Chapter 6

Status of biodiversity in Papua New Guinea

    Miller, S.; Hyslop, E.; Kula, G., and Burrows, I.


      Papua New Guinea probably harbours more than five percent of the world's biodiversity within some of the world's most biologically diverse ecosystems. Many of these organisms are endemic; that is, they are found only in Papua New Guinea or on the island of New Guinea. This chapter reviews the status of knowledge of Papua New Guinea's biodiversity from a taxonomic perspective, for example, by group of organism. Chapter 7 reviews the extraordinary range of environments which exist in Papua New Guinea. Many of the same processes that have promoted the evolution and maintenance of Papua New Guinea's biodiversity have also provided the diversity of habitats that we see today, so there is some overlap between Chapters 6 and 7.

      The term "diversity" (or biological diversity) means many things, and many measures have been proposed to quantify diversity (Cousins 1991; Holloway and Stork 1991; Vane-Wright et al. 1991). At least four major types of diversity measurement can be recognised:

        • species richness — the absolute number of species in a given area;
        • a combination of species richness and abundance — taking account of rarity and balance within the species found in a given area;
        • genetic heterogeneity — variability within species; and
        • taxonomic distinctness — incorporating an historical (evolutionary distance) component.


      This chapter focuses on species richness because very few data are available for the other categories for Papua New Guinea.

      This chapter is based heavily on Volume 2 of the Papua New Guinea Conservation Needs Assessment (Beehler 1993), which should be referred to for more detailed background, discussion, and bibliographies. We have also gathered data from the staff and libraries of the University of Papua New Guinea, Papua New Guinea University of Technology, Wau Ecology Institute, Christensen Research Institute, Bishop Museum, and elsewhere.

      In many cases, summary data are available only at the level of the island of New Guinea. In some cases, specialised collecting or analysis techniques have been applied only in the western half of the island of New Guinea, so data from Irian Jaya must be extrapolated to Papua New Guinea. In Chapters 6 and 7, the term New Guinea will be used to include the island of New Guinea and associated smaller islands, including all of Papua New Guinea (the region called Papuasia by botanists). Moreover, the Papua New Guinea biota cannot be understood in isolation because much of the biota is shared with Irian Jaya, the Solomon Islands, and Australia.


      Status of Biological Knowledge

      The many institutions interested in Papua New Guinea’s terrestrial biodiversity have organised over 100 expeditions to areas throughout the country (Frodin and Gressitt 1982; Allison 1991). These areas were generally chosen on the basis of their biological interest (for example, high diversity, high endemism, and interesting geography). Although major gaps remain in coverage (especially in marine environments), knowledge of Papua New Guinea’s biodiversity is much better than that of many other tropical countries. Papua New Guinea is much better known than Irian Jaya for most (but not all) organisms — a factor which complicates analyses of endemism and distribution patterns.

      Frodin and Gressitt (1982) presented an excellent history of the terrestrial biological exploration of New Guinea, extended for botany by Stevens (1989) and Frodin (1990), so only the major elements will be repeated here. Biological exploration of New Guinea by Western scientists started in the mid-1800s. The earliest expeditions collected relatively few specimens, however, and these specimens usually have relatively non-specific locality data. These expeditions provided the first specimens of many species that are endemic to New Guinea, but more comprehensive collections must be consulted to understand distribution patterns. The major period of exploration by foreign-based scientists occurred from about 1920 to 1950. Some of the most important work was associated with the first (1933-34) through seventh (1964), and especially the third (1938-39) Archbold Expeditions. Starting in the 1950s, local scientific institutions began to be built in Papua New Guinea and a great deal of research was done by scientists in residence at those institutions. Various government departments, especially in agriculture, fisheries, forestry, and public health, developed research programs and scientific collections. The Lae Herbarium, now part of the Forest Research Institute, stands out as a major, world-class centre for botanical studies. Universities were developed at Port Moresby and Lae, with field research stations at Motupore Island and elsewhere. In 1961, the Bishop Museum established a field station at Wau. This evolved into the independent Wau Ecology Institute in 1972 and continues to be a centre of biological research and conservation programs. Because of sampling intensity, Wau has become one of the best known places in the country for terrestrial biodiversity. The recent establishment of the Christensen Research Institute near Madang has dramatically increased knowledge of marine biodiversity.

      The result of the activities in biological research in New Guinea is a host of resources within Papua New Guinea, and at major international research centres (especially Sydney, New York, London, Leiden, Honolulu, Canberra, Boston, and Bogor). Overall, a very large body of data exists, but it is scattered through scientific literature (journals and books) that has accumulated over the last 200 years and collections located around the globe. For example, it has been estimated that some 200 000 reptile and amphibian specimens (Allison 1993) and 450 000 plant specimens (Stevens 1989) exist from New Guinea. Bringing these data together in a modern information management system is a major challenge, but it is possible and will provide the knowledge base for resource management (Miller 1993). However, even a comprehensive compilation of existing knowledge would still have major gaps, so additional sampling and research is desperately needed. Even for plants, the overall density of sampling remains poor and is one of the lowest in the Malesian botanical region (Stevens 1989; Frodin 1990). The tremendous geographical diversity in Papua New Guinea requires adequate samples on which to base analysis of variation within and amongst species to correctly define those species. Increasing the absolute knowledge of Papua New Guinea's biota will be accomplished only by improving in-country scientific infrastructure (especially collections) and human resources.

      The gaps in existing biological knowledge include the following (see the Conservation Needs Assessment for more details): many areas within Papua New Guinea have never been adequately sampled (see map and list in Chapter 7); many organisms that are small in size or live in habitats that are difficult to sample have been inadequately collected (especially fungi, soil invertebrates, and some marine invertebrates); and many of the most diverse organisms have never been subjected to modern taxonomic analysis (especially some insects and marine invertebrates).


      How can Papua New Guinea fill these gaps in knowledge of biodiversity? The most important step is to create an information management system within Papua New Guinea that can receive, manage, and disseminate biodiversity knowledge. Such a centre does not necessarily have to own and maintain all the data, but could serve a coordinating and networking role. Without such a centre, data that flow from national and international institutions is effectively lost. The processes of the Conservation Needs Assessment and this Biodiversity Country Study demonstrated how difficult it is to find biodiversity information on Papua New Guinea within the country. Additional processes needed are:

        • strengthening national institutions involved in biodiversity and their human resources;
        • collaborative projects with international research centres to make New Guinea data available; and
        • additional field surveys, taxonomic research, and ecological research.


      National Species Diversity, Endemism, and Status

      Table 6.1 summarises knowledge of major taxonomic groups of organisms. Classification in the table generally follows Parker (1982) with the use of five kingdoms (Margulis and Schwartz 1988). Some artificial categories of convenience are used (for example, marine molluscs). Freshwater categories generally include brackish water dwellers. Because data are often available for New Guinea rather than Papua New Guinea, we have specified which figure is provided. For some marine invertebrates, faunal data are only available for specific regions (for example, Madang, Motupore Island, or the Torres Strait).

      Endemism refers to species unique to a certain area. Endemism tends to follow biogeographic boundaries determined by geographic factors and evolutionary history. It is important to note that biogeographic boundaries do not correspond with the political boundaries of Papua New Guinea. The western half of the island of New Guinea is the Indonesian State of Irian Jaya. The island of Bougainville is part of Papua New Guinea, but is biogeographically most similar to the Solomon Islands. The same conflict between biogeographic and political boundaries arises across the Torres Strait between the southern part of Papua New Guinea and the northern tip of Queensland, Australia (Kikkawa et al. 1981). Another factor which affects apparent endemism is the level of study which has taken place on particular taxonomic groups, for example, the relatively visible bird species are well-documented, whereas only limited information exists on many invertebrates. Thus, some organisms are known only from a particular place simply because they have not yet been identified elsewhere. At individual sites, high species diversity (absolute number of species) does not necessarily correlate with high numbers of endemic species. Additional information on species which are endemic to Papua New Guinea may be found in the Conservation Needs Assessment report and maps.

      As noted previously, there is a vast literature on the biota of New Guinea, but it is scattered in an array of books and journals in many languages. Outside of the vertebrates, there are very few synthetic papers which review entire groups for any particular region of interest. We have used the Conservation Needs Assessment and literature that could readily be gathered to construct Table 6.1. We have not undertaken a detailed assessment of the scattered data in the taxonomic literature (for example, Papua New Guinea records within taxonomic studies on a world basis). This would produce a great deal of data and should be done, but it will be a time consuming activity. Ewers (1973) provides a window on a small part of the taxonomic literature. A search of the computer database version of Biological Abstracts showed some 4 000 papers on biology (including ecology and some medical subjects) of New Guinea from 1970 to 1994, an average of some 166 titles per year. This is, in fact, an underestimate, because it under-samples the taxonomic literature (for example, New Guinea records within monographs covering larger areas). There is also a vast number of additional records that are found only in collections and have never been mentioned in the literature. These, too, could be compiled into a very useful database by gathering data from the collections in Papua New Guinea and combing the major museums and herbaria of the world for Papua New Guinean specimens.


      Viruses, Bacteria, and Algae

      Very poor data are available for unicellular organisms such as viruses and bacteria outside of those with direct economic importance in agriculture (Shaw 1984; Muthappa 1987) and the health of humans and domestic animals. Algae, both freshwater and marine, are poorly documented beyond a few studies (especially Vyverman 1991a, 1991b).



      Shaw (1984) listed 2 390 fungi from Papua New Guinea. Hawksworth (1994 personal communication) suggests a ratio of 1:6 between vascular plants and fungi. Using a conservative estimate of 15 000 vascular plants yields an estimate of 90 000 fungal species. Lichens, which are symbiotic combinations of fungi and algae, were reviewed by Streimann (1986) and Lambley (1991). Bryophytes, the mosses and liverworts are the subject of active research that has not yet been synthesised (Grolle and Piippo 1984; Piippo 1994).



      Protozoans are almost entirely unstudied, with the exception of marine foraminifera, which have been censused at Motupore Island (57 species — Haig 1979) and Madang (182 species — Langer 1992).



      The vascular plants probably include 15 000 to 20 000 species of ferns and flowering plants. There is a recent list of genera (Hoeft 1992), but modern species-level treatments exist for only a small portion of the flora (for example, Womersley 1978; van Royen 1980; Henty 1981). Most of Papua New Guinea is within the region covered by the ongoing Flora Malesiana project. The overall status of knowledge of plants has been reviewed recently by Johns (1993), Stevens (1989), and Frodin (1990). Orchids are particularly diverse, with well over 3 000 species (van Royen 1979; Howcroft 1992). Seagrasses and seaweeds are important in marine ecosystems.



      Knowledge of the taxonomy and distribution of invertebrates in the New Guinea area is much less focused than is the case for vertebrates. The information to a large extent reflects the interests of individual taxonomists who have worked on particular groups often in circumscribed areas. The marine invertebrates, other than molluscs and crustaceans, are very poorly known. Some groups have been studied at Motupore Island and Laing Island, and many are now under study at Madang, but much remains to be done. A large number of animal phyla are represented by marine invertebrates that are effectively unknown in Papua New Guinea but are probably major components of the marine biodiversity. The situation can be illustrated with corals, based on studies in progress at Madang by D. Potts and colleagues.

      The total number of coral species in Papua New Guinea is unknown although about 300 species have been reported. No established coral taxonomist has worked extensively in New Guinea, and serious scientific collections, suitable for assessing biodiversity, are extremely limited. The only intensive study of a single site by an experienced coral taxonomist listed 285 living species from Motupore (Veron and Kelley 1988). On the north coast, four collections have been made near Madang (J. Oliver in 1988; B. Hoeksema in 1992; J. Pandolphi; and D. Potts in 1994); an extensive study at Laing Island is now being published (Claerebout); and largely undescribed collections have been made by geologists working on uplifted fossil reefs of the Huon Peninsula. Veron (1993) recognises more than 650 reef-building coral species in the entire central Indo-Pacific region (northern Australia to Japan), based mainly on his own work in Australia, Japan and the Philippines. Most have ranges from Australia to Japan. Although he includes only one Papua New Guinean site (Motupore), it is likely that most of these species also exist in Papua New Guinea and that other undescribed species are also common. Thus, the total number of reef-building coral species in Papua New Guinea is likely to be at least 700 (D. Potts, personal communication, 1994).

      Yet, coral biodiversity in Papua New Guinea may be much greater than 700 species. Despite their abundance, large size, and geological and biological importance, coral taxonomy is very poorly known. The world has only seen about 25 serious coral taxonomists over the last century, and most of these have been geologists more concerned with fossils than living forms. Species recognition is notoriously difficult even for experts, and especially in the most diverse genera and families (for example Acropora, Montipora, Porites, and Faviidae). There has been a strong tendency to group morphologically similar forms into a relatively small number of highly variable species with cosmopolitan distributions (for example, nearly 300 species of world Porites were described before 1900, but only about 30 are now accepted). Recent genetic work demonstrates that species names are not being applied consistently in adjacent localities and that some presumably widely distributed "cosmopolitan" species may consist of suites of locally endemic species (Potts, in preparation). Independent taxonomic studies of the same individual corals using live tissues, genetics, micromorphometrics, and traditional skeletal morphology show that the traditional characters have the least ability to discriminate species consistently (Potts et al. 1993). For these and other reasons, it seems likely that future coral taxonomy will recognise many more species than at present, and that many will be endemics restricted to local regions such as Papua New Guinea. Thus, total Papua New Guinean coral biodiversity may exceed 1 000 reef-building species, with a similar number of smaller, cryptic, often deepwater species of non-reefbuilders (which have never been seriously collected anywhere in Papua New Guinea).

      Studies of other invertebrate groups, such as sea pens (Pennatulacea), nudibranchs, and crustaceans indicate that the Madang lagoon and vicinity may have the highest species richness of any site in the world. Hoeksema (1993) found this was also true for mushroom corals (Fungiidae). He found 36 of the 42 species previously described worldwide, and also described a new species. It is also likely that other species exist in the area. Moreover, as with corals, genetic data are calling into question the entire concept of shallow water "cosmopolitan" species in many invertebrate groups (for example, jellyfish — Greenberg et al., in press).


      Other faunal studies on marine invertebrates include sponges (25 common species from Motupore Island — Kelly Borges and Bergquist 1988), various coelenterate groups (Bismarck Sea — Bouillon 1984; Bouillon et al. 1986; Pages et al. 1989), sea anemones (18 species in Madang vicinity — Fautin 1988), octocorals (83 species from Bismarck Sea — Verseveldt and Tursch 1979), and echinoderms (177 species from Torres Strait — Clark 1921).

      Some work has been done on marine crustaceans (crabs, etc.), but much work remains to be done. Morgan (1988) reported 198 species of marine and freshwater decapod crustacea from the Madang region. Marine molluscs are probably better known than most other invertebrates because of the popularity of their shells with collectors. However, the existing knowledge is superficial and does not penetrate the very high diversity that exists in Papua New Guinea. A popular guide to shells of Papua New Guinea includes 950 species (Hinton 1979), but there are more than 800 prosobranch gastropods (snails) alone in New Guinea (Wells 1990), 268 of which are found at Motupore Island (Signor et al. 1986). Ongoing studies of nudibranchs in Madang Lagoon have recorded at least 600 species and are expected to reach an eventual total of 700-800 species (T. Gosliner, personal communication).

      The status of terrestrial, freshwater, and brackish molluscs (mostly snails) was reviewed by Cowie (1993). Some 650 species have been described but they are in great need of synthetic revision and many species remain unknown.

      Terrestrial and freshwater invertebrates other than insects and molluscs have been very poorly sampled and studied. Several recent studies include earthworms (at least 42 species — Easton 1984; Nakamura 1992), leeches (at least five species — Van der Lande 1994), flatworms (Sluys and Ball 1990), and onychophorans (seven species — Van der Lande 1993). Freshwater decapod crustaceans (crabs, crayfish, etc.) include at least 87 species, with at least 31 endemic species (Eldredge 1993). Nematodes, except those that parasitise crops, livestock, and humans, are almost unknown in Papua New Guinea, although there must be a very large number of species. Freshwater rotifers include at least 135 species, all widespread outside Papua New Guinea (Segers and De Meester 1994).

      Information on New Guinea insects and related terrestrial arthropods is in drastic need of collation, and the quality and quantity of knowledge is uneven for different groups. Generally, levels of diversity and of endemism are high amongst the insects. The New Guinea fauna is primarily derived from the oriental region, but Australian elements are present also, especially in southern savannahs. The general status of knowledge of insects in Papua New Guinea was reviewed by Miller (1993). The bibliography by Gressitt and Szent-Ivany (1968) has been the only broad attempt to synthesise knowledge of Papua New Guinean insects, although reviews exist for some groups (for example, flies in Evenhuis 1989). Given ratios of the New Guinea fauna to the world fauna for groups that are well known, rates of description of new taxa, and the size of the Australian insect fauna (CSIRO 1991), there may be 300 000 species of insects in New Guinea, but this figure may be high or low by 100 000 species. Despite the relatively small area, Papua New Guinea ranks twelfth amongst world nations in terms of endemism of large butterflies (Papilionidae, Pleridae, Nymphalidae, 56 of 303 species are endemic) (Sisk et al. 1994). The mites of New Guinea are very poorly known, but have the potential for being a megadiverse group.



      Existing knowledge of the fishes of New Guinea has been well synthesised in field guides (Allen 1991; Allen and Swainston 1993) and in a checklist (Kailola 1987-1991), but more research remains to be done. There are well over 3 000 fishes in the region, including over 300 found in freshwater. The native freshwater fish fauna is derived from the marine fauna, and primary freshwater ostariophysian fishes are absent with the exception of Scleropages spp. (Roberts 1978; McDowell 1981). All species naturally occurring in freshwater are either diadromous or descendants of marine families. This has been noted to have an adverse effect on species diversity and fish stock abundance resulting in low fisheries yields (Coates 1985). Of the 329 species occurring in freshwater in New Guinea (Allen 1991), 13 species are non-indigenous, and about 102 species are believed to have a marine larval stage and are relatively widespread outside New Guinea. This leaves 214 native fish species that are limited to freshwater, of which 149 (70%) are endemic to New Guinea. Two closely related families are unique to Australasia — Melanotaeniidae (Rainbowfishes) and Pseudomugilidae (Blue-eyes), and this fauna further differs to that of other continental tropical regions which are dominated by cichlids and primary division Ostariophysan fishes (carps, barbs, loaches, characins, and catfishes). The Ostariophysan assemblage is represented in New Guinea-Australia only by plotosid and ariid catfishes. All the freshwater fishes except the lungfish (Neoceratodus), bony tongues (Osteoglossus), and possibly galaxiids are considered to be derived from marine ancestors.

      The freshwater ichthyofauna can be clearly divided into two zoogeographic regions. Freshwater bodies to the south of the central cordillera have an ichthyofauna closely allied with that of northern Australia, reflecting a former land connection. While several of those species with diadromous habits can be found in both southern and northern rivers, the fish permanently inhabiting freshwater in the north are invariably different species from those in southern water bodies. Apart from the land barrier formed by the central cordillera, northern rivers are much younger than southern rivers. Of those fish families common to both northern and southern rivers, species diversity is invariably lower in the north (Coates 1987b).



      Of the three orders of amphibians, neither caecilians nor salamanders occur in Papua New Guinea. Frogs (Anura: five families) are well represented, with 197 species described at present, and new species being recognised as current research proceeds (Allison 1993). The five families are: Bufonidae (the introduced cane toad, Bufo marinus), Hylidae, Leptodactylidae, Microhylidae, and Ranidae (Allison 1993). Not all species are aquatic — a large number are forest dwellers which burrow beneath the surface, or live beneath the leaf litter. The majority of species are endemic to either Papua New Guinea or the island of New Guinea. A southern group having its origins from Australia can be recognised, as can a group of species originating from the Solomon Islands to the south-east of Papua New Guinea. The surrounding islands have, in general, a depauperate amphibian fauna in comparison with the adjacent mainland. Much taxonomic work remains to be done, and the rich variation in specialised habitat requirements which is already known, suggests that many more species are yet to be described. The only toad in Papua New Guinea (Bufo marinus) is a classic example of a poorly planned biological control introduction.



      The reptilian fauna of New Guinea consists of representatives of all four main groups. The turtles and tortoises (Chelonia), number thirteen species in total, seven associated with freshwater and six with marine habitats (Allison 1993). Three freshwater species are endemic in the broad sense, of which one, Chelodina parkeri, is restricted to the Fly River system.

      The bulk of the reptiles are lizards (Squamata) with approximately 195 species of lizards and 98 species of snakes (Allison 1993). Most of the lizard species (71%) belong to the family Scincidae with the Agamidae and Gekkonidae both represented by more than ten species. The overall endemism is about 60 percent. The snakes belong to seven families, all of which are shared with Australia. Among the snakes, the level of endemism is considerably less than that found for lizards. At least 32 species (33%) can be considered endemic to the island of New Guinea and northern Australia.

      There are two described species of crocodile in Papua New Guinea — the New Guinea or freshwater crocodile (Crocodylus novaeguineae) and the saltwater crocodile (Crocodylus porosus). Both species are still found in relatively large numbers and are heavily exploited for hides and meat. However, freshwater crocodile populations along the south coast probably represent an undescribed species (Allison 1993: 165).



      The birds of Papua New Guinea are relatively well studied compared to other animals (Beehler et al. 1986; Coates 1985, 1990; Osborne 1987), and show a high degree of endemism. To date, a total of 762 species of bird have been recorded from Papua New Guinea, of which 405 (53%) are endemics (Papua New Guinea Bird Society Checklist, in preparation). This figure is higher than some other recent figures (for example, 725 species in New Guinea — Beehler et al. 1986; and 740 species in New Guinea — Coates 1985-1990) because of the inclusion of vagrants, taxonomic changes, and additional islands covered. Of these endemics, 289 (39%) are confined to mainland Papua New Guinea and/or the D'Entrecasteaux group of islands. Sixty (8%) are endemic to the Bismarck Archipelago and/or Admiralty Islands with nine of these extending into the Solomon Islands. Thirty (4%) are endemic to the Solomon Islands. Thirty-eight species are shared between Papua New Guinea and Northern Queensland while the 357 non-endemics are generally more widely spread. On a world basis, Papua New Guinea ranks fifth in terms of the number of restricted-range bird species and seventh in the number of endemic bird areas as mapped by the International Council for Bird Preservation (Bibby et al. 1992).



      Papua New Guinea is impoverished in terms of the range of mammalian orders compared to South-East Asia (nine orders of mammals found there which are lacking in Papua New Guinea). At least two of the existing four mammalian groups — marsupials and rodents — show a high degree of endemism. However, with 187 indigenous mammals (Flannery 1990; Menzies 1991), the New Guinea region has only slightly fewer mammal species than Australia, although the surface area of the former is approximately ten percent that of Australia. In addition, some 24 marine mammals (cetaceans) and the dugong occur within Papua New Guinean waters (extrapolated from distribution maps in Jefferson et al. 1993).

      Approximately 71 species of marsupials have been recorded from the New Guinea area, of which 60 (84%) can be considered as endemics (not occurring in Australia). Two species of monotremes occur in New Guinea and the long-beaked echidna (Zaglossus) is endemic to the island of New Guinea. Seventy-five bat species belonging to six families are known. Rodents, belonging exclusively to the Muridae, have radiated extensively in the New Guinea region.


      Overall Species Diversity

      Estimates of the total number of species in the world vary tremendously, especially because estimates of insects range from several million to fifty million. Briggs (1994) presented an estimate of 12 300 000 terrestrial species and 200 000 marine species. However, Briggs did not include fungi, which would number about 1 500 000 species (250 000 vascular plants x 6, following Hawksworth), yielding a world total for multicellular species of approximately 14 000 000.

      What is the total number of species in Papua New Guinea? No exact figure can possibly be determined, but a very rough estimate can be derived in two ways. In some of the the better known groups of organisms (for example, flowering plants, and birds), about five percent of the world's species occur in New Guinea. This is remarkable in itself, given that New Guinea has less than one percent of the world's land area. Five percent of the putative world figure of 14 000 000 is 700 000. This is likely on the high side, because Briggs (1994) probably overestimates the number of insect species (Basset et al. 1995). Alternatively, working up from the species, numbers in Table 6.1 yield about 90 000 fungi, 20 000 plants, 5 000 invertebrates, 300 000 insects, and 4 000 vertebrates for something more than 400 000 species. However, this does not take into account the probably large, but totally unknown, numbers of nematodes and mites. A reasonable working estimate for total species numbers in New Guinea is in the range of 300 000 to 500 000 for fungi, plants, and animals. The major problems in this estimate come from the megadiverse groups — fungi, nematodes, insects, and mites. These statistics are for New Guinea as a whole — most of these species (probably at least two-thirds) will occur within Papua New Guinea, but it is impossible to guess how many.

      If approximately 400 000 species occur in New Guinea, how many of these are endemic? Statistics vary tremendously between groups (see Table 6.1), from very low for many marine groups to fairly high for many vertebrates. Vascular plants are likely 60 percent endemic (Johns 1993). However, endemicity will be much lower for Papua New Guinea, because of biotic overlap with Irian Jaya, Australia, and the Solomon Islands.


      Conservation Status of Species

      Table 6.2 lists species that are legally protected or recognised as threatened by Papua New Guinean laws, CITES or the IUCN. Animals are organised by phylum, class, and order, then alphabetically by genus and species. IUCN status data come from Collar et al. (1994) for birds, and Groombridge (1993) for other animals. Important background information on IUCN status can also be found in Thornback and Jenkins (1982, mammals), Seri (1992, mammals), Groombridge (1982, reptiles), and Wells et al. (1983, invertebrates). The birds follow the new IUCN criteria (Mace and Stuart 1994; Collar et al. 1994): Extinct (Ex), Extinct in the wild (Ew), Critically endangered (c), Endangered (En), Vulnerable (V), Data deficient (D), and Near-threatened (N). The other groups follow the traditional IUCN categories (for example, Wells et al. 1983): Extinct (Ex), Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), and Insufficiently known (K). Marsupials listed as potentially vulnerable by Seri (1992; see also Kennedy 1992:vi) were upgraded to vulnerable by Groomsbridge et al. (1993). Species in the IUCN categories classified as indeterminate, insufficiently known, data deficient and near-threatened are not included unless they are also listed by Papua New Guinea or CITES. Latin and English names generally follow Beehler and Finch (1985, birds), Flannery (1990, mammals) and Allen (1991, fishes). Three birds protected by Papua New Guinean law do not actually occur in Papua New Guinea — Goura cristata, Circus melanoleucus and Astrapia nigra (Coates 1985: 118, 304, 1990: 543, 546; Beehler, personal communication, 1994). Chapter 20 reviews national legislation, as well as international conventions and agreements that provide for the management and protection of Papua New Guinea's biological diversity (see also Seri 1992). The listing of entire higher taxa under CITES makes them rather difficult to tabulate — there are probably over 3 000 species of orchids in Papua New Guinea, all recognised under CITES Appendix II, although many remain unknown to science.

      There is a considerable lack of concordance between the listings of Papua New Guinea, CITES, and IUCN — both in species included and their status. Except for the IUCN bird list (Collar et al. 1994), there is little or no current survey data backing up the listings. Except for the birds and marsupials, there has been very little consultation with managers or biologists in Papua New Guinea in formulating the CITES and IUCN lists. All the lists, especially the IUCN one, are very uneven and heavily influenced by information that is available opportunistically. While this can be useful, the taxa that are absent may simply be absent because of lack of information, not lack of conservation problems. Even for mammals, many of the listings are based on data that are over 20 years old (for example, comments in Seri 1992).

      Care should be taken to promote opportunities for conservation through farming or ranching (Parsons 1991). For species that are endangered by habitat loss, the best way often to protect them in Papua New Guinea is to get local people to value the species and therefore the habitat. Thus, it is important that protected species listing and permitting processes are flexible enough to allow for the sale of individuals cultivated by appropriate means. The permit system that has worked for crocodile farming, should be considered for application to other organisms, including birdwing butterflies. Once a species has been listed as ‘protected’ under Papua New Guinean law, there does not seem to be an efficient mechanism for permitting trade (or delisting, if necessary). In summary, lists of threatened species are useful as a starting point, but need continual refinement based on modern field surveys and other assessments by experts.

      According to the IUCN criteria applied in the Birdlife International World Checklist of Birds (Collar et al. 1994), 31 Papua New Guinean endemic birds are listed as threatened, including the Victoria Crowned-pigeon (Goura victoria), Southern Crowned-Pigeon (Goura scheepmakeri), New Guinea Harpy Eagle (Harpyopsis novaeguineae), and three species of Bird of Paradise. These are important species from the conservation standpoint, both as indicator and flagship species. A further 32 endemics are listed as near-threatened while 20 others are listed as data deficient, despite birds being the best known faunal group in Papua New Guinea. The status of monotreme and marsupial mammals in Papua New Guinea was recently reviewed by Seri (1992). However, comprehensive survey data are available for only a very few species of any animals. Even the best known threatened invertebrate species in Papua New Guinea, the Queen Alexandra's Birdwing Butterfly (Troides alexandrae) and the Manus Green Tree Snail (Papustyla pulcherrima) have not been the subject of adequate status surveys (Miller 1993; Cowie 1993).

      Creating a list of rare or threatened plants for Papua New Guinea would be very difficult. Many species are known from only one or a few collections. Their "rarity" relates more to lack of sampling or taxonomic attention than to actual conditions in the field. Some of the actual narrow endemics might be threatened by forestry or mining, but others are totally protected by inhospitable remoteness. The same problem applies to most invertebrates. Some orchid populations have suffered from overcollecting, and sandalwood and ebony may be endangered by overexploitation (J.R. Croft, personal communication, 1994).


      Role of CITES on Papua New Guinea's Biodiversity Endowment

      Papua New Guinea signed the Convention on International Trade in Endangered Species (CITES) in 1976. Detailed data on the volume of export trade of CITES listed organisms by species or derivative product are not available for recent years. General data on the volume of export of crocodiles and butterflies are discussed in Chapter 12. In 1992, only 11.4 percent (8 464 specimens) of the butterflies exported through the Insect Farming and Trading Agency were CITES listed (P.B. Clark, personal communication, 1994). Most exports of CITES listed crocodiles and butterflies are from ranches, so presumably the trade is sustainable, but detailed data are not available. There is anecdotal evidence that illegal exports are taking place in regard to orchids (Kores 1977; Bandisch 1992), butterflies, reptiles, and birds, but no firm statistics are available. In some cases, it is clear that wildlife was exported from New Guinea without proper permits, but it is not clear if it originated in Papua New Guinea or Irian Jaya.


      Ex-Situ Conservation Infrastructure

      Organised ex situ conservation programs in Papua New Guinea are limited to a relatively few species. However, there are significant genetic resources maintained by various institutions (see also Chapter 12). The largest botanical garden is the National Botanical Garden in Lae (also the site of the national herbarium). Smaller botanical gardens exist at the University of Papua New Guinea (Waigani) and elsewhere. A major living collection of orchids is maintained at the Lipizauga Botanical Sanctuary, Mount Gahavisuka (Goroka). Germplasm collections of agricultural cultivars exist at various research sites around the country (for example, Bourke 1985; Levett et al. 1985; Sowei 1992). The Insect Farming and Trading Agency (Bulolo and elsewhere) maintains butterfly colonies and, along with Wau Ecology Institute, Christensen Research Institute, and Unitech, encourages butterfly ranching thoughout the country. Several small collections of captive vertebrates exist, with the largest at Wau Ecology Institute. Crocodile farms exist throughout the country. The Department of Agriculture maintains cultures of microbes for research and identification. Some species are maintained at zoos and botanical gardens outside of Papua New Guinea.

      There are several significant collections of voucher specimens that are crucial to documenting biodiversity within the country (Frodin 1985; Sakulas 1985). The national herbarium is at the Forest Research Institute, Lae, but smaller herbaria are held at Wau Ecology Institute, Christensen Research Institute, University of Papua New Guinea, Unitech, and the Department of Agriculture. Two national insect collections exist at the Department of Agriculture (Konedobu) and at the Forest Research Institute (Lae). Smaller insect collections are maintained at Wau Ecology Institute, Christensen Research Institute, and various agricultural research stations. Vertebrate collections are maintained at the National Museum, Department of Environment and Conservation, Department of Fisheries, Wau Ecology Institute, and Christensen Research Institute.

      Flagship Species

      Flagship species are those which may serve to generate support for conservation action from which a broader component of biodiversity would benefit. Usually flagship species are endemic to the area, and often are threatened in a conservation sense. The uniqueness of the monotreme and marsupial mammals is such that a number of species could serve as flagship species (for example, Zaglossus bruijni, Dendrolagus dorianus, D. goodfellowi, and D. scottae). Another is the Dugong (Dugong dugon).

      The Victoria Crowned-Pigeon (Goura victoria), Southern Crowned-Pigeon (Goura scheepmakeri), Vulturine Parrot (Psittrichas fulgidus), and New Guinea Harpy Eagle (Harpyopsis novaeguineae) could draw attention to the effects of lowland rainforest destruction by logging. Birds of Paradise are recognised worldwide as being almost wholly New Guinean. The Ribbon-Tailed Bird of Paradise (Astrapia mayeri), Black Sicklebill (Epimachus fastuosus), Goldie's Bird of Paradise (Paradisea decora), and Blue Bird of Paradise (Paradisea rudolphi) are all attractive and have international appeal.

      Possible reptile flagship species include the turtle (Carettachelys insculpta); the barred python (Liaisis boa); Boelen’s python (Python boeleni); Varanus salvadori, probably the longest lizard in the world; V. prasinus, the green monitor, with its striking colourations; Corucia zebrata, an unusual skink species with a prehensile tail, which may be the largest skink species in the world; and frogs of the family Microhylidae, which are unusual in being terrestrial breeders with unusual habits.

      Among fish, the Pseudomuqilidae (for example, Pseudomuqil connieae) and Melanotaenidae (for example, Melanotaenia lacustris), which are endemic to the Australia-New Guinea region, are obvious candidates for flagship species. A number of colourful coral reef fish species could attract attention to the extensive reefs of Papua New Guinea.

      Flagship invertebrate species must include the birdwing butterflies (Troides, formerly Ornithoptera) and, in particular, T. alexandrae which is the largest butterfly in the world with a wingspan in excess of 25 centimetres. Other possibilities include the weevil (Gymnopholus lichenifer), which has well-studied mutualistic relationships with a variety of plants and animals, and the Manus green tree snail (Papustyla pulcherimma).


      Population Status of Species at Risk

      Because of the high biodiversity of Papua New Guinea and the logistical difficulties of field surveys, detailed information on population sizes, over time, is available for only a few species. Crocodiles are perhaps the best known example in Papua New Guinea (Goldstein 1991). Once threatened with extinction by hunting for the skin trade, they are now managed in the wild and also in captivity. Seri (1992) provided recent data on selected mammals, and the Conservation Needs Assessment provides additional examples and references.

      Saltwater and freshwater crocodile populations declined during the late 1950s and 1960s, through indiscriminate hunting. In 1969, the Crocodile Trade (Protection) Act (Chapter 213) was implemented which, to protect breeders, placed a ban on trade in skins greater than 51 centimetres belly width. This halted any further decline in crocodile numbers, as indicated by a steady export level during the 1970s. The Act also allows for the control of the crocodile industry on a systematic basis. Regulations under this Act control crocodile farming and the purchase and export of skins. In 1981, a ban was placed on trade in skins smaller than 18 centimetres. This ban was established because Papua New Guinea was in a position to ranch crocodiles on a large scale. By 1984, although the number of skins exported was the same as in previous years, 30 percent were from ranched animals and consequently were of higher grade and greater size. Both species of crocodile are listed in Appendix 2 of CITES which means that they are regarded as vulnerable, but trading is allowed to continue. For many years, Papua New Guinea has been the only country allowed by CITES to trade in C. porosus. In 1982, extensive monitoring of both species commenced especially in the Ambunti District of East Sepik Province.


      Invasive and Introduced Species

      Non-indigenous species represent a major economic threat. Other than a few of the most important agricultural pests, data do not exist to address these issues in any detail. The recent report prepared for the United States (OTA 1993) provides ample examples of the problems and solutions.


      The aquatic plants — water hyacinth (Eichhornia crassipes) and salvinia (Salvinia molesta) — have been major problems in the Sepik River and elsewhere, illustrating the damage that invasive non-indigenous species can cause, and the potential of biological control (see Box 6.1). Introduced freshwater fish provide another case study of positive and negative environmental aspects of introductions. The recent establishment of the apple snail in the Lae area (Laup 1991) also underscores the importance of pest exclusion and detection programs (Kanawi et al. 1994).

      Other species have been introduced for biological control of agricultural pests and weeds. The early history of biological control introductions in Papua New Guinea is reviewed by Wilson (1960). Modern biological control programs against insects, snails, and weeds in Papua New Guinea are reviewed by Waterhouse and Norris (1987). While biological control programs can be very beneficial, poorly implemented programs can have serious impacts on non-target organisms (Howarth 1991).

      Herington (1977) reviewed animal species that are known to have been intentionally introduced to Papua New Guinea for agriculture, hunting, and biological control. Some data on insect introductions for biological control was compiled by Wilson (1960) and Waterhouse and Norris (1987), but documentation for non-indigenous insects remains very poor. There has not been a thorough review of non-indigenous plants in Papua New Guinea, although many are included in the review of weeds by Henty and Pritchard (1975). Box 6.2 summarises non-indigenous animals in Papua New Guinea, but there is no comprehensive program for monitoring the presence of non-indigenous species. Therefore, general statistics on distribution, population sizes and rates of change, environmental impacts, as well as the effectiveness and costs of control measures, are not available.


      Box 6.1: Invasive Aquatic Weeds: Salvinia (Salvinia molesta) and Water Hyacinth (Eichhornia crassipes).

      Two aquatic weeds, Salvinia molesta and Eichhornia crassipes, are now widespread in the low-lying wetlands of Papua New Guinea. Salvinia molesta was first recorded in Papua New Guinea in 1977 at Wau, and on the Sepik River where it was probably introduced in 1971-72 (Mitchell 1978-1979, 1979). By 1979, salvinia covered 80 square kilometres and the physical impact of the weed was reflected in the decline in fish catches, crocodile hunting, and sago gathering, and also in the disruption to the lives of Sepik villagers. People in a number of villages were unable to reach markets to sell produce and children were prevented from attending school (Mitchell et al. 1980; Coates 1982, 1987a). A control program was instituted in 1979 (Thomas 1979), and physical, chemical, and biological control methods were tested (Thomas 1985). Biological control using the South American weevil Cyrtobagous salviniae was spectacularly successful (Thomas 1985; Laup 1985; Thomas and Room 1986). The initial development of the weevil population introduced in 1982 was nitrogen limited (Room and Thomas 1985, 1986a, 1986b), but once established on plants enriched with fertiliser, the populations became self-sustaining. By June 1985, the weevil had destroyed an estimated two million tonnes of weed which had covered 250 square kilometres. The local people have now resumed their former lifestyles (Thomas and Room,1986). Eichhornia crassipes was first recorded in 1962 when it was found near Bulolo in old gold mining dredge ponds. Despite warnings (Mitchell 1978-1979; Osborne and Leach 1984), it has become widespread throughout lowland areas of Morobe, Madang, and East Sepik Provinces, and has recently spread to the wetlands west of Port Moresby, in Central Province. It has also been reported from Manus, New Ireland, North Solomons, West New Britain, Eastern Highlands and Western Provinces (Laup 1986a). Attempts at biological control using the weevil Neochetina eichhorniae have produced some promising results. This weed now covers the majority of Waigani Lake and has severely disrupted the once productive fishery based on Oreochromis mossambica. Pistia stratiotes is widespread throughout the lowlands of Papua New Guinea, and although it is a weed in other parts of the world, it has not reached weed proportions here (Laup 1986b). The same is true of Hydrilla verticillata which has only been recorded near Wau, Madang, Port Moresby, and at Lake Kutubu (Leach and Osborne 1985).




      Introduced Freshwater Fish in Papua New Guinea


      Twenty-one species of freshwater fish representing 19 genera, 11 families, and all six continents have been introduced into Papua New Guinea for various reasons (West 1973; West and Glucksman 1976; Glucksman et al. 1976). The reasons include sport, aquaculture, ecological manipulation, control of pests, ornamentation, and improvement of subsistence welfare. Most introductions have been unsuccessful or were never released into the wild. Nine to eleven are thought to be established in Papua New Guinea (Allen 1991; Osborne 1993). Of the successful introductions, most have had a negligible impact as either food fishes or in the control of mosquitoes (Allen 1991). Oreochromis mossambica is an exception as it now provides the major subsistence source of protein to villagers living along the Sepik River, and it was, prior to infestation of the lake with water hyacinth, the basis of a thriving commercial fishery on Waigani Lake near Port Moresby.

      Allen (1991) regards most of the earlier introductions as having had a negative impact through competition for space and limited food resources, or by feeding on the native species. Even the popular Oreochromis mossambica has adversely affected the environment, creating turbid conditions in formerly clean lakes and overcrowding the indigenous fauna because of its prolific breeding. On the positive side, the number of established introductions is relatively few, and Allen (1991) states that the Fly River appears to be free of introductions. Coates (in litt.), however, indicates that common carp (Cyprinus carpio) occur in the Fly River and, furthermore, that he recorded the climbing perch (Anabas testudineus) from the Fly River in 1985.


      Four major negative effects may result from fish introductions in Papua New Guinea:

      • predation and competition — introduced species can lead to a reduction in native fish diversity either through competition for a common resource or through increased predation pressure;
      • introduction of parasitic or pathenogenic organisms, not previously found in the region, to native fish species;
      • stunting, particularly in cichlids. (Introduced fish species may undergo explosive population expansion resulting in precocious maturity and reproduction. This, in turn, leads to a habitat filled with large numbers of small fish which encroach on the living space of indigenous species); and
      • environmental effects, particularly in common carp and tilapine cichlids.


      The introduced species may alter the habitat to such an extent through their activities that native species are unable to survive.

      Two species of trout, Salmo trutta and Oncorhynchus mykiss, have been introduced into the highland regions of Papua New Guinea, with an introduction of Salvelinus fontinalis having been unsuccessfully attempted. Only Oncorhychus mykiss seems to have become established and widespread. Their potential impact, well-recognised in Australia and New Zealand, is as predators and competitors with native fish species, but as Allen (1991) notes, as most introductions have taken place above 2 000 metres in Papua New Guinea where there are few native fish species, their impact on other fish is likely to be minimal, but they will have an impact on other species and on general stream ecology.

      The Cyprinidae dominate the ichthyofauna of other tropical regions and Cyprinus carpio has been widely introduced throughout the world. This species is well-established in the Sepik system in Papua New Guinea and is also found in certain Southern Province rivers. The species is known to increase water turbidity, and directly and indirectly destroy rooted vegetation. This is often accompanied by a decline in native fish populations and spread and build-up of carp populations (see Cadwallader 1978; Taylor, Coutney and McCann 1984).

      The family Poeciliidae are live-bearing ovoviviporous fishes originating in Central and Southern America. Two species —Poecilia reticulata and Xiphiphorus helleri — are somewhat localised within the country, whereas Gambusia affinis introduced for its supposed ability to control mosquitoes, is more widely distributed. All three species have been recognised as having a detrimental effect on small surface feeding native fish species belonging to the genera Melanotaenia, Pseudomugil, Craeterocephalus and Retropinna in Queensland. Members of the same four genera also occur widely in Papua New Guinea (Arthington et al. 1981). Introduction of two or more poecilid species into the same water body would seem to have a synergystic effect upon the disappearance of small surface dwelling native species (McKay 1984). There is little evidence to support the view that poecilids are more capable of controlling mosquito populations than native surface dwelling species. The possibility that Gambusia may be associated with introduced parasitic species in Papua New Guinea is currently under investigation (see also Mungkaje 1986).

      Introduction of the Mozambique tilapia, Oreochromis mossambicus, into Papua New Guinea is relatively well-documented (Glucksman et al. 1976) and appears to have been based upon 250 fish brought to Port Moresby in 1954 from either Malaya or Java. Within the next five or so years the species became established in rivers in the vicinity. At present this species is widely spread throughout the country with particular importance in the Sepik system (Coates 1985). In that river system, it has been shown to constitute between 30 to 50 percent of the weight of gill net catches. Recent studies in rivers in the Port Moresby region (Kawei and Menzies, unpublished; Hyslop, Mala and Soare 1994) have shown that this species in conjunction with the Gourami, Trichogaster pectoralis, make up the bulk of catches. In the lower reaches of the Angabanga River, Oreochromis mossambicus constituted between six to 90 percent of the fish catch by weight (Hyslop et al. 1994). The Mozambique tilapia was also noted by Berra et al. (1975) as the species collected from most localities in the Laloki River.

      Nelson and Eldridge (1991) and Maciolek (1984) document introductions of Mozambique tilapia throughout the Pacific Basin and note its propensity to dominate native fish faunas and its poor acceptance as a food fish in various places including Kiribati, the Cook Islands, and Guam. In Fiji, the species is considered as a pest because of its tendency to dominate ichthyofaunas to the exclusion of native species and the trend towards stunted populations. Allen (1991) associates the species with increasing turbidity levels because of their feeding method, and domination of bottom space by nest building males. The ability of this species to switch its dietary habits according to its habitat means that it is very likely to compete with native fish species in a range of locations. Moreover, the response of the tilapia population to fluctuations in environmental conditions is to mature early and reproduce out of season resulting in a rapid build up of numbers which, because of limitations of food/space never attain significant size. The implications of this are twofold — tilapia rarely reach marketable size in sufficient numbers, and native species may be edged out because of pressure for food and/or space.

      The family Belontiidae contains two species of gourami — Trichogaster pectoralis and T. trichopterus. The former is widely distributed, particularly in Central and Gulf Provinces. Two studies (Kawei and Menzies, unpublished; Hyslop et al. 1994) have shown this species to be extremely abundant in Southern Highlands Province rivers. Hyslop et al. (1994) quantified the contribution of T. pectoralis to the catch from the lower Angabanga River at between three and 82.5 percent by weight. In many places, especially in areas of swamp, this species in conjunction with O. mossambicus constituted the entire catch. Furthermore most individuals of both species were of small size (five to six grams), perhaps indicative of stunting.

      The family Anabantidae is characterised by possession of an accessory air breathing organ, which enables species to tolerate low oxygenation and periods out of water. Anabas testudensis, the Indian climbing perch, has been introduced into Papua New Guinea from Irian Jaya and the spread of this species has been the subject of various press releases from the Department of Fisheries and Marine Resources. The earliest record of the species is from 1976 in the Morehead River. Present observations indicate that it is widespread throughout the Fly/Ok Tedi system and also the lower Strickland River. It seems to be particularly abundant in shallow weedy habitats (Hyslop et al., unpublished). Anecdotal evidence indicates that Anabas may cause the death of predatory fish species, file snakes and even crocodiles by becoming lodged in the pharynx of these species by means of its spines on the opercula and fins.

      Several features are shared by introduced fish species in Papua New Guinea which increases their chances of survival and proliferation over those of indigenous species:

      • the lack of radiation/specialisation of the native ichthyofauna ensures that introduced species which are often "generalists" can find a niche to establish themselves in the community;
      • superior environmental tolerance: most successful introduced fish species are able to tolerate a wide range of conditions, such as low oxygen levels, via adaptations including accessory air breathing organs and tolerance of low levels of dissolved oxygen;
      • enhanced reproductive capacity allowing rapid population build up: the Poecilidae are all live bearers (ovoviviporus) ensuring a high survival rate of offspring as young are produced at an advanced stage. High fecundity occurs. For example, a five kilogram carp may produce one million oocytes (Merrick and Schmida 1984). Early non-seasonal maturity and mouth brooding of eggs in tilapia and gouramis ensures good survival of offspring and fast increase in population size;
      • dispersal abilities: carp, Mozambique tilapia, climbing perch and gouramis can survive out of water for some time if kept moist, allowing them to survive transportation between water bodies by humans. Others have high tolerance of salinity enabling spread between water bodies;
      • ecological adaptability: opportunistic feeding behaviour and omnivory are typical of most introduced freshwater fish species in Papua New Guinea; and
      • aggressive behaviour: gambusia and male oreochromis possibly harass other fish species.

      The freshwater fish fauna of New Guinea is particularly susceptible to the effects of introduced fish species because of the lack of specialisation. Nine species of introduced fish are widely established in Papua New Guinea and, while there is limited direct evidence to substantiate the case, these species have resulted and are likely to result in loss of biodiversity in the freshwater ichthyofauna. Other prominent examples of deleterious effects from non-indigenous species in Papua New Guinea include the cane toad (Bufo marinus), the African giant snail (Achatina fulica), water hyacinth (Eichhornia crassipes), and salvinia (Salvinia molesta).


      Box 6.2: Animals Introduced to Papua New Guinea (based on Herington 1977; Eldredge 1994).

      Although general statistics on non-indigenous animals in Papua New Guinea are not readily available, there are data on the various species of molluscs, insects, fish, amphibians, birds, and mammals that have been introduced. The following summary is based on Herington (1977) and Eldredge (1994).


      MOLLUSCS (Eldredge 1994; Mead 1961, 1979; Hadfield et al. 1993)

      • Giant African snail (Achatina fulica), introduced during Japanese times; well-established.
      • Predatory snail (Gonaxis quadrilateralis), New Britain, New Ireland; status unknown.
      • Predatory snail (Gonaxis kibweziensis), status unknown.
      • Predatory snail (Euglandia rosea), introduced in 1958.
      • Freshwater snail (reported as Pomacea lineata, but probably P. canaliculata).
      • Unidentified pearl oysters (probably Pinctada maxima), 7 000 shells from Kuri Bay, Western Austalia to Port Moresby in 1977.

      INSECTS (See Wilson 1960; Herington 1977; Waterhouse and Norris 1987 for data on biological control introductions). Only more recent introductions are listed here. Many additional introductions have probably not been recorded in the formal literature.

      • Parasitic fly (Sturmiopsis inferens), introduced from India in 1981 to control sugarcane borer (Sesamia grisescens); probably not established; Ismay and Dori 1985.
      • Parasitic flies (Trichopoda pennipes and Trichopoda pilipes), introduced from Hawaii in 1977 and 1980-1981 to control green stink bug (Nezara viridula); probably not established; Ismay and Dori 1985.
      • Parasitic wasp (Apanteles flavipes), introduced from India in 1981 to control sugarcane borer (Chilo terrenellus); Ismay and Dori 1985.
      • Parasitic wasp (Paraceraptrocerus nyascius), introduced from Queensland in 1981 to control white wax scale (Ceroplastes destructor); Ismay and Dori 1985.
      • Parasitic wasps (Opius importatus and O. phaseoli), introduced from Hawaii in 1980-1981 to control beanfly (Ophiomyia phaseoli); Ismay and Dori 1985.
      • Parasitic wasp (Trissolcus basalis), introduced from Western Australia in 1978 to control green vegetable bug (Nezara viridula); Ismay and Dori 1985.
      • Psyllid (Heteropsylla spinulosa), introduced in 1992 to control the weed Mimosa invisa; Kuniata 1993.

      FISH (Allen 1991)

      Freshwater fishes—26 species introduced (15 established); most notably — two species of tilapia, common carp, mosquito fish.

      • Blenny (Petroscirtes breviceps) found in continuous flow bilge water at Port Moresby; from Northwestern Australia.

      AMPHIBIANS (Allison 1993)

      • Marine toad (Bufo marinus) to Papua New Guinea in 1937 from Hawaii and Australia (Zug et al. 1975).

      BIRDS [also imported, ducks, geese, turkeys, pigeons, and cage birds] (Lever 1987; Coates 1990)

      • Red jungle fowl (Gallus gallus).
      • House sparrow (Passer domesticus, unsuccessful, 1976).
      • Common starling (Sturnus vulgaris, considered vagrants from Australia).
      • Common myna (Acridotheres tristis, Bougainville).

      MAMMALS (Lever 1985; Flannery 1990)

      • Domestic dog (Canis familiaris).
      • Domestic pig (Sus scrofa).
      • Domestic cat (Felis catus).
      • Timor deer (Cervus timoriensis).
      • Axis deer (Axis axis, restricted).
      • Fallow deer (Dama dama, may be extinct).
      • Water buffalo (Bubalus bubalis).
      • Domestic cattle (Bos taurus).
      • Domestic Horse (Equus caballus).
      • Domestic goat (Capra hircus).
      • Polynesian rat (Rattus exulans).
      • Black rat (Rattus rattus, confined to lowlands).
      • Brown rat (Rattus norvegicus, from major seaports).
      • Ricefield rat (Rattus argentiventer, rare, few records).
      • Himalayan rat (Rattus nitidus, from Vogelkop).
      • House mouse (Mus musculus, serious pest).

      Available on the internet on April 1999