Pacific Science (1999), vol. 53, no. 4: 346-356 © 1999 by University of Hawai'i Press. All rights reserved

The Freshwater Ichthyofauna of Bougainville Island, Papua New Guinea!

J. H. POWELL AND R. E. POWELL2

ABSTRACT: Tailings disposal from the Bougainville Copper Limited open-cut porphyry copper mine on Bougainville Island, Papua New Guinea (1972-1989) impacted the ichthyofauna of the Jaba River, one of the largest rivers on the island. To assess the 'extent of this impact, comparative freshwater ichthyologi­ cal surveys were conducted in five rivers on the island during the period 1975­ 1988. Fifty-eight fish species were recorded, including one introduction, Oreo­ chromis mossambicus. The icthyofauna is dominated by euryhaline marine spe­ cies consistent with that of the Australian region, but more depauperate. There are more than 100 species present on mainland New Guinea that are absent from Bougainville streams. Oreochromis mossambicus was the most abundant species in the sampled streams, accounting for 45% of the catch. The most abundant native fishes were the mainly small Gobiidae and Eleotridae. There were few native fish of potential value as food and these were restricted to an eleotrid gudgeon (Ophieleotris aporos), tarpon (Megalops cyprinoides), eel (An­ guilla marmorata), and snappers (Lutjanus argentimaculatus and Lutjanus fus­ cescens). Fish production in the rivers is limited by the morphology of the streams and the depauperate ichthyofauna. Fish yield from the Jaba River in its premining state is estimated to have ranged from 7 to 12 t/yr. The popula­ tion living in the Jaba ,catchment in 1988 (approximately 4,600 persons) shared this resource, resulting in an extremely low per-capita fish consumption rate of less than 3 kg/yr. '

THE FRESHWATER ICHTHYOFAUNA of the Aus­ most seven-eighths of the world's primary tralian region, which includes Australia, freshwater fishes (Darlington 1957), so the mainland New Guinea, and associated island absence of the group from the New Guinea groups, is unique because the dominant mainland has left the freshwater ichthyo­ groups of primary freshwater fish (Ostario­ fauna depauperate (Haines 1983) and limited physi) are not represented (Roberts 1978). in capacity for freshwater fisheries produc­ This has been confirmed by a number of tion (Coates 1985). ichthyological surveys of several major rivers It is well known that faunal diversity on on the New Guinea mainland, including the islands (including the freshwater fauna) is Fly (Roberts 1978), Laloki (Berra et al. naturally depressed, depending in part on 1975), Angabanga (Hyslop 1996), and Purari island area (MacArthur 1972) and the dis­ Rivers (Haines 1983) to the south; the Gogol tance of the island from the nearest continen­ (Parenti and Allen 1991), Ramu (Allen et al. tal land mass (Usinger 1963). The combined 1992), Upper Yuat (Jenkins 1997), and Sepik effects of reduced area, remote location, and Rivers (Allen and Coates 1990) to the north; the absence of ostariophysan fishes suggest as well as rivers in Irian Jaya (Allen and severe reduction of the freshwater ichthyo­ Boeseman 1982). fauna of the Papua New Guinea island The ostariophysan fishes account for al- groups. It is surprising that there has been rela­ tively little freshwater ichthyological study of 1 Manuscript accepted 10 February 1999. 2 Environmental Management and Planning Services the New Guinea islands except for records Pty. Ltd., P.O. Box 406, Samford, Queensland, 4520, provided by Munro (1967), Kailola (1975), Australia. and Allen (1989, 1991) for both marine and 346 The Freshwater Ichthyofauna of Bougainville Island, Papua New Guinea-PoWELL AND POWELL 347 freshwater fishes. Gressitt (1982) excluded source of high-quality protein to the indige­ freshwater fishes from his treatment of the nous inhabitants of the island is also assessed. ecology of New Guinea, and Choy (1986) concluded that freshwater ecology in the Pa­ Study Area cific Islands had been largely overlooked. Environmental issues associated with in­ CSIRO (1967) described the landforms of dustrial development in Papua New Guinea Bougainville and Buka Islands. The islands have imposed a requirement for management lie at the northern extremity of the Solomon and political decisions in the absence of de­ Island Archipelago on the western edge of tailed knowledge of local freshwater envi­ the Pacific Plate (Briggs 1987). The two is­ ronments. An example on the Papua New lands together are 238 km (150 miles) long Guinea mainland is the Ok Tedi copper and by 63 km (40 miles) wide, with a total area of gold mine (Allen 1991) and the Bougainville 8568 km2 (3475 square miles). The climate is Copper mine on Bougainville Island (Ellis wet tropical with an annual rainfall of ap­ 1988). Bougainville Copper Limited operated proximately 3000 mm (Ruppin 1988). Recent the large open-cut porphyry copper mine on volcanism has covered Bougainville Island Bougainville Island (6 0 00' S, 1550 30' E) with a mantle of ash often more than 1 m from 1972 until 1989, when civil unrest deep (Speight 1967), which, together with forced premature mine closure. The mine was continuing tectonic instability and rainfall, situated at an altitude of 900 m in Panguna has produced both high relief and a high rate in the headwaters of the Jaba River in the of erosion and deposition. Crown Prince Range of Bougainville Island, The configuration of Bougainville has in­ North Solomons Province, Papua New hibited development of very large stream Guinea (Figure 1). catchments (Speight 1967), the largest being 2 During the period 1972-1989, and with the Luluai catchment (464 km ). The rivers government approval, Bougainville Copper are short (generally < 35 km long) and tor­ Ltd. discharged tailings at rates up to rential. The headwaters and upstream areas 140,000 tjday into the JabajKawerong River of the rivers within the Crown Prince Range system. The tailings were discharged into the are boulder-strewn torrents passing into pool headwaters of the Kawerong River, and ap­ and riffle zones as stream gradient decreases. proximately 60% of the input entered the sea On the coastal lowlands the streams become 35 km downstream. Sedimentation on this meandering, swiftly flowing, generally single­ scale severely changed the morphology of the channel rivers. All the rivers surveyed are river and resulted in total loss of ichthyo­ nontidal; river flow is unidirectional at the fauna in the main channel and partial loss mouth, and the salt water mixing zone is off­ in tributaries. The Jaba River tributaries shore. Ryan (1991) described streams of sim­ were not directly influenced by sedimenta­ ilar morphology in Fiji. tion, but were isolated from the sea by high Detailed hydrological information is only suspended-solid loading in the main channel. available for the JabajKawerong system, To evaluate the extent of this impact and which has a catchment area of 440 km2 and to assist Bougainville Copper Ltd. with on­ a mean annual discharge of 44 m 3 sec-I going negotiations for loss of natural re­ (Ruppin 1988). sources resulting from its operations, ich­ thyological surveys of five streams on Bougainville Island were conducted during MATERIALS AND METHODS the period 1975-1988. The purpose of this paper is to document Fish were collected from a total of 22 sites the freshwater ichthyofauna of Bougainville distributed along five rivers: the Pinei, Ara­ Island in a biogeographical context and pro­ kawau and Luluai Rivers on the eastern side vide information about a largely unknown of the Crown Prince Range, and the Mari­ resource on a remote tropical Pacific Island. ropa and Pangara Rivers on the western side The importance of freshwater fishes as a (Figure 1). The Pangara River collections in- PAPUA NEW GUINEA ". c:o:' • a~ -,s)~"• ~ •' I O'tj~ 'It.. Bougainville IJ~ .'

_60S -¥'--f'>-._-..,.~------..,.,.-_r'_------

MOTUPENA OINT

o 50 KILOMETRES

15511E I

FIGURE 1. Bougainville Island showing river sampling locations and position of Panguna mine site in the head­ waters of the Jaba River. The Freshwater Ichthyofauna of Bougainville Island, Papua New Guinea-PoWELL AND POWELL 349

TABLE 1

SURVEY DATES AT FIVE RIVERS ON BOUGAlNVlLLE ISLAND DURING THE PERIOD 1975-1988

RIVER 1975 1976 1977 1978 1981 1982 1984 1986 1987 1988

Pinei X XX XX X X Arakawau X X X XX Luluai XX X X X Pangara X X X X X X X Mariropa X X X X cluded visits to two tributaries (the Orei [site fuscescens also was recorded previously from PA4] and Nunopa Rivers [sites PA3 and Bougainville Island by Allen and Talbot PA5]) in the upper reaches of its catchment. (1985) as were Stenogobius hoesei (Watson At least three sites were sampled in each 1991), Sicyopterus sp. 2 (Allen 1991), and river, one site in each major river zone: the Awaous ocellaris (Watson 1992). Several of upper boulder rapid zone, the middle pool the other species in Table 2 (Caranx sexfas­ and riffle zone, and the lower single-channel ciatus, Sicyopterus ouwensi, and Rhyacichthys zone. aspro) were also recorded by Kailola (1975), Each river was surveyed on several occa­ leaving 51 new records from the surveys re­ sions between 1975 and 1988 (Table 1). Fish ported here. All of these new records refer to were collected using a variety of methods, species that require access to marine or including 100-mm multifilament gill net (30 brackish environments to complete their life by 3.2 m), lO-mm knotless bait seine net (20 histories. by 5 m), 37-mm monofilament gill net (20 by Sites have been grouped into those repre­ 1 m), white lights and knives, diving and sentative of the lower reaches (approximately spearing, hook and line (angling and set 0-5 km from the mouth), middle reaches lines), and fish traps (drums). (approximately 6-10 km upstream), and up­ The fishing methods used by local land­ per reaches (approximately 11-20 km from owners are mainly traditional (white lights the mouth). Longitudinal zonation is appar­ and knives, spears, and hook and line). Gill ent, with the largest number of species (47) netting and seine netting are rarely, if ever, being recorded from the lower reaches and conducted. fewer species (30-32) from the middle and Captured fish were identified to species upper reaches. wherever possible, and the standard length Some species such as the eel (Anguilla and weight of individual fish were recorded. marmorata), grunter (Mesopristes cancella­ A complete reference fish collection was tus), flagtails (Kuhlia marginata and K stored at Bougainville Copper Ltd.'s Pan­ rupestris), and some representatives of the guna mine site at the time of mine closure. Gobiidae and Eleotridae occur throughout Only a small selection of duplicate specimens the river systems. Others occur mainly in the was lodged at the Queensland Museum, lower reaches: the tarpon (Megalops cypri­ Brisbane. noides), glass perchlets (Ambassis miops and A. nalua), sawfish (Pristis microdon), trevallys (Carangidae), and snappers (Lutjanidae). The upper reaches are inhabited by the ma­ RESULTS AND DISCUSSION jority of representatives of the Gobiidae and A species checklist and the sites at which Eleotridae in addition to the eels and flag­ they have been recorded are shown in Table tails. 2. A complete list of fishes including dates Most of the fish that attain larger size (and and authors of original descriptions is avail­ are therefore prized as food) inhabit the low­ able from the authors on request. Lutjanus er reaches of the streams (e.g., the tarpon, TABLE 2: A CHECKLIST OF THE FRESHWATER ICHfHYOFAUNA ON BOUGAINVILLE ISLAND SHOWING

LOWER REACHES MIDDLE REACHES

TAXA ARI LUi MAl MA2 MA3 PAl PH AR2 LU2 LU3 MA4 PA2 PI2 PI3

Pristidae Pristis microdon Megalopidae *Megalops cyprinoides 7 Anguillidae Anguilla marmorata 2 2 4 6 2 9 2 Opichthidae Lamnostoma orientalis Engraulidae Stolephorus tri 47 Syngnathidae *Microphis brevidorsalis *Microphis leiaspis 2 2 Ambassidae Ambassis urotaenia 15 *Ambassis interruptus 6 5 Ambassis macracanthus 4 13 21 86 7 2 Ambassis miops 14 9 1 Ambassis nalua 144 392 7 Terapontidae *Terapon jarbua 1 *Mesopristes argenteus 1 1 2 2 *Mesopristes cancel/atus 2 4 3 3 3 Kuhliidae *Kuhlia marginata 3 28 83 3 1 17 6 7 2 51 13 *Kuhlia rupestris 3 3 3 13 2 4 6 Carangidae Caranx sexfasciatus 2 2 Trachinotus russelli 1 Leiognathidae *Leiognathus equulus 3 Lutjanidae *Lutjanus argentimaculatus 2 4 Lutjanus fulviflamma 1 *Lutjanus fuscescens 7 6 Gerridae Gerres filamentosus Pomadasyidae Pseudopristipoma nigra 2 Mullidae Upeneus vittatus Toxitidae Toxotesjaculatrix 2 Scatophagidae *Scatophagus argus 4 CicWidae Oreochromis mossambicus 96 772 40 2 Mugilidae *Cestraeus goldiei 1 *Crenimugil heterocheilus 18 1 5 17 2 23 7 *Liza subviridis 3 2 10 6 10 6 Valamugil buchanani 7 Valamugil seheli Polynemidae *Polydactylus plebeius 2

352 PACIFIC SCIENCE, Volume 53, October 1999

TABLE 2 (continued)

LOWER REACHES MIDDLE REACHES

TAXA ARI LUI MAl MA2 MA3 PAl PIl AR2 LU2 LU3 MA4 PA2 PI2 PI3

Gobiidae *Awous melanocephalus 53 34 4 2 9 4 5 46 3 *Awous ocellaris 1 6 1 *Glossogobius celebius 1 1 7 6 3 1 1 Glossogobius giuris 1 8 1 Goby sp. 2 *?Schismatogobius sp. *Sicyopterus ouwensi 2 6 4 21 3 7 *Sicyopterus sp. 2 (Allen 1991) 2 *Sicyopterus sp. A 2 *Sicyopterus sp. B 13 *Sicyopterus sp. C *Stenogobius hoesei 2 *Stiphodon elegans *Stiphodon sp. Eleotridae Belobranchus belobranchus Butis butis I *Hypseleotris guentheri 160 254 3 126 *Ophieleotris aporos 1 22 49 3 4 5 19 49 15 Ophiocara porocephala 5 *Oxyeleotris gyrinoides 2 10 3 3 3 33 6 Oxyeleotris sp. 1 1 12 13 Rhyacicthyidae *Rhyacichthys aspro 9 Tetraodontidae Chelanodon patoca 1 Total no. of species 13 23 2 17 23 9 21 16 14 7 11 13 17 13

• Lodged at Queensland Museum, Brisbane, Australia. eels, and snappers). Food fish are less plenti­ island in 1959 by the Department of Agri­ ful in the middle and upper reaches of the culture, Stock, and Fisheries in an attempt to streams. However, these larger species are increase food production (Glucksman et al. not abundant (total number captured: tar­ 1976). Carp were not collected in this series pon, 10; eels, 40; and snappers, 24), indicat­ of surveys and seem not to have become ing that the rivers are probably not a plenti­ established. However, tilapia escaped from ful source of fish for the local inhabitants. ponds in the catchments of several rivers and Allen (1991) reported that the freshwater were very abundant in the Pangara River. fauna of New Guinea (papua New Guinea The Pangara catchment is isolated from the and Irian Jaya) consisted of 329 species, of sea as a result of tailings disposal in the Jaba which 13 were introduced forms and 102 River. Because of this isolation, some im­ were marine species with life history stages portant piscivorous species are absent from requiring salt water. In contrast a total of 58 the Pangara catchment: the grunters (Meso­ species (5960 individuals) was collected from pristes cancellatus and M argenteus), the Bougainville (18% of the mainland fauna), jungle perch (Kuhlia rupestris), and the one­ the most abundant being the introduced spot perch (Lutjanus fuscescens). Tilapia has African species Oreochromis mossambicus proliferated in the absence ofthese predators. (tilapia), which accounted for 45% of the The native ichthyofauna as an assemblage catch (2742 individuals). Tilapia and carp is typical ofthe Tropics, being composed of a (Cyprinus carpio) were introduced to the reasonably large number of species (57) none The Freshwater Ichthyofauna of Bougainville Island, Papua New Guinea-PoWELL AND POWELL 353

UPPER REACHES TOTAL SIZE RANGE TAXA AR3 LU4 LU5 MA5 PA3 PA4 PA5 PI4 NO. (SL IN ,\1M)

Gobiidae *Awous melanocephalus 3 3 2 2 2 174 20-130 *Awous ocellaris 1 9 13-84 *Glossogobius celebius 6 4 2 37 57-180 Glossogobius giuris 3 1 16 42-136 Goby sp. 2 5 41-96 *?Schismatogobius sp. 1 1 *Sicyopterus ouwensi 73 117 2 24 4 7 40 313 39-122 *Sicyopterus sp. 2 (Allen 1991) 1 1 4 39-122 *Sicyopterus sp. A 1 3 3 12 2 24 62-65 *Sicyopterus sp. B 14 1 4 7 39 34-67 *Sicyopterus sp. C 1 34-40 *Stenogobius hoesei 6 58-72 *Stiphodon elegans 1 *Stiphodon sp. 1 40 Eleotridae Belobranchus belobranchus 2 82-142 Butis butis 1 87 *Hypseleotris guentheri 53 26 625 32-65 *Ophieleotris aporos 15 8 4 4 202 48-230 Ophiocara porocephala 5 59-122 *Oxyeleotris gyrinoides 12 2 3 1 80 75-266 Oxyeleotris sp. 6 1 36 44-141 Rhyacicthyidae *Rhyacichthys aspro 12 3 36 2 5 71 61-198 Tetraodontidae Chelanodon patoca 1 134 Total no. of species 22 4 14 14 8 8 11 14 58

of which is particularly abundant. Ten spe­ of Bougainville Island is consistent with that cies account for 84% of the catch, and most of the Australian region, but more depau­ ofthese, except for the flagtails (K marginata perate because of the absence of representa­ and K rupestris), belong to smaller bait-sized tives from the families Plotosidae, Ariidae, representatives of the families Ambassidae, Melanotaenidae, Clupeidae, Osteoglossidae, Gobiidae, and Eleotridae. Forty of the 57 Atherinidae, Lobotidae, Apogonidae, Solei­ native species are rare (less than 20 indi­ dae, Hemiramphidae, Belonidae, Kurtidae, viduals for each of these species captured and Cynoglossidae, a total exceeding 100 from all surveys). species (Roberts 1978, Allen and Boeseman There also seems to be no freshwater fish 1982, Allen 1991). endemic to Bougainville Island, although en­ Munro (1967) listed about 220 species of demicity might be revealed by revision of the native freshwater fish from Papua New Gobiidae and Eleotridae. Most of the species Guinea alone, and McDowall (1981) consid­ belong to the categories defined by Lowe­ ered that there are about 165-170 species of McConnell (1975) as (1) representatives of freshwater fish in Australia. The low number marine families (e.g., the tetradont puffers); of native fish species on Bougainville con­ (2) diadromous fishes (e.g., the anguillid eels); firms expectation and results from a combi­ and (3) occasional visitors that are euryhaline nation of remote location, small area, and representatives of marine families (e.g., Pris­ recent geological development. The 57 native tidae, Carangidae, and the Mugilidae). species on Bougainville Island are more con­ In this regard, the freshwater ichthyofauna sistent with the 80 species reported from Fiji 354 PACIFIC SCIENCE, Volume 53, October 1999 by Ryan (1980 and 1991), but the Fiji Island tion to the influence of sea level changes on group extends over 4 degrees of latitude, the morphology of streams and the effects of whereas Bougainville extends over just 2 de­ volcanism on water quality. Volcanic ash grees of latitude, from 5° to 7° S. production coupled with high rainfall, high Most of the species recorded from Bou­ river discharge, and turbidity are additional gainville are widely distributed throughout factors that would not favor development of the Indo-Pacific (see Weber and de Beaufort a prolific freshwater ecology. 1953, Fowler 1959, Masuda et al. 1984, Smith and Heemstra 1986). For example, Fish Yield Lutjanus argentimaculatus and Lutjanus ful­ viflamma extend from eastern Africa to the There is no commercial freshwater fishery western Pacific including Fiji, Samoa, and on Bougainville. However, because mining New Caledonia, and Lutjanus fuscescens is activities have disrupted some of the streams, found in freshwaters of China, the Philip­ an assessment of the potential for freshwater pines, Indonesia, and Papua New Guinea fish production has been attempted to deter­ (Allen and Talbot 1985). Bougainville Island mine the extent of fish loss. is the easternmost record for this species to Coates (1985) prepared estimates of fish date. yield for the Sepik River, mainland Papua Although Bougainville lies west of the an­ New Guinea, and concluded that a realistic desite line, it can probably be considered an fish yield figure for the Sepik might be 3000­ oceanic island in the zoogeographical sense 5000 t/yr. The catchment area of the Jaba of having acquired its freshwater ichthyo­ River is 440 km2 (Ruppin 1988), 0.56% that fauna by overseas dispersal. Many of the of the Sepik (catchment area 78,000 km2 marine euryhaline genera represented (e.g., [Coates 1985]). Discharge figures provide a Butis, Megalops, Gerres, Pristis, Scato­ similar ratio (Jaba River mean discharge, phagus, and Ambassis) confirm Pacific Plate 40 m 3 sec-I; Sepik River average discharge, marginal records listed by Springer (1982). 8000 m 3 sec-I). Based on the ratio of catch­ The dominance of Gobiidae and Eleotridae ment areas and discharge, the yield from the combined (21 of 57 species [37%]) is also Jaba River would be unlikely to exceed 25 consistent with patterns presented by Ryan t/yr (range, 17-28 t/yr), provided the same (1991) for Fiji, Vanuatu, and Samoa. The species were present. A more realistic yield presence of the amphidromous sicydiine figure for the Jaba River will be less than this gobies Stiphodon from the Pacific and because the ichthyofauna is even more de­ Sicyopterus from the Indo-Pacific reflects pauperate than that of the Sepik. Bougainville's transitional position between The average catch rate from the 100-mm these two major regions and reinforces the gill net in Bougainville streams was 9.5 g/m2 link between ocean basins and biogeography per set, which is approximately 60% of the suggested by Parenti (1991). catch rate (12.7-22.6 g/m2 per set) quoted by However, the island has also been influ­ Coates (1985). If 60% is applied as a mod­ enced by the added effects of volcanism and ifying factor to account for the depauperate plate tectonic activity. Weber (1973) con­ ichthofauna of Bougainville rivers, then a re­ firmed earlier reports by Stoddart (1969) of alistic fish yield for the Jaba River might be widespread mass mortality of reef corals in 10-17 t/yr. These estimates can be further the central Solomon Islands. Stoddart con­ reduced by at least 30% to eliminate the sidered that a contributing factor in coral contribution made by tilapia to the catch in mortality was tectonic activity resulting in the Sepik (Coates 1985), resulting in a yield noneustatic changes in sea level. Therefore, for native species in the Jaba River of 7-12 in addition to remote location and limited t/yr. A similar figure is obtained « 6 t/yr) if niches associated with island biogeography the world average for freshwater fish yield (Simmons 1979), evolution of the ichthyo­ (1.0-1.5 g/m2 [Lowe-McConnell 1975]) is fauna on Bougainville has included adapta- applied to the combined water surface area The Freshwater Ichthyofauna of Bougainville Island, Papua New Guinea-PoWELL AND POWELL 355 of the channels within the Jaba River system thyological survey of the Sepik River, Pa­ (approximately 4 x 106 m 2). pua New Guinea. Rec. West. Aust. Mus. The population density of the Jaba catch­ Suppl. 34: 31-117. ment was 10.45 persons/km2 (basin area, ALLEN, G. R., and F. H. TALBOT. 1985. 440 km2; population, 4600), whereas the Indo-Pacific fishes No. 11. Review of the population density ofthe Sepik was only 1.15 snappers of the genus Lutjanus (pisces: 2 2 persons/km (basin area, 78,000 km ; popu­ Lutjanidae) from the Indo-Pacific, with lation, 68,000 [Coates 1985]). It is quite pos­ the description of a new species. Bishop sible that the population density of the Jaba Museum Press, Honolulu. system (being nine times that of the Sepik) ALLEN, G. R., L. R. PARENTI, and D. exceeds the biological capacity of the river to COATES. 1992. Fishes of the Ramu River, support even a subsistence fishery, so that an Papua New Guinea. Ichthyol. Explor. average per-capita fish consumption rate of Freshwaters 3: 289-304. < 3 kg/yr is likely. BERRA, T M., R. MOORE, and L. F. REY­ In conclusion, these preliminary estimates NOLDS. 1975. The frehwater fishes of the of fish yield, taken in conjunction with pop­ Laloki River System of New Guinea. Co­ ulation figures, appear to support the obser­ peia 1975: 316-326. vation made by Oliver (1973) that fishing is BRIGGs, J. C. 1987. Biogeography and plate an "occasional pastime which does not add tectonics. Developments in Palaeontology significantly to diet." and Stratigraphy, 10. Elsevier, Amsterdam. CHOY, S. 1986. Freshwater ecology: A ne­ glected science in the Pacific Islands. SPC Fish. Newsl. 36: 25-26. ACKNOWLEDGMENTS COATES, D. 1985. Fish yield estimates for the We thank Bougainville Copper Ltd. for Sepik River, Papua New Guinea, a large logistic support and funding throughout the floodplain system east of "Wallace's surveys. Sponsorship for preparation of this Line." J. Fish BioI. 27: 431-443. paper was arranged by Pam Ruppin in her CSIRO. 1967. 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