12. New Caledonia – (Rennell I.) – New Guinea
Notes on the main text
Many groups show affinities between New Caledonia/Vanuatu and Papua New Guinea, especially northern PNG and the Bismarck Archipelago, but have no records from the Solomon Is. In some cases this will be due to lack of adequate collecting, but often there does seem to be a direct New Caledonia – New Guinea connection (sometimes with a Solomon Islands vicariant filling the gap).
The New Caledonia – New Guinea track is documented in many scattered publications and in an earlier track analysis Morrone (2002) proposed New Caledonia – New Guinea as the ‘Neoguinean region’, one of just 12 for the world.
The disjunction between the Vogelkop and the Bismarck Archipelago that occurs in Phyllanthus bourgeosii (Fig. 6) is also seen in forms of Osmoxylon (Aral.) and is related to the important connections between the Moluccas and New Britain, and between Waigeo and New Ireland (Heads, 2001, 2003: Fig. 73). At its three disjunct localities P. bourgeoisii grows on the edge of mountain torrents as a rheophyte. Webster and Airy Shaw (1971) described the distribution as ‘curious’. A New Caledonia and New Guinea disjunction also occurs in Phyllanthus pancherianus.
Sampling of Zosterops species in the Moyle et al. (2009) study was not complete, but it is clear that Z. rennellianus is not directly allied with nearby species of the southern Solomon Is. or Vanuatu. In Phillimore’s (2006) sample of New Zealand, Lord Howe, Norfolk and Vanuatu populations, Z. rennellianus was linked with Z. tenuirostris of Norfolk I.
The New Caledonia – Santa Cruz connection in the Pteropus pselaphon group (Fig. 7) also occurs in plants (Austrobuxus sp.; Picrodendr.; van Balgooy & Franken, 1984). (Politically the Santa Cruz group is in the Solomon Is., biogeographically it is allied closely with Vanuatu, Fiji and, as here, New Caledonia). A Solomon Is. – Moluccas connection similar to that of the chrysoproctus group occurs in Odonata (Rhinocypha libereta of Guadalcanal and its allies in the Moluccas; Polhemus et al., 2008) and in podocarps (Dacrydium magnum and Podocarpus spathoides; Heads, 2003: Figs. 80, 87).
In fungi, Mycena irritans is in New Caledonia and New Guinea (Horak, 1983).
In lichens, Pseudocyphellaria clathrata is known from Norfolk I., New Caledonia, New Guinea, western Malesia, Africa and South America (Galloway, 1994).
The liverworts Plagiochilon theriotanus and P. braunianus are in New Caledonia, New Guinea, and further west in Malesia etc. (Schuster, 1979). Frullania heteromorpha, Jungermannia hirticalyx and Telaranea kogiana are all in New Caledonia and eastern New Guinea. Cololejeunea wightii is in New Caledonia, New Guinea, and Penang (Peninsular Malaysia). (The disjunction between New Guinea and Peninsular Malaysia seen in the last species is standard, cf. Heads, 2003). Marchantia lecardiana is in New Caledonia and New Britain, M. multiloba is in New Caledonia, New Guinea and the Philippines. Herbertus leratii is a rare endemic from southern Grande Terre, compared with a species of New Guinea/Indonesia by So (2003). Porella maxima is in New Caledonia and Papua New Guinea (So, 2002). Porella acutifolia has a similar connection: in the South Pacific and Australasia there are two varieties of this species, var. acutifolia in eastern and south-east Asia, New Guinea and Hawaii, and var. linguifolia of New Caledonia (So, 2002). Plagiochila inflata is in New Guinea, New Caledonia and Norfolk (So, 2000).
Examples in mosses include Bescherellia elegantissima of New Caledonia, Loyalty Is., and the PNG Highlands (Sastre-De Jesús, 1987). Miller et al. (1978) recorded Calymperes tahitense var. denticulatum, Fissidens laxiretis, and Megalostylium papuanum all in New Caledonia and New Guinea. Hymenodon tenellus (also Goodenough I.) and Taxithelium nitidulum (also d’Entrecasteaux Is.) have similar ranges and Rhizogonium novaehollandiae var. minus is recorded from I. des Pins and New Guinea. Trematodon novae-caledoniae is in New Caledonia and the Bismarck Archipelago, T. hannoverae is in New Caledonia and New Hanover (off northern New Ireland). Spiridens vieillardii is in Lord Howe, New Caledonia and New Guinea. Leskeodon acuminatus is in New Caledonia, New Guinea, Java and Borneo.
In ferns, Ctenopteris subsecundo-dissecta is restricted to New Guinea and New Caledonia. Schizaea wagneri of New Guinea and Malesia is closest to S. intermedia of New Caledonia (Selling, 1946). S. inopinata of Malesia east as far as western New Guinea and Micronesia is closest to S. laevigata of New Caledonia and S. melanesica of New Caledonia, Fiji and Tonga. Coryphopteris fasciculata (Thelypterid.) is in New Caledonia, New Guinea, and Sulawesi (Holttum, 1977).
The best-known plant group restricted to New Guinea and New Caledonia is Nothofagus subgen. Brassospora (Nothofag.). It comprises 19 species of large trees in New Caledonia and New Guinea and is the only Nothofagus subgenus present in these countries. Fossils of the group are known from New Zealand, Australia, Antarctica and South America. The extant range of Brassospora may represent the original centre of diversity for the group which has become extinct in secondary outliers (Heads, 2006). The other subgenera have their respective cores in different localities, and so probably all formed as vicariants. This explains the notable absence of the New Zealand and Australian subgenera from the mountains of New Guinea and New Caledonia.
Direct New Caledonia – New Guinea connections, not involving the Solomon Is. or Vanuatu, also occur in the following taxa.
In monocots, Sciaphila densiflora (Triurid.) occurs in New Caledonia, New Guinea (including Milne Bay islands), Malesia and Sri Lanka. S. corallophyton is in New Caledonia, north-eastern New Guinea, and eastern Caroline Is. (Ponape) (van Meerendonk, 1984).
The sea grass Enhalus (Hydrocharit.) is in New Caledonia, New Guinea and Torres Strait, and further west (den Hartog, 1970).
Rhuacophila (previously treated in Dianella) (Hemerocallid.) is in Malesia, New Guinea, New Caledonia and Fiji (Clifford, Henderson & Conran, 1998).
In dicotyledons, Nepenthes (Nepenth.) has closely related forms in New Caledonia (N. vieillardii) and western New Guinea (N. lamii; Cheek & Jebb, 2001).
Alyxia (Apocyn.) is widespread from India to south-eastern Polynesia but has its major centres of diversity in New Guinea and New Caledonia (Avé, 1984). Ser. Laxiflorae comprises A. leucogyne of New Caledonia and species of New Guinea, Sulawesi, and the Philippines (Markgraf, 1977).
Dicarpellum (Hippocrat.) of New Caledonia is closest to Brassiantha of New Guinea and Sarawakodendron of Borneo (Aubréville et al., 1967-).
Coode (1987) proposed that Dubouzetia (Elaeocarp.) of New Caledonia, New Guinea, and the Northern Territory forms a group with Peripentadenia of north-eastern Queensland and Crinodendron of temperate South America, with all three showing clear vicariance. Dubouzetia comprises three sections (Coode, 1987):
sect. Spongosperma: New Guinea,
sect. Dubouzetia: New Caledonia,
sect. Oligovula: New Guinea/Moluccas, the Northern Territory (Australia), and New Caledonia.
For Coode this represents a ‘peculiar picture’ as it is not compatible with ‘island-hopping’ along the Melanesian arc – ‘The basic problem seems to be that D. elegans (sect. Oligovula) is found on both New Caledonia and the New Guinea mainland (but not the New Hebrides, Solomon Is. or Bismarck Archipelago [or Australia])’. Coode thought that ‘perhaps it is a New Guinea species which reached New Caledonia relatively late…’, but the distribution of the sections instead indicates that the New Guinea – New Caledonia sector is fundamental in the genus as a whole and not a secondary development.
Hunga (Chrysobalan.) is restricted to New Caledonia (five species) and PNG (three species) (Prance, 1979). In PNG the genus occurs in the Papuan Peninsula (the ‘tail’ of the country) on and east of a line: Bakaia/Sogeri, and also in the Louisiade Islands (Milne Bay) with H. longifolia, endemic to Misima I. Prance regarded H. longifolia as most closely related to H. lifouana of New Caledonia, not to the other PNG species. This pattern is repeated in Loranthaceae: an ‘extreme variant’ of the south-west Pacific Amyema artensis occurs on Rossel I. (Louisiade Is.) and New Caledonia (Barlow, 1992). A. scandens is only known from New Guinea and New Caledonia. Barlow suggested that both A. artensis and A. scandens reached New Caledonia by dispersal from New Guinea, but this does not explain the absence from the Solomon Is., Vanuatu, and Australia.
The closest relative of Hooglandia (Cunon.) from New Caledonia (north-eastern Grande Terre) ‘may prove to be Aistopetalum’, a New Guinea genus (McPherson & Lowry, 2004).
Sloanea sect. Antholoma (Elaeocarp.) (van Balgooy, 1971) is restricted to New Caledonia and New Guinea. Thorne (1965) cited Pelma (Orchid.) as a New Guinea – New Caledonia group. In Aubréville et al. (1967-) Pelma is synonymised under Bulbophyllum, but its only New Caledonian species, B. (P.) neocaledonicum, is described as related to ‘Indonesian’ species and so the New Caledonia – New Guinea affinity may remain.
In Myrtaceae, Xanthomyrtus occurs in New Caledonia, southern New Ireland, New Guinea (most species), Borneo and the Philippines. It may be significant that Scott (1979a) keyed out the only New Caledonian species, X. hienghenensis, next to X. schlechteri, widespread through New Guinea to Fergusson I. and New Ireland. Uromyrtus artensis of New Caledonia may be closest to U. novoguineensis of the Cyclops Mts. (above Jayapura, north central New Guinea) (Scott, 1979b).
In groups with New Caledonia – Bismarck Archipelago connections, Heliconia indica var. austrocaledonica (Helicon.), of southern New Caledonia and western Vanuatu, keys out with H. i. micholitzii of New Ireland and Manus, rather than with H. i. denisiana of the Solomon Is. or H. i. rubricarpa of north-eastern PNG and New Britain (Kress, 1990). Meiogyne glabra (Annon.) of central New Britain is keyed out next to M. ‘spec. 1’ of New Caledonia by van Heusden (1994). (Glabrous forms found around Suva, Fiji, may also belong here; pers. obs. 2005). Arthrophyllum (Aral.) occurs in New Caledonia, the Bismarck Archipelago, New Guinea, and through Malesia to south-east Asia (Philipson, 1979; van Balgooy, 1993a; Frodin, 2001). Dolichandrone spathacea (Bignon.) has a similar distribution but extends to Sri Lanka and India (van Steenis, 1963).
Many species of coral are in New Caledonia and New Guinea (and other areas) but do not occur in Australia (Veron, 2000), despite the presence of suitable habitat on the Great Barrier Reef. The gastropod Xenophora granulosa is in New Caledonia, New Britain, the Philippines and Mauritius (Ponder, 1983).
In lobsters, Ibacus brevipes is in New Caledonia, south-western New Guinea and the Philippines, Acanthacaris tenuirama is in New Caledonia, south-western New Guinea and further west (Holthuis, 1991).
In spiders, Desis maxillosa (Desidae) is in New Caledonia and New Guinea. Argyrodes amboinensis (Theridiidae) is in New Caledonia, New Guinea, the Moluccas, and Sulawesi. Cyclosa albopunctata (Araneidae) is in New Caledonia, New Guinea, and Africa.
In Diptera, Elephantomyia subgen. Elephantomyia (Tipulidae) is known from New Caledonia and western New Guinea. Blepharipa sugens (Tachinidae) is in New Caledonia, PNG, and Malesia. Allactoneura (Mycetophilidae) is in New Caledonia, western New Guinea, Sulawesi, Borneo and further west. In Tipulidae Limonia (Thripticomyia) subsaltens is in New Caledonia, Fiji, Samoa and PNG, and Gonomyia subgen. Gonomyia is widespread in the tropics but in Australasia/Oceania only at: New Caledonia, Fiji, eastern and western New Guinea.
In beetles, the eucnemid Mesogenus austrocaledonicus of New Caledonia is sister to M. cavifrons of New Guinea (Muona, 1991) and in chrysomelids, Stethotes bertiae of New Caledonia seems closest to S. minuta of New Guinea (Jolivet et al., 2007a). Ottistira (Curculionidae) is in New Caledonia, New Guinea, and Indonesia (Kuschel, 2008).
In butterflies, the New Caledonian Papilio montrouzieri and Polyura clitarchus have their closest relatives in the Papuan region (Holloway & Peters, 1976). In other Lepidoptera, Holloway (1979) cited six New Caledonian endemics that have their closest relatives in New Guinea: Epiplema lateritica (Uraniidae), Anisozyga caledonica, Chloroclystis macroaedeagus (Geometridae), Agrius fasciatus (Sphingidae), Nycteola canoides and Veia pectinata (Noctuidae). Lasioceros aroa (Notodontidae) is in New Caledonia and New Guinea, with a distinct subspecies in Fiji. Polyacme dentata (Geometridae) is on New Caledonia/ Loyalty Is., and its two closest related species are on New Guinea.
Leatherback turtles (Dermochelys coriacea) are widespread in the world’s seas. Two animals tagged with satellite tracking devices were found to swim between New Caledonia and Morobe, PNG (Anon., 2002).
In birds, Vuilleumier & Gochfeld (1976) suggested that eight New Caledonian endemic species (in Accipiter, Ducula, Charmosyna, Coracina, Aplonis, Corvus, Zosterops and Erythrura) had a ‘presumed origin’ in New Guinea, reflecting putative affinities. In at least some of these the relationships are doubtful. For example, Corvus (= Physocorax) moneduloides may have no direct affinity with birds of Oceania, Australia or New Guinea (Berlioz, 1962) and, in particular, its extraordinary tool-making abilities set it apart (Weir, Kenward & Kacelnik, 2004).
The moth Pareromene cheesmani (Pyralidae) of New Caledonia and northern and southern Vanuatu has its closest relatives in New Zealand (three species) and Rennell I. (P. fusca) (Gaskin, 1974). This group follows the trend of the Norfolk – New Caledonia – Rennell – Papua ridge/trench system and biogeographic connection cited in the main text. Other Rennell I. connections are shown in the following passerines.
Gerygone flavolateralis (Acanthizidae) is in New Caledonia (one subspecies), the Loyalty Is. (three subspecies), northern Vanuatu (one subspecies), and Rennell I. (one subspecies) (mapped in Mayr & Diamond, 2001).
Myiagra caledonica (Monarchidae) is in New Caledonia and the Loyalty Is. (two subspecies), southern Vanuatu, northern and central Vanuatu, and Rennell I. It is related to species of the south-eastern Solomon Is. (San Cristobal; M. cervinicauda), Santa Cruz – Fiji (M. vanikorensis), and the Solomon Is. (M. ferrocyanea).
13. New Zealand – New Caledonia – New Guinea
Notes on the main text
Mearnsia (Myrt.) is sister to Carpolepis of New Caledonia, and Metrosideros s. str. of New Caledonia, New Zealand and the central Pacific islands (east and south-eastern Polynesia to Hawaii, but not in New Britain, New Guinea or the Philippines). The Pacific Metrosideros/Mearnsia clade with its internal vicariance is sister to another vicariant, Tepualia of South America/South Africa, and this whole group (tribe Metrosidereae) is in turn sister to tribe Backhousieae of eastern Australia, yet another vicariant (Wilson et al., 2005). Central and western Australia are occupied by other tribes.
Wilson (1996) compared the distribution of Mearnsia with that of Xanthomyrtus (Myrt.), cited above. In Metrosideros s. str. there is clear-cut vicariance between two main groups (Wright et al., 2000), with M. cherrieri and its allies in New Caledonia, Fiji, the Solomon Is., New Ireland, and northwest in the Bonin Is., while M. excelsa and its allies occur in New Zealand, Lord Howe I., the Kermadec Is., Vanuatu, Fiji and east through Polynesia. Wright et al. noted the ‘clear geographic separation’, but rather than accepting Wilson’s (1996) vicariance explanation they regarded the non-New Caledonian species of the first group as ‘apparent vagrants’ from New Caledonia and the species of the second group as dispersed from New Zealand. Nevertheless, this does not account for the most fundamental aspect of the pattern – the striking allopatry of the two Metrosideros groups, nor does it explain the presence of Mearnsia but not Metrosideros in New Guinea and the Philippines, or the absence of both from Australia.
Wilson (1996) noted that the presence of Metrosideros on the Bonin Is. is ‘remarkable’ and that ‘standard long-distance dispersal cannot account for its occurrence’ there. He also referred to similar distributions in other taxa such as the palm Clinostigma (ranging from the Bonin and Caroline Is. to Fiji). Wright et al. (2000) found that the Bonin Islands Metrosideros is closest to a Fijian species, confirming a pattern similar to that of Clinostigma.
Fungi showing the track (from Horak, 1983) include Marasmius cylindraceo-campanulatus: New Zealand, New Caledonia, PNG, Java; and Cuphocybe: New Zealand, New Caledonia, New Guinea. This track also appears in the range of Rozites: New Zealand, New Caledonia, New Guinea, Chile, southern Argentina (12 species), and two species in the Northern Hemisphere; and Entoloma haastii: New Zealand, New Caledonia, PNG, Chile, southern Argentina, Europe.
In lichens, Megalospora sulphureorufa of Lord Howe and New Caledonia is related to M. bartlettii of New Zealand (North Cape) and M. granulans of New Guinea (Fig. 5; Sipman, 1983). The only Lord Howe specimen is placed in the same species as the New Caledonian one, but shows similarities with the New Guinea species. Pseudocyphellaria poculifera is in northern New Zealand, Lord Howe I., Norfolk I., New Caledonia, Fiji, New Guinea (Morobe), and also Peninsular Malaysia and Uganda (Galloway, 1994).
The liverwort group Acromastigum sect. Acromastigum is in New Zealand, New Caledonia, and Hawaii, while the related A. subgen. Triandrophyllopsis is endemic to Mt. Wilhelm, PNG (Schuster, 1969).
The moss group Hypnodendron sect. Sciadocladus occurs in New Zealand, New Caledonia and the Solomon Islands (not in New Guinea) (Tangney, 1990).
The fern ally Tmesipteris (Australia – Pacific) is represented in PNG by a single species on Bougainville I. (northern Solomon Is.), Goodenough I. (D’Entrecasteaux Is.) and Isuarava (mainland PNG north-east of Mt. Albert Edward). It appears to be related to T. sigmatifolia of New Zealand and New Caledonia (Johns & Bellamy, 1981).
Hypericum gramineum shows a ‘morphological and geographical trend’ along the line: New Zealand, New Caledonia, New Guinea, Taiwan (Robson, 1981).
Lagenophora lanata (Compositae) is in New Zealand (north of Auckland), New Caledonia, New Guinea (widespread through the mountains), Malesia and India (Koster, 1966).
Westermann (2000) described marine faunal realms of the Mesozoic (based largely on invertebrates) and for the Indo-Pacific Realm proposed as ‘an acceptable minimum range’: New Zealand, New Caledonia, New Guinea, and the Himalayas.
The landsnail Papulaoma monticola (Punctidae) and its immediate relatives occur in New Zealand, New Caledonia and New Guinea (F.M. Climo, pers. comm., 20 – v – 2007).
The spider Orsinome (Tetragnathidae) is in New Zealand, New Caledonia, New Guinea, and east to Madagascar. In Diptera, Thoracochaeta brachystoma (Sphaeroceridae) is in New Caledonia, New Zealand, PNG, and the Holarctic.
Outer arc groups absent in New Caledonia
Examples include the palms Metroxylon, Gulubia, Calamus, Physokentia and Hydriastele s. lat., Paramapania (Cyper.), the family Dichapetalaceae, Melastomataceae tribe Astronieae, Spiraeanthemum s. str. (Cunon.) (Fig. 2; Pillon et al., 2009), Myristica (Myristica.), Eurya (Thea.), Gunnera (Gunner.; montane in the tropics), Coriaria (Coriaria.; montane in the tropics), Parasponia (Ulm.), Haplolobus (Burser.), Dracontomelon (Anacard.), Pometia (Sapind.), Melicope sect. Lepta ‘dioecious group’ (Rut.) (Hartley, 2000), Atuna (Chrysobalan.), Endospermum (Euphorb.), Dillenia (Dillen.), Burckella and Palaquium (Sapot.), Weinmannia sect. Fasciculata (Cunon.), Anacolosa (Olac.), Osmoxylon (Aral.) and Dolicholobium (Rub.).
Many of these are also absent in Queensland (Metroxylon, Gulubia, Calamus, Paramapania, Gunnera, Coriaria, Parasponia, Astronieae, Eurya, Haplolobus, Dracontomelon, Pometia, Atuna, Burckella, Weinmannia, Anacolosa, Osmoxylon and Dolicholobium). Gunnera and Coriaria, both on the outer arcs (and in New Zealand), are only found in montane sites in the tropics and may be absent from New Caledonia for ecological reasons.
Solem (1958) cited ‘very great differences and only a few similarities’ between the land snails of New Caledonia and Vanuatu. For example, two high level taxa of the ‘Palaeo-Oriental fauna’ (Pupinidae, Trochomorphinae) and two of the ‘Pacific Ocean fauna’ (Partulidae, Microcystinae) reach Vanuatu but not New Caledonia.
In Homoptera, Fennah (1969) observed an ‘abrupt discontinuity’ between New Caledonia and Vanuatu, with some groups present in New Caledonia, Australia etc. but not on the islands to the north-east (Vanuatu etc.). Other groups of Vanuatu, the Solomon Is., and Fiji are absent from New Caledonia.
In Tipulidae (Diptera), Hynes (1993) highlighted several notable absences from New Caledonia, for example, Styringomyia didyma, otherwise widespread in the Pacific in Melanesia (New Guinea, Solomon Is., Vanuatu, Fiji), Polynesia (Samoa, Hawaii, French Polyesia) and Micronesia. It is replaced on New Caledonia by two endemic Styringomyia species. Other examples of fauna present on the outer Melanesian arc but absent from New Caledonia include the coral genus Zoopilus (Hoeksema, 1989), the cone shells Conus glaucus, C. nimbosus, C. proximus, and C. gloriamaris (Röckel et al., 1995), the Pacific landsnail family Partulidae (Caroline Is. east to the Marquesas) (Cowie, 1992), and the cicada subtribe Cosmopsaltriina (Duffels & Turner, 2002). Examples in Diptera include the genera Japenoides (Tabanidae) (Mackerras, 1971), Acropsilus (Dolichopodidae), Dissoptera (Syrphidae), Actinochaetopteryx and Cavillatrix (Tachinidae), and Zygothrica (Drosophilidae), the polymorphic species Mimegralla albimana (Micropezidae; Hennig, 1966), and in mosquitoes Uranotaenia, the Mansonia crassipes group and the Aedes (subgen. Stegomyia) scutellaris group (the last found throughout the South Pacific except New Caledonia and New Zealand) (Belkin, 1962; Taylor & Maffi, 1978). Examples in Homoptera include Lamenia and Phaciocephalus (Fennah, 1969), in beetles the elateroid Maelodorus (Muona, 1991) and the tribe Crinotarsini (Cerambycidae; Gressitt, 1961), and in spiders Conothele (Ctenizidae) and Nihoa (Barychelidae).
Taxa include, in fishes, the blenniids Blenniella caudolineata and Istiblennius bellus (Springer & Williams, 1994), the clupeids Amblygaster sirm and Sardinella melanura (Clupeidae) (Whitehead, 1985) and the lutjanids Lutjanus boutton, L. johnii, and L. timorensis (Allen, 1985); in birds, the kingfisher Todiramphus (Halcyon) chloris: Red Sea to Indonesia, New Guinea, the Solomon Is., Vanuatu (south to Aneityum) and Fiji/Tonga/Samoa, and Aerodramus (Collocalia) vanikorensis (Apodidae), widespread from the Philippines and through Melanesia to Vanuatu (del Hoyo et al., 1999), and in bats, the globally widespread families Hipposideridae and Molossidae (Nowak, 1999). The absence in New Caledonia of genera such as Chaerophon (Molossidae) and Emballonura (Emballonuridae), both widespread in the south-west Pacific including Vanuatu and Fiji, may perhaps reflect insufficient collecting (Parnaby, 2002) or they may be endemic to the outer arc track.