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New Caledonia/outer arc vicariants

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14. New Caledonia/outer arc vicariants
Seed plants

Gonystylus (Thymel.) occurs through Malesia from Sumatra to the Solomon Is. and Fiji, and its sister, Arnhemia, is endemic to the Northern Territory of Australia. The clade is absent in New Caledonia but is replaced there by its sister group, comprising Deltaria and Solmsia both endemic to New Caledonia. The sister of this whole clade is Lethedon of New Caledonia (15 species) and Queensland (Beaumont et al., 2009).

Corynocarpus similis (Corynocarp.) of New Guinea, the Bismarck Archipelago, the Solomon Is. and Vanuatu vicariates neatly with a clade in New Caledonia (C. dissimilis) and New Zealand (C. laevigatus) (Wagstaff & Dawson, 2000). Arytera brackenridgei (Sapind.) of the Solomon Is., Vanuatu, and Fiji/Tonga/Samoa vicariates with three New Caledonian endemics (A. gracilipes, A. lepidota, A. arcuata) (van Balgooy et al., 1996). In Guioa (Sapind.), a clade on the Solomon Is., Vanuatu, Fiji and Tonga (G. sufusana etc.) vicariates with one on Australia, Lord Howe and New Caledonia (G. microsepala etc.) (van Balgooy et al., 1996).

Morierina (Rub.), endemic to New Caledonia, is closely related morphologically to Badusa of Fiji, Tonga, Vanuatu, Solomon Is., Biak I. (off north-western New Guinea), Palau I. and the Philippines (Jansen, 1984) (Motley et al., 2005, instead had Morierina in a clade with New Caledonian endemic species of Bikkia but the only Badusa in their sample was the Palau species).
15. New Caledonia – Vanuatu
Notes on the main text

The two species of Tinadendron (from Achille, 2006) are:

T. noumeanum: Noumea Peninsula, New Caledonia, thicket and sclerophyll forest on calcareous terrain at the edge of the sea, up to 50 m.

T. kajewskii of the Loyalty Is. (Maré) and southern Vanuatu (Erromango and Aneityum), at both localities in dense humid forest on uplifted coral, up to perhaps 300 m.

Examples of the Loyalty Is. – Vanuatu connection described recently include the landsnail Sturanya sublaevigatoides (Helicinidae) of the Loyalty Is. (all three islands) which is most similar to S. sublaevigata of Vanuatu (Tanna north to Banks and Torres Is.) (Richling, 2009) and the dipteran Bactrocera trilineola (Tephritidae) known only from the Loyalty Is. (Lifou, Mare) and Vanuatu (Mille, 2008).

Lower plants

New Caledonia/Vanuatu liverworts include: Acromastigum filum (in Vanuatu on Tanna), Bazzania bernieri (I. des Pins, Aneityum) and Lepidozia aubertii (in Vanuatu on Tanna). Schistochila integerrima is known from New Caledonia and Aneityum (So, 2003). Mosses restricted to the two areas include Campylopus verrucosus (in Vanuatu on Santo), Euptychium neo-caledonicum (Aneityum), E. dumosum, Fissidens subacutissimus, Glossadelphus obscurus (Santo), Leucobryum neo-caledonicum, Macromitrium pulchrum, Mittenothamnium nano-operculatum (Pentecost), Pogonatum circinatum (Aneityum, Santo), Pterobryella rigida (Aneityum), Rhizogonium medium (I. des Pins, Vanuatu), Symphysodontella cylindracea var. attenuata, Trichosteleum neo-caledonicum, and T. subrhinophyllum (Banks). Rhizogonium trachypelma is in the Kermadec Is., New Caledonia and Vanuatu (Malekula).

In ferns Sphenomeris deltoidea, Trichomanes assimile, T. laetum, Humata pusilla, Thelypteris herusiana, Cyathea vieillardii, Blechnum obtusatum, and B. gibbum (all from Aubréville et al., 1967-) and Sphaerostephanos richardsii (Holttum, 1977) are all restricted to New Caledonia and Vanuatu. Orthiopteris firma is in these islands and there are also possible records from Fiji (western Viti Levu).
Seed plants

Heliconia indica var. austrocaledonica (Helicon.) is in southern New Caledonia, the Loyalty Is., and western Vanuatu (mapped by Kress, 1990). Orchids with similar patterns include the following examples from Aubréville et al. (1967-) and Lewis & Cribb (1989).

Goodyera subregularis: New Caledonia (southern Grande Terre) and Vanuatu (throughout the east, north to the Santa Cruz Is.

Megastylis: endemic to New Caledonia (7 species) and Vanuatu (one species, on Aneityum).

Moerenhoutia grandiflora: New Caledonia and Vanuatu (Aneityum and Efate).

Dipodium punctatum var. squamatum: New Caledonia (widespread on Grande Terre and I. des Pins) and Vanuatu (Aneityum and Erromango).

Spathoglottis petri: north-eastern New Caledonia and Vanuatu (widespread).

Diplocaulobium ouhinnae: New Caledonia (north-eastern Grande Terre) and Vanuatu (northern and central).

Bulbophyllum neocaledonicum: New Caledonia (disjunct: southern and north-eastern Grande Terre) and Vanuatu (Santo).

B. atrorubens: New Caledonia (central and north-eastern Grande Terre) and Vanuatu (Santo).

Zeuxine vieillardii: New Caledonia (widespread on Grande Terre) and Vanuatu (Tanna and Santo).

Phaius robertsii: New Caledonia (disjunct: south and north-eastern Grande Terre) and Vanuatu (Banks Is.: Vanua Lava).

In eudicots, Ficus ser. Austrocaledonicae (Moraceae) is in New Caledonia, Loyalty Is. and Vanuatu (Tanna, Aneityum, Erromango, and Ambrym) (Corner, 1970). Arthroclianthus (Leguminosae) is in New Caledonia and Vanuatu (Lewis et al., 2005). Dizygotheca (Aral.) (= Schefflera ‘Pacific’ clade, subgroup Dizygotheca) is in New Caledonia and southern Vanuatu (Erromango, Tanna) (Plunkett et al., 2005; Lowry, 1989). Other New Caledonia/Vanuatu groups are ‘Strobilopanax’ (Aral.; = Meryta pro parte, Tronchet et al., 2005; Thorne, 1969) and Oxera (Labiatae) (de Kok & Mabberley, 1999). Polyscias schmidii (Aral.) of southern Vanuatu (Erromango) appears to be closely related to the New Caledonian P. nitida and P. nothisii (Lowry, 1989; Eibl et al., 2001). Phyllanthus myrianthus (Phyllanth.) of southern Vanuatu appears to be most closely related to New Caledonian species (Webster, 1986).

Santalum austrocaledonicum (Santal.) is in New Caledonia, the Loyalty Is. and southern Vanuatu (Aneityum, Erromango). Vitex collina (Labiatae) is restricted to New Caledonia and southern Vanuatu (Aneityum, Erromango, and Efaté) (Mabberley, 1998), as are Sarcomelicope simplicifolia subsp. neo-scotica (Rut.; in Vanuatu on Aneityum and Erromango) (Hartley, 1982) and Drypetes deplanchei subsp. deplanchei (in Vanuatu on Aneityum) (Green, 1990). Dysoxylum bijugum (Mel.) is on Norfolk I., New Caledonia, and Vanuatu (Mabberley, 1998). Other New Caledonia – Vanuatu species include Trachymene cussonii (Umbellif.) (Green, 1979), Alyxia podocarpa (Apocyn.; Middleton, 2002), Arytera neoebudensis (Sapind.; van Balgooy et al., 1996), and Lotus anfractuosus (Legum., Kramina & Sokoloff, 2004; in New Caledonia only on the Loyalty Is. and I. des Pins), Haloragis prostrata (Halorag.), Euphorbia pancheri (Euphorb.), and Trichospermum inmac (Tilia.) (the last three from Aubréville et al., 1967-)


In nudibranchs, Phyllidiopsis neocaledonica of New Caledonia and P. vanuatuensis of Vanuatu are sister species, separated by a trench 6000-7000 m deep ‘that might have caused fragmentation in the original range and subsequent speciation of the two’ (Valdés, 2001a). P. holothuriana is on the Norfolk Ridge 150 km southeast of Grande Terre, and southern Vanuatu (Aneityum) (Valdés, 2001b). The mesogastropod Eatonina heliciformis is in New Caledonia and central Vanuatu (Ponder & Yoo, 1980).

Solem (1958) stressed the great difference between the snail faunas of New Caledonia and Vanuatu (the ‘Pacific’ families of Mollusca that reach Vanuatu but not New Caledonia are cited above). Among the ‘few similarities’ he found was Draparnaudia singularis, only known from the two countries.

In spiders, Tangaroa dissimilis (Opell, 1983), Argiope caledonia (Levi, 1983), Arctosa konei (Lycosidae) and Dolomedes titan (Pisauridae) are restricted to New Caledonia and Vanuatu (in A. konei New Caledonia has one subspecies, Vanuatu the other). In Diptera, Idiohelius and Limonia (Goniodineura) apicifusca (Tipulidae) and Brachytarsina rouxi (Streblidae) have the same distribution, and the Tripteroides caledonicus group of mosquitoes occurs in New Caledonia, the Loyalty Is., Vanuatu, Santa Cruz Is., and Rotuma (north of Fiji) (Taylor & Maffi, 1978).

In Hemiptera, the marine water strider Halobates katherinae is in New Caledonia and Vanuatu (sister to a Fiji and Tonga pair of species) (Andersen, 1991), as is the genus Gnostocoris, which belongs to a Tasman Sea family of ‘obviously… ancient, primitive relicts’ (Monteith, 1979).

New Caledonia – Vanuatu Diptera include Dolichopus melanesiana (Dolichopodidae): New Caledonia and Vanuatu (Bickel, 2008).

Examples from beetles include the weevils Mauia subnotata (Anthribidae) (New Caledonia, Loyalty Is., Vanuatu, the genus Baladaeus (Curculionidae): New Caledonia, Loyalty Is., Vanuatu, and the genus Tetragynetes (Curculionidae) (Kuschel, 2008). Tetragynetes comprises two species, T. dispar: Loyalty Is. (Lifou, Mare), I. des Pins, Vanuatu (Eromango, Aneityum, Tanna), and T. onychialis: Vanuatu (Ambrym). Kuschel wrote that the genus ‘is only remotely related to Platysimus’, a central Pacific genus (see section 22 below) but is placed next to it in his treatment.

In Hymenoptera, Orasema koghisiana (Heraty, 1995), Pseudofoenus ritae (Jennings & Austin, 2002) and Theronia simillima (Gupta, 1962) are restricted to New Caledonia and Vanuatu. In beetles, the buprestid Metataenia subgen. Mroczkowskia is in New Caledonia and Aneityum (R. Holynski, pers. comm. 18 – viii – 2005).

In butterflies, Anaphaeis java peristhene (Papilionidae) is in Norfolk I., New Caledonia, the Loyalty Is., and Vanuatu, Danaus hamata moderata (Danaidae) is in New Caledonia and Vanuatu, D. pumila is in both these plus the Loyalty Is. and I. des Pins, Euploea sylvester tristis (Danaidae) is in New Caledonia and Vanuatu (plus Santa Cruz Is.), E. nemertes iphianassa has the same range, E. treitschkei jessica is in New Caledonia and Vanuatu, Luthrodes cleotas excellens and Nacaduba biocellata armillata (Lycaenidae) are in New Caledonia, Loyalty Is. and Vanuatu (Holloway & Peters, 1976). In other Lepidoptera, Ozola plana (Geometridae) of New Caledonia forms a ‘rather isolated section’ with O. achrophaea and O. hesperias of Vanuatu (Holloway, 1979). Cleora immemorata (Geometridae), Cephonodes lifuensis (Sphingidae), Agape lifuensis, Asota alienata (Hypsidae), Calathusa basirufa, Parallelia cunelineata (Noctuidae) are all on New Caledonia and Vanuatu, and (with the exception of Asota alienata) the Loyalty Is. (Holloway, 1979). Pareromene cheesmani (Pyralidae) is on New Caledonia and Vanuatu (Gaskin, 1974).

In amphidromous freshwater fishes, Lentipes kaaea (Gobiidae) is in New Caledonia and Vanuatu ( as are the bleniid reef fishes, Ecsenius tessera and E. isos (Bleniidae) (Springer, 1999); the tripterygiid Enneapterygius rhothion is in New Caledonia and southern Vanuatu (Erromango) (Fricke, 2002).

In birds, Ptilinopus greyii (Columbidae) is in New Caledonia, Loyalty Is., Vanuatu, and the Santa Cruz Is. Lichmera incana (Meliphagidae) is in New Caledonia (one subspecies), Loyalty Is. (two subspecies), and Vanuatu (two subspecies).

In Clytorhynchus (Fig. 10), C. pachycephaloides (Monarchidae) is in New Caledonia and Vanuatu (Erromango north to Banks Is). The other three species in the genus are C. vitiensis of Fiji, Rotuma, Tonga and Samoa, C. nigrogularis of Fiji and Santa Cruz, and C. hamlini of Rennell I. Relatives of Clytorhynchus include Neolalage, endemic to Vanuatu (Efate to Banks Is.), and Mayrornis of Fiji and Santa Cruz Is. These ranges all indicate an important break by Santa Cruz and Banks Is. (Distributions of Vanuatu birds are taken here from Bregulla, 1992, rather than Dickinson, 2003).

Other birds restricted to New Caledonia and Vanuatu include Accipiter fasciatus vigilax (Accipitridae), Chalcophaps indica sandwichensis (Columbidae) (including Banks and Santa Cruz Is.), Turdus poliocephalus pritzbueri (Lifu and Tanna), Lalage leucopyga simillima (Loyalty Is. and southern Vanuatu) and Coracina caledonica (Grande Terre, Loyalty Is., Erromango, Malekula and Santo). Phylidonyris (Guadalcanaria) undulata (Meliphagidae) of New Caledonia and P. notabilis of Vanuatu (central and Santo) are closely related (Diamond & Marshall, 1976) and sometimes separated as genus Glycifohia (Dickinson, 2003); (Phylidonyris is then restricted to Australia).
16. New Caledonia – Vanuatu – Fiji
Takhtajan (1986) wrote that a ‘curious connection’ exists between the floras of New Caledonia and those of Vanuatu and Fiji. This area of endemism is ‘curious’ because it would not be predicted using the model of dispersal from Asia and Australia that Takhtajan and most authors have followed.
Lower plants

The moss Ectropothecium pacificum and the fern Leptopteris wilkesiana are both restricted to New Caledonia, Vanuatu, and Fiji/Samoa.

Seed plants

The following orchids are all endemic to New Caledonia, Vanuatu and specified localities in the Fiji/Samoa/Tonga area (Lewis & Cribb, 1989): Microtatorchis schlechteri (Fiji; Kores, 1991, treated the Fiji material as a separate species), Erythrodes oxyglossa (Fiji, Wallis and Futuna Samoa, Tonga), Goodyera rubicunda var. triandra (Fiji, Samoa), Earina valida (Fiji, Samoa), Bulbophyllum samoanum (Fiji, Samoa), Pristiglottis montana (Samoa), Coelogyne lycastoides (Fiji, Samoa, Tonga; Clayton, 2002), Peristylus novoebudarum (Fiji/Niuafou), and Didymoplexa micradenia (Fiji, Samoa, Tonga; Kores, 1991; Lewis & Cribb, 1989, also included a Javanese species in the synonymy).

Examples from dicots include the following.

Geissois s. str. (Cunon.): New Caledonia, Vanuatu (Vanua Lava to Aneityum), Santa Cruz Is., and Fiji (Pillon, 2008).

Balanops sparsifolia (Balanop.): New Caledonia, keyed with B. pedicellata of Vanuatu and Fiji (Carlquist, 1980; the only other locality for the genus is in the Cairns area, north-eastern Queensland).

Ficus scabra (Mor.): New Caledonia, Vanuatu, and Fiji/Samoa/Tonga (Smith, 1979-1996).

Piliocalyx (Myrt.): New Caledonia, southern Vanuatu (Erromango), and Fiji (Smith, 1979-96; Craven, 2001; Craven et al., 2006).

Leucopogon septentrionalis (Eric.): New Caledonia, Vanuatu, and Fiji (Smith, 1979-1996).

Manilkara dissecta (Sapot.): New Caledonia, Vanuatu, and Fiji/Samoa/Tonga (Smith, 1979-1996). Acacia simplex (Legum.) has the same range (Smith, 1979-1996).

Schefflera ‘Pacific clade’, ‘subgroup Gabriellae’ (Aral.): New Caledonia, southern Vanuatu (Aneityum), and Fiji (Plunkett et al., 2005; Lowry, 1989).

Cyclophyllum (Rub.): New Caledonia, Vanuatu, and Fiji (Morat, Jaffré & Veillon, 2001).

Plectranthus forsteri (Labia.): New Caledonia, Vanuatu, and Fiji/Tonga (Smith, 1979-1996).

Insects in New Caledonia, Vanuatu and Fiji include, in Diptera Bactrocera (Bactrocera) curvipennis (Tephritidae), in Homoptera Plestia oceanica (also Samoa; Fennah, 1969), in beetles Cnemidothrex (Curculionidae (Kuschel, 2008), in butterflies Hasora khoda khoda (Hesperidae), a related group of subspecies in Appias albina (including A. a. psyche) (Papilionidae) (also Samoa/Tonga), Euploea tulliolus forsteri (southern Vanuatu) and Jamides carissima susana (Lycaenidae) (Holloway & Peters, 1976), and in moths Gymnoscelis sara and Scardamia eucampta (Geometridae), Parallelia prisca (also Samoa/Tonga), Hypena fijiensis (Noctuidae) and Xenothictus/Xeneda (Tortricidae; the two may be congeneric) (Holloway, 1979).


Taxa endemic to New Caledonia – Vanuatu – Fiji (and often including nearby Samoa and Tonga) include the reef fishes Helcogramma hudsoni (Tripterygiidae) (also Samoa and Kiribati; Fricke, 2002), Ecsenius fourmanoiri (also Lord Howe I. and southern Tonga, in Fiji only in the Lau Group; Springer, 2002), and the bird Rhipidura spilodera (Rhipiduridae) (northern and central Vanuatu). In bats, Notopteris is a ‘very ancient, long-isolated lineage’ (Flannery, 1995) that occurs in northern New Caledonia (N. neocaledonica), southern Vanuatu and Fiji (N. macdonaldi) and Tonga (as subfossils).

17. New Caledonia – Fiji/Samoa/Tonga
Lower plants

In mosses, Trichostomum insulare is in New Caledonia and Fiji (Viti Levu), Leucobryum tenuifolium and Distichophyllum vitianum are in New Caledonia, Fiji and Samoa. In liverworts, Lophocolea parva, Megaceros monospirus, Schusterella parhami and Telaranea bisetula are in New Caledonia and Fiji, Lopholejeunea renistipula is known only from I. des Pins and Fiji. The ferns Ctenopteris crassifrons and Dictymia mckeei are both in New Caledonia and Fiji, as are Lindsaea moorei (in Fiji only on Mt. Korobaba) and Belvisia melanesica (in Fiji only on Mt. Tomanivi). Asplenium carruthersii of Fiji is very close to A. oligolepidum of New Caledonia (Brownlie, 1977). Christella pacifica ranges in New Caledonia, Fiji, and Samoa (Holttum, 1977), Schizaea melanesica is in New Caledonia, Fiji, and Tonga (Brownlie, 1977).

Seed plants

Hibbertia lucens (Dillen.) is restricted to New Caledonia and Fiji. Smith (1978, 1979-1996) regarded this distribution as ‘very unusual’ in plant species. He wrote that ‘Because of the presumably independent origin of the two archipelagoes in the remote past, it seems likely that in this case one or more disseminules travelled from New Caledonia to Fiji comparatively recently (in terms of geological time) and that there has not yet been time for independent evolutionary directions to have become apparent’. In fact, the geological history of New Caledonia and Fiji may not have been as independent as Smith suggested (see main text).

Thorne (1969) cited Buraeavia (Picrodendr.) (currently placed in the more widespread Austrobuxus) as a New Caledonia – Fiji group but Smith (1979-1996) and Airy Shaw (1980) regarded the Fijian species as closer to the New Guinea species than to any in New Caledonia (cf. section 14, above).

Acmopyle (Podocarp.) is extant in New Caledonia and Fiji (there are fossil records from Australia, Antarctica and Chile; Hill & Brodribb, 1999, also New Zealand; Pole, 2007). The orchids Anoectochilus imitans, Octarrhena oberonioides, Phreatia pachyphylla, and Nervilia platychila are all restricted to New Caledonia and Fiji (Kores, 1991). Hetaeria whitmeei and Bulbophyllum pachyanthum are on New Caledonia and Fiji, Samoa and Tonga (Kores, 1991). Planchonella sphaerocarpa had some weak support as sister of P. membranacea of Fiji (Swenson et al., 2007). Polyscias joskei (Aral.) of Fiji is sister to a group of New Caledonian species (Eibl et al., 2001). Diospyros pancheri (Eben.) of New Caledonia was regarded as closest to D. foliosa of Fiji by Kostermans (1977). Passiflora barclayi (Passiflor.) is only known from New Caledonia and Fiji. (De Wilde, 1972, synonymised the species under the more widespread P. aurantia, while Green, 1972, Smith, 1979-1996, and Jaffré et al., 2001, recognised P. barclayi as distinct). The genus Pytinicarpa (Compositae) is restricted to New Caledonia (the species were treated under Brachycome in Jaffré et al., 2001) and Fiji (treated under Keysseria in Smith, 1979-1996) (Nesom, 2001). In a broad sample of Pacific Planchonella species (Sapot.), P. sphaerocarpa of New Caledonia was placed with P. membranacea of Fiji (Swenson, Munzinger & Bartish, 2007; New Guinea species were not widely sampled and some might be involved here).

In other examples, Podocarpus colliculatus (Podocarp.) of the I. des Pins is closest to P. pallidus of Tonga (de Laubenfels, 2005). Typical Nicotiana fragrans (Solan.) occurs on New Caledonia, the Loyalty Is. and Tonga (there is a distinct form endemic to the Marquesas).


The spider crab Guinotinia comprises one species in New Caledonia, the other in Fiji and Tonga (Richer de Forges & Ng, 2009).

In craneflies (Tipulidae), Limonia fijiana is endemic to New Caledonia, Fiji and Samoa, L. ochricapilla to New Caledonia and Fiji, in dolichopodid flies; Cymatopus baravikai of Fiji is ‘most similar in appearance’ to C. neocaledonicus of New Caledonia (Evenhuis, 2005b). In beetles, the staphylinid Megarthrus has two sister species in New Caledonia and Fiji, respectively (Cuccodoro, 1998).

In fulgoroid Homoptera, Suva albonotata is is New Caledonia, Fiji, Samoa and Sri Lanka (Fennah, 1969). In Lepidoptera, Anisodes lautokensis (Geometridae) is restricted to New Caledonia and Fiji, Poecilasthena leucodrya (Geometridae) is on these two and also Samoa (Holloway, 1979). Nola lichenosa (Nolidae) is in New Caledonia, Fiji and Rotuma; specimens from Vanuatu have a similar facies but may differ. Polyura gamma (Nymphalidae) of New Caledonia is related to a Fiji species (Holloway & Peters, 1976).


Drepanoptila (Columbidae) is a monotypic genus endemic to New Caledonia (and ‘without doubt the most beautiful bird of the country’ – Delacour, 1966). Goodwin (1983) regarded it as ‘very similar’ to the orange doves of Fiji (Ptilinopus victor superspecies). Gymnomyza (Meliphagidae) occurs in New Caledonia (‘Leptomyza’), Fiji and Samoa, with one endemic species in each.
18. New Caledonia – Samoa
Lower plants

Taxa endemic to New Caledonia – Samoa include liverworts cited by Miller et al. (1983): Bazzania falcifolia, Cololejeunea arrectifolia, Frullania angustistipa, Heteroscyphus jackii, and Lophocolea explanata, and So (2000) adds Plagiochila bialata. Other examples include the moss Syrrhopodon polytrichoides and the fern Humata brackenridgei.

Seed plants

The tree Crossostylis (Rhizophor.) has one clade in New Caledonia, Samoa and French Polynesia, and a vicariant in Fiji, northern Vanuatu and the Solomon Is. (Setoguchi et al., 1998).


Examples include the spiders Tamasesia (Mysmenidae) and Argyrodes samoensis (Theridiidae), the dolichopodid fly Helixocerus (Bickel, 2002), and the cone shell Conus exiguus (Röckel et al., 1995).

In related patterns, the spider Argyrodes gracilis is in Lord Howe Is., New Caledonia, and Samoa. Thalassomya maritima (Chironimidae) is in New Caledonia and Samoa, and disjunct in Micronesia (Marshall Is, Palau, Guam, northern Mariana Is.) and Hong Kong. These connections probably run south of Fiji, as in the Crossostylis clade cited above. Nevertheless, a northern track may be shown by the moth Nola samoana (Nolidae) of New Caledonia, Vanuatu, Rotuma (north of Fiji) and Samoa. It is replaced in Fiji by N. fijiensis (Holloway, 1979).

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