Palau’s Forest Resources, 2003 Joseph A. Donnegan, Sarah L. Butler, Olaf Kuegler, Brent J. Stroud, Bruce A. Hiserote, and Kashgar Rengulbai Resource Bulletin PNW-RB-252 January 2007 United States Department of Agriculture Forest Service Pacific Northwest Research Station The Forest Service of the U.S. Department of Agriculture is dedicated to the principle of multiple use management of the Nation’s forest resources for sustained yields of wood, water, forage, wildlife, and recreation. Through forestry research, cooperation with the States and private forest owners, and management of the National Forests and National Grasslands, it strives—as directed by Congress—to provide increasingly greater service to a growing Nation. The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of race, color, national origin, age, disability, and where applicable, sex, marital status, familial status, parental status, religion, sexual orientation, genetic information, political beliefs, reprisal, or because all or part of an individual’s income is derived from any public assistance program. (Not all prohibited bases apply to all programs.) Persons with disabilities who require alternative means for communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, 1400 Independence Avenue, SW, Washington, DC 20250-9410 or call (800) 795-3272 (voice) or (202) 720-6382 (TDD). USDA is an equal opportunity provider and employer. Authors Joseph A. Donnegan is an ecologist, Sarah L. Butler is an ecologist, Olaf Kuegler is a statistician, and Bruce A. Hiserote is an information management specialist, Forestry Sciences Laboratory, 620 SW Main Street, Suite 400, Portland, OR 97205-3028. Brent J. Stroud was the quality assurance technician for the Portland Forestry Sciences Laboratory, but now works for Interior West Forest Inventory and Analysis. Kashgar Rengulbai was a forester on Palau. Abstract Donnegan, Joseph A.; Butler, Sarah L.; Kuegler, Olaf; Stroud, Brent J.; Hiserote, Bruce A.; Rengulbai, Kashgar. 2007. Palau’s forest resources, 2003. Resour. Bull. PNW-RB-252. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 52 p. The Forest Inventory and Analysis Program collected, analyzed, and summarized field data on 54 forested plots on the islands in the Republic of Palau. Estimates of forest area, tree stem volume and biomass, the numbers of trees, tree damages, and the distribution of tree sizes were summarized for this statistical sample. Detailed tables and graphical highlights provide a summary of Palau’s forest resources and a comparison to 1987 data. Keywords: Palau, biomass, damage, FIA, forest inventory, volume, land cover. Summary The Forest Inventory and Analysis (FIA) Program conducted a systematic inventory of the forests of Palau in 2003 to estimate forest area, tree stem volume, biomass, carbon storage, tree damages, and the percentage cover of understory vegetation species. Fifty-four permanent field plots were installed, 11 in limestone forest and 43 in volcanic forest. Land cover was mapped into five broad classes, forest, nonforest vegetation, urban, barren, and inland water, by using recently acquired high-resolution satellite imagery. Soil survey information was used to classify forest into limestone and volcanic types. Our estimates for this inventory are derived from a sample based on 110,028 acres in Palau, with about 16 percent of the land classified as limestone forest and 67 percent classified as volcanic forest. About 2 percent of the landscape was classified as urban land, the majority on Koror and Babeldaob. We estimated gross tree stem volume to be about 302 million cubic feet for all size classes including seedlings and saplings. Aboveground dry biomass for tree stems 5 inches and greater was estimated to be 4.7 million tons. Approximately 13 percent of the trees sampled in the inventory had some form of damage. The growth of damaging vines in the crowns of trees was the most prevalent damage type, with conks and lost apical dominance following in prevalence. The most frequently identified damaging agents were other vegetation, disease, and weather. Palau’s forests are relatively dense and have high species diversity. A total of 128 tree species and 132 understory species were measured on FIA plots. The average number of tree species per sixth-acre plot was 12 in volcanic forest and 10 in limestone forest. Palauan foresters and ecologists were especially helpful with species identification in the field. Palau’s Forest Resources, 2003 Introduction This summary of forest resources on the islands of Palau (fig. 1) was based on a forest inventory conducted in 2003 by the USDA Forest Service, Pacific Northwest Forest Inventory and Analysis (FIA) Program in cooperation with Pacific Island foresters. The inventory is a subset of the national FIA effort and was designed to help answer local and national questions about the status and trends in tropical forested ecosystems and to share forestry skills among cultures and agencies. The fieldwork for this inventory was conducted by a multinational crew including foresters from Palau, American Samoa, and mainland U.S. foresters and ecologists. This systematic, sample-based field inventory on Palau is an update to prior inventory work (Cole et al. 1987, MacLean et al. 1988) estimating forest type area, tree size distribution, and stem volume. New information is included for all forests on biomass, and damages for living and dead trees. The inventory was designed to provide resource managers with information about the current situation so they can better manage their forested and nonforested lands, and manage or mitigate any changes in the resource. Empirically based knowledge of the status and trends in forest vegetation can help managers plan sustainable land use practices, plan sustainable supplies of wood, control invasive species, control erosion, and manage disturbances such as fire and animal damage. Objectives • • • • Estimate the current area of forest land by forest type group and stand size class. Estimate the volume, biomass, and carbon storage for tree species by diameter class. Estimate the numbers of trees affected by damaging agents, such as insects and diseases, and estimate the numbers of dead trees. Share measurement and analysis techniques among groups involved in the inventory. Methods Site Description Palau is well known for its tremendous diversity and concentration of both terrestrial and marine life along the network of coral reefs and the chain of more than 300 raised limestone and volcanic islands. The chain of islands runs from the northeast to southwest for a distance of about 125 miles, approximately 470 miles east of the Philippines. The islands range in size from the relatively large, volcanic island RESOURCE BULLETIN PNW-RB-252 Figure 1—The Republic of Palau is in the North Pacific Ocean approximately 470 miles east of the Phillipines. Land cover was mapped from 2005 QuickBird satellite data. Palau’s Forest Resources, 2003 of Babeldaob, at about 92,000 acres, to coral rocks topped with trees, only a few yards across. The tremendous marine diversity is supported by the convergence of oceanic currents and the filtering and nutrient effects of the dense terrestrial forests. The topography of the four volcanic islands is marked by steep slopes and rolling hills, with mangroves often found on coastal flats. The limestone coral islands or “Rock Islands” support a diversity of tree species on often surprisingly shallow soils deposited in the recesses of the rugged karstic topography. The highest point in Palau is the approximately 700-foot-tall Mount Ngerchelchuus on Babeldaob (Palau Conservation Society). The climate of Palau is hot and humid with little annual variation in the mean maximum temperature of 87.9 °F and the mean minimum of 75.6 °F (fig. 2). Total annual precipitation averages 147.3 inches. In the wettest months, June and July, monthly precipitation averages 17.04 inches, whereas during the driest months, February, March, and April, monthly precipitation averages 8.84 inches (data from Western Regional Climate Center 2004). During an El Niño year (warm phase of the Southern Oscillation), rainfall tends to be above average for that summer and below average in the following autumn and winter months (Climate Prediction Center 2005). Palau borders the typhoon belt in the western tropical Pacific and is subject to occasional typhoon damage. Prior to European contact, extensive terracing was practiced on Babeldaob in upland areas. Paleoenvironmental evidence suggests terracing may have contributed to landcover change from forest to savanna and fernland, and accelerated erosion as much as 2,500 years before present (Athens and Ward 2002). Additional recent disturbances have included mining of bauxite, military action during World War II (which denuded much of Peleliu and Angaur), and conversion of forest for agriculture. Forest was cleared during the recent construction of a new capitol on Babeldaob and a road to encircle the island. Barren volcanic soils pose high risk of erosion with frequent, intense rainfall and are often a corridor for colonization by invasive species. Vegetation Types The flora of Palau shares many common elements with that of east-Asian and Indo-Malesian floras (Mueller-Dombois and Fosberg 1998). Species composition in Palau is distinctive depending on the parent material the soil is derived from, with volcanic soils found in the north (Babeldaob, Arakabesang, Malakal, and Oreor), and limestone soils on Peleliu, Angaur, and the Rock Islands to the south. Where volcanic soils are deposited along coastal flats, mangrove and freshwater swamps are common. The volcanic islands also support a shrubby, strand vegetation, a RESOURCE BULLETIN PNW-RB-252 Figure 2—(A) The annual maximum and minimum temperatures in Palau do not fluctuate markedly. However, (B) precipitation is highly variable throughout the year with a distinct summer wet season and a dry early spring (Western Regional Climate Center 2004). Palau’s Forest Resources, 2003 closed-canopy interior volcanic forest, ravine and riparian forest, and savanna and fernland on degraded soils (Mueller-Dombois and Fosberg 1998). The southern, raised limestone islands support evergreen, closed-canopy forest that is generally quite species rich and variable from island to island (Cole et al. 1987). Note that the current report does not describe the vegetation of the Southwest Islands of Palau, approximately 400 miles southwest of Babeldaob. Additionally, the FIA inventory does not provide sufficient plot density to reliably classify vegetation types in these highly diverse tropical forests, but provides plot-level data to help refine remotely sensed estimates of forest type acreage and developmental stage. Some of the species named in the following vegetation descriptions (from Cole et al. 1987, and Mueller-Dombois and Fosberg 1998) were not identified on the 2003 sample of FIA field plots, either owing to their absence, their understory cover not meeting the 3 percent threshold, or possible misidentification. The FIA and local island staff were trained by regional botanical experts, and when species could not be identified in the field, they were sent for expert identification. However, misidentification in the field remains a possibility. Volcanic Forest Types Mangrove— Mangroves play a vital role in buffering the effects of storms and waves along coastal areas. They also provide habitat for marine life and filter runoff exiting terrestrial ecosystems. The filtration that mangroves provide helps to sustain coral reef and fish habitat by trapping sediment. The mangroves of Palau are a particularly species-rich mangrove type that extends along a narrow strip of the tidal zone. The species found here include Rhizophora mucronata, Rhizophora apiculata, Bruguiera gymnorrhiza, Ceriops tagal, Sonneratia alba, Avicennia marina, Lumnitzera littorea, Xylocarpus granatum, Scyphiphora hydrophyllacea, Dalbergia candenatensis, Derris trifoliata, Nypa fruticans, Acrostichum aureum, and Nephrolepis acutifolia (see table 1 for common and scientific names with authorities). Mangroves are very dense forest types where left undisturbed. Swamp forest— This forest type tends to occur slightly inland of mangrove in areas of fresh or slightly brackish water. Horsfieldia amklaal, Cynometra ramiflora, Barringtonia racemosa, Campnosperma brevipetiolata, and Heritiera littoralis are often found in the intact freshwater swamps, whereas Hibiscus tiliaceus is found in disturbed areas. Scientific name Common name Aglaia palauensis Kaneh. Allophylus ternatus (Forst.) Radlk. Allophylus timorensis (DC.) Bl. Alphitonia carolinensis Hosok. Annona reticulata L. Areca catechu L. Artocarpus mariannensis Trec. Astronidium palauense (Kaneh.) Mgf. Avicennia marina (Bl.) Bakh. Badusa palauensis Valeton Barringtonia racemosa (L.) Spreng. Bruguiera gymnorrhiza (L.) Lam. Buchanania palawensis Lauterb. Calophyllum inophyllum L. Calophyllum pelewense P. F. Stevens Calophyllum soulattri Burm. f. Calophyllum sp. L. Campnosperma brevipetiolata Volk. Canarium hirsutum Willd. Casearia cauliflora Volk. Casearia sp. Jacq. Casuarina equisetifolia L. Cerbera floribunda K. Schum. Cerbera manghas L. Cocos nucifera L. Commersonia bartramia (L.) Merr. Cordia micronesica Kaneh. & Hatus. Cyathea lunulata (Forst. f.) Copel. Diospyros ferrea (Willd.) Bakh. Diospyros sp. L. Dodonaea viscosa (L.) Jacq. Dracaena multiflora L. Drypetes nitida Kaneh. Dysoxylum sp. Blume meseoes chebeludes ebeludes, chebeludes elebiob, elebiong ngel ra ngebard buuch, betel nut ebiei, wild breadfuit meskui dadaiit ralm koranges kadeges, denges, adeges geloagl laurel chesemoroch olebtaches, chesemoroch kelela charm, kiu mesechaues kevert ngas, ironwood emeridech emeridech lius, coconut aspitan, eremallueang eluu, tree fern keuelt Number measured 23 3 1 50 1 6 4 32 2 29 51 43 10 34 20 6 6 72 11 1 7 6 2 19 26 7 1 46 10 10 6 14 4 13 Estimated number (≥1 inch diameter) 847,803 36,989 12,330 1,531,290 12,330 73,977 48,518 1,641,189 24,659 897,973 3,407,461 1,095,193 405,607 1,184,578 608,511 356,489 61,247 1,133,048 553,209 12,330 84,906 24,259 24,659 362,987 318,967 165,915 12,330 634,436 393,478 544,171 73,177 878,093 187,882 1,572,844 Standard error for estimated number (±) 431,384 37,027 12,342 803,629 12,342 74,055 35,710 590,773 24,685 582,173 2,020,608 671,454 253,723 579,030 363,048 332,521 45,601 372,006 439,687 12,342 93,158 17,855 17,251 187,580 175,394 153,955 12,342 212,213 357,921 302,309 34,397 397,272 180,779 901,519 Gross volume (≥5 inches diameter) 1,783,505 206,199 29,280 4,434,129 62,771 292,165 5,140,163 1,169,439 1,422,693 3,048,028 2,986,606 5,542,897 1,308,999 5,436,901 1,040,719 480,790 1,910,972 22,836,047 1,072,970 157,802 626,648 1,436,905 281,010 2,378,467 8,416,588 29,882 312,315 1,985,641 384,300 717,026 1,642,371 459,125 373,021 64,752 Standard error for estimated volume (±) 1,359,572 206,415 29,311 1,662,961 62,836 292,472 4,028,666 460,600 1,424,186 2,276,642 1,838,290 2,800,425 1,155,922 3,643,184 654,142 342,400 1,635,495 6,496,431 1,081,278 157,967 687,549 1,536,233 234,815 906,798 4,303,993 29,913 312,643 633,206 270,515 458,131 831,416 279,546 309,627 64,820 RESOURCE BULLETIN PNW-RB-252 Table 1—Scientific names, common names, estimated number and gross volume, and standard errors for estimated totals of species measured as trees on Palau Table 1—Scientific names, common names, estimated number and gross volume, and standard errors for estimated totals of species measured as trees on Palau (continued) Scientific name dekemerír roro mesekerrak, java plum rebotel kesiil ksid oseked tilol tilol kalngebard ra belau blancheos belau chebouch, akaboek dokou demailei, ebouch cheramal emeklachel, eumail ersachel ersachel keam, boui dort ngemoel, mekekad Number measured 1 49 1 23 20 65 7 1 19 2 9 21 12 40 30 3 26 3 22 49 6 3 9 3 5 11 25 25 24 101 5 1 20 1 12,330 1,419,038 12,129 974,799 381,553 3,017,755 648,639 153,586 172,613 165,915 1,096,866 1,388,568 995,490 2,457,435 650,797 36,989 1,565,282 178,245 825,997 1,027,314 73,977 460,757 97,036 36,388 60,647 700,649 2,709,590 1,003,150 1,002,188 3,852,198 626,672 12,330 381,553 12,330 Standard error for estimated number (±) 12,342 530,051 13,308 753,514 251,122 1,068,888 313,824 153,747 76,502 153,955 936,957 780,722 741,721 969,319 298,347 37,027 776,102 166,261 495,311 537,789 51,752 461,240 106,466 39,925 66,541 641,212 2,248,657 563,395 567,747 1,132,578 614,824 12,342 267,583 12,342 Gross volume (≥5 inches diameter) 64,508 6,648,512 600,183 1,960,664 4,324,582 6,011,868 130,011 — 2,248,390 409,933 3,187,214 751,255 725,013 2,520,347 2,603,796 261,379 1,201,069 93,117 4,368,123 4,916,499 1,315,262 — 518,759 360,357 1,714,517 404,770 375,843 5,112,032 10,735,352 13,722,763 42,993 29,390 8,903,896 127,232 Standard error for estimated volume (±) 64,575 2,094,414 658,512 1,472,100 2,660,022 3,482,031 76,791 — 1,098,577 410,364 3,429,467 355,672 655,318 1,145,401 1,020,943 261,653 508,131 65,804 2,646,783 1,349,345 959,938 — 569,175 395,379 1,881,143 332,830 273,510 4,154,971 9,640,055 5,211,646 43,038 29,421 6,152,703 127,365 Palau’s Forest Resources, 2003 Elaeocarpus graeffei Seem. Elaeocarpus joga Merr. Erythrina fusca Lour. Eugenia cumini (L.) Druce Eugenia javanica Lam. Eugenia reinwardtiana (Bl.) DC. Eugenia sp. L. Evodia sp. Fagraea ksid Gilg and Bened. Ficus sp. L. Ficus tinctoria Forst. f. Flacourtia rukam Zoll. & Moritzi Garcinia matudai Kaneh. Garcinia rumiyo Fosb. Garcinia sp. L. Gironniera celtidifolia Gaudich. Glochidion ramiflorum Forst. Glochidion sp. J.R. & G. Forst. Gmelina elliptica Sm. Gmelina palawensis H. J. Lam. Gmelina sp. L. Guettarda speciosa L. Gulubia palauensis (Becc.) Moore & Fosb. Hernandia sonora L. Hernandia sp. L. Heterospathe elata (Becc.) Becc. Hibiscus tiliaceus L. Horsfieldia amklaal Kaneh. Horsfieldia novo-guineensis Warb. Horsfieldia palauensis Kaneh. Horsfieldia sp. Wild. Inocarpus fagifer (Park.) Fosb. Intsia bijuga (Colebr.) O. Ktze. Lumnitzera littorea (Jack) Voigt Common name Estimated number (≥1 inch diameter) Scientific name Common name Macaranga carolinensis Volk. Manilkara udoido Kaneh. Manilkara zapota (L.) P. v. Roy. Morinda citrifolia L. Morinda latibracteata Val. Morinda pedunculata Val. Morinda sp. L. Mussaenda frondosa L. Neonauclea forsteri Merrill Nephelium lappaceum L. Osmoxylon oliveri Fosberg & Sachet Osmoxylon sp. Pandanus aimiriikensis Mart. Pandanus dubius Spr. Pandanus kanehirae Mart. Pandanus palawensis Mart. Pandanus tectorius Park. Pandanus utiyamai Kaneh. Pangium edule Reinw. ex Bl. Parinari corymbosum (Bl.) Miq. Parinari laurina Gray Parinari sp. Pinanga insignis Becc. Pithecellobium dulce (Roxb.) Benth. Pongamia pinnata (L.) Merr. Pouteria calcarea (Hosok.) Fosb. Pouteria obovata (R. Br.) Baehni Pouteria sp. Aubl. Premna obtusifolia R. Br. Pterocarpus indicus Willd. Ptychosperma palauensis (Kaneh.) Moore & Fosb. Rhizophora apiculata Bl. Rhizophora mucronata Lam. Rhus taitensis Guill. bedel udeuid sapodilla ngel kesengelengel, ngel kesengelengel rambutan, rambotang ertochet beku, ongor buuk ongor ongor riamel bkau, apgau eritem, gritin ebouch, demailei kamatsiri, opiuma kisaks elangel elangel osem las esbuuch bngaol tebechel eues Number measured 49 29 6 8 5 1 3 1 3 1 3 9 25 21 6 1 13 1 2 63 56 4 240 4 1 3 37 1 9 2 8 129 31 40 Estimated number (≥1 inch diameter) 5,074,421 283,579 73,977 946,172 61,648 12,330 36,989 153,586 460,757 12,330 178,245 675,990 3,130,866 810,575 202,904 153,586 559,393 12,330 24,659 2,008,183 2,086,261 331,830 11,655,307 465,410 151,094 36,989 855,954 12,330 381,148 24,659 239,892 3,657,133 365,084 714,045 Standard error for estimated number (±) 1,760,927 173,439 62,652 352,952 50,610 12,342 37,027 153,747 461,240 12,342 155,148 639,395 1,232,969 608,016 190,886 153,747 274,312 12,342 24,685 672,025 1,251,377 223,617 2,337,956 510,641 165,778 37,027 637,145 12,342 309,108 24,685 240,144 2,280,225 327,871 251,240 Gross volume (≥5 inches diameter) 671,260 2,085,556 1,296,562 132,139 211,608 81,626 127,267 — — 100,828 65,896 145,875 272,360 897,251 332,031 — 463,602 1,050,209 237,335 21,934,439 5,980,513 66,714 8,329,960 80,161 — 160,981 4,146,921 59,315 246,764 917,560 329,740 8,538,981 2,265,232 5,850,777 Standard error for estimated volume (±) 357,840 1,209,721 1,181,412 99,769 165,154 81,712 127,400 — — 100,934 65,965 86,232 167,696 621,269 262,594 — 464,088 1,051,312 237,584 7,016,031 3,010,760 48,675 1,802,335 87,951 — 161,150 1,701,642 59,377 165,406 918,523 330,086 5,332,588 1,681,146 2,434,616 RESOURCE BULLETIN PNW-RB-252 Table 1—Scientific names, common names, estimated number and gross volume, and standard errors for estimated totals of species measured as trees on Palau (continued) Table 1—Scientific names, common names, estimated number and gross volume, and standard errors for estimated totals of species measured as trees on Palau (continued) Scientific name Rinorea carolinensis Kaneh. Semecarpus venenosus Volk. Sonneratia alba J. E. Sm. Stemonurus ammui (Kaneh.) Sleumer Swietenia macrophylla King Swietenia mahagoni (L.) Jacq. Symplocos racemosa Roxb. Tecoma stans (L.) Juss. ex HBK. Terminalia catappa L. Theobroma cacao L. Timonius subauritus Fosberg & Sachet Tournefortia argentea L. f. Trichospermum ledermannii Burret Vitex cofassus Reinw. ex Bl. Xylocarpus granatum Koen. Unknown 0 Unknown 1 Unknown 2 Unknown 3 Unknown 5 Unknown 11 Unknown, other Total Common name tonget urur ngmui honduras mahogany mahogany beach almond, miich suklatei, cocoa rirs, tree heliotrope elsau bars, beokel meduulokebong Number measured Estimated number (≥1 inch diameter) Standard error for estimated number (±) Gross volume (≥5 inches diameter) Standard error for estimated volume (±) 1 58 52 5 5 8 19 1 4 5 19 4 11 9 16 13 6 2 4 7 1 7 12,129 2,266,927 5,959,545 61,648 202,904 381,148 829,575 153,586 49,318 61,648 1,595,842 48,918 252,222 110,966 338,528 188,082 331,630 307,171 24,659 85,906 12,330 36,989 13,308 689,153 3,934,040 61,712 179,561 345,648 509,114 153,747 38,757 61,712 1,090,534 36,301 167,347 111,082 292,559 168,017 215,243 307,493 17,251 67,207 12,342 20,875 44,758 6,877,953 11,503,833 3,215,714 327,606 414,720 224,606 — 805,001 617,845 925,779 327,509 1,067,694 5,134,682 1,399,499 120,479 37,791 — 228,639 513,431 126,165 1,657,879 49,108 2,306,382 11,491,789 3,219,089 235,724 307,348 143,427 — 657,530 618,494 614,886 270,923 945,028 5,140,070 1,140,977 85,328 32,354 — 188,570 452,329 126,298 1,471,197 2,248 96,594,950 6,639,026 268,511,801 26,355,589 Palau’s Forest Resources, 2003 RESOURCE BULLETIN PNW-RB-252 Interior volcanic forest— Occurring primarily on volcanic substrates, interior volcanic forest encompasses distinct subtypes of forest in undisturbed ecosystems. Moving inland from the mangroves and swamps, the dominant species include Casuarina equisetifolia and Pandanus tectorius. Farther inland, the forest grades into a mixed-forest canopy that includes Campnosperma brevipetiolata, Horsfieldia amklaal, H. palauensis, H. novo-guineensis, Myristica insularis, Parinari laurina, Parinari corymbosum, Serianthes kanehirae, Glochidion macrosepalum, G. ramiflorum, Alphitonia carolinensis, Calophyllum inophyllum var. wakamatsui, C. soulattri, C. pelewense, Symplocos racemosa var. palauensis, Cerbera floribunda, Manilkara udoido, and Fagraea ksid. The forest is dense, multilayered, and structurally complex here. Ravine and riparian forest— In the complex terrain of Palau’s savannas, ravines often break up the open forest and grassland with narrow strips and sheltered hillsides of more dense and diverse forest vegetation occurring in relatively moist areas. Barringtonia racemosa and the poison tree (causes a severe skin rash) Semecarpus venenosus are common components of the ravine and riparian type. Limestone forest type On the limestone substrate of Peleliu, Angaur, and the Rock Islands, organic matter from the vegetation forms a thin layer in places over the coral rock. The limestone substrate is often steep, porous, and extremely rugged. Rock that borders the ocean is often undercut by waves and bioerosion, making access difficult. A species-rich and structurally diverse forest forms here with the dominant species Intsia bijuga, Psychotria spp., Clerodendron inerme, Eugenia reinwardtiana, Morinda latibracteata, Garcinia matudai, G. rumiyo var. clacicola, Rinorea sp., Cycas circinalis, Flacourtia rukam var. micronesica, Aidia cochinchinensis, Meryta senfftiana, Polyscias grandifolia, Geniostoma sessile, Premna serratifolia, Cyrtandra todaiensis, Guettarda speciosa, Badusa palauensis, Psychotria hombroniana, Ixora casei, and Tarenna sambucina. Field Methods The Palau inventory was based on the FIA inventory design that was implemented across the mainland United States beginning in 2000. Adaptations were made to the national design to include additional branching and rooting forms, using topography to define site productivity or drought resilience (in perhumid climates), additional tree crown measurements, and including special-interest species ranging from invasive plants to pathogens to culturally or economically important species 10 Palau’s Forest Resources, 2003 of various life forms. In the mainland FIA Program, plots are spaced within forest land on a 3.3-mile grid. With the assistance of the Government of Palau, Bureau of Agriculture, plots were spaced across all vegetation types at approximately 1.9-mile intervals, yielding a triple intensification of the mainland inventory plot grid. The FIA plot-cluster is composed of four 24-foot-radius subplots (fig. 3). Three of those subplots are equally spaced, as if on spokes of a wheel, around the central subplot. The distance from the middle of the central subplot to the middle of each subplot on the three spokes was 120 feet. Forest Inventory and Analysis Pacific Islands Plot Design Overall plot footprint: ~1.5 acres Distance between subplot centers: 120 feet Distance from subplot center to microplot center: 12 feet Key 6.8-foot-radius microplot: seedlings + saplings 24-foot-radius subplot: all trees (5 inches diameter at breast height) Plot 2 Subplot Microplot N 1 4 3 Figure 3—Forest Inventory and Analysis Pacific Islands plot design. 11 RESOURCE BULLETIN PNW-RB-252 A variety of information was collected at the plot, subplot, and tree levels (USDA Forest Service 2003). Differences in forest-type conditions are also mapped. For example, roads that intersect subplots are mapped, as are clear boundaries in forest tree size classes. The primary variables collected include plot location, slope, aspect, elevation, subplot slope position and shape, tree species, diameters, heights, damages, branching and rooting forms, decay, epiphytic loadings, crown characteristics, tree locations, and regeneration information. The fieldwork for this inventory was performed February to April 2003. Analysis Methods The FIA estimates of forest land are based on a system that uses aerial photography or satellite imagery to define different types of land (strata) across the landscape. The simplest stratification is separating land into forest and nonforest strata. However, stratifications can be assisted or refined by using ancillary data such as topography, soil information, life zone or climatically based information, and prior inventories of vegetation groups. The Palau stratification for estimating numbers of trees, volume, biomass, and carbon was conducted via a classification of 2003 IKONOS satellite data. The initial land-cover classification divided the landscape into forest, urban, nonforest vegetation, barren, and water cover types. At least 10 percent cover of trees was used as the basis for the forest-land classification, and includes both agroforests and mangrove forests. Nonforest vegetation includes other vegetation types with less than 10 percent cover of trees. A geographic information system (GIS) was used to sum acreage for each type. Forest was further classified into volcanic and limestone forest types by overlaying a soils layer (USDA Natural Resources Conservation Service 1983) on the land-cover-classification layer in a GIS. We used the GIS to individually sum the acreage of limestone and volcanic forest types. From this limestone/volcanic stratification, field plot condition types were expanded to the landscape level. The acreage each field plot represents was derived by dividing the total acreage of forest in each of the limestone and volcanic soil types by the number of field plots that fell within the geographic bounds of that specific type. Average stand size was expanded from the plots to the landscape level by using the same expansion factors. Land cover was further refined into more detailed classes through an ongoing cooperative project with the USDA Forest Service, Pacific Southwest Region, Remote Sensing Laboratory. Both the 2003 IKONOS and 2005 QuickBird satellite imagery were used for the detailed mapping and served as the basis for land cover estimates. 12 Palau’s Forest Resources, 2003 Wood volume was estimated for individual trees by using tree height and two stem diameter measurements. These measurements are expanded to tree-level volume estimates by using equations for sections of a cone. Both gross stem volume and net stem volume estimates were calculated. Net stem volume subtracts damage and rotten defects from gross stem volume. Biomass for individual tree stems was estimated by using the specific gravity for known species (30 out of 129 species measured on Palau have known specific gravities). For species where specific gravity was not known, an average specific gravity, according to forest type, was used. These estimates of aboveground tree biomass are derived from bole volume and include only biomass for the main stem, excluding branches, roots, and foliage. An additive combination of relative density and relative basal area (importance value) was used to classify forest types and assess the species dominance in a stand. Traditional site productivity estimates require forest stand age, derived from the annual rings of forest trees. Because tropical trees do not produce consistent annual rings, a modified topographic relative moisture index (TRMI) (Parker 1982) was examined as a proxy for site productivity. The TRMI used a weighted, additive combination of slope steepness, slope shape, and slope position to assess the potential moisture retention in a forested stand. Remeasurement data from these plots at the next inventory cycle will provide a better estimate of productivity than TRMI. In very wet environments like Palau, TRMI may be best considered a drought resilience proxy, predicting where soil and water accumulates, owing to topography, across a landscape. We also recognize moisture is not likely to be the only factor limiting tree growth and that a prolonged excess of moisture can be detrimental to the growth of many tree species. Reliability of FIA Data The area of forested land cover types classified from the IKONOS satellite imagery was assumed to be accurate and was used as the basis for the expansion of the numbers of trees, tree volume, and tree biomass from the plot scale to the foresttype scale. Possible sources of error not accounted for in our estimates include errors in the land cover map owing to incorrect interpretation of the image, errors from rounding when working with pixel-based imagery, and measurement errors on field plots. Standard errors for the expansion of our estimates from field plots to the forested landscape were calculated according to the proportion of area occupied by either volcanic or limestone forest types. Volcanic and limestone forest areas were treated as known rather than estimated, and variance was calculated by using methods in Cochran (1977). Using one standard error as our basis for evaluation gives a 68 percent chance that the true total gross tree stem volume on Palau lies between 13 RESOURCE BULLETIN PNW-RB-252 275,061,708 and 329,711,230 cubic feet. There is a 68-percent chance that the true number of trees (>1 inch diameter) on Palau lies within the range of 89,955,926 to 103,233,978. Readers are cautioned to examine the standard errors associated with species-level estimates for the number of individuals and volume by tree species in table 1. Resource Highlights Land Cover Palau is currently about 82 percent forested, which includes agroforest and secondary forest vegetation (table 2). About 2 percent of the landscape was classified as urban land and about 14 percent was classified as nonforest vegetation including savanna and agricultural lands. The area for land cover estimates totaled 110,028 acres. A prior vegetation survey (Cole et al. 1987) delineated vegetation types by using 1976 aerial photography, and estimated forest cover to be 80 percent of total land, including secondary vegetation and agroforest (table 2). Nonforest vegetation was estimated to be about 20 percent of the total land cover. Total acreage calculated from the Cole et al. (1987) maps that FIA recently scanned and digitized was 101,851 acres. The methodology and the area sampled for land cover estimates differ from those used for prior estimates. The Cole et al. (1987) work surveyed and mapped less limestone forest and defined forested lands on the basis of a canopy cover of approximately ≥ 30 percent. The recent effort used a forest-land canopy threshold of ≥ 10 percent to remain consistent with national FIA standards and the recent FIA work in American Samoa and Guam. In the current survey we mapped roads as urban land. To compare similar sample areas on Palau, we summarized land cover area on an island-by-island basis for Babeldaob, Peleliu, Angaur, and Koror (fig. 4). Although the difference in canopy cover thresholds between 1987 and 2005 still exists, trends suggest there may be some areas of Babeldaob where nonforest vegetation has reverted to forest. On Peleliu and Koror, it appears that forest land is being converted to urban and nonforest vegetation uses. These observations are tentative pending a thorough analysis of land cover change for Palau. 14 Palau’s Forest Resources, 2003 Table 2—Estimated area by land cover, 1976 a and 2005 Land cover 1976 Limestone forestb Volcanic forest 9,626 71,948 Total forest land c Acres 17,316 73,369 81,574 90,685 1,550 18,250 339 139 2,433 15,329 1,103 64 414 20,277 19,343 101,851 110,028 Nonforest and other areas: Nonforest urban Nonforest vegetation Barren lands Unclassified Water Total nonforest and other land Total area in survey 2005 a Figures for maps derived from 1976 aerial photography differ slightly from those published by Cole et al. (1987). Figures here were derived by scanning the historical maps and aggregating area totals with a geographic information system. b The sample area for forest cover, especially limestone types, was increased between the 1976 and 2005 remotely sensed imagery. c Forest land includes agroforest and secondary forest vegetation. 60,000 40,000 1988 2005 2,000 1,500 500 0 0 F N ore o n st ve fo ge re t a st tio Ba n r re n U rb an W at er U nk no w n To ta l Land cover 2,000 2,500 1988 2005 2,000 Acres 3,000 Land cover Peleliu 1,500 1,000 500 0 0 Land cover Angaur 1988 2005 F N ore o n st ve fo ge re t a st tio Ba n r re n U rb an W at er U nk no w n To ta l 1,000 F N ore o n st ve fo ge re t a st tio Ba n r re n U rb an W at er U nk no w n To ta l Acres 4,000 1988 2005 1,000 20,000 5,000 Koror 2,500 F N ore o n st ve fo ge re t a st tio Ba n r re n U rb an W at er U nk no w n To ta l Acres 80,000 3,000 Babeldaob Acres 100,000 Land cover Figure 4—Forest land on Babeldaob is maturing and may be encroaching slightly on nonforest vegetation. On Peleliu, Koror, and Angaur, forest land is being lost to urban and nonforest vegetation land uses. Conversion of forest to urban land use is also expected on Babeldaob upon completion of the circumnavigating “Compact Road” in 2006, that will open areas previously inaccessible. 15 Productivity and Drought Resilience Site productivity estimates are difficult to obtain in tropical environments owing to a lack of consistent annual tree rings. The standard premise for site productivity hinges on determining how height or diameter changes with respect to tree age under particular site conditions. Remeasurement of individual trees at the next inventory cycle will provide better estimates of site productivity by using growth over the specific period. For the present work, we attempted to estimate productivity as an additive index based on a site’s potential moisture accumulation owing to local topography (TRMI) (Parker 1982). However, in a region of such high precipitation, as in Palau, the TRMI estimate of productivity may be much less reliable than productivity estimates based on detailed soil mapping. For example, differences among forest, savanna, and fernland topsoils are dramatic with respect to soil organic matter, cation exchange capacity, and potential aluminum toxicity for plant growth, with a succession of soil degradation occurring from forest soils to savanna and fernland soils through processes of land clearing, repeated burning, and subsequent erosion and leaching of topsoil nutrients (Smith and Babik 1988). The topographic relative moisture index may have utility in very wet environments as a predictor of resilience to occasional drought, because TRMI predicts moisture accumulation (and likely sediment/soil accumulation as well) based on topography. Using TRMI in this context, we classified approximately 58 percent of the forested lands in Palau as medium high to highest drought resilience (fig. 5). This relatively high percentage owes to much of the land on Babeldaob being gently sloping volcanic soils, and thus drought resilience on these lands is expected to be relatively high. Babeldaob represents approximately 80 percent of our inventory area. On the limestone and limestone-derived soils of the Rock Islands, resilience to drought would be expected to be less based on lower moisture retaining capacities of these well-drained, often shallow soils and the complex karst topography. However, aluminum toxicity to plants is not expected on limestone-derived soils, whereas it is likely to limit plant growth in degraded volcanic-derived soils. Forest Structure Forest Inventory and Analysis estimates stand size class on forested field plots to capture the predominant diameter of live trees in forest stands. In Palau the dominant size class for volcanic forest types tends to be small, in the 5- to 11-inch diameter category (table 3, fig. 6). Limestone forest types tend to be dominated by slightly larger trees, in the 11- to 20-inch category (fig. 6). No plots were sampled on limestone soils that included stands in the smallest diameter category of 1 to 5 inches. 16 Forest land potential drought resilience (Percentage of total forest area and acreage) 10% 10,055 6% 5,940 13% 12,260 Low Medium low Medium 29% 28,546 23% 21,973 Medium high High Highest 19% 17,915 Figure 5—A topographic relative moisture index was used to predict drought resilience in the perhumid climate of Palau. On the gentle slopes and relatively deep volcanic soils of the largest island, Babeldaob (approximately 80 percent of forested acreage on Palau), drought resilience is expected to be higher than in the raised limestone Rock Islands. The diameter distribution for trees on Palau is heavily weighted toward small-diameter trees (fig. 7). This “reverse-J” distribution of tree diameter is typical where regeneration is abundant. Through time, many small trees die before they grow into the larger diameter classes. For trees at least 5 inches in diameter, tree heights are predominantly in the 30- to 50-foot height classes (fig. 8). The tallest tree on an FIA plot, a Horsfieldia sp., measured 44 inches in diameter and approximately 114 feet tall. Wood volume (fig. 9, tables 4, 5, and 6), biomass (fig. 10, table 7), and carbon storage (table 8) follow a similar distribution as the “reverse-J” seen for the number of trees according to diameter (fig. 7). Most of the volume, biomass, and carbon is accounted for in the smaller diameter classes owing to the relatively high number of trees in those size classes. (Text continues on page 33.) 17 18 Table 3—Estimated number of live trees on forest land by species and diameter class Diameter class (inches) Species Aglaia palauensis Allophylus ternatus Allophylus timorensis Alphitonia carolinensis Annona reticulata Areca catechu Artocarpus mariannensis Astronidium palauense Avicennia marina Badusa palauensis Barringtonia racemosa Bruguiera gymnorrhiza Buchanania palawensis Calophyllum inophyllum Calophyllum pelewense Calophyllum soulattri Calophyllum sp. Campnosperma brevipetiolata Canarium hirsutum Casearia cauliflora Casearia sp. Casuarina equisetifolia Cerbera floribunda Cerbera manghas Cocos nucifera Commersonia bartramia Cordia micronesica Cyathea lunulata Diospyros ferrea Diospyros sp. Dodonaea viscosa Dracaena multiflora Drypetes nitida Dysoxylum sp. Elaeocarpus graeffei Elaeocarpus joga Erythrina fusca Eugenia cumini Eugenia javanica Eugenia reinwardtiana Eugenia sp. Evodia sp. ≤5.0 5.0–6.9 7.0–8.9 9.0– 10.9 614,342 — — 1,075,099 — — — 1,382,270 — 606,866 3,064,235 614,342 307,171 916,529 460,757 307,171 — 307,171 455,773 — — — — 153,586 — 153,586 — 153,586 307,171 458,265 — 767,928 151,094 1,535,855 — 916,529 — 755,468 151,094 2,434,941 611,850 153,586 122,495 — 12,330 197,272 12,330 61,648 — 184,943 — 121,295 135,224 135,625 73,777 36,588 61,447 12,330 — 123,095 36,588 — 60,647 12,129 12,330 61,447 — 12,330 — 258,920 49,318 49,118 12,330 73,377 12,330 36,989 12,330 159,083 — 109,765 36,388 255,319 36,788 — 36,989 36,989 — 135,625 — 12,330 — 36,989 — 24,259 86,106 172,613 — 61,247 61,648 12,330 24,459 98,636 24,459 12,330 12,129 — — 36,989 — — — 147,954 36,989 24,459 12,129 24,459 12,129 — — 171,813 — 48,718 48,518 145,554 — — 49,318 — — 36,989 — — — 36,989 12,330 84,906 61,047 86,307 12,330 36,588 — — 12,129 86,307 12,129 — 12,129 — — 61,648 85,706 — — 49,318 — — 12,129 — — — — 12,330 — 12,129 60,647 97,036 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 — — — 61,648 — — — — — 24,259 48,718 24,659 — 48,518 24,659 24,659 — 147,954 12,129 — — — 12,330 24,659 171,813 — — 24,659 — 12,330 — 12,330 — — — 86,106 — 24,459 12,129 48,518 — — — — — — — — 12,129 — — 24,259 12,129 49,318 — 12,129 — — — 98,636 — — — — — 12,330 49,118 — 12,330 — — — — — 12,330 — — 36,388 — 12,129 12,129 12,129 — — Number of trees 24,659 — — — — — 24,659 — — — — — — — — — — — 12,129 — — — — — — — 24,259 — — — — — — 12,330 86,307 73,977 — 12,129 — — — — — — — — 12,330 — — 12,330 — — — — — — — — — — 24,259 — — — — — — — — — 36,788 — — 12,129 12,129 — 24,259 12,129 12,129 12,129 — — — — 19.0– 20.9 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ — — — — — — 12,129 — — — — — — 36,588 — — — 36,989 — — — — — — — — — — — — — — — — — — — — 12,129 — — — — — — — — — — — — — — — 12,330 — — — 12,330 36,989 — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — 12,129 — — — — — — 12,129 — — — 36,989 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,129 — 12,330 — — — — — — — — — — — — 12,129 — — — — — — — — — — — — — — — — 12,129 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — All classes 847,803 36,989 12,330 1,531,290 12,330 73,977 48,518 1,641,189 24,659 897,973 3,407,461 1,095,193 405,607 1,184,578 608,511 356,489 61,247 1,133,048 553,209 12,330 84,906 24,259 24,659 362,987 318,967 165,915 12,330 634,436 393,478 544,171 73,177 878,093 187,882 1,572,844 12,330 1,419,038 12,129 974,799 381,553 3,017,755 648,639 153,586 Table 3—Estimated number of live trees on forest land by species and diameter class (continued) Diameter class (inches) Species 19 Fagraea ksid Ficus tinctoria Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Guettarda speciosa Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo–guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata Morinda pedunculata Morinda sp. Mussaenda frondosa Neonauclea forsteri Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus palawensis Pandanus tectorius Pandamus utiyamai ≤5.0 5.0–6.9 7.0–8.9 — 1,072,607 1,228,684 921,513 2,150,197 307,171 — 1,369,810 153,586 606,866 460,757 — 460,757 — — — 614,342 2,610,954 757,960 767,928 2,895,698 614,342 — 151,094 — 4,914,737 — — 921,513 — — — 153,586 460,757 — 153,586 614,342 3,069,219 604,374 153,586 153,586 460,757 — 49,318 — 110,565 36,989 135,425 221,131 24,659 85,506 24,659 60,847 234,261 — — 97,036 12,129 24,259 61,648 49,318 146,954 86,307 478,850 12,330 12,330 12,129 — 110,565 197,272 24,659 — 61,648 12,330 36,989 — — — 24,659 61,648 36,989 181,942 — — 73,977 — — 12,129 36,989 12,330 85,706 49,118 12,330 85,506 — 24,259 135,625 24,659 — — 12,129 — 24,659 49,318 36,788 36,989 208,201 — — 48,518 — 36,989 36,989 — 24,659 — — — — — 12,330 — — 12,330 24,259 49,318 — 24,659 — 9.0– 10.9 61,648 — — 12,330 49,118 48,718 — 12,129 — 24,259 86,106 12,330 — — — 12,129 — — — 49,318 159,283 — — 24,259 12,330 12,129 12,330 12,330 — — — — — — — — — 12,330 — — — — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 24,659 — 12,330 — 36,989 24,659 — 12,330 — 24,259 49,118 12,330 — — 12,129 12,129 — — 12,330 — 48,918 — — 12,129 — — — 12,330 — — — — — — — — — — — — — — — 12,330 — — 12,330 — — — — — 24,259 36,989 12,330 — — — — — — 36,788 12,330 12,129 — — 24,259 — — 12,330 — — — — — — — — — — — — — — — — Number of trees 24,659 — — — — — — — — — — — — — — — — — 24,459 12,129 — 24,459 — 12,330 — — — — — — — — — — — — — — 12,330 12,330 — — — — — — 24,259 24,259 — — — — 12,330 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 19.0– 20.9 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 — — — — — — — — — 12,330 — — — — — — — — — 12,330 12,129 — — 12,129 — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,129 — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,129 — — — — — — — 12,129 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — 12,129 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — 29.0+ — 12,129 — — — — — — — — — — — — — — — — 12,330 12,330 24,659 — — 12,129 — — — — — — — — — — — — — — — — — — — All classes 172,613 1,096,866 1,388,568 995,490 2,457,435 650,797 36,989 1,565,282 178,245 825,997 1,027,314 73,977 460,757 97,036 36,388 60,647 700,649 2,709,590 1,003,150 1,002,188 3,852,198 626,672 12,330 381,553 12,330 5,074,420 283,579 73,977 946,172 61,648 12,330 36,989 153,586 460,757 12,330 178,245 675,990 3,130,866 810,575 202,904 153,586 559,393 12,330 20 Table 3—Estimated number of live trees on forest land by species and diameter class (continued) Diameter class (inches) Species Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pongamia pinnata Pouteria calcarea Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni Symplocos racemosa Tecoma stans Terminalia catappa Theobroma cacao Timonius subauritus Tournefortia argentea Trichospermum ledermannii Vitex cofassus Xylocarpus granatum Unknown 0 Unknown 1 Unknown 2 Unknown 3 Unknown 5 Unknown 11 Unknown, other Total ≤5.0 — 1,379,778 1,520,904 307,171 9,522,303 453,281 151,094 — 453,281 — 307,171 — 153,586 2,288,831 — 307,171 — 1,689,441 5,836,250 — 153,586 307,171 767,928 153,586 — — 1,510,936 — 153,586 — 153,586 151,094 307,171 307,171 — — — — 5.0–6.9 — 184,743 282,778 24,659 2,083,686 — — 36,989 134,424 — 49,318 — 86,307 782,687 242,589 86,307 — 271,049 12,330 — 36,989 49,318 49,318 — — 12,330 36,388 12,129 36,989 — 61,648 36,989 12,129 — 12,330 24,259 — 12,330 7.0–8.9 12,330 73,777 86,106 — 36,989 12,129 — — 134,224 12,330 24,659 — — 401,472 85,506 98,636 12,129 135,224 24,659 12,330 — 12,330 12,330 — 12,330 24,659 24,259 12,129 24,659 12,330 24,659 — 12,330 — — 36,989 — — 9.0– 10.9 12,330 36,989 98,436 — 12,330 — — — 73,177 — — — — 110,565 12,330 61,648 — 73,177 — — 12,330 12,330 — — 24,659 — 12,129 12,330 24,659 — 36,989 — — — — 24,659 12,330 12,330 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 — 61,648 48,918 — — — — — 36,388 — — — — 36,788 24,659 61,648 — 49,118 12,330 — — — — — 12,330 24,659 — — 12,330 — 12,330 — — — 12,330 — — — — 86,307 — — — — — — — — — — — 36,788 — 61,648 — 12,129 — — — — — — — — — 12,330 — 12,330 36,989 — — — — — — — Number of trees — — 12,330 12,330 36,989 — — — — — — — — — — — — 12,129 — — — — 12,330 — — — — — — — 24,659 12,330 — — 12,129 — 12,330 — 12,330 12,330 — — — — — — — — — — — — 12,129 — — — — — 24,659 12,330 — 12,330 — — — — — — — — — — — — — — 19.0– 20.9 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 — 12,330 — — — — — — 12,330 — — — — — — — — — — 24,659 — — — — — — — — — — — — — — — — — — — 24,659 — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — 24,659 — — — — — — — — — — — 12,330 49,318 12,330 12,129 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — 24,659 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — 29.0+ All classes — 24,659 49,318 2,008,182 — 2,086,261 — 331,830 — 11,655,307 — 465,410 — 151,094 — 36,989 — 855,954 — 12,330 — 381,148 — 24,659 — 239,892 — 3,657,133 — 365,084 — 714,045 — 12,129 12,330 2,266,927 36,989 5,959,545 — 61,648 — 202,904 — 381,148 — 829,575 — 153,586 — 49,318 — 61,648 — 1,595,842 — 48,918 — 252,222 — 110,966 — 338,528 — 188,082 — 331,630 — 307,171 — 24,659 — 85,906 — 12,330 12,330 36,989 76,110,228 10,058,255 4,047,132 2,219,238 1,558,645 822,076 551,827 318,767 220,731 160,084 122,295 135,024 73,777 196,872 96,594,950 Palau’s Forest Resources, 2003 70,000 57,612 60,000 Area (acres) 50,000 Volcanic Limestone 40,000 30,000 20,000 10,000 0 14,202 12,091 6,272 6,511 Less than 5 5–10.9 Stand size class (inches) 11–19.9 Figure 6—The trees that characterize volcanic forests tend to be somewhat small, in the 5- to 11-inch category. Forested stands on limestone soils tend to be characterized by larger trees in the 11- to 20-inch category. 60 Trees (millions) 50 Number of trees by size class Total trees in all classes: 94,598,338 40 30 20 10 1. 0– 2 3. .9 0– 4 5. .9 0– 6 7. .9 0– 9. 8.9 0– 11 10 .0 .9 – 13 12. .0 9 – 15 14 .0 .9 – 17 16 .0 .9 – 19 18. .0 9 – 21 20. .0 9 – 23 22 .0 .9 – 25 24. .0 9 – 27 26. .0 9 –2 8. 9 29 .0 + 0 Diameter (inches) Figure 7—Small trees are very common in Palau, indicating regeneration is abundant. 21 RESOURCE BULLETIN PNW-RB-252 Number of trees (millions) 7 6 Number of trees by height class 5 (>5 inches diameter) 4 3 2 1 + 9 10 0 –9 90 9 70 –7 9 80 –8 9 –6 9 60 –5 9 50 –4 9 40 –3 30 10 –1 9 20 –2 9 <1 0 0 Height (feet) 90 80 Volume by size class 70 Total volume in all classes: 268,020,715 60 50 40 30 20 10 5. 0– 9 40 + 0 .9 10 .0 –1 4. 9 15 .0 –1 9. 9 20 .0 –2 4. 9 25 .0 –2 9. 9 30 .0 –3 4. 9 35 .0 –3 9. 9 Gross volume (million cubic feet) Figure 8—For trees greater than or equal to 5 inches in diameter, the most common heights attained range from 30 to 50 feet tall. Diameter (inches) Figure 9—As with numbers of trees, gross volume of wood tends to be concentrated in the smaller diameter classes. However, many larger diameter trees contribute to the substantial volume in those size classes. 22 Palau’s Forest Resources, 2003 Table 4—Estimated gross volume of all live trees on forest land by forest-type group and diameter class Diameter class (inches) Forest-type group Less than 5 5–10.9 11–19.9 Limestone forest Volcanic/ravine forest 6,248,751 28,116,999 28,309,959 62,596,123 Cubic feet 28,665,545 60,913,733 19,013,470 68,521,890 82,237,724 220,148,745 34,365,749 90,906,081 89,579,278 87,535,360 302,386,469 Total 20+ All sizes Table 5—Estimated net volume of all live trees ≥5 inches in diameter on forest land by forest-type group and diameter class Diameter class (inches) Forest-type group Limestone forest Volcanic/ravine forest 11–19.9 20+ All sizes 28,285,444 62,311,497 Cubic feet 28,570,500 18,577,815 60,297,635 67,289,502 75,433,759 189,898,634 90,596,941 88,868,135 265,332,393 85,867,317 1,400 1,200 Biomass by size class 1,000 Biomass total, all classes: 4,686,793 800 600 400 200 5. 0– 9 40 + 0 .9 10 .0 –1 4. 9 15 .0 –1 9. 9 20 .0 –2 4. 9 25 .0 –2 9. 9 30 .0 –3 4. 9 35 .0 –3 9. 9 Stem biomass (thousand tons) Total 5–10.9 Diameter (inches) Figure 10—Biomass is also concentrated in the smaller size classes. 23 24 Table 6—Estimated gross volume of all live trees on forest land by species and diameter class Diameter class (inches) Species 5.0–6.9 358,196 Aglaia palauensis — Allophylus ternatus 29,280 Allophylus timorensis 735,731 Alphitonia carolinensis 62,771 Annona reticulata 211,184 Areca catechu — Artocarpus mariannensis 577,119 Astronidium palauense — Avicennia marina 453,193 Badusa palauensis 452,159 Barringtonia racemosa 524,400 Bruguiera gymnorrhiza 243,769 Buchanania palawensis 188,139 Calophyllum inophyllum 204,921 Calophyllum pelewense 63,170 Calophyllum soulattri — Calophyllum sp. Campnosperma brevipetiolata 410,952 116,651 Canarium hirsutum — Casearia cauliflora 300,526 Casearia sp. 33,773 Casuarina equisetifolia 50,830 Cerbera floribunda 261,427 Cerbera manghas — Cocos nucifera 29,882 Commersonia bartramia — Cordia micronesica 683,380 Cyathea lunulata 128,287 Diospyros ferrea 187,930 Diospyros sp. 34,597 Dodonaea viscosa 204,143 Dracaena multiflora 47,105 Drypetes nitida 64,752 Dysoxylum sp. 64,508 Elaeocarpus graeffei 662,676 Elaeocarpus joga — Erythrina fusca 383,104 Eugenia cumini 127,922 Eugenia javanica 1,003,419 Eugenia reinwardtiana 130,011 Eugenia sp. 161,826 Fagraea ksid — Ficus sp. — Ficus tinctoria 7.0–8.9 217,203 206,199 — 1,129,350 — 80,981 — 208,231 — 211,682 527,080 1,038,683 — 500,540 498,086 79,559 164,982 750,845 119,701 157,802 118,182 — — 272,907 — — — 708,128 256,014 277,568 67,158 115,524 74,774 — — 1,399,992 — 338,074 369,502 1,093,777 — — — 56,385 9.0– 10.9 448,937 — — 467,218 — — — 364,270 114,276 816,673 711,656 871,613 140,678 392,480 — — 192,860 973,757 142,942 — 207,941 — — 719,166 1,364,708 — — 306,007 — — 163,789 — — — — 95,607 — 160,248 677,261 1,425,442 — 499,547 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 — — — 1,269,881 — — — — — 379,532 1,053,700 397,703 — 704,449 337,712 338,061 — 2,465,508 195,799 — — — 143,752 529,306 4,923,463 — — 277,547 — 251,528 — 147,157 — — — 1,623,421 — 380,749 190,441 930,288 — 481,374 — — — — — — — — 440,417 — — 578,607 229,549 852,655 — 194,770 — — — 2,491,979 — — — — — 267,966 1,481,204 — 312,315 — — — — — 251,142 — — 1,376,416 — 298,384 297,325 382,084 — 277,505 — — 759,169 — — 834,761 — — — — — 608,341 — — — 748,863 — — — 3,277,299 — — — — — 294,836 — — — — — — 846,599 — — — — 1,568,153 — 400,104 740,797 560,575 — 822,261 — — 17.0– 18.9 19.0– 20.9 Cubic feet — — — — — — — — — — — — — 635,097 — — — — — — — — — — — — — 1,760,060 — — — — 309,745 — 4,050,260 1,937,832 497,877 — — — — — — — — — — — 647,213 — — — — — — — — — — — — — — — — — — — — — — — 600,183 — — — 497,380 426,207 616,284 — — — — — — 409,933 — — 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ — — — — — — — — — — — — 924,553 — — — 1,243,385 3,038,783 — — — — — — — — — — — — 530,228 — — — — — — — — — — — — — — — — — — — 1,278,815 — — — — — — 947,598 — — — 3,494,230 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 2,134,102 — 1,308,417 — — — — — — — — — — — — 1,403,132 — — — — — — — — — — — — — — — — 1,092,791 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 1,857,843 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 2,606,279 All classes 1,783,505 206,199 29,280 4,436,940 62,771 292,165 4,488,431 1,149,620 1,422,693 3,048,028 2,974,144 5,542,897 1,308,999 5,436,901 1,040,719 480,790 1,910,972 22,891,446 1,072,970 157,802 626,648 1,436,905 194,582 2,345,608 8,416,588 29,882 312,315 1,975,061 384,300 717,026 1,642,371 466,823 373,021 64,752 64,508 6,726,266 600,183 1,960,664 4,419,626 6,011,868 130,011 2,242,513 409,933 2,662,664 Table 6—Estimated gross volume of all live trees on forest land by species and diameter class (continued) Diameter class (inches) Species 25 Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo-guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata Morinda pedunculata Morinda sp. Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus tectorius Pandanus utiyamai Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pouteria calcarea 5.0–6.9 7.0–8.9 399,756 181,521 547,736 937,863 89,179 296,385 93,117 218,166 860,819 — 518,759 48,678 72,835 223,139 157,229 604,676 303,272 1,795,603 61,419 29,390 60,415 — 328,807 753,108 95,965 — 211,608 81,626 127,267 — 65,896 145,875 136,228 677,136 — 261,072 — — 689,431 1,056,482 66,714 7,832,438 — 160,981 205,465 80,400 734,850 459,661 172,200 533,580 — 159,556 888,819 275,888 — 93,646 — 181,631 191,147 248,982 254,896 1,549,305 — — 420,363 — 186,047 207,350 — 142,104 — — — 100,828 — — 51,506 220,116 332,031 188,207 — 73,623 506,454 521,650 — 303,291 80,161 — 9.0– 10.9 — 155,242 507,022 605,835 — 139,993 — 363,533 771,909 190,681 — — 120,735 — — — 744,724 1,962,502 — — 256,073 127,232 167,974 226,930 119,687 — — — — — — — 84,627 — — — — 163,712 431,678 925,184 — 133,263 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 146,034 — 730,739 579,278 — 231,111 — 401,804 732,695 268,382 — 227,398 144,215 — — 270,510 — 870,606 — — 176,722 — — — 177,080 — — — — — — — — — — — — — 1,097,576 962,513 — — — — — 307,851 — — — — — 534,904 1,121,009 159,607 — — — — — 1,020,728 344,493 242,883 — — 609,515 — — 225,960 — — — — — — — — — — — — — — 1,990,397 — — — — — — — — — — — — 701,063 — — — — — — — — 396,646 — — — 1,153,528 — — 288,213 — — — — — — — — — — — — — — 421,904 1,115,534 — — — — 17.0– 18.9 19.0– 20.9 Cubic feet — — — — — — — — — — — — — — 694,331 513,192 673,159 — 420,704 — — — — — — — — — — — — — 806,216 930,642 — 856,766 — — — — 1,228,373 633,825 — — — — 383,995 — — 328,616 — — — — — — — — — — — — — — — — — — — — — — — — — — 852,296 607,489 — — — — — — — — — — 21.0– 22.9 23.0– 24.9 — — — — — — — — — — — — — — — — — — — — 742,335 — — — 575,215 — — — — — — — — — — — — — 1,822,127 — — — — — — — — — — — — — — — — — 1,376,732 — — — — — — — 1,089,139 — — — — — — — — — — — — — — — — — 1,386,894 1,399,150 — — — — 25.0– 26.9 27.0– 28.9 — — — — — — — — — — — — — — — 897,498 — — — — — — — — — — — — — — — — — — — 1,138,744 — — — — — 1,507,134 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 1,050,209 — — — 3,444,684 1,289,904 — — — — — — — — — — 29.0+ — — — — — — — — — — — — — — — 2,995,219 6,929,268 5,239,828 — — 1,462,131 — — — — — — — — — — — — — — — — — 7,877,115 — — — — — All classes 751,255 725,013 2,520,347 2,582,637 261,379 1,201,069 93,117 4,484,048 5,048,410 1,315,262 518,759 369,722 1,714,517 404,770 348,375 5,140,116 10,710,158 13,656,236 61,419 29,390 9,339,552 127,232 682,828 2,085,556 1,296,562 142,104 211,608 81,626 127,267 100,828 65,896 145,875 272,360 897,251 332,031 449,279 1,050,209 237,335 22,417,949 5,980,513 66,714 8,268,992 80,161 160,981 26 Table 6—Estimated gross volume of all live trees on forest land by species and diameter class (continued) Diameter class (inches) Species Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni Symplocos racemosa Terminalia catappa Theobroma cacao Timonius subauritus Tournefortia argentea Trichospermum ledermannii Vitex cofassus Xylocarpus granatum Unknown 0 Unknown 1 Unknown 3 Unknown 5 Unknown 11 Unknown, other Total 5.0–6.9 541,147 — 127,669 — 314,678 3,089,759 815,800 288,555 — 1,155,137 23,568 — 175,805 179,811 128,478 — 30,258 168,555 22,145 129,573 — 178,267 120,479 35,180 44,577 67,712 — 50,997 7.0–8.9 9.0– 10.9 11.0– 12.9 13.0– 14.9 15.0– 16.9 1,054,853 59,315 119,095 — — 864,486 — — — — 712,852 — — — — — — — — — — — — 229,537 — Cubic feet 595,227 392,699 — — — — — — — — — 1,461,139 — 312,069 — — — — — — — — 143,626 — 319,190 554,708 — — — — — — 495,846 — 526,363 538,634 633,032 — — — — — 438,429 — — 714,086 — — — — — — — 560,851 — — — 808,378 — — — — — — — — 435,761 239,527 — — — — — — 2,545,438 758,392 725,964 44,758 852,343 169,976 70,618 — 71,296 96,128 165,490 187,894 187,252 44,889 288,139 91,031 113,859 — 48,232 — 235,500 — — 1,320,834 183,142 586,929 — 954,517 — — 151,802 163,613 — 340,579 — 131,543 116,849 387,762 — 368,179 — — — 210,219 126,165 140,256 612,144 507,898 1,176,660 — 891,534 149,403 — — — — 298,932 399,693 — — 262,220 — 178,624 — — 184,062 — — — 882,643 — — 17.0– 18.9 — — 19.0– 20.9 — — — — — — 1,703,686 — — — — — — — — — — — — — — — — 21.0– 22.9 23.0– 24.9 — — — 488,213 — — — — — — — — — — — — — — — — — — — 1,255,579 — — — — — — — — — — — — — — — — — — — — — — — — 1,024,803 — — — — — — — 25.0– 26.9 — — — — — — — 27.0– 28.9 29.0+ — — — — — — — — — — — — — — — — 1,367,032 — — 2,270,776 — — — — — — — — — — — — — — 924,646 — — — — — — — — — — — — — — — — — — — — — — — 1,364,929 8,351,476 — — — — — — — — — — — — — — — — 1,440,229 All classes 4,161,263 59,315 246,764 717,750 314,678 8,450,819 2,265,232 5,295,943 44,758 7,423,924 11,503,833 3,215,714 327,606 414,720 224,606 805,001 617,845 925,779 327,509 1,067,694 4,765,096 1,633,164 120,479 83,412 228,639 513,431 126,165 1,631,482 37,070,470 28,333,126 25,502,485 29,412,054 19,963,042 19,114,573 14,917,757 11,136,046 10,620,417 11,997,362 12,725,013 7,104,057 40,124,318 268,020,719 Table 7—Estimated aboveground dry stem weight of all live trees on forest land by species and diameter class Diameter class (inches) Species 27 Aglaia palauensis Allophylus ternatus Allophylus timorensis Alphitonia carolinensis Annona reticulata Areca catechu Artocarpus mariannensis Astronidium palauense Avicennia marina Badusa palauensis Barringtonia racemosa Bruguiera gymnorrhiza Buchanania palawensis Calophyllum inophyllum Calophyllum pelewense Calophyllum soulattri Calophyllum sp. Campnosperma brevipetiolata Canarium hirsutum Casearia cauliflora Casearia sp. Casuarina equisetifolia Cerbera floribunda Cerbera manghas Cocos nucifera Commersonia bartramia Cordia micronesica Cyathea lunulata Diospyros ferrea Diospyros sp. Dodonaea viscosa Dracaena multiflora Drypetes nitida Dysoxylum sp. Elaeocarpus graeffei Elaeocarpus joga Erythrina fusca Eugenia cumini Eugenia javanica Eugenia reinwardtiana Eugenia sp. Fagraea ksid Ficus sp. Ficus tinctoria 5.0–6.9 7.0–8.9 9.0– 10.9 5,722 — 468 11,753 1,003 3,374 — 9,219 — 7,240 7,223 8,377 3,894 3,426 3,274 1,009 — 6,565 1,491 — 5,953 896 812 4,176 — 477 — 10,917 2,869 4,203 553 3,261 753 1,034 1,031 10,586 — 7,955 2,656 20,836 2,700 3,775 — — 3,470 3,294 — 18,041 — 1,294 — 3,326 — 3,382 8,420 16,593 — 9,115 7,957 1,271 3,426 11,995 1,530 2,521 2,341 — — 4,360 — — — 11,312 5,726 6,208 1,073 1,845 1,195 — — 22,365 — 7,020 7,673 22,712 — — — 703 7,172 — — 7,464 — — — 5,819 1,826 13,046 11,369 13,924 2,247 7,147 — — 4,005 15,556 1,827 — 4,119 — — 11,489 21,801 — — 4,888 — — 2,617 — — — — 1,527 — 3,328 14,063 29,599 — 11,652 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 19.0– 20.9 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes — — — 20,286 — — — — — 6,063 16,833 6,353 — 12,828 5,395 5,401 — 39,386 2,502 — — — 2,296 8,456 78,652 — — 4,434 — 5,625 — 2,351 — — — 25,934 — 7,906 3,955 19,317 — 11,228 — — — — — — — — 7,036 — — 9,243 3,667 13,621 — 3,547 — — — 39,809 — — — — — 4,281 23,662 — 5,287 — — — — — 4,012 — — 21,988 — 6,196 6,174 7,934 — 6,473 — — 12,128 — — 13,335 — — — — — 9,718 — — — 13,637 — — — 52,355 — — — — — 4,710 — — — — — — 13,524 — — — — 25,051 — 8,308 15,383 11,640 — 19,179 — — Tons — — — — — — — — — — — — — — — — 6,432 64,703 6,363 — — — — — 10,339 — — — — — — — — — — — 4,792 — 10,328 12,797 — — — — — — — — — — 10,146 — — — — — — 32,051 — — — 30,957 — — — — — — — — — — — — — — — — — — — — 8,850 — — — 4,190 — — — — — — — — — — — — — 14,770 — — — 25,819 48,545 — — — — — — — — — — — — 8,470 — — — — — — — — — — — — — — — — — — — 20,429 — — — — — — 17,256 — — — 55,820 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 34,092 — 20,902 — — — — — — — — — — — — 37,211 — — — — — — — — — — — — — — — — 22,692 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 29,679 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 32,474 28,491 3,294 468 70,880 1,003 4,667 71,703 18,365 22,728 48,692 47,512 88,548 20,911 99,006 16,625 7,681 39,681 365,691 13,713 2,521 12,414 38,107 3,108 37,471 134,455 477 5,287 31,552 8,595 16,036 26,237 7,458 5,959 1,034 1,031 107,452 4,792 40,713 91,774 124,836 2,700 52,307 4,190 33,177 28 Table 7—Estimated aboveground dry stem weight of all live trees on forest land by species and diameter class (continued) Diameter class (inches) Species 5.0–6.9 7.0–8.9 9.0– 10.9 Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo-guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata Morinda pedunculata Morinda sp. Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus tectorius Pandanus utiyamai Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pouteria calcarea 6,386 4,349 13,124 22,471 1,425 4,735 1,488 3,485 13,752 — 8,287 451 1,164 3,565 2,863 9,660 4,845 28,685 981 470 1,177 — 5,253 12,031 1,533 — 3,380 1,304 2,033 — 1,053 2,330 2,176 10,817 — 4,171 — — 16,739 22,952 1,449 125,123 — 2,572 3,282 1,926 17,607 11,013 2,751 8,524 — 2,549 14,199 4,407 — 868 — 2,902 3,481 3,977 4,072 24,750 — — 8,193 — 2,972 3,312 — 2,270 — — — 1,611 — — 823 3,516 5,304 3,007 — 1,176 12,297 11,333 — 4,845 1,281 — — 3,720 12,148 14,516 — 2,236 — 5,807 12,331 3,046 — — 1,929 — — — 11,897 31,351 — — 4,991 2,033 2,683 3,625 1,912 — — — — — — — 1,352 — — — — 2,615 10,481 20,100 — 2,129 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 19.0– 20.9 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes 2,333 — 17,509 13,880 — 3,692 — 6,419 11,705 4,287 — 2,107 2,304 — — 4,321 — 13,908 — — 3,444 — — — 2,829 — — — — — — — — — — — — — 26,649 20,911 — — — — — 7,376 — — — — — 8,545 17,908 2,550 — — — — — 16,306 5,503 3,880 — — 11,879 — — 3,610 — — — — — — — — — — — — — — 48,327 — — — — — — — — — — — — 11,199 — — — — — — — — 6,336 — — — 22,482 — — 4,604 — — — — — — — — — — — — — — 10,244 24,235 — — — — Tons — — — — — — — 11,092 10,754 6,721 — — — — — — 12,879 — — — 23,941 — — 6,134 — — — — — — — — — — — — — — 20,694 — — — — — — — — — — — — 8,198 — — — — — — — — 14,867 13,687 — — 12,353 — — — 5,250 — — — — — — — — — — — — — 14,750 — — — — — — — — — — — — — — — — — — — — — — — — — 14,468 — — — 9,189 — — — — — — — — — — — — — 44,241 — — — — — — — — — — — — — — — — — 21,993 — — — — — — — 21,227 — — — — — — — — — — — — — — — — — 33,674 30,397 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 16,777 — 83,637 — — — — — — — — — — — — 14,338 — — — — — — — — — 18,191 — — 29,374 — — — — — — — — — — — — — — — — — 31,319 — — — — — — — — — — — — — — — — — — — — 47,849 110,695 83,706 — — 28,497 — — — — — — — — — — — — — — — — — 191,256 — — — — — 12,001 17,371 60,388 61,880 4,176 19,187 1,488 71,633 80,648 21,011 8,287 3,425 27,389 6,466 6,344 82,113 171,095 218,158 981 470 182,028 2,033 10,908 33,317 20,713 2,270 3,380 1,304 2,033 1,611 1,053 2,330 4,351 14,334 5,304 7,177 16,777 3,791 544,308 129,927 1,449 132,097 1,281 2,572 Table 7—Estimated aboveground dry stem weight of all live trees on forest land by species and diameter class (continued) Diameter class (inches) Species Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni Symplocos racemosa Terminalia catappa Theobroma cacao Timonius subauritus Tournefortia argentea Trichospermum ledermannii Vitex cofassus Xylocarpus granatum Unknown 0 Unknown 1 Unknown 3 Unknown 5 Unknown 11 Unknown, other Total 5.0–6.9 7.0–8.9 9.0– 10.9 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 8,645 — 2,040 — 5,027 49,359 13,032 4,610 — 18,453 376 — 2,358 3,505 2,052 — 483 2,693 354 2,070 — 2,848 1,925 562 712 1,082 — 815 16,851 1,384 1,903 — — 40,663 12,115 11,597 715 13,616 2,715 1,128 — 1,390 1,536 3,119 3,002 2,991 717 4,603 1,454 1,819 — 771 — 3,762 — — 13,810 — — — — 21,100 2,926 9,376 — 15,248 — — 2,036 3,189 — 6,420 — 2,101 1,867 6,195 — 5,882 — — — 3,358 2,015 2,241 11,388 — — — — 9,779 8,114 18,797 — 14,242 2,387 — — — — 5,635 6,385 — — 4,189 — 2,854 — — 2,940 — — — — — — — — 14,100 — 23,342 — 4,985 — — — — — — — — 2,294 — 5,099 8,861 — — — — — — — — — 3,814 — — — 7,921 — 8,409 8,605 10,113 — — — — — 7,004 — — 11,408 — — — — — — — 630,759 486,266 445,152 508,209 347,497 335,342 Tons 9,509 6,273 — — — — — — — — — — — — 8,960 — — — — — — — 12,914 27,216 — — — — — — — — — — — — — — — — 6,961 — 3,826 — — — — — — — — — — — — — 250,139 19.0– 20.9 188,788 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes — — — 8,112 — — — — — — — — — — — — — — — — 20,058 — — — — — — — — — — — — — — — — — — — — — — — — — — — 16,371 — — — — — — — — — — — — — — — — 21,838 — — — — — — — — — — 14,771 — — — — — — — — — — — — — — — — — 36,276 — — — — — — — — — — — — — — — — — — — — — — — — — — 21,805 133,415 — — — — — — — — — — — — — — — — 23,008 66,476 1,384 3,942 11,925 5,027 135,002 36,187 84,603 715 118,597 183,774 51,371 4,395 8,083 3,588 15,174 9,870 14,789 5,232 17,056 76,122 26,090 1,925 1,333 3,653 8,202 2,015 26,063 193,672 217,167 251,921 129,498 702,384 4,686,793 29 30 Table 8—Estimated carbon mass of all live trees on forest land by species and diameter class Diameter class (inches) Species 5.0–6.9 7.0–8.9 9.0– 10.9 Aglaia palauensis Allophylus ternatus Allophylus timorensis Alphitonia carolinensis Annona reticulata Areca catechu Artocarpus mariannensis Astronidium palauense Avicennia marina Badusa palauensis Barringtonia racemosa Bruguiera gymnorrhiza Buchanania palawensis Calophyllum inophyllum Calophyllum pelewense Calophyllum soulattri Calophyllum sp. Campnosperma brevipetiolata Canarium hirsutum Casearia cauliflora Casearia sp. Casuarina equisetifolia Cerbera floribunda Cerbera manghas Cocos nucifera Commersonia bartramia Cordia micronesica Cyathea lunulata Diospyros ferrea Diospyros sp. Dodonaea viscosa Dracaena multiflora Drypetes nitida Dysoxylum sp. Elaeocarpus graeffei Elaeocarpus joga Erythrina fusca Eugenia cumini Eugenia javanica Eugenia reinwardtiana Eugenia sp. Fagraea ksid Ficus sp. Ficus tinctoria 2,804 — 229 5,759 491 1,653 — 4,518 — 3,547 3,539 4,105 1,908 1,679 1,604 494 — 3,217 730 — 2,917 439 398 2,046 — 234 — 5,349 1,406 2,060 271 1,598 369 507 505 5,187 — 3,898 1,302 10,210 1,323 1,850 — — 1,700 1,614 — 8,840 — 634 — 1,630 — 1,657 4,126 8,131 — 4,466 3,899 623 1,679 5,877 750 1,235 1,147 — — 2,136 — — — 5,543 2,806 3,042 526 904 585 — — 10,959 — 3,440 3,760 11,129 — — — 344 3,514 — — 3,657 — — — 2,851 895 6,393 5,571 6,823 1,101 3,502 — — 1,962 7,622 895 — 2,018 — — 5,629 10,683 — — 2,395 — — 1,282 — — — — 748 — 1,631 6,891 14,504 — 5,709 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 19.0– 20.9 — — — 9,940 — — — — — 2,971 8,248 3,113 — 6,286 2,644 2,646 — 19,299 1,226 — — — 1,125 4,143 38,540 — — 2,173 — 2,756 — 1,152 — — — 12,708 — 3,874 1,938 9,466 — 5,502 — — — — — — — — 3,447 — — 4,529 1,797 6,674 — 1,738 — — — 19,507 — — — — — 2,098 11,595 — 2,591 — — — — — 1,966 — — 10,774 — 3,036 3,025 3,888 — 3,172 — — 5,943 — — 6,534 — — — — — 4,762 — — — 6,682 — — — 25,654 — — — — — 2,308 — — — — — — 6,627 — — — — 12,275 — 4,071 7,538 5,704 — 9,398 — — Tons — — — — — — — — — — — — — 4,971 — — — — — — — — — — — — — 15,705 — — — — 3,152 — 31,704 15,169 3,118 — — — — — — — — — — — 5,066 — — — — — — — — — — — — — — — — — — — — — — — 2,348 — — — 5,061 4,337 6,271 — — — — — — 2,053 — — 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes — — — — — — — — — — — — 7,237 — — — 12,651 23,787 — — — — — — — — — — — — 4,150 — — — — — — — — — — — — — — — — — — — 10,010 — — — — — — 8,455 — — — 27,352 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 16,705 — 10,242 — — — — — — — — — — — — 18,233 — — — — — — — — — — — — — — — — 11,119 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 14,543 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 15,912 13,961 1,614 229 34,731 491 2,287 35,134 8,999 11,136 23,859 23,281 43,388 10,247 48,513 8,146 3,764 19,444 179,189 6,719 1,235 6,083 18,672 1,523 18,361 65,883 234 2,591 15,460 4,211 7,858 12,856 3,654 2,920 507 505 52,652 2,348 19,949 44,969 61,170 1,323 25,630 2,053 16,257 Table 8—Estimated carbon mass of all live trees on forest land by species and diameter class (continued) Diameter class (inches) 31 Species 5.0–6.9 7.0–8.9 9.0– 10.9 Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo-guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata Morinda pedunculata Morinda sp. Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus tectorius Pandanus utiyamai Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pouteria calcarea 3,129 2,131 6,431 11,011 698 2,320 729 1,708 6,738 — 4,061 221 570 1,747 1,403 4,733 2,374 14,056 481 230 577 — 2,574 5,895 751 — 1,656 639 996 — 516 1,142 1,066 5,300 — 2,044 — — 8,202 11,247 710 61,310 — 1,260 1,608 944 8,627 5,397 1,348 4,177 — 1,249 6,957 2,160 — 425 — 1,422 1,706 1,949 1,995 12,128 — — 4,015 — 1,456 1,623 — 1,112 — — — 789 — — 403 1,723 2,599 1,473 — 576 6,025 5,553 — 2,374 627 — — 1,823 5,953 7,113 — 1,096 — 2,846 6,042 1,493 — — 945 — — — 5,830 15,362 — — 2,446 996 1,315 1,776 937 — — — — — — — 662 — — — — 1,282 5,136 9,849 — 1,043 — — 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 1,143 — 8,579 6,801 — 1,809 — 3,145 5,735 2,101 — 1,032 1,129 — — 2,117 — 6,815 — — 1,688 — — — 1,386 — — — — — — — — — — — — — 13,058 10,246 — — — — — 3,614 — — — — — 4,187 8,775 1,249 — — — — — 7,990 2,697 1,901 — — 5,821 — — 1,769 — — — — — — — — — — — — — — 23,680 — — — — — — — — — — — — 5,488 — — — — — — — — 3,105 — — — 11,016 — — 2,256 — — — — — — — — — — — — — — 5,019 11,875 — — — — Tons — — — — — — — 5,435 5,269 3,293 — — — — — — 6,311 — — — 11,731 — — 3,006 — — — — — — — — — — — — — — 10,140 — — — — — 19.0– 20.9 — — — — — — — 4,017 — — — — — — — — 7,285 6,707 — — 6,053 — — — 2,572 — — — — — — — — — — — — — 7,227 — — — — — 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes — — — — — — — — — — — — — — — — — — — — 7,089 — — — 4,503 — — — — — — — — — — — — — 21,678 — — — — — — — — — — — — — — — — — 10,777 — — — — — — — 10,401 — — — — — — — — — — — — — — — — — 16,500 14,894 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 8,221 — 40,982 — — — — — — — — — — — — 7,025 — — — — — — — — — 8,914 — — 14,393 — — — — — — — — — — — — — — — — — 15,346 — — — — — — — — — — — — — — — — — — — — 23,446 54,241 41,016 — — 13,964 — — — — — — — — — — — — — — — — — 93,716 — — — — — 5,881 8,512 29,590 30,321 2,046 9,402 729 35,100 39,518 10,296 4,061 1,678 13,421 3,168 3,109 40,236 83,836 106,898 481 230 89,194 996 5,345 16,325 10,149 1,112 1,656 639 996 789 516 1,142 2,132 7,023 2,599 3,517 8,221 1,858 266,711 63,664 710 64,728 627 1,260 32 Table 8—Estimated carbon mass of all live trees on forest land by species and diameter class (continued) Diameter class (inches) Species 5.0–6.9 7.0–8.9 9.0– 10.9 Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni Symplocos racemosa Terminalia catappa Theobroma cacao Timonius subauritus Tournefortia argentea Trichospermum ledermannii Vitex cofassus Xylocarpus granatum Unknown 0 Unknown 1 Unknown 3 Unknown 5 Unknown 11 Unknown, other 4,236 — 999 — 2,463 24,186 6,386 2,259 — 9,042 184 — 1,156 1,717 1,006 — 237 1,319 173 1,014 — 1,395 943 275 — 349 530 399 8,257 678 932 — — 19,925 5,937 5,683 350 6,672 1,331 553 — 681 752 1,529 1,471 1,466 351 2,255 713 891 — 378 — — 1,843 — 6,767 — — — — 10,339 1,434 4,594 — 7,472 — — 998 1,563 — 3,146 — 1,030 915 3,035 — 2,882 — — 988 — 1,646 1,098 5,580 — — — — 4,792 3,976 9,211 — 6,979 1,169 — — — — 2,761 3,129 — — 2,053 — 1,398 — — — 1,441 — — — — — — — 6,909 — 11,437 — 2,443 — — — — — — — — 1,124 — 2,499 4,342 — — — — — — — — — 1,869 — — — 3,881 — 4,120 4,216 4,955 — — — — — 3,432 — — 5,590 — — — — — — — 309,072 238,270 218,124 249,022 170,274 164,318 Total 11.0– 12.9 13.0– 14.9 15.0– 16.9 17.0– 18.9 19.0– 20.9 Tons 4,659 3,074 — — — — — — — — — — — — 4,390 — — — — — — — 6,328 13,336 — — — — — — — — — — — — — — — — 3,411 — 1,875 — — — — — — — — — — — — — 122,568 92,506 21.0– 22.9 23.0– 24.9 25.0– 26.9 27.0– 28.9 29.0+ All classes — — — 3,975 — — — — — — — — — — — — — — — — 9,828 — — — — — — — — — — — — — — — — — — — — — — — — — — — 8,022 — — — — — — — — — — — — — — — — 10,701 — — — — — — — — — — 7,238 — — — — — — — — — — — — — — — — — 17,775 — — — — — — — — — — — — — — — — — — — — — — — — — — 10,684 65,373 — — — — — — — — — — — — — — — — 11,274 32,573 678 1,932 5,843 2,463 66,151 17,732 41,455 350 58,113 90,049 25,172 2,153 3,961 1,758 7,435 4,836 7,247 2,564 8,358 37,300 12,784 943 653 988 1,790 4,019 12,771 94,899 106,412 123,441 63,454 344,168 2,296,529 Palau’s Forest Resources, 2003 Number of Canopy and Understory Species In addition to counting and measuring overstory trees, understory vegetation cover and layer heights were estimated for shrubs, forbs, vines, and grasses on FIA subplots where a species occupied at least 3 percent cover on that subplot (table 9). Tree seedlings that are less than or equal to 1 inch in diameter are also estimated as understory vegetation cover (table 10). Special interest species (e.g., rare, endangered, medicinal, or invasive) identified by island foresters were also noted when found. However, if a species covered less than 3 percent of a subplot and it was not listed as special interest, it was not enumerated. Table 9—Average nontree understory cover a on FIA field subplots by species and forest type Forest type Volcanic Scientific name Cover Achyranthes aspera L. Adiantum philippense L. Alpinia carolinensis Koidz. Alpinia pubiflora K.Schum. Alpinia purpurata (Vieill.) K. Schum. Alpinia sp. Roxb. Angiopteris evecta (J.R. Forst.) Hoffman Antrophyum plantagineum (Cav.) Kaulf. Asplenium nidus L. Asplenium pellucidum Lam. Asplenium polyodon G. Forst. Bambusa vulgaris Schrad. ex J.C. Wendl. Blechnum orientale L. Bolbitis sp. Schott. Caesalpinia major (Medik.) Dandy & Exell Caesalpinia sp. L. Calamus sp. L. Cassytha filiformis L. Centotheca lappacea (L.) Desv. Cheilanthes tenuifolia (Burmann) Swartz Clerodendrum sp. L. Clidemia hirta (L.) D. Don Clidemia sp. D. Don Combined ferns Costus speciosus Smith Cyperus sp. L. Cyrtosperma chamissonis (Schott) Merrill Dalbergia palauensis Hosokawa Davallia solida (Forster fil.) Swartz. Schrad Davallia sp. Sm. Decaspermum raymundii Diels Decaspermum sp. J. R. Forst. & G. Forst. Derris sp. Lour. Derris trifoliata Lour. Percent 2.0 4.8 9.6 5.0 3.3 3.2 3.7 — 1.6 — 1.0 2.0 4.6 3.3 1.0 2.0 1.0 6.0 1.3 1.0 1.0 1.3 1.0 1.0 2.2 50.0 8.3 3.0 1.7 3.0 2.0 2.0 35.0 19.0 Number of subplots 1 4 14 3 3 13 3 — 65 — 1 1 16 3 1 1 1 3 3 1 2 4 1 2 5 1 3 1 3 5 1 2 4 5 Limestone Standard deviation Cover — 4.1 17.6 4.4 3.2 3.0 2.5 — 1.02 — — — 6.1 4.0 — — — 7.8 0.6 — 0.0 0.5 — 0.0 1.1 — 10.1 — 1.2 1.9 — 1.4 12.9 13.4 Percent — — — — — — — 1.0 1.4 2.0 2.1 — — — — — 5.0 — — — — — — — — — — — 1.5 — — — — — Number of subplots — — — — — — — 2 23 1 10 — — — — — 2 — — — — — — — — — — — 4 — — — — — Standard deviation — — — — — — — 0.0 0.7 — 2.1 — — — — — 4.2 — — — — — — — — — — — 0.6 — — — — — 33 RESOURCE BULLETIN PNW-RB-252 Table 9—Average nontree understory cover a on FIA field subplots by species and forest type (continued) Forest type Volcanic Scientific name Cover Dianella sp. Lam. Dioscorea alata L. Dioscorea bulbifera L. Dioscorea sp. L. Donax cannaeformis (Forst. f.) K. Schum. Donax sp. Lour. Eriachne pallescens R. Br. Eurya japonica (Korth.) This.-Dyer Eustachys petraea (Sw.) Desv. Ficus sp. L. Ficus tinctoria Forst. f. Flagellaria gigantea Hook f. Freycinetia villalobosii Martelli Gleichenia linearis (Luers.) Fosberg Gliricidia sp. (Blank) Hanguana malayana Merrill Hedyotis cornifolia Kaneh. Hedyotis korrorensis (Valeton) Hosok. Hedyotis sp. L. Heterogonium pinnatum (Copel.) Holtt. Hippobroma longiflora (L.) G. Don Hyptis capitata Jacq. Ipomea sp. L. Ischaemum polystachyum J. S. Presl Ixora casei Hance Ixora sp. L. Lantana camara L. Lindsaea obtusa J. Sm. Hook. Lycopodium cernuum L. Lycopodium phlegmaria L. Lygodium auriculatum (Willd.) Alston Lygodium circinnatum (Burm. f.) Sw. Machaerina mariscoides (Gaud.) J. Kern Macrothelypteris torresiana (Gaud.) Medinilla blumeana Mansfeld Melastoma malabathricum L. Merremia peltata (L.) Merrill Merremia sp. Dennst. Ex Endl. Microsorum scolopendria (Burm. f.) Mimosa pudica L. Mussaenda frondosa L. Nepenthes mirabilis Druce Nephrolepis acutifolia (Desv.) J. Christ Nephrolepis biserrata (Sw.) Schott Nephrolepis hirsutula (J.R. Forst.) K. Presl Pandanus aimiriikensis Martelli Pandanus sp. L. f. Paspalum conjugatum Berg. Pennisetum polystachyon (L.) J.A. Schultes Phaleria nisidai Kaheh. Phragmites karka (Retz.) Trin. ex Steud. Percent 1.7 5.0 1.5 3.0 11.0 24.0 45.0 7.8 4.7 1.5 6.5 1.0 23.8 47.1 15.0 19.0 2.0 1.0 3.6 2.0 1.0 1.5 — 17.7 4.7 2.8 2.5 1.1 10.9 1.0 1.7 6.6 5.3 2.5 1.0 1.5 25.3 1.5 2.2 2.0 2.0 1.2 2.0 2.0 9.9 10.7 4.6 11.4 1.0 2.0 25.3 34 Number of subplots 3 4 2 1 9 3 4 10 3 4 2 1 6 20 1 4 7 1 5 1 1 2 — 3 3 37 2 14 11 1 9 5 19 2 2 16 19 2 5 1 1 25 1 4 7 9 68 5 1 1 3 Limestone Standard deviation Cover 1.2 1.6 0.7 — 11.9 23.3 5.8 9.9 0.6 1.0 2.1 — 29.3 38.3 — 14.5 1.2 — 3.7 — — 0.7 — 19.7 2.9 3.1 0.7 0.3 9.0 — 0.9 3.2 7.3 0.7 0.0 0.9 28.8 0.7 1.6 — — 0.4 — 0.8 15.8 24.2 5.9 8.3 — — 22.4 Percent — — — — — — — — — — — — — — — — — — — — — — 1.0 — — 5.0 — 2.0 — — — — — — — — 5.0 — 2.7 — — — — 24.7 — — 2.0 — — — — Number of subplots — — — — — — — — — — — — — — — — — — — — — — 1 — — 1 — 1 — — — — — — — — 1 — 19 — — — — 13 — — 1 — — — — Standard deviation — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 3.3 — — — — 30.2 — — — — — — — Palau’s Forest Resources, 2003 Table 9—Average nontree understory cover a on FIA field subplots by species and forest type (continued) Forest type Volcanic Scientific name Phyllanthus sp. L. Piper fragile Benth. Piper sp. L. Pityrogramma calomelanos (L.) Link Polypodium punctatum (Raddi) Hook. Polyscias grandifolia Volk. Portulaca oleracea L. Pteris ensiformis Burm. f. Pteris plumula Desv auct. Non Retz. Pteris tripartita Sw. Pteris vittata L. Rhaphidophora carolinensis (Volk.) Fosberg Rhaphidophora sp. Hassk. Rhaphidophora versteegii Engl. & K. Krause Rhynchospora sp. Vahl Scaevola sericea Vahl var. taccada (Gaertn.) Thieret & B. Lipscomb Schefflera odorata Merr. & Rolfe Schefflera sp. J.R. & G. Forst. Schizaea dichotoma (L.) Sm. Scleria levis Willd. Spathoglottis carolinensis Schlecht. Thelypteris forsteri Morton Thelypteris parasitica (L.) Reed. Thelypteris sp. Schmidel Thelypteris unita (L.) C.V. Morton Vernonia sp. Schreb. Vitex sp. L. Vitex trifolia L. Wikstroemia elliptica Merr. Unknown annual grass 1 Unknown fern 0 Unknown fern 1 Unknown fern 2 Unknown fern 3 Unknown fern 4 Unknown forb 1 Unknown forb 2 Unknown grass 1 Unknown orchid 0 Unknown perennial grass 1 Unknown perennial grass 1a Unknown perennial grass 2 Unknown shrub 0 Unknown shrub 2 Unknown vine 1 Unknown vine 2 Unknown vine 3 a Cover Number of subplots Limestone Standard deviation Cover 0.6 — — 0.0 0.0 19.4 13.2 — — 0.7 0.0 1.4 1.0 2.7 — Percent — — 5.5 — — — — — — — — — — 2.5 — Percent 1.3 1.0 1.0 1.0 1.0 22.5 25.0 2.0 2.0 1.6 1.0 2.0 1.5 2.5 25.0 3 1 1 2 2 4 3 1 1 8 2 2 4 23 1 5.0 1 — 17.3 1.0 1.2 3.0 1.7 6.0 2.0 3.0 6.2 2.0 — 25.0 2.4 4.3 1.0 12.8 8.0 1.0 1.0 3.6 3.3 1.0 2.8 13.6 1.0 45.5 3.1 5.0 2.8 3.0 25.0 3 2 20 8 3 1 3 1 24 3 — 1 5 4 1 6 4 1 2 11 3 1 5 13 1 2 7 1 9 1 1 14.2 0.0 0.5 2.1 0.6 — 1.0 — 14.2 1.0 — — 1.7 4.3 — 28.0 9.1 — 0.0 2.7 1.5 — 4.0 18.8 — 62.9 5.2 — 4.6 — — Number of subplots Standard deviation — — 6 — — — — — — — — — — 11 — — — 5.9 — — — — — 2.0 — — — — — — 1.0 — 5.6 — — — — 1.5 2.0 — — 3.0 2.0 — 1.0 — — — 1.6 — 2.0 1.0 — — 2 — — — — — — 1 — 7 — — — — 4 2 — — 1 2 — 1 — — — 11 — 5 1 — — 0.0 — — — — — — — — 10.8 — — — — 0.6 0.0 — — — 1.4 — — — — — 0.7 — 0.7 — — — — — — — — — — 1.3 — Percentage cover of “1” indicates cover less than or equal to 1 percent. Cover estimates are averaged among subplots where each species was found. A total of 188 subplots was surveyed for vegetation cover. The number of subplots where a species was found and the standard deviation for cover estimates provide an idea of spatial variability for each species across the island group. 35 RESOURCE BULLETIN PNW-RB-252 Table 10—Average understory tree cover a on FIA field subplots by species and forest type Forest type Volcanic Scientific name Cover Aglaia palauensis Kaneh. Aidia cochinchinensis Lour. Allophylus ternatus (Forst.) Radlk. Alphitonia carolinensis Hosok. Alphitonia zizyphoides A. Gray Areca catechu L. Astronidium palauense (Kaneh.) Mgf. Astronidium sp. A. Gray Badusa palauensis Valeton Barringtonia asiatica (L.) Kurz Barringtonia racemosa (L.) Spreng. Bruguiera gymnorrhiza (L.) Lam. Buchanania engleriana Volk. Buchanania palawensis Lauterb. Buchanania sp. Spreng. Calophyllum inophyllum L. Calophyllum pelewense P. F. Stevens Campnosperma brevipetiolata Volk. Canarium hirsutum Willd. Casuarina equisetifolia L. Cerbera manghas L. Cerbera sp. L. Cocos nucifera L. Commersonia bartramia (L.) Merr. Cyathea lunulata (Forst. f.) Copel. Diospyros discolor Willd. Diospyros ferrea (Willd.) Bakh. Diospyros sp. L. Dracaena multiflora Tentative Drypetes nitida Kaneh. Dysoxylum sp. Blume Elaeocarpus joga Merr. Elaeocarpus sp. L. Elattostachys falcata (Seem.) Radik. Eugenia cumini (L.) Druce Eugenia javanica Lam. Eugenia reinwardtiana (Bl.) DC. Eugenia sp. L. Evodia sp. Lam. Fagraea ksid Gilg and Bened. Ficus sp. L. Ficus tinctoria Forst. f. Flacourtia rukam Zoll. & Moritzi Garcinia matudai Kaneh. Garcinia rumiyo Fosb. Percent 2.4 — 2.0 3.3 2.0 3.0 1.4 1.0 2.0 — 5.9 1.6 1.0 3.0 5.5 1.5 2.3 2.8 7.0 — 2.8 1.0 3.8 1.0 4.3 2.5 2.5 2.5 — — 2.4 2.2 1.0 1.0 2.0 — 6.6 1.9 2.0 1.0 2.3 5.3 2.8 6.8 8.2 36 Number of subplots 21 — 1 6 1 3 11 1 1 — 7 15 1 1 2 2 4 9 2 — 4 1 9 1 4 2 2 2 — — 7 5 1 1 1 — 7 8 1 1 3 19 5 4 19 Limestone Standard deviation Cover 1.6 — — 2.7 — 2.0 0.7 — — — 6.5 1.4 — — 3.5 0.7 1.5 2.9 1.4 — 1.7 — 4.3 — 3.9 0.7 0.7 0.7 — — 1.0 1.3 — — — — 8.4 1.0 — — 0.6 4.4 1.5 9.0 12.5 Percent 2.3 1.5 — — — — — — 2.8 1.0 3.0 — — — — 2.2 — — — 2.0 — — 34.0 — — — — — 3.0 3.0 — 8.0 — — 4.8 1.8 4.3 3.3 — — — 2.4 — — — Number of subplots 4 2 — — — — — — 4 1 1 — — — — 5 — — — 1 — — 3 — — — — — 1 1 — 1 — — 4 8 9 3 — — — 7 — — — Standard deviation 0.5 0.7 — — — — — — 0.5 — — — — — — 0.8 — — — — — — 29.5 — — — — — — — — — — — 2.8 0.7 2.8 1.5 — — — 1.3 — — — Palau’s Forest Resources, 2003 Table 10—Average understory tree cover a on FIA field subplots by species and forest type (continued) Forest type Volcanic Scientific name Garcinia sp. L. Glochidion ramiflorum Forst. Glochidion sp. J.R. & G. Forst. Gmelina elliptica Sm. Gmelina palawensis H. J. Lam. Gulubia palauensis (Becc.) Moore & Fosb. Heterospathe elata (Becc.) Becc. Hibiscus tiliaceus L. Horsfieldia amklaal Kaneh. Horsfieldia novo-guineensis Warb. Horsfieldia palauensis Kaneh. Horsfieldia sp. Wild. Inocarpus fagifer (Park.) Fosb. Macaranga carolinensis Volk. Manilkara udoido Kaneh. Manilkara zapota (L.) P. v. Roy. Melochia aristata A. Gray Morinda citrifolia L. Morinda pedunculata Val. Myristica sp. Gronov. Neonauclea forsteri Merrill Omalanthus sp. A.Juss. Ormosia calavensis Azaola ex Blanco Osmoxylon oliveri Fosberg & Sachet Osmoxylon pachyphyllum (Kanehira) F.R. Fosberg & M.-H. Sachet Osmoxylon sp. Miq. Pandanus aimiriikensis Mart. Pandanus dubius Spr. Pandanus kanehirae Mart. Pandanus peliliuensis Kaneh. Pandanus tectorius Park. Parinari corymbosum (Bl.) Miq. Parinari laurina Gray Parkia parvifoliola Hosok. Pinanga insignis Becc. Pithecellobium dulce (Roxb.) Benth. Polyscias nodosa (Bl.) Seem. Pongamia pinnata (L.) Merr. Pouteria calcarea (Hosok.) Fosb. Pouteria obovata (R. Br.) Baehni Premna obtusifolia R. Br. Rhizophora apiculata Bl. Rhizophora mucronata Lam. Rhus taitensis Guill. Cover Number of subplots Limestone Standard deviation Cover Percent 7.8 2.0 1.0 2.0 — 2.0 2.0 — 2.5 — 2.2 — — 5.5 — — — — — — — — — — Percent 4.3 3.0 — — 1.5 9.3 4.2 5.0 3.0 4.5 4.3 2.5 2.0 3.1 2.0 1.7 2.0 1.5 21.5 2.0 4.0 11.5 2.0 5.0 4 4 — — 2 3 6 8 2 15 45 2 2 17 5 3 1 4 2 1 1 4 1 4 3.9 1.4 — — 0.7 5.1 2.0 5.1 — 3.1 10.6 0.7 1.4 3.0 1.4 0.6 — 0.6 26.2 — — 12.3 — 5.0 3.0 1 — 3.0 3.5 — 1.5 — 3.1 8.0 30.8 1.0 19.9 — — — — 1.5 5.0 13.9 1.5 1.9 10 10 — 2 — 8 4 12 1 103 — — — — 2 1 15 2 8 3.3 2.8 — 0.7 — 2.4 11.4 27.5 — 20.6 — — — — 0.7 — 23.9 0.7 1.4 Number of subplots Standard deviation 5 2 1 1 — 1 1 — 2 — 10 — — 2 — — — — — — — — — — 6.9 — — — — — — — 0.7 — 0.8 — — 3.5 — — — — — — — — — — — — — 1.0 27.0 2.5 — 1.5 — — 2.0 — 1.5 1.8 3.0 2.0 1.0 1.6 — 1.0 — 2.0 1 3 6 — 2 — — 1 — 4 4 1 1 1 7 — 4 — 1 — 37.5 1.6 — 0.7 — — — — 0.6 0.5 — — — 0.5 — — — — 37 RESOURCE BULLETIN PNW-RB-252 Table 10—Average understory tree cover a on FIA field subplots by species and forest type (continued) Forest type Volcanic Scientific name Cover Semecarpus venenosus Volk. Sonneratia alba J. E. Sm. Stemonurus ammui (Kaneh.) Sleumer Swietenia mahagoni (L.) Jacq. Symplocos racemosa Roxb. Tecoma stans (L.) Juss. ex HBK. Terminalia catappa L. Terminalia sp. L. Timonius subauritus Fosberg & Sachet Trichospermum ledermannii Burret Xylocarpus granatum Koen. Unknown 0 Unknown 1 Unknown 20 Percent 2.2 2.8 40.0 6.0 2.3 3.0 3.0 20.0 — 1.0 1.6 1.3 2.0 2.0 Number of subplots 31 8 1 2 6 1 1 1 — 2 8 4 1 1 Limestone Standard deviation Cover 1.6 3.1 — 2.8 0.8 — — — — — 1.4 0.5 — — Percent 3.0 — — — — — — — 8.5 — — — — — Number of subplots 1 — — — — — — — 2 — — — — — Standard deviation — — — — — — — — 9.2 — — — — — a Percentage cover of “1” indicates cover less than or equal to 1 percent. Cover estimates are averaged among subplots where each species was found. A total of 188 subplots was surveyed for vegetation cover. The number of subplots where a species was found and the standard deviation for cover estimates provide an idea of spatial variability for each species across the island group. For the 2003 inventory, 128 tree species and 132 understory species were measured on FIA plots. The volcanic forest area sampled was about four times the area sampled for the limestone forest. Because we expect the number of species counted to increase with increasing area, it is not surprising that over twice the number of tree species and over four times the number of understory species were found in volcanic forest compared to limestone forest (fig. 11). On a per-plot basis (approximately one-sixth acre), the mean number of tree species found on volcanic plots was 12, with an average of 10 tree species found per limestone plot (fig. 12). The one limestone plot where only two species occurred (fig. 12) was a mangrove plot where 90 trees (>5 inches) were measured, 65 Rhizophora apiculata and 25 R. mucronata. The area of mangrove forest could not be accurately delineated from limestone forest with this small sample, thus separate expansion factors were not calculated for mangrove species. Mangrove species numbers may be falsely high owing to this expansion. However, because mangrove species tend to dominate the extensive mangroves on Palau, and only five partial plots contained mangrove species, the estimate may be reasonable. Additional mapping data are forthcoming and will help refine our estimate for mangrove species. The urgent need to better understand mangroves as buffers to coastal disturbance (storms, typhoons, and tsunami) suggests the need for a separate, detailed mangrove inventory. 38 Palau’s Forest Resources, 2003 140 Number of species 120 Volcanic 100 Limestone 80 60 40 20 0 Tree species Understory species Number of plots Figure 11—One hundred and twenty-eight tree species and 132 understory species were measured on 54 forested plots in Palau. 10 Number of plots 9 8 Volcanic 7 Limestone 6 5 4 3 2 1 0 1–2 3–4 5–6 7–8 9–10 11–12 13–14 15–16 17–18 19–20 Number of tree species on FIA plots Figure 12—The average number of tree species found per sixth-acre plot was 12 for volcanic forests and 10 for limestone forests. 39 RESOURCE BULLETIN PNW-RB-252 Tree Damage and Mortality Approximately 13 percent of the individual trees on Palau show some sign of damage (table 11). The most prevalent damage types are vines in crowns, conks (shelf fungus indicating rot), and lost apical dominance (damage to the primary growing leader) (fig. 13). The prevalence of vines in tree crowns was high, 41 percent. Although the vines are common, their effect on their supporting tree hosts was not quantified. Effects ranged from some additional weight to rare instances of strangulation. For the identifiable damaging agents, damage by other vegetation, disease, and weather ranked as the most prevalent primary damage agents (fig. 14). Less than 5 percent of the trees sampled during the inventory were dead (fig. 15). Of those dead, nearly one-fourth could not be identified to species level. The most prevalent, identifiable dead trees were Cyathea lunulata (7.3 percent of total dead), Calophyllum inophyllum, Calophyllum pelewense, and Pinanga insignis (each accounting for 6.1 percent of total dead). Epiphytes— In the moist environment of the tropics many plant species are found growing on longer lived trees. These epiphytes use the trees primarily for support, but they also use space, moisture, and nutrients that might otherwise be used by the tree. When the epiphytes accumulate over many years, their weight can be excessive, leading to the breakage of branches and occasionally the stems of trees. In Palau we rated epiphytic loading on trees as a summation of loading on the bole, branches, and canopy. We estimate that about 65 percent of the trees in Palau had few to no epiphytes, and 45 percent of the trees had moderate to high amounts of epiphytes (table 12). Forest Dynamics— Forest structure and species composition change through time as a result of forest succession and disturbance dynamics. The forests of Palau have been strongly influenced by a long history of disturbance, both human-caused and climatic disturbance. Human populations and land use on Palau have changed, resulting in differing influences on forest vegetation. Within the last century, the most important land use changes have involved the conversion of forest to agriculture and its subsequent abandonment and reversion to grassland, and eventually forest. Endress and Chinea (2001) documented the abandonment of agricultural lands and reversion to forest by comparing 1947 and 1992 aerial photography for approximately 20,500 acres in the Ngeremeduu Bay drainage area on Babeldaob. These authors noted the most dramatic change occurred during the period 1947 to 1976, using the Cole et al. (1987) map as an intermediate comparison point. (Text continues on page 49.) 40 Table 11—Number of trees by primary damage type and species for all trees (≥5 inches diameter at breast height; includes dead trees) Species Broken Broken Broken Canker/ bole branches roots gall Conks Cracks/ Discolored Excessive Lost apical Open seams foliage branching dominance wound Number of trees — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 171,813 36,989 12,330 456,192 12,330 73,977 48,518 258,920 12,330 254,719 330,896 468,521 98,436 293,308 197,072 49,318 36,588 — — — — — — — — — — — — — — — — 24,659 — — — — — — — — — 12,129 — — — 12,129 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — 12,330 — — — 12,330 12,330 12,129 12,129 — — — 12,330 — — 714,911 — 24,659 12,330 — 24,659 — 72,777 12,330 84,906 60,647 24,659 135,425 306,637 12,330 12,330 542,498 98,636 73,577 73,177 98,036 36,788 36,989 — 453,391 — — — — — — — — — — — — — — — — — — — — — 12,129 — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,129 — — — 24,659 — — — — — — — — — — — — — — — — — — Vines in crown All damages — — — — — — — — — — — — — — — — — 12,330 — — — — — — 12,330 — 12,129 — — — — — — — — — — 12,330 — — — — — — — — — 24,259 — — — 49,318 — — 12,330 — — — — — 12,129 36,989 — — — — — 12,129 61,648 — — 36,989 — — — 24,659 12,330 48,518 49,118 12,330 — 36,388 12,330 — 36,788 — — — — 61,648 123,295 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — 12,330 12,330 — — — — 12,330 — — — — — — — — — — — 12,330 12,330 — — — — 61,648 12,330 — — — — — — — — — 12,330 24,459 24,659 — — 12,129 — 73,977 12,330 — — 12,330 — 12,330 — 12,129 — — 12,330 73,777 Palau’s Forest Resources, 2003 41 Aglaia palauensis Allophylus ternatus Allophylus timorensis Alphitonia carolinensis Annona reticulata Areca catechu Artocarpus mariannensis Astronidium palauense Avicennia marina Badusa palauensis Barringtonia racemosa Bruguiera gymnorrhiza Buchanania palawensis Calophyllum inophyllum Calophyllum pelewense Calophyllum soulattri Calophyllum sp. Campnosperma brevipetiolata Canarium hirsutum Casearia cauliflora Casearia sp. Casuarina equisetifolia Cerbera floribunda Cerbera manghas Cocos nucifera Commersonia bartramia Cordia micronesica Cyathea lunulata Diospyros ferrea Diospyros sp. Dodonaea viscosa Dracaena multiflora Drypetes nitida Dysoxylum sp. Elaeocarpus graeffei Elaeocarpus joga No damage (continued) Species No damage Erythrina fusca Eugenia cumini Eugenia javanica Eugenia reinwardtiana Eugenia sp. Fagraea ksid Ficus sp. Ficus tinctoria Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo-guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata 12,129 207,201 181,942 570,685 36,788 110,966 12,330 24,259 135,425 49,318 294,908 343,626 36,989 134,224 — 182,342 431,133 36,989 109,165 12,129 36,388 73,977 49,318 220,531 221,931 845,935 — 12,330 181,942 12,330 122,695 234,261 49,318 — 24,659 Broken Broken Broken Canker/ bole branches roots gall Conks — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — 12,129 — 12,129 — — — — — — — — — 24,459 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 24,259 — — 12,330 — — — — 12,330 — — 12,330 — 24,659 61,648 — — 12,129 12,129 — — 12,330 12,330 36,588 12,330 — 36,388 — 12,330 — — 12,330 12,330 Cracks/ Discolored Excessive Lost apical Open seams foliage branching dominance wound Number of trees — — — — — — — — — — — — — — — — 12,129 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 24,659 — — — — 12,330 — 36,989 — — — — — 12,330 12,330 — 24,659 — — — — — — 12,330 — — — — 24,259 24,259 — — — — — — — — — — — — 36,788 12,330 — — 12,129 — — — — 12,330 — — 12,129 — — — — — — Vines in crown All damages — 12,129 — — — 36,989 — — — — 12,330 — — 36,788 12,330 — — 24,659 — 12,129 — 12,330 36,989 — — 49,118 — — — — 36,989 36,989 12,330 12,330 24,659 — 12,129 48,518 24,259 — 61,648 — — 24,459 24,659 24,659 — — 61,247 24,659 36,788 135,425 36,989 — 24,259 24,259 12,330 49,318 24,659 12,330 147,154 12,330 — 48,518 — 49,318 49,318 24,659 24,659 36,989 RESOURCE BULLETIN PNW-RB-252 42 Table 11—Number of trees by primary damage type and species for all trees (≥5 inches diameter at breast height; includes dead trees) Table 11—Number of trees by primary damage type and species for all trees (≥5 inches diameter at breast height; includes dead trees) (continued) Species No damage Broken Broken Broken Canker/ bole branches roots gall Conks Cracks/ Discolored Excessive Lost apical Open seams foliage branching dominance wound Vines in crown All damages Number of trees Morinda sp. — — — — — — — — 12,330 — 24,659 24,659 61,648 145,554 61,648 86,307 12,330 12,330 554,428 516,039 24,659 2,071,357 12,129 36,989 353,555 12,330 36,989 24,659 86,307 1,343,843 377,214 357,556 12,129 528,168 172,613 61,648 49,318 36,989 36,989 12,330 24,659 84,906 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — 12,129 — 12,330 — — — — — — 12,330 — — — — — — — — 48,518 — 12,330 — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 49,318 24,659 — — — — 24,659 — — — — 12,129 — 12,330 — 12,330 — — — — — — 24,659 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — — — — — — — — — — — — — — — — — 12,330 — — — — — 12,129 — — — — 12,330 — — — — — — — — — — 12,129 — 12,330 — 12,330 — — — — 12,330 — 12,330 — 12,330 — — — — — — — — — — 12,330 — — — — — — — — — — 12,330 — 12,330 — — — — — — — — — — — — 12,330 — 36,989 — — — — — 12,330 12,330 24,659 — 123,295 — — 24,459 — 49,318 — — — — 12,330 — 24,659 — — — 36,989 — 36,989 — 24,259 24,659 — 12,330 — 36,989 — 60,647 — 12,330 — 12,330 110,966 49,318 — 123,295 — — 49,118 — 49,318 — — 48,718 — 73,977 — 49,318 — — — 36,989 24,659 36,989 36,989 24,259 Palau’s Forest Resources, 2003 43 Morinda pedunculata Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus tectorius Pandanus utiyamai Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pouteria calcarea Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni Symplocos racemosa Terminalia catappa Theobroma cacao Timonius subauritus 12,330 (continued) Species Tournefortia argentea Trichospermum ledermanni Vitex cofassus Xylocarpus granatum Unknown Unknown 0 Unknown 1 Unknown 3 Unknown 5 Unknown 10 Unknown 11 Unknown 30 Unknown, other Total No damage 36,588 98,636 61,648 147,954 12,330 134,824 24,659 49,318 85,906 12,330 12,330 12,129 73,977 18,714,675 Broken Broken Broken Canker/ bole branches roots gall Conks — — — — — — — — — — — — — — — — — — 12,330 — — — — — — — 24,659 195,872 — — — — — — — — — — — — — 97,836 — — — — — — — — — — — — — 12,330 — — 24,659 — — 24,459 — — — — — 12,330 12,330 687,652 Cracks/ Discolored Excessive Lost apical Open seams foliage branching dominance wound Number of trees — — — — — — — — — — — — — — — — — — — — — — — — — — 12,129 12,330 — — — — — — — — — — — — — — — — — — — — — — — — — — 12,330 295,308 — — 12,330 12,330 — — — — — — — — — Vines in crown All damages — — 36,989 — — — — — — — — — — 12,330 — 49,318 36,989 — 12,330 24,459 — — — — — 12,330 281,778 1,144,644 2,776,866 RESOURCE BULLETIN PNW-RB-252 44 Table 11—Number of trees by primary damage type and species for all trees (≥5 inches diameter at breast height; includes dead trees) Palau’s Forest Resources, 2003 <1% 4% 1% <1% <1% <1% Tree damages 7% Vines in crown Conks 10% Lost apical dominance 41% Open wound Broken branches Broken roots Broken bole 11% Canker/gall Discolored foliage Excessive branching Cracks/seams 25% Figure 13—Vines in the crowns of trees are very common in Palau. Damage from vines was recorded when the vines occupied 20 percent or more of the crown. Rot, indicated by the fruiting shelf fungi (conks) was also a common damage. 2% 3% <1% <1% 11% Primary damage agent Vegetation Disease Unknown 11% Weather 53% Physical Insect Fire Human 19% Figure 14—Disturbance by other vegetation and disease were the most common identifiable damaging agents for trees. 45 RESOURCE BULLETIN PNW-RB-252 25 Dead trees (percent) 20 15 10 5 Ca Cy All u lop ath nk n ow Ca hyll ea l o p u m lu n n u hy i llu nop lata hy m Pi p e l e l l u m na we n Ca So ga in nse su n s a r n er i g n i na is Al ati ph aa ito equ l b n is e a P ia c ti f B a a r i na a r o l o l i a i rri n r i c n en Ho gton or ym sis ia r r a bos As sfiel tr o d i a c em a nid o p ium alau sa pa ens El ae l i o c uae s ns ar is R pu Rh izo hus s jog Ca mp Ti pho taite a no mo ra a nsi s p n iu p ic s er m a s s u u la t br bau a Di evip ritus os e py tiol ata Ma r o s c a G ar c i n fer r ra e n i Pa ga c a ru a a r m iy nd oli an n o Pr us k ens em an i eh s B a na du o b i r a e sa tus i p Eu C al fo l i ge alop auen a n ia hy sis G u r e i n l lu m Rh lub war sp. izo ia p dti a ph a or lau na a m en u c s is ro na ta 0 Species Figure 15—Identifying dead trees proved to be difficult in Palau. Nearly one-fourth of those dead could not be identified. Table 12— Estimated number of trees on forest land by epiphyte loading (amounts of nontree vegetation in the canopy and branches) and species (includes dead trees) Epiphyte loadings 46 Species None Aglaia palauensis Allophylus ternatus Allophylus timorensis Alphitonia carolinensis Annona reticulata Areca catechu Artocarpus mariannensis Astronidium palauense Avicennia marina Badusa palauensis Barringtonia racemosa Bruguiera gymnorrhiza Buchanania palawensis Calophyllum inophyllum 36,388 — — 135,625 — — 24,259 12,330 — 97,036 97,436 184,943 — 60,647 Low 123,095 12,330 — 160,284 — 36,989 — 135,625 12,330 145,554 134,624 234,261 49,118 146,154 Moderate Number of trees 61,648 12,330 12,330 110,966 12,330 12,330 24,259 73,977 12,330 36,388 110,966 36,989 24,659 73,577 High 12,330 12,330 — 86,307 — 24,659 — 61,648 — 24,259 36,989 24,659 24,659 49,318 All loadings 197,072 36,989 12,330 357,556 12,330 73,977 24,259 271,249 24,659 206,201 282,578 295,908 98,436 269,049 Palau’s Forest Resources, 2003 Table 12— Estimated number of trees on forest land by epiphyte loading (amounts of nontree vegetation in the canopy and branches) and species (includes dead trees) (continued) Epiphyte loadings Species None Calophyllum pelewense Calophyllum soulattri Calophyllum sp. Campnosperma brevipetiolata Canarium hirsutum Casearia cauliflora Casearia sp. Casuarina equisetifolia Cerbera floribunda Cerbera manghas Cocos nucifera Commersonia bartramia Cordia micronesica Cyathea lunulata Diospyros ferrea Diospyros sp. Dodonaea viscosa Dracaena multiflora Drypetes nitida Dysoxylum sp. Elaeocarpus graeffei Elaeocarpus joga Erythrina fusca Eugenia cumini Eugenia javanica Eugenia reinwardtiana Eugenia sp. Fagraea ksid Ficus sp. Ficus tinctoria Flacourtia rukam Garcinia matudai Garcinia rumiyo Garcinia sp. Gironniera celtidifolia Glochidion ramiflorum Glochidion sp. Gmelina elliptica Gmelina palawensis Gmelina sp. Gulubia palauensis Hernandia sonora Hernandia sp. Heterospathe elata 24,659 36,989 — 61,447 36,588 — — 12,129 — 12,330 12,330 — — 49,318 — 36,588 12,129 73,377 24,659 36,989 — 85,706 12,129 36,388 24,259 218,330 — 24,659 — — — 12,330 36,989 48,718 — 24,659 12,330 72,777 184,943 — 109,165 — — — Low 86,106 12,330 48,918 332,897 12,129 12,330 84,906 — 24,659 98,436 159,083 12,330 — 98,636 49,318 49,318 48,718 24,459 12,129 — — 269,649 — 97,836 121,295 340,025 — 24,659 — 12,129 85,906 36,989 85,706 233,260 24,659 85,306 — 72,777 221,331 36,989 — — 60,647 73,977 Moderate Number of trees 36,989 — 24,459 271,249 36,388 — — 60,647 — 61,648 110,565 — — 234,261 49,318 — — 12,330 — — 12,330 122,695 — 72,977 84,906 36,588 24,459 123,295 12,330 12,129 49,318 12,330 122,895 36,989 12,330 73,377 — 61,247 110,966 12,330 — 36,388 — 12,330 High 61,648 — — 172,613 12,330 — — — — 36,989 36,989 — 12,330 172,613 — — 12,330 — — — — 49,118 — 12,129 — — 12,330 — — — 24,659 12,330 73,977 24,659 — 12,129 12,330 12,330 49,318 24,659 — — — — All loadings 184,743 12,330 73,377 776,759 60,847 12,330 84,906 60,647 24,659 197,072 306,637 12,330 12,330 505,510 98,636 49,318 61,047 36,788 12,129 — 12,330 441,462 — 182,942 206,201 376,613 36,788 147,954 12,330 24,259 159,883 61,648 282,578 294,908 36,989 170,813 12,330 146,354 381,615 73,977 — 36,388 60,647 86,307 47 RESOURCE BULLETIN PNW-RB-252 Table 12— Estimated number of trees on forest land by epiphyte loading (amounts of nontree vegetation in the canopy and branches) and species (includes dead trees) (continued) Epiphyte loadings Species Hibiscus tiliaceus Horsfieldia amklaal Horsfieldia novo-guineensis Horsfieldia palauensis Horsfieldia sp. Inocarpus fagifer Intsia bijuga Lumnitzera littorea Macaranga carolinensis Manilkara udoido Manilkara zapota Morinda citrifolia Morinda latibracteata Morinda pedunculata Morinda sp. Nephelium lappaceum Osmoxylon oliveri Osmoxylon sp. Pandanus aimiriikensis Pandanus dubius Pandanus kanehirae Pandanus tectorius Pandanus utiyamai Pangium edule Parinari corymbosum Parinari laurina Parinari sp. Pinanga insignis Pithecellobium dulce Pouteria calcarea Pouteria obovata Pouteria sp. Premna obtusifolia Pterocarpus indicus Ptychosperma palauensis Rhizophora apiculata Rhizophora mucronata Rhus taitensis Rinorea carolinensis Semecarpus venenosus Sonneratia alba Stemonurus ammui Swietenia macrophylla Swietenia mahagoni 48 None Low — 12,330 — 182,742 — — 145,554 — 12,330 98,636 — — — — 24,659 — — — — 109,165 12,330 49,318 — — 73,577 49,118 12,330 110,966 12,129 — 60,647 — — — — 1,145,970 303,236 110,966 12,129 73,377 135,625 — 24,659 — 49,318 159,083 86,307 355,755 12,330 12,330 84,906 12,330 110,565 110,966 36,989 — — — 12,330 — 24,659 — 36,989 84,906 24,659 36,989 — — 197,272 269,649 12,330 1,269,939 — 24,659 134,824 12,330 12,330 — 24,659 246,590 24,659 234,261 — 294,708 36,989 12,330 24,659 36,989 Moderate Number of trees — 49,118 98,636 282,178 — — — — 36,788 36,989 24,659 24,659 36,989 — — — — 24,659 24,659 12,129 24,659 12,330 — 12,330 172,613 123,295 — 505,510 — 12,330 109,365 — 36,989 — 61,648 — 49,318 61,648 — 135,425 — — — 24,659 High All loadings 49,318 24,659 49,318 172,413 — — — — 12,330 36,989 12,330 — 24,659 12,330 — 12,330 — 36,989 — — — — 12,330 12,330 221,931 123,295 — 308,238 — — 97,836 — 36,989 24,659 — — — 24,659 — 73,977 — 49,318 — 12,330 98,636 232,860 234,261 810,347 12,330 12,330 84,906 12,330 159,683 184,943 73,977 24,659 61,648 12,330 12,330 12,330 24,659 61,648 61,648 97,036 49,318 49,318 12,330 24,659 591,816 516,239 12,330 2,083,686 — 36,989 342,025 12,330 86,307 24,659 86,307 246,590 73,977 320,567 — 504,109 36,989 61,648 24,659 73,977 Palau’s Forest Resources, 2003 Table 12— Estimated number of trees on forest land by epiphyte loading (amounts of nontree vegetation in the canopy and branches) and species (includes dead trees) (continued) Epiphyte loadings Species Symplocos racemosa Terminalia catappa Theobroma cacao Timonius subauritus Tournefortia argentea Trichospermum ledermannii Vitex cofassus Xylocarpus granatum Unknown Unknown 0 Unknown 1 Unknown 3 Unknown 5 Unknown 10 Unknown 11 Unknown 30 Unknown, other Total None Low 49,318 12,330 12,330 24,259 24,259 — — 110,966 12,330 73,177 24,459 24,659 24,259 — — — 36,989 12,330 12,330 12,330 36,388 12,330 73,977 24,659 73,977 — 36,989 — 12,330 61,648 — — 12,129 — 5,182,700 8,742,199 Moderate Number of trees — — 36,989 48,518 12,330 24,659 24,659 — — 12,330 12,330 12,330 — 12,330 12,330 — 36,989 4,794,504 High All loadings — 24,659 — — — — 61,648 — — 24,659 12,330 — — — — — 12,330 12,330 36,989 49,318 84,906 24,659 98,636 110,966 73,977 — 73,977 24,659 24,659 61,648 12,330 12,330 12,129 49,318 2,772,138 16,308,841 Comparing the inventory data from the present study to data from the MacLean et al. (1988) timber inventory suggests that the trend of forest recovery and maturation continues on Palau. With the exceptions of forest loss owing to roads and urban uses, forests appear to be maturing. On average, trees have been getting larger (fig. 16), and as they increased in size, some trees died during the “thinning phase” (Oliver and Larson 1996) of forest succession. We found fewer stems per acre, and greater basal area and volume per acre on forested field plots compared to the MacLean et al. (1988) data (table 13). The MacLean et al. (1988) work was restricted to forested lands on Babeldaob, whereas the current inventory included additional forest types in limestone forest and the Rock Islands. Although our current sample of plots in limestone forest was relatively small (11 out of 54 plots total), the 2003 data suggest much higher net wood volume in limestone (4,107 cubic feet per acre) than in volcanic forests (2,424 cubic feet per acre). Much of the limestone forest is in difficult terrain where access and human populations are limited. The limestone forests were probably less impacted by agricultural development and thus are likely to be in a later phase of maturity than their volcanic counterparts. Soil fertility may also play a role in wood volume differences owing to less aluminum toxicity in limestone versus the volcanic-derived soils. 49 RESOURCE BULLETIN PNW-RB-252 70 Percentage of all trees by diameter class All trees (percent) 60 50 40 30 20 1985 2003 10 >2 9 7 27 –2 9 –2 5 25 –2 23 23 21 – 15 –1 7 17 –1 9 19 –2 1 .9 9. –1 0. 9 11 .0 –1 2. 9 13 –1 5 –8 7. 0 5. 0– 6. 9 0 Diameter (inches) Figure 16—The diameter distribution for the percentage of total trees measured on field plots appears to have changed between the 1985 and 2003 inventories. Trees measured in the latest inventory were, on average, larger than those measured in 1985, contributing to higher basal area and volume per acre values. Table 13—Estimated number of trees per acre, basal area per acre, net volume per acre, and standard errors (SE) for 1985 and 2003 Trees per acre 2 Basal area (ft per acre) Volume (ft3 per acre) 50 1985 SE 2003 SE 1,282 136 102 — 856 146 68 — 1,740 188 2,744 296 Palau’s Forest Resources, 2003 Acknowledgments The FIA Pacific Island inventories are the result of cooperative effort between a number of groups and agencies. The USDA Forest Service, State and Private Forestry in the Pacific Southwest Region has been a strong supporter of natural resource assessments in the Pacific Islands. Many thanks to the Pacific Islands Imagery Consortium partners for sharing costs on imagery and interpretation. Thanks to Margie Falanruw with the Institute of Pacific Islands Forestry, Lynn Raulerson at University of Guam, and Agnes Rinehart for species identification and training; Lisa Fischer and Zhafeng Liu (Leo) at the Remote Sensing Lab in the Pacific Southwest Region and Julie Anderson with PNW-FIA for imagery interpretation and mapping support; Pua Michael, Larry Mamis, Pamela Skilang, Dale Baer, Falaniko Mika, Ritofu Lotovale for field assistance and logistical support. We also thank Dale Weyermann, John Chase, Chuck Veneklase, and Ron Wanek with FIA for GIS, programming, and technical support. Tom Cole, Robin DeMeo, Bob Gavenda, and Tarita Holm offered very helpful comments on prior drafts. Metric Equivalents When you know: Inches Feet Miles Acres Cubic feet Tons Tons per acre Cubic feet per acre Trees per acre Degrees Fahrenheit Multiply by: 2.54 .305 1.609 .405 .028 907 2.24 .07 2.471 (F - 32)/1.8 To find: Centimeters Meters Kilometers Hectares Cubic meters Kilograms Tonnes or megagrams per hectare Cubic meters per hectare Trees per hectare Degrees Celsius Literature Cited Athens, J.S.; Ward, J.V. 2002. Holocene paleoenvironmental investigations on Ngerekebesang, Koror, south Babeldaob, and Peleliu islands. Honolulu, HI: International Archaeological Research Institute, Inc. Climate Prediction Center. 2005. Composite rainfall of ENSO (El Niño/Southern Oscillation) for Pacific Island stations. http://www.cpc.ncep.noaa.gov/pacdir/ ENSO11.shtml. (April 1). Cochran, W.G. 1977. Sampling techniques. New York: John Wiley & Sons. 428 p. 51 RESOURCE BULLETIN PNW-RB-252 Cole, T.G.; Falanruw, M.C.; MacLean, C.D.; Whitesell, C.D.; Ambacher, A.H. 1987. Vegetation survey of the Republic of Palau. Resour. Bull. PSW-RB-22. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 13 p. Endress, B.A.; Chinea, J.D. 2001. Landscape patterns of tropical forest recovery in the Republic of Palau. Biotropica. 33: 555–565. MacLean, C.D.; Cole, T.G.; Whitesell, C.D.; McDuffie, K.E. 1988. Timber resources of Babelthuap, Republic of Palau. Resour. Bull. PSW-RB-23. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 8 p. Mueller-Dombois, D.; Fosberg, F.R. 1998. Vegetation of the tropical Pacific Islands. New York: Springer-Verlag. 733 p. Oliver, C.D.; Larson, B.C. 1996. Forest stand dynamics. New York: Wiley. 520 p. Parker, A.J. 1982. The topographic relative moisture index: an approach to soilmoisture assessment in mountain terrain. Physical Geography. 3: 160-168. Smith, C.W.; Babik, N.R. 1988. Properties and management considerations of some acid soils on the islands of Palau. In: Demeterio, J.L.; DeGuzman, B., eds. Management and utilization of acid soils of Oceania. Guam: University of Guam: 39–61. U.S. Department of Agriculture, Forest Service [USDA FS]. 2003. Field instructions for the inventory of the Pacific Islands. Portland, OR: Pacific Northwest Research Station, Forestry Sciences Laboratory, Forest Inventory and Analysis. 256 p. U.S. Department of Agriculture, Natural Resources Conservation Service. 1983. Soil survey of Islands of Palau, Republic of Palau. Washington, DC: Soil Conservation Service. 120 p., 72 plates. Western Regional Climate Center. 2004. Koror, Palau, period of record monthly climate summary, 1953-2004. http://www.wrcc.dri.edu/cgi-bin/cliMAIN. pl?pikoro. (September 12). 52 Pacific Northwest Research Station Web site http://www.fs.fed.us/pnw/ Telephone (503) 808-2592 Publication requests (503) 808-2138 FAX (503) 808-2130 E-mail pnw_pnwpubs@fs.fed.us Mailing address Publications Distribution Pacific Northwest Research Station P.O. Box 3890 Portland, OR 97208-3890 U.S. Department of Agriculture Pacific Northwest Research Station 333 SW First Avenue P.O. Box 3890 Portland, OR 97208-3890 Official Business Penalty for Private Use, $300