Professional Documents
Culture Documents
R. Thomas Becker
Museum fur Naturkunde, Invalidenstr. 43, D-10115 Berlin, Germany
e-mail: thomas.becker@rz.hu-berlin.de
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Figure 1 Global (pantropical) distribution of Praemeroceras petterae (Petersen) in UD II-E (rhomboidea Zone), originally only known from the Canning Basin. Slightly altered
reconstruction from Heckel and Witzke (1979), NZ. = New Zealand, Ma. = Malaysia, B. = Burma, le. = Indochina, S. CH. = South China, TB. = Tibet, TM. = Tarim,
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KAZ. = Kazakhstan, N. CH. = North China, J. = Japan, MAD. = Madagascar, KO. = Kolyma, N. EUR. = Northern Europe, S. EUR. = Southern Europe, N. AM. = /'0
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North America, S. AM. = South America.
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Western Australian ammonoid palaeobiogeography 387
discussed for Frasnian conodonts by Klapper (1995). known in the world, compnsmg a total of 165
In addition, the detailed time resolution of species-level taxa (excluding ten species of
ammonoid data gives prospects for a much refined bactritids) in the Frasnian (Upper Devonian = UD l-
history of palaeobiogeographical changes. Thus, the A/B) to middle part of the Famennian (UD IV-B).
relative proportion of endemism has been analyzed This forms the basis for a re-appraisal of faunal
zone by zone in Canning Basin faunas. links with other regions as well as for a regional
A global diversity analysis of all Upper Devonian diversity analysis. An overview of Canning Basin
ammonoids at the generic level and based on the ammonoid zones and their correlation with
highest available time resolution has been chronostratigraphic units and conodont zonations
conducted by Becker (1993a). This study gave clear is given in Table 1.
relationships between eustatic changes and times of
diversification and extinctions ("species area Methodological aspects
effect"). Major extinctions were associated with Canning Basin ammonoid taxa fall into three
anoxic events and subsequent rapid regressions but categories concerning their spatial distribution: (a)
ammonoids could flourish in times of long-lasting taxa known from several to many widely separated
global sealevel lowstands. So far, there have been areas (semicosmopolitan - cosmopolitan in the
no published investigations at the species level and warm water biome = pantropical), (b) endemic taxa,
on regional scales, in order to elucidate differences and (c) "spot taxa" with patchy distribution in two
between lower and major taxonomic levels, or to or three widely separated regions, without evident
establish differences between regional and global intermediate occurrences. The latter are especially
diversity patterns. Of special interest are links interesting for the establishment of faunallinks and
between diversity changes and the degree of give evidence that an incomplete fossil record can
endemism since both were influenced by sealevel artificially increase the number of supposed
changes. endemics. Westermann (2000) has recently
proposed the term "didemic" for taxa with rather
disjunct occurences but this includes only
CANNING BASIN AMMONOID distribution in precisely two regions.
PALAEOBIOGEOGRAPHY Many endemic taxa are also closely related to
European, North African, or American forms and it
Previous work and data base is difficult to judge whether these were true
Principles of Devonian ammonoid allopatric species in a biological sense or
palaeobiogeography have been established by morphologically distinctive regional morphotypes/
House (1964, 1973, 1981). A recent review of general subspecies. Tests for genetic isolation by
trends in the Devonian biogeographic history of the geographically intermediate populations are not
group was given by Becker (in Becker and available. For this reason, the current analysis does
Kullmann 1996). Palaeobiogeographic relationships not distinguish between species and subspecies.
of ammonoids from the Famennian of Western Thus, for a cautious faunal comparison, closely
Australia (Canning Basin) were discussed by related taxa are also taken into consideration (Table
Teichert (1943) and Petersen (1975; see also Becker 2).
1993b). They found strong similarities with other, Many widespread Devonian ammonoid species
widely separated regions that formed part of an are still the subject of detailed taxonomic revision
Upper Devonian equatorial Prototethys (Figure 1). and a number of species names used in the
Based on the congruent evolution of monographs by Glenister (1958) and Petersen (1975)
5poradoceratidae in widely distant regions, will have to be changed due to the restudy of old
Petersen (1975: 11) postulated a permanent gene type material. For example, Maeneceras biferum
flow between European and Australian populations (Phillips) is a late Hembergian (UD IV) taxon and
in the Famennian. But genetic isolation must have cannot be used any more as an Upper Nehdenian
existed in other goniatite and clymenid groups at (UD II-G) zonal index, as shown by Becker (1993a,
the same time and along the same route of potential b) and Becker and House (1997). Updated faunal
faunal exchange. Differences in the degree of lists and range charts will be published elsewhere.
endemism of higher taxa may provide insights into Regardless of nomenclatoral aspects, the
palaeoecological aspects which enabled or restricted comparison of distant faunas is based on as
the formation of pantropical genepools. consistent taxonomic concepts as possible. For some
Intensive new field work in the Canning Basin regions (e.g., North China, Algeria, Urals)
since 1989 has greatly expanded the knowledge of published data had to be re-interpreted without the
ammonoid taxonomy and stratigraphy of the region opportunity to examine type material. Comparison
(Becker et al. 1993, Becker and House 1997). By now, of most key regions, however, is based on recent
it is the region with the most diverse and most detailed studies including the author (e.g., Becker
detailed Upper Devonian ammonoid succession 1992, 1993b, Becker and House 1993, 1994, House
388 R.T. Becker
E2 Ponticeras discoidale
E1 Probeloceras lutheri lutheri 5
D ?no record
C2 Manticoceras Sp. -
4 transitans
C1 Timanites angustus Timan
B3 Koenenites n.sp. 3
falsi-
B2 "Protimanites" pons 2 ovalis ?Genundewa
c--
B1 Chutoceras n.sp.
Table 1 Regional ammonoid zonation of the Canning Basin and its correlation with conodont zones and
chronostratigraphic units. UD = Upper Devonian.
Western Australian ammonoid palaeobiogeography 389
Table 2 Frasnian/Famennian faunal similarities between the Canning Basin and twenty other basins/crustal blocks,
based on the number of identical species and on the number of additional closely related species.
Region Frasnian: Frasnian: Frasnian: Famennian: Famennian: Famennian:
identical related "total identical related "total
species species similarity" species species similarity"
NW Canada 2 2 3 2 5
W. United States 1 1 3 3
E. United States 18 9 27 2 0 2
SW England 12 6 18 3 3
Armorican Massif 1 1 3 3
Ardennes 16 1 17 5 5
Germany 32 19 51 27 14 41
Montagne Noire 15 5 20 11 2 13
Kantabrian Mts. 6 6 5 1 6
North Africa 14 8 24 22 5 27
Poland 4 3 7 21 9 30
Ural 9 3 12 18 7 25
Timan 11 7 18 1 1
Novaya Zemlya 4 5 9 6 3 9
Chios, Greece 6 1 7
Kazakhstan 6 6
Turkestan 2 2
Rudnyi Altai 7 1 8
North China 8 1 9
South China 5 3 8 1 1
Canning Basin 99 69
and Kirchgasser 1993, Becker et al. 1997, 2000, Qualitative analysis: "Spot Taxa" and other
unpublished work in Morocco, North America, indicators
Germany and other regions).
Faunal comparisons have been made with twenty Frasnian
other regions (Table 2) which include some blocks Typical Frasnian cosmopolitan species are
with rather limited scientific investigation (e.g., Manticoceras cordatum (Sandberger and Sandberger),
Chios, North China, Turkestan, Rudnyi Altai, M. lamed (Sandberger and Sandberger),
Novaya Zemlya, Urals: Bashkiria) or with rather Aulatornoceras auris (Quenstedt), and Tornoceras
poor faunas in terms of abundance and diversity typum (Sandberger and Sandberger). The latter is
(e.g., Frasnian of Poland and western United States; commonly but wrongly quoted in the literature as
Canada, South China; early to middle Famennian of T. simplex (v. Buch), which represents a Middle
the Ardennes Shelf, of England and of Kazakhstan). Devonian nomen dubium, probably a
No distinction has been made between German holzapfeloceratid (Becker 1993b). The listed taxa
regions south (Saxothuringia) and north (Rhenish seem to be both very widespread and rather
Massif/Harz Mountains) of a supposed "Rheic bradytelic but future taxonomic revision may show
Ocean", since no intra-German biogeographic that populations from various regions and strata
differences have been recognized at all in the Upper can also be separated. Species of Manticoceras (in a
Devonian. The restricted Kazakhstan data base wider sense), especially M. intumescens (Beyrich),
excludes the prolongation of the Urals into the have been frequently misidentified.
western part (Aktyubinsk region/Mugodzhar Significant Frasnian "spot taxa" of the Canning
Mountains) of the country and leaves the Basin are Prochorites alveolatus (Glenister) and
Karaganda Basin and Semipalatinsk regions as one Probeloceras lutheri (Clarke) (suggesting a link to
entity. Records from other regions are so incomplete New York State), the Costamanticoceras koeneni
and sporadic that they were omitted. These include (Roemer) and "Maternoceras" prumiense (Steininger)
the Frasnian of Kolyma (NE Siberia), of the Groups (link to the Rhenish Massif), Manticoceras
Kuznetsk Basin, and of Kirgisia as well as the evollltllm Petter (link to North Africa), and rare M.
Famennian records from Bohemia, Moravia, Bolivia, latisellatum Yanishewsky and M. solnzevi
the Caucasus, and Taimyr. Faunas from the Carnic Bogoslovskiy (link to the Timan). Ponticeras
Alps and Iran are under current detailed study and discoidale described by Glenister (1958) from
interpretations have to await the forthcoming Western Australia was reported by Gatley (1983) to
results. occur also in the Ardennes (Belgium). Much richer
\;J
TIMANITES
BELOCERAS
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15
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60
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s
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Figure 2 Frasnian distribution of lower Frasnian Timanites (squares) and of middle to upper Frasnian Beloceras (dots) showing significant differences in their distribution ..,11>
""
~
patterns. Timanites is lacking in the western Prototethys region, Beloceras along the equatorial Transarctic Route.
Western Australian ammonoid palaeobiogeography 391
middle Frasnian Ponticeras assemblages occur in the confirms the necessity for a rigorous and consistent
Timan. Southern Chinese species of Mesobeloceras, taxonomy and the suspicion that true endemism
Beloceras, and Splweromanticoceras were partly was perhaps somewhat lower than is apparent from
described under different generic names but are current data.
very close, if not conspecific, with members of the Famennian examples of "spot taxa" are Tornia
genera from the Canning Basin, Germany, and n.sp. and the Cheiloceras postinversllm-semiinversllm
North Africa. Group (indicating links to Poland), Protactoclymenia
Conclusive is the distribution of early Frasnian krasnopolski (Tschernyscheff) (link to the Urals),
(UD I-C) Timanites (Figure 2), which was spread Karaclymenia n.sp. juv. (link to Polar Russia), and
along the equator from the Canning Basin via the Maeneceras milleri (Famennian link to New York;
Urals (Bashkiria, Tartar Republic, western slope of Figure 3). The last, however, is related to M.
Middle Ural), Timan, Polar Ural and Novaya aClltolaterale (Sandberger and Sandberger) and M.
Zemlya to Western Canada (Transarctic Route of sedgwicki Wedekind from Germany and Iran. The
House 1973 and Becker 1993a, b). For unknown late middle Famennian (UD IV) Raymondiceras is
reasons, and neglecting false records of rare but occurs as widely separated as in the
homoeomorphic late Givetian Eobeloceratidae (see Canning Basin (R. inceptllm Petersen), Southern
Becker and House 1993), the genus did not inhabit Urals (R. aktllbense Bogoslovskiy), and in Montana
the Western European, North African, and eastern (R. simplex (Raymond». Pseudoclymeniidae can be
North American regions. The late middle to upper very common in lower Hembergian (UD Ill-B) beds,
Frasnian Beloceras shows a partly reverse but show a patchy distribution in Germany, the
distribution excluding the Transarctic Route and Urals and Western Australia. Similar irregular
New York but with wide distribution on North patterns were recognized in the Prolobitidae but
Gondwanan crustal blocks such as North Africa new records of endemic genera from Morocco and
(South Morocco, Southwest Algeria and eastern the Canning Basin expand our
Moroccan Meseta), the Montagne Noire, Pyrenees, palaeobiogeographical knowledge of the family
Cantabrian Mountains, West Iberia, Carnic Alps, considerably.
Eastern Iran, and South China (Guangxi, Yunnan)
as well as in the surrounding of the "Rheic Ocean" Simple quantitative analysis
including SW England, Belgium/Northern France, There are strong biogeographic differences
and Germany (Rhenish Massif, Harz Mountains, between taxonomic levels. Australia has no endemic
Thuringia). Another branch of distribution led Devonian ammonoid family and at the generic level
around Kazakhstan to the Rudnyi Altai and to the the relative rate of endemism is also low (around
Kusnetsk Basin. The westerly restricted Beloceras 5% of all genera). At the species level, endemism is
distribution is even more remarkable in the light of significant and lies around 50% and over, both in
the fact that ancestral members of its lineage the Frasnian and in the Famennian. Analyzed zone
(Naplesites, Mesobeloceras) occur in New York State by zone, the average rate of endemism is around
and on Novaya Zemlya (also in Poland). Beloceras 40% in both stages. This rather high rate seems to
williamsi Wells of New York was re-assigned by reflect the wide spatial distance between eastern
House and Kirchgasser (1993) to the triainoceratid and western parts of the Prototethys (Figures 1-2).
genus Wellsites. An indication that Beloceras ranged Since investigations of Canning Basin faunas have
into the eastern part of the Ural seaway is based on proceded much farther than in other regions, it can,
an unconfirmed note in Tiasheva (1961: 99) that the however, be expected that some Australian
genus co-occurs with Manticoceras faunas in supposed endemics will eventually be discovered
Bashkiria. elsewhere, as has happened in the past (see above:
Praemeroceras petterae and Pan ticeras discoidale).
Famennian Despite the high number of endemic species-level
Early Famennian (UD II) goniatites have been taxa it is not suggested that Canning Basin faunas
intensively revised by Becker (1993b), which allows belong to a clearly defined eastern Gondwana
a detailed evaluation of Canning Basin province but subprovince status (see Westermann
cheiloceratid, tornoceratid, dimeroceratid and 2000; chorotype Canning Basin, chronotype =
sporadoceratid faunas. Praemeroceras petterae, Famennian) may be granted. The low-level
originally described by Petersen (1975) as an distinction is based on the fact that there are no
endemic form of Western Australia, turned out to endemic genera in the studied faunas which are
be globally widespread (Becker 1993b). It was common as well as very distinctive.
previously illustrated under different names or was The comparison of the Canning Basin with twenty
subsequently reported from Poland, Germany, other regions (Table 2) gives a clear picture of
Morocco, Chios (eastern Greek Agais), the faunal similarities that bears no easy correlation
Canadian Rocky Mountains and (new record with estimated distances between crustal blocks or
herein) from Eastern Iran (Figure 1). This example with their postulated spatial configuration. There is
392 R.T. Beeker
._,
'I
,-'.
' '
Figure 3 Maeneceras milleri (Flower and Caster), a typical "spot taxon" known from a single specimen in eastern North
America (3A-C, natural size, from Miller 1938: PI. 37, Figures 5-7) and abundantly from the late UD 11 of the
Canning Basin (D-E; GSWA 109145/ AI, "Syncline Gully", Bed K, Mt. Pierre area, natural size).
Western Australian ammonoid palaeobiogeography 393
also no fundamental palaeobiogeographic East Asian plates to account both for pantropical
difference between the two Upper Devonian stages. genepools of pelagic forms and for diverging outer
The global Kellwasser Crisis at the end of the shelf facies differences.
Frasnian had no lasting effect on principle fauna1 Assemblages from South China (Hunan, Guangxi,
exchange although there are some interesting trends Guizhou, and the Baoshan region of Yunnan) are
in detail. also rather rare and of low-diversity although
The greatest faunal similarities during all of the typical cephalopod facies were present in some
investigated time-span are with the classical faunas depressions between wide reefal platforms. The
from Germany. In the Frasnian there are 32 poor affinities between the Canning Basin and
common species (33%) and further 19 similar taxa, Southeast Asian regions is again thought to reflect
giving a "total similarity" of more than 50(10. In the mostly differences in shelf facies rather than the
Famennian there are 27 common species (39%) and presence of biogeographical barriers. It is possible
the wider similarity is close to 60% (41 taxa). that detailed future work will produce richer
Frasnian ammonoids show also clear relationships Frasnian faunas from South China. A few
("total similarity" between 20 and 30%) with the unpublished goniatites have been seen by the
Timan, North Africa, the Montagne Noire, the author in regional museum collections at Guilin and
Ardennes Shelf, SW England, and with Eastern Nanning.
North America (mostly New York State). These The Famennian picture differs insofar that
regions all formed one larger realm within or ammonoid faunal similarities with North America
adjacent to the western Prototethys. Close affinities and the Timan were lost. This is not based on
with the Canning Basin emphasize that there was differences in faunal composition but, again, on the
no general biogeographic barrier, neither too deep simple fact that early to middle Famennian
oceanic nor too shallow, between eastern and ammonoids are almost lacking in the two regions.
western Prototethys regions plus the Uralian- After the Upper Kellwasser Event, typical
Transarctic seaway. This important conclusion is in ammonoid biofacies did not re-develop for a long
some conflict with the plate tectonic reconstructions period. The influence of regional structural and
of Scotese and McKerrow (1991) which shows an facies evolution was the dominant factor controlling
extensive central Prototethys ocean that should at a restricted ammonoid distribution, not increased
least have produced a clearer separation of Russian spatial distance or the establishment of migrational
faunas. Migrations from and to Europe and North barriers. A reverse process can explain the increased
Africa could have taken place along the equatorial Famennian faunal similarity between the Canning
northern margin of Gondwana but this is Basin, the Urals (18 common species) and the Holy
contradicted by the absence of Frasnian Cross Mountains (21 common species). At the same
intermediate faunas from Tibet, Pakistan, time poorly studied faunas appeared in regions
Afghanistan, Iran or Turkey and by the lack of lacking Frasnian goniatites completely: Greek
Timanites in the western Prototethys. In the Scotese Agais, North China (Great Khingan), Caucasus,
and McKerrow reconstruction, Beloeeras faunas Turkestan, and Kazakhstan. Transgression
from the Rudnyi Altai and Kusnetsk Basin also produced new regions hospitable for ammonoids
would be completely isolated. The Devonian but spreading events, such as the one associated
reconstruction of Kent and Van der Voo (1990) with the global Annulata Event (Becker and
shows an even wider ocean without intermediate Kullmann 1996: Figure 4), did not reach all basins.
plates between Australia and Europe and is anyway The ammonoid analyses seems to contradict to
dismissed here because of the highly unlikely wide some extent palaeogeographic results obtained from
ocean separating North Africa and Europe. the study of Asian and Australian vertebrates. Rich
Crustal blocks positioned in the neighbourhood and Young (1996) observed that fish faunas of both
of northwest Australia, especially former parts of regions became more similar at some stage in the
Western Indonesia-Malaysia-Thailand, Vietnam- Upper Devonian and suggested a narrowing of the
Laos-Cambodia, Burma, and Tibet have no marine barriere between both crustal units, fitting
recorded Upper Devonian ammonoid faunas at all; the Metcalfe (1996) reconstruction. Palaeomagnetic
ammonoid facies never developed there or did not data (Zhao et al. 1996), however, indicate that
occupy significant areas. Southeast Asian blocks Chinese blocks were pulling away from the western
seem to have had a different shelf-facies history. If part of Australia as early as between the Early
placed just west of Australia, as in the Silurian and Middle Devonian. Furthermore,
reconstruction of Metcalfe (1996) and to a lesser Johanson and Ritchie (1999) recently pointed out
extent (W jNW) also in Scotese and McKerrow that latest Devonian fish faunal similarities between
(1990), these plates (Figure 4) would have restricted South China and eastern Australia were not as close
the faunal exchange to the west more than evident as originally thought and during most of the Upper
from the data. The reconstruction of Heckel and Devonian there were close affinities with
Witzke (1979) leaves more space between South Euramerica. Stratigraphical data in Rich and Young
394 R.T. Becker
30N
LATE DEVONIAN
308
Figure 4 Rejected plate tectonic reconstructions for the Upper Devonian by (A) Scotese and McKerrow (1990) and by
(B) Metcalfe (1996) showing too wide a central Prototethys ("Palaeotethys") and blocking of the Canning
Basin (black dots) by an array of Southeast Asian crustaI blocks. WB = West Burma, S = Sibumasu (parts of
Indonesia, Malaysia and Thailand), L = Lhasa, QI = Qiangtang, WC = Western Cimmerian Continent (ca.
Turkey to Pakistan), NC = North China, I = Indochina (parts of Vietnam, Laos/Cambodia), T = Tarim, Q =
Qaidam, SC = South China. The Scotese and McKerrow reconstruction is especially in conflict with the
Transarctic and Kazakhstan distribution of Timanites and Be/oeeras (Figure 2).
(1996: Figure 2) show that faunal links between Canning Basin DD IV-assemblages (Piker Hills)
eastern Australia and China appeared late in the were found in a rather unusual hemi-pelagic
Famennian when they were no more ammonoids in shallow black shale facies with many gastropods,
Western Australia. brachiopods, and plant remains. New South Wales
faunas (Jenkins 1966, 1968) contain both endemic
"Intra-Australian" relationships species (e.g., "Genuclymenia" keepitensis) and forms
In Famennian II and IV, the Canning Basin and known from Western Europe and North Africa
New South Wales have not a single species in (Erfoudites, Pia tyclymen ia). Generally, western and
common. Eastern Australian ammonoid source eastern Australian Devonian ammonoid faunas
areas are part of terranes which were attached later represent completely disjunct basins and
to the craton. There was certainly again a facies subprovinces that both had some permanent faunal
influence on different faunal composition since exchange with western parts of the Prototethys.
Western Australian ammonoid palaeobiogeography 395
Migration from Eastern Australia via the types of morphology and paleoecology. Only a
Protopacific (Panthalassia) is ruled out because of more general study might clarify whether
the lack of similar faunas in western North representatives of certain morphotypes tend to be
America. more endemic or cosmopolitan than others.
species LKW
EVENT
15
BUGLE GAP
EVENT
UKW
EVENT
ANNULATA
EVENT
10
CONDROZ
EVENTS
B c E F G K GAB CAB
UPPER DEVONIAN I UPPER DEVONIAN 11 UO III UDIV
FRASNIAN FAMENNIAN
Figure 5 Species level total extinctions in the Upper Devonian of the Canning Basin showing three major Frasnian
extinctions (Bugle Gap, Lower Kellwasser and Upper Kellwasser Events) and some Famennian phases of
high faunal overturn (Condroz Events, UD II-F IG, extinction subsequent to the Annulata Event).
those from German or Polish submarine transgressive episodes associated with the
seamount sections positioned in deeper parts of Genundewa and Timan Events.
shelf margin basins. Several diversity peaks b) Small scale extinction at the end of VD I-C
clearly correlate with transgressive episodes, but corresponding with the global Koenenitidae
not all eustatic rises (see Becker and House 1997) decline near the end of MN Zone 4 (= transitans
allowed diversification or the spread of more Zone).
species into the Canning Basin. This indicates a
c) Important regional diversification, with a peak
significant influence of regional structural
in VD I-F2 (Gogoceras nicolli Zone; ca. MN Zone
evolution. The maximum diversity with 31
6 = upper part of punctata Zone), correlating
species/subspecies was reached in the
with the regional main development of hypoxic
Virginoceras erraticum Zone (VD I-J 2). Plots of total
goniatite shales.
species level diversity and of total extinctions
(Figures 5-6) give the following stages of d) Bugle Gap Event (named herewith): Significant
diversity development (for event sequence and regional extinction associated with a retreat of
terminology see Becker et al. 1993 and Becker and hypoxic facies and regression at the end of VD
House 1997): I-F2 (within MN Zone 6) in the Bugle Gap area.
a) Early Frasnian (VD I-B/C) immigration and This event may be an important and
speciation of low-diverse faunas, probably characteristic regional signature of the Canning
corresponding with the international Basin.
Western Australian ammonoid palaeobiogeography 397
I endemiC
•
LKW
EVENT
PRESERVED SPECIES LEVEL
TOTAL DIVERSITY
CANNING BASIN, W. AUSTRALIA
~
cosmopolitan/
NEHDEN
BUGLE GAP RADIATION
EVENT
SEMICH. CONDROZ
EVENT EVENTS
UKW
EVENT
o ANN.
penes EVENT
B C o El G H I K AIBCDE FIG
I HI AB CAB
UPPER DEVONIAN I UPPER DEVONIAN 11 UD III UDIV
FRASNIAN FAMENNIAN
Figure 6 Preserved species level total diversity in the Upper Devonian of the Canning Basin and the total number of
endemic species. Diversifications occurred in the early middle Frasnian (UD I-ElF), with the semichatovae
Transgression, in the upper Frasnian, lower Famennian (Nehden Radiation), early in UD 1II and with the
global Annlllata Event.
e) Stable and moderately low diversity in UD I-C/ j) Diversification in the latest Frasnian Crickites
H (MN Zones 7-10, = ca. hassi Zones). lindneri Zone (UD I-L ln , upper part of MN Zone
f) Important wave of immigration and speciation 13, = upper part of Late rhenana Zone) which is
at the base of UD I-I corresponding with the characterized by several regional transgressive
international "semichatovae Transgression" pulses.
(early MN Zone 11, jamieae Zone to lowest k) Upper Kellwasser Event: A regional decline of
part of Early rhenana Zone). ammonoid faunas started at the base of the
g) Stable high diversity in UD 1-1 2 and I-J l (MN lingllifomlis Zone (= topmost part of MN Zone
Zones 11-12, = Early rhellalla Zone), in 13, UD I-Llb)' characterized by a short-termed
agreement with the globally high sealevel. regression with a spread of neritic faunas (e.g.,
h) Maximum regional diversity (31 species/ diverse gastropods). The main extinction lies
subspecies) in the Virgilloceras erraticllm Zone above a subsequent transgressive level
(UD I-J 2, upper part of Zone 12, upper part of (Manticoceras gllppyi Bed) within the linglliformis
Early rhellalla Zone), followed by a massive Zone (see Becker et a/. 1991, Becker and House
extinction (27 species) during a significant 1997) which correlates with the Upper
regressive phase (top of MN Zone 1 It Kellwasser level but which is not hypoxic. The
corresponds in time with the global extinction last Mallticoceras range up to the last centimeter
just prior to the Lower Kellwasser level. of the Frasnian. The regional extinction pattern
i) Very low diversity and very rare faunas in the is more gradual than in Europe and North
lower part of MN Zone 13 (UD I-K, = lower Africa and this may be related to the complete
part of Late rhellalla Zone). The regional lack of lack of hypoxic Kellwasser facies. The final
faunas from this interval is intriguing but may sudden extinction of manticoceratids must have
partly reflect undersampling. been caused by factors such as temperature and
398 R.T. Becker
number
at species
LKW %endemics
30 • EVENT 100
NEHDEN
RADIATION
BUGLE GAP
EVENT
20
SEMICH. .'
EVENT
10
"
:
Figure 7 Canning Basin relationships between preserved total species-level diversity and the relative (percentage) rate
of endemism (interrupted lines), Low-diverse/highly endemic faunas alternate with species-rich/more
cosmopolitan faunas, However, there is also a smaller-scale positive correlation between changes in diversity
and endemism.
salinity that are not very obvious in the marginifera Zone) and in the lower part of VD
sedimentary record. Ill-A. The low diversity of VD-G 2 and VD III-
1) Earliest Famennian beds (VD II-A/B, Pa. A 3 reflects inadequate sampling. The global
triangularis to Middle crepida Zones), spanning and regional transgression at the base of VD
ca. 2 ma, have no ammonoids at all. Regionally, II-G (classical do liB, Maeneceras subvaricatum
there were no ammonoid survivors at all. Zone) is marked by the immigration of the
m) The rapid Nehden Adaptive Radiation (VD 1I- oldest sporadoceratids but not by a
CID, Vpper crepida Zone to lowermost diversification.
rhomboidea Zone) produced the second regional p) The immigration of pseudoclymenids took place
diversity peak, which correlates well with a during a minor diversification phase (VD Ill-B)
global and regional sealevel high. and their sudden extinction (within the Lower
n) The Lower Condroz Event at the end of VD lI- trachytera Zone) represents a highly interesting
D (Oxytornoceras n.sp. Zone) led to a significant small-scale global extinction event that is not
extinction which, however, was balanced by reflected by a change in lithofacies.
new forms entering in VD I-E (Praemeroceras q) The immigration of the oldest clymenids (base
petterae Zone, middle part of rhomboidea Zone). of VD III-C, ca. base of Vpper trachytera Zone)
0) Diversity remained moderately high during and prolobitids produced another minor
the higher parts of VD 11 and in the lower diversity increase but faunas remained of very
part of VD III (upper part of rhomboidea Zone low diversity in comparison with those from the
to Vppermost marginifera Zone), but there Vrals, Poland, and Germany. At the end of the
were several faunal overturns, for example regional Protactoclymenia euryomphala Zone
between VD I-F and I-G (within basal there was a small-scale extinction, correlating
Western Australian ammonoid palaeobiogeography 399
Famennian boundary interrupted the Canning II) von Europa und Nord-Afrika. Courier
Basin ammonoid record totally for ca. 2 ma, but Forschungsinstitut Senckenberg 155: 1-405.
caused no fundamental biogeographical change. Becker, RT. (1995). Taxonomy and Evolution of Late
8) All known global extinctions of the Frasnian to Famennian Tornocerataceae (Ammonoidea). Berliner
middle Famennian, even small-scale events geowissenschaftliche Abhandlungen, E 16: 607-643.
such as the koenenitid and pseudoclymenid Becker, RT. and House, M.R (1993). New early Upper
extinctions, can be recognized in the Canning Devonian (Frasnian) goniatite genera and the
evolution of the "Gephurocerataceae". Berliner
Basin. Magnitudes and phases, however, were
geowissenschaftliche Abhandlungen, E 9: 111-133.
strongly influenced by regional factors. The
Becker, RT. and House, M.R (1994). Kellwasser Events
Bugle Gap Event at the end of VD I-E 2 (upper
and goniatite successions in the Devonian of the
part of punctata Zone) was a significant regional Montagne Noire with comments on possible
extinction. causations. Courier Forschungsinstitut Senckenberg 169:
9) Analyzed zone by zone, there was an 45-77.
alternation of low diverse/endemic and highly Becker, RT. and House, M.R (1997). Sea-level changes in
diverse/cosmopolitan faunas. "Lazarus Phases" the Upper Devonian of the Canning Basin, Western
of endemic forms and a small-scale positive Australia. Courier Forschungsinstitut Senckenberg 199:
correlation between diversity trends and the 129-146.
relative rate of endemism suggest that many Becker, R.T. and Kullmann, J. (1996). Paleozoic
forms may episodically have lived outside the Ammonoids in Space and Time. In Landman, N.,
studied sections. True endemism was probably Tanabe, K. and Davis, RA. (eds.) Ammonoid
somewhat lower than is evident from current Paleobiology, Topic in Geobiology 13: 711-753.
data. Becker, RT., House, M.R, Kirchgasser, W.T. and
Playford, P.E. (1991). Sedimentary and faunal
changes across the Frasnian/Famennian boundary in
the Canning Basin of Western Australia. Historical
ACKNOWLEDGEMENTS Biology 5: 183-196.
This study would have been impossible without Becker, RT., House, M.R and Kirchgasser, W.T. (1993).
all the support and co-operation of M.R House Devonian goniatite biostratigraphy and timing of
(Southampton). He as well as G. Klapper and K. fades movements in the Frasnian of the Canning
McNamara (Perth) read and corrected the Basin. Geological Society of London, Special Publication
manuscript. Field work in the Canning Basin was 70: 293-321.
financed by grants of NERC (to M.R. House) and by Becker, RT., House, M.R and Klapper, G. (1997). Event
the DFG. It was conducted jointly with W.T. sequence and sealevel-changes in the Upper Givetian
and Frasnian of the Tafilalt (Southern Morocco). In C.
Kirchgasser (Potsdam, New York), G. Klapper
Brett (ed.) The Amadeus Grabau Symposium:
(Iowa City), P.E. Playford (Perth), RT. Brown International Meeting on Cyclicity and Bioevents in the
(Canberra), R Feist (Montpellier), R Crick Devonian System, Program and Abstracts: 18.
(Arlington) and others. The Geological Survey of Becker, RT., House, M.R., Menner, V.V. and
Western Australia generously assisted in providing Ovnatanova, N.S. (2000). Revision of ammonoid
vehicles, equipment and other support that enabled biostratigraphy in the Frasnian (Upper Devonian) of
extensive fieldwork. the Southern Timan (Northeast Russian Platform).
Acta Geologica Polonica 50: 67-97.
Gatley, S. (1993). Frasnian (Upper Devonian) goniatites from
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