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22. New Caledonia – central South Pacific
Notes on the main text

New Caledonia has been seen as a centre of origin for many other groups; for example, Holloway (1979) wrote that the tortricine moths Xenothictis/Xeneda ‘appear to have radiated in New Caledonia and spread to other areas of the Pacific’, and in Cupaniopsis (cited in the main text), Adema (1991) proposed New Caledonia as a centre of origin for Pacific species. In Meryta (Aral.), Tronchet et al. (2005) suggested that because the New Caledonian M. denhami lies phylogenetically among other volcanic island taxa, it is derived by recent back-migration from a centre of origin held by these. Instead, the diversity of Meryta in New Caledonia may represent a series of accretions of arcs and floras onto the continental crust. Although one member of the central Pacific clade (M. denhami) has already been accreted onto New Caledonia, the other members survive on and around central Pacific arcs and volcanic centres (Micronesia – Vanuatu – eastern Polynesia) and remain unaccreted, at least for now.


Lower plants

Examples in mosses include Racopilum spectabile: Malesia, Micronesia (Caroline Is., Mariana Is.), Solomon Is., Vanuatu, New Caledonia, Fiji/Samoa with the typical subspecies, and in New Caledonia and Tahiti, each with endemic subspecies, Rhizogonium setosum: New Guinea, Vanuatu, New Caledonia, Fiji/Samoa, and the Society Is., Macromitrium tongense: New Guinea, New Caledonia (I. des Pins), Lord Howe, Fiji/Tonga and Henderson I. (near Pitcairn), Orthorrhynchium cylindricum: Loyalty Is., Vanuatu, Fiji/Tonga/Samoa, Australs, Society Is., and Marquesas, Papillaria helictophylla: New Caledonia, Vanuatu, Wallis & Futuna, the Society Is., and the Marquesas, Syrrhopodon mammillatus: New Caledonia, Fiji/Samoa, and the Society Is. (Raiatea), S. banksii: New Caledonia, Fiji and Samoa to Micronesia and the Marquesas (Fisher, 2006), Vesicularia aperta: New Caledonia, Vanuatu, Wallis & Futuna, and Tahiti (with a queried record from Hawaii). Thuidium obtusifolium var. neocaledonicum is endemic to New Caledonia, the type variety is in southern Vanuatu (Erromango), Austral, Society, Tuamotu, Gambier (incl. Pitcairn and Henderson) and Marquesas Is.

Examples in liverworts include Dendroceros tahitiensis: New Caledonia and the Society Is. (Tahiti and Moorea), Cheilolejeunea cookiensis: Louisiade Archipelago (south-eastern New Guinea), New Caledonia, Fiji/Samoa, Cook Is. (Rarotonga) and Society Is. (Tahiti) and Porella viridissima: Norfolk I. and New Caledonia east to Tahiti (So, 2002).

In pteridophytes, Tmesipteris lanceolata of New Zealand and New Caledonia is related to T. norfolkensis of Norfolk I. and T. gracilis of the Society Is. and Marquesas (Chinnock, 2003). Examples in ferns (records from Aubréville et al., 1967-) include Culcita straminea: New Caledonia, Vanuatu, Fiji and central Polynesia, Antrophyum alatum: New Caledonia, Fiji/Samoa and Tahiti, Teratophyllum wilkesianum: New Caledonia, Samoa and Tahiti, Ctenopteris blechnoides: New Caledonia to Polynesia, Paesia rugosula: Philippines (Luzon), New Caledonia and Tahiti (it resembles P. scaberula of New Zealand), Grammitis neocaledonica of New Caledonia, which is very close to G. hookeri of ‘Polynesia’, and Cyathea alata of New Caledonia, which is related to C. decurrens of Vanuatu, Fiji/Samoa, the Cook Is. and Society Is. (Holttum, 1964).


Seed plants

In Trimenia (Trimen.), the tree species range from Sulawesi through New Guinea, the Solomon Is., and Fiji to the Marquesas. As mentioned, the single New Caledonian species is a tree. It may belong to a Coral Sea track as its pollen shows affinities with the liane species, found in north-eastern New South Wales (the MMO) and disjunct in PNG (Philipson, 1986).

Monocot examples of New Caledonia – central Pacific endemism include the orchid genera Earina: New Zealand (in the three main islands plus Chatham Is.; two species), New Caledonia (three species), and one species each in Vanuatu, Fiji, Samoa and Tahiti, and Microtatorchis: New Caledonia, New Guinea, Philippines, the Caroline Is., Fiji, and Samoa to the Society Is. (Smith, 1979-1996).

Other monocots with a New Caledonia – central Pacific range include Cenchrus calyculatus (Gram.): New Caledonia, Micronesia, east to the Marquesas, Oberonia equitans (Orchid.): New Caledonia and Vanuatu east to the Tuamotu Is., and Taeniophyllum fasciola (Orchid.): New Caledonia east to the Society Is.

Other examples include the following eudicots.

Ficus prolixa (Mor.): Micronesia, Vanuatu and New Caledonia (not the Solomon Is.), east to south-eastern Polynesia (Smith, 1979-1996).

Schleinitzia (= Prosopis insularum, Mimos.): Philippines, New Guinea, Solomon Is., Vanuatu, New Caledonia, and Fiji – south-eastern Polynesia (de Vogel, 1975; Lewis et al., 2005).

Homalanthus nutans (Euphorb.): Caroline Is., New Caledonia, Vanuatu and Society Is.

Pisonia sect. Paucistaminatae (Nyctagin.): New Guinea (4 species), and one species in each of the Solomon Is., New Caledonia, Society Is. and Hawaii (Stemmerik, 1964).

Capparis nummularia (Capparid.): Mariana Is., Palau and other Caroline Is., Mussau I. north of New Ireland (but not New Ireland, New Britain, or New Guinea), Solomon Is., New Caledonia, and Fiji to the Tuamotu Is. and Pitcairn I. (St. John, 1965).

Canavalia sericea (Legum.): from the Caroline Is. to Fiji/Samoa/Tonga and east to the Society Is., with southwestern outliers at New Caledonia (I. des Pins) and Queensland (Sauer, 1964).

Weinmannia sect. Leiospermum: New Zealand, New Caledonia, Vanuatu, Bismarck Archipelago (and Sulawesi), east to the Marquesas (Hopkins & Bradford, 1998).

Crossostylis (Rhizophor.): New Caledonia, northern Vanuatu, Solomon Is., and Fiji east to the Marquesas. Morat et al. (1984) suggested Crossostylis and Earina had a ‘Pacific island origin’. Both genera show notable allopatry among their respective species and also with related genera and so a vicariance origin of the pattern is likely.

Grewia crenata (Tilia.): New Caledonia, the Loyalty Is., Vanuatu, Fiji, and east to the Society Is.

Morinda myrtifolia (Rub.): New Caledonia (four varieties) east to the Tuamotu Is. and Marquesas (Smith, 1979-1996).

Hedyotis foetida (Rub.): New Caledonia to the Austral Is. (Smith, 1979-1996).

Nicotiana fragrans (Solan.) of New Caledonia (southern Grande Terre, Loyalty Is.), Tonga and the Marquesas.

In Alstonia (Apocyn.) a large clade comprises 12 New Caledonian endemics, plus A. costata of New Caledonia, Vanuatu and the Solomon Is., east to the Marquesas (Sidiyasa, 1998). This central Pacific group is sister to a species of eastern Australia.


Invertebrates

In freshwater shrimps a New Caledonia – south-eastern Polynesia clade was recovered in the genus Caridina (Page et al., 2007). These authors cited ‘surprisingly good dispersal abilities’ to explain patterns in this genus but, again, chance processes cannot explain the central Pacific endemism and its standard boundary at the old plate margin: another clade of Caridina is in Okinawa, Malesia, and Australia, vicariant to the west of the central Pacific group.

In spiders, the genus Tangaroa (Uloboridae) comprises one species in the Caroline Is. (Yap), one in New Caledonia/Vanuatu, and a third in south-eastern Polynesia (Tahiti, Rapa) (Opell, 1983). A central Pacific clade of Dicaea (Thomisidae) in New Caledonia, New Zealand, Fiji, Tonga and the Austral Is. (including Rapa) vicariates neatly with Mimusenops: Society Is., Marquesas, Hawaii, and the Americas (Garb & Gillespie, 2006). The authors suggested that the pattern is due to island-hopping across the Pacific, but this does not account for the very precise east Pacific/west Pacific split around a node in south-eastern Polynesia.

Examples in Diptera include the Bactrocera atra-passiflorae complex: New Caledonia, Tuvalu, and Fiji east to French Polynesia (Michaux & White, 1999), Abbemyia baylaci (Dolichopodidae): New Caledonia, Vanuatu, east to French Polynesia (Bickel, 2002), vicariating with the other two congeners in eastern Australia, and Limonia subgenus Doaneomyia (Tipulidae): New Caledonia, Vanuatu, Fiji, and the Society Is.

Examples from beetles (all Curculionidae, from Kuschel, 2008) include Platysimus: Solomon Is. and Kiribati to New Caledonia, Kermadec Is., Austral and Society Is., Viticis: southern Ryukyu Is. (Ishigaki I. and Iriomote I., off Taiwan) (this species keys out first suggesting it may be a separate clade), New Caledonia, Loyalty Is., Vanuatu (Santo to Aneityum), Fiji, and one species in the Cook Is. (Mangaia) and Marquesas (Nuku Hiva), and Elytroteinus geophilus: north-eastern New Caledonia (Balade), Vanuatu (Epi), Fiji (Lakemba, Moce only), Wallis, Samoa (Upolu), Tonga (Tongatapu, Vavau), and Cook Is. (Rarotonga, Aitutaki). The species is close to E. vitiensis: Fiji (Nandarivatu, 1000m).

Examples in butterflies include Badamia atrox (Hesperidae): New Caledonia, Vanuatu, Fiji and Marquesas, Euploea lewinii: New Caledonia, Vanuatu and the Santa Cruz Is., east to the Society Is., and Hypolimnas antilope lutescens (Nymphalidae): New Caledonia, Vanuatu, Fiji/Samoa/Tonga and the Cook Is. (all from Holloway & Peters, 1976). In other Lepidoptera, Anisodes samoana (Geometridae) is in New Caledonia, the Loyalty Is., Vanuatu, Fiji/Samoa/Tonga, and the Society Is. (Holloway, 1979).


Vertebrates

In amphidromous freshwater fishes, the genus Lentipes (Gobiidae) is in Africa, Indonesia, and the Pacific, where it occupies the Ryukyu Is. and Hawaii in the north, and New Guinea, New Caledonia, Vanuatu and the Marquesas in the south, each with endemic species (www.fishbase.org).

The petrel Pseudobulweria rostrata has breeding records from New Caledonia and the Solomon Is. east to the Marquesas.

Other Central Pacific endemic birds include the parrot Cyanoramphus (New Zealand – New Caledonia – Society Is.) (see main text and notes, section 11).


23. New Caledonia – Polynesia – Hawaii
Lower plants

Liverwort examples include Acromastigum sect. Acromastigum: New Zealand, New Caledonia, and Hawaii, Cololejeunea ceatocarpa: Réunion, New Caledonia and Hawaii, Lophocolea autoica: New Caledonia and Hawaii, and Bazzania inaequabilis: New Caledonia, Samoa, and Hawaii (with a queried record in the Philippines). The moss Holomitrium seticalycinum is only recorded in New Caledonia and Hawaii.


Seed plants

In Freycinetia sect. Freycinetia (Pandan.) Stone (1973) recognised a group of species from New Zealand and Norfolk I. (F. baueriana), New Caledonia (F. longispica), Rarotonga (F. wilderi), Rapa I. (F. rapensis), and Hawaii (F. arborea). This central Pacific affinity is distributed very differently from two other Freycinetia clades in New Caledonia, sect. Solmsiella of New Caledonia and New Guinea (cf. secton 11) and sect. Warburgiella: New Guinea, Solomon Is., Vanuatu, and New Caledonia (section 19) (Stone, 1981).

In a sample of spiny solanums (Solanum sect. Leptostemonum), the New Caledonian S. pancheri is sister to two Hawaiian species in a strongly supported clade that is sister to and vicariant with Australian species (Levin et al., 2006).

The Hawaiian species of Melicope sect. Pelea (Rut.) are closest to the New Caledonian M. vieillardii (Hartley, 2000); this clade’s relatives occur in south-eastern Polynesia. (New Caledonian species were not sampled in the molecular study of Harbaugh et al., 2009.)

In a sample of Pacific Pittosporum species (Pittospor.), New Caledonian species formed a clade with species of Fiji, Tonga and Hawaii (Gemmill et al., 2002).

Lysimachia mauritiana (Primul.) occurs in the central Pacific only on New Caledonia, Rapa I., and Hawaii. It is also in China, Japan and the Mariana Is., and disjunct in Mauritius and Réunion in the south-west Indian Ocean (van Balgooy & Bentvelzen, 1984).

In an example of character, rather than taxon, geography, leaf heteroblasty – pronounced changes between juvenile and adult leaves – is especially common in plants of Hawaii, New Caledonia, New Zealand and the Mascarenes (Burns & Dawson, 2006). The authors also observed that the heteroblasty in New Zealand and New Caledonia species is similar despite the different climates, ‘and the phenomenon likely results from some aspect of their shared geological history’.


Invertebrates

Examples in molluscs include the cone shell Conus smirna: New Caledonia, Kermadec Is., and Hawaii (Röckel, 1995) and the cowry Blasicrura rashleighana, with one subspecies on New Caledonia and the only other one on Hawaii (Lorenz & Hubert, 1993). The hypogeal shrimp Ligur uveae is known from the Loyalty Is., Fiji, Tuvalu, Hawaii, Moluccas/Philippines and the Aldabra Is. north of Madagascar (Maciolek, 1983).

In the cosmopolitan beetle genus Rhantus (Dytiscidae), the ‘pacificus group’ is definitely known from New Zealand, New Caledonia, the Solomon Is., New Guinea, and Hawaii (Balke et al., 2007b; certain species from Chile/Argentina, Assam, and West Africa may perhaps also belong here, M. Balke, in litt. 8 – ii – 2007).
Vertebrates

In the morwongs cited above, Cheilodactylus francisi of Lord Howe, New Caledonia and the Kermadec Is. is most closely related to C. vittatus of Hawaii, and these two are related to an Easter I. species. The monotypic snapper genus Randallichthys is known from New Caledonia, Hawaii and Okinawa (Allen, 1985).




24. New Caledonia (/Melanesia) – South America
Some documentation for a New Caledonia – tropical South America track was given in a previous paper (Heads, 2006).
Lower plants

In fungi, Amparoina spinosissima is recorded from New Caledonia, Argentina, Colombia (Horak, 1983) and Japan (as Mycena spinosissima; Takahashi, 1998) giving records in the four corners of the Pacific, but not in Australia.

In liverworts, Dendroceros brasiliensis is in New Caledonia, Brazil, and the West Indies (Cuba, St Vincent, Guadeloupe) and Frullania bicornuta in New Zealand (North Island), New Caledonia, Juan Fernandez, Chile, Patagonia and Fuegia. In mosses, Himantocladium bauerlenii is in New Guinea, New Caledonia, North and South America. In ferns, Schizaea intermedia of New Caledonia and S. wagneri of New Guinea/Malesia are related to S. germani and relatives in tropical America (Selling, 1946). In this group the Old World and New World taxa show parallel clines, with the largest spores in both occurring in the south.

In the tree fern Cyathea, the ‘Cyathea clade’ (Large & Braggins, 2004) is in New Guinea, eastern Australia, New Caledonia east to Frrench Polynesia, and in tropical South America.


Seed plants

The podocarp Retrophyllum Page (= Podocarpus sect. Polypodiopsis, = Decussocarpus de Laub. sect. Decussocarpus ) (extant and fossil) has a very similar Melanesia – South America pattern, with extant species in New Caledonia, Fiji, New Guinea, Moluccas, and tropical South America (Peru to Colombia and Venezuela), and fossil species in Australia, New Zealand and Chile (Herbert et al., 2002).

In Hernandia subgen. Hernandia (Hernand.; Kubitzki, 1970) the moerenhoutiana Group ranges: New Caledonia, Vanuatu, Solomon Is., Fiji, east to the Society Is., and also in Nicaragua, Jamaica and Hispaniola. This Pacific group is vicariant with the bivalvis Group, endemic to the McPherson-Macleay Overlap.

The orchid Erythrodes ranges in north-eastern India, China, south-east Asia, Malesia, New Guinea, Vanuatu, New Caledonia, and Fiji/Tonga/Samoa, and is most closely related to Microchilus, widespread in the Galapagos and tropical America (Ormerod & Cribb, 2003b).



Astelia subgenus Asteliopsis (Astelia.) comprises sect. Isoneuron in northern New Zealand/New Caledonia, sect. Periastelia in Hawaii/Marquesas/Rapa (these two sections form group A), and sect. Micrastelia of Patagonia/Falkland Is. (Skottsberg, 1937). (The mainly South Pacific genus Astelia is abundant in many forests as well as alpine communities. A detailed revision is long overdue and would be of great interest).

Heliconia (Helicon.) occurs in the south-west Pacific (1-2 species; New Caledonia, Vanuatu, Solomon Is., New Guinea, Moluccas, and Fiji, Samoa) and in tropical America (50-200 species; Mexico, West Indies to southern Brazil) (Kress, 1990). Citronella (Icacin.) is in Malesia, eastern Australia, New Caledonia, Fiji/Samoa/Tonga, central Chile (30-38ºS), Brazil, north to Ecuador (Guayaquil), and disjunct in Panama/Costa Rica (de Vogel, 1975).

In eudicots, the family Oncothecaceae, monotypic in New Caledonia, and the Metteniusaceae of Costa Rica to Ecuador, mainly in Colombia, form the first and second basal branches (a trans-Pacific sequence) in a cosmopolitan clade of Lamiales, Solanales etc. or alternatively are sisters (a trans-Pacific clade) (González & Rudall, 2007, González et al., 2007).

The legumes include at least two examples (Lewis et al., 2005). Castanospermum of north-eastern Australia, New Caledonia, and Vanuatu is sister to Alexa of Venezuela, the Guianas, and Amazonian Brazil. Schleinitzia, known from the Philippines to New Caledonia and Society Is. (including the Mariana Is. but not Australia), is in a well-supported clade with Kanaloa of Hawaii (Kaho’olawe) and Desmanthus, in warm America and the Caribbean.

In Proteaceae, Sleumerodendron of New Caledonia is sister to Euplassa with 20 species of tropical South America (Willis, 2007; cf. Virot in Aubréville et al., 1967-). Kermadecia of New Caledonia and Turrillia of Vanuatu and Fiji may also belong here (cf. Weston & Barker, 2006) but were not sampled by Willis; their exact position does not affect the trans-tropical Pacific disjunct track in Sleumerodendron/Euplassa and they may even belong to it.

In the diverse, pantropical genus Diospyros (Eben.), ‘clade Q’ of Duangjai et al. (2006; equivalent to clade XI of Duangjai et al., 2009) has a trans-tropical Pacific distribution in south-east Asia, New Caledonia, and tropical America including the Caribbean.

Lagenophora (Compos.) is recorded from India to New Zealand, New Caledonia, Hawaii, Juan Fernandez Is./southern Chile/Falkland Is., then disjunct at southern Guatemala, Panama and north-western Venezuela (near Tachira) (van Balgooy, 1975).

Nothofagus subgen. Brassospora is extant only in New Caledonia – New Guinea, where it is abundant and diverse, and the only member of Nothofaus present. There is also fossil material of Brassospora in New Zealand, Australia and Antarctica, but this shows less diversity than the extant group (Heads, 2006). Its sister group is subgen. Nothofagus, extant on South America (the main massing of all species is South America) and with fossils in Australia.

Stereocaryum (Myrt.) is in New Caledonia and the Malay Peninsula (Scott, 1980) and represents an ‘odd’, ‘unexplained’ and ‘phytogeographically enigmatic’ distribution (Johnson & Briggs, 1984). The genus is related to the very large, mainly South American Eugenia, although it ‘can hardly be a late derivative of Eugenia itself’ (Johnson & Briggs, 1984). Stereocaryum has also been compared with the South American – West Indian Calycorectes (Briggs & Johnson, 1979). Craven (2001) suggested that Stereocaryum belongs in Eugenia s. str., which is centred in Central and South America but also has rich representation in New Caledonia (36 species) and Sri Lanka (22 species), with lesser concentrations elsewhere in the Old World.

Rhizophora mangle (Rhizophor.) is in New Caledonia, Fiji/Samoa/Tonga, Galapagos, Central America (west coast) and West Africa (Ding Hou & van Steenis, 1963). Halophila decipiens var. pubescens (Hydrocharit.) is in New Caledonia, Tahiti and the Caribbean (and Sri Lanka). It is replaced by the typical variety in Australia, Malesia and Asia (den Hartog, 1966).
Invertebrates

In millipedes, Jeekel (1986) recognised that Agathodesmus of New South Wales, Queensland and New Caledonia (Mesibov, 2009) includes a species from Jamaica. Some authors had suggested human transport to New Caledonia and Jamaica, but the New Caledonian species is endemic (Mesibov, 2009) and Hoffmann (1999) reported a second Jamaican species in the genus. Hoffmann wrote ‘while a multiple transport of rare and localised species from the Antipodes to a single West Indian island is not impossible, it does appear improbable’. The New Caledonia – Jamaica connection is instead related here to the Pacific history of the Caribbean plate and the large igneous plateaus in the central Pacific.

In spiders, Aucana (Pholcidae) is in New Caledonia and central Chile (Antofagasta to Bío-Bío) (Huber, 2000).

In Trichoptera, the tribe Grumichellini comprises Triplexa of southeastern Australia, Gracilipsodes of New Caledonia, and Atanatolica, Grumichella and Amazonatolica of tropical America: north-westernmost Argentina (Jujuy) and southeastern Brazil north to Costa Rica, and in the Lesser Antilles (Dominica) (Holzenthal & Oliveira Pes, 2004; Malm & Johanson, 2008).

The scale insect Diaspis casuarinae is endemic to New Caledonia and is an ‘unusual’ species; it ‘bears some relationship’ with D. chilensis of Chile (Williams & Watson, 1988; www.sel.barc.usda.gov/scalenet/scalenet.htm). In Lepidoptera, Holloway (1993) cited a possible South American connection between the New Caledonian sphingid Compsulyx and the neotropical Adhemarius.

In chrysomelid beetles Zira of New Caledonia is closer to Lioplacis and other Brazilian genera than to any Australia/New Zealand taxa (P. Jolivet, pers. comm. 17-xi-2006).


Vertebrates

The Meiolaniidae are extinct turtles up to 3m long with cranial horns and frills and a club tail. These bizarre, fossil, marine animals show a trans-tropical Pacific distribution similar to that of the extant, terrestrial plants cited above. Meiolania is known from the Miocene and Pleistocene of the Northern Territory, eastern Australia, Lord Howe I. (where species survived until 2000 years ago), and New Caledonia, including the Loyalty Is. and Walpole I. (south-east off Grande Terre). The rest of the family comprises two other genera restricted to Queensland and one genus from the Eocene of Argentina (Gaffney, 1996). The Walpole I. specimens seem to belong to a locally endemic form and Gaffney suggested the family represented a relatively archaic Pacific biota that survived on young oceanic islands. He suggested that this took place by the process of biogeographic ‘hopskotch’ (McKenna, 1983), in which populations on volcanic islands survive in situ by colonising nearby new islands as the older ones erode and subside. The process is probably fundamental for Pacific biogeography.

In snakes, the subfamily Boinae occurs in Madagascar (2 genera), Mauritius (2 genera), Melanesia (Candoia, formerly Enygrus) and tropical America (4 genera). Candoia ranges in Sulawesi, Palau, New Guinea, Bismarck Archipelago, Solomon Is., Vanuatu, New Caledonia (Loyalty Is. only) and Fiji/Samoa/Tonga. Burbrink (2005) found Candoia to be the sister group of the Madagascar genera and this whole clade sister to the American genera, giving either trans-Atlantic connections, trans-Pacific connections, or both. A trans-tropical Pacific link would involve the usual New Caledonia/Fiji – central America/Antilles disjunction. In any case, Burbrink (2005) regarded the group as relictual and the biogeographic pattern derived from distributions established prior to Gondwana breakup.

The skink Emoia cyanura also has a trans-tropical Pacific range, occurring on the Loyalty Is. (not Grande Terre), Vanuatu, Solomon Is. and Bismarck Archipelago east to Polynesia and Peru (Bauer & Sadlier, 2000). Three birds, each usually treated in its own family, are often regarded as related: Rhynochetos of New Caledonia, its sister Aptornis (fossil) of New Zealand, and the sister of these two, the sunbittern Eurypyga of tropical America (Guatemala – Brazil). Livezey (1998) found a sister relationship between a clade comprising Rhynochetos and Aptornis and one comprising Eurypyga and Messelornis of the Eocene of the north Atlantic (Wyoming and Germany), indicating both trans-Pacific and trans-Atlantic links.


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