long or more, fruit ellipsoidal, beaked, beak tending to be hooked.

A form with broad leaves and beak of fruit only curved, is often separated as P. magnoliaefolia (Jacquin) A. Dietrich. Micronesian material seen is sterile. The Rota plant could be this, but the two are here not separated as species.

MARIANAS ISLANDS.—Rota: Songsong village, planted, 5–10 m, Evans 2261 (US).

Guam: In pot, 1963, seen by Fosberg but not collected.

MARSHALL ISLANDS.—Likiep Atoll: Likiep I. planted in pot, Fosberg 36724 (US).

Peperomia palauensis C. de Candolle

Peperomia palauensis C. de Candolle, Bot. Jahrb., 56:505, 1921 [type from Palau, Koror, Ledermann 14102].—Kanehira, Enum. Micr. Pl., 304, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:20–21, 1938; 22:104–105, 1959—Glassman, Bishop Mus. Bull., 209:54, 1952.

Decumbent fleshy plant, rooting at nodes, stems moderately to closely appressed puberulent, rarely more prominently hairy; leaves mostly alternate, oval to obovate, apex obtuse to acutish, base mostly obtuse, blade 2–4 cm long, sparingly puberulent below, obscurely so above, rarely prominently so on both faces (Stone 4552), petiole 3–7 mm long; spikes pedunculate terminal and opposite upper leaves, up to 7 cm long, peduncle to 1 cm, puberulent, rachis obscurely puberulent or glabrous; fruit globose or subglobose, 0.8 mm long.

The type was collected on “Korror, bei 10–100 m ü.M., auf Kalkfelsen (Ledermann n. 14102)” in 1914. No mention was made of where it was deposited, but it was doubtless lost in the bombing of Berlin in World War II. A fragment in the de Candolle Herbarium, Geneva, has 2 leaves, a piece of stem, and a part of a spike. The leaves are oval, slightly acuminate, very minutely appressed puberulent and finely brown puncticulate, especially beneath, ciliate toward apex; petiole and stem strongly appressed pubescent; the spike has the rachis only very slightly puberulent.

Peperomia palauensis C. de Candolle var. palauensis

Stems appressed puberulent, leaves broadly ovate, usually obtuse at base, rachis of spike puberulent, rarely glabrous or almost so.

Probably endemic to Palau, found commonly on limestone islands. Reported from Saipan but these records are here assigned to P. mariannensis. Reported also from Ponape, but Ponape records are assigned to P. glassmanii. Yuncker (1959:104) refers Kanehira 4644 to P. leptostachya, but the appressed pubescence suggests P. palauensis.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Palau: Koidzumi in 1915 (TI); Todaiyama, Kanehira 4644 (FU); Makarakol, Hosokawa 9235 (A). Babeldaob: Arumizu, Hosokawa 9083 (A); Gatulel-tô, Ailai-son, Hosokawa 7305 (A, part). Koror: Tuyama in 1939 (TI); C. de Candolle, 1921:505 (citing Ledermann 14102, type, fragment in G seen by us); Armijt, Tuyama in 1939 (TI); in insulis in Sinu Iwayama, Tuyama in 1939 (TI). Aulupse’el (Oropusyakaru-tô, Aurapushekaru): Aulupse’el I., just south of Koror, south side, Maluker Bay, Risong, 2 m, Fosberg 47566 (US, BISH); Du’ebachel Beach, 10 m, Evans 586 (US, UH, Fo) (unusually densely pubescent); Hosokawa7465 (BISH, A), 9112 (BISH, A); northwest end of island; just south of Koror, Stone 4552 (UG). Urukthapel: Magaiald (north arm of Urukthapel I.), southwest shore of Malakal Harbor, 1 m, Fosberg 25869a (Fo); west peninsula, Fosberg 32210a (US). Ngeanges (Gaiangas): west of southern point of Urukthapel, 2–25 m, Fosberg 25843 (US, BISH, Fo, NY, L, DPU). Eil Malk: Makarakaru, Tuyama in 1937 (TI).

The puberulence on the rachis of the spike seems to be almost always present, but Fosberg 32210a, in other respects this species, seems to lack it.

Peperomia palauensis var. occidentalis Fosberg, new variety

Caulis prostratus, ramis erectis, conspicue puberulis trichomatibus non valde adpressis, spicarum rachibus glabris.

Main stem prostrate, rooting at nodes, leafy branches erect, stems prominently hirtellous with spreading to somewhat inwardly curving hairs, leaves broadest at middle or distally, ciliolate only toward the obtuse to acutish apex, acute at base, petiole to 10 mm long; spikes with hirtellous peduncle, glabrous rachis, not very densely floriferous, quite sparsely so toward base.

Characterized by spreading to somewhat incurved hairs on stems, leaves acute at base, petioles 6–10 mm long; sparsely flowered glabrous or almost glabrous spikes.

VERNACULAR NAMES.—

Rumriuwafathu (Sonsorol: Berry 18)

rterül (Palau: Salsedo 40)

CAROLINE ISLANDS.—Palau: Koror: Limestone rock island in Iwayama Bay, Salsedo 40 (US); Ngarmid, 30 m, Fosberg 42486 (US, UH, Fo, K, MO, L).

Sonsorol I.: Northern part, on rocks, Berry 18 (US, type).

Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth

Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth, Nov. Gen. et Sp. Pl., 1:64, 1815.—Merrill, Phil. Jour. Sci. Bot., 9:72, 1914.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Kanehira, Enum. Micr. Pl., 304, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:9–10, 1938; 22:97–98, 1959.—Tuyama, Kagaku Nanyô 4:16, 1941 — Bryan, Plants of Guam [unpub., 1946?].—Glassman, Bishop Mus. Bull., 209:54, 1952—Stone, Micronesica, 6:245, 1971.

Piper pellucidum L., Sp. Pl., 30, 1753 [type from “America calidiore”].

Peperomia bilineata sensu Volkens, Bot. Jahrb., 31:401, 1901 [non (Blume) Miquel, Nova Acta Acad. Nat. Cur., 19 (Suppl. 1):485, 1843].

Peperomia pellucida var. obtusifolia Koidzumi, Bot. Mag. Tokyo, 29:248, 1915 [nom. nud.].

Peperomia yapensis C. de Candolle, Bot. Jahrb., 56:504, 1921 [type from Yap, Volkens 342, not seen by us].—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Kanehira, Enum. Micr. Pl., 304, 1935.—Hosokawa, Jour. Jap. Bot., 13:201, 1937; Bull. Biogeogr. Soc. Jap., 7:188, 1937b.

Piper bilineata Miquel ex Kanehira, Enum. Micr. Pl., 305, 1935 [in syn.; error by Kanehira].

Peperomia lineata Miquel ex Yuncker, Occ. Pap. Bishop Mus., 22:99, 1959 [probably an error by Yuncker for P. bilineata Miquel].

Slender erect glabrous, stems translucent; leaves thin, cordate, mostly opposite, strongly nerved, on slender petioles; spikes pedunculate, terminal and in upper axils, elongate, strongly exceeding leaves; fruits longitudinally ribbed.

Native of tropical America, but widely introduced in the Old World tropics; in Micronesia in Guam, the high, and a few low, islands of the Carolines, and Jaluit in the Marshalls. On mossy rocks, moist soil, and the bases of coconut trees, mainly in lowlands, common around villages and in plantations, disturbed shaded places.

VERNACULAR NAMES.—

podpod lahe (Guam: Costenoble 1197)

rtertil (Palau: Tuyama, 1941, citing A. Krämer)

rtertiiel (Palau, Babeldaob I.: Fosberg 32446)

makel ni pan (Yap: Falanruw 755)

rafung (Yap: Wong 318)

rum (Sonsorol: Berry 20)

pukuson (Truk: Wong 178)

opul kongkong (Eauripik: Fosberg 47128)

USES.—In Sonsorol the plant is pounded and the juices are used on cuts, Berry 20.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MICRONESIA.—Kamiya Shinsaburo (Matsutaka) 132 (TI).

MARIANAS ISLANDS.—Guam: Costenoble 1197 (US); G.E.S. 234 (US, BM, NY); Merrill, 1914:72; Agaña, Seale in 1900 (BISH).

CAROLINE ISLANDS.—Palau: Kanehira & Nisida 3 (FU); Yamada in 1925 (TI); G. Koidzumi in 1915 (TI); Akoru-kaigan, Hosokawa 7101 (BISH, A); Arukoron-son, Konrai, Hosokawa 7082 (BISH, A) (both cited by Hosokawa, 1937a: 201 as P. yapensis). Babeldaob (Babeltaob, Babelthuap): Koidzumi in 1915 (TI); Aimiriik, Tuyama in 1937 (TI); west coast of island, Ngeremetengel, in village on bare rocks near stream, 1 m, Fosberg 32446 (US, BISH); Molegojok (Meligeok), Tuyama in 1939 (TI); Ngarsul, Tuyama in 1937 (TI); Aimion, Tuyama in 1937 (TI); Ngiual, Hosaka 3384 (US, BISH, DPU). Koror: Yuncker, 1938:9–10. Angaur: 25 m, Fosberg 25910 (US, BISH, Fo, DPU, NY); Koidzumi s.n. in 1915 (TI); s. coll. s.n. in 1930 (TI); s. coll. s.n. (TI).

Sonsorol: Village area along the tracks, Berry 20 (US).

Yap: Wong 318 (US, BISH, Fo, NY, L, DPU); Kanehira & Hatusima 4327 (FU); Kamiya? s.n. (TI); Volkens, 1901:401 (citing Volkens 312 as P. bilineata); de Candolle, 1921:504 (citing Volkens 342 as type of P. yapensis); Mabo, Hosokawa 8824 (Fo); Tornii I., 100 ft [30 m], Hosaka 3257 (US, BISH); in oppido Dogol, Tuyama in 1939 (TI), Tuyama 7217 (UG); Kanif, Takamatsu 1910 (BISH); near Catholic Mission, Blackburn 243 (US); Balabat, 50 ft [15 m], Falanruw 755 (US); Tabunfi village, southeast Yap, 5 m, Cushing & Mitag 553a (US, UH); Ma’lai Village, Mun. Kanifay, 3 m, Fosberg 46343 (US, Fo).

Eauripik: Eauripik Islet, 1–2 m, Fosberg & Evans 47128 (Fo).

Truk: 6 ft [2 m], Wong 178 (US, Fo, BISH). Dublon: 50 ft [15 m], Hosaka 2758 (US, BISH, Fo, NY, L, DPU). Tol: Hosokawa, 1937a:201, and 1937b: 188 (citing Hosokawa 8305 as P. yapensis); Amatyan (Amatang), Hosokawa 8305 (BISH).

Ponape: Kolonia, Hatusima 10740 (FU); vicinity of Colonia, shade of banana tree, Glassman 2584 (US, BISH); Not District, Colonia, Wong 60 (US, BISH, Fo, NY, L, DPU); Net village, Salomon & George 49 (US).

Kusaie: Hosokawa, 1937a:201 (citing Hosokawa 6460 as P. yapensis); Lele, Hosokawa 6460 (A), Takamatsu 332 (BISH); Lele Islet, Glassman 2709 (US, BISH), 15 m, Wong 79 (US, BISH, Fo, NY, DPU), 1–5 m, Fosberg 26552 (US, BISH, Fo, NY, DPU).

MARSHALL ISLANDS.—Jaluit: Koidzumi in 1915 (TI); Koidzumi, 1915:248 (as var. obtusifolia).

Peperomia ponapensis C. de Candolle

Peperomia ponapensis C. de Candolle, Bot. Jahrb., 56:504, 1921.—Kanehira, Enum. Micr. Pl., 304, 1935.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935; Jour. Jap. Bot., 13:200, 1937a; Kudoa, 5:95, 1937b—Yuncker, Occ. Pap. Bishop Mus., 14:17, 1938; 22:102–103, 1959.—St. John, Pac. Sci., 2:108, 1948.—Glassman, Bishop Mus. Bull., 209:54, 1952.—Fosberg, Atoll Res. Bull., 39:7, 1955 [as Peperomia sp.]; Phytologia, 13:240–241, 1966.—Stone, Micronesica, 1:155, 1964 [as Peperomia].

Herb up to 40 cm tall, leaves and branching opposite and alternate, usually on same plant, where alternate with a tendency to an appearance of a branch opposite to a leaf, though this may be merely a matter of the original shoot being crowded aside by an axillary branch, often with a leaf-scar opposite an apparently alternate leaf; leaf blades broadly elliptic to obovate, up to 3.5 cm wide and 5.5 cm long, 5 or 3 nerved, when 5, the outer pair weaker, all originating at base, apex obtuse to bluntly acute or slightly acuminate, petiole up to 15 mm long, lower leaves often reduced and tending to be orbicular; spikes pedunculate, terminal and in upper axils of opposite leaves, leaf-opposed to alternate leaves, with some tendency to paniculate arrangement, up to 7 cm long, more loosely flowered toward base, some tendency toward earlier flowering at base of spike; fruit globose, notably papillate-glandular.

This species is here given a much broader circumscription than by previous authors. The reduction of P. volkensii C. de Candolle, P. gibbonsii C. de Candolle, and P. trukensis Yuncker has been discussed elsewhere (Fosberg, 1966). Since that paper was published, we have examined a fragment of the type of P. gibbonsii in the de Candolle Herbarium at Geneva (Gibbons 1072). It is a single spike with several immature and mature fruits. The latter are globose. Several stigmas are bifid, the character on which P. gibbonsii was founded. Other stigmas appear entire, but these are very poorly developed. Our conclusions as to uniting this species with P. ponapensis remain unchanged.

Also at Geneva we examined a fragment of Ledermann 13986, from Ponape. This is not the type of P. ponapensis, nor was it cited with the original description, but it was so named by C. de Candolle. It appears to be this species as usually understood, with alternate, slightly obovate acutish leaves.

We have not seen authentic material of P. volkensii C. de Candolle, which was based on Volkens 2 from Kusaie and Finsch 1 from Ebon, Marshall Islands, but we have material from the exact type-locality, Lele Islet Kusaie, Fosberg 26538, which agrees well with this species. We consider that Yuncker (1959) by stating that P. volkensii is endemic to Kusaie has effectively lectotypified it.

Endemic to the eastern Carolines—Truk, Ponape, Pingelap, and Kusaie—and the central to southern Marshalls—Lae, Ailinglapalap, Arno, Jaluit, and Ebon atolls. The records from the Marianas admitted by Yuncker are here referred to the very similar P. mariannensis C. de Candolle. A specimen from Alamagan, Northern Marianas, Falanruw 1923, keys to this species and was referred there at first by us. On reconsideration, however, it could be an aberrant individual of P. mariannensis and is cited under that species. Two varieties may be distinguished.

Key to Varieties of Peperomia ponapensis

Stems completely glabrous ...............................................var. ponapensis

Stems more or less subappressed hirtellous ........................................var. trukensis

Peperomia ponapensis C. de Candolle var. ponapensis

Peperomia ponapensis C. de Candolle, Bot. Jahrb., 56:504–505, 1921 [type from Ponape, Ledermann 13914, not seen by us].—Stone, Micronesica, 1:155, 1964.

Peperomia pallida sensu Schumann, Bot. Jahrb., 9:198, 1888 [non (Forster f.) A. Dietrich, Spec. Pl., ed. 6, 1:153, 1831].

Peperomia gibbonsii C. de Candolle, Bot. Jahrb., 56:504, 1921 [type from Marshall Is., Ailinglapalap, Gibbons 1072].—Kanehira, Enum. Micr. Pl., 303, 1935.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:10–12, 1938; 22:104, 1959.—Fosberg, Phytologia, 13:241, 1966.

Peperomia volkensii C. de Candolle, Bot. Jahrb., 56:503, 1921 [type from Kusaie, Volkens 2, not seen by us].—Kanehira, Enum. Micr. Pl., 304, 1935.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:119, 1935.—Yuncker, Occ. Pap. Bishop Mus., 14:17, 1938; 22:102, 1959.—Fosberg, Phytologia 13:240–241, 1966.

Endemic to the Eastern Caroline and Marshall islands. Found mainly at very low elevations, growing on limestone and basalt rocks, frequently on walls of ruins.

USES.—“The plants are used medicinally” (Yuncker, 1959:22:103).

VERNACULAR NAMES.—

warin (Pingelap: St. John 21475, Glassman 2650)

rabikiaga (Lae: Fosberg 34027)

rapisrage (Ailinglapalap: de Candolle, 1921:504; Gibbons 1072)

drebijdreke = holding rocks (Arno: Stone 1082, Anderson 3745, Hatheway 849)

rebijrege (Jaluit: Fosberg 26734)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Ponape: s.l. Ledermann 13986 (G); de Candolle, 1921:504–505 (citing Ledermann 13914, from Matalanim, type, not seen by us); Matalanim District, Sabera, Glassman 2760? (US); Nanmatol I., Fosberg 26379 (US, BISH, Fo, NY, L, DPU), 1–3 m, Fosberg 26377 (US, BISH, Fo, NY, DPU); Nanmatol ruins, Matalanim, Hatusima 11124 (FU), Kanehira 858 (FU); Jokaazi Rock, near Kolonia, Hatusima 10976 (FU), 10977 (NY).

Pingelap: On coral stone wall in moist forest, near lagoon beach, St. John 21475 (US, BISH); Pingelap I., Glassman 2650 (US, BISH).

Kusaie: Kamiya 267 (TI); de Candolle, 1921:503, Yuncker 1938:17, 1959:102 (all as P. volkensii, citing Volkens 2, type from Lele I., not seen by us); Lele I., 1–5 m, Fosberg 26538 (US, BISH, Fo, NY, L, DPU); Rero-Taonsak, Hosokawa 6177 (A), near Maarem, 200 m, Hatusima 11144 (FU).

MARSHALL ISLANDS.—Lae: Lae I., very rare, one colony on rocks in Barringtonia forest, Fosberg 34027 (US, BISH, Fo, NY, L, DPU).

Ailinglapalap: de Candolle, 1921:504, Yuncker, 1938:10–12, 1959:104 (all as P. gibbonsii, citing Gibbons 1072, type); Gibbons 1072 (G), fragment seen by us.

Arno: West side Taklepeg I. (E. Taklep), Stone 1032 (Fo); Kijbwe I., Anderson 3745 (US, BISH, Fo, NY, L, DPU); Arno I., Hatheway 791 (US, BISH, Fo, NY, L); Takleb Ej I., Hatheway 849 (US, BISH, Fo, NY, L).

Mili: Stone, 1964:155, casual record, probably this species.

Jaluit: Imruj (Imrodj) I., Fosberg 26734 (US, BISH, Fo, NY, L, DPU).

Ebon (Boston): Schumann, 1888:198 (as P. pallida, citing a Finsch specimen not seen by us); de Candolle, 1921:503; Kanehira 1935:304 (both as P. volkensii, citing Finsch n.l, not seen by us).

Peperomia ponapensis var. trukensis (Yuncker) Fosberg

Peperomia ponapensis var. trukensis (Yuncker) Fosberg, Phytologia 13:241, 1966 [type from Truk, Tol I., Takamatsu 21].—Hosokawa, Jour. Jap. Bot., 13:200–201, 1937a; Bull. Biogeogr. Soc. Jap. 7:188, 1937b [both as P. ponapensis C. de Candolle].

Peperomia trukensis Yuncker, Occ. Pap. Bishop Mus., 14:23–24, 1938; 22:108, 1959.

Stems, petioles, leaf blades and peduncles more or less subappressed hirtellous, blades ciliate almost all around.

Some specimens, such as Fosberg 24550, are so densely hirtellous that they could be mistaken for P. palauensis, but the rachis is glabrous.

Found in Truk and rarely on Ponape, on rocks at low to middle altitudes.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Truk: Yuncker, 1938:23–24, 1959:108; Hosokawa 1937a:201, 1937b: 188 (all as P. trukensis). Moen (Wara): Wara Mt., Tukuman, Hosokawa 8473 (BISH, A). Dublon (Natusima): Hosokawa 8360 (US, BISH, A); upper ridges and top of Mt. Tolomen (Tolowan), 200–360 m, Fosberg 24550 (US, BISH, Fo, DPU). Tol: s.l., Stone 5336 (BISH); Suyoto, Takamatsu 21 (BISH, type); Uriribot, Hosokawa 8283 (BISH, A); 8251 (BISH, A).

Ponape: Yuncker, 1959:108 (as P. trukensis); Jokaj (Gyokazi, Jokaazi), Hosokawa 6054 (BISH, A); Hatusima 10977, part (FU).

Piper L.

Piper L., Gen. Pl., ed. 5, 18, 1754 [=1753].

Chavica Miquel, Syst. Pip., 222, 1843.

Macropiper Miquel, Comment. Phytogr. 35, 1840.

Shrubs or climbers, mostly aromatic, with alternate, simple, frequently stipulate leaves with expanded bases, scars extending around nodes; spikes or racemes axillary or leaf-opposed (may be paniculate or umbellate outside Micronesia); flowers bisexual or unisexual and then often dioecious, surrounded by 3 scale-like or peltate bracts, these separate or connate; stamens several; stigmas 2–4 or more, fruit a fleshy drupe, often immersed in the swollen fleshy rachis or connate with the bracts.

Pantropical, a few species in Micronesia. Common shrubs in undergrowth and small openings in forest, or climbing on tree trunks and creeping on ground. Several species are of economic importance, including the black pepper of commerce, cubebs, and the betel pepper.

Key to Micronesian Species of Piper

1. Shrub with axillary spikes .......................................................2

2. Spike 1 in an axil ..............................................P. guahamense

2. Spikes usually 2–5 in an axil ..................................................P. latifolium

1. Climbers, or shrubs with leaf-opposed spikes ..........................................................3

3. Leaves with one of the pairs of principal nerves arising well above base, climbers ..............4

4. Leaves with one strong pair of nerves arising slightly above base, usually a pair of faint ones at base, a second strong pair about 1 cm or so above the base, lower leaf-surface minutely black-punctate ..................................P. nigrum

4. Leaves with 2 pairs of principal nerves arising basally, lower leaf-surface not strongly punctate ...........................................................P. betle

3. Leaves with all principal nerves arising basally, or near base ....................................5

5. Principal nerves 5 or 7, climbers rooting at nodes .......................................6

6. Leaves commonly longer than wide, never peltate, if almost as broad as long (juvenile) then slightly puberulent beneath near base and on petiole, mature spikes commonly as long as or longer than leaves, peduncle usually longer than petiole ..... P. ponapense

6. Leaves as broad as long, often peltate, never puberulent beneath, mature spikes much shorter than leaves, especially pistillate ones, peduncle commonly shorter than petiole ............................................................................................P. fragile

5. Principal nerves 9–13, shrubs or climbers .....................................................7

7. Shrubs with orbicular-cordate leaves, rather shortly pointed, equilateral at base, spikes less than 10 cm long ..................................................P. methysticum

7. Climber with leaves broadly oblong-ovate to ovate, inequilateral at base, prominently pointed, spikes about 15 cm or more long ...................................P. hosokawae

Piper betle L.

Piper betle L., Sp Pl. 28, 1753 [type from Ceylon, Hermann, not seen by us].—Endlicher, Ann. Wien. Mus. Naturgesch., 1:164, 1835.—Christian, The Caroline Islands, 337, 350, 1899.—Volkens, Bot. Jahrb., 31:461, 1901.—Safford, Contr. U.S. Nat. Herb., 9:224, 353–354, 1905.—von Prowazek, Deutschen Marianen, 115, 1913.—Merrill, Phil. Jour. Sci. Bot. 9:72, 1914.—Kanehira, Enum. Micr. Pl., 304–305, 1935.—Okabe, Nettai Sangyô Kenkyû-sho ihô, 5:2, 1940; Jour. Anthrop. Soc. Nippon, 56:424, 1941.—Tuyama, Kagaku Nanyô, 4(1): 16, 1941.—Bryan, Plants of Guam [unpub., 1946?].—Glassman, Bishop Mus. Bull., 209:54, 1952 — Yuncker, Occ. Pap. Bishop Mus., 22:90–91, 1959.—Fosberg, Phytologia, 13:234, 1966.—Fosberg, Falanruw, and Sachet, Smithsonian Contr. Bot., 22:20, 1975.

Chavica bette (L.) Miquel, Syst. Pip., 228, 1843.

Piper betel L. ex Endlicher, Ann. Wien. Mus. Naturgesch., 1:164, 1835 [sphalm.].

Climber with broadly cordate to ovate-oblong leaves, aromatic and somewhat acrid to taste when crushed, main nerves 9, the outermost pair much weaker than the others, the innermost pair diverging from the midrib 1–2 cm above the base, more or less alternate, the rest basal; flowers dioecious, in oppositi-folious pedunculate spikes that are usually shorter than the leaves, staminate spikes with bracts imbricate, pistillate spikes much shorter and thicker than staminate, densely tomentose, about 6–8 mm thick, stamens 2, stigmas 5–7; fruits coalescent and somewhat embedded in rachis.

Found generally in the Indo-Pacific region from India as far east as the betel nut (Areca catechu L.) is chewed, that is, to New Guinea, Western Melanesia and Western Micronesia; usually planted or persisting from plantings, around old dwelling sites and gardens.

This is a very complex or variable species, as examination of the material in almost any large herbarium of tropical plants will show. No modern attempts to analyze and classify the variations have come to our attention. We have not tried to do this for the species as a whole, but have admitted three forms in Micronesia, all of which have been previously recognized as separate species or varieties, but which were placed in two varieties by Yuncker (1959:90–91). Of the two species that were based on Micronesian material, one has been reduced to lower rank, to include the broad-leafed Marianas specimens, while the other is placed in the synonymy of the typical form.

Key to Micronesian Forms of Piper betle

1. Leaves somewhat oblong-ovate, with base oblique, not deeply cordate .........................f. betle

1. Leaves broadly ovate-cordate, acuminate, base deeply cordate, not conspicuously oblique.

2. Leaves puberulent beneath, at least on nerves ..................................f. densum

2. Leaves glabrous beneath .........................................................f. marianum

Several references, as well as one specimen, examined years ago, which cannot be referred to one of the three forms, are listed below under the species. Local uses and vernacular names, likewise, are listed together under the species, with no attempt to segregate them by forms.

USES.—“Natives chew a combination of fresh P. betle leaves, a piece of the fragrant seed of the Areca palm, and a pinch of quicklime. Betle-chewing has not developed to any extent in Polynesia, where the P. methysticum beverage called kava apparently takes its place” (Yap: Yuncker, 1959: 91). “The fresh green leaves (manaon) are chewed by the natives wrapped around a fragment of Areca nut together with a pinch of quicklime” (Guam: Safford, 1905:354). “Used with lime and Areca-nut” (Guam: Perez in 1962). “Used in betel chewing” (Rota: Fosberg 31877).

VERNACULAR NAMES.—

pupúlo (Marianas: von Prowazek, 1913; Safford, 1905; Merrill, 1914)

pupúlu (Marianas: von Prowazek, 1913)

pupúlu (Guam: Seale in 1900, Safford, 1905)

pupulo (Guam: Whiting P. 10, Perez in 1962)

populo (Guam: B. C. Stone 5126)

haboi or kaboi (Sonsorol: Berry 90)

a-kabui (Palau: Okabe, 1940)

kabúi (Palau: S. Walleser in Tuyama, 1941)

kebui (Palau: D. Otobed, 1967)

gabúi (Yap: Volkens, 1901; Yuncker, 1959; Wong 484)

gabusy (Yap: Cushing 660)

gogoth (Yap: Christian, 1899)

habui (Fais, Evans 357)

tuh (Ponape: Glassman, 1952)

betel pepper: English

MARIANAS ISLANDS.—Endlicher, 1835:164 (Lesson specimen); von Prowazek, 1913:115. Sarigan: Kanehira 2161 (FU).

Piper betle L. f. betle

Piper betle L., Sp. Pl., 28, 1753 [sensu stricto].

This is very variable in leaf shape but there seems to be no order to the variation, so it is difficult to separate the glabrous Marianas plants as Opiz did in his P. marianum (vide infra), except on leaf width and prominence of cordate base. The typical form is here restricted to plants with rather oblong-ovate leaves, shallowly cordate at base.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Yuncker, 1959:90 (as P. betle).

Agrigan: Hosokawa 8009 (A), 8011 (A).

Alamagan: Anderson 418 (US); “Chalan nitiput,” s.s.w. coast, Falanruw 1929 (US).

Sarigan: Hole in savanna plateau above village, 350–375 m, Evans 2371 (US, UH, Fo, K).

Saipan: Hosokawa 6658 (A); Kanehira 892 (BISH); Kanehira, 1935:305 (as P. betle); Mt. Tagkochao, Stone 5424 (US).

Rota: Slopes above As Malote, south side of island, Fosberg 31877 (US).

Guam: G.E.S. 190 (US, BISH); Mt. Tenjo, 300 m, Bryan 1248 (US, BISH, FU); Mangilao, Perez in 1962 (UG).

CAROLINE ISLANDS.—Yap: Gagil, Cushing 660 (US, Fo).

Piper betle f. densum (Blume) Fosberg

Piper betle f. densum (Blume) Fosberg, Phytologia, 13:235, 1966.

Piper densum Blume, Verh. Batav. Gen. 11:193, fig. 18, 1826 [type from Java, Blume, not seen by us].

Piper betle var. densum (Blume) C. de Candolle in de Candolle, Prodr., 16(1):360, 1869.—Yuncker, Occ. Pap. Bishop Mus., 22:90–91, 1959.

Leaves puberulent beneath on nerves.

Found in Micronesia in the Marianas (Alamagan and Guam) and in the Carolines (Sonsorol, Palau, Yap, Fais), and said to be on Ponape.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Yuncker 1959:90–91 (as P. betle var. densum); Gaudichaud 77 bis (P).

Alamagan: Partido village, Fosberg 31667 (US, BISH, Fo, DPU); SSW coast, 500 ft [150 m], Falanruw 1914 (US).

Guam: Nelson 49 (BISH); Whiting P10 (Fo, US); Agaña, Seale in 1900 (BISH); Umatac water point, 100 m, Fosberg 35412 (US, BISH, Fo, DPU); Mt. Santa Rosa, Glassman 149 (Fo); Dededo, 100 m, Evans 721 (US).

CAROLINE ISLANDS.—Kanehira, 1935:305; Yuncker, 1959:91 (as P. betle var. densum).

Sonsorol: Berry 90 (US, UH) (puberulence only evident on very young leaves, leaf bases somewhat oblique).

Palau: Babeldaob: Ogiwaru, east coast, Takamatsu 1429 (BISH); Marikyoku (Melakiok), Kanehira 453 (FU). Angaur: East side, 3–5 m, Fosberg 31995 (US).

Yap: Volkens, 1901:461 (citing Volkens 303, not seen by us); 40 ft [12 m], Wong 484 (US, Fo); Gorror I., central plateau, 100 ft [30 m], Hosaka 3307 (US, BISH).

Fais: Lochochoy, 5 m, Evans 357 (US, Fo) (leaves not very deeply cordate).

Ponape: Glassman, 1952:54 (plant observed, not collected).

Piper betle f. marianum (Opiz) Fosberg

Piper betle f. marianum (Opiz) Fosberg, Phytologia, 13:235, 1966.

Piper marianum Opiz in Presl, Rel. Haenk., 1:159, 1828 [type from Guam, Haenke, not seen by us].—Endlicher, Ann. Wien. Mus. Naturgesch., 1:164, 1835.—Kunth, Linnaea, 13: 574, 1839; Ann. Sci. Nat. II, Bot., 14:179, 1840.—Safford, Contr. U.S. Nat. Herb., 9:354, 1905.

Piper potamogetonifolium Opiz in Presl, Rel. Haenk., 1:156, 1828 [type from Guam, Haenke, not seen by us].—Endlicher, Ann. Wien. Mus. Naturgesch., 1:164, 1835.—Safford, Contr. U.S. Nat. Herb, 9:335, 1905.—Merrill, Phil. Jour. Sci. Bot., 9:72, 1914.—Kanehira, Enum. Micr. Pl., 305, 1935.—Yuncker, Occ. Pap. Bishop Mus., 22:83, 1959.—Fosberg, Phytologia, 13:235, 1966.

Macropiper potamogetonifolium (Opiz) Miquel, Syst. Pip., 218, 221, 1843–1844.

Piper betle δ mariannum C. de Candolle, Prodr., 16(1):360, 1869 [based on P. mariannum Opiz, which is a misspelling of P. marianum Opiz].

A broad-leafed, deeply cordate glabrous form. This has seldom been collected, at least in Micronesia. Its range extends to the Bonins and probably to other parts of the area of the species.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Endlicher, 1835:164 (Haenke specimen of P. potamogetonifolium).

Saipan: Hosokawa 6658 (A).

Guam: Opiz, 1828:156, 159 [citing Haenke, types of P. marianum and P. potamogetonifolium, not seen by us].—Endlicher, 1835:164 [Haenke specimen of P. marianum].—Kunth, 1839:574, 1840, 179; Miquel, 1843–1844:218; Merrill, 1914:72 (“Collected in Guam by Haenke …”); Barrigada Village, Stone 5126 (US, BISH, UG).

Piper fragile Bentham

Piper fragile Bentham in Hooker, London Jour. Bot., 2:284, 1843 [type from New Guinea, Hinds, not seen by us].—Kanehira, Enum. Micr. Pl., 305, 1935.—Yuncker, Occ. Pap. Bishop Mus., 22:92–93, 1959—Stone, Pac. Sci., 13:100, 1959.

Piper carolinense Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:117, 1935 [type from Palau, Hosokawa 7505].—Kanehira, Kagaku Nanyô, 4:16, 1941.

Piper trukense Hosokawa, Jour. Jap. Bot., 13:200, 1937a [type from Truk, Hosokawa 8459]; Bull. Biogeogr. Soc. Japan, 7:188–189, 1937b.

Glabrous vine, rooting at nodes; leaves orbicular to broadly oval or broadly elliptic, somewhat acuminate, base rounded to cordate or peltate, peltate and cordate or rounded bases on same plant, peltate more often on sterile branches, nerves usually 7, lowest pair reduced, lowest two pairs basal or sub-basal, third pair emerging from midrib slightly above base; flowers dioecious, in spikes usually shorter than leaves, opposite them, staminate spikes slender, linear, several cm long, pistillate much shorter, oblong or linear oblong; stamens 2, anthers apparently trilocular (bilocular according to Bentham); stigmas 4–5, spreading to recurved; fruit globose, more or less fused with bracts and rhachis.

A rather uniform species. Piper trukense Hosokawa from Truk, and P. carolinense Hosokawa, from Palau, do not seem to differ significantly.

Known from New Guinea (Hinds, type), Borneo (?), Philippines, and Micronesia; in Micronesia known from the Western and Central Carolines, including Truk, Palau, and many of the atolls. Common, covering ground and tree trunks in dense lowland forest.

USES.—Used as “baby medicine,” for bruises, as tonic; small green flowers for maremar [garlands] (Ifaluk: Abbott and Bates 86).

VERNACULAR NAMES.—

kesebebúi (Palau: Tuyama, 1941:16)

walitogu (Woleai: Alkire 81)

hatogobwei (Woleai: Alkire 17)

gatogobwei (Ifaluk: Abbott and Bates 86)

hata ho bue (Ifaluk: Fosberg 47216)

atobuei (Satawal: Fosberg 46907)

adogobwe (Namonuito: Stone 2079)

anes (Nomwin: Evans 1115)

atukupui (Nomwin: Fosberg 24580)

atukupwi (Nomwin: Evans 1066)

ootikk (Truk: Hosokawa, 1937b: 189) (Moen, Dublon, Udot, Tol).

ötik (Truk: Moen: Pelzer 85)

enes (Truk: Wong 170)

ëres (Truk: Pis: Fosberg 24688)

atopui (Etal: Anderson 2063); (Satawan: Anderson 1104)

atoopwei (Namoluk: Marshall 15)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Palau: Kanehira 466 (FU); Kasioru, Takamatsu 1500 (BISH). Babeldaob: Airai, ¼ mi [0.4 km] behind airport terminal, Salsedo 223 (US). Koror: Kanehira 341 (FU); Tuyama in 1939 (TI); Meiung, Salsedo 313 (US). Urukthapel (Urktable): “in a forest on coral islet of Urktable (Todai-san),” Hosokawa 7505 (BISH, A, isotypes of P. carolinense Hosokawa, cited by Hosokawa, 1935:117); Tôdai-yama, Kanehira & Hatusima 4671 (FU). Malakal: Ngerungsiil, Hardy 57 (US, Fo). Ng’edobel: North of Eil Malk I., Fosberg 50566 (US, BISH, Fo, NY, Y). Bairakaseru (North of Peliliu): Koch 1152 (Fo). Peliliu: Hosokawa 9219 (A, BISH), 9212 (BISH); Kanehira & Hatusima 4782 (FU); Takamatsu 1789 (BISH). Angaur: Takamatsu 1806 (BISH); ridge on northwest corner of island, Fosberg 25948 (US, BISH, Fo, NY, DPU).

Sonsorol: Berry 9 (US).

Woleai: Utagal I., 12 m, Wong 21 (US, BISH, Fo, NY, DPU); Falalis I., Alkire 81 (US, Fo), Alkire 17 (US).

Faraulap: 2 m, Fosberg & Evans 47330 (US, Fo).

Ifaluk: Falarik, east coast, Abbott and Bates 86 (US, BISH); Ifaluk I., 2 m, Fosberg 47216 (US, UH, Fo, NY, L), Fosberg 47214 (US, BISH, Fo, NY, L, P, TI, A, QLD, K, G).

Satawal: interior of island, 2 m, Fosberg 46907 (US, Fo).

Puluwat: 1–3 m, Niering 764 (US).

Namonuito: Ulul I., Stone 2079 (BISH); Ono, village and vicinity, 0–3 m, Evans 1013 (US); 1024 (US); Piseras Island, beach and rampart, 0–3 m, Evans 910 (US, UH, Fo, K); main taro pit, 3–5 m, Evans 888 (US, BISH, Fo, MO); Magur, center of island, 3–5 m, Evans 949 (US, UH, Fo); 941 (US, UH, Fo, K); Onari I., village and vicinity, 0–3 m, Evans 984 (US).

Murilo: Rui I., interior of island, 3–5 m, Evans 1198 (US).

Nomwin: Nomwin I., 1–2 m, Fosberg 24580 (US, BISH, DPU); woods behind village, 0–3 m, Evans 1066 (US); Fananu I., interior of island, 3–5 m, Evans 984 (US).

Truk: Wong 170 (US, Fo); Huyôtô, Kanehira 4262 (FU); Melot, Hosokawa 8355 (A); Tadiu, Hosokawa 8396 (A, BISH). Moen (Wara): Mt. Tukuman, Hosokawa 8459 (US, BISH, A, isotypes of P. trukense, cited by Hosokawa, 1937a:200, 1937b: 188–189 as from Mt. Tratyau); Pelzer 85 (US, BISH, Fo); east end of island, 100–200 m, Evans 774 (US, UH, Fo), 778 (US), 791 (US, BISH), 792 (US, UH, Fo); west coast, Muam village, 15 m, Fosberg & Pelzer 26019 (US, BISH, Fo, NY, L, DPU). Dublon (Natusima): Hosokawa 6521 (A); Mt. Troman, Trowasi, Hosokawa 8500 (US, A, BISH). Tol: Hosokawa 8293 (cited by Hosokawa 1937b: 189, not seen by us); Mt. Tumuital (Uiniboet), 200–460 m, Fosberg 24451 (US, BISH, Fo, DPU). Udot: Mt. Witonnap, Hosokawa 8355 (cited by Hosokawa, 1937b: 189, not seen by us, perhaps the same as specimen of the same number cited above). Pis: 1–2 m, Fosberg 24688 (US, BISH, Fo, NY, DPU); center of island, 3–5 m, Evans 844 (US), 846 (US), 836 (US).

Nama: center of I., 3–5 m, Evans 1319 (US) (juvenile).

Namoluk: Leor village, Anderson 913 (US, BISH, Fo, DPU); 5 m, Mac Marshall 15 (US).

Etal: Etal I., Anderson 2063 (US, BISH, Fo, DPU).

Lukunor: Piafö I., Anderson 2198 (US, BISH, Fo, DPU); Saponöch I., Anderson 2204 (US, BISH, Fo, NY, DPU).

Satawan: Satawan I., Anderson 1104 (US, BISH, Fo, NY, L, DPU); Moch I., Anderson 1106 (US, BISH, Fo, NY, DPU).

Piper guahamense C. de Candolle

Piper guahamense C. de Candolle in de Candolle, Prodr., 16(1):336, 1869 [type from Marianas, doubtless Guam, Gaudichaud 16].—Safford, Contr. U.S. Nat. Herb. 9:354— 355, 1905—Merrill, Phil. Jour. Sci. Bot., 9:71, 1914.—Kanehira, Fl. Micr., 418, 1933.—Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:117, 1935.—Kanehira, Enum. Micr. Pl., 305, 1935.—Bryan, Plants of Guam [unpub., 1946?].—Yuncker, Occ. Pap. Bishop Mus., 22:86–87, 1959.—Fosberg, Phytologia, 13:235–236, 1966.—Fosberg, Falanruw, and Sachet, Smithsonian Contr. Bot., 22:21, 1975.

Piper latifolium Gaudichaud ex Kunth, Linnaea, 13:575, 1839 [type from the Marianas, Gaudichaud 163] [non L.f., Suppl. Pl., [468], 1781].

Macropiper latifolium sensu Miquel, Syst. Pip., 219, 1843–1844 [non (Forster f.) Miquel, Comment. Phytogr., 36, 1840].

Macropiper methysticum Miquel, Syst. Pip., 217, 1843–1844 [non (Forster f.) Miquel, Comment. Phytogr., 36, 1840].

Piper macgillivrayi sensu von Prowazek, Deutschen Marianen, 115, 1913 [non C. de Candolle in Seemann, Fl. Vit., 262, 1868].

Erect to somewhat reclining shrub 1–3 m long, nodes prominent; leaves large, orbicular-cordate, slightly acuminate, 9- or 11-nerved from base, petioles shorter than blades, lower half expanded-sheathing; flowers dioecious or rarely monoecious, in solitary axillary pedunculate spikes, these usually shorter, sometimes longer, than the leaves, the staminate more slender than the pistillate, occasional spikes mixed, staminate and pistillate, peltate bracts imbricate when spikes are young; stamens 3, oval, bilocular; ovaries subtruncate at apex, crowded on spikes, rarely a few on staminate spikes (Rodin 613), stigmas 3–4, fleshy, tomentose; fruits connate, red.

Marianas records of Piper macgillivrayi C. de Candolle by von Prowazek (1913), Piper latifolium Gaudichaud by Kunth (1839), Macropiper latifolium Miquel by Miquel (1843–1844), and Macropiper methysticum Miquel by Miquel (1843–1844) are doubtless based on specimens of P. guahamense, although we have not seen these specimens, except that of Gaudichaud. None of these species is otherwise known from the Marianas.

The species is endemic to the Marianas, where it grows as an undergrowth shrub, in and around the edges of wet forests.

Two forms are known, one with leaves prominently puberulent beneath especially on nerves, petioles, peduncles, and young stems also puberulent, the other glabrous except the stigmas.

Key to Forms of Piper guahamense

Leaves glabrous beneath ...........................................................f. glabrum

Leaves puberulent beneath ..........................................................f. guahamense

The following uses and vernacular names are listed with no attempt to segregate them to apply to particular forms.

USES.—Root used to cure gonorrhea (Rota: Yuncker, 1959:87).

VERNACULAR NAMES.—

pupúlon anite (Rota: Fosberg 25013)

pupulan-aniti (Guam: Fosberg 46252)

pupúlo aneti (Guam: Seale in 1900)

pupúlo aniti (Guam: Nelson 5)

pupúlu ito (Guam: Seale in 1900)

Piper guahamense f. glabrum (Yuncker) Fosberg

Piper guahamense f. glabrum (Yuncker) Fosberg, Phytologia, 13:236, 1966.

Piper guahamense var. glabrum Yuncker, Occ. Pap. Bishop Mus., 22:87–88, 1959 [type from Rota, Fosberg 25036].

Leaves glabrous beneath.

Found on Agrigan, Tinian, Rota, and very rarely on Guam.

Although the ranges of this and the puberulent form are partially distinct, the trivial nature of the distinction suggests that the category of forma would be more appropriate than varietas. One sheet of G.E.S. 387 in the U. S. National Herbarium is puberulent, and was determined by Yuncker as P. guahamense; another in the Bishop Museum is glabrous and was a part of the basis for Yuncker’s var. glabrum. The latter sheet is the only basis for ascribing var. glabrum to Guam. However, there seems no reason to doubt the locality of the sheet.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Yuncker, 1959:87–88 (as P. guahamense var. glabrum).

Agrigan: Hosokawa 8011 (BISH).

Tinian: Ohtani 136 (FU).

Rota: Kanehira 1753 (FU), 1788 (FU), 3619 (FU); Necker R113 (US); 50 ft [15 km], Grether 4425 (US); ¾ mile [0.6 km] east of Sabana, 2nd terrace from top, 420–450 m, Fosberg 25036 (BISH, type, US, Fo, NY, DPU, isotypes); near Sinapal, Hosokawa 7568 (A, BISH); 2nd main terrace from top, 320 m, Fosberg 31855 (US, BISH, DPU); between Rota and Tataacho Pt., Fosberg 25013 (US, DPU); Shinapatu, 150 ft [45 m], Hosaka 3045 (US, BISH, DPU); Sabana, Kondo in 1952 (BISH): 400 m, Evans 2237 (US, BISH, Fo, K); main road to Sabana from airport road, 200–400 m, Sachet 1772 (US, Fo). Guam: G.E.S. 387 part (BISH).

Piper guahamense C. de Candolle f. guahamense

Piper guahamense C. de Candolle in de Candolle, Prodr., 16(1):336, 1869.

Leaves and younger parts prominently puberulent.

Confined to Rota and Guam, so far as known, and the common form on Guam.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—s.l. Gaudichaud 163 (P, type).

Rota: Middle level plateau above Tataacho Point, 150–250 m, Evans 2058 (US, BISH, Fo, K).

Guam: s.l., LeGuillou in 1839 (P); Hombron in 1839 (P, 2 sheets); G.E.S. 387 part (US); Nelson 5 (US, BISH); Marche 101 (P); Merrill, 1914:71; Mt. Lamlam, summit, 400 m, Anderson 144 (US, BISH), 318 (US, BISH, Fo, DPU); Harmon-Barrigada road, west slope, Barrigada Hill, Stone 3782 (UG); east edge of Barrigada Point area, 400–600 ft [120–180 m], Moore 7 (US); Ritidian Point, 400 ft [120 m], Hosaka 3089 (US, BISH); Necker 159 (US); near Talisay, 150 m, Fosberg 35263 (US, BISH, Fo, NY, DPU), 150 m, Fosberg 35264 (US, BISH, Fo, DPU); Yigo, Whiting P10a (US, Fo); headwaters of Ylig River, Rodin 613 (US); Agaña, Seale in 1900 (BISH); Talofofo area, Pedrus 23 (BISH); Naval Magazine, Almagosa Springs Road, Stone 4105 (UG); Canada, Bryan 1074 (US, BISH); Maguao, 15 m, limestone, Bryan 1036 (US, BISH); Mogfog, 400 ft [120 m], Gressitt 2007 (US); southwest of Anao Point, 200 m, Fosberg 31938 (US, BISH, Fo, NY, L, DPU); one mile south of Taguac, 110 m, Fosberg 35478 (US, BISH, Fo, NY, L, DPU); Mataguac, Reid Moran 4438 (BISH, US, Fo, UC, MO); Anigua, 4 m, Fosberg 31214 (US, BISH, Fo, NY, L, DPU); near Mt. Tenjo, Swezey in 1936 (BISH); Tailalo, Reid Moran 4581 (US, BISH, Fo, UC); Andersen Air Force Base, Reid Moran 4404 (US, BISH, Fo, UC); north of northwest strip, Conover & Wagner 563 (BISH); Tarague Beach, Andersen Air Force Base, 0–25 m, Evans 731 (US, BISH, Fo, K); Lujuna, just off route 15, 180 m, Evan 1579 (US, UH, Fo, K, Mo, L); Dededo Well Field, 110 m, Evans 1692 (US, Fo, UH, K, P); Mt. Lamlam, track from Jumullong Mangio, 300–405 m, Evans 237 (US, Fo, P, L, BISH, MO); Mt. Lamlam, 370 m, Fosberg 46252 (US, Fo, UH, A, L); 46263 (US, Fo, UH, P, L, MO).

Piper hosokawae Fosberg

Piper hosokawae Fosberg, Phytologia, 13:237, 1966.

Piper decumanum var. palauense Hosokawa, Trans. Nat. Hist. Soc. Formosa, 28:153–154, 1938 [type from Peliliu, Hosokawa 9226] [non Piper palauense Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:118, 1935].—Yuncker, Occ. Pap. Bishop Mus., 22:89, 1959.

Piper majusculum sensu Kanehira, Enum. Micr. Pl., 305, 1935 [non Blume, Verh. Bot. Gen., 11:210, 1826].

Coarse glabrous climber; leaves ovate or ovate-oblong, strongly and usually obliquely cordate at base, acuminate at apex, main nerves 9–11 (12–13 according to Hosokawa), diverging from midrib basally or subbasally, petiole shorter than the longer of the basal lobes of the blade, somewhat winged to about half way up; spikes pedunculate, from nearly as long as to much longer than leaves; ovary attenuate to a neck in upper part, stigmas 3, tomentulose.

This Palau plant seems sufficiently distinct from either of the two forms, one American, the other Malaysian, that were originally included in P. decumanum L.

Endemic to Palau, so far as known, where it grows in forests throughout the group, but not common.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Palau: Kanehira, 1935:305 (as P. majusculum); Hosokawa 1938:153–154, Yuncker, 1959:89 (both as P. decumanum var. palauense); Akoru-kaigan, Hosokawa 7098 (BISH, A); Todaiyama, Kanehira & Hatusima 4646 (FU); Kanehira 2027 (FU). Babeldaob: Airai, Takamatsu 1163 (BISH); Marikyoku [Melekiok], Kanehira 392 (FU); Ogiwal, Hosokawa 9192 (cited by Hosokawa, 1938:153–154, not seen by us). Peleliu: Hosokawa 9226 (BISH, A, isotypes); Takamatsu 1784 (BISH), 1763 (BISH); Hatusima 4742 (FU). Angaur: Kanehira 568 (FU).

Piper latifolium L.f.

Piper latifolium L.f., Suppl., 91 [as P. methysticum corrected to P. latifolium in Emendanda, p. [468] at end of book] 1781 [type from Tahiti, no collection mentioned].—Fosberg, Phytologia, 13:236–237, 1966.

Piper micronesiacum Hosokawa, Trans. Nat. Hist. Soc. Formosa, 32:287, 1942 [type from Kusaie, Hosokawa 9498]; Yuncker, Occ. Pap. Bishop Mus., 22:86–87, 1959 [sub. P. guahamense].

Large shrub, to 2.5 m, leaves orbicular, to 17 × 15 cm, thin, base deeply cordate, apex acuminate, main veins 9–13 from base, slightly puberulent beneath, petiole 5–9 cm long, ⅓ to ⅔ invaginate; spikes axillary, 1-several in an axil, about 10 cm long on peduncles 1 (−4?) cm long.

In habit similar to P. methysticum but with axillary spikes; differing from P. guahamense in the usually more than one spike in an axil. Found from Kusaie south to the New Hebrides and east to Tahiti and the Marquesas.*

CAROLINE ISLANDS.—Kusaie, Mt. Keres, Mallens, Hosokawa 9498 (A, US, isotypes of P. micronesiacum).

Piper methysticum Forster f.

Piper methysticum Forster f., Pl. Esc. Ins. Oc. Austr., 76, 1786 [type from cultivation in the Society, Friendly, or Sandwich islands, collected by Forster, not seen by us].— Christian, Caroline Islands, 188–193, 329, 348, 1899.—Kanehira, Fl. Micr., 81–82, pl. 14, 1933; Enum. Micr. Pl., 305, 1935—Hoshino, Guntô no Sangyô [1934]:[1–11, 1934].— Smith, Jour. Am. Arb., 24:353, 1943.—Glassman, Res. Rev., 1950:16–18, 1950; Bishop Mus. Bull., 209:55, 1952.—Massai and Barrau, S. Pac. Comm. Techn. Pap., 94:44, 1956.— Yuncker, Occ. Pap. Bishop Mus., 22:88–89, 1959.—Fosberg, Phytologia, 13:238, 1966.

Macropiper methysticum (Forster f.) Miquel, Comment. Phytogr., 36, 1840.

Large shrub (to 50 ft [15 m] according to Glassman 1950), nodes prominent; leaves large, orbicular-cordate, slightly acuminate, nerves 11 or 13, from base except inner pair which issue from midrib 1–2 cm above base, petioles and veins beneath very shortly but densely puberulent, margin ciliolate, petiole 2–3 cm long, not wing-expanded, only the extreme base sheathing, stipules lanceolate, 4–5 cm long, caducous; flowers dioecious, in pedunculate spikes mostly less than half the length of leaves, pistillate only known from Micronesia, bracts strongly imbricate when young, their margins erose or even fimbriate; ovaries obovoid, stigmas 3, flattish; fruit not seen.

New Guinea, Melanesia, Polynesia; in Micronesia planted or persisting from planting, possibly spontaneous, on Ponape and Kusaie. We have seen no specimens from Kusaie, but do not doubt that it occurs there or at least did formerly. Guam records probably apply to the superficially similar P. guahamense. Christian (1899:348), gives the names langil and thlangil from Yap, saying they apply to “a variety of the wild kava,” but since we have no record of P. methysticum from there, we assume that these names must apply to something else. We also doubt the authenticity of the records from Palau and Truk, as we have seen no material from either group.

ETHNOBOTANY.—Christian (1899:188–193) discusses kava-drinking, says it exists only on Ponape and Kusaie in Micronesia; he regards it as having great medicinal value. A watery infusion of the pulverized root is a widely used beverage (Yuncker, 1959:89). Account of preparation and use of “sakau” (kava), made from this plant; cites beliefs in medicinal virtues, also; banned in Kusaie by missionaries in 1850 (Ponape: Glassman, 1950:18). “There are several centres of kava drinking in the area: Ponape in the Carolines …” (Ponape: Massal and Barrau, 1956:44). Roots are source of beverage kava (Ponape: Glassman 2725). Root used to make drink (Ponape: Kanehira, 1935:305). Observed in common use on Ponape in 1973 (Sachet, ined.).

VERNACULAR NAMES.—

chakau (Ponape: Christian, 1899:329)

choko (Ponape: Christian, 1899:329)

sakaw (Ponape: Gabung 1)

sakau (Ponape: Not District, Glassman 2725; Yuncker, 1959, p. 89)

sawkaw (Ponape: Fosberg 26318)

shakau (Ponape: Kanehira, 1935:305; Kiti District, Glassman, 1952:55)

seka (Kusaie: Christian, 1899:329; Kanehira, 1933: 82)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Guam: Marche 10 (P) (placed here with doubt by Yuncker but probably better referred to P. guahamense).

CAROLINE ISLANDS.—Palau: Babeldaob (Babelthuap): Melekiok (Marukyoku), Yuncker, 1959:88–89, citing Kanehira 453 which we refer to P. betle f. densum.

Truk: Christian, 1899:189, reports, as hearsay, that kava grows in “Hogolu or Ruk.”

Ponape: Christian, 1899:188–193, 329; Metaranium (Matalanim), Kanehira 835 (FU, BISH); Gabung 1 (US); Colonia, Kanehira 671 (NY); Mt. Asama-san, 500–200 m, Hosokawa 9601 (A); Mt. Seletereh, 500 ft [150 m], Glassman 2725 (US, BISH); south base of Tolenot Pk., Not District, 3–10 m, Fosberg 26318 (US); Anapeng-pa, Takamatsu 708 (BISH); Tolomail, cultivated, Takamatsu 970 (BISH); Kuporujo, cultivated, Takamatsu 676 (BISH).

Kusaie: Christian, 1899:188–189; Kanehira, 1933: 82; Glassman, 1952:55.

Piper nigrum L.

Piper nigrum L., Sp. Pl., 28, 1753.—Saflord, Plant World, 5: 196, 1902; Contr. U.S. Nat. Herb., 9:282, 355, 1905.—Okabe, Nettai Sangyô Kenkyu-sho ihô, 5:2, 1940.—Bryan, Plants of Guam [unpub., 1946?].—Yuncker, Occ. Pap. Bish. Mus., 22: 90, 1959.

Glabrous climber with ovate to broadly elliptic leaves, obtuse to rounded at base, acuminate at apex, minutely but densely black-punctate beneath, 7-nerved, the outer pair very faint, arising at base, a second pair, much stronger, just above base, the other pairs arising well above the base, well separated, the upper strongly alternate, petiole 1–2 cm long, expanded only at extreme base; spikes pedunculate, less than 10 cm long, not equaling leaves, pistillate flowers rather loosely arranged on rachis, ovary broadly ovoid, stigmas linear, 3, spreading, tending to be raised on an extremely short style; fruit globose, free, neither connate nor sunken in rachis (description mostly from Philippine specimens).

Widely cultivated in southeast Asia and Malesia, known to be or have been cultivated in Micronesia, on several of the high islands.

ETHNOBOTANY.—This is the plant that yields the black pepper and the white pepper of commerce. It has been planted on an experimental scale in several places in Micronesia, e.g., Palau (fide Okabe, 1940) and Guam (Safford, 1902); some of excellent quality is produced for sale in Ponape, called there, “Sele Soal” fide Salomon and George.

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

MARIANAS ISLANDS.—Yuncker, 1959:90 (“No specimens seen in Micronesian collections examined”).

Guam: Mangilao, Dept. Agriculture Nursery, Falanruw & Barcinas 998 (UG, US).

CAROLINE ISLANDS.—Palau: Koror, cultivated, Cheatham 168 (US); Blackburn 161 (US).

Ponape: Ponsakir, Salomon & George 9 (US).

Piper ponapense C. de Candolle

Piper ponapense C. de Candolle, Bot. Jahrb., 56:502, 1921 [type from Ponape, Ledermann 18429a, not seen by us].— Kanehira, Enum. Micr. Pl., S05, 1985.—Hosokawa, Jour. Jap. Bot., 13:199, 1937a; Bull. Biogeogr. Soc. Jap., 7:188, 1937b; Kudoa, 5:95, 1937c; Trans. Nat. Hist. Soc. Formosa, 33:120, 1943.—Riesenberg, S. W. Jour. Anthrop., 4:428, 1948.—Glassman, Bish. Mus. Bull. 209:54–55, 1952; Pac. Sci. 7:303, 1953.—Yuncker, Occ. Pap. Bish. Mus., 22:93–95, 1959.—Stone, Pac. Sci., 13:100, 1959.

Piper kusaiense Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:118, 1935 [type from Kusaie, Hosokawa 6324].—Kanehira, Enum. Micr. Pl., 305, 1935.

Piper palauense Hosokawa, Trans. Nat. Hist. Soc. Formosa, 25:118, 1935 [type from Palau, Hosokawa 7361]—Okabe, Jour. Anthr. Soc. Nip., 56:426, 1941; Nankyô, 2:21, 1943 [as P. palawense].

Glabrous or subglabrous aromatic climber, rooting at nodes; leaves almost orbicular to broadly lanceolate, strongly acuminate, base rounded to cordate, petioles commonly about 1 cm long on mature plants, juvenile plants with generally broader, more cordate leaves and longer petioles, these and basal parts of nerves often slightly puberulent, nerves usually 7, diverging at base or almost so, outer pair somewhat weaker, rarely an extra very weak outer pair; flowers dioecious, in linear elongate pedicillate spikes, usually equaling or exceeding leaves; bracts of staminate spikes imbricate; stamens 2, anthers elliptic or oblong, bilocular; stigmas 3 or 4, spreading, pubescent, easily rubbed off in age; ovaries obconical, subtruncate at apex; fruit obovoid, red, fleshy, not greatly enlarging.

Extremely variable, but P. kusaiense and P. palauense do not appear to differ in any essential respect. Broad leafed or juvenile forms, when sterile, distinguished with difficulty from P. fragile. A slight trace of puberulence on the petiole and under sides of the veins at the base of the leaf in juvenile material seems confined to P. ponapense. The petioles are also generally shorter in P. ponapense, but there is some overlapping in young specimens.

Found throughout the Caroline Islands (Palau, Truk, many of the atolls, Ponape, Kusaie) from lowlands to middle altitudes, in forests, thickets, and openings, creeping on ground and climbing on rocks and tree trunks.

USES.—Plant parts are used to alleviate stomach aches (Ponape, Truk, Ant: Glassman, 1952:55).

VERNACULAR NAMES.—

kasibibui (Palau: Fosberg 47448)

ksebebui (Palau: Fosberg 25801)

kesibibui (Palau: Okabe, 1941:425)

keshbibbui (Palau: Okabe, 1943:21)

tolboy (Lamotrek: Fosberg & Evans 46786)

elek (Satawal: Fosberg 46897)

enes (Truk: Wong 152)

ennes (Truk: Pelzer 47)

eras (Truk: Fosberg 24554)

eresi (Truk: Hosokawa, 1937b: 188)

anek (Namoluk: Marshall 89)

anük (Etal: Anderson 2062)

anük (Lukunor: Anderson 2102)

anük (Satawan: Anderson 1107)

ahnuck (Ant: Glassman, 1953:303)

kon’ok (Ponape: Riesenberg 40)

ko’nok (Ponape: Yuncker, 1959:95)

konok (Ponape: Glassman, 1952:54, 55)

kai-fuhl (Kusaie: Glassman 2718)

GEOGRAPHIC RECORDS AND SPECIMENS EXAMINED

CAROLINE ISLANDS.—Palau: Babeldaob: Aimiriik, Kanehira & Okamoto 18 (FU), Kanehira 285 (FU, BISH); Hosokawa 7207 (A); in a thicket along a valley near Mt. Unkeshiu, Hosokawa 7138 (cited by Hosokawa, 1935:118, not seen by us); Arumonogui, near Arumatan, Hosokawa 6748 (BISH, A); Malukyoku, Galdok (Ngardok) Kaisyalu I., Hosokawa 7361 (A, isotype of P. palauense); Garamiscan Colony, Fosberg 25763 (US, BISH, Fo, NY, DPU, L); Kamsetsu, Takamatsu 1120 (BISH); Kasioru, Takamatsu 1507 (BISH); Kaiguru, Takamatsu 1590 (BISH). Koror: Kanehira 196 (FU); in a forest on coral at Almizu, Hosokawa 7407 (cited by Hosokawa, 1935:118, not seen by us). Ngarakabesang (Arakabesan): Takamatsu 1246 (BISH); south side of west peninsula of island, 20–30 m, Fosberg 32132 (US, BISH, Fo, DPU). Aulupse’el: Ngar Malk, 1–10 m, Fosberg 32616 (US, BISH, Fo, NY, DPU), Fosberg 32617 (US, BISH, DPU); Du’ebachel beach, east end of island, 2 m, Fosberg 47448 (US, UH, Fo, MO, K, NY); Ngeremengii, Trail from bauxite mining area to school, Falanruw, Fosberg, et al. 1058 (US). Ngeanges (Gaiangas): Island west of south point of Urukthapel, Fosberg 25801 (US, BISH, Fo, DPU). Peliliu: Takamatsu 1788 (BISH); Kanehira & Hatusima 4779 (FU). Angaur: East side, Fosberg 31989 (US).

Lamotrek Atoll, Lamotrek Islet, interior of island, 1–3 m, Fosberg & Evans 46786 (US).

Satawal: interior of island, 2 m, Fosberg 46897 (US, BISH, Fo).

Puluwat Atoll: 1–3 m, Niering 768 (US).

Namonuito: Ulul I., Stone 2078 (BISH).

Murilo: Murilo I., 3–5 m, Evans 1241 (US), 1240 (US); Ruo I., interior of island, 3–5 m, Evans 1193 (juvenile) (US), 7202 (US); main taro pit, 1176 (US, UH, Fo, K, MO); observed, not collected (Stone, 1959:100).

Nomwin: Nomwin I., observed, not collected (Stone, 1959:100).

Truk: Wong 152 (US); Tadiu, Hosokawa 8401 (BISH, A, US); Akisima, Kanehira 3872 (FU), 3877 (FU); Uoala, H. F. Moore 114 (US). Moen: Kanehira 1716 (FU); Pelzer 47 (US); 48 (US); near Baker Dock, 3 m, Fosberg 24704 (US, BISH, Fo, DPU), Fosberg 24697 (US, BISH, Fo, DPU); south slope of Mt. Tonaachau, Fosberg 26053 (US, BISH, Fo, DPU, NY, L); Daniells 100 (US); track leading to highest point on island, 100–400 m, Evans 1265 (US, BISH, Fo); 1284 (US, BISH). Dublon (Trowasi, Natusima): Hosokawa 8362 (BISH); Mt. Tolomen (Tolowan), 200–360 m, Fosberg 24554 (US, BISH, Fo, NY, DPU); Mt. Tumuital (Uiniboet), 200–460 m, Fosberg 24444 (US, BISH, Fo, NY, DPU). Tol: Hosokawa 8327 (A, BISH); 300 m, Kanehira 1288 (FU); Uriribot, Hosokawa 8269 (BISH, A), 8292 (BISH, A).

Losap: Seen but not collected by Anderson in 1949; Losap I., interior of island, 3–5 m, Evans 1342 (US, UH).

Namoluk: Seen but not collected by Anderson in 1949; Namoluk Islet, 2–3 m, Marshall 89 (US).

Etal: Etal I., Anderson 2062 (US, BISH, Fo, DPU).

Lukunor: Lukunor I., Anderson 2167 (US, BISH, Fo, NY, DPU); Oneop I., Anderson 2102 (US, BISH, Fo, NY, L).

Satawan: Moch I., Anderson 1107 (US, BISH, Fo, NY, L, DPU).

Ant: observed as an epiphyte by Glassman (1953: 303).

Ngatik (Natik): Kanehira 4120 (FU).

Ponape: Ledermann 13266 (BISH); Kanehira 672 (BISH); Hosokawa 5472, 5538, 5585, 5598, 5721, 5858, 5919, 6032, 6148 (all cited by Hosokawa, 1943: 120, not seen by us); Lederman 13429a, type (cited by de Candolle, 1921:502, not seen by us); Riesenberg 40 (BISH); H. F. Moore 108 (US), 107 (US); Colonia, Kanehira 672 (US, NY, BM, FU); Palikir, Kanehira 721 (NY); Kiti, Kanehira 1539 (US, NY, FU); outlet of Tawensokola River, Glassman 2444 (US, BISH); Parkier, Kanehira 706 (FU); Mt. Kankauzan, Hosokawa 5505 (A); Nanipiru, 200 ft [60 m], Hosaka 3539 (US, BISH, Fo, DPU); Mt. Tolotom, Takamatsu 1059 (BISH); Param, Takamatsu 633 (BISH); Net Village, along river bank, Salomon, S. & T. George 57 (US).

Kusaie: Kanehira 1350 (FU, P); Matante, Kanehira 4199 (FU); Mt. Matante (Mt. Buache), Takamatsu 496 (BISH); lowest slope and foot of south side, 1–50 m, Fosberg 26566 (US, BISH, Fo, NY, DPU); Mt. Buache, Hosokawa 6261 (A), Taonsakku, Hosokawa 6324 (BISH, A, isotypes of P. kusaiense); Yasibayasi-Taontakku, Hosokawa 6215 (A); Lele I.: Fosberg 26543 (US, BISH, Fo, NY, L, DPU), Glassman 2718 (US, BISH); south slopes and ridges of Mt. Tafeyet, south of the Lele Harbor, Fosberg 26668 (US); Hill 541, 500 ft [150 m], J. F. G. Clarke s.n. (US); Mt. Fenkol (Mt. Crozer, Mt. Hinkorn, Hinkoln), 100 m, Takamatsu 385 (BISH); Kanehira 1388 (FU), 4193 (FU).

Synonyms and Excluded or Misapplied Names

Chavica Miquel. See Piper L.

Chavica betle (L.) Miquel. See Piper betle L.

Macropiper Miquel. See Piper L.

Macropiper potamogetonifolium (Opiz) Miquel. See Piper betle f. marianum (Opiz) Fosberg

Macropiper latifolium (Forster f.) Miquel. See Piper guahamense C. de Candolle for Marianas records

Macropiper methysticum (Forster f.) Miquel. See Piper methysticum Forster f., but see Piper guahamense C. de Candolle for Marianas records

Micropiper Miquel. See Peperomia Ruiz & Pavón

Peperomia bilineata (Blume) Miquel. See Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth, at least for Yap record

Peperomia gibbonsii C. de Candolle. See Peperomia ponapensis C. de Candolle var. ponapensis

Peperomia guamana C. de Candolle. See Peperomia mariannensis C. de Candolle f. mariannensis

Peperomia guamana var. saipana (C. de Candolle) Yuncker. See P. mariannensis f. saipana (C. de Candolle) Fosberg

Peperomia hoeferi C. de Candolle. See P. mariannensis C. de Candolle f. mariannensis

Peperomia ladronica Hosokawa. See P. mariannensis C. de Candolle f. mariannensis

Peperomia lineata Miquel ex Yuncker. See Peperomia pellucida (L.) Humboldt, Bonpland & Kunth

Peperomia membranacea Hooker & Arnott. See Peperomia mariannensis C. de Candolle for Micronesian record

Peperomia pacifica Hosakawa. See Peperomia mariannensis f. saipana (C. de Candolle) Fosberg

Peperomia pacificicola Hosokawa. See Peperomia mariannensis C. de Candolle f. saipana (C. de Candolle) Fosberg

Peperomia pallida (Forster f.) A. Dietrich. See Peperomia ponapensis C. de Candolle f. ponapensis for Marshall Islands record

Peperomia pellucida var. obtusifolia Koidzumi. See Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth

Peperomia pellucidopunctulata C. de Candolle. See Peperomia mariannensis C. de Candolle for Marianas record

Peperomia saipana C. de Candolle. See Peperomia mariannensis f. saipana (C. de Candolle) Fosberg

Peperomia tinianensis Hosokawa. See Peperomia mariannensis C. de Candolle f. mariannensis

Peperomia trukensis Yuncker. See Peperomia ponapensis var. trukensis (Yuncker) Fosberg

Peperomia volkensii C. de Candolle. See Peperomia ponapensis C. de Candolle var. ponapensis

Peperomia yapensis C. de Candolle. See Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth

Piper betel L. ex Endlicher. See Piper betle L.

Piper betle var. densum (Blume) C. de Candolle. See Piper betle f. densum (Blume) Fosberg

Piper betle var. mariannum C. de Candolle. See Piper betle L. f. marianum (Opiz) Fosberg

Piper bilineata Miquel ex Kanehira. See Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth

Piper carolinense Hosokawa. See Piper fragile Bentham

Piper decumanum var. palauense Hosokawa. See Piper hosokawae Fosberg

Piper densum Blume. See Piper betle f. densum (Blume) Fosberg

Piper guahamense var. glabrum Yuncker. See Piper guahamense f. glabrum (Yuncker) Fosberg

Piper kusaiense Hosokawa. See Piper ponapense C. de Candolle

Piper latifolium Gaudichaud ex Kunth. See Piper guahamense C. de Candolle

Piper macgillivrayi C. de Candolle ex Seemann. See Piper guahamense for Marianas records

Piper majusculum Blume. See Piper hosokawae Fosberg for Micronesian records

Piper mariannum Opiz. See Piper betle f. marianum (Opiz) Fosberg

Piper marianum Opiz. See Piper betle f. marianum (Opiz) Fosberg

Piper micronesiacum Hosokawa. See Piper latifolium L. f.

Piper obtusifolium. L. See Peperomia obtusifolia (L.) A. Dietrich

Piper palauense Hosokawa. See Piper ponapense C. de Candolle

Piper palawense Okabe. See Piper ponapense C. de Candolle

Piper pellucidum L. See Peperomia pellucida (L.) Humboldt, Bonpland, & Kunth

Piper potamogetonifolium Opiz. See Piper betle f. marianum (Opiz) Fosberg

Piper trukense Hosokawa. See Piper fragile Bentham.

MYRICACEAE

Shrubs and trees, usually aromatic; leaves simple, alternate or rarely opposite, exstipulate or with very early caducous stipules; flowers dioecious or monoecious, in catkins, subtended by bracts, reduced to small clusters of stamens or to a solitary naked pistil, ovary 1-celled, with one ovule, style very short, stigma cup-shaped, produced into two spreading flat lobes, usually regarded as two stigmas; fruit a drupe covered by waxy or fleshy peltate processes or warty, containing a single seed.

Myrica L.

Myrica L., Gen. Pl., ed. 5, 449, 1754 [=1753]; Sp. Pl., 1024, 1753.

Characters of the family.

An almost cosmopolitan genus. One species introduced into Guam from East Asia, not collected recently.

Myrica rubra Siebold & Zuccarini

Myrica rubra Siebold & Zuccarini, Abh. Akad. Muenchen IV, 3:230, 1846.—Bryan, Plants of Guam [unpub., 1946?].— Stone, Micronesica 6:245, 1971.

Tree to 15 m tall, essentially glabrous, leaves narrowly obovate to oblanceolate or somewhat spatulate, 4–12 cm long, 1–3 cm wide, apex obtuse to rounded, base cuneate-contracted, margins entire to obscurely crenate-serrate toward apex, coriaceous, under surface punctate with sessile yellow resinous glands, venation not prominent, 6–8 main veins on a side, network closely reticulate, petiole 1 cm or less long; staminate flowers in simple or branched catkins, 1-several in a leaf axil, 1–3 cm long, anthers very plump, about 1.5 mm long and wide, sessile, several together subtended by concave glandular-ciliolate scales, pistillate catkins shorter, to 1.5 cm, solitary, minutely hirsute, flowers reduced to an ovary surmounted by an expanded cup-shaped, greatly bilobed stigma, subtended by a glandular obtuse concave bract, fruit a subglobose drupe about 10 × 7–8 mm, hirsute, closely beset with erect peltate processes that become fleshy and red when mature, endocarp hard.

MARIANAS ISLANDS.—Guam: Bryan [1946?].—Stone, 1971:245.

Said to have been introduced in 1911 (fide Stone), no Micronesian collections available, possibly no longer extant there. Description from Japanese and Ryukyu specimens.

Literature Cited

Fosberg, F. R., and M.-H. Sachet

1975.    Flora of Micronesia, 1.: Gymnospermae. Smithsonian Contributions to Botany 20:1–15.

Lanjouw, J., and F. A. Stafleu

1964.    Index Herbariorum, Part I: The Herbaria of the World. 5th edition. Regnum vegetabile, 31:1–251.

Otobed, D. O.

1967.    Partial List of Some Plants and Trees of the Palau Islands. Manuscript, files of F. R. Fosberg, Department of Botany, National Museum of Natural History, Smithsonian Institution. [Duplicated and circulated as Guide List of Plants of the Palau Islands. 25 pages. Koror, Palau, 1971].

Rumphius, G. E.

1743.    Herbarium Amboinense. Volume 3, 218 pages, 141 plates. Amsterdam.

Sachet, M.-H., and F. R. Fosberg

1955.    Island Bibliographies, v + 577 pages. Washington, D.C.

1971    Island Bibliographies Supplement. ix + 427 pages. Washington, D.C.

Thomas, P.

1745.    A … Journal of a Voyage to the South Seas … 347 + 39 pages. London.

Index

Accepted names are in roman type. Synonyms, missapplications, and page numbers of principal treatments are in italics.

Areca catechu, 17

Casaarina, 1

litorea, 4

Casuarina, 1

equisetifolia, 4

equisetifolia var. souderi, 6

litorea, 4

litorea var. souderi, 6

mertensiana, 4

Casuarinaceae, 1

Chavica, 16

betle, 17

Macropiper, 16

latifolium, 20

methysticum, 20, 23

potamogetonifolium, 18

Micropiper, 6

Myrica, 27

rubra, 27

Myricaceae, 27

Peperomia, 6

argyreia, 6

bilineata, 13, 14

breviramula, 7, 8, 9

caperata, 6, 7, 8

gibbonsii, 14, 15

glassmanii, 7, 8, 9, 12

guamana, 9, 10, 11

guamana var. saipana, 10, 11, 12

hoeferi, 10

kraemeri, 7, 8, 9, 10

kusaiensis 7, 9

ladronica, 10

leptostachya, 7, 9, 12

lineata, 13

magnoliaefolia, 12

mariannensis, 7, 8, 9, 10, 12, 15

mariannensis f. mariannensis, 10, 12

mariannensis f. saipana, 7, 10, 11

membranacea, 9, 10

obtusifolia, 6, 7, 12

pacifica, 11, 12

pacificicola, 11

palauensis, 7, 8, 10, 12, 16

palauensis var. occidentalis, 7, 13

palauensis var. palauensis, 12

pallida, 15

pellucida, 7, 13

pellucida var. obtusifolia, 13, 14

pellucidopunctulata, 9, 10, 11

ponapensis, 7, 10, 14, 15

ponapensis var. ponapensis, 15

ponapensis var. trukensis, 7, 15, 16

saipana, 11, 12

sandersii, 6

tinianensis, 10

trukensis, 14, 16

volkensii, 14, 15, 16

yapensis, 13, 14

Piper, 6, 16

betel, 17

betle, 16, 17, 18

betle f. betle, 17, 18

betle f. densum, 17, 18, 23

betle var. densum, 18

betle δ mariannum, 18

betle f. marianum, 17, 18

bilineata, 13

carolinense, 19

decumanum, 22

decumanum var. palauense, 22

densum, 18

fragile, 16, 19, 24

guahamense, 16, 20, 21, 22, 23

guahamense f. glabrum, 21

guahamense var. glabrum, 21

guahamense f. guahamense, 21

hosokawae, 16, 22

kusaiense, 24, 25

latifolium, 16, 20, 22

macgillivrayi, 20

majusculum, 22

mariannum, 18

marianum, 18, 19

methysticum, 16, 17, 22, 23

micronesiacum, 22

nigrum, 16, 23

obtusifolium, 12

palauense, 22, 24, 25

palawense, 24

pellucidum, 13

ponapense, 16, 24

potamogetonifolium, 18, 19

trukense, 19, 20

Piperaceae, 6

* Since the above was written, Professor A. C. Smith has shown us photographs indicating that the hairs along the nerves on the underside of the leaf blade of P. micronesiacum are similar to those of P. guahamense rather than to those of P. latifolium. He is inclined to associate P. micronesiacum with P. guahamense. Our inclination would be, on the basis of this observation, to regard both P. micronesiacum and P. guahamense as varieties of P. latifolium, but we prefer to give the matter further study.