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Home , Ardei, Caloenas, Casuariidae, Collocalia, Dromornis, Dromornithidae

https://doi.org/10.24199/j.mmv.1975.36.05 27 May 1975

ANTARCTIC DISPERSAL ROUTES, WANDERING CONTINENTS, AND THE ORIGIN OF 'S NON-PASSER1FORM AVIFAUNA

By Pat Vickers Rich *

* The Museum, Tech University, Lubbock, Texas, U.S.A. 79409; present temporary address, The National Museum of Victoria, Russell Street, Melbourne, Victoria.

Introduction passeriform families. In this way, I hope to clarify the certainties and uncertainties that In 1858, P. L. Sclater recognized Australia accompany such determinations for each family, on the basis of its living avifauna, as a unique and to estimate which dispersal route (Antarctic biogeographic unit, distinct from the Oriental or Indomalaysian) seems most probable for fauna that characterized the Asian mainland. initial dispersal of each non-passeriform family These two faunas complexly intermingle on the between Australia and the remaining world. islands of the Indonesian archipelago, a situa- In order to evaluate the availability of the two tion reflected not only by the but by other routes for avian dispersal throughout the vertebrate and invertebrate groups as well. Mesozoic and Cenozoic, I have summarized: A. R. Wallace in a number of papers (1863, the timing of break-up and ( 1 ) data regarding 1869, 1876) initially described the avifaunas separation of those continental plates closely encountered in this transitional zone; his he associated with Australia during that time work was followed by numerous refinements period; (2) paleoclimatological data available (Stresemann, 1927-34, 1939-41; Rensch, for the Antarctic and Indomalaysian dispersal culminating with those which led Mayr 1931) routes, as well as for Australia during post- 1944a-b, 1945a-b, 1972) to conclude (1941, Paleozoic time; and (3) data available on major part of Australia's avifauna was that the phylogenetic relationships, world-wide diversity, Southeast Asia. Although Darling- derived from endemism, distribution, and palaeontological ton (1957) agreed with this thesis and Gentilli record of all the non-passeriform avian families hinted at a dual origin, Serventy ( 1972, ( 1949) having a Cenozoic record in Australia. 1973) was the first to seriously consider multiple geographic origins of the Australian Dispersal Routes To and From Australia avifauna in light of the accumulating geological evidence suggestive of dynamic, not static, Geological Evidence relationships of the continents during the Meso- Antarctic Dispersal Route (Operative, zoic and Cenozoic. Australia had not always Mesozoic arid early Cenozoic) been a neighbour of Asia. Serventy concluded, data collected primarily in the however, that excepting a few Australian Geophysical strongly support the idea that a families which might have used an Antarctic last decade dispersal route between Australia and dispersal route, Mayr was correct in assuming southern certain other Gondwana continents was present that Southeast Asia had spawned most of the of the Mesozoic, and between Australasian avifauna. Cracraft (1972, 1973) during much Australia and some southern continents as late strongly supported an austral dispersal route as the early Tertiary (Allan, 1969; Creer, for the , , and possibly some Francheteau and Sclater, 1969; Smith and suboscine passeriforms but other- 1970; and Hallam, 1970; and others). Such a route wise subscribed to the thesis of northern origin was most likely of an archipelagic nature for most other avian groups. (similar to the linkage between Asia and In the following paper, I wish to investigate that Australia today) as geologic data suggest that the initial assumptions and the reasoning complete terrestrial continuity probably has have been used in previous studies to determine not existed between South America, West the source area for each of the Australian non-

63 64 PAT VICKERS RICH

Antarctica, East , 1 and New Zealand of geologic evidence. Despite the controversies

(Tedford, 1973a; Jardine and McKenzie, that still attend the Africa-Madagascar, Mada- 1972) during the mid to late Mesozoic and gascar-India, Africa-India, Australia-India, Cenozoic. East-West Antarctica continental pairs, general Time of the break-up of the many contin- agreement exists among most supporters of ental segments, originally part of Gondwana- plate tectonic theory on the time of fragmenta- land, can be estimated by using several types tion of many of Gondwanaland's several seg- ments. Such fragmentation did not occur 1 Present-day Antarctica can no longer be con- simultaneously, however, and original rupture sidered a single, terrestrially continuous unit, but is between continent-pairs have begun long instead composed of at least two segments, East and may West Antarctica, each with a distinct geologic history before actual terrestrial continuity between land (Hamilton, 1964, 1967; Dalziel and Elliot, 1971, masses was lost (e.g. East Africa at present). 1973; Elliot, 1972). At present West Antarctica is Thus, a distinction needs to be made between comprised of a number of islands separated from the continental East Antarctica by basins with sub-sea evidence indicating initial rifting or break-up level depths. (B in Table 1), and that indicative of active

TABLE 1

Evidence available for timing the break-up and dispersal of Gondwanaland during the Mesozoic and Cenozoic (see Figure 1 for summary). B, break-up occurring (involving displacements of less than 100 km.); S, large-scale separation of continental masses underway.

Continental Pair Time of Break-up /Dispersal Evidence SOUTH AMERICA- Post- to Polar wander curves. AFRICA mid- (7, 13, 14, 17, 37, 38) B Aptian (27, 39) Salt deposits in coastal West Africa and coastal Brazil. B 106-136 m.y. (late - Occurrence of massive tholeiitic basalt early Cretaceous) (7, 8) flows dated by K/ A method on both sides of Atlantic; Serra Geral Fm. of Brazil; Kaoko volcanics of Southwest Africa; Lupata volcanics of Africa. S 70-75 m.y. (26, 29, 46) Magnetic banding (oldest anomaly, 31). B Post early Cretaceous Non-marine ostracod faunas virtually (10) identical (at specific level) in Brazil and West Africa. S Albian (mid-Cretaceous) Fully marine conditions indicated by (10, 18, 28) sediments over broad areas on either side of South Atlantic. S Late Turonian Fully marine conditions along western (12, 27, 28) Africa. Ammonite faunas of North and South Atlantic similar. SOUTH AMERICA- S > 20-25 m.y. but WEST > Magnetic banding in the Drake Passage ANTARCTICA late Mesozoic (22) for minimum date; similarity of tecto- (probably archipelagic nostratigraphic provinces until end of during Jurassic to recent) Andean Orogeny (-early Tertiary) for maximum date. S Post-Cretaceous (24) Paleomagnetism of volcanics and intru- sives on indicate major bending of Peninsula occurred post-Cretaceous.

WEST ANTARCTICA- S 80 m.y. NEW ZEALAND (4, 5, 23, 26) Magnetic banding between Campbell (late Cretaceous) Plateau and West Antarctica; oldest (probably archipelagic, anomaly, 32. Jurassic to Recent) ANTARCTIC DISPERSAL ROUTES 65

Continental Pair Time of Break-up/ Dispersal Evidence 78 m.y. (19) Beginning of activity on Endeavour Fault (late Cretaceous) (fault active between 47-78 m.y.). Mid-Jurassic (3, 10) Volcanics. Tasmanian dolerites, Ferrar dolerites (Antarctica).

4. EAST ANTARCTICA- Mid-Cretaceous-Tertiary Polar wander curves. AUSTRALIA (3, 13) (probably archipelagic Jurassic to Recent) S 45-50 m.y. (1, 2, 4, 6. 10, Magnetic banding between Australia and 14, 16, 19, 26) East Antarctica; oldest anomaly, 18. S Mid-late (1, 9, 10, Marine transgression in southern Aus- 30) tralia; open marine limestones. S Late (33, 36) Erosional unconformity (marine ) evi- dencing establishment of circum-Antarctic current. B Mid-Jurassic (42) Dolerites in and East Ant- arctica. B Late Jurassic to Rifting in southern Australia. Cenomanian (43)

5. EAST ANTARCTICA- B Triassic-Jurassic (11) Onset of rifting, southern Mozambique. AFRICA B Mid-Triassic to mid- Volcanics. Extrusion of Stormberg, Jurassic Karoo, Swaziland basalts. Intrusives in 200-155 m.y. (8, 10, 20) Rhodesia, South America. S Post mid-Jurassic- Polar wander curves. pre mid-Cretaceous (13) S Pre late Cretaceous Magnetic banding between Africa and 140 m.y. (2, 12) East Antarctica. S Pre Neocomian (15) Marine sediments. (early Cretaceous) S Pre Valanginian (10) Marine sediments. (early Cretaceous)

Polar wander curves. 6. EAST ANTARCTICA- S Mid-Jurassic-mid- INDIA Cretaceous (13, 14, 21) S Pre-Cenomanian (10) Marine sediments between Antarctica and (late Cretaceous) India. B Albian, 100-105 m.y. Volcanics. Extrusion of Rajmahal Traps. (early Cretaceous) (14, 21)

AFRICA-MADAGASCAR S Mid-Jurassic- Polar wander curves. (probably archipelagic mid-Cretaceous (14) Cretaceous to Recent) B ? (35) Rifting. B Early Jurassic (32) Volcanics. S Jurassic-late Cretaceous Volcanics, rifting, sediments (by Turo- marine conditions between (31); pre early Cretaceous nian open (45) Madagascar and Africa). B Cretaceous (11) Rifting in East Africa. S Late Mesozoic (10) Faunal. present in East Africa and S Late Cretaceous (10) Volcanics Madagascar. sediments in Mozambique S Late Eocene (34) Marine Channel.

mid-Cretaceous (14) Polar wander curves for India and 8. INDIA-MADAGASCAR S Post Madagascar. oldest anomaly, 30. S 70-75 m.y. (41) Magnetic anomalies; between India and S Campanian (10) Marine sediments (late Cretaceous) Madagascar. B Late Cretaceous-early Volcanics. Deccan Traps of western Tertiary, 59-65 m.y. India. (10, 14, 21, 40) faunas (see over). S Post late Cretaceous (10) Marine 66 PAT VICKERS RICH

Continental Pair Time of Break-up /Dispersal Evidence 9. INDIA/MADAGASCAR- S Mid-Jurassic- Polar wander curves. AFRICA mid-Cretaceous (10) S 70-75 m.y. (41) Magnetic anomalies; oldest anomaly, 30. S Pre latest Cretaceous, Magnetic banding in NW sector of Indian 67 m.y. (15) Ocean. S 40 m.y. (26) Magnetic banding. S Post late Cretaceous Faunas (marine invertebrates) of Coro- (10) mandel Coast of India quite similar to those of Assam and South Africa in late Cretaceous but distinct from those from Narbada Valley in NW India. Faunas of NW India most similar to those of North Africa and Europe. Evidence suggests non-marine barrier between northern and southern Africa, eastern and western India.

10. AUSTRALIA-INDIA B Late Jurassic-early Volcanics. Rajmahal Traps (NE India),

Cretaceous, 100-105 m.y. Banbury ( Perth Basin ) and Ashmore (9, 12, 14, 15) Reef (Aust.) basalts, Broom Beds (tuffs, Canning Basin, Aust.). S Early Cretaceous Marine sediments. First occurrence of (9, 12, 15) marine sediments (during Mesozoic) in Perth Basin. Thick sequence of non- marine sediments as young as Jurassic in subsurface, southern Perth Basin. S Pre early Tertiary Sediments on either side of Ninetyeast (3) Ridge in Indian Ocean 'show a unified history and little or no relative motion since early Tertiary'. S Pre mid to upper Submarine erosion occurring in Tasman Oligocene (33, 36) sea evidenced by thinness or absence of Oligocene sediments in this area and the scoured oceanic surface below such sediments; mid to upper Oligocene sedi- ments, when, present often intensely current-bedded, suggesting presence of submarine current. B Latest - Rifting in . earliest Cretaceous (43) B Permian-Neocomian (42) Vulcanism, Banbury Basalt, latest Neo- comian tholeiite. S Late Jurassic (in NW Magnetic banding in Indian Ocean and part of Australia) to initiation of marine transgression in late Neocomianw\ptian Western Australia. (in SW Australia) (43) S Pre latest Jurassic Sediments (oldest) overlying oceanic (Tithonian) (44) crust near Western Australian coast. 11. AUSTRALIA- S 80 m.y. NEW ZEALAND (4) Magnetic banding between Campbell Plateau and W. Antarctica, Australia, (probably archipelagic and E. Antarctica indicates New Zealand Cretaceous to Recent) moving north while Australia still attached to E. Antarctica. -Eocene Volcanics. Beginning of significant vol- 51 m.y. (3) canic activity in eastern Australia that continue through much of the Tertiary. ANTARCTIC DISPERSAL ROUTES 67

The following papers are referred to in Table 1 occurrence of magnetic anomalies in the inter- under their respective numbers: vening ocean basins, (3) the first differences in 1. Jones, 1971 2. LePichon and Heirtzler, 1968 the polar wander paths of segments previously 3. Wellman, McElhinny, and McDougall, 1969 part of one continental plate, and (4) differ- 4. Griffiths and Varne, 1972 ences in the tectonostratigraphic provinces of 5. Pitman, Herron and Heirtzler, 1968 6. Weissei and Hayes, 1971 two previously geologically similar and con- 7. Creer, 1970 tiguous continental plates (see Smith and 8. Dietz and Holden, 1970 Hallam, 1970; and particularly Tarling and 9. Veevers, 1971 10. Smith and Hallam, 1970 Tarling, 1971 for more detail on the dating 11. Sowerbutts, 1972 methods mentioned above). Such data evidenc- 12. Tarling, 1971 ing break-up and dispersal of the Gondwana 13. Francheteau and Sclater, 1969 14. McElhinny, 1970 continents since the Triassic are summarized 15. Veevers, Jones, and Talent. 1971 in Table 1 and Figure 1 for each continental 16. McElhinny and Wellman, 1969 17. Vilas and Valencio, 1970 pair, except East Antarctica-West Antarctica 18. Burke, Dessauvagie, and Whiteman, 1971 (see discussion below). Figure 1 specifically 19. Christoffel and Ross, 1970 indicates the variety of the data used to esti- 20. Dietz and Sproll, 1970 21. McDougall and McElhinny, 1970 mate the time of break-up of each continental 22. Dalziel and Elliot, 1971 pair, while Table 1 details that data, indicating 23. Griffiths, 1971 whether break-up or active dispersal was 24. Hamilton, 1964 25. Maxwell, et al, 1970 underway. 26. Heirtzler, et al, 1968 Fragmentation within Gondwanaland began 27. Reyment, 1969 as early as the Triassic, but actual severance in 28. Funnell and Smith, 1968 29. Dickson, Pitman, and Heirtzler, 1968 terrestrial continuity between those parts of the 30. McGowran, 1971 (ms) southern continents connected in the late 31. von der Borch, 1972 32. Flores, 1970 Palaeozoic (approx. 225 m.y. B.P.) was a late 33. Kennett, et al, 1972 Mesozoic and even early Tertiary phenomenon 34. Simpson, et al, 1972 (approx. 70-130 m.y. B.P.; see figures 1-2, 35. Kent, 1972 36. Carter and Landis, 1972 7-10, and Table 1). The first complete separa- 37. Creer, Embleton, and Valencio, 1969 tion was apparently between Africa and East 38. Larson and LaFountain, 1970 Antarctica in the late Jurassic or early Creta- 39. Leyden, Bryan, and Ewing, 1972 40. Wellman and McElhinny, 1970 ceous (Heirtzler, et a/., 1968, Francheteau and 41. McKenzie and Sclater. 1971 Sclater, 1969; Dietz and Sproll, 1970; Smith 42. Harrington, et al, 1973 and Hallam, 1970; Dietz and Holden, 1970; 43. Veevers and Evans, 1973 44. Heirtzler, et al, 1973 Dingle and Klinger, 1971; Heirtzler and Bur- 45. Green, 1972 roughs, 1971; Tarling, 1971; Sowerbutts, 1972; 46. Valencio and Vilas, 1972 47. Jones, 1972 Jones, 1972; Dingle, 1973). Although not so 48. Dingle, 1973 well understood, the severance of any close alliance between India and Australia probably occurred in the early Cretaceous (Heirtzler, et al, 1968; Veevers, 1971; Tarling, 1971; separation of continental masses (S in Table 1) Veevers, Jones, and Talent, 1971; Crawford, 2 with concomitant loss of non-marine dispersal 1971 ), certainly not later than late Mesozoic pathways. Widespread occurrence of vulcanism (von der Borch, et aL, 1972). Thus the shortest and extensional faulting, sometimes accom- dispersal route between Australia and the re- panied by evaporite deposits, can be the first maining continents during the late Mesozoic indication of continental break-up. Similarly, and much of the was across Antarc- active separation between continental blocks is 2 Crawford argues that during the Phanerozoic evidenced by: (1) the first widespread occur- India was closely apposed to Africa with a seaway rence of marine sediments after a long period separating India and Australia. Just how broad such of continental sedimentation, (2) the first a water gap would have been is uncertain at present. 68 PAT VICKERS RICH

SOUTH AMERICA

Fig. 1 —Evidence available for timing the post-Paleo- at this time to South America, as to New dispersal Gondwanaland. zoic break-up and of Zealand, appears to have been archipelagic. The following symbols were used: m, mag- Similarly Antarctican archipelago may netic anomalies on the seafloor; p, polar a West wander curves; s, widespread occurrence of have lain between South America and New marine sediments after long period of non- Zealand (Fleming, 1962; Tedford, 1973a; marine sedimentation and other sedimento- logical data; v, occurrence of widespread Elliot, 1972). Yet unresolved and critical to vulcanism; f, faunal (both marine and ter- late Mesozoic and Cenozoic reconstructions, restrial) similarities and differences; r, onset however, is the precise relationship of West of major extensional rifting; t, similarity of tectonostratigraphic provinces; st, occurrence Antarctica to South America and East Antarc- of widespread salt deposits. tica. Many workers suggest that during much of the Mesozoic West Antarctica and New tica. Asia, at this time, was as much as 30-50° Zealand were all part of one plate's compres- north of Australia (Dietz and Holden, 1970; sive margin (Elliot, in press, 1972), and must Tedford, 1973a; Jardine and McKenzie, 1972; have developed in close proximity to a con- Raven and Axelrod, 1972). tinental sediment source, East Antarctica and In the late Mesozoic (see figure 4), East Australia (van der Linden, 1969), with the Antarctica and Australia seemingly formed a southern Andean Cordillera and the Antarctic continuous land mass (Jones, 1971; Le Pichon Peninsula, although not necessarily a linear and Heirtzler, 1968; Smith and Hallam, 1970; chain (Dalziel and Elliot, 1971, 1973), forming McElhinny, 1970; Heirtzler, et al., 1968; a link between these two continents. Severe Veevers, 1971; McGowran, 1971). Connection disruption of this perhaps already archipelagic ANTARCTIC DISPERSAL ROUTES 69

c«ou» INDIA Of*

AFRICA

SOUTH AMERICA

$200myBP early Cretaceous x but pre-Valanginian *

AUSTRAUA OLlfloe< EAST nearly Eocene ANTARCTICA 50-55myBP

Probably archipelagic (post-Paleozoic) /80my6P NEW ZEALAND

Fig. 2—Estimate of time of most recent severence and that deformation of the Scotia Arc occurred of land connections between presently distinct prior to the late Mesozoic (Katz, 1973). Elliot continental masses, once part of Gondwana- land. (pers. comm., 1974) has pointed out that this does not necessarily mean that insular linkage cordillera may have begun prior to 82 m.y. B.P. was lacking between the two areas during the (Dalziel and Elliot, 1971, 1973) with transform Mesozoic and part of the Cenozoic, however. or transcurrent faulting rotating the Ellsworth More general agreement attends the positioning Mountains (now situated between East Antarc- of New Zealand decidedly closer to Marie Byrd tica and the Antarctic Peninsula) (Dalziel and Land in West Antarctica until about 80 million Elliot, 1971, 1973; Elliot, 1972; Schopf, years ago, when it apparently began its north- 1969). During the late Mesozoic and early ward drift with the foundering of the Campbell Cenozoic, South America and West Antarctica Plateau and opening of the Tasman Sea may have approached one another more closely (Griffiths, 1971; Griffiths and Varne, 1972). than at present (Dalziel and Elliot, 1973). The opinion of most geologists dealing with Some workers, however, suggest that East and these areas, however, despite the differing West Antarctica were independent entities dur- details of their reconstructions, is that any ing the Mesozoic, becoming closely apposed dispersal route involving West Antarctica from only in the Cenozoic (Beck, 1972). Others mid-Mesozoic to the Recent would probably further suggest that the post-Jurassic tectonic have been archipelagic (Elliot, 1972; Jardine histories of southern South America and West and McKenzie, 1972; Dalziel, pers. comm., Antarctica have been decidedly independent 1973). 70 PAT VICKERS RICH

late Fig. 3—Continental arrangement during the LATE JURASSIC 135 my. Jurassic (135 m.y.) as modified from Ted- ford, 1973a. In such reconstructions, the earth's surface is divided into a series of crustal plates that move relative to each other. Three types of boundaries separate these plates: (1) transform faults (light lines) where the plates slide laterally past one another; (2) ridges (dark lines), where new molten rock is added to the edges of two plates and they subsequently move away from one another; and (3) trenches (hatched lines), where one plate slides beneath an- other. Arrows indicate direction of relative motion.

Fig. 4—Continental arrangement during the late Cretaceous (65 m.y.) as modified from Tedford, 1973a. ANTARCTIC DISPERSAL ROUTES 71

Fig. 5—Continental arrangement during the Oligo- cene (35 m.y.) as modified from Tedford, 1973a. Fig. 6—Continental arrangement at present. 1

72 PAT VICKERS RICH

Only in the Tertiary, the early to mid-Eocene Throughout the Cenozoic the area between (50-55 m.y. B.P.), and certainly by the late Southeast Asia and Australia has probably been Oligocene (Kennett, et aL, 1972), did Australia dotted with islands, may of which have been sever its connection with East Antarctica (Le ephemeral, occasionally disappearing down along with Pichon and Heirtzler, 1 968; Jones, 1 97 oceanic trenches or submerging Veevers, 1971; Smith and Hallam, 1970 older parts of ocean basins. In some senses a Davies and Smith, 1971; McGowran, 1971 dispersal route of sorts probably existed across Vogt and Conolly, 1971; Griffiths and Varne, this archipelago in the Paleogene, but distances 1972; Weissel and Hayes, 1972) and begin its between islands as well as the total length of northward journey, which is still in progress the route were much greater than at present (see figure 9). From a position of 60-70° south and prohibitive in all but the rarest cases of latitude (measured from Canberra) in the Cre- non-marine vertebrate dispersal. Most of central taceous, Australia moved to within 10° of its and southern New Guinea, the leading edge of present position (see figure 10) by the the Australian plate in the Paleogene, was (Wellman, McElhinny, and McDougall, 1969; underwater until the late Oligocene-early Mio- Veevers, Jones, and Talent, 1971; Raven and cene (Raven and Axelrod, 1972; Davies and Axelrod, 1972). New Zealand continued separ- Smith, 1971; Tcdford, 1973a; Audley-Charles ating from both Australia and West Antarctica, et aL, 1972). During the Miocene, a southward increasing the isolation that presently charac- migrating island arc system collided with the terizes it. Thus, during the late Mesozoic and northern edge of the Australian crustal plate early Tertiary, the separation of Australia and (sec figure 7 for explanation), adding the final, New Zealand from their present Asian and most northern segment to form modern New Guinea Indonesian neighbours was great with close (Hamilton, 1972b) and causing considerable apposition occurring only after the Oligocene uplift. By late Cenozoic, therefore, extensive (see Hamilton, 1972a-b; Haile and McElhinny, emergent lowland areas and developing high- 1972; Stauffer and Gobbett, 1972; contra Ridd, lands were present in New Guinea. In the Indo- 1971, and Audlcy-Charles et al, 1972). malaysian area (extremely complex geologi- cally) much of the Malay Peninsula, and at least parts of Sumatra, Java, Borneo, and Sula- Indonesian Dispersal Route (Operative, wesi were emergent during the early Cenozoic Late Cenozoic) (Grabau, 1926; van Bemmelen, 1949; Kum- The mid-Cenozoic close approach of Aus- mel, 1961). Major parts of the Andaman- tralia to Asia (including Southeast Asia, see Sumatra-Java-Timor-Outer Banda-Seram chain Haile and McElhinny, 1972 and closely ap- of islands, however, are composed of uplifted posed Indonesia, see Ben-Avraham and Uyeda, subduction melanges (a jumble of sediments,

1 1 973 ; Hamilton, 972a-b ) , presented new volcanics, and metamorphics typifying an area possibilities for dispersal to and from Palae- where one major plate of the earth's crust is arctica. With Antarctica's movement south of underthrusting another) produced during the the Australo-Antarctic Ridge and concomittant Miocene and uplifted sometime later (Hamil- deterioration in its climate accompanying posi- ton, 1972a-b; Katili, 1971). During the late tioning closer to the pole, it was further isolated , folding further uplifted many of the from Australia. Thus, its importance as a dis- previously homogeneous lowland areas in both persal route declined as that of Indonesia in- Indonesia and New Guinea (Kummel, 1961), creased. By the Miocene, the principal dispersal providing a mountainous, temperate dispersal route for terrestrial organisms to and from route (Raven and Axelrod, 1972) between Australia was the Indomalaysian Archipelago Asia and Australia, probably for the first time (McKenna, 1973), including the Philippines, in the Mesozoic-Cenozoic. Further emergence much of which was subaerial by this time during the concurrent with develop- (Hamilton, 1972a; van Bemmelen, 1949; ment of ice sheets elsewhere in the world, Grabau, 1926). shortened water gaps, and made the Indo- ANTARCTIC DISPERSAL ROUTES 73 malayan area a very probable route for dicotyledons in these floras have non-entire dispersal of terrestrial organisms. margins and small leaves, more typically reminiscent of temperate plant living in mild to cool conditions at present (Axelrod Paleoclimatological Evidence and Bailey, 1968). Again, the Mora of East

Antarctica is not as well known, with only two Antarctic Dispersal Route lower Tertiary localities: one in the McMurdo Based on palcobotanical data, the climatic Sound area (Cranwell et al., 1960; Mclntyre regime of the Antarctic Peninsula and East and Wilson, 1966) and a second near Prydz Antarctica was much less severe during the late Bay (Kemp, 1972a-b) on the Australian side Mesozoic and early Tertiary than at present. of East Antarctica. Much of the pollen from During the Jurassic, scouring rushes (Equise- these localities is similar to that from West tales), ferns (Filicales), cycadophytes (Cyca- Antarctica. The McMurdo sample derived from dophyta), and conifers are known from several Eocene erratic boulders and recycled sediments localities in Grahamland, West Antarctica in the Ross Sea includes conifer, palm, Poda- (Plumstead, 1962, 1964; Bibby, 1966). Un- carpus, and Nothofagus pollen, among others. fortunately, only one post-Triassic, pre-Tertiary Probable in situ late Oligocenc sediments in the flora is known from East Antarctica (Victoria Ross Sea have recently produced many of the

Land); it contains a cycadophyte as well as same elements with Nothofagus and Podo- three types of conifers very similar to those carpaceae dominant and minor percentages of known from India, East Africa, and Australia Proteaceae and Myrtaceae. Noticeably absent in strata of early Jurassic age. Such assemblages is pollen of groups presently found on sub- in both East and West Antarctica suggest antarctic islands such as Gramineae, Cypera- equable, moist conditions, warmer than at ceae, Umbelliferae, or Compositae (E. Kemp., present, particularly in order to accommodate pers. comm., 1974). The Prydz Bay pollen flora the cycadophytes, which were experiencing derived from Cretaceous-Eocene sediments is latitudinal restriction in the Northern Hemi- decidedly more varied, containing at least 21 sphere during the Jurassic (Plumstead, 1964). genera including Nothofagus (fusca, menziesii,

, Podocarpidites, Gleicheniiditcs, The presence of freshwater actinopterygian fish and brassi ) (Filicales) in lower Jurassic sediments of East Antarctica Osmundacites, and Cyanthidites (Schaeffcr, 1972) lends corroborative evidence reminiscent of dominant taxa in the early Certainly as diverse for more equable, certainly non-glacial Meso- Tertiary Australian flora. incompatible with a zoic conditions in parts of Antarctica. Mega a flora as this seems and microfloral evidence indicates that at least continent-wide glaciation prior to the Oligocene. floras from East and West two families of ferns (Cyatheaceae and Schiz- In summary, admittedly rare, suggest aeaceae), several conifers (Araucaria, Agathis, Antarctica, although Podacarpus) (Andrews, 1961) and angio- a temperate climate for parts of this region sizeable trees and sperms including the Southern Beech (Notho- capable of supporting a diverse assemblage of plants during much of fagtis), mustards (Cruciferae), Myrtaceae, possibly as late as the Miocene Proteaceae, Loranthaceae, and possibly Drimys the Mesozoic, Quaternary on the Falkland (Winteraceae) were growing on Seymour Island and even the Islands, which support no forests today (Axel- in West Antarctica during the Cretaceous of this same flora that (Cranwell, 1959). Likewise, floras very tenta- rod, 1960). Much occurred in Antarctica was also present in tively dated as early to mid-Tertiary from King America, Kerguelen Island, George and Seymour Islands, also in West southern South Zealand, and parts of Australia (Axelrod, Antarctica, contain scouring rushes, ferns, New as well as a 1960) during the early to mid-Tertiary. This conifers (including Araucaria) , flora was apparently rather similar to the variety of angiosperms, including Nothofagus temperate and cool temperate rainforests of (Adie, 1953; Cranwell, 1959; Plumstead, present-day southern Chile, Patagonia, south- 1964). Approximately ninety per cent of the 74 PAT VICKERS RICH eastern Australia, Tasmania, New Zealand, and "consolidated glacial deposits and extensive New Caledonia. areas of glacially scoured bedrock" (Newell, Additional information of paleoclimatic signi- 1972, p. 64). Even though much of West ficance has been derived from geologic and Antarctica (particularly the Antarctic Penin- fossil invertebrate data. Recent electron micro- sula) supported Nothofagus dominated forests graph studies of sand grain surface features perhaps as late as the Miocene, giaciation may based on submarine core samples from the have been underway in limited areas as early Pacific, has been interpreted to as the Eocene. The evidence for widespread indicate that giaciation, although probably of early Tertiary giaciation, however, based on limited extent, was underway in parts of Ant- such sedimentological and volcanic data has arctica as early as the Eocene (Geitzenauer recently been challenged by Mercer (1973), et ai, 1968; Gram, 1969; Margolis and Ken- who believes the botanical evidence for both nett, 1970, 1971a-b; Kcnnett and Brunner, segments of Antarctica suggest nothing more 1973). Apparently glacially transported sand than local giaciation until 3-4-6 m.y. B.P. Cer- grains were raited away from the Antarctic as tainly by 4-5 m.y. B.P. giaciation on a contin- part of icebergs and deposited as the ice melted. ental scale is suggested by widespread erosion Accompanying the first occurrence of such surfaces over much of the Ross Sea continental sands, Mandra (1969, 1971; also in Mandra shelf, evidently produced by a grounded ice and Mandra, 1970) noticed a drop in silico- shelf (Hayes, et al, 1973), but glacial history flagellate species diversity, which was inter- of Antarctica prior to this time has yet to be preted as indicative of cooling of the southern resolved. oceans at that time. In the early and middle Unfortunately, owing to the lack of post-

Oligocene cores, ice rafted sand is no longer Jurassic paleomagnetic data from East Antarc-

a dominant feature, if present at all, and silico- tica (primarily because of the lack of post-

flagellate diversity in such sediments is higher Jurassic rocks cropping out above the present than in immediately older sediments, indicat- ice cover, as well as the paucity of samples ing warmer seas than in the earlier Tertiary, even from West Antarctica) there is as yet no perhaps owing to the establishment of the coherent picture of the South Pole position circum-Antarctic current at about this time. with respect to Antarctica during the late Additional geologic evidence mustered to Mesozoic and early Tertiary. A number of suggest an Eocene giaciation in West Antarctica papers (Francheteau and Sclater, 1969; McEl- has been recovered from Marie Byrd Land. hinny and Wellman, 1969; Crawford, 1971; LcMasuricr (1970) has noted the occurrence McElhinny, 1970; among others) summarizing of hyaloclastitcs, volcanic rocks known to form available paleomagnetic evidence derived from at present only in subglacial or subaqueous continents exclusive of Antarctica, including eruptions of lava. Accumulations of hyaloclas- Australia, present no concensus on East Antarc- titcs up to 400 metres thick with no evidence tica's mid to late Mesozoic or Early Tertiary of subaerial lava flows have been interpreted latitudinal position. Data reported by McEl- to indicate ice at least that thick in some places hinny and Wellman (1969) and Francheteau during the Eocene, providing, of course, that and Sclater (1969) do not exclude the possi- such volcanics belong to a single flow and pro- bility of a high latitude (yet outside the viding that Le Masurier is correct in assuming Antarctic Circle) dispersal route along the that this area in West Antarctica was above Victoria Land and Wilkes Land coast of East sea level when such lavas were extruded. Other Antarctica during the early Tertiary. Almost all hyaloclastite accumulations in the same area reconstructions, however, place West Antarctica indicate giaciation from the Oligocene to the within the Antarctic Circle during late Mesozoic Pleistocene where ice thickness may have and Cenozoic, and yet the evidence of Notho- reached 1200 metres by the Pliocene. Lava /flgw^-dominated forests in this area is incon- flows dated as late Miocene in the Jones Moun- trovertible. Either such forests were able to tains of West Antarctica are known to overlie withstand a yearly cycle with three months of ANTARCTIC DISPERSAL ROUTES 75 perpetual darkness or parts of East and West number of presently tropical taxa, with first Antarctica lay at a lower latitude. This only occurrences of such forms as Sapindaceae, points out the need for additional geophysical Stercuiiaceae, and definite Anacardiaceae data before the latitudinal position of both among others. To the north of Southeast Asia, major segments of Antarctica during the however, cool to warm temperate assemblages Mesozoic and Cenozoic is clear. had differentiated by the early Cenozoic. Floras of northern Thailand (Endo and Fujiyama, Indomalaysian Dispersal Route 1966), Siberia, Manchuria, Korea, and Japan to cool, temperate In marked contrast the (Barbour, 1929; Chaney, et al, 1963; Endo, floras of Paleogene Antarctica, are floras from 1964; Endo and Fujiyama, 1966) during early the late Mesozoic and Cenozoic Indomalaysian to mid-Cenozoic, commonly contained such area. Although no thorough, current summary forms as Sequoia, Taxodium, Pinus, Alnus, of the Southeast Asian and Indonesian fossil Populus, Betulaceae, Ulmaceae, Platanaceae, floras is available, a number of early papers juglandaceae, Ericaceae as well as some (Barbour, 1929; Colani, 1917, 1920; Edwards, families (e.g. Fagaceae, Aceraceae) held in 1923; Goppert, 1854; Holden, 1916; Krausel, common with the more southerly areas. 1922, 1925, 1926; Seward, 1941; Andrews, Undoubtedly, a detailed re-evaluation of 1961 among others) discussing leaf, wood, and much of the early work on Indomalaysian fruit evidence and more recently papers paleofloras must be carried out before the (Chaney, et al., 1963; Endo, 1964; Endo and floral history of this area will be understood. Fujiyama, 1966; Germeraad, et ah, 1968; Jones However, with recent palynological investiga- et a/., 1966; Kon'no, 1966; Manten, 1969) tion confirming many of the earlier identifica- describing the pollen, allow an estimation of tions of Cenozoic tropical forms based on the late Mesozoic and Cenozoic vegetation of leaves, wood, and fruit, apparently several this area. By the Cretaceous, several elements families of plants presently restricted to the now characteristic of the warm, humid tropical tropics occurred in Southeast Asia as early as lowlands of Southeast Asia and Indonesia were the Cretaceous and Paleogene. During the entire present there. Germeraad, et al (1968) re- history of the modern avian families, and cer- ported Nypa palm, now restricted to the man- tainly during the Neogene, then, much of the grove swamps of the Indomalaysian region, Indomalaysian area probably supported a from late Cretaceous sediments of Sarawak. warm, humid tropical flora, not so different Paleogene floras of northern Viet Nam (Colani, (at the family level, at least) from that 1920), Borneo (Krausel, 1925), Sumatra occupying the same area today. (Krausel, 1922), and Java (Goppert, 1854) contain many of the tropical elements found Australia in floras of these areas today: Palmae, Panda- Instrumental in the development of the naceae, Ficus, Quercus (closely related to Australian avifauna were the dramatic changes tropical oaks in this ), Piperaceae, Ana- in the climatic regime affecting Australia dur- cardiaceae or Annonaceae, Moraceae, Musa- ing post-Palaeozoic times. Raven and Axelrod ceae, Dipterocarpaceae, Rhizophora (man- (1972), Gill (1961), Dorman (1966), and grove), among others. Some of these early Burbidge (1960) summarize much of the geo- floras contain only tropical forms, but others chemical, floral, faunal, pedologic, and litho- exhibit a mixture of these and more temperate logic data that are employed to reconstruct elements such as Taxus, Phoebe, Betula, Ben- Australia's past climate. zoin (Colani, 1920), characteristic of the Most important as temperature indicators Himalayas or more northern areas in Asia, are radioisotope, paleobotanical, and faunal and probably represent elements of upland data. Paleotemperature determinations, based 18 16 floras of Indonesia and Southeast Asia. Mio- on oxygen isotope studies (0 /0 ratios) of cene and younger floras from the Indomalay Cretaceous belemnites from Australia and New region are more diverse, containing a large Zealand (Stevens and Clayton, 1971), indicate 76 PAT VICKERS RICH decreasing near shore water temperatures from that contained such additional taxa as Casuar- an early Cretaceous (ca. 110 m.y. B.P.) high ina> Melaleuca, Hakea, and Eremophila. Most to an Albian-Cenomanian (ca. 100 m.y. B.P.) of this mid to late Cenozoic flora appears to low (N.Z. samples: from 42° S. present lati- have been (and still is) adapted to the drier, tude, ca. 16° C, comparable to averages N.Z. more seasonal conditions that became prevalent temperatures at 39-40° S.; Australian samples on a continental scale as Australia moved into from Lake Eyre region at 28° S. present lati- the high pressure "horse latitudes" with their tude, ca. 15° C, comparable to temperatures accompanying xeric climates. Sclerophyll forest, at 41° S. at present). Temperatures rose again savannas, and more arid vegetation covered in the Coniacian-Santonian (ca. 18° C. from much of Australia, while rain forests were samples taken at 42° S. present latitude in present along the eastern, mountainous edge of N.Z., comparable to temperatures at 36-37° S. the continent (Tedford, 1973a). at present) but began dropping again during In New Guinea, with the emergence of the Campanian (ca. 70-75 m.y. B.P.), con- extensive lowlands in the late Oligocene-early tinuing into the Paleocene. Oxygen isotope Miocene, a rich tropical flora with strong studies on shells of several different molluscan Malaysian affinities and quite distinct from genera (collected from localities in southern that of mainland Australia gained a foothold. Victoria between the present 38-39° S. lati- Only much later in the Pliocene and Pleisto- tudes) suggest a continued drop in tempera- cene, accompanying the tectonism that pro- tures into the late Eocene (to ca. 12-16° C.) duced extensive temperate, montane environ- followed by a rise to a Cenozoic maximum in ments as well as xeric conditions in areas of the mid-Miocene (22-25° C), probably accom- rain shadow (Raven and Axelrod, 1972), was panying the onset of Australia's northward drift there much exchange between New Guinea from Antarctica. Since the Miocene, tempera- and Australia. At this time, too, northern tures have continued to drop, with minor temperate forms (e.g. Castanopsis, Rhododen- fluctuations, to the present (12° C. for southern dron) were able to enter New Guinea and Victoria) (Dorman, 1966; Dorman and Gill, Australia from Asia for the first time along 1959a-b). the temperate uplands created by the mid to Cretaceous and early Tertiary floras indicate late Cenozoic tectonism. widespread temperate conditions evidenced by Invertebrate faunas (particularly foramini- mixed forests of austral gymnosperms and ever- fcra, mollusca, and echinoids) indicate, by green angiosperms (Podocarpaceae, Araucaria- faunal composition and diversity, a Tertiary ceae, Araliaceae, Myrsinaceae, Protaceae, temperature maximum in the early Miocene. 3 Winteraceae, Atherospermataceae, Epacrida- Only at this time were any of the larger ceae, Malvaceae, Loranthaceae, Sapindaceae, foraminifera (Trybliolepidina, Cycloclypeus) Casuarinaceae, Myrtaceae, and Nothofagus present in the southern Australian Bass Strait (Fagaceae); Raven and Axelrod, 1972). Dur- Province. Likewise, the first major reef building ing this time Araucaria, now restricted in episode in Australia since the Palaeozoic began Australasia to areas in New Guinea and coastal in the Miocene when Old World reef com- eastern Australia, where average temperature of munities, as well as other tropical invertebrates the coldest month is 11° C, grew in Tasmania and southern Victoria. Nothofagus brassi, pre- 8 The mid-Tertiary temperature rise in Australia, sently restricted to altitudes above 3000 feet in indicated by invertebrate faunas and oxygen isotope ratios, correlates New Guinea, was widespread in the early Ter- well with the drift of this continent northward into lower, more tropical latitudes. tiary of southern and eastern Australia. Mid- Throughout most of the world, temperatures steadily Tertiary floras, on the other hand, indicate re- (with minor fluctuations) declined throughout the striction of such temperate forests (Nothofagus- Cenozoic. Once Australia neared its present position sometime in Araucaria dominated) to the southern and the Miocene and its northward drift slowed, its climate deteriorated in synchrony with northern margins of the continent and the dom- the rest of the world, climaxing in the low temperatures inance of the Eucalyptus-A carta assemblage of the Pleistocene. ANTARCTIC DISPERSAL ROUTES 77

(Palmiere, 1971), colonized the Australian from probable mid-Miocene sediments and continental shelf (Newell, 1971, 1972). In include , several taxa of diprotodontids Victoria, many groups of marine molluscs (extinct family closely related to the (cones, cowries and volutes in particular) ), , phalangers, thylacoleo- exhibited increased diversity in the early to nids, monotremes, and even porpoises among mid-Miocene, as well as a marked increase in others (see Stirton, Tedford, and Woodburne, Indopacific taxa (Darragh, 1973; Darragh, 1968; Tedford, 1973a-b). A number of smaller 1974). By the late Miocene (Cheltenhamian), forms including dasyurids, phalangerids, and however, temperatures began dropping as indi- peramelids were also present. The occurrence cated by the striking reduction both in number of koalas, as well as the diversity of of individuals and in diversity of both molluscs in these mid-Tertiary faunas, suggests a more and echinoderms. By Kalimnan times (about heavily vegetated, more humid environment 6 m.y. B.P.), the marine invertebrate faunas during the mid-Tertiary than presently exists in the continental shelf areas around Australia in these areas today, a suggestion certainly were essentially modern (Brown, Campbell, corroborated by other vertebrate as well as and Crook, 1968), indicating temperatures not floral evidence. so different from those today. Vertebrate faunas, too, indicate changes in Summary humidity and temperature in much of Australia During the Mesozoic and as late as mid- during the Cenozoic. Lungfish, confined to a Eocene (ca. 50 m.y. B.P.), geological and small area of southeastern in the paleobotanical data suggest that a southern late 19th century (Grigg, 1964), are present dispersal route across East Antarctica and in Cretaceous sediments in southern Victoria archipelagic West Antarctica was the most (Schaeffer, 1969) and at a number of localities likely connection between Australia and other of Tertiary age in the Australian interior. continents. Southeast Asia during this period Teleost fish are known from most of the was probably far to the north, perhaps as much localities producing lungfish, yet many of these as 30-50° of latitude. At least many of the areas no longer support permanent streams. islands making up West Antarctica, as well as Crocodilians, now rarely found further south parts of East Antarctica, supported substantial than Rockhampton on the northeast coast and forests dominated by several kinds of gymno- the Ord River (Gill, 1961) in the northwest, sperms, including Araucaria and Podocarpus ranged into northern and Vic- as well as a number of angiosperms including toria in the Tertiary, demonstrating greater Nothojagus. Part of this route, possibly along humidity and yearly minimum temperatures the Victoria Land and Wilkes Land coast of than encountered today in those areas. Diverse East Antarctica and appropriate islands in West fossil avifaunas from the mid to late Tertiary Antarctica, may have lain north of the Antarctic of interior South Australia and the Northern circle, thus providing a route having some Territory (see figures 3-4, table II, and later daylight hours throughout the entire year. By section of paper) include (Phoeni- the end of the Miocene, Australia had severed copteridae) and even anhingas (Miller, 1963b, its connections with East Antactica and drifted 1966a), the latter restricted to warm regions north to within 10° of its present position, with permanent streams and abundant stream- strengthening the dominance of the Indomalay- side vegetation today and probably for some sian dispersal route, which has been most in- time in the past (Rich, 1972, 1974). Fossil fluential during the latter part of the Cenozoic. mammals, similarly, suggest a more humid Simultaneous with northward drift, climatic environment during the Tertiary (see Tedford, changes affecting the Australian continent 1973a for discussion of the fossil marsupial brought about a shift from the early Cenozoic temperate flora faunas of Australia) . In interior South Australia cool dominated by proteads and the a diverse assemblage and Nothojagus to the mid to late Cenozoic of medium to large-sized mammals are known (including the present) zoned vegetation com- 78 PAT VICKERS RICH

posed of arid (in interior Australia), tropical ments in southeastern Victoria have produced (in New Guinea and Northern Australia), and (Talent, et al, 1966; Waldman, 1970) cool temperate (in southeastern Australia and and record an early presence of birds on the Tasmania) elements. Such marked changes in Australian continent. These, together with a

Australia's geographic position, its climate and few Eocene-Oligocene bones from floras, and its links with the rest of the world, southeastern Australia (Simpson, 1946, 1957, undoubtedly have had a tremendous effect on 1959, 1965, 1970; Gill, 1959a-b; Finlayson,

its Cenozoic vertebrate faunas, including its 1938; Anonymous, 1959), a single record birds. of an unidentified bone (now lost) from early Tertiary sediments near Sunnybank, southeastern Queensland (Hill and Denmead, Origin of the Non-passeriform Avifauna of Australia 1960, p. 349; Houston, 1967, p. 85), and a impression as well as a possible fore- Australian Avifaunal Composition limb of a bird from near Wannon, western The living terrestrial and fresh-water non- Victoria (Chapman, 1910; David, 1950), con- passeriform avifauna of Australia and New stitute the early record of birds in Australia. Guinea is composed of some 44 families, of By mid-Miocene, however, a moderately di- which the only endemics are the and verse assemblage of modern avian families (Casuariidae: and Casu- was present, known from several localities (see

arius) and the collared hemipode (Pediono- Table 3 ) in the interior of Australia and midae) (see Table 3). If those birds restricted includes: 4 to the Australia-New Guinea-Southwest Pacific (mihirung birds) (Rich, (excluding New Zealand) area are considered, 1973) then the megapodes (Megapodiidae) and owlet Dromaiinae () (Rich, 1973) to frogmouths (Aegothelidae) can be added Pelecanidae () (Miller, 1966b) the list of endemic families. In addition, a few Phalacrocoracidae () (Rich, distinct subfamilies are endemic to Australia 1973) and/or New Guinea and the Southwest Pacific, (an eagle) (Rich, 1973) including the Cereopsinae, , Cacatu- () (Rich, 1973) inae, and Micropsittinae, all except the first, Phoenicopteridae (flamingos) (Miller, members of the Psittacidae (parrots). At the 1966a) generic level, 34 taxa (exclusive of those in cf. Gruidae (cranes) (Stirton, et al, 1968) the above-mentioned families and subfamilies) cf. Rallidae (rails) (Stirton, et al, 1968) are endemic to Australia and/or New Guinea , undetermined family alone (see Table 2) and an additional 28 are (shorebirds) (Rich, 1973) endemic to Australia and/or New Guinea and Laridae () (Stirton, et al., 1968) the Southwest Pacific (again excluding New Burhinidae (thick knees, stone curlews) Zealand). The fossil record adds two families (Stirton, et al, 1968; Rich, 1973) not present in Australia today: the Dromorni- (nightjars, frogmouths) thidae, composed of large to truly gigantic (Rich, 1973). birds endemic to Australia, and the Phoenicopteridae (flamingos), both of which The avifauna, even at this time, was strikingly survived into the Pleistocene in Australia; modern with only the dromornithids (4-5 flamingos, of course, yet survive in several parts genera in the Miocene) and the flamingos of the world. (2 genera in the Miocene) forming an odd

4 History of the Australian Non-Marine To complete the list of Australian Miocene fossil Avifauna (Figures 7-8) birds, Wilkinson, H. E.. 1969, reported an (Diomedeidae) from Victoria. Since this is a marine Early Cretaceous (perhaps late Jurassic; bird, however, it will not be considered in the above J. Warren, pers. comm., 1974) lacustrine sedi- discussion. ANTARCTIC DISPERSAL ROUTES 79

TABLE 2 Macropygia (, Columbinae) Chcdcophaps (Columbidae, Columbinae) Endemics to Australia-New Guinea-Southwest Phaps (Columbidae, Columbinae) Pacific (after Peters 1931-1945) Ocyphaps (Columbidae, Columbinae) Lophophaps (Columbidae, Columbinae) t Australia and/or New Guinea Geophaps (Columbidae, Columbinae) % Australia and/or New Guinea and Southwest Pacific Histriophaps (Columbidae, Columbinae) * Australia and/or New Guinea, Southwest Pacific, Leucosarcia (Columbidae, Columbinae) Asia Geopelia (Columbidae, Columbinae) Gymnophaps (Columbidae, Columbinae) FAMILY LEVEL Reinwardtoena (Columbidae, Columbinae) f Casuariidae (including iCasuarius and Gatltcolumba (Columbidae, Columbinae) ~\Dromaius) Henicophaps (Columbidae, Columbinae) t Pedionomidae (tPedionomus) Trugon (Columbidae, Columbinae)

if Megapodidae ($Megapodius, iLeipoa, \Alectura, Otidiphaps (Columbidae, Columbinae) §Telegalla, §Aepypodius) Caloenas (Columbidae, Columbinae) * Podargidae (*Podargus) Goiira (Columbidae, Goiirinae) % Aegothelidae ($Aegotheles) Edectus (Psittacidae, Psittacinae) * Hemiprocnidae (*Hemiprocne) Geoffroyus (Psittacidae, Psittacinae) Polytelis (Psittacidae, Psittacinae) SUBFAMILY LEVEL Aprosmictus (Psittacidae, Psittacinae) t Ceropsinae (~\Cereopsis) AUsterus (Psittacidae. Psittacinae) JLoriinae {tTrichoglossus, ^Pseiiteles, Platycercus (Psittacidae, Psittacinae) iGlossopsitta, $Opopsitta, -fLathamus, Purpureicephalus (Psittacidae, Psittacinae)

tChalcopsitta, \Pseudeos, $DomicelIa t Psephotus (Psittacidae, Psittacinae) #, Wreopsittacus, iNeopsittacus, Neophema (Psittacidae, Psittacinae) Melopsittacus (Psittacidae, Psittacinae) %Psittaculirostris ) ftCacatuinae {tProbosciger, Walyptorhynchus, Pezoporus (Psittacidae, Psittacinae) Geopsittacus (Psittacidae, Psittacinae) tCallocephalon, %Cacatuat \Eolophus, jNymphicus) Psittrichas (Psittacidae, Psittacinae) % Micropsittinae ($Micropsitta) Psittacella (Psittacidae. Psittacinae) Loriculus (Psittacidae, Psittacinae) GENERIC LEVEL Scythrops (Cuculidae, Cuculinae) %Zonerodius (Ardeidae) Ninox (Strigidae) * Xenorhynchus (Ciconiidae) Uroglaux (Strigidae) t Anseranas (Anatidae, Plectropterinae) Eurostopodus (Caprimulgidae) t Stictonetta (Anatidae, ) Dacelo (Alcedinidae, Daceloninae) t (Anatidae, Anatinae) Tanysiptera (Alcedinidae, Daceloninae) tSalvadorina (Anatidae, Nyrocinae) Melidora (Alcedinidae, Daceloninae) tChenonetta (Anatidae, Anserinae) Clytoceyx (Alcedinidae, Daceloninae) t (Anatidae, Oxyurinae) % Renicopernis (Accipitridae, Elaninae) f Lophoictinia (Accipitridae, Milvinae) element leaving no living representatives in (Accipitridae, Milvinae) t Haemirostra Australia. Both of the latter groups survived * (Accipitridae, Milvinae) t Erithrotriorchis (Accipitridae, ) into the Pleistocene but not the Recent. t Harpyopsis (Accipitridae, Buteoninae) By the Pliocene (Casuarius) had %Synoicus () differentiated and were present in New Guinea t Anurophasis (Phasianidae) tEulabeorms (Rallidae) (Plane, 1967) while a third genus of , * Rallina (Rallidae) Phoeniconaias, occurred for the first and only $Tribonyx (Rallidae) time in Australia (Phoeniconotius is not known * Amaurornis (Rallidae) * Poliolimnas (Rallidae) from sediments younger than Miocene, while t Rallicula (Rallidae) Phoenicopterus persisted into the Pleistocene). %Gymnocrex (Rallidae) Pleistocene faunas add the first known Aus- t Megacrex (Rallidae) t Peltohyas (Charadriidae) tralian occurrences of the Podicepedidae (Podi- f Cladorhynchus (Recurvirostridae) ceps, ), the (spoon- * Esacus (Burhinidae) atratus (Black Swan), the tStiltia (Glareolidae) bills), cf. Cygnus tProcelsterna (Laridae) Megapodiidae (mound builders, including a (Laridae) %Gygis gigantic form nearly twice the size of any * Ptilinopus (Columbidae, Treroninae) from several localities in t Megaloprepia (Columbidae, Treroninae) living , * Ducula (Columbidae, Treroninae) eastern Australia), the Phasianidae (quail), Lopholaimus (Columbidae, Treroninae) t the Falconidae (falcons), the Turnicidae f Petrophassa (Columbidae, Columbinae) 80 PAT VICKERS RICH

A PLIOCENE MIOCENE O OUGOCENE + EOCENE- PAL EOCENE ©MESOZOIC

Fig. 7—Tertiary and Mesozoic localities producing (Tyto, barn ), and several passeriforms. A fossil birds in Papua-New Guinea and Aus- number of additional groups reported by tralia: Awe (1); Bullock Creek (2); Rivers- DeVis leigh (3); (4); Peak Downs (5); (1888a-b, 1889a-b, 1891a-c, 1905; in Gregory, Chinchilla Lake Kanunka (6); (7); Lake 1906) and Longman (1945) need restudy to Palankarinna (8, 9); Eurinilla Creek and ascertain their taxonomic Lake Pinpa (10); Port Noarlunga (11); positions (Phala- Christie's Beach (12); Pritchard Brothers' crocoracidae, Ciconiidae, Anatidae, Rallidae, Quarry (13); Mt. Gambier (14); Devil's Den 5 Otididae ). (15); Spring Creek (Minhamite) (16); Beaumaris (17); Endurance Pit (18); Lake Thus, at least 23 of the 44 families consti- Ngapakaldi (19); Lake Pitikanta (20); Red- tuting the modern non-passeriform avifauna of bank Plains (21); Well (22); Australia have mid to late Cenozoic histories Bugaldi (near Coonabarabran (23); Redruth, Wannon (24); and Koonwarra (25). there, while two additional families have Mio- cene to Pleistocene records but no modern Turnix, ( quail ) , the Pedionomidae representatives in the Australian avifauna. (plains wanderers), the Scolopacidae (sand- Most certainly the lack of a Tertiary record pipers), the Chionididae (Chionis, sheathbill), for many other families is owing to the small the Columbidae (pigeons, doves), the Psitta- 3 Taxa named in boldfaced type indicate earliest cidae (parrots and allies) , the Tytonidae known occurrence in Australia. ANTARCTIC DISPERSAL ROUTES 81

vertebrate producing Pig. 8—Quaternary localities producing fossil birds number of Cenozoic Australia: Dia- in Papua-New Guinea and localities presently known in Australia. mantina River (no specific locality known) Owing to the limited nature of the fossil (1); Cassidy Locality (2); Warburton River Localities (3); Cooper's Creek Localities (4); record in Australia, first occurrences of avian Thorbindah Cud- Lake Callabonna (5); (6); families and genera have no special significance die Springs (7); Ashford Caves (Bone Cave) regarding which dispersal route was being used, (8); Bingara (9); Canadian Lead (Gulgong) (10); Walli and Wellington Caves (11); or in what direction movement occurred. By Caves (Guineacor) (12); Lake Wombeyan the Miocene, when the first extensive avifaunas Menindee (13); Lake Tandou (14); Baldina in Australia, Southeast Asia and Creek (15); Normanville (Salt Creek) (16); are known Kangaroo Island (17); Henschke's Cave and Australia were close enough to one another for Penola (19); Mt. Gam- Victoria Cave (18); avifaunal interchange to have been underway. bier (20); King Island (21); Madura Cave nothing of the (22); Scott River (23); Mammoth Cave Unfortunately we know almost (24); Darling Downs (Kings Creek, War- earlier Australian terrestrial avifaunas, which wick) Drover's Cave (26); Gore (27); (25); could have marked significance regarding South Buchan Caves (28); Weeke's Cave (29); and affinities, the existence Brother's Island, Pt. Lincoln (30) ; American enhanced by Lancefield (31). of an Antarctic dispersal in the early Tertiary. PAT VICKERS RICH

TABLE 3

Summary of the Avifaunas from Australian Cenozoic Localities 1

(if reference not cited, from Rich, 1973) TERTIARY EOCENE Queensland Interbedded sediments in unnamed basalt overlying Darra Fm. and underlying Sunnybank Fm. at Sunnybank, Aves, undetermined (now lost). Cribb, et al., 1960.

South Australia Christie's Beach, 'Transitional Marl' Member, base of Blanche Point Marls, Palaeeudyptes cf. antarcticus (Spheniscidae, penguin). Simp- son, 1946, 1957.

South Australia Port Noarlunga Jetty. Banded Marl Member of Blanche Point Marls, Palaeeudyptes cf. antarcticus. Simpson, 1946, 1957. OLIGOCENE South Australia Mount Gambler. Gambier Limestone. Spheniscidae (penguin). Simp- son, 1946, 1957.

South Australia Pritchard Brothers' Quarry. Gambier Limestone, 1\ miles WNW of Mt. Gambier, Palaeeudyptinae (penguin). Simpson, 1946, 1957. MIOCENE

South Australia Dam Locality. Moloorina Station, cf. Dromornithidae (mihi- rung bird, shell). M. Woodburne, M. Archer, pers comm., 1973.

South Australia Lake Pinpa. Unnamed unit; Phalacrocorax sp. (Phalacrocoracidae, ), Anatidae (), Burhinidae (thick-knee, stone curlew).

South Australia Eurinilla Creek. Unnamed unit; Anatidae (duck), cf. Phoenicopteridae (flamingo).

South Australia Lake Palankar'mna. Etadunna Formation, Ngapakaldi fauna (includ- ing localities LFM Loc. 9, U.C.M.P. Locs. V-5762, V-5763, V-5764, V-5765, V-5770, V-5771, Mammalon Hill); Pelecanus tirarensis (Pelecanidae, ), Phalacrocoracidae or Anhingidae (cormorant or darter), Phoenicopteridae, gen. et sp. undet. (flamingo), Phoeni-

conotius eyrensis ( Phoenicopteridae), (undet.), Oxyura (Anatidae, duck), cf. Gruidae (), Rallidae (), Burhinidae (thick-knee), Charadriiformes (shorebirds), Laridae (), Stirton, Tedford, and Woodburne, 1967.

South Australia Lake Pitikanta. Etadunna Formation, Ngapakaldi fauna (including Discovery Basin U.C.M.P. V-5774 and U.C.M.P. V-6150); Phoeni- copterus novaehollandiae (Phoenicopteridae, flamingo). Miller, 1963b.

Queensland Riversleigh. Carl Creek Limestone, Riversleigh fauna; Dromornithidae (new gen., new sp.) (mihirung bird).

Victoria Devil's Den (Glenelg River). Glenelg Group, Balcomian Stage, north of Dartmoor; Anthropodxptes gilli (Spheniscidae, penguin). Simpson, 1959.

South Australia Kangaroo Well. Unnamed formation. Kangaroo Well fauna, Aves, undetermined. Stirton, Tedford, and Woodburne, 1967.

South Australia Lake Ngapakaldi, Leaf Locality. Wipijiri Formation, Kutjamarpu fauna (U.C.M.P. V-6213); Dromaius (Casuariidae, emus), Drom- ornithidae (mihirung bird); Ngapakaldi 2 (U.C.M.P. V-6213), Dromornithidae. Stirton, Tedford, and Woodburne, 1967.

Northern Territory Bullock Creek. Camfield Beds, Bullock Creek fauna; Dromornithidae (1 new gen., 2 new sp.) (mihirung birds).

1 Those taxa prefaced by (x) were last studied by DeVis, and further study is needed to ascertain their correct taxonomic position. ANTARCTIC DISPERSAL ROUTES 83

Northern Territory Alcoota. Waite Formation, Alcoota fauna (including Paine Quarry U.C.M.P. V-6345, Rochow Locality U.C.M.P. V-6349); cf. Dromaius (Casuariidae, emu), Dromornithidae (3 new gen., 3 new sp.) (mihirung bird), Accipitridae (eagle).

New South Wales Bugaldi, diatomaceous earth deposit, cf. Aegothelidae (owlet-- mouth).

Victoria Beaumaris. Sandringham Sands, Cheltenhamian Stage, Beaumaris fauna; Spheniscidae (penguin), IPseudaptenodytes macraei (Sphenis- cidae, penguin ) , IP. minor, Diomedea thyridata (Diomedeidae, albatross). Simpson, 1965, 1970; Wilkinson, 1969.

Victoria Spring Creek, Minhamite. ?Cheltenhamian Stage; Pseudaptenodytes macraei (Spheniscidae, penguidae). PLIOCENE South Australia Lake Palankarinna, Lawson-Daily Quarry (Lawson Quarrv), Mampu- wordu Sands, Palankarinna fauna (U.C.M.P. V-5769); dromaius ocypus (Casuariidae, emu), Dromornithidae (mihirung bird). Miller, 1963a.

New Guinea Awe. Otibanda Formation, Awe fauna; Casuarius sp. (Casuariidae, cassowar). Plane, 1967.

South Australia Lake Kanunka. Katipiri Sands, Kanunka fauna (U.C.M.P. V-5772, V-5773, V-5855); Dromaius (Casuariidae, emu), Pelecanus (Pele- canidae, pelican), Phafacrocorax (Phalacrocorax, cormorant), (2 sp.), Anhinga novaehollandiae (Anhingidae, anhinga, snake bird), Phoeni- conaias gracilis (Phoenicopteridae, flamingo), Phoenicopterus ruber (Phoenicopteridae) Anatidae (duck), cf. Aythya (Anatidae, duck). Stirton, Tedford, and Woodburne, 1967.

Queensland Peak Downs. australis (Dromornithidae, mihirung birds). Owen, 1874, 1879b.

Queensland Chinchilla. Chinchilla Sands, Chinchilla fauna; Dromaius (Casuarii- dae, emu), Pelecanus conspicillatus (=r P. proavus) (Pelecanidae, pelican), Haliaetor melanoleucos (Phalacrocoracidae, cormorant),

(x) Xenorhynchus nanus (Ciconiidae, ), (x) Anas elapsa ( = Nettion elapsum, Brodkorb) 1 (Anatidae, duck), (x) Biziura exhumata (Anatidae, duck), (x) Dendrocygna (Anatidae, duck), (x) Nyroca reclusa (= reperta, = Aythya reclusa, Brodkorb) (Anatidae, duck),

(x) Nyroca robusta ( — Aythya robusta, Brodkorb) (Anatidae, duck), Gallinula (= Tribonyx) mortierii reperta (Rallidae, rail) (Olson, 1974), Fulica atra (Rallidae, rail) (Olson, 1974), DeVis, 1888-1905. PALEOGENE(?) Victoria Redruth (Wannon River). Ironstone, 'wing of a struthious bird', Aves, undetermined (David, 1950), feather impression (Chapman, 1910). MIDDLE TERTIARY

Tasmania Endurance Pit. cf. Dromornithidae (mihirung birds), footprints. (Rich and Green, 1974). PLEISTOCENE QUEENSLAND Darling Downs Dromaius cf. novaehollandiae (Casuarridae, emu) Anhingidae (darter, J. van Tets, pers. comm., 1973) (x) Palaeopelargus nohilis (Ciconiidae, stork) (x) Platalea Isubtenuis (Threskiornithidae, spoonbill) (x) Dendrocygna validipennis (Anatidae, duck) (x) Nyroca sp. (Anatidae, duck) (x) Necrastur alacer (Accipitridae, , eagle) (x) Taphaetus brachialis ( = Uroaetus = Aquila) (Accipitridae, eagle) Progura gallinacea (Megapodiidae, mound builder) (van Tets, 1974a) Gallinula mortierii reperta (Rallidae, rail) (Olson, 1974) — ? Warwick (x) Lithophaps ulnaris (Columbidae, pigeon) DeVis, 1888-1905.

provided by Brodkorb, 1963, 1964, 1967, 1971a. 1. Generic reallocation most recently 84 PAT VICKERS RICH

Gore Progura naracoortensis (Megapodiidae, mound builder). Longman, 1945; van Tets, 1974a.

King's Creek, Darling Downs (x) Dromaius patriots (? — D. novaehollandiae) (Casuariidae, emu) (x) Taphaetus brachialis { — Uroaetus — Aquila) (Accipitridae, hawk, eagle). DeVis, 1888-1905.

Diamantina (general locality; could be any of several places along the Diamantina River) (Dromornithidae, mihirung bird).

Thorbindah, Paroo River Dromaius (Casuariidae, emu) Genyornis, cf. newtoni (Dromornithidae, mihirung bird). NEW SOUTH WALES Ashford Caves (Bone Cave) (U.C.M.P. V-67237, U.C.M.P. V-5545) Aves, undetermined.

Bingara Dromaius (Casuariidae, emu). L. Marcus, pers. comm., 1973. Canadian Lead Dromornithidae (mihirung bird). Etheridge, 1889; Owen, 1879b.

Cuddie Springs Genyornis cf. Newtoni (Dromornithidae, mihirung bird). Anderson and Fletcher, 1934.

Lake Menindee (including Rifle Range Loc. U.C.M.P. V-67233, U.C.M.P. V-5371, V-67185, V-67186, V-67187) Dromaius (Casuariidae, emu). Lake Tandou Dromaius (Casuariidae, emu) (Podicepedidae, ) cf. Ciconiiformes (, , ibises) cf. Cygnus atratus (Anatidae, swan) cf. Columbidae (pigeon).

Wellington Caves (including Bone Cave, Mitchell's Cave, Phosphate Mine (in part U.C.M.P. V-5538)) (x) Dromaius patricus {ID. novaehollandiae) (Casuariidae, emu) Casuarius lydekkeri (Casuariidae, cassowary) Dromornithidae (mihirung bird) Progura gallinacea (Megapodiidae, mound builder). Mitchell, 1839; Miller, 1962; Rich, 1973; van Tets, 1974a.

Walli Caves Progura gallinacea (Megapodiidae, mound builder), van Tets, 1974a.

Wombeyan Cave (Guineacor) (in part U.C.M.P. V-5537) Aves, undetermined Dromaius (Casuariidae, emu) Progura gallinacea (Megapodiidae, mound builder), van Tets, 197 4a. VICTORIA

Buchan Cave (J. van Tets, pers. comm., 1973) Rallidae (rails) Coturnix sp. (Phasianidae, quail) Alcedinidae (kingfishers) Passeriformes (song birds, perching birds).

Lancefield Dromornithidae (mihirung bird). TASMANIA King Island Dromaius minor (Casuariidae, emu). Spencer and Kershaw, 1910. ANTARCTIC DISPERSAL ROUTES 85

SOUTH AUSTRALIA

Baldina Creek (near Burra) Dromornithidae (mihirung birds).

Brother's Island, Pt. Lincoln Dromornithidae (mihirung bird).

Cooper's Creek (includes several localities collected primarily by J. W. Gregory and later parties from the University of California, Berkeley) (Katipiri Sands, Malkuni Fauna).

University of California Localities (Miller, 1963-1966; Rich, 1973)

U.C.M.P. Site 2 (U.C.M.P. V-5379) Unkumilka Waterhole Dromaius sp. (Casuariidae, emu) Phalacrocorax sp. (Phalacrocoracidae, cormorant).

U.C.M.P. Site 3 (U.C.M.P. V-5378) Between White Crossing and Site 2 Dromaius sp, (Casuariidae, emu) Phalacrocorax sp. (Phalacrocoracidae, cormorant; middle-sized sp., lg. sp.).

U.C.M.P. Site 4 (U.C.M.P. V-5380) Anhinga novaehollandiae (Anhingidae, darter/snake bird) Phalacrocorax sp. (Phalacrocoracidae, cormorant; middle-sized sp., lg. sp.) Anatidae (duck).

U.C.M.P. Site 5 (U.C.M.P. V-5381) Pirranna Soakage Dromornithiade (mihirung birds) Phalacrocorax (Phalacrocoracidae, cormorant; middle-sized sp.) Anatidae (small sp.) (duck).

U.C.M.P. Site 7 (U.C.M.P. V-5859) Phalacrocorax sp. (Phalacrocoracidae, cormorant; middle-sized sp.) Pelecanus conspicillatus (Pelecanidae, pelican) Anatidae (duck) sp. (crane).

U.C.M.P. Site 8 (U.C.M.P. V-5860) Dromaius sp. (Casuariidae, emu) Dromornithidae (mihirung birds) Phalacrocorax sp. (lg. sp.) Pelecanus conspicillatus (Pelecanidae, pelican) Anatidae (swan)

U.C.M.P. Site 9 (U.C.M.P. V-5861) Katipiri Waterhole (= Kuttipirra) Dromaius sp. (Casuariidae, emu) Dromornithidae (mihirung birds) Phalacrocorax sp. (Phalacrocoracidae, cormorant, middle-sized sp.) Anatidae (swan).

U.C.M.P. Site 10 (U.C.M.P. V-5869) Anhinga novaehollandiae (Anhingidae, darter/snake bird) Phalacrocorax sp. (Phalacrocoracidae, cormorant) Platalea sp. (Threskiornithidae, spoonbill).

U.C.M.P. Site 14 (U.C.M.P. V-5866) Anhinga novaehollandiae (Anhingidae, darter) Phalacrocorax sp. (Phalacrocoracidae, cormorant) Phoenicopterus ruber (Phoenicopteridae, flamingo).

U.C.M.P. Site 16 (U.C.M.P. V-5868) Phalacrocorax sp. (Phalacrocoracidae, cormorant) Pelecanus conspicillatus (Pelecanidae, pelican) Anatidae (swan). 86 PAT VICKERS RICH

U.C.M.P. Site 18 (U.C.M.P. V-6147) Dromaius sp. (Casuariidae, emu) Phalacrocorax sp. (Phalacrocoracidae, cormorant) Pelecanus conspicillatus (Pelecanidae, pelican) Anatidae (duck).

Malkuni Waterhole (U.C.M.P. V-5382) Pelecanus conspicillatus (Pelecanidae, pelican).

Markoni Locality (U.C.M.P. V-5382) Dromornithidae (mihirung birds) Podiceps sp. (Podicepedidae, grebes) Phalacrocorax sp. (Phalacrocoracidae lg. sp.) Tyto (Tytonidae, barn owl).

J. W. Gregory Localities (DeVis 1888-1905; in Gregory, 1906)

East of Pirani (x) Biziura exhumata (Anatidae, duck).

Ernu Camp (= Malkuni waterhole)

cf. Genyomis newtoni (Dromornithidae, mihirung bird). (x) Phalacrocorax gregorii (Phalacrocoracidae, cormorant) (x) P. vetusius (x) Archaeocygnus lacustris (= Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Biziura exhumata (Anatidae, duck)

(x) Chenopsis nanus ( = Cygnus lacustris, Brodkorb) (Anatidae, swan).

Lower Cooper Aniiinga novaehollandiae (Anhingidae, darter) (x) Phalacrocorax gregorii (Phalacrocoracidae, cormorant) (x) P. vetustus cf. Dromaius novaehollandiae (Casuariidae, emu) (x) Xenorhynchopsis tibialis (Ciconiidae, stork)

(x) Anas gracilipes ( = Nettion gracilipes, Brodkorb) (Anatidae, duck) (x) Archaeocygnus lacustris (= Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Chenopsis nanus (= Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Nettapus eyrensis (Anatidae, duck).

Lower Cooper Locality 2 Dromaius sp. (Casuariidae, emu) Dromornithidae (mihirung birds).

Lower Cooper Locality 3

Dromaius sp. (Casuariidae, emu).

Lower Cooper Locality 4

(x) Dromaius patricus (cf. D. novaehollandiae) (Casuariidae, emu).

Lower Cooper Locality 5 (x) Asturaetus furcillatus (= Plioaetus jurcillatus Richmond) (Falconi- dae, falcon).

Lower Cooper Locality 6 (x) Baza gracilis (= Aviceda gracilis, Brodkorb) (Falconidae, falcon).

Cooper Creek Anhinga laticeps (Anhingidae, darter) Pelecanus grandiceps (Pelecanidae, pelican).

Patteramordu (= Red Bluff Locality of U.C.M.P.)

(x) Anas strenua ( — Nettion strenum, Brodkorb) (Anatidae, duck). ANTARCTIC DISPERSAL ROUTES 87

Unduwampa

(x) Xenorhynchopsis minor (Ciconiidae, stork) (x) Archaeocygnus lacustris {— Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Chenopsis nanus ( — Cygnus lacustris, Brodkorb) (Anatidae, swan). Wankamaminna (x) Phalacrocorax gregorii (Phalacrocoracidae, cormorant) (x) Archaeocygnus lacustris (— Cygnus lacustris, Brodkorb) (Anatidae, swan). Wurdulumankula (Wurdulmankula)

(x) Dromaius patricus (? cf. D. novaehollandiae) (Casuariidae, emu) (x) Phalacrocorax gregorii (Phalacrocoracidae, cormorant) P. carbo (J. van Tets, pers. comm., 1973) (x) P. vetustus (x) Xenorhynchopsis minor (Ciconiidae, stock) (x) Xenorhynchus nanus (Ciconiidae) (x) Ibis conditus (~ Carphibis condita, Brodkorb) (Threskiornithidae, ibis) (x) Archaeocygnus lacustris (= Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Chenopsis nanus (— Cygnus lacustris, Brodkorb) (Anatidae, swan) (x) Nyroca effodiata (= Aythya effodiata, Brodkorb) (Anatidae, duck) (x) Leucosarcia proevisa (Columbidae, pigeon).

Henschke's Quarry Cave, Naracoorte

Progura naracoortensis (Megapodiidae, mound builder), van Tets, 1974a.

Kangaroo Island Dromaius novaehollandiae diemenianus (Casuariidae, emu). (Condon, 1973).

Lake Callabonna (Lower level)

Dromaius sp. (Casuariidae, emu) Genyornis newtoni (Dromornithidae, mihirung bird).

Lake Callabonna (Upper level)

Phalacrocorax sp. (Phalacrocoracidae, cormorants).

Lake Eyre (exact locality unknown) (x) Ocyplanus proeses (Laridae, gull). DeVis in Gregory, 1906. Mount Gambier Range Dromornithidae (mihirung birds).

Normanville (= Salt Creek)

Dromornithidae, cf. Genyornis newtoni (mihirung bird).

Penola

Dromornithidae, cf. Genyornis newtoni (mihirung bird).

Victoria Cave, Naracoorte (van Tets and Smith, 1974) Dromaius sp. (Casuariidae, emu) Progura naracoortensis (Megapodiidae, mound builder) Leipoa ocellata (Megapodiidae, mound builder) Coturnix pectoralis (Phasianidae, quail) C. australis Coturnix sp. (Phasianidae, quail) Turnix varia (Turnicidae, button quail) Pedionomus torquatus (Pedionamidae, plains wander) philippensis (Rallidae, rail) Peltohyas australis (Charadriidae, ) Tringa glareola (Scolopacidae, sandpiper, stilt, curlew) Gallinago hardwickii (Scolopacidae, sandpiper, stilt, curlew) )

KK I'M VK KIRS RICH

Calidris ruficollis (Scolopacidae, sandpiper, stilt, curlew) Chionis minor (Chioniidac, sheath-bill) Pezoporus wallicus (Psittacidae, ) Tyto novaehollandiae (Tytonidae, barn owl) Gymnorhina tibicen (Passeriformes, songbirds). Gymnorhina tibicen (Passeriforms, song birds)

Warburton River Localities (Katipiri Sands, Malkuni Fauna)

( assidy Locality (U.C.M.P. V 5539) cf. Genyornis (Dromornithidae, mihirung bird).

Green Bluff Locality (U.C.M.P. V 5775) Dromaius sp. (Casuariidae, emu) Phalacrocorax (Phalacrocoracidae, cormorant; lg. sp.) Anserinae (goose Accipitridae (hawk, eagle).

Kalamurina Dromaius sp. (Casuariidae, emu) Dromornithidae, cf. Genyornis (mihirung bird) (x) Phalacrocorax gregorii (Phalacrocoracidae, cormorant) /'. ( x ) veiustus Brodkorb) (Anatidae, duck) (x) Anas gracilipes ( Nation gracilipes, lacertosus (van lets, 1974c) (x) Tapltaetus lacertosus ( Icthyophaga (Accipitridae, eagle)). DeVis, 1888-1905; Rich, 1973,

I ookout Locality (U.C.M.P. V-5776) Phalacrocorax sp. (middle-sized sp., lg. sp.) (Phalacrocoracidae, cormoranl ).

Marcus Locality Phalacrocorax sp. (Phalacrocoracidae, cormorant; middle-sized sp., lg. sp.) cf. Pelecanus sp. (Pelecanidae, pelican).

Pundrakadarintva Soakage (U.C.M.P. V-5777) Phalacrocorax sp. (Phalacrocoracidae, cormorant).

Wai bin ton River (x) Pelecanus vatidipes (Pelecanidae, pelican). Etheridge, 1894.

Weeke's Cave (Nullarbor Plain) Platibls flavipes (Threskiornithidae, spoonbill) fasciatus (Accipitridae, ) Falco cenchroides (Falconidae, falcon) Turnix sp. (Turnicidae, bustard quail) Cinclorhamphus cruralis, C. mathewsi, Poephila guttata> ArtamwV leucorhynchus (Passeriformes, song birds), van lets, 1974b.

Wis 1 1 KN AUSTRALIA

I abrinth Cave Aves, undetermined.

Madura ( 'ave Aves, undetermined.

Mammoth ( lave Dromornithidae (mihirung bird).

Scott River ?Dromornithidae (mihirung bird), egg. Butler, 1969; Rich, 1973. ,

ANTARCTIC DISPERSAL ROUTES 89

Previous Opinions Regarding Origin of the major routes may have existed that non-marine Australian Avifauna vertebrates could have employed in reaching or leaving Australia during the late Mesozoic In studying the Australo-Papuan avifauna and Ccnozoic: the Antarctic as well as the three decades ago, Mayr (1941, 1944a-b, Indomalaysian. With this additional informa- 1945b) concluded that it had been derived tion provided by geology, one of Mayr's basic from the north, primarily southeastern Asia, assumptions, that of continental stability, is with colonization occurring throughout the undermined, and his ideas regarding the origin

Cenozoic ( see figure 9a ) . He recognized of the Australian avifauna require re-examina- several groups of invaders, which he believed tion. to represent different colonizations, oldest to Such re-evaluation has been initiated in most recent. The oldest invasion (early to mid- papers by Serventy (1972, 1973), Cracraft Tertiary at the latest, Mayr, 1944a) was re- (1972, 1973), Schodde and Calaby (1972), flected by the most unique groups in the living and Mayr (1972) himself. Serventy suggested non-passeriform avifauna of Australia, includ- such forms as Dromaius (emus), possibly ing the Casuariidae (containing Casuarius Anseranas (Magpie Goose), the Megapodiidae and Dromaius) , Megapodiidae, Loriinae, Caca- (mound builders), the Podargidae (frog- tuinae Kakatoeinae), Platycercinae, (= and mouths), as well as several parrot subfamilies Podargidae. Mayr noted that the nearest rela- (Loriinae, Cacatuinac, Platycercinae (in- tives of these six groups were uncertain, but cluded in the Psittacinae) ) among the non- argued that none were any closely related more passeriforms might well have utilized a to South American families than to Old World southern, rather than a northern route betweea families. His second and more recent wave of Australia and the remaining world. Mayr, in colonists (collared included the Pedionomidae commenting on Serventy's paper, tentatively hemipodes) along with several passeriform considered the ratites, the platyccrcine parrots, families. third The wave consisted of a number and possibly the flamingos as Antarctic dis- genera of endemic to Australo-Papua, which perses but left the question open. Cracraft, he believed were Miocene or Pliocene arrivals, on the other hand, strongly supported a included: Synoicus and (Phasianidae), Geo- southern dispersal route for the ratites, mega- pelia (Columbidae), Irediparra (included in podes, and parrots, but agreed with both Ser- Jacana, Jacanidae), Notophoyx (included in venty and Mayr that the majority of Australian A rdea, Ardeidae ) , Dupetor ( Ardeidae ) avian genera (including most of the non-pas- =* Erythrotriorchis ( Accipitridae ) , Uroaetus ( serifonus) were spawned by Southeast Asian Aquila, Accipitridae), Lophoictinia (Milvinac), faunas. Is this really the case, however, and Syma (included in Halcyon, Alccdinidae), what sort of reasoning and assumptions are Dacelo (Alcedinidae), Tanysiptera (Alccdini- employed to determine each family's route of dae), Cacomantis (Cuculidae), Misocalius (in- dispersal between Australia and the rest of the cluded in Chrysococcyx, Cuculidae), Chaleites world? The following section deals in detail (included in Chrysococcyx, Cuculidae), and with these questions. Eudynamis (Cuculidae). Fourth and fifth Re-evaluation of Probable Immigration Routes waves included taxa that are presently only Into Australia: Assumptions, Reasoning, specifically, or subspecifically distinct, or indis- and Data tinguishable from southeast Asian forms (Plio- cene to Recent arrivals, according to Mayr). Zoogeography is often not a very satisfying Such ideas viewed against a backdrop of stable endeavour owing to the frequent uncertainty continents were reasonable and remained so of conclusions one is able to reach concerning into the late 1950's. With the development areas of origin and early radiation of certain during the last fifteen years of the concepts of organisms. Despite this, however, many workers continental drift, sea-floor spreading, and plate have attempted to understand the distributional tectonics, it has become apparent that two histories of organisms including birds, although 90 PAT VICKERS RICH

j^&Er ^ %Ss

Fig. 9 (A) Cartoon expressing — the classic ideas that together with Mayr's ideas may have concerning origin of the Australian avifauna, been important in shaping the composition of eloquently detailed by Mayr (1944a-b); (B) Australia's bird fauna. cartoon depicting an additional component ANTARCTIC DISPERSAL ROUTES 91 others seriously question many of the criteria composition of continental flora, and history of utilized in such attempts (Rotramel, 1973, connection to other land masses. Darlington among others). The following section sum- further stressed other problems peculiar to marizes briefly assumptions and reasoning deciphering avian zoogeography including the utilized by ornithologists and zoogeographers (1) rapidity with which an avian taxon can who have tried to determine origins of elements disperse, thus obscuring the direction from comprising the Australian avifauna. In addition, which it came; (2) possibility of contempor- it outlines assumptions employed in this paper aneous multiple dispersals; (3) rarity of avian to re-evaluate the immigration and emigration ; and (4) lack of understanding of the pathways of all the non-passeriform families, significance of non-breeding ranges of migratory presents pertinent historical, distributional, and birds. However, after so carefully and precisely diversity data for each family considered, and outlining these assumptions and clues, most briefly outlines the reasoning employed in the were not evaluated when Darlington individu- determination of the dispersal route(s) utilized ally considered a number of "dominant" avian by each Australian group between Australia families; instead diversity and endemism were and the remaining world. usually the only clues considered. Darlington's In his treatise on zoogeography, Darlington judgement on the relative significance of these (1957) outlined several working principles and few clues was not consistent; on the one hand a number of practical clues used in determining he believed that the Phasianidae originated, or the area of origin of different organisms includ- at least underwent a major radiation, in the ing (1) diversity, (2) degree of differentiation Oriental region based on their great diversity 6 (including degree of endemism), (3) extent of there (in comparison to its low diversity area inhabited (whether restricted or wide- elsewhere), but was unwilling to readily accept spread), (4) continuity of area inhabited, the equal probability that the parrots originated (5) distribution of related, competing, and or underwent an early radiation in Australia associated families, and (6) fossil record. based on the same type of distributional and 7 How much weight to give each of these clues, diversity information. Besides and particularly in cases where information is fi "It [the Phasianidae] is the first large family of this limited, was not made clear. Similarly, if some survey which shows an apparent pattern of radiation are available for each clue category, it was data from one main center, the Oriental Region. The true not made clear which clue or clues had priority pheasants, jungle , and peacocks are almost all in over others if multiple interpretations were the Orient, and from there the ancestors of Afropavo Peacock), and of the guinea fowls may possible. Darlington suggested that degree of (the Congo have reached Africa; the ancestors of the differentation (clue 2) should have priority (Tetraoninae) may have gone northward and around fact that information over diversity (clue 1 ), in the northern part of the world, eventually differen- pertinent to the remaining clues (clues 3, 4, tiating most in North America; and the quails may have radiated over the whole world, with a secondary 5, 6) could be used to question the significance evolution center in southern North America. The of diversity. He quite appropriately pointed out -like birds probably [italics mine] have relatively might not only that high diversity in an area low powers of dispersal, so their present distribution there, but ." signal the long history of a group may reflect something of their early history . . might also be due to a number of other factors Darlington (1957, p. 270).

7 "Parrots . . . are almost equally numerous in such as the size and diversity of the environ- species in Old and New Worlds but are much more inhabited (i.e. number of different habitats ment diverse in the former, all the New World forms available), and the amount of competition a belonging to one of the several Old World sub- group encountered from other organisms during families; this suggests an Old World origin of parrots. But this does not carry the analysis far enough. It is its history in an area. To this list Keast (1972) only in the Australian Region and adjacent island factors affecting group diversity added such areas that parrots occur in great diversity. Elsewhere topographic as size of total area inhabited, in the Old World parrots are relatively few and all diversity, percentage of area within the tropics, belone to one subfamily, the same one that is in America. Does the unique diversity of parrots in the total latitudinal range on continent, vegetational Gl ,

92 PAT VICKERS RICH

with in its present geographic position, parrots, Darlington examined several other mained slowly accelerating as more of the avian groups that occur in Australia (Anatidae, colonization emerged during the Ceno- , Strigiformes, Ardeidae, Chara- Malay Archipelago With such a foundation, Mayr's conclusion driidae, Scolopacidae, Rallidae, Cuculidae, zoic. avifauna consisted of several Caprimulgiformes, Apodidac, Alcedinidae) (that Australia's "waves" of northern colonists) was attempting to determine their centres of origin different reasonable. However, as a previous sec- and distributional histories, but could reach no quite this paper has established, present further firm conclusions for any other non- tion of strongly suggests that Aus- passeriform group. He believed that most of geological evidence been anything but stable during the the groups which he considered were old, had tralia has having drifted as much as complex distributional histories, and that areas history of birds, 30-50° of latitude from an Eocene connection of origin as well as early patterns of radiation nearly to its present position could not be determined. He noted that the with Antarctica Thus Mayr's second Caprimulgiformes (nightjars, frogmouths, and by the late Miocene, assumption is seriously challenged. If his first allies) probably had experienced three major re-examined, although most cer- radiations, one in the Australian-Oriental area assumption is have resulted from at least (Podargidae, Aegothelidae), another in tropical tainly true, it could interchange, i.e. (1) Aus- America (Nyctibiidae, Steatornithidae), and a two types of faunal receiving most of its avifauna from Asia third in an undesignated area that produced tralia recent biotic interchange be- the Caprimulgidae, but he was non-commital or (2) the most of the world being about what dispersal paths had been employed tween Australia and the rest Asia. If the latter is the case, similarities during this tripartate diversification. It is with with and Australia would have resulted this background of uncertainty about the dis- between Asia the two faunas, not a one way persal histories of birds that Darlington made from a mixing of Asia. Because of the present the following statement: "Almost all the ascer- immigration from of avian distributional tainable relationships of Australian land birds lack of understanding the conclusion that Asia are toward Asia. The few supposed African histories, however, Australia's avifauna seems relationships, for example that of the honey spawned most of conclusion far too specific for eaters with Promerops of South Africa, are premature, a limited data available. doubtful, and there is no sure, direct relation- the accepted the recent ship between any Australian and South Ameri- Serventy (1972, 1973), concensus of geologists supporting continental can land birds" (1957, p. 262). Thus with this old, endemic elements statement and others regarding the origins of drift and re-examined the avifauna, attempting to point out the parrots and pigeons, Darlington emphasized in Australia's relationships of some Australian birds the "Asiatic" character of the Australian avi- the closer rather than Asian forms. fauna, essentially agreeing with Mayr's ideas. to South American His acceptance of ratite monophyly (as most Mayr's ( 1 944a-b) reasoning, that most 1963;Meise, 1963; Australian birds were derived from Oriental recently supported by Bock, Sibley, 1960) allowed forms, relied heavily on the two observational Parks and Clark, 1966; strongly a southern dispersal assumptions that (1) the Australian avifauna him to support group, and along with additional more closely resembled that of the Oriental route for this not some of the region than that of any other area and (2) data, led him to state: "May present Australasian during the history of birds, Australia had re- older elements of the fauna have reached Australia via Africa and Antarctica during the earlier stages of Gond-

Australian Region reflect place of origin, or with- wanaland?" Serventy did not extend his con- drawal of some groups of parrots from other parts of siderations to other, less unique Australian bird answer this the world, or local radiation? 1 cannot groups, because he believed they had immi- question, but it seems to me that parrots need not have to their lack of close ." grated from Asia due originated in the Australian Region . . Darlington similarity to South forms. (1957. p. 271). American ANTARCTIC DISPERSAL ROUTES 93

Cracraft (1972, 1973) likewise assumed time to evolve quite separately and diverge that continental drift had occurred during the from one another. Relationships between living Mesozoic and Cenozoic (in fact he presented forms that might be used in support of such a review of continental drift literature) and an early Tertiary dispersal route should thus attempted to elucidate the dispersal histories of be sought at higher taxonomic levels, not avian taxa based primarily on phyletic relation- specific and generic. ships of those taxa (using a method discussed In this paper a number of assumptions will by Hennig, 1966; Nelson, 1969; and others). be made and clues used in an attempt to He emphasized that all other clues regarding decipher the dispersal history of Australian origin of avian groups should be subordinate avian families. Assumptions include: to phyletic clues, but then noted how pitifully (1) the geographic arrangement of con- poor present understanding of phyletic relation- tinents has been continually altered ships was for most avian families and thus, throughout the history of birds and I assume, how little is understood of original quite profoundly in the positioning of dispersal routes if that information is derived Australia during the Cenozoic (i.e. from phyletic studies. He discussed those few Australia has drifted from an early groups whose relationships he believed were Tertiary position adjacent to Antarctica, best understood, including taxa with Australian perhaps as much as 30-50° of latitude

histories : (Spheniscidae) , ratites, northwards), nearing its present position galliforms (including Megapodiidae), parrots by the late Miocene;

( Psittacidae pigeons ( Columbid ae , and ) , ) (2) if the geographic position of a continent and allies (Cuculiformes) among the relative to others has changed during non-passeriforms. Yet this limited with con- the history of a group of organisms, sideration as a basis, he was willing to conclude then faunal composition on any one in another paper (Cracraft, as 1972, p. 173, continental mass may reflect several well as 1973) that "the vast majority of Aus- episodes of exchange with other geo- tralian families undoubtedly have relationships graphic areas; those areas most recently to Asian families and therefore sup- can be in contact or closely apposed to one posed to have come from the north. This another should be expected to exhibit includes most non^ [italics mine] and the greatest faunal similarity. all oscines." Cracraft, like Serventy, did not present data that would support such a state- With these assumptions the practical clues ment, and thus presumably accepted much of examined in an attempt to determine the the reasoning previously articulated by Mayr dispersal routes to and from Australia utilized and Darlington. Unfortunately, all of these during the history of any one avian family workers have not stated the important point include: that similarity of the Australian and Asian avifaunas could well be due to a two-way, (1) phyletic relationships of the family to not just a one-way, exchange of old endemic others (accompanied by distributional faunas, and that present-day avifaunal similarity and degree of differentiation data for need only signal the latest, not necessarily the "other" families); sole, episode of interchange between Australia (2) fossil history of family, worldwide; and the rest of the world. Why not investigate (3) degree of endemism of family both the Australian nature of the Oriental avifauna within and outside of Australia (see as well as the Asian nature of the Australian figure 5); avifauna? It, also, should be realized that (4) diversity of family both within and out- because linkage between Australia and other side of Australia (see figure 5); southern continents was broken during or (5) nature of distribution: cosmopolitan, before the Eocene, birds on any of these pantropical, restricted; continuous or gondwanic continents today have had ample segmented. G2 5M PAT VICKERS RICH

How each of the above clues are interpreted extreme cases, i.e. where (1) a family is highly in the following study needs brief explanation: diverse, having a large number of endemic Phyletic relationships. If a group (or groups) genera/subfamilies in Australia but is un- can be identified as being closely related to a represented or has only a scant record else- family with Australian representatives, and where and (2) a family is highly diverse information is available concerning intragroup outside Australia but is represented in Australia phyletic relationships, then those data will by a single genus or species, often conspecific sometimes favour one dispersal route over with extra-Australian populations. another between Australia and the rest of the Case 1 may represent a group that has had world, although not always allowing resolution a long history in Australia and just recently of the direction of movement along that route. emigrated elsewhere, especially if the extra- An explanation of the observed distribution of Australian forms are closely (conspecific or at the group members (including the Australian the most congeneric) related to Australian taxa. taxa) attempts to determine the minimum The high diversity and endemism in Australia number of dispersals necessary (see Nelson, cannot be explained by that continent's greater 1969, for excellent discussion of this pro- environmental diversity in comparison to that cedure) to produce the present array of birds of its nearest continental neighbour today, the with the constraints imposed by understanding Palearctic-Oriental region. Certainly a greater of phylogenetic relationships and knowledge of topographic and climatic diversity, a decidedly Mesozoic and Cenozoie paleogeography. Such greater land area (ca. 3 million miles2 in explanations, however, offer only "parsimoni- Australia; ca. 21 million miles 2 in the Oriental- 2 ous" first approximations that may well be Palearctic; and ca. 33 million miles in the rclincd, and perhaps found to be decidedly Old World when Africa is included (Golen- more complex, if understanding of the phylo- paul, I960)), as well as a greater tropical

geny of certain avian groups is improved and/or land area characterizes the Palearctic-Oriental a more geographically and temporally complete Realms when compared with the Australian fossil record of birds materializes in the future. (excluding New Zealand). As previously men- Recent and fossil material are of equal import- tioned (also see below), the effect of differ- ance when constructing phylogenics, each ential competition encountered by an avian limited by incompleteness of specimens (or group in either area cannot be measured and study thereof) or of geographic representation. remains unknown in this consideration. Thus Geographic position of the most primitive taxa original entry of groups in this category into

(especially if based on living forms alone) is Australia via an Antarctic dispersal route not a priori assumed to be indicative of the should be considered a possibility along with centre of origin or early radiation of a group an Indomalaysian route. If the extra-Australian but must be examined in light of the distribu- forms are quite distinct (although few in num- tional patterns of other more primitive and ber), however, the situation may be more advanced members of the group. complex and will be discussed further under Unfortunately, this clue is of limited use each family affected. because of the poor understanding of phylo- Case 2 apparently represents the opposite genetic interrelationships of many of the non- extreme, a group that has only recently arrived passeriform taxa. This is well demonstrated by in Australia and has not yet differentiated to diversity of opinions expressed today concern- any marked degree. In this case the Indo- ing interfamilial relationships of birds, many malaysian or transoceanic route is most prob- of which are cited in the following section. able paths of original entry. Degree of endemism and diversity. Darling- Of course, these two cases represent the ton (1957) has posed some serious objections extremes of a spectrum of possibilities; many to the use of endemism and diversity (see avian families are moderately diverse, with previous section of this paper), but in this many endemic taxa in several areas of the study, such clues do seem helpful in two world, including Australia. Unless other data ANTARCTIC DISPERSAL ROUTES 95

are available, it may be impossible, as Darling- late Cenozoic in particular) have produced ton (1957) has suggested, to determine the sufficient fossil material to allow partial recon- most probable route of initial entrance for these struction of Cenozoic avifaunas. To some families at present. extent, early Tertiary European avifaunas can Nature of distribution. Distributional clues be used to suggest what avian taxa might have have been used as discussed by Darlington simultaneously lived in Southeast Asia, but (1957), i.e. the continuity or discontinuity of such would only be extrapolation in need of an area inhabited by a particular group can be confirmation, since barriers to overland dis- used to determine if a group has recently or persal may well have separated the two areas long ago invaded an area, where a disjunct during parts of the Tertiary (see Szalay and distribution is indicative of a group that has McKenna, 1971; McKenna, 1972a). Just be- invaded long enough in the past to have experi- cause such early Tertiary faunas are not pre- enced fragmentation into disjunct populations sently known, however, one should not con- separated by considerable distances. demn the value of fossils for the future. Were Two other clues that Darlington (1957) the records more complete, particularly in South employed in his considerations of avian zoo- America and Australia, questions regarding the geography are not particularly useful when antiquity of many groups in these areas could considering the origin of the Australian avi- be answered, as could those concerning which fauna because of the total lack of or rarity dispersal route between Australia and the rest of data for each: the distribution of "com- of the world was most likely for those groups. peting" families and the composition of fossil Were the record more complete, it could be avifaunas. used to evaluate parsimonious conclusions In the former case I know of no careful and based solely on phyletic relationships and dis- sufficiently broad-based studies designed to tribution of modern taxa, which may be gross 3 determine what avian families (or for that oversimplifications of reality. In making such a matter what other organisms, not just birds) statement, I am not advocating a cessation of are "competing" with certain Australian taxa research on avian phytogeny, which I think can both within and outside of Australia. How much (or how little) competition a taxon B A group that well exemplifies such oversimplifica- tion by parsimonious arguments based on the living encounters in different geographic areas may fauna alone is the mammalian order Perissodactyla well be the primary factor accounting for represented by the tapirs (tropical America, South- differential diversity in certain areas, but how east Asia, and Indonesia), the rhinoceroses (Africa, Asia, and Indonesia), and the horses and can this be recognized and measured? Darling- eastern allies (central and eastern Asia and Africa) in the ton very appropriately pointed this (1957) modern fauna (Walker, 1964). Of these living forms kept in mind to problem out, and it should be the tapirs are the most primitive and the horses most temper all conclusions regarding location of advanced or derived (Simpson, 1945). Thus the most early centres of radiation, as well as antiquity parsimonious explanation of origin and dispersal of this group would require an Old World origin of the of families in certain areas, particularly in the equids and the rhinoceroses and an Old or New where the early Tertiary fauna case of Australia World origin of the tapirs. The fossil record, however, may well have been impoverished and unbal- indicates the "reality" of perissodactyl dispersal was anced and where there is no fossil record to far more complex than the modern record intimates. confirm what groups were actually there. A wide variety of equids and rhinoceroses as well as a number of now extinct perissodactyl groups The fossil record, also, is not particularly inhabited North America and Europe, as well as dispersal helpful at present in determining what those areas now inhabited by this order at various route was used by birds present in Australia times during the Cenozoic. Without necessarily alter- during the Cenozoic because of the incomplete- ing the hypothesized phyletic relationships of the living perissodactyls, the fossil evidence certainly increases ness of the Paleogene record in Australia (see the possibilities for area of origin and early radiation America (Pascual and Table 3), South of both the horses and rhinoceroses, challenging the Rivas, 1971), and Southeast Asia. Only Europe specificity implied by the original parsimonious and to a lesser extent North America (mid to argument. 96 PAT VICKERS RICH

PLIO. PLEI. CRET. PALEOCENE EOCENE OLfGOCENE MIOCENE

... . . — i a 1 M e oe S i" g O CASUARIIDAE - DRO POD1CEPEDIDAE PELECANIDAE

PHALACR OCORACIDAE ANHINGIDAE ARDE1DAE C1CONIIDAE THRESKIORNITHIDAE PHOENICOPTERIDAE ANATIDAE ACCIPITRIDAE

F ALCONIDAE - MEGAPODIIDAE

1

PHA3IAN1DA I 1 TURNICIDAE PEDIONOMIOAE GRUIDAE RALLIDAE OTIOIDAE JACANIDAE HAEMATOPODIDAE CHARADRMDAE ARENARIIDAE

SCOLOPACIDAE RE 3URVIROSTRIDAE PHALAROPIDAE — - 1 1 8URHINI0AE | GLAREOUDAE —

ROS COLUMBIDAE PSITTACIDAE

TYTON1DAE- PODARQtO \E — AE09TMELIDAE CAPRIMULGIDAE APODIDAE 1 HEMIPROCNIDAE

BU(JEROTIDAE ALCEDINIDAE-

Fig. 10—Geologic ranges (world-wide) for those non- estimate is more complex) that cannot be passeriform avian families represented in the confirmed by, or deduced from, the modern Cenozoic of Australia. Horizontal axis cali- brated in millions of years, in general follow- avifauna alone (see Cracraft and Rich, 1972, ing Berggren, 1969, 1971. concerning the early radiation of the Cathar- also Nelson, only help in understanding of avian distribution, tidae in Europe; 1969). Despite the rarity of avian fossils in most but I am criticizing those who would condemn parts of the world, the palaeontological record the fossil record as a useless, even misleading item of information for zoogeographic con- is important to this study in one way. It demon- siderations. A good fossil record combined with strates that at least twenty of the forty-six data from the living fauna and continental avian families (see figure 10) that make up the reconstructions for the Mesozoic and Cenozoic, Cenozoic avifauna of Australia had probably on the contrary, can offer alternatives (and differentiated by the Eocene (data mainly from often a closer estimate of reality, even if that Brodkorb, 1963, 1964, 1967, 1971a) and thus ANTARCTIC DISPERSAL ROUTES 97

could have utilized the Antarctic dispersal route (see above), palaeontological data from Brod- to initially disperse between Australia and the korb (see above and 1971b) and Olson remaining world. (1974), and phylogenetic information from Brodkorb, Mayr and Amadon, Storer (1960, Summary of Phyletic Relationships, Diversity, 1971 ) , Woolfenden, Brown and Amadon, Endemism, Distribution, and Palaeontological Bock, and Sibley and Ahlquist (1972). The Record of Avian Families Having Australian latter reference summarizes most of the impor- Representatives tant works on avian phylogenetic studies and Using five the clues (phyletic relationships, will not be duplicated here. diversity, endemism, distribution, fossil record) Casuariidae (Dromiceidae, Casuariidae of Mayr stressed in the previous section, an attempt is and Amadon, 1951; Brodkorb, 1963; Casuarii- made in this section to determine for each avian dae of Sibley and Frelin, 1972). Emus and family with a Cenozoic record in Australia cassowaries; Miocene-Recent; Dromaius re- what the most probable route of entry has been. stricted to Australia, Casiiarius to Australia, Palaeontological data are given to emphasize New Guinea, and some Pacific islands; two which families are known to have differentiated genera in family, Dromaius and Casiiarius: by the early Tertiary, thus indicating the real greatest endemism and diversity in Australia; nearest extra-Australian relatives, elephant possibility that the Antarctic dispersal route birds and rheas-, not and kiwis; could have been used. Data for each family nearest relatives, the dromornithids (Rich, are presented in the following order: technical 1973). and common names; temporal range (both for world and Australia; if coincident only one The emus and cassowaries form a unique, range listed); geographic range; total number quite distinct and moderately diverse ratite' of living and fossil genera (if different from group in Australia whose nearest extra-Aus- following category); total number of living and tralian (New Guinea and Southwest Pacific fossil genera in Australia-New Guinea; geo- included) relatives occur in South America graphic area of greatest diversity; geographic and Africa (see Rich, 1973; Cracraft, 1973; area of greatest endemism (at generic, sub- Sibley and Frelin, 1972; Jehl, 1971; Glenny, familial levels); phyletic relationships to other 1965; Sibley, 1960). The family is also present, avian families. A brief analysis follows each set but neither as diverse nor endemic, in New of data for every family and attempts to estab- Guinea and on some southwest Pacific islands. lish which dispersal route, Antarctic and/or The only ratite known from Asia, the Indomalaysian, has been used. In many cases (Struthio), first recognized in Europe and both routes may have been utilized at different Africa in the late Miocene (Rich, 1972), is times during the history of an avian family far too specialized to have given rise to the while in other cases one route seems more Australian ratites, however. Casuariid differen- probable. In several cases, a choice of routes tiation had occurred in Australia by the Mio- cannot be made based on available data. cene, and therefore a Paleogene or earlier The familial classification employed follows differentiation is necessitated. More advanced that of Peters (1931, 1934, 1937, 1940, 1945) ratites were present in South America during with synonomies of the Mayr and Amadon the Eocene (Brodkorb, 1963; Rich, 1973; (1951) and the Brodkorb (1963," 1964, 1967, Cfacraft, 1973), and thus forms that could 1971a) classifications also listed. Generic have been ancestral to Australian ratites must assignment to each family follows Peters (see have differentiated even before, and an early above) except where modified by Bock (1956), dispersal of this group across a southern route

Woolfenden ( 1961 ) , Brown and Amadon must be considered a possibility. (1968), Rand and Gilliard (1968), Sibley Dromornithidae. Mihirung birds; Miocene- and Frelin (1972), Rich (1973) and Condon Pleistocene (presently extinct); restricted to (in press). Data on diversity and endemism Australia; 5-6 genera, including Genyornis were derived primarily from Peters and Condon (Stirling, 1896; Stirling and Zietz, 1896; 1900; 98 PAT VICKERS RICH

1905; 1913), Dromomis (Owen, 1872; 1874; and a second fossil genus, Liptornis (mid- 1879a-b) and 3 new genera (Rich, 1973); Miocene, South America); nearest extra- nearest extra-Australian relatives probably Australian relatives probably within the Pele- elephant birds (Aepyornithidac), rheas (Rhei- caniformes: Lanham (1947) believed the dae); nearest relatives the Casuariidae. Phaethontidae and Fregatidae, with recent pantropical distributions, were the most primi- The Mihirung Birds (Rich, 1973) or "giant tive members of the , and thus emus" appear to be most closely related to more primitive but closely related to the Pele- the Casuariidae (Rich, 1973) and both in turn canidae; other possible close relatives are the arc closer morphologically to the more primi- fossil families Odontopterygidae (Eocene of tive elephant birds and more advanced rheas- England), Pscudontornithidae (Miocene of Brazil or ostriches than to the moas-kiwis (Rich, 1973). Germany?, Miocene of North Amer- ica, Pliocene of New Zealand), Pelargornithidae The same biogcographic arguments that are (Miocene of Europe), Cladornithidae (Oligo- valid for the casuariids hold for the Dromorni- cene of South America), Cyphornithidae thidae, and thus an Antarctic route needs to (Miocene of North America). be considered a possibility for the ancestors of both groups. More diverse in the past (2 genera in the Miocene), the pelicans are represented by a Podicipedidae. Grebes; Miocene-Recent; Pleis- single genus in the living avifauna is tocene-Recent in Australia; cosmopolitan dis- that most varied in tribution; 6 genera in family (2 extinct); 1-3 the Old World (Africa-Europe-Asia). genera (Podiceps, ?Polhccphalus\ ?Tachyhap- One species occurs in, and is endemic to, tus) having cosmopolitan distribution, with Australia, as is another in the New World remaining genera in New World; nearest extra- (out of a total of seven species). Because of Australian relatives (at the family level) not the lack of distinct endemics (at the generic certain, perhaps the (Gaviidac) with an level) restricted to Australia and/or Australia- entirely Holoarctic distribution (first fossil Old World and the low diversity of the genus occurrences Paleocene of Europe, Oligocene of Pelecanus in Australia, entrance via the Indo- North America), Enaliornithidae (extinct, malaysian route or even oceanic dispersal Cretaceous of England), Lonchodytidae (ex- is more probable. Also, if tinct. Cretaceous of North America), Baptor- the Pelecanidae really were nithidae (extinct, late Cretaceous of North derived from forms similar to tropic birds America), Charadriiformes, Sphcniscidae, Pele- and frigate birds with tropical distributions, caniformes, Proccllariformcs, or . Indomalaysian dispersal would be favoured. Pelecanus tirarensis, the mid-Miocene form Three species of grebes (Podiceps ruficollis, from interior Australia, may well be close to P. cristatits, and P. polioccphcdus) are present the ancestral form that first entered Australia in the recent Australian avifauna. The first two (as it ncared Southeast Asia) and gave rise to have ranges extending into most of the Old the living P. conspiclllatus. World, and only P. poUocephalus is endemic Phcdanocoracidae. (Phalacrocoracinae of Mayr to Australia. For a number of reasons initial and Amadon, 1951). Cormorants; Paleocene- entrance via southward dispersal along the Recent; Miocene-Recent in Australia; family Indomalaysian route seems most probable: cosmopolitan; 2-3 living, 4 extinct genera in- (1 ) greatest generic diversity within the family cluding Graculavus ( Paleocene of North as well as specific diversity within Podiceps is America), Actiomis (Eocene of England), in the New World; (2) probable nearest rela- PUocarbo (Pliocene of Europe), and Plotop- tives to the family occur in Holarctica (includ- terum (Miocene of North America); one genus,

Phalacrocorax t with a ing North America); and (3) the family is cosmopolitan distribution, only member of family in represented by only one genus in Australia, Australia; fossil record of Phalacrocorax extends into early a genus that is not endemic or particularly to mid-Oligocene; living members diverse there. of family equally diverse and endemic in New and Old Pelecanidae. Pelicans; Miocene-Recent; one Worlds; nearest relatives the specialized An- genus {Pelecanus) with world-wide distribution hingidac (see below): closest relatives of the ANTARCTIC DISPERSAL ROUTES 99

cormorants-anhingas possibly contained in the reported from the Pleistocene of Australia suborder Sulae (see Brodkorb, 1963; in agree- (DeVis, 1905; confirmed by Miller, 1966a). ment with Lanham, 1947) including the Because of their present and past (see Rich, Elopterygidae (Cretaceous-Eocene of Europe) 1972, 1974) restriction to warm, wet climates, and the Sulidac (first occurrence in the Oligo- an lndomalaysian dispersal route seems most cene of Europe, Miocene of North America; probable. cosmopolitan at present). At no time during the late Mesozoic or early Tertiary could the Antarctic route have Although Phalacrocorax carbo suggests by had a climate that was more equable than its cosmopolitan distribution that an Indo- temperate. The warm, humid tropical climate malaysian or oceanic route has been used most presently characteristic of some parts of recently by the cormorants, direction of move- northern Australia and New Guinea is a mid ment along that route is not certain. The Phala- to late Tertiary innovation as the Australian crocoracidae are quite an old group, first recog- plate drifted into the lower latitudes. The nized in the late Mesozoic/Paleogene of North occurrence of Protoplotus, thought to be an America and Europe, as is Phalacrocorax, early member of the Anhingidae (Lambrccht, is first which known in the mid-Oligocene. 193 1 ), in Paleogene sediments of Sumatra does Unfortunately, none of the practical clues used not contradict the above conclusion. in this study favour one route over another. Ardeidac. Herons and allies; early Eocene to Phyletic relationships of the phalacrocoracids, Recent; no fossil record in Australia; family including Phalacrocorax, to other birds (other cosmopolitan; 27 living, 8 extinct genera: 9 than to the more specialized tropical anhingas) living genera in Australia-New Guinea, none are not well understood, diversity and endem- of which are endemic; one-third of genera in ism are about uniform the world over, and family with cosmopolitan distribution; greatest diversity and endemism in both distribution is nearly continuous along con- New and Old Worlds, similar in both areas; nearest relatives tinental margins and somewhat inland over perhaps the cosmopolitan Ciconiidae (Ligon. most of the world except in the far north and 1967. would disagree) (first appear in Eocene far south. To further complicate matters, the in circum-Mediterranean area; Oligocene in fresh-water cormorants are not restricted to South America; Miocene in North America; environments, and oceanic dispersal could Pliocene in Asia; Pleistocene in Australia) or easily have played an important part in their the nearly cosmopolitan Threskiornithidae (first dispersal with regard to Australia. occur in Eocene in Europe; Miocene in South

Thus, with present evidence, it appears im- America; and Pleistocene in North America, Africa, Australia). possible to determine the route initially used and by the cormorants between Australia and the Herons and allies have had a long history, rest of the world. the family certainly being differentiated by the Anhingidae. (Anhinginae, Mayr and Amadon, early Tertiary (Eocene of Europe and North 1951 ). Anhingas, darters, or snake birds; America; Eocene-Oligocene of Africa). The ?Eocene-Recent; Miocene-Recent in Australia; most recent movement of this group has been 2 genera in the family (Protoplotus, restricted across the lndomalaysian route as exemplified to ?Eocenc of Sumatra); one genus, Anhinga, by the (Bubulcus = Ardeola) that in Australia, has pantropical distribution; family has reached Australia from Asia in the last presently with similar diversity and endemism few years (Crosby, 1972). The remaining throughout tropics of the world; nearest rela- ardcids, except for Zonerodius (restricted to tives the Phalacrocoracidae, with a cosmopolitan N.G.-SW Pac.)< are all wide ranging genera, distribution. most of which are decidedly more diverse The one species of anhinga (A. melano- outside of Australia. Diversity and endemism gaster), occurring in Australia, also extends of the family is about equal in both the New throughout tropical Asia and tropical Africa, and Old World while low in Australia, all of being distinct from the New World A. anhinga. which suggests that the Ardeidae dispersed A second distinct species, A. laticeps, was southward from Palearctica into Australia 100 PAT VICKFRS RICH

into Pleistocene; greatest diver- and/or may have utilized an oceanic route. terus persisted sity of family presently in South America and Ciconiidae. .Storks; Eocene-Recent; Pliocene- Africa but, when fossils considered, also in Recent in Australia; family cosmopolitan except Australia; nearest relatives the Palaelodidae for New Zealand; 1 I living, 9 extinct genera (Miocene-Pliocene of Holarctica), Agnopteri- only one of which {Xctiorhyncluts) is known dae (Hocene-Oligocene of Europe); Telma- Australia; greatest diversity and endemism in batidac (early Eocene of South America), family Old World (Africa and Asia in Of in Scaniornithidae (early Palcocene of Europe), particular, although three genera known in Torotigidac (late Cretaceous of Holarctica), New World); nearest relatives probably Ardei- and more distantly the Ciconiidae and Thres- dae or Threskiornithidae (see above), both of kiornithidae, and probably the Anseriformes that are nearly, which have recent distributions (IXlacour and Mayr, 1945; Hopkins, 1949) and long fossil records if pot, cosmopolitan and Ciconiiformes (Mayr and Amadon, 1951; Recent), or Cathartidae ( Ligon, (Eocene to Glennys, 1955; Sibley, 1960) (both orders, 1967). Mamardi, 1902, 1963) all of which have cosmopolitan ranges but differing degrees of Owing to the low diversity (one endemic diversity and endemism in different parts of genus with one species) of this family in the world. Australia and its high diversity and endemism in neighbouring Asia (as well as the remaining Although not members of the living avifauna Old World), entrance of the Pleistocene-Recent of Australia, during the Tertiary the flamingos members of this group from Southeast Asia via formed a moderately diverse group on that the Indomalaysian route seems most probable. continent, including one endemic genus {Phoe- group Threskiomithidae* (-- Plataleidae of Brodkorfc, niconotius). During the Pleistocene, this 1963). Ibises and spoonbills; Hocenc-Rcccnt; became extinct in Australia (Miller, 1963b), Pleistocene-Recent in Australia; 17 living, 3 but survived in other parts of the world in extinct genera; 3 living genera in Australia; varied climatic zones from tropical to tem- diversity and family cosmopolitan; greatest perate. All of the Australian genera, except in Old World with 7 genera known endemism Phoeniconotius, are also represented in the from New World; nearest relatives, Ciconi'ulae Old World {Phoeniconaias restricted Africa), or Ardeidae, both with nearly or completely or over most of the world in tropical to tem- Cosmopolitan distributions and long fossil his- perate climates {Phoenicopterus). At present tories (Eocene to Recent). the family does not occur in Southeast Asia repre- The three genera of threskiornithids and the East Indies, being restricted further Australia have either cosmopolitan sented in west in Eurasia and Africa in the Old World. (Plegadis) or wide ranges in the Old World Because the family had differentiated by the is endemic (Platalea, Threskiornis), and none early Tertiary in both Europe and South specifically diverse in, Aus- to, or any more America, and is at present equally diverse and such low tralia than elsewhere. Because of endemic in both geographic areas, both the diversity and the lack of endemic generic or Indomalaysian and Antarctic dispersal routes suprageneric taxa in Australia at present, an must be considered seriously. The Indomalay- Indomalaysian route for dispersal of those sian or an oceanic route has probably been forms in Australia appears most probable. utilized most recently, accounting for the Phoenicoptaridae. Flamingos; late Eocene- Pliocene-Pleistocene similarity at the generic Recent; Miocene-Pleistocene in Australia (pre- level of Phoenicopterus (in fact Miller (1963b) sently evtmet); nearly cosmopolitan; 3 living, suggested specific synonomy with P. ruber in extinct fossil genera {i'Jornis, Hocene-Oligo- 3 the Pleistocene) as well as close resemblance eene of Europe; TiUornis, Oligoeene of South of the Australian Pliocene Phoeniconaias and America; Phoen'uonotius, Miocene of Aus- the same genus occurring in Africa today. tralia); no living phoenicopterids in Australia Whether this was brought about strictly by a but an endemic genus (PfweriicanQtius) and during the Miocene (or Phoemcoplerus present during Miocene, Phoeni- northward movement in the case of Phoenicopterus conaias in the late Pliocene, while Phoenicop- slightly before) ;

ANTARCTIC DISPERSAL ROUTES 101

(first recorded in Europe during the early primitive member of the Anserinae (swan- Miocene) and then or perhaps later in the goose subfamily). Others would disagree, how- case of Phoeniconaias, or whether movement ever, and ally it with the sheldrakes (Tador- was southwards from Eurasia, or both (different nini) (Delacour and Mayr, 1945, 1946). for each genus), is at present unclear. Since Cereopsis, like Anseranas, is restricted to the extinct European Tertiary forms Elornis Australia presently, with a probable Pleistocene and Tiliornis have not been thoroughly re- record of the family in New Zealand (evidenc- examined since the late 19th century, it is ing ability of this group to cross water barriers uncertain how closely cither is related to the in the past if interpretations of the New Zealand Recent and other fossil genera from Europe, form are correct). The Dendrocygnini (tree- Africa, Australia, and the New World. At ducks) are considered more advanced toward present, then, which was the initial route of the goose-swan condition than Cereopsis but dispersal between Australia and the remaining more primitive in many characters than the world has not been determined. Cygnini and Anserini themselves. Within the tribe of tree-ducks (Woolfendcn, 1961) one Anatidac (including Anscranatidae of Woolf- very distinct genus, enden, 1961). Ducks, geese, swans; Eocene- Stictonetta, is at present Recent; Miocene-Recent in Australia-New restricted to Australia, while Dendrocygna Guinea; 51 living, 23 extinct fossil genera, occurs pantropically, including Australia. The 13 genera in Australia-New Guinea of which tribe is first recorded in Miocene sediments of Anseranas, Stictonetfa, Malacorhynchus, Cere- South Dakota, first known in Australia in the opsis, Chenonetta, and Biziura are endemic; Pleistocene. Frith (1964a-b), on the other family cosmopolitan and only slightly more hand, has allied this form with the swans and diverse in Old World than elsewhere; many thus removed it from the Dendrocygnini. Since genera with Holarctic distribution; nearest the most advanced subgroups within the Ana- relatives probably the Anhimidae (screamers), tidae proper had differentiated by the end of presently restricted to South America. the Paleogcne, ancestors of more primitive Entrance of the many different groups of members of the Anseriformes discussed above Anatidae into Australia has probably been must have been present even earlier, and thus

complex, possibly involving both the Antarctic a southern dispersal of this group is a possi- and the Indomalaysian routes. Differentiation bility. Then, too, such forms could have been of the family occurred early, at least by the relict in Australia after utilizing an Indo- Eocene and probably earlier. The most closely malaysian route or even oceanic dispersal. related family to the anatids is the Anhimidae, The true geese (Anserini), are presently present only in South America today with a restricted to Holarctica, apparently never Pleistocene record on that continent. In addi- reaching Australia, while the remaining tribe tion, many anseriforms in the Australian avi- of the Anserinae, the swans (Cygnini) have a fauna are primitive members of the order. nearly cosmopolitan distribution. The Cygnini The magpie geese, (Woolfenden, are first known from Oligocene sediments in 1961; Johnsgard, 1961, 1962; Sibley and Europe, from the Pleistocene elsewhere. This Ahlquist, 1972) or Anscranatinae of many early occurrence in Europe coupled with the classifications (Mayr and Amadon, 195 1 group's maximum diversity in Eurasia presently Delacour, 1954), presently restricted to Aus- and its probable derivation from a strictly tralia, are anseriforms somewhat advanced over Holarctic group (Anserini) all suggest south- the anhimid condition but not so highly ad- ward dispersal via the Indomalaysian route. vanced as the true ducks-gcese-swans (Wool- Members of several tribes of the Anatinae fenden, 1961; Verheyan, 1953, 1955; Sibley (Tadorini, Anatini, Aythyiini, Oxyurini of (1960) has noted their distinctiveness from Woolfendcn (1961)) including both primitive other anatids). The (Cere- and advanced forms are present in Australia opsis) is considered by some (Woolfenden, today. This subfamily was distinct at least by 1961; Johnsgard, 1961, 1962) to be the most the late Oligocene when it was present in 102 PAT VICKERS RICH

Eurasia, in Africa by the early Miocene. there is apparently relatively short, and again Primitive members of the Anserinae that prob- Indomalaysia seems to be the only major ably most closely resemble the common ances- dispersal route utilized. tor of the Anserinae-Anatinac (Woolfenden, The fourth anatine tribe, the Aythyiini, is 1961) occur in Australia (Cereopsini) and represented by only the cosmopolitan Aythya pantropically (or nearly so: Dendrocygnini). (by one endemic species) in Australia, a genus

It is interesting that one of the more primi- much more diverse elsewhere in the world. tive tribes of the Anatinae (Woolfenden, 1961), The tribe is most diverse in Asia, most endemic the Oxyurini (stiff-tailed ducks), is most diverse in the Old World, and its earliest fossil occur- in South America (Heteronetta, Nomonyx, rence is in the early Miocene of Europe. Oxyura) while Africa and Australia each have Woolfenden (1961) believed that the Mergini an endemic genus (M.-Thalassornis; Aust.- (restricted both at present and in the past to Biziura), and both share Oxyura, the only form Holarctica) were the nearest, more primitive present in Holarctica. Such a distribution could relatives of the Aythyiini. All these data strongly be explained either (1) by initial use of the suggest that the Aythyiini entered Australia via Antarctic and then a northward dispersal Indomalaysia. during the Miocene into Asia and across In summary, the entrance of the Anatidae tropical southern Asia, Europe, and North into Australia has probably been complex and Africa into the present Ethiopian realm before may have involved both the Antarctic and restriction of the tropics in the late Tertiary, Indomalaysian dispersal routes. The most recent or (2) by southern dispersal into Australia exchange, which is still in progress, undoubtedly from southeastern Asia in the mid-Tertiary involves only the Indomalaysian route. and differentiation since that time. At present Accipitrichte. Hawks and eagles; Eocene-Recent; it difficult is to make a choice between the two Miocene-Recent in Australia: family cosmo- possibilities. politan; 63 living, 24 extinct genera; 17 living Other tribes of the Anatinae, however, have genera in Australia-New Guinea of which apparently arrived in Australia via southward Lophoietinia, Hamirostra (all Milvinae), movement along the Indomalaysian route. The Erithrotriorchis (Accipitrinae), and Harpyopsis Tadornini (sheldrakes), have a cosmopolitan (Buteoninae) are endemic; Henicopernis (Per- ninae) is endemic to New Guinea and the distribution, but no one genus is cosmopolitan Southwest Pacific, Haliastur to Australia-New (2 endemic to South America, 1 to Africa, Guinea-Southwest Pacific-Asia, and Butastur to 1 to Eurasia, and 2 occur over most of the New Guinea-Asia-Africa; most closely related Old World including Australia). Low diversity families Falconidae, Pandionidae, also with and lack of endemism in Australia, as com- cosmopolitan distribution. pared to that of the Old World, indicate that the group has moved south, instead of north The Falconiformes is the most generically along the Indomalaysian route. diverse order of non-passeriform birds, contain- The Anatini are also cosmopolitan, although ing some 81 genera in the Peters (1931) classi- several genera are endemic to different parts of fication and slightly less in that of Brown and the world including Malacorhynchus (whose re- Amadon (1968). Within this order, the Acci- lationships are poorly understood, R. Schodde, pitridae is the most diverse family with more pers. comra., 1974) and Chenonetta in Aus- than 60 genera (63, Brown and Amadon, 1968; tralia; Cheniscus is endemic to Southeast 68, Peters, 1931), equalled in intrafamilial Asia-Southwest Pacific-Australia-New Guinea. diversity only by the parrots and pheasants. Diversity and endemism in the Old and New About twenty-five per cent of known accipitrid World are nearly equal, with that in Australia genera occur in Australia and come from several being extremely low when compared with the of the subfamilies within this family including other continents. Because of such lack of three types of kites (Perninae, Elaninae, Mil- diversification in Australia in comparison to vinae), the fish eagles (Valiaeetus), the har- the rest of the world, the history of the Anatini riers (Circinae), and the hawks and eagles ANTARCTIC DISPERSAL ROUTES 103

(Accipitrinae). Conspicuously absent in this N.G.-Aust.-SW Pac), Machaerhamphus (Af.- avifauna, however, are any of the primarily As.-N.G.-SW Pac), and the Australian endemic scavenging forms in the subfamily Gypaetinae, Henicopernis (N.G.-SW Pac). Three other widespread in the Old World. genera are endemic to the New World and a Most intriguing with regard to Australian fourth occurs in the Old World and the South- are the milvine kites, thought to west Pacific. As with the milvines, an Antarctic be some of the most primitive accipitrids route for the subfamily should not be excluded (Brown and Amadon, 1968). Of a total of from consideration, but neither should the seven genera known, four occur in Australia, Indomalaysian route. The subfamily is about two of which {Lophoictinia and Hamirostrd) equally diverse in Southeast Asia and Aus- are endemic to that continent. Haliastur occurs tralia-New Guinea-Southwest Pacific, although only in Australia - New Guinea - Southwest Aviceda is much more diverse specifically in Pacific-Southeast Asia, while the fourth genus, Southeast Asia than in Australia. , is found over most of the Old World Only one genus, Circus, represents the Cir- in addition to Australia and the Southwest cinae in Australia. This genus has a cosmo- Pacific. The remaining three genera are re- politan distribution, and is most diverse in the stricted to the New World. Thus, the main Old World, which suggests use of an Indo- diversity of this group occurs in the New World malaysian dispersal route, with birds moving and Australia with only two genera of the seven southward from Asia. reaching Southeast Asia and one reaching The Accipitrinae are represented by seven Europe. The fossils are consistent with the genera in Australia and New Guinea: those recent form with Milvus represented in the with (1) a nearly or entirely cosmopolitan early Miocene of Europe and two presently distribution (Accipiter, Spizaetus, Aquila); extinct genera restricted to the New World. (2) an Old World-Australian and/or New Certainly more advanced members within the Guinean distribution (Buteastur, Hieraaetus); Accipitridae had differentiated by the Eocene and (3) a range restricted to Australia and/or (Brodkorb, 1964), thus signalling an even New Guinea (Harpyopsis, Erythrotriorchis). earlier differentiation of ancestors. Such More primitive members of the subfamily evidence suggests that an Antarctic route for (Brown and Amadon, 1968), including the entrance of the milvines should not be dis- harriers and harrier hawks, are cosmopolitan. counted. The Indomalaysian route is presently Members of the subfamily Circaetinae, prob- in use and could have been as early as the ably most closely related to, yet more primitive Miocene. Movement across this route, however, than, the accipitrines are restricted to the Old may just as well have been northwards from World. Since the accipitrines have very similar Australia, rather than southwards from Eurasia. diversities (more than ten genera each out of Elanus, with a nearly cosmopolitan distribu- a total of more than 30) in South America, tion, represents the only member of Elaninae Africa, and Asia but has lower diversity and (also thought to be primitive accipitrids, Brown little endemism in Australia, an Old World and Amadon, 1968) in Australia. Two other origin seems most plausible for the Australian genera are in this subfamily, one genus endemic forms. The Indomalaysian archipelago may to the New World, and a second endemic to well have been the only route utilized by Africa. Owing to the cosmopolitan nature of accipitrines now known in Australia between Elanus and the lack of any more than specific that continent and the rest of the world. of this widespread genus in differentiation Pandionidae. Ospreys; Pleistocene-Recent; only scriptus), southward Australia (E. notatus, E. Recent in Australia; one genus (Pandion), dispersal along the Indomalaysian route seems which has nearly cosmopolitan distribution probable. including Australia-New Guinea: family prob- The third group of primitive accipitrids, the ably most closely related to Accipitridae pernine kites, include seven genera, three of (Hudson, 1948; Jollie (1953) uncertain about with falconiforms; Compton which occur in Australia: Aviceda (Af.-As.- relationships .

104 PAT VICKERS RICH

(1938) allies with Cathartidae), which likewise to Australia and the southwestern Pacific. They has cosmopolitan distribution. are thought by some workers to be primitive this most closely related to The osprey (Pandion haliaetus) is the only within order and chachalachas (; member of this family and has a nearly cosmo- the South American et al. 1959, politan distribution, but does not occur in New Cracraft, 1972, 1973; Hudson f suggested, Zealand. Its record extends into the Pleistocene 1966; Chandler, 1916). Others have of Europe and North America but no further. on the contrary, that megapodes are specialized (Clark, or that cra- Due to its widespread nature and singular lack phasianids 1960, 1964a~b) cids are closely related to pheasants than of diversity, it might be thought of as a more Ahlquist, relatively recent development (Ricklers and either is to the megapodes (Sibley and Cox, 1972; Mayr, 1963). Thus, the Indo- 1972). Providing that megapodes are actually malaysian dispersal route has most probably primitive galliforms, they may be assumed been involved and not the Antarctic. Whether to have differentiated by the Eocene (Brod- the direction of movement was north or south korb, 1964) because pheasants had appeared can be questioned, and until the relationships by this time. In such a case, an early southern of the pandionids are more clearly understood, dispersal of the mound-builders would have an Australian origin cannot be ruled out. been a distinct posibility. On the other hand, south across Falconidae. Falcons/ caracaras. Miocene-Recent; the group could also have moved ?Pleistocene-Recent in Australia; family cosmo- the Indomalaysian route and been isolated there

politan; 1 1 living, 3 fossil genera; 1 living at some time during or after the mid-Tertiary. genus (Falco, nearly cosmopolitan) and pos- Both hypotheses are viable but neither is more 9 sibly 1 endemic fossil genus (Plioaetus ) in probable than the other in light of available Australia; most closely related family, Accipi- data. tridae (also cosmopolitan in distribution) or In Australia two living and one extinct (a even the Strigiformes (Sibley and Ahlquist, giant Pleistocene form, Progura (van Tets, 1 972) (Eocene-Recent, cosmopolitan) 1974a)) genera are endemic. Three other The Falconidae, most diverse in the New genera (Megapodius, Telegalla, Aepypodius) World, is represented in Australia by a single occur in Australia and/or New Guinea-South- living genus and possibly one endemic form, west Pacific, while Macrocephalon and Eulipoa thought by most workers to be congeneric with occur only in the Southwest Pacific. Falco. Falco is quite diverse world-wide, with Phasicin'tdae. ( of Mayr and Ama- only seven species in Guinea. Australia-New don, 1951; Brodkorb, 1964). Quail, pheasants; Thus, it has most probably utilized the Indo- early Oligocene-Recent; Pleistocene-Recent in malaysian route, moving southwards from Asia. Australia/New Guinea; two subfamilies—Phasi- Megapodiidae, Mound builders, brush turkeys; aninae, restricted to the Old World and Austral- Pleistocene-Recent; family restricted to Aus- asia; Odontophorinae, restricted to the New tralia, New Guinea, and the Southwest Pacific; World; 59 living, 8 extinct genera; 4 living

7 living, 1 fossil genera; 6 living genera (2 genera in Australia-New Guinea, of which only endemic: Leipoa and Alectura) and 1 fossil Anttrophasis is endemic (restricted to New (Progura) from Australia; family with about Guinea at present but has Pleistocene record in equal diversity and endemism in Australia- south-eastern Australia); closely related to the New Guinea and the Southwest Pacific; most Numinidae (guinea fowl, restricted to Africa closely related family Cracidae (curassows and presently, Pleistocene record in Europe), the chachalacas) of tropical America (Hudson et Holarctic Tetraonidae (grouse, first occurs in ah, 1966, 1969; Holman, 1964; Cracraft, 1972, early Miocene of North America, Pleistocene 1973) although some would dispute this rela- in Europe), and the North American Melea- tionship (see refs. in Sibley and Ahlquist, gridae (turkeys, first occur in Miocene of 1972). North America); more primitive but related to all the above are the Cracidae-Megapodiidae The mound-builders are galliforms endemic (see above). 9 Named by DeVis and needs restudy to ascertain its taxonomic position. The family Phasianidae is first known in the ANTARCTIC DISPERSAL ROUTES 105 early Oligocene of North America and Europe which dispersal route (s) may have been util- (different subfamilies on each continent). In ized by ancestral turnicids or pedionomids. Europe it was taxonomically diverse in the mid Similarity between the Turnicidae of the Old to late Cenozoic and is most diverse in Asia World and Australia could be accounted for and islands of the Southwest Pacific today. by a mid-late Tertiary interchange, either All of the genera of phasianids occurring at northwards or southwards, along the Indo- present in Australia belong to the Old World malaysian route, and the differences in diversity subfamily, Phasianinae. One endemic genus between Australia and Africa are not great (Anurophasis) has diverged from the Old enough to significantly favour either possiblity. World forms, but Coturnix, Synoicus, and Gruidae. Cranes; Eocene-Recent; Miocene- Excalfactoria also occur in the Old World Recent in Australia: family nearly cosmopolitan and Southwest Pacific. Since the Australian except for South America and New Zealand; diversity and endemism are so low, the Asian 4 living, 8 extinct genera; 1 genus (Grits) in and Southwest Pacific diversity so high, coupled Australia; family most diverse and with greatest with the conspecificity of three of the four endemism in Africa; nearest relatives within the , possibly Australian genera with Old World forms, dis- the Aramidae (limp- kins) of the New World (first recorded in persal southwards along the Indomalaysian Oligocene of North America (R. Ernry, S. route is the most probable explanation for Olson, pers. comm., 1972), the Psophiidae the pheasants and quail presently known in (trumpeters, presently restricted to South Australia. America, with no fossil record), or the Turnicidae. Bustard quail; Pleistocene-Recent; Eogruidae (early Oligocene of Asia (see Cracraft, 1 2 genera (Turnix, Ortyxelos), only Turnix 969 ) ) , Rallidae, Heliornithidae, occurs in Australia; family restricted to Old Eurypygidae. World, Southwest Pacific, and Australia/New The family Gruidae had differentiated by the Guinea with a fossil record in Asia and early-mid Eocene in Holarctica and apparently Australia; most closely related family, the Aus- was present in Australia by the mid-Miocene tralian endemic Pedionomidae, nearest relatives (Stirton, Tedford, and Woodburne, to birds outside of Turnicidae-Pedionomidae 1968). uncertain (see Bock and McEvey, 1969) but In the Recent fauna, the Gruidae are most most likely lie within the Gruiformes. diverse in the Old World, with only the genus Pedionomidae. Collared hemipode; no fossil Grus extending into the New World and Aus-

record; 1 genus and species present in Aus- tralia. Grus is most varied in Asia, with only tralia; most closely related family, the Turni- two of the total nine species in the genus cidae. occurring in Australia-New Guinea (including

the endemic G. rubicund) . Thus, based on the The bustard quail (Turnicidae) are generi- low endemism and diversity of the Gruidae in cally most diverse in Africa where both genera Australia, the high diversity in the Old World, in this family occur. Turnix is the only genus and lack of conflicting evidence from the fossil that reaches Australia and is itself most diverse record, southward movement over the Indo- there and in the Southwest Pacific. With this malaysian route would best explain the present evidence alone, the most likely path of inter- record of cranes in Australia. change between Australia and the rest of the Rallidae. Rails; Cretaceous or Eocene-Recent10 world is Indomalaysian. But in which direction ; probably mid-Miocene-Recent in Australia; did the movement occur? The Pedionomidae, family cosmopolitan; 51-52 living, 27 extinct endemic to Australia, may well be the most genera; 13-14 living genera in Australia; pos- closely related, primitive yet perhaps more sibly one extinct fossil genus (Stirton, et ah, group (retains hallux; Bock and McEvey, phylo- 10 1969), to the Turnicidae. With lack of If Tetmdtornis included in the Rallidae (fide genetic analyses that point to the group of Brodkorb (1967)), family first known in Cretaceous; Cracraft (1969) has presented convincing arguments birds most closely related to the Turnicidae- for placing this genus in a separate family, and thus Pedionomidae, it is not possible to suggest the first record of the Rallidae is Eocene. ,

106 PAT VICKERS RICH

1968) in the mid-Miocene of central Australia; early Tertiary (no fossil record corroborates greatest diversity in Southwest Pacific region this, however) that did not successfully cross with slightly less diversity in Old World and the Antarctic sweepstakes route into Australia. New World; greatest number of endemic genera Otididae. ; Eocene-Recent; late Plio- in South America, but nearly as many in cene-Recent in Australia; family restricted to Southwest Pacific; closest relatives of Rallidae Old World and Australia-New Guinea; 11 lie within the Gruiformes, perhaps the Gruidae, genera in recent avifauna, 1 extinct fossil Aramidae, Psophiidae, Turnicidae, Heliornithi- genus; 1 living genus (Eupodotis — Ardeotis) dae, Eurypygidae (see above). in Australia; most closely related family prob- ably the extinct Gryzajidae (early Pliocene of At least nine genera of rallids are present in eastern Europe); other closely related forms the living avifauna of Australia, of which three probably within the Gruiformes, perhaps the are endemic to Australia and/or New Guinea: Cariamidae (seriemas, Pleistocene-Recent, South Tribonyx ( — Gallimda in Condon, 1973; America). rarely a vagrant to New Zealand), Rallicula, and Megacrex. To this list can be added Eula- Bustards, restricted to the Old World and beomis and Gymnocrex that are endemic to Australia and particularly diverse in Africa, Australia-New Guinea and some islands in the are first recognized in the middle Eocene of Southwest Pacific. All of these genera contain Europe. Modern genera first appear in the only one species in Australia, except Rallicula Miocene (Chlamydotis) and Pleistocene (Otis, with four. Three other genera range through Tetrax). Only one species of one genus (Eupo- Australia-New Guinea-Southwest Pacific-Asia dotis australis) is known in Australia, and its

(Rallina, Amaurornis, Poliolimnas) , while the entrance seems most probably to have been via remaining forms have cosmopolitan (Rallus, southward expansion along the Indomalaysian Porzana, Gallimda, Fulica) or nearly cosmo- route perhaps as early as mid-late Pliocene. politan (Crex, Parphyrio; not found in the Jacanidae. Jacanas; Miocene-Recent; no fossil New World ) distributions. Although genera record in Australia; family pantropical; 5 living, that occur in the living avifauna of Australia 1 extinct genera; 1 genus (Jacana) living in have very brief (Pliocene or younger) fossil Australia at present; greatest diversity and records, other members of the family are known endemism in Old World; closest relatives prob- far back into the Tertiary. ably the Phegminornithidae (early Miocene, If identifications are correct, well differen- North America); other close relatives within the Charadriiformes (possibly the Rostratulidae tiated rails were present in Europe during the (Jehl, 1968), first recognized in Eocene of Eocene and Oligocene when Australia was Europe, at present cosmopolitan except North still attached to Antarctica. This information, America or the Scolopacidae (see below)). coupled with the low degree of endemism and lack of speciation of endemic forms in Aus- Only one genus (and one species) of Jacani- tralia, suggests that the Indomalaysian route or dae is present in Australia at present. Peters a transoceanic route utilized late in the Ceno- (1934) recognized an endemic genus, Iredi- zoic was probably responsible for rails presently parra (L gallinacea), for the Australian-South- known in Australia. The great diversity and west Pacific, but Slater (1971) included this high endemism in the Southwest Pacific is form in lacana, which otherwise has a range probably due to geographic isolation imposed restricted to the New World (where it has a by the insular nature of the region as well as Pleistocene to Recent record). a long history in this area, most of which has Restudy of the Australian form is in order, been closely associated with Asia through the however, to determine if it really is closer to Cenozoic; the diversity in South America may the New World or Old World forms. It appears be due to a combination of environmental that the Indomalaysian route was employed diversity as well as a long history of the group (with southward movement) in the initial en- there. Perhaps the rallids were one of the trance of this family into Australia because of groups present in South America during the the low diversity in Australia and the greater ANTARCTIC DISPERSAL ROUTES 107

diversity in the Old World. Also the present not occur in the New World), and all except restriction of the family to the warm, humid Vanelius have only winter ranges in Australia. tropics argues against the previous use of a Only Peltohyas, with one species, is endemic to cool, temperate high latitude dispersal route. Australia. The high diversity and endemism of Haematopodidae. (Haemalopodinae in Scolo- this group in the Old World in comparison to pacidae of Brodkorb, 1967). Oyster-catchers; that of Australia suggest that the plovers most Miocene-Recent; no fossil record in Australia; probably dispersed south along the Indomalay-

family cosmopolitan; I genus; closest relatives sian route or across oceanic barriers rather late probably one of the following: Palacotringinae in the Cenozoic. The Paleogene occurrence of (Cretaceous-Eocene, North America and Eur- the charadriids in Holarctica does not contra- ope), Scolopacidae (cosmopolitan at present; dict this conclusion. first occurs in Eocene of Europe), Aphrizidae (= Arenariidae; no fossil record, nearly cos- Arenariidae (often included in the Charadriidae mopolitan but not in Africa), Charadriidae or Scolopacidae). Turnstones; no fossil record;

(cosmopolitan, first occurs in early Oligocene family cosmopolitan; 2 living genera; 1 genus of North America and Europe), Phalaropidae (Arenaria) in Australia; most diverse and (Holarctic, no fossil record), Rostratulidae endemic in North Ameriea; nearest relatives (sec above), Burhinidac, and Recurvirostridac possibly the Charadriidae or Phalaropodidae (see below). (Jehl, 196S) (see above).

The oyster-catchers are represented by two Since the main centre of diversity (which is species in the only genus of the family, tlaema- low) of this family is in North America and topus, in the Australian avifauna. One species only one cosmopolitan species (Arenaria inter- (H. juliginosus) is endemic to Australia, while pres) occurs in Australia, dispersal to Australia the other is nearly cosmopolitan. The group is most likely occurred as a southward expansion first recognized in the early Miocene of North of the group along the lndomalaysian route. America. Certainly the fndomalaysian route Seoiopaeidae. (Scolopacinae of Mayr and Ama- accounts for the present cosmopolitan nature don, 1951; Brodkorb, 1907). Snipe, sandpipers; of Haematopus, but what the history of the Eocene-Recent; no fossil record in Australia.; family cosmopolitan; 27 living, 3 extinct family has been and where it originated are not

genera; I 1 living genera in Australia; New and clear. The appearance of the genus in North Old World with about equal diversity and America in the Miocene could be owing to endemism, including Australia and the South- origin there or somewhere else, even Australia, west Pacific; about one-third of genera in and thus whether movement along the Indo- family cosmopolitan; closest relatives probably malaysian route of Haematopus has been north among the following: Palaeotringidae (see, or south, and which route was used by the above), Rostratulidae (cosmopolitan except for ancestor of the oyster-catchers remains to be North America, first known in Eocene of determined. Europe) ; Charadriidae, Aphrizidae, Phalaro- podidae, or Haematopodidac (see above); Charadriidae. (Charadriinae of Mayr and Jehl (1968) suggests the Ciallinagoninae and Amadon, 1951; Brodkorb, 1 . Plovers; 967) Calidrinac. Oligocene-Reccnt; no fossil record in Australia; family cosmopolitan; 31 living, 3 extinct genera; The Scolopacidae are first known in the 5 living genera in Australia, most of which arc Eocene of Europe. Of those genera in Aus-

winter residents only, with one endemic genus, tralia at present, Litnosa is first known from Peltohyas; greatest diversity in Old World; the Eocene 11 of Europe, Numenius from the greatest endemism in Old World but for a mid-Miocene of Europe, Bartramia from the single continent in South America; nearest mid-Pliocene of North America, Calidris from relatives as in Haematopodidac (see above), the early Pliocene of North America, Philo- possibly closest to the Glareolidae (Jehl, 1968). machus from the early Pleistocene of Asia. Four of the five genera of plovers and dot- 11 All of the early Tertiary forms ncc<\ re-evalua- terels occurring in Australia arc cosmopolitan tion to establish the presence of living genera during in distribution (except Vanellus, which does the Paleogene. RICH 108 PAT VICKERS

movement (north or south) along All of the genera occurring in Australia are Cenozoic Indomalaysian route or oceanic dispersal. only boreal winter visitors, most not breeding the of Mayr and in Australia, except for Scolopax. Six of the Phalaropodidae, (Phalaropinae 1967). Phalaropes; eleven genera have cosmopolitan (or nearly so) Amadon, 1951; Brodkorb, record; family cosmopolitan with distributions, two arc primarily New World no fossil breeding ranges exclusively in Holarctica, genera that occasionally wander to Australia, winter ranges in southern hemisphere; Australasia and the Old boreal three are restricted to probably one 1 living genus; closest relatives World. of following: Palaeotringidae, Rostratulidae, Indomalaysian route is in Quite clearly the Scolopacidae, Charadriidae, Haematopodidae genera following this use at present, with many (see above); Jehl (1968) suggests the Tringini route south during the boreal winter. Some and Prosobonini. however, forms ( Tryngites, Bartramia), species, two of occasionally visit Australia using neither the Phalaropus includes three and a Antarctic nor the Indomalaysian route but which have nearly cosmopolitan ranges spending cross broad oceanic barriers. What the history third that breeds in the New World, Because of the wide ranging Scolopacidae has been is its non-breeding season in Australia. single genus that uncertain, as is its place of origin. What can the family is represented by a anywhere be deduced from the above data is that many is neither very diverse nor endemic rather of the genera presently in Australia have in the world, dispersal has probably been in probably made use of, and are still using, the recent. In the case of P. tricolor that breeds Indomalaysian route. North America, neither Antarctic nor Indo- routes have been utilized, but in the Recurvirostridae. (Recurvirostrinae of Mayr malaysian along stilts; Eocene- case of the other two species, movement and Amadon, 1 95 1 ) . Avocets and prob- Recent; no fossil record in Australia; family the Indomalaysian route seems the most cosmopolitan; 3 living, 2 extinct genera; all able explanation for present distributions. living genera occur in Australia; most diverse Determination of direction of movement is more (Clado- in Australia; one genus to Australia difficult. Because Australia is used only as a related family prob- rhynchus); most closely boreal winter refuge by birds in this family, Phalaropodidae (see ably one of the following: some workers have suggested that the group below), Jacanidae, Haematopodidae (Jehl, has originated elsewhere and expanded into 1968), Charadriidae, Aphrizidae, Scolopacidae, Australia. Darlington (1957), however, rightly Rostratulidae, or Palaeotringidae (see above). has raised objection to such interpretations

Australia is the only continent that supports (see above). all three genera (Himantopus, Cladorhynchus, Burhinidae. Thicknees, stone curlews; Miocene- Recurvirostra) of avocets and stilts; Clado- Recent; family cosmopolitan; 1-2 living genera rhynchus is restricted to Australia. Himantopus (including Orthorhamphits in Esacus; some contains only one species (and that is cosmo- (Condon, 1 973 ) consider only one genus, Australia; politan), while Recurvirostra contains four that Burhinus, present) which occurs in genus (Milnea) from the Miocene are each restricted in range, one being endemic one extinct of Europe; greatest endemism and diversity in to Australia. With the fossil record of the group Asia and Australia; closest relatives perhaps being as old as it is (if Presbyornis and Cohonia Jacanidae, Haematopodidae (see above; also are accepted as members of the family (Brod- Jehl (1968)), Glareolidae (see below), Droma- history begins at least by the korb, 1967)), its didae (coast and islands of north and west side early Eocene). Combined with the above data some other family within the Charadriifcrmes, on endemism and diversity of the group, an of Indian Ocean, no fossil record) or perhaps Antarctic route cannot be discounted for initial entrance into Australia. Similarly, a south- The burhinids are most endemic and diverse wards dispersal along the Indomalaysian route in the Old World and Australia (Esacus, cannot be discounted. The present cosmopoli- Burhinus), with only one genus (Burhinus) tan distribution undoubtedly is due to late extending into the New World. Some workers ANTARCTIC DISPERSAL ROUTES 109

(Peters, 1934) have suggested that a third ; Oligocene l ~-Recent; probably Miocene-

genus (Orthorhamphus) , endemic to Australia, Recent in Australia; family cosmopolitan; 15 should be recognized. The family is not a living, 6 extinct genera; 8 living genera in relatively recent derivation but has a history Australia; possibly 1 extinct endemic genus in Australia, described by DeVis highest extending at least into the early Miocene of (1905); diversity in New World, although not decidedly Europe and mid-Miocene of Australia. Simi- greater than in Old World; half of genera of larity of forms between the Old World and Larinae cosmopolitan, 2 endemic to Holarctiea, Australia are most probably due to dispersal 1 to Galapagos; more than half of genera of across the Indomalaysian route, but what the Sterninae cosmopolitan, 2 endemic to South direction of movement (north or south) has America, 2 to Australia and/or New Guinea been is ambiguous. If the movement has been and Southwest Pacific; nearest relatives probably north, which dispersal route was employed by Stercorariidae (skuas, oceanic birds breeding in burhinid ancestors? Arctic and Antarctic, wintering on all oceans, known as fossils only from Pleistocene of North Glareolidae. Pratincoles or swallow plovers, America) (Hudson et al., 1969); the highly courses; no fossil record; family restricted to specialized Rhynchopidae (skimmers, North the Old World-Australia-New Guinea-South- America;, South America, Africa, and Asia, no west Pacific; 5 living genera; 2 living genera fossil record), or perhaps birds within Brod- in Australia (both in the subfamily Glareo- korb's Scolopacidae (Palaeotringinac, Rostra- linae, the pratincoles); most diverse in Africa; tulinae, Scolopacinae, Aphrizinae, Charadriinae, one endemic genus each in Africa and Phalaropodinae, Haematopodinae), Recurviros- Australia while remaining genera range over tridae, Jacanidae, Burhinidae, Dromadidae, Old World or Old World-Australasia (excluding Glareolidae, Thinocoridae, Chionididae, or any N.Z.); nearest relatives probably among the other members of the Charadriiformes. following: Charadriidae (Jehl, 196S), Burhini- dae, Dromadidae (see above), Thinocoridae The terns are more diverse generically in (temperate South America, no fossil record), Australia (as well as the rest of the world) Chionididae (southern South America and than are the gulls. Only one genus of larine, Antarctic Islands, possibly Pleistocene record Larus, with three species, is known in Australia, in southeastern Australia, J. van Tets, pers. a very low diversity when compared with that comm., 1972), Rostratulidae, Haematopodidae, Scolopacidae, Recurvirostridae, Phalaropidae of the group elsewhere in the world. In con- (see above and below). trast, terns (Sterninae) are quite diverse, with seven of the nine genera in the subfamily repre- pratincoles are represented The by two sented in the Australian avifauna. Five of the genera in Australia, one endemic {Stiltia) and seven genera are nearly, if not completely, cos- another that is widely distributed throughout mopolitan. The remaining two (Procelsterna, diverse the Old World (Glareola), though most Gygis) are endemic to Australia and/or New in Africa. The one species of Glareola (G. Guinea and some of the Southwest Pacific maldivarum) that occurs in Australia is only islands. related to the a non-breeding visitor. Closely Gulls are first recognized in the mid-Oligo- pratincoles are the Cursoriinae, considered by cene of Europe, and Larus had differentiated most workers to be the more primitive sub- by the early Miocene in Europe and North are entirely family in the Glareolidae, that America. Since the subfamily is so little differ- restricted to the Old World and particularly entiated and not diverse in Australia even today as whole is diverse in Africa. The family a but is diverse outside of Australia, present decidedly more diverse and endemic in the members of this group most probably arrived Old World than in Australia, and this accom- in Australia via southward dispersal along the panied by the phyletic conclusions favours Indomalaysian route relatively recently. The dispersal of the glareolids southward along the 12 Indomalaysian route. HaJcyornis, from the Eocene London Clay fauna, once thought to be the oldest Laridae, has recently Laridae. (Sterninae and Larinae of Mayr and been transferred to the (Harrison and Amadon, 1951; Brodkorb, 1967). Gulls and Walker, 1972). .

no PAT VICKERS RICH subfamily's occurrence in the Paleogene of the group, it is impossible to determine which Europe does not argue against this conclusion. case is more probable. In addition to the living The Sterninae are first known in Miocene forms, an extinct genus has been reported from sediments of Europe and occur in the mid- the middle Eocene of Germany (which needs Miocene of Australia (Stirton, Tedford, and re-examination to ascertain its identity). If Woodburne, 1968). They are most diverse in correctly identified, it has significance in signal- Australia and the Southwest Pacific today, but ling the differentiation of the family at an early most genera are cosmopolitan. Based on the date, for nothing is known of the group's subfamily's endemism, diversity, and distribu- world-wide distribution at that time. Thus, tion no clear decision concerning their area of distributional history of the rostratulids is un- origin can be made. decipherable at present; although the Indo- Thus, at present, southward movement along malaysian route has probably been used most the Indomalaysian route most probably ac- recently, direction of that dispersal is quite counts for the presence of the known Larinae, uncertain. but area of origin and route employed by the Columbidae. Doves and pigeons; Miocene- Sterninae is uncertain. The most recent episode Recent; Pleistocene-Recent in Australia; family of dispersal of the terns has undoubtedly been cosmopolitan; 58 living, 3 extinct genera; 15-27 Indomalaysian or oceanic, but in what direction living and 1 extinct genera in Australia-New (north or south) and what route was utilized Guinea (15 living genera endemic to Aust- N.G.-SW Pacific plus possibly endemic fossil by the ancestors of the terns, Antarctic, Indo- genus described by DeVis that needs re-evalua- malaysian, or oceanic? tion); most diverse and endemic in Australia Rostratulidae. (Rostratulinae of Mayr and and the Southwest Pacific; nearest relatives the Amadon, 1951; Brodkorb, 1967). Painted snipe; extinct and solitaires (Raphidae) of the Eocene-Recent; no fossil record in Australia; Mascarene Islands in the Indian Ocean and family cosmopolitan except for New Zealand possibly the sand grouse (Pteroclidae) at pre- and North America; 2 living, 1 extinct genera; sent occurring in much of the Old World (first only 1 genus (Rostratula) in Australia; one recognized in the late Eocene-early Oligocene genus endemic to South America, the other to of Europe; Storer (1971) following Maclean the Old World and Australia; nearest relatives (1967) has placed the sand grouse in the probably one of the following: Palaeotringidae, Charadriiformes; Sibley (1966), suggests rela- Scolopacidae, Aphrizidae, Charadriidae, Phala- tionship to both groups); relationships of the ropodidae, Haematopodidae, or Jacanidae (see Columbiformes uncertain, perhaps showing above) closest affinity with the Psittacidae, more distant to the Charadriiformes or . At present a single species of painted snipe, Rostratula bengalensis, occurs in most of the At present the pigeons form a group that is Old World (excluding Europe) as well as in extremely diverse and highly endemic in the

Australia and New Zealand. Although it has Southwest Pacific and Australia, and only probably recently dispersed across the Indo- moderately so in South America, Asia, and malaysian route, the direction of dispersal of Africa. All four subfamilies (Treroninae, Col- this family or of its ancestors is not obvious. umbinae, Goiirinae, Didunculinae or Ptilino- The only other living rostratulid is the South pinae, Columbinae, Macropygiinae, Turturinae Amerfcan Nycticryphes. The Recent disjunct of Condon (1973) are represented in the distribution of the family suggests that pre- Southwest Pacific and/or Australia, but only viously it may have been much more wide- two (Treroninae, Columbinae) are represented spread, or that by chance one population elsewhere. The fossil record of the group is so

accidentally established itself in either South incomplete as to be useless (except that it America or the Old World, while the main indicates the group had differentiated at least population remained in the homeland that by the Miocene and occurred in Holarctica at spawned the founder. Because phyletic relation- that time), as is the understanding of what ships have not been adequately determined for avian group is most closely related to the ANTARCTIC DISPERSAL i s ROU I 111

pigeons. However, due to such great diversity be given the possibility of an origin and/or and endemism at both the generic and the early radiation on the southern continents specific levels in Australia when compared to and an Antarctic dispersal of the group or the pigeon faunas of the rest of the world, an perhaps even an origin within Australia. Un- origin of the group and/or a long history on doubtedly the Indomalaysian route has most that southern continent should not be dis- recently been used in assuring similarity of the counted, nor should the possibility of an African and Asian forms with those of the Antarctic dispersal of the group originally. Southwest Pacific and Australia. Recent movement along the Indomalaysian Cucutidae. Cuckoos; Eocene-Recent; no fossil route accounts for generic similarity between record in Australia; family cosmopolitan, 37 Old World and Australian forms, but such living and 2 extinct fossil genera; 9-10 living similarity in no way demands a northern origin genera in Australia; one (Microdynamis) of for the group—it demands only interchange which is endemic and 3 of which are endemic between the two land masses. There may, in to Australia -New Guinea - Southwest Pacific fact, have been movement both north and south {Seythropx, Rhmipho mantis, Calieehthrus)\ greatest diversity of along this route when individual genera are family in Southwest Pacific and Asia considered. with twice as many endemic genera in the New World as the Old; most closely related Psit taeniae. Parrots; Miocene-Recent; Pleisto- family probably the (Musophagidac) cene-Recent in Australia; family with pan- now restricted to Africa but with a Tertiary tropical distribution at present. Tertiary repre- record (Eocene-Miocene) in Europe. sentatives in Europe; 81 living, 4 extinct genera; 37 living genera in Australia of which Cuculids are represented by 9-10 living 37 are endemic to Australia-New Guinea-South- general in Australia, only one of which is west Pacific; subfamilies Loriinac, Cacatuinac, endemic. This is in marked contrast to the Micropsittinae endemic to Australia-New great diversity and high endemism of the family Guinea-Southwest Pacific; besides 2 endemic in the Southwest Pacific and Asia. As the subfamilies in New Zealand, only subfamily nearest relatives of the cuculids are probably occurring outside of Australia (also occurs in the musophagids (turacos), living in Africa Australia) is the Psittacinae; Condon (1973) has recently proposed the designations Cac- today (and had a history in early to mid- tuidae (Proboscigerinae, Calyptorhynchinae, Tertiary Europe), this group probably dispersed Cacatuinae), Psittacidae (Palacornithinac), into Australia southwards along the Indo- Loriidac, Apopsittidae, Polytelidae, Platycerci- malaysian route. Endemism among the cuculids dae (Pczoporinae, Lathaminae, Platycercinae) is highest in the New World and the group had for Australian parrots; greatest diversity and differentiated there by the mid-Eocene; so, an endemism in Australia-New Guinea-Southwest Antarctic dispersal route cannot be completely Pacific while South America has moderate ruled out but seems less likely based on known diversity; nearest relatives are uncertain but evidence owing to the group's low endemism in may be among the Columbiformcs (pigeons, Australia. Because faunal interchange sand grouse) or the Cuculiformes (turacos, between cuckoos, and allies). North America and Europe was high as late as the early Eocene (McKcnna, 1972a), cucu-

Although the relationships of this group to lids might well be expected in Europe if present others is not sufficiently well understood, the in North America; thus histories in both the great diversity and endemism of the group in Old and the New World have probably been Australia, New Guinea, and the Southwest long, and could account for such great diversity Pacific, in comparison to the strikingly low and endemism in these areas. Cuculids may be diversity of that group in Asia and Africa one group, that although present in South (with only two and four genera respectively), America at a time when the Antarctic dispersal does not obviously suggest a northern origin route was open, did not have the right sweep- for the group. Instead, I believe, it strongly stakes ticket! suggests that more serious consideration should Marchant's ( 1 972 ) suggestion that the 112 PAT VICKERS RICH

genera Cacomantis and Chrysococcyx were the Indomalaysian route. Otherwise, greater derived from a common ancestor in Antartica endemism and diversity would certainly be and then spread north into Australia and Africa expected on a continent with such a small respectively, does not take into account the number of nocturnal avian carnivores. probable prc-carly Cretaceous (ca. 200 m.y. Tytonidae (including the Phodilidae, which B.P.) or early Cretaceous separation of Africa Brodkorb (1971a) isolates as a separate family; and East Antarctica (see figs. 1-2) and in no Tytoninae of Mayr and Amadon, 1951). Barn way invalidates the above discussion. This ; Miocene-Recent; Pleistocene-Recent in timing of the Afro-Antarctic break-up is prob- Australia; family cosmopolitan; 2 living genera; ably too far in the past to have permitted 1 genus (Tyto) in Australia; Tyto cosmo- dispersal between Australia and Africa of the politan, but other genus, Phodilus, endemic to Asia and the Southwest Pacific; modern avian families, let alone closely related greatest diver- sity and endemism of family in Asia and the genera. Certainly the warm tropical vegeta- Southwest Pacific; closest relatives probably tional belt that lined much of southern Asia the Strigidae or the Protostrigidae (see above). and the Mediterranca area during the early and into the mid-Tertiary would have provided an Because the greatest diversity (both generic excellent corridor for distribution between and specific) of this group occurs in the Old Australia and Africa, once Australia had moved World and Southwest Pacific (only Tyto far enough north to allow such exchange to reaches Australia and Tyto alba reaches the take place (by the Miocene). Isolation imposed New World), southward movement through when such a corridor was gradually destroyed Indomalaysia seems the most likely route be- during the Neogene by increasingly cooler tween Australia and the rest of the world. world-wide climate, could easily explain the Phodilus in addition to Tyto occurs in the Old Cacomantis-Chrysococcyx distribution seen World; another extinct genus in the subfamily today. Phodilinae occurred in the Miocene of Europe. Strigidae. (Striginae of Mayr and Amadon, Podargidae. Frogmouths; no fossil record; re- 1951). Owls; Eocene-Recent; no fossil record stricted to Asia-Southwest Pacific-Australia- m Australia; family cosmopolitan, 28 living and New Guinea; 2 living genera, Podargus (Aust.- 3 extinct fossil genera; 2 genera in Australia; N.G.-SW Pac.) and Batrachostomus (As.-SW many genera cosmopolitan with endemism Pac.); nearest relatives within the Caprimulgi- nearly the same in both New and Old Worlds; forrnes including Caprimulgidae, Aegothelidae group most diverse in South America, South- (see below), Aegialornithidae (Eocene-Oligo- west Pacific, and Asia; nearest relatives prob- cene of Europe), Nyctibiidae (restricted to ably the Tytonidae (barn owls; Ford, 1967 American tropics with a Pleistocene record in et aL) also cosmopolitan in distribution (known South America), or the Steatornithidae (re- as fossils from the Miocene of Europe, else- stricted to northern South America, no fossil where only Pleistocene or later), and the Pro- record). tostrigidae (Eocene, North America) and more Aegothelidae. Owlet frogmouths; Pleistocene- distantly the Caprimulgiforrnes (see below), Recent; Miocene-Recent in Australia; at least and perhaps the Falconiformes. 1 genus (Aegotheles); presently restricted to the Southwest Pacific-New Guinea-Australia Only two genera of typical owls occur in the but has Pleistocene record in New Zealand; living Australian avifauna, while the group is most diverse specifically in New Guinea; nearest quite diverse elsewhere, particularly in the relatives in the Caprimulgiforrnes (see above, Southwest Pacific, Asia, and the New World. including Podargidae). Apparently Ninox (with four Australian-New Caprimttlgidae. Nightjars; Pleistocene-Recent; no fossil record in Australia; family Guinean species) and Uroglaux (1 species in cosmopoli- tan; 19 living genera; 2 genera New Guinea; genus restricted to New Guinea { Eurostopodas and Caprimulgus; Condon (1973) recognized and the Southwest Pacific) or forms that gave only the latter) in Australia-New Guinea; rise to them, dispersed southwards from Asia greatest diversity in the New World, particu- and the Pacific Islands to Australia along larly in South America; largest numbers of ANTARCTIC DISPERSAL ROUTES 113

endemics in the New World, particularly in it is impossible to determine their geographic South America; closest relatives within the range; primitive caprimulgiforms might well Caprimulgiformes (see above). have been more widespread, and if such were The Caprimulgiformes, including the three the case, the Antarctic route should not be previously mentioned families ( Podargidae, dismissed. Aegothelidae, and Caprimulgidae), are first Hemiprocnidae. Crested swifts; Pleistocene- represented in the Eocene/Oligocene of Europe Recent; no fossil record in Australia; 1 genus; by the Aegialornithidae. An aegothelid has family restricted to New Guinea-Southwest been recovered from Miocene sediments in Pacific-Asia; most diverse and endemic in Australia, but the only other record of the the Southwest Pacific; closest relatives prob- ably the swifts order is in the Pleistocene from several different (Apodidae; but Sibley, 1960, thought this due to convergence localities in many parts of the world. The order but later (Sibley and Ahlquist, 1972) noted close simi- is composed of two families (Podargidae, larity) that have a cosmopolitan distribution Aegothelidae) endemic to Australia-New (see below). Guinea-Southwest Pacific (and Asia in the case The crested of the Podargidae), two endemic to tropical swifts form a small group (one genus with South America (Nyctibiidae, Steatornithidae), three species) that is most diverse in the Southwest and one with a cosmopolitan distribution Pacific. It is probably most (Caprimulgidae). closely related to the swifts (Apodidae) that occur throughout In the case of the cosmopolitan Caprimul- the world. Where the group originated is uncertain, gidae, entrance into Australia appears to have but its dispersal between Australia and the been via southward movement along the Indo- rest of the world seems intimately tied with malaysian route. Only two genera, one cosmo- the Indomalaysian region. At present politan (Caprimulgus) and another shared with either a northward or southward dispersal could have occurred, depending on the Old World (Eurostopodus) , occur in Aus- area of origin. tralia. The family is much more diverse with more endemic species in the Old World than Apodidae. Swifts; Eocene or Oligocene-Recent; no fossil record in Australia, but most diverse and endemic in Australia; family cosmo- politan; 16 living, 1 extinct genera; 3-5 in the New World. Such data would seem best living genera in Australia; diversity is low in Europe explained by a southward dispersal along the and New Zealand but about the same for all Indomalaysian route. other continents; closest relatives, the Hemi- Dispersal routes utilized by the other two procnidae (see above), perhaps the Trochilidae families or their ancestors are uncertain, how- (, with a New World, primarily ever. They are either entirely (Aeogthelidae), South American, distribution and a Pleistocene or nearly (Podargidae), restricted to and most record there; Sibley, 1960; Cohn, 1966, how- diverse in Australia-New Guinea-Southwest ever, has suggested the two groups convergent); and perhaps Pacific including New Zealand. Only the Podar- more distantly the Caprimulgi- formes (see above). gidae range into Asia. What relationships they have to the Caprimulgidae and the American All of the genera of swifts that occur in tropical forms (Nyctibiidae, Steatornithidae) Australia-New Guinea are monotypic or con- are uncertain, and so too is the route into tain only a few species (Hirundapus, Apus,

Australia used by both groups or their ances- Collocalia, Chaetura, Meamsio); only the first tors. Did they develop in Australia and/or the three are recognized by Condon (1973) and Southwest Pacific and spread north along the are more diverse in the Southwest Pacific Indomalaysian route? If so, did the ancestral and the Old World. No endemic genera occur stock arrive originally via the Antarctic or the in Australia. The fossil record indicates that Indomalaysian route? Although caprimulgi- the family was present in the Old World by the forms are known only from the Eocene/Oligo- Eocene or Oligocene, and both Apus and Col- cene of Europe, because of generally poor localia differentiated there by early Miocene. records in the rest of the world at this time, The recent distributional and diversity data 114 PAT VJCKERS RICH suggest that southward movement over the Bucerotidae. Hornbills; Eocene-Recent; no fossil record Australia; family occurs in Old World- lndomalaysian route into Australia is most in Pacific-New 12 living, 3 probable. Southwest Guinea; extinct genera; 1 living genus (Aceros), reaches

( Brodkorb, Alcedinidae. Halcyonidae of New Guinea, greatest endemism in Southwest 1971a). Kingfishers; Hocene/Oligocene-Rccent; Pacific; nearest relatives probably among the no fossil record in Australia; family cosmo- following: Coraciidae, Upupidae, Alcedinidae, politan; 14 living, 1 extinct genera; 7 living Motmotidae, Todidae, or Meropidae (see genera in the Australian-New Guinean region, above). one of which is endemic to New Guinea (ClyWceyx) and 3 of which are endemic to The three families mentioned above (the Australia-New Guinea-Southwest Pacific (Meli- bee eaters, rollers, and hornbills) have in dora, DacelOi Tanysiptera); greatest diversity common a great diversity only in the Old in Australia - New Guinea - Southwest Pacific World, in contrast to a single genus repre- region; similar endemism in Australia-South- sentation in Australia and/or New Guinea. west Pacific and Old World regions; nearest In addition the coraciids and bucerotids have relatives among the following: Todidae (todies a long history (Eocene/Oligocene-Recent) in of the Greater Antilles, Pleistocene record the Old World. Thus, it seems most likely that there), Moimotidae (motmots, restricted to the southward movement along the lndomalaysian American tropics with a Pleistocene record there), and Meropidae (see below). route accounts for their presence in the Aus- tralia-New Guinea avifauna.

The kingfishers, first known in the Eocene In summary, there are several avian families or early Oligoccne of Europe and later there (or their ancestors) in the Australian-New in the Miocene, probably dispersed along the Guinean fauna whose initial dispersal to or lndomalaysian route. Diversity of the group in away from that continent may well have been the New World is so low (two genera in a via either an Antarctic or an lndomalaysian single subfamily Ccrylinae) as to suggest a sweepstakes route, or in some cases via oceanic rather recent arrival there, which thus argues dispersal. These families include (see Table against an Antarctic dispersal unless diversity 4): in the past is revealed in the fossil record of Casuariidae the group when better known. Dromornithidae Phalacrocoracidae Meropidae. Bee caters; Pleistocene-Recent; no fossil record in Australia; distrihuted over much Phocnicopteridae of Old World, Southwest Pacific-Australia; 7 Anatidac (in part: Anseranatidae, Cereop-

living genera; 1 genus {Merops) in Australia; sini, Dendrocygnini, Oxyurini) greatest diversity and endemism in Africa; Accipitridac (in part: Milvinae, Perninae) closest relatives among the Alcedinidac, Todi- Megapodiidae dae, Motmotidae (see ahovc), and Upupidae Pedionomidae (hoopoes, restricted to warm parts of Old Turnicidae World, Pleistocene record in Old World) and Hacmatopodidac Coraciidae and Bucerotidae (restricted to Old Rccurvirostridae World and Australasia, with greatest diversity Burhinidae in Old World and Hocene-Oligocene record in Europe), Laridae (in part: Sterninae) Coraciidae. Rollers; Hocene/Oligocene-Rccent; Rostratulidae no fossil record in Australia; family restricted Columbidae to Old World and Australia-New Guinea-New Psittacidae

Zealand; 5 living. I extinct genera; 1 genus Podargidae {Euryst&mus) in Australia; greatest diversity Acgothelidae. and endemism in Africa (including Madagas- car); nearest relatives probably Bucerotidae The remaining families not listed above have (see below), Upupidae, Alcedinidae, Motmo- most probably moved south into Australia tidae, Todidae, or Meropidae (sec above). across the lndomalaysian route. :

ANTARCTIC DISPERSAL ROUTES 115

TABLE 4 a much longer interchange of birds between Australia and the rest of the world. Compilation of dispersal routes probably used between Australia and the remaining world by Discussion and Conclusions those non-passeriform families comprising the Since Mayr's (1944a) classic paper suggest- Australian avifauna. ing that the major part of the Australian non- marine avifauna had been derived from the INDOMALAYSIAN ROUTE (southward move- north, ideas concerning continental stability ment) : Podicipedidae, Anhingidae, Ardeidae, Ciconiidae, Threskiornithidae, Anatidae (in part: have radically changed. In the 1940's and as Cygnini, Tadornini, Anatini, Aythiini), Accipi- late as the 1950's, prevailing opinion among tridae (in part: Elaninae, Circinae, Accipitrinae), Pandionidae (?N., ?S.), Falconidae, Phasianidae, geologists, as well as biogeographers, demanded Gruidae, Rallidae, Otididae, Jacanidae, Chara- that continents had maintained their present driidae, Arenariidae, Phalaropodidae, Scolopaci- positions with respect to one another through- dae, Glareolidae, Laridae (in part: Larinae), Cuculidae, Strigidae, Tytonidae, Caprimulgidae, out the Phanerozoic. Only with the pioneering Hemiprocnidae (?N., ?S.), Apodidae, Alcedini- work of Vine and Matthews (1963) on ocean dae, Meropidae, Coraciidae, Bucerotidae. floor magnetism did consensus begin shifting

ROUTE UNCERTAIN (Antarctic, Indomalaysian, in favour of the currently popular theories and in some cases Oceanic dispersal possible) encompassing continental drift, ocean floor Casuariidae, Dromornithidae, Phalacrocoracidae, spreading, and plate tectonics. Seen in the light Pelecanidae, Phoenicopteridae, Anatidae (in part: Anseranatidae, Cereopsini, Dendrocygnini, of new and convincing evidence, Australia Oxyurini), Accipitridae (in part: Milvinae, (including New Guinea) is no longer thought Perninae), Megapodiidae, Pedionomidae, Turni- to have remained forever bounded by 45° South cidae, Haematopodidae, Recurvirostridae, Bur- hinidae, Laridae (in part: Sterninae), Rostra- latitude and the equator, but instead is viewed tnlidae, Columbidae, Psittacidae, Podargidae, as a truly incredible Tertiary voyager (a Noah's Aegothelidae. Ark, McKenna, 1973), moving north from a connection with the Antarctic some 50-55 m.y.

For many non-passeriform families ( 19 of the ago to its present position, a journey of some 46 families, five of these only in part) of the 30-50 degrees of latitude. During its northward obligate terrestrial birds in the living, as well trek, profound changes in climatic regimes as the Tertiary avifauna of Australia, at least affecting Australia brought about the switch two hypotheses are available regarding route of from a once cool, temperate vegetation cover- original entrance. In many cases there is not ing much of the continent to the present varied enough information to permit a choice of one and more zoned flora with the restricted cool route in preference to another, and a most temperate assemblages in the south, the tropical important point is that such a choice cannot vegetation in the north (probably a Miocene in- be made at present, but does exist. Many of novation), and the widespread arid assemblages the non-passeriform families have utilized the that characterize much of Australia today. Indomalaysian route most recently (Miocene Undoubtedly synchronous changes occurred in and more recently). The mixing of the Aus- those faunas inhabiting the island continent, tralian and Old World avifaunas much more including the birds. Unfortunately, only die thoroughly than the mammalian faunas of these closing phases of Australia's avifaunal history two areas has increased the difficulty of dis- are known, and thus such change cannot pre- tinguishing Australian and Asian elements. sently be documented for birds as it has been This late Cenozoic utilization of the Indo- for the floras. What the late- Mesozoic and early malaysian route should not, however, be used Tertiary terrestrial faunas were like is yet a to suggest a northern origin for all or most mystery. of the Australian non-passeriform avifauna. From a review (see first section of paper) The striking similarity of the Australian-New of late Mesozoic and Cenozoic geologic history Guinean fauna to that of Asia and the South- of Australia, other southern continents, and west Pacific tells only of the latest episode in the ocean basins now separating them, it is 116 PAT VICKERS RICH apparent that during the last 80 to 90 million If the modern families of birds, like the years, two possible land routes (both probably majority of modern placental mammalian sweepstakes routes in Simpson's (1940, 1953) families, had first developed and radiated and McKenna's (1972a-b, 1973) terminology) mainly during the late Paleogenc or Neogene between Australia and the rest of the world in Holarctica, probably no choice of routes have been available to terrestrial vertebrates: between Australia and the rest of the world the Antarctic and the Indomalaysian. The used by different groups of birds would Antarctic route was the only path not requiring be necessary; the Indomalaysian route would dispersal across broad ocean basins that lay be the only major stepping stone to Australia between Australia and the rest of the world as suggested by Mayr (1944a) for the birds from the late Mesozoic until the Eocene. and by Matthew (1915) for vertebrate faunas, Between that time and sometime during the primarily mammals. Unlike modern placental Miocene, broad water barriers on the north and mammalian families, however, a large number an increasingly hostile climate accompanied by of the living avian families apparently differ- an ever widening water barrier on the south entiated during the Paleogene (see figure 10). essentially isolated Australia from the world's Twenty of the forty-six avian families that remaining land masses. During this period of make up the Cenozoic avifauna of Australia, profound isolation (that lasted perhaps as long had developed by the Eocene, some possibly as 20-30 million years), the avifauna, as well as early as the Paleocene (Phalacrocoracidae), as the rest of the non-marine fauna, developed a few more by the Oligocene (Phasianidae, without much interference from the outside, Charadriidae, Laridae), and most of the re- only plagued, or advantaged, by the profound maining families at least by the Miocene climatic changes that occurred as Australia (excepting Megapodiidae, Turnicidae, Pedio- drifted northward. In the mid to late Tertiary, nomidae, Arenariidae, Phalaropodidae, Glareo- as Australia neared its present position, the lidae, Podargidae, Aegothelidae, Caprimulgidae, Indomalaysian route became the only major Hemiprocnidae, and Meropidae that have only dispersal route that could be utilized by non- a Pleistocene or Recent record). Differentiation marine vertebrates. The Antarctic route was undoubtedly occurred sometime before the inactivated by the ever increasing breadth of the groups first appeared in the fossil record, and water barrier and the expansion of glaciation thus probably more than just the twenty families to a present continent-wide scale. with lengthy records also had records extending Paleoclimatological evidence suggests that well back into the Paleogene. So, when con- the two dispersal routes had decidedly different sidering the origin of the Australian avifauna, climates throughout their entire existence. both Indomalaysia and Antarctica should be During the late Mesozoic and early Tertiary considered as possible routes of dispersal (see the Antarctic route was covered in many places figure 9). with sizeable forests composed of Nothofagus, In summary, then, because of the early Araucaria, other primitive angiosperms, gym- differentiation of modern bird families and the nosperms, ferns, and scouring rushes, among availability of two dispersal routes to and from

others, that is, a cool temperate flora much Australia, one cannot assume that most, if not like that covering Australia during the early all, of the Australian non-marine avifauna Tertiary. The climate was never tropical but originated in the north. Most probable route was in no way the severe one affecting Ant- of initial entrance of ancestors of the following arctica today. The Indomalaysian route, on the families or the families themselves is impossible other hand, has apparently supported a tropical to determine at present: Casuariidae (including

vegetation throughout the Cenozoic, and seem- Dromaius and Casuarius) , Dromornithidae,

ingly its climate has remained much as it is Phalacrocoracidae, Pelecanidae, Phoenicopteri- today during the entire time it has served as a dae, Anatidae (in part: Anseranatidae, Cere- major route between Australia and the rest of opsini, Dendrocygnini, Oxyurini), Accipitridae the world. (in part: Milvinae, Perninae), Megapodiidae, ANTARCTIC DISPERSAL ROUTES 117

Pedionomidae, Turnicidae, 13 Haematopodidae, Antarctica. It was this 'changed' fauna that Scolopacidae (in part), Recurvirostridae, Bur- probably began dispersing northward across hinidae, Laridae (in part: Sterninae), Rostra- the Indomalaysian route as Asian forms tulidae, Columbidae, Psittacidae, Podargidae, spread south when the Australian 'ferryboat' and Aegothelidae. Determination of the most (McKenna, 1973) neared its Asian 'dock'. probable route of initial dispersal for such In conclusion, then, I would reopen con- groups will have to await further studies on sideration of the origins of the non-passeriform avian phylogeny and a more complete fossil avifauna of Australia and call for a re-evalua- record from the Australian late Mesozoic and tion of over 40% of that fauna as data in- Paleogene. A point to be stressed here, how- creases, particularly that regarding phylogeny ever, is that such Australian birds should be of avian groups and the fossil record in Aus- recognized as groups of uncertain origin and tralia. Then, and only then, can probabilities be should not be lumped with those forms that estimated that favour either an Indomalaysian most probably entered Australia from either or an Antarctican route for those groups in the north or the south. At the species level question. approximately 43% ($s well as 19 of the 46 Acknowledgements families present in Cenozoic of Australia) of Many people are to be thanked for thought- the non-passeriform, non-marine living avi- provoking discussions that enhanced this paper, fall in the category 'groups in particular W. Bock, M. McKenna, and R. of unknown origin', hardly a small segment of Tedford, as well as D. Bohaska, J. Cracraft, the Australian bird fauna! The remaining I. Dalziel, T. Darragh, D. Dorwood, D. Elliot, groups, on the other hand, most probably E. Gaffney, W. Hamilton, G. and J. Hope, E. arrived via the Indomalaysian dispersal route Kemp, A. McEvey, B. Nielsen, S. Olson, M. or in a few cases as long distance travellers Plane, P. Raven, T. Rich, B. Schaeffer, R. across the Pacific or Indian oceans (Charadrii- Schodde, J. van Tets, and J. Warren. R. Gooris formes in particular). rendered figures 7, 8 and 10 and H. Galiano Certainly arguments using the close similarity figures 9A-B. Financial support for this study of the Southeast Asian and Australian avifaunas was provided by an American Association of as an indicator that only the Indomalaysian University Women Fellowship, an N.D.E.A. route has served birds dispersing between Aus- Title IV Fellowship, and an Australian-Ameri- tralia and the remaining world must be ques- can Educational Foundation Grant for 1973- tioned. Such a similarity reflects the recent, 1974. Research facilities for the writing of this close apposition of the Palearctic and Australian paper were kindly provided by The American continental masses and is certainly indicative Museum of Natural History, New York, The of interchange, but movement of faunas could National Museum of Victoria, Melbourne, and have been northward as well as southward. The Bureau of Mineral Resources, Canberra. Such close similarity does not rule out the i: possibility of an Antarctic dispersal at some- *Such severe climatic changes taking place in the Australian Tertiary may serve as yet another factor, time in the past. Such southern dispersal would in addition to those mentioned by Keast (1972), that have occurred at a time much more distant in accounts for the differences in the diversity and endemism characteristic of the South American, the past than that across Indomalaysia. The African, and Australian avifaunas. Both the South fauna that arrived in Australia via an Antarctic American and the African land masses have remained route would certainly have evolved during the at nearly the same latitudinal positions throughout the Tertiary with concomittant stability of climatic 40 million years since their arrival there, especi- regimes affecting those continents. Australia, on the ally as they were probably in extreme isolation other hand, has quite noticeably moved some 30-50° for 20 million years. Owing to the climatic of latitude northwards. Perhaps such changes, and possibly accompanying , account in part taking Australia drifted north- changes place as for the lower endemism in Australia when com- ward, undoubtedly there were concomitant pared to that characterizing South America as well as such factors as smaller land area, less changes, perhaps even large-scale extinctions, topographic diversity, decidedly smaller areas of tropical environ- in the faunas that had reached Australia across ments, etc. suggested by Keast. 118 PAT VICKERS RICH

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