Origins and evolution of the Antarctic biota

322 Pages·1989·58.89 MB·English

by  J. Alistair Crame

Origins and Evolution of the Antarctic Biota
Geological Society Special Publications
Series Editor K. COE
GEOLOGICAL SOCIETY SPECIAL PUBLICATION NO 47
ORIGINS AND EVOLUTION OF THE ANTARCTIC BIOTA
EDITED BY
J. A. CRAME
British Antartic Survey
Natural Environment Research Council
Cambridge
1989
Published by
The Geological Society
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British Library Cataloguing in Publication Data
Origins and the evolution of the Antarctic
1. Antarctic. Fossils
I. Crame, J. A. (James Alistair) 1949-
560.998'9
ISBN 0-903317-44-3
Printed in Great Britain at the Alden Press, Oxford
Contents
J. AL1STAIR CRAME, Origins and evolution of the Antarctic biota: an introduction 1
L. R. M. COCKS, Antarctica's place within Cambrian to Devonian Gondwana 9
FRANCOISE DEBRENNE & PETER D. KRUSE, Cambrian Antarctic archaeocyaths 15
GERALD F. WEBERS & ELLIS L. YOCtlELSON, Late Cambrian molluscan faunas and the origin
of the Cephalopoda 29
G. C. YOUNG, The Aztec fish fauna (Devonian) of southern Victoria Land: evolutionary and
biogeographic significance 43
SHERR1 L. DEFAUW, Patterns of evolution in the Dicynodontia with special reference to
austral taxa 63
W. G. CHALONER & G. T. CREBER, The phenomenon of forest growth in Antarctica: a review 85
MARY E. DETTMANN, Antarctica: Cretaceous cradle of austral temperate rainforests? 89
ROSEMARY A. ASKIN, Endemism and heterochroneity in the Late Cretaceous (Campanian)
to Paleocene palynofloras of Seymour Island, Antarctica: implications for origins, dispersal
and palaeoclimates of southern floras 107
T. H. RICH, P. V. RICH, B. WAGSTAFF, J. McEWEN-MASON, C. B. DOUTHITr & R. T.
GREGORY, Early Cretaceous biota from the northern side of the Australo-Antarctic rift
valley 121
R. E. MOLNAR, Terrestrial tetrapods in Cretaceous Antarctica 131
G. R. STEVENS, The nature and timing of biotic links between New Zealand and Antarctica
in Mesozoic and early Cenozoic times 141
PETER DOYLE & PHILIP HOWLETT, Antarctic belemnite biogeography and the break-up of
Gondwana 167
RODNEY M. FELDMANN & DALE M. TSHUDY, Evolutionary patterns in macrurous decapod
crustaceans from Cretaceous to early Cenozoic rocks of the James Ross Island region,
Antarctica 183
SANKAR CHATTERJEE & BRYAN J. SMALL, New plesiosaurs from the Upper Cretaceous of
Antarctica 197
JUDD A. CASE, Antarctica: the effect of high latitude heterochroneity on the origin of the
Australian marsupials 217
K. BIRKENMAJER & E. ZASTAWNIAK,L ate Cretaceous-early Tertiary floras of King George
Island, West Antarctica: their stratigraphic distribution and palaeoclimatic significance 227
J. T. EASTMAN (cid:127) L. GRANDE, Evolution of the Antarctic fish fauna with emphasis on the
recent notothenioids 241
ANDREW CLARKE & J. ALISTAIR CRAME, The origin of the Southern Ocean marine fauna 253
R. EWAN FORDYCE, Origins and evolution of Antarctic marine mammals 269
ELLEN THOMAS, Development of Cenozoic deep-sea benthic foraminiferal faunas in Antarctic
waters 283
LES WATLING & MICHAEL H. THURSTON, Antarctica as an evolutionary incubator: evidence
from the cladistic biogeography of the amphipod Family Iphimediidae 297
Origins and evolution of the Antarctic biota: an introduction
J. A. CRAME
British Antarctic Survey, Natural Environment Research Council, High Cross, Madingley
Road, Cambridge CB3 0ET, UK
Within the last 25 years there has been a dra- in this volume L. R. M. Cocks provides a brief
matic increase in our knowledge of the fossil but succinct review of Antarctica's place within
record of Antarctica. Improved access to the Cambrian to Devonian Gondwana. It is perhaps
remotest parts of the continent, the advent of not generally appreciated that the greater part
offshore drilling and intensive study of early of Antarctica's extensive Early and Middle
expedition collections have all led to the ac- Cambrian sedimentary record was laid down
cumulation of a vast amount of data that under tropical conditions. The extensive
stretches back nearly 600 Ma to the beginning of Shackleton Limestone of the Transantarctic
the Cambrian period. No longer can Antarctica Mountains, and its correlatives, contains abun-
be dismissed from our view of the history of life dant shelly faunas with Australian and Chinese
on earth simply because so little is known about affinities, and part of the margin of what we
it; it is fast becoming another crucial reference now know as the Gondwana supercontinent
point for global palaeontological syntheses. formed a reef belt that protruded 10~ ~ into
If, today we have an image of Antarctica as a the Northern Hemisphere. The most abundant
remote, inhospitable continent that supports and characteristic early Palaeozoic fossils of
little life, we now know that such a view cannot Antarctica are in fact archaeocyaths and F.
be projected back indefinitely through time. Debrenne & P. D. Kruse show how, in con-
Abundant plant and animal fossils from a variety junction with certain skeletal cyanobacteria and
of periods point to much more benign climates algae, these primitive coral-like organisms were
and immediately raise a series of interconnected the primary framework-builders of the reefs.
questions: where did such organisms come from, Taxonomic revisions presented by these authors
how long did they persist, and precisely when confirm the particularly strong faunal links be-
(and how) did they become extinct? Can our tween Antarctica and South Australia in the
most southerly continent throw further light on Early Cambrian.
the long-term role of climate in driving evol- Amongst the inarticulate brachiopods, tri-
utionary trends (e.g. Valentine 1967; Vrba lobites, primitive molluscs and other marine
1985)7 invertebrates that inhabited the Early Cambrian
It was with points such as these in mind that a reefs and carbonate platforms, a particularly
mixed group of palaeontologists, biologists, ge- important series of monoplacophorans and gas-
ologists and geophysicists gathered together tropods has been identified by G. F. Webers &
for an international discussion meeting on the E. Yoeheison. These occur in the Late
'Origins and evolution of the Antarctic biota' at Cambrian (Dresbachian) Minaret Formation of
the Geological Society, London on 24 and 25 the Ellsworth Mountains and comprise some 20
May 1988. A further workshop day (26 May taxa. The most important of these is the mono-
1988) was held at the British Antarctic Survey, placophoran, Knightoconus antarcticus, which
Cambridge and altogether over 100 scientists could well have been a direct ancestor of the
from some 15 different countries took part in cephalopods. Its 7cm long, curving, cap-like
the discussions. shell bears multiple internal septa along at least
Obviously, in such a short space of time, it one-third of its length and these are perforated
was not possible to cover all aspects of Antarctic by a rudimentary connecting tube (siphuncle).
palaeontology, and some subjects were necess- The stratigraphic position of this species can be
arily reviewed in greater detail than others. As demonstrated to be slightly beneath that of
Palaeozoic biotas and biogeography were to be Plectronoceras, the first true cephalopod (from
the subject of a complementary symposium China).
('Palaeozoic biogeography and palaeogeogra- In his review, Cocks points out that there is a
phy'; Oxford, 14-19 August 1988), they were very limited Ordovician and Silurian sedimen-
not examined exhaustively here. Nevertheless it tary and palaeontological record in Antarctica;
was felt to be important to spend some time it is likely that this was a time of extensive up-
considering Antarctica's first macrofossils, and lift and erosion. However, it is apparent that,
From Crame, J. A. (ed.), 1989, Origins and Evolution of the Antarctic Biota,
Geological Society Special Publication No. 47, pp. 1-8.
2 J.A. CRAME
during the later part of early Palaeozoic time, phyllolepid placoderms, xenacanth sharks and
profound changes were occurring in the dis- bothriolepid antiarchs.
tribution of supercontinents and the large Collision of the Gondwana and Baltica/
Gondwana plate slid progressively over the con- Laurentia blocks in the latest Devonian to form
temporary south pole. By the early Middle the single supercontinent of Pangea radically
Devonian the pole lay under either S. Africa or alters our view of the subsequent evolution of
S. America, and Antarctica came to occupy a life in the southern high latitudes. It has tra-
mid- to high-latitude position (35~176 de- ditionally been felt that the classic 'Gondwana'
pending on the reconstruction used). It has sequences in these regions, comprising Permo-
essentially maintained this ever since. Carboniferous non-marine glacial facies
Although palynomorphs of probable Early overlain by coal measure facies containing a
Devonian age are known from the Beacon Glossopteris flora and Triassic non-marine facies
Supergroup of southern Victoria Land (Kyle with a Lystrosauras fauna and Dicroidium flora,
1977), the most characteristic fossils of this were the product of supercontinent isolation.
age comprise a series of marine invertebrates However, viewed in terms of the single super-
from the Horlick Mountains, Ohio Range, continent model, it would now appear that
Transantarctic Mountains. These clearly have these classic sedimentary facies and biotas can
Malvinokaffric Province (i.e. cool-water) af- be re-interpreted as austral components of a
finities (Doumani et al. 1965) and testify to Pangean super-sequence (Veevers 1988). The
the comparatively high-latitude position of basal glacial successions reflect the mid- to high
Antarctica in the Early Devonian. Some Lower southern latitudes and succeeding beds contain
Devonian fish fossils have been recorded from a provincial biota separated from boreal Pangea
both the Horlick and Ellsworth mountain ranges on the east by the Tethyan gulf and on the west
and Middle-Late Devonian plant remains are by a tropical land zone (Veevers 1988, fig. 2c).
known from the Ruppert Coast of Marie Byrd Although Veevers dates the coalescence of
Land (Grindley et al. 1980); however, these are Gondwana and Euramerica slightly later (mid-
overshadowed in both stratigraphical and bio- Carboniferous, c. 320 Ma), it is clear from his
geographical importance by the Givetian (late reconstruction that the classic Glossopter& flora
Middle Devonian) Aztec Siltstone fish fauna now has a distinctly austral aspect.
from southern Victoria Land. Dicynodonts, a widespread, morphologically
In a comprehensive review, G. C. Young diverse group of mammal-like reptiles prevalent
shows that the Aztec fauna comprises not only during the Permian and Triassic, are the subject
the oldest but also one of the most diverse fossil of a comprehensive review by S. L. De Fauw.
vertebrate assemblages yet found in Antarctica. Essentially composed of herbivores, this group
Over 30 taxa have so far been identified and appears to have undergone two significant
these belong to four major groups of jawed adaptive radiations: one in the Late Permian
fishes -- the placoderms (armoured fishes), (Tatarian) and the other in the early Middle
acanthodians (spiney-finned fishes), chon- Triassic (Anisian). Both radiations were un-
drichthyans (sharks and their relatives) and doubtedly aided by the development of homo-
osteichthyans (boney fish) -- and one group of geneous Gondwana floras (the Glossopteris and
agnathans (primitive jawless vertebrates; the Dicroidium floras, respectively), but it is import-
thelodontids). The fauna has its strongest bio- ant to emphasize that many genera achieved
geographic links with eastern Australia and very widespread distributions in Pangea. The
indicates that a distinctive East Gondwana ver- equatorial barrier does not appear to have been
tebrate faunal province persisted until at least impenetrable to terrestrial vertebrates (see also
the end of the Middle Devonian (Young 1981). Chatterjee 1987), and of the 35 known dicyno-
By the latest Devonian the vertebrate faunas of dont genera approximately 11 are known from
Euramerica and East Gondwana are much more two or more cratonic regions. Lystrosaurus, the
similar, particularly in the composition of their famous dicynodont from the Fremouw For-
respective freshwater assemblages. Young links mation of Antarctica, is now known to have
this change to major palaeogeographic recon- been particularly widely dispersed. Semi-
structions and suggests that this may have been aquatic and probably able to forage for a variety
the time when the ocean separating these two of food types, it may have been the most suc-
supercontinents disappeared. The presence cessful mammal-like reptile of all time.
of a significant number of primitive taxa (or One of the most significant palaeontological
paraphyletic stem groups) in Antarctica and discoveries yet made in Antarctica is that of a
Australia suggests that this extensive continental series of fossil forests. Characterized by both
region was the site of origin of major groups of petrified logs and in situ stumps bearing clearly-
INTRODUCTION 3
defined annual growth rings, they are now occurs in later Cretaceous beds) and northwards
known from the Permian and Lower Triassic through South America. Similarly, the gymno-
of the Transantarctic Mountains, the Lower sperm Dacrydium balansae/D, bidwillii alliance
Cretaceous of Alexander Island and the Upper is now distributed from South America, through
Cretaceous-lower Tertiary of the nothern New Zealand, to certain Pacific islands and
Antarctic Peninsula region (e.g. Creber & Malaysia. Its history can be traced by the pollen
Chaloner 1984). So striking is their abundance genus Lygistepollenites and the oldest occur-
and diversity, and so mild the climatic deduc- rences (Coniacian-Santonian) shown to be
tions made from them, that it was thought at on the Antarctic Peninsula and in southeast
one time that they may indicate periods of Australia.
fundamental change in the growing conditions Particularly important in confirming the high-
in polar regions (perhaps related to shifts in the latitude origins of a number of angiosperm
tilt of the earth's axis; e.g. Jefferson 1983). lineages has been the elucidation of the
However, W. G. Chaloner & G. T. Creber Cretaceous-Tertiary boundary section on
have been interested in this problem for some Seymour Island by R. A. Askin. Data from
time, and in their review of the phenomenon of here are important in confirming that species
forest growth in Antarctica they suggest that from a range of families, which includes
the only factor which inhibits tree growth in the the Fagaceae, Myrtaceae, Proteaceae,
highest latitudes at the present day is tempera- Winteraceae, Casuarinaceae, Gunneraceae,
ture. It is now apparent that the annual input of Bombacaceae and Loranthaceae, have late
solar energy to the polar regions is sufficient to Cretaceous-early Tertiary first appearances
produce large annual growth rings and it has somewhere along the southern Gondwana mar-
been demonstrated that many evergreen trees gins. It should be emphasized, however, that
can tolerate long, dark winters. Studies on the not all these families necessarily originated in
composition of both living and fossil forests this region. Myrtaceous pollen, for example, is
have shown that the low angle of sunlight does known from early Campanian and Maastrichtian
not cause excessive mutual shading of trees. sediments on the Antarctic Peninsula; this oc-
It has long been suspected that many of the currence clearly predates Paleocene records in
key elements of today's Southern Hemisphere Australia and New Zealand, but postdates those
humid and perhumid forests could be traced from undifferentiated 'lower Senonian' in
well back into the Mesozoic period. This is Borneo and Santonian in Gabon. In her paper,
particularly so for certain lycopods, ferns, Dettmann speculates that a number of austral
araucarians and podocarps which may belong to angiosperm taxa may actually have had northern
lineages originating as early as the early Jurassic, Gondwanan or even Laurasian origins and then
or even the late Triassic (Fleming 1963, 1975). migrated via a South America-Antarctica-
Certainly, elements of the extensive Late Australia route.
Jurassic evergreen coniferous rainforests that It seems to be possible to conclude that
covered much of the southern Gondwana mar- Antarctica was both an important source
gins have persisted to the present day (Jefferson area and dispersal corridor (during the early
1983; Dettman 1986). Cretaceous-early Tertiary period) for austral
If much of the early evidence for the antiquity plants now living at mid- to low latitudes. Quite
of austral forest floras came from macro- why a number of major groups should either
palaeobotany, much of the latest information originate in, or disperse through, the highest
stems from a proliferation of palynological in- southern latitudes is not readily apparent, but,
vestigations. Utilizing both on- and offshore as Askin points out, the climate during this
successions, and a wide variety of palynoflora period was for the most part mild and equable.
taxa, it is now possible to establish at least Indeed, it could well be that an unusual com-
the Cretaceous ancestry of a number of living bination of climatic conditions (comprising a
groups. M. E. Dettmann, for example, shows polar winter/summer light cycle, low light angles
how the fern Lophosoria, which is restricted to and relatively mild and wet weather at the
South and central America, had a much wider poles) provided an unique stress impetus for
distribution in the Cretaceous and Tertiary. diversification of a wide variety of taxa. We
Traced by its unique spore, Cyatheacidites will return to polar origination events and the
annulatus (Dettman 1986), this genus can time-discrepancy of high- and lower-latitude
be shown to have its earliest stratigraphic oc- occurrences of taxa (often referred to as
currence (in basal Cretaceous strata) in the heterochroneity) below.
Antarctic Peninsula-South America region; it We know that for much of the Cretaceous
then migrated eastwards to Australia (where it Antarctica was covered by thick forests,
4 J.A. CRAME
but which animals may have inhabited them? New Caledonia, Papua New Guinea and South
Although the Cretaceous terrestrial and fresh- America. However in the Jurassic, eastern
water faunal record from the continent is still Gondwana rotated in a direction away from
poor, T. H. Rich, P. V. Rich, B. Wagstaff, the South Pole and this led directly to a sub-
J. McEwen-Mason, C. B. Douthitt & R. T. stantial improvement in global climates.
Gregory maintain that it is possible to specu- Simultaneously, new shallow-water migration
late on the composition of at least the Early routes were established around the southern
Cretaceous biotas by close comparison with Gondwana margins and by the Middle Jurassic
southeastern Australia. At this time the two warm-temperate Tethyan faunas were able
continents were of course still joined and it is to spread to Australia, New Caledonia, New
estimated that the terrestrial vertebrates pre- Zealand and West Antarctica. Such a scenario
served within the Otway and Gippsland basins is certainly borne out by the belemnite as-
could have lived at as high a palaeolatitude as semblages described by P. Doyle & P. J.
85~ Hypsilophodont dinosaurs are particularly Howlett, for in the Middle-Late Jurassic a
common in these Victoria coast assemblages, distinctive Belemnopsis- Hibolithes- Duvalia
and there is at least one theropod, Allosaurus fauna can be traced southwards from European
sp.. Turtles are common too, although they Tethys to the Gondwana margins. Local en-
seem to belong to primitive types, and there is a demic centres can be identified at this time,
variety of fish (including ceratodont lungfish such as in the embryonic trans-Gondwana sea-
and the unique Australian koonwarrids), lepido- way, but they are only discernible at the species
saurs, pterosaurs, plesiosaurs (presumably level.
freshwater) and birds, together with a possible By the Early Cretaceous, extensive rifting
labyrinthodont amphibian. That dinosaurs had occurred across Gondwana and the proto-
could have lived in southeastern Australia (and Indian and South Atlantic oceans were begin-
thus Antarctica too) is not altogether surprising ning to form. This in turn led to a reversal of the
for they have been known for some time from rotation of the eastern Gondwana block and the
Late Cretaceous deposits on the North Slope of re-introduction of its southern margins into
Alaska. Rich et al. believe it unlikely that the the highest latitudes. Here, a cool-temperate
herbivorous hypsilophodontids would have molluscan fauna developed, with one of its most
migrated vast distances to areas of winter day- important components being the endemic
light and suggest instead that a large brain and belemnite family, the Dimitobelidae. Further
eyes may have pre-adapted them to the low opening of the Atlantic and Indian oceans oc-
light conditions of polar habitats. curred throughout the Late Cretaceous and it
By comparing assemblages from both was at this time that New Zealand's land links
Australia and southern South America, R. E. with the rest of Gondwana were severed by
Molnar has suggested that the one dinosaur the formation of the Tasman Sea. However,
known from the Late Cretaceous of Antarctica as Stevens points out, there were still good
(an ankylosaur from James Ross Island; Olivero Late Cretaceous shallow-marine connections
et al. in press) was part of a terrestrial fauna between New Zealand, West Antarctica and
that included hypsilophodontids, pterosaurs, South America. A cool-temperate Weddellian
ratites, sphenodontians, leiopelmatid frogs and Province can be established between these local-
ceratodontid lungfish. In reviewing austral ities until well into the Paleogene (Zinsmeister
Mesozoic terrestrial vertebrate faunas, he was 1982).
struck by the fact that three of the fifteen The abundant early Campanian-late Eocene
known genera from the Jurassic-Cretaceous marine invertebrate faunas of the James Ross
of Australia (Allosaurus, Austrosaurus and Island region have been investigated by R. M.
Siderops) seem to have been relicts. Coupled Feldmann & D. M. Tschudy. They are particu-
with the occurrence of the apparent labyrinth- larly interested in an unusually rich decapod
odontid amphibian, this could be taken to indi- crustacean fauna which occurs in association
cate one of two things: either there was some with ammonites, bivalves, gastropods, echino-
form of geographical barrier protecting the derms and brachiopods. Concentrating on the
Australian region (and Antarctica too?), or macrurous decapods (lobsters), they have been
certain taxa may have preferentially survived in able to identify four species in the shallow-water
polar regions (cf. Vermeij 1987). Campanian-Paleocene Lopez de Bertodano
Returning to the marine realm, G. R. Stevens Formation. Hoploparia stokesi is extremely
indicates that a distinctive cool-temperate abundant, but shows no clear-cut evolutionary
Maorian province can be detected within the trends in shape over a 20 Ma period. Metane-
Triassic invertebrate assemblages known from phrops jenkinsi, a probable derivative of H.