The MMO has been recognised as significant by many biogeographers, such as Croizat (1964) and Holloway (1979).
For Melicope and relatives (see main text), Poon et al. (2007) found that Melicope vitiflora from Fiji grouped with Pitaviaster. but did not sample the New Caledonian plants.
Direct New Caledonia – north-eastern Queensland connections are exemplified by the genus Austrovelia (Hemiptera; Andersen & Polhemus, 2003). A connection between New Caledonia and Cape York Peninsula in northern Queensland occurs in the spider Argiope lobata (Levi, 1983) and the Blediuscircularis group of staphylinid beetles (Herman, 1986).
The lichen Pseudocyphellaria argyracea occurs around the Indian Ocean, and from New Zealand, New Caledonia and New Guinea east through the Pacific, but is in Australia only at the MMO (Galloway, 1994).
The liverwort Acrolejeunea securifolia has one subspecies endemic in New Caledonia, one in the Philippines, New Guinea, northern Solomon Is.; one at the MMO, Norfolk I., north-eastern New Zealand; and one in the Cook Is. and French Polynesia (Gradstein, 1975). The moss Thuidium protensulum is in New South Wales/Queensland, Lord Howe I., and New Caledonia.
In Hernandia subgen. Hernandia (Hernand.; Kubitzki, 1970), the ‘moerenhoutiana Group’ of New Caledonia, Vanuatu, the Solomon Is., and east to the Society Is. and central America/West Indies is vicariant with the ‘bivalvis Group’, endemic to the MMO.
The Cunoniaceae are especially diverse in New Caledonia. Vesselowskya is endemic to the MMO (between the Hunter and Clarence Rivers in north-eastern New South Wales) and is sister toa clade Pancheria (New Caledonia), Cunonia (New Caledonia, South Africa) and Weinmannia (Mascarenes, Malesia, New Zealand, Pacific islands, South America) (Hufford & Dickison, 1992; Bradford, 2002).
Some New Caledonia – MMO groups also extend north in Australia. Neorapinia collina (Labiatae) of Norfolk I., New Caledonia and southern Vanuatu forms a group (Neorapinia) with Vitex lignum-vitae of the MMO/north-eastern Queensland and V. lucens of northern New Zealand (Mabberley, 1998).
In a related pattern, Duboisia (Solan.) comprises four more or less vicariant species: D. arenitensis in northern Australia (Arnhem Land), D. hopwoodii in central and western Australia, D. leichhardtii inland by the MMO, and D. myoporoides in coastal eastern Australia (Queensland-New South Wales) and New Caledonia (Haegi & Symon, 1984).
Geissois s. lat. (Cunon.) is at the MMO, disjunct in north-eastern Queensland, also (Geissois s. str.) in New Caledonia (centre of diversity), Vanuatu, and Fiji (Hoogland, 1984; Hopkins, 2006). Xylosma (Flacourt.) is in south-east Asia (5 species), Malesia (4 species), MMO (two locally endemic species; Jessup, 1982), New Caledonia (20 species), and South America (c. 70 species). Canavalia sericea (Legum.) ranges from the Caroline Is. to Fiji/Tonga/Samoa and east to the Society Is., with south-western outliers at New Caledonia (I. des Pins) and between the MMO and north-eastern Queensland (Sauer, 1964).
In Trimenia (Trimen.), the single New Caledonian species is a tree, as are the species of Sulawesi, New Guinea, Solomon Is., and Fiji to the Marquesas, but through its pollen the New Caledonian species has affinities with the high-climbing liane species, known from the MMO, north-eastern Queensland and PNG (Philipson, 1986).
The mesogastropod Pseudopisinna gragaria rigifera is in New Caledonia and at the MMO (Ponder & Yoo, 1980). The cowry Nesiocypraea langfordi moretonensis is in New Caledonia and at the MMO, and the other subspecies in the species, N. l. langfordi, is disjunct at the Philippines/southern Japan (Lorenz & Hubert, 1993) (see below for this disjunction). The lobster Parribacus caledonicus follows a linear distribution: MMO, New Caledonia, Vanuatu, Fiji, and Samoa (Holthuis, 1991).
The angelfish Centropyge flavissima is at Christmas and Cocos-Keeling Is. in the eastern Indian Ocean, Micronesia to eastern Polynesia, Melanesia and New Caledonia with an outlier at the MMO (Allen, Steene & Allen, 1998).
8. New Caledonia – Lord Howe I./Norfolk I. Notes on the main text
Page et al. (2005) assumed that Paratya is able to undertake long-distance trans-oceanic dispersal, but only because it occurs on isolated oceanic islands, and they cited endemic Paratya species on the Bonin Is. and the Chatham Is. east of New Zealand as support for this. The Bonin Is. occur at a subduction zone (the Mariana Trench) where islands have been coming and going since long before the modern islands existed. The Chatham Is. include Mesozoic terranes of the New Zealand basement and have a record of persistent volcanism. The biota includes many old New Zealand taxa. It is suggested that they also preserve an old, central Pacific biota transported on the former islands (now seamounts) of the Hikurangi Plateau to the Chatham subduction zone in the Cretaceous (Heads, 2009b).
Page et al. (2005) proposed that ‘The first biogeographical issue to consider is the geographical origin of the genus Paratya’, but there is no need to assume that there was a centre of origin located within the current range of the genus, and that the other species somehow migrated out from here. There is no evidence for this and it seems improbable given the precise mosaic patterns of endemism and vicariance in the genus. The (New Caledonia (Norfolk + Lord Howe)) arrangement is just one of these.
Keast (1966) cited relationships of the New Caledonia biota with Norfolk and Lord Howe Is. and also Australia. He described the mist-shrouded summit plateau of Lord Howe I. with dense moss forest and relict taxa such as the hemipteran family Peloridiidae (see track 3, above). Lillemets & Wilson (2002) described the ‘surprising’ diversity of terrestrial isopods on Lord Howe. They interpreted the fauna and the land as ‘contracted remnants’, and the taxa (including poorly dispersing groups) as ‘ancient relicts’ of a formerly more widespread biota. Australiodillo is one example, being endemic to Lord Howe (five species), New Caledonia (two species) and Queensland/New South Wales (one species) (see notes on New Caledonian isopods at the end of the Appendix).
Taxa endemic to Lord Howe and Norfolk Is. include the fern Cephalomanes bauerianum, the seed plants Elymus multifllurus var. kingianus (Gram.), Dianella intermedia (Herocallid.), Lagunaria patersonia subsp. patersonia (Malv.), Solanum bauerianum (Solan.) and Calystegia affinis (Convolvul.) (all from Green, 1994), and fauna such as the anemone fish Amphiprion mccullochi (Fautin & Allen, 1992).
The fern Doodia media subsp. media is in eastern Australia, Lord Howe, Norfolk and New Caledonia, and plants from the last three localities are most similar (Green, 1994).
Taxa connecting the Lord Howe/Norfolk centre with New Caledonia include Korthalsella taenioides f. emersa (Visc.) on Lord Howe, Norfolk, and New Caledonia (Molvray, 1997) and Oberonia titania (Orchid.)on Lord Howe, Norfolk, New Caledonia and Vanuatu (Lewis & Cribb 1989). Tylophora biglandulosa (Asclepiad.) has the same range plus Fiji (Green, 1994). The shorefish Plesiops insularis is at Lord Howe, Norfolk, New Caledonia and the Is. Chesterfield (north-west of New Caledonia) (Mooi, 1995).
Taxa with eastern Australia (especially MMO) – New Caledonia connections via Lord Howe and Norfolk Is. include the orchid Bulbophyllum argyropus (Orchid.) of the MMO, Lord Howe and Norfolk, very closely related to B. corythium of New Caledonia (Green, 1994). Celtis paniculata (Ulm.) is widespread in coastal eastern Australia, Lord Howe, Norfolk, and New Caledonia (Green, 1994). In Sarcomelicope (Rut.) there are five species endemic on New Caledonia and one other. This has one subspecies in Queensland/New South Wales, Lord Howe and Norfolk Is., another in New Caledonia and Vanuatu, and a third in Fiji (Hartley, 1982). Elaeodendron australe (Celastr.) of eastern Australia is related to E. curtipendulum of Lord Howe, Norfolk and New Caledonia (Green, 1994).
An interesting arc south of New Caledonia is displayed by Zanthoxylum pinnatum (Rut.): Lord Howe, Norfolk, southern Vanuatu (Erromango), Fiji/Tonga, with the closely related Z. pancheri on New Caledonia (Green, 1970).
The beetle genus Dematochroma (Chrysomelidae) has most species in New Caledonia, but has two endemics on Lord Howe and two on Norfolk (and one on Timor) (Jolivet et al., 2007a,b). The genus is absent from Australia and New Zealand but may possibly occur in other parts of Melanesia.
The spectacular angelfish Chaetodontoplus conspicillatus occurs from the Great Barrier Reef south to central New South Wales, and also on reefs around Lord Howe I., Norfolk I., and New Caledonia (Allen et al., 1998). Similar connections are seen in other reef fishes, such as the tripterygiid Enneapterygius rufopileus of Lord Howe I., Norfolk I., New Caledonia, and Fiji/Tonga (Fricke, 2002) and the gobiid Eviota hoesi endemic to Lord Howe I., reefs on the Lord Howe Rise (Middleton and Elizabeth Reefs), Norfolk I., and New Caledonia (Gill & Jewett, 2004).
9. New Caledonia – Lord Howe Island Lower plants
In lichens, Megalospora sulphureorufa is in Lord Howe and New Caledonia (southern Grande Terre, I. des Pins, and the Loyalty Is.) (Fig. 5; Sipman, 1983). The mosses Ctenidium pubescens, Entodon pancherianus and Fissidens arcuatus, and the fern Blechnum contiguum are known only from Lord Howe and New Caledonia.
Pandorea pandorana subsp. austrocaledonica (Bignon.) is on Lord Howe, New Caledonia and southern Vanuatu (Aneityum) (Green, 1994), Rottboellia coelorachis (Gram.) is also on Lord Howe, New Caledonia and southern Vanuatu (Tanna and Erromango) (Green, 1990). Zygogynum (incl. Bubbia, Belliolum and Exospermum) (Winter.) ranges in Lord Howe I., New Caledonia, the Solomon Is., PNG, and north-eastern Australia (Vink, 1985). The Guioa microsepala clade (Sapind.) occurs in Australia, Lord Howe and New Caledonia (van Balgooy et al., 1996). Polyscias cissodendron (Aral.) is on Lord Howe, New Caledonia, Vanuatu, and the Solomon Is. (Eibl et al., 2001).
The spider Argyrodes gracilis (Theridiidae) is only on Lord Howe, New Caledonia, and Samoa. The deep sea crustacean Kimbla comprises one species of New Caledonia and one from Taupo Seamount, south-west of Lord Howe I. (E 156° 10’, S 33° 15’) (Richer de Forges, 1993).
In morwongs (perciform fishes), Cheilodactylus francisi occurs around Lord Howe, New Caledonia, the Kermadec Is. and Hawaii (Burridge & Smolenski, 2004). The rail genus Tricholimnas (= Galliralluslafresnayanus) is only on Lord Howe and New Caledonia.
10. New Caledonia – Norfolk Island The islands along the Norfolk Ridge have high terrestrial endemism and the marine fauna on the seamounts along the ridge also show high diversity and endemism (Richer de Forges, Koslow & Poore, 2000).
Holloway (1996) wrote that ‘Evidence for the geological youth of Norfolk Island and its isolation since emergence through the volcanic events commencing 3 million years ago appears to be strong… and may be sufficient to justify the assumption that the island acquired its biotas by dispersal from the three nearest source areas, Australia, New Zealand and New Caledonia’. But this approach of relying solely on current geographic areas overlooks the possibility that there may have been other land in the region, as was later shown in this case.
The liverworts Anthoceros elegans and Radula robinsonii are both known only from New Caledonia and Norfolk. Plagiochila norfolkiensishas the same range, and So & Grolle (2001) compared it with a New Guinea endemic. P. inflata is in New Guinea, New Caledonia and Norfolk (So, 2000).
The thirteen New Caledonian Araucaria species form a clade that is sister to the Norfolk species A.heterophylla (Setoguchi et al., 1998). Euphorbia norfolkiana (Euphorb.) of Norfolk is very close to E. kanalensis of New Caledonia (Green, 1994). Exocarpus phyllanthoides (Santal.), Korthalsella disticha (Visc.) and Zehneria baueriana (Cucurbit.) are all endemic to New Caledonia and Norfolk I. (Green, 1979, 1994). Dysoxylum bijugum (Mel.), Euphorbia obliqua (Euphorb.), Tropidia viridifusca (Orchid.) and Phreatia paleata (Orchid.) are all on Norfolk I., New Caledonia and Vanuatu (Mabberley, 1998; Green, 1994; Lewis & Cribb, 1989).
In papilionid butterflies, Papilio ilioneus is on New Caledonia and Norfolk, Anaphaeis java peristhene is at these localities plus Vanuatu (Holloway & Peters, 1976). In other Lepidoptera, Holloway (1993) cited Norfolk – New Caledonia sister group relationships in Hydrillodes (related next to Fiji/Tonga, Kermadec Is.), Pyrrhorachis (next to Vanuatu), Agathia (next to New Guinea and Fiji), Ericeia (next to Vanuatu), Thalassodes (next to New Guinea and Bismarck Archipelago) and Cleora (New Caledonia/Vanuatu sister to Norfolk, then this group sister to Society and Rapa I.). In addition, Trigonistis toroensis (Noctuidae) of New Caledonia is sister to T. andersoni of Norfolk, and Luceria cooki (Noctuidae) is restricted to Norfolk and New Caledonia, Trissernis greeni (Noctuidae) is on Norfolk, New Caledonia and Vanuatu, and Ericeia hirsutitarsus (Noctuidae) is on Norfolk, New Caledonia, the Loyalty Is. and Vanuatu (Holloway, 1979).
11. New Caledonia – New Zealand Notes on the main text
Iredale (1912, p. 81) commented that the New Zealand – New Caledonia connection ‘has been constantly overlooked by most New Zealand writers’.
Chambers et al. (2001) interpreted the New Caledonia – New Zealand connection as an indication of dispersal to or from New Caledonia, but the molecular clock calibrations that they based their conclusions on are not accepted here. New Zealand – New Caledonia distribution is probably not due to dispersal from one country to the other (either prior to geological separation or after separation), but could be due to the vicariance of a widespread ancestor which also gave rise to Australian or Pacific relatives.
Based on the 3 Ma date as a maximum age for the Kermadec Is. and Norfolk species, Buckley & Simon (2007) inevitably concluded that Maoricicada (and the rest of the New Zealand alpine biota) is ‘very young’. (But even with calibration points which are too young, a Miocene age for the group was calculated and this is before the main rise of the New Zealand Southern Alps. Buckley & Simon (2007) referred to ancestral Maoricicada as being ‘preadapted’ to the alpine habitat and this would make sense as the group has survived uplift so successfully while maintaining largely allopatric distributions. It also indicates that the age of the mountains cannot be used to calibrate the age of taxa endemic there).
The basal position of the New Caledonian members in Cyanoramphus and the ideas of Herzer et al. (1997) on a ‘landbridge’ between New Caledonia and New Zealand led Boon et al. (2001) to propose an Australian origin for Cyanoramphus, followed by dispersal to New Caledonia and then to New Zealand. They suggested that the New Zealand – New Caledonia dispersal occurred within the last 500 000 years, assuming a molecular clock and rates inferred for another bird group (the basis for this rate was not given). This age is much younger than the proposed land bridge, and so the ‘land bridge theory of colonisation… may not be a viable explanation’ (Boon et al., 2001).
Former islands such as the one described from near Norfolk may have supplied both southern New Caledonia and northern New Zealand with biota as they came into existence; southern New Caledonia + northern New Zealand (Poor Knights/Hen and Chickens Is.) is a centre of endemism for groups such as the monocot family Xeronemataceae (Heads, 2009a).
In liverworts, Goebeliella is monotypic in New Zealand (the subantarctic Auckland Islands,
the main islands and the Chathams) and New Caledonia, and is sister to a large, more or less cosmopolitan group: Radulaceae, Frullaniaceae, Jubulaceae plus Lejeuneaceae (Heads, 2009a).
Schuster (1969) referred to a ‘fair number’ of liverwort species with the New Caledonia – New Zealand distribution. Miller et al. (1983) cited Chiloscyphus physanthus, Marchantia pileata, Riccardia plana, Schusterellamicroscopica, Tetracymbaliella comptonii and Zoopsis ceratophylla from New Caledonia and New Zealand only, Frullania squarrosula, Radula dentifolia and Riccardia bistrata from New Caledonia and North I., and Geocalyx caledonica from New Caledonia, South I. and Stewart I. Lopholejeunea colensoi is in New Caledonia and Campbell I. Schistochila caledonica is restricted to New Caledonia and the Paparoa Range, an important biogeographic centre in the South Island (So, 2003). In mosses, Eucamptodon inflatus is restricted to New Caledonia and New Zealand.
The pteridophyte Tmesipteris lanceolata is endemic to New Zealand and New Caledonia (Chinnock, 2003). The New Caledonian fern Adiantum novae-caledoniae is closely related to A. fulvum of New Zealand and Norfolk I. Molecular studies showed that Hymenophyllum paniense of New Caledonia is closest to H. scabrum of New Zealand (Ebihara et al., 2003).
In podocarp trees, Philippi in Aubréville et al. (1967-) wrote that Prumnopitys ferruginoides (Podocarp.) of New Caledonia has a conspecific form in New Zealand, but this actually refers to the closely related ‘corresponding species’ in New Zealand, P. ferruginea(de Laubenfels, 1978, D. de Laubenfels, pers. comm. 15 – v – 2008). These podocarps are the largest trees in many forests and their New Caledonia – New Zealand affinity is of special interest.
The monocots Astelia sect. Isoneuron (Astel.; Skottsberg, 1937) and Xeronema (above) are both only known from northern New Zealand and New Caledonia. The New Zealand endemic orchid Danhatchia (northern North Island, north-western South Island) is closely related to the New Caledonian Gonatostylis (Ormerod & Cribb, 2003a). Corynocarpus laevigatus (Corynocarp.) of northern New Zealand (including the Chatham and Kermadec Is.) is sister to C. dissimilis of New Caledonia (Wagstaff & Dawson, 2000). The New Caledonian dicot Strasburgeria is sister to the New Zealand Ixerba (Strasburger.). Schefflera sect. Schefflera (Aral.) is in New Zealand, New Caledonia, Vanuatu, and Fiji/Samoa (Plunkett et al., 2005) (cf. the liverwort Schusterella, in New Zealand, New Caledonia and Fiji).
The angiosperm Gymnostoma (Casuarin.) has a wide South Pacific distribution in Malesia, north-eastern Queensland, the Solomon Is., New Caledonia and Fiji, and, fossil, in the rest of eastern Australia, New Zealand, and South America. The earliest known fossils are Paleocene. In the species they sampled, Steane, Wilson & Hill (2003) found two clades, one in Malesia/Australia (six species) and one in New Caledonia (four species), and suggested that long-distance dispersal from New Zealand to New Caledonia might explain the New Caledonian presence of Gymnostoma. They reached this conclusion on the basis of fossil-based dates and proposed as ‘significant’ the fact that ‘while the Casuarinaceae have a fossil record in New Zealand that dates back to Paleocene … it does not extend back to the time when New Zealand is believed to have separated from Gondwanaland’. They concluded that this ‘suggests a requirement for dispersal… or a poorly known fossil record’. As the second alternative is very likely, the first is probably unnecessary.
In Proteaceae, Garnieria of New Caledonia (southern and north-western Grande Terre) is sister to Toronia of New Zealand (North Cape to East Cape), and the unrelated Knightia comprises two species in New Caledonia and one in northern New Zealand (Weston & Barker, 2006).
The plants Phelline (Phellin.), Paracryphia (Paracryph.) and Amphorogyne (Santal.) were previously only known from New Caledonia but fossil leaf material has been described from southern New Zealand (Pole, 2009).
The long history of the New Zealand – New Caledonia region as a biogeographic centre is indicated by many Mesozoic fossil groups endemic to it, for example, the gastropod Protodolium, known only from the Cretaceous of New Zealand, the Chatham Is., and New Caledonia (Stilwell, 1994).
The bivalve mollusc Monotis is one of the most characteristic taxa of the Upper Triassic. Of the five subgenera (Grant-Mackie, 1978), Monotis subgen. Inflatomonotis is in New Zealand, New Caledonia, and British Columbia, and M. subgen. Maorimonotis is restricted to New Zealand and New Caledonia. Overall, invertebrate faunas of the New Zealand and New Caledonian Triassic show ‘remarkable similarity’ (Campbell, 1994), with many taxa only known from the two localities (Campbell & Grant-Mackie, 1995; Begg & Grant-Mackie, 2003).
In molluscs, Athoracophoridae are a family of terrestrial slugs endemic to the lands and islands around both Coral Sea and the Tasman Sea (see section 5). They have most of their species in New Zealand (including the Subantarctic and Chatham Islands; Burton, 1963, 1980) and New Caledonia; others occur in Vanuatu, the Solomon Is., the Bismarck Archipelago, Papua New Guinea mainland, and eastern Australia (north-eastern Queensland to Sydney; not in Tasmania) (Heads, 2009a). The sister group of Athoracophoridae is the cosmopolitan family Succineidae.
The gastropods Alcithoe and Ataxocerithium are both known from New Zealand and New Caledonia, a distribution explained by the historically continuous distribution of the genera on the Norfolk Ridge (Bouchet & Kantor, 2003).
In spiders, Cambridgea (Stiphidiidae) is only in New Zealand (including Chatham Is.) and New Caledonia, as is Desis marina (Desidae). Hypodrassodes (Gnaphosidae) is in New Zealand, Lord Howe, and New Caledonia.
In beetles, Astetholea (Cerambycidae) is restricted to New Zealand and New Caledonia (Lu & Wang, 2005).
In cerambycid beetles, tribe Enicodini is restricted to New Zealand and New Caledonia (Gressitt, 1961). In staphylinid beetles, the tribe Nesoneini has the same range, with the related tribes in Australia, New Zealand and southern South America (Silphotelini and Austrothysini), and Australia and New Zealand (Anepiini). This is, ‘consistent with an ancient origin and vicariance during breakup of Gondwana’ (Newton, 2004).
In anthribid beetles, Androporus, Dasyanthribus, Helmoreus and Micranthribus are all restricted to New Caledonia and New Zealand (Holloway, 1982).
In Diptera, Anomalomyia, Tetragoneura, Sigmoleia (all Mycetophilidae), Aphopeas (Tabanidae) and Nervijuncta (Ditomyiidae) are all restricted to New Caledonia and New Zealand. Parentia (Dolichopodidae) is ‘a Gondwanan genus confined to Australia, New Zealand and New Caledonia’ in which the New Caledonian species appear to be close to New Zealand forms (Bickel, 2002). Also in Dolichopodidae, the distinctive xanthurus group of Paraclius comprises seven species in New Caledonia and one in North Island, New Zealand (Bickel, 2009).
In birds, the snipe Coenocorypha (Scolopacidae) is extant on the New Zealand subantartctic islands (Snares, Auckland, Campbell, Antipodes and Chatham Is.), with fossil records on North and South Islands of New Zealand, Norfolk I., New Caledonia and Fiji (Worthy, Miskelly & Ching, 2002). The extinct rail genus Tricholimnas of New Caledonia and Lord Howe is closest to Capellirallus of North Island, Cabellus of Chatham Is., and Vitirallus of Fiji (all extinct; Worthy 2004).
The bird Rhynochetos (Rhynochetidae) of New Caledonia is sister to the fossil Aptornis of New Zealand (Livezey, 1998).