Records of the Western Australian Museum Supplement No. 58: 385-401 (2000).

Palaeobiogeographic relationships and diversity of Upper Devonianammonoids from Western Australia

R. Thomas BeckerMuseum fur Naturkunde, Invalidenstr. 43, D-10115 Berlin, Germany

e-mail: thomas.becker@rz.hu-berlin.de

Abstract - Upper Devonian ammonoids from the Canning Basin of WesternAustralia represent one of the most diverse faunas globally known. It consistsof cosmopolitan (pantropical), endemic and "spot" taxa (with disjunctdistribution in few widely separated basin). Endemism is low at the genericbut very significant (ca. 50%) at the species level. Linked with regional facieschange and eustatic influences, there were alternating episodes with low­diverse, relatively highly endemic or with species-rich and rathercosmopolitan faunas. Faunal similarities both in the Frasnian and Famennianwere closest with Germany, slightly less with North Africa, SW England, theArdennes, and the Montagne Noire. Frasnian faunal links with the Timanand eastern North America were severed after the Frasnian-Famennianboundary whilst relationships with the Urals and Poland became closer.Faunal similarities were clearly more dependent on regional faciesdevelopments of plates than on their spatial distance. The regional diversitycurve reflects both global extinctions and radiations as well as effects ofCanning Basin structural evolution. Three major extinctions, the Bugle Gap,Lower Kellwasser and Upper Kellwasser Events occurred in the Frasnian.Not a single ammonoid species regionally survived the Frasnian-Famennianboundary and the basal Famennian lacked ammonoids all over Australia. Thepost-Annulata Event regression regionally initiated a strong final decline ofammonoid faunas.

INTRODUCTIONThe spatial distribution patterns of Devonian

ammonoids are still rather poorly studied althoughpalaeobiogeographic differences of pelagic groupsmay give distinctive signals for routes of faunalexchange between open shelf areas of major crustalblocks. In the Devonian, the plate tectonicconfiguration of continents and of smaller crustalplates is still controversal if the world maps ofHeckel and Witzke (1979), Scotese and McKerrow(1990), Kent and Van der Voo (1990), Bachtadse etal. (1995) and Metcalfe (1996) are compared. IGCP421 intends to use consistent taxonomic andbiogeographic data as well as regional event andextinction patterns to fingerprint former blocks ofNorthern Gondwana. Such signatures shall becompared with palaeomagnetic and tectonic data inorder to revise the reconstruction of Mid-Palaeozoicplate movements. In this context, intensive work inrecent years, jointly with M.R. House and W.T.Kirchgasser, will be used here for an analysis ofammonoid data from Western Australia. These holdimportant clues for the general understanding ofUpper Devonian ammonoid biogeographies and forplacing the famous Canning Basin reef complexesin a plate tectonic framework (Figure 1).

The interpretation of Mid-Palaeozoic ammonoiddistribution data has to recognize their activenektonic lifestyle, either in the open water columnor near to the seafloor, as well as the smallersignificance of postmortem drift in comparison toRecent nautilids (House 1987). Ammonoid shellswere flooded much faster and lived mostly indeeper quiet outer shelf settings lacking strongcurrents. Many taxa were pan(sub)tropical. Intheory, both very shallow marine seaways and deepoceanic areas may have acted as barriers forDevonian ammonoids. This follows frompalaeobiological constraints for maximum verticalmigration to the food-bearing seafloor by maximumimplosion depths (see Hewitt 1996) and fromsensitiveness to high water agitation (e.g., bystorms). Trends of endemic evolution in wide openseas, therefore, may be a more significant indicationof plate tectonic isolation of regions than endemismin benthic and nearshore assemblages subject tolarger influence of local environmental factors.However, in recent years, facies control onDevonian ammonoid distribution has become moreand more evident. This study intends to investigatethe importance of facies versus distance /barriercontrol on distribution patterns as it has been

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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 alteredreconstruction from Heckel and Witzke (1979), NZ. = New Zealand, Ma. = Malaysia, B. = Burma, le. = Indochina, S. CH. = South China, TB. = Tibet, TM. = Tarim,KAZ. = Kazakhstan, N. CH. = North China, J. = Japan, MAD. = Madagascar, KO. = Kolyma, N. EUR. = Northern Europe, S. EUR. = Southern Europe, N. AM. =North America, S. AM. = South America.

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Western Australian ammonoid palaeobiogeography

discussed for Frasnian conodonts by Klapper (1995).In addition, the detailed time resolution ofammonoid data gives prospects for a much refinedhistory of palaeobiogeographical changes. Thus, therelative proportion of endemism has been analyzedzone by zone in Canning Basin faunas.

A global diversity analysis of all Upper Devonianammonoids at the generic level and based on thehighest available time resolution has beenconducted by Becker (1993a). This study gave clearrelationships between eustatic changes and times ofdiversification and extinctions ("species areaeffect"). Major extinctions were associated withanoxic events and subsequent rapid regressions butammonoids could flourish in times of long-lastingglobal sealevel lowstands. So far, there have beenno published investigations at the species level andon regional scales, in order to elucidate differencesbetween lower and major taxonomic levels, or toestablish differences between regional and globaldiversity patterns. Of special interest are linksbetween diversity changes and the degree ofendemism since both were influenced by sealevelchanges.

CANNING BASIN AMMONOIDPALAEOBIOGEOGRAPHY

Previous work and data basePrinciples of Devonian ammonoid

palaeobiogeography have been established byHouse (1964, 1973, 1981). A recent review of generaltrends in the Devonian biogeographic history of thegroup was given by Becker (in Becker andKullmann 1996). Palaeobiogeographic relationshipsof ammonoids from the Famennian of WesternAustralia (Canning Basin) were discussed byTeichert (1943) and Petersen (1975; see also Becker1993b). They found strong similarities with other,widely separated regions that formed part of anUpper Devonian equatorial Prototethys (Figure 1).Based on the congruent evolution of5poradoceratidae in widely distant regions,Petersen (1975: 11) postulated a permanent geneflow between European and Australian populationsin the Famennian. But genetic isolation must haveexisted in other goniatite and clymenid groups atthe same time and along the same route of potentialfaunal exchange. Differences in the degree ofendemism of higher taxa may provide insights intopalaeoecological aspects which enabled or restrictedthe formation of pantropical genepools.

Intensive new field work in the Canning Basinsince 1989 has greatly expanded the knowledge ofammonoid taxonomy and stratigraphy of the region(Becker et al. 1993, Becker and House 1997). By now,it is the region with the most diverse and mostdetailed Upper Devonian ammonoid succession

387

known in the world, compnsmg a total of 165species-level taxa (excluding ten species ofbactritids) in the Frasnian (Upper Devonian = UD l­A/B) to middle part of the Famennian (UD IV-B).This forms the basis for a re-appraisal of faunallinks with other regions as well as for a regionaldiversity analysis. An overview of Canning Basinammonoid zones and their correlation withchronostratigraphic units and conodont zonationsis given in Table 1.

Methodological aspectsCanning Basin ammonoid taxa fall into three

categories concerning their spatial distribution: (a)taxa known from several to many widely separatedareas (semicosmopolitan - cosmopolitan in thewarm water biome =pantropical), (b) endemic taxa,and (c) "spot taxa" with patchy distribution in twoor three widely separated regions, without evidentintermediate occurrences. The latter are especiallyinteresting for the establishment of faunallinks andgive evidence that an incomplete fossil record canartificially increase the number of supposedendemics. Westermann (2000) has recentlyproposed the term "didemic" for taxa with ratherdisjunct occurences but this includes onlydistribution in precisely two regions.

Many endemic taxa are also closely related toEuropean, North African, or American forms and itis difficult to judge whether these were trueallopatric species in a biological sense ormorphologically distinctive regional morphotypes/subspecies. Tests for genetic isolation bygeographically intermediate populations are notavailable. For this reason, the current analysis doesnot distinguish between species and subspecies.Thus, for a cautious faunal comparison, closelyrelated taxa are also taken into consideration (Table2).

Many widespread Devonian ammonoid speciesare still the subject of detailed taxonomic revisionand a number of species names used in themonographs by Glenister (1958) and Petersen (1975)will have to be changed due to the restudy of oldtype material. For example, Maeneceras biferum(Phillips) is a late Hembergian (UD IV) taxon andcannot be used any more as an Upper Nehdenian(UD II-G) zonal index, as shown by Becker (1993a,b) and Becker and House (1997). Updated faunallists and range charts will be published elsewhere.Regardless of nomenclatoral aspects, thecomparison of distant faunas is based on asconsistent taxonomic concepts as possible. For someregions (e.g., North China, Algeria, Urals)published data had to be re-interpreted without theopportunity to examine type material. Comparisonof most key regions, however, is based on recentdetailed studies including the author (e.g., Becker1992, 1993b, Becker and House 1993, 1994, House

388 R.T. Becker

CHRONOSTRAT. AMMONOlDS CONODONTS EVENTS

UD B Protoxyclymenia sp. posteraIV A Protactoclymenia eurylobica f-- Annulata

C Protactoclymenia euryomphalaUpper trachytera

B2 Falcitornoceras n.sp. aft. bilobatumLower trachytera

UD B1 Pseudoclymenia australisIII

UppermostA3 Sporadoceras angustisellatum

A2 Pernoceras delepineimarginifera

Z A1 Protornoceras sp.et 12 Cycloclymenia n.sp.- Upper marginiferaZ 11 Dimeroceras n.sp. aft. padbergenseZW H Sporadoceras teicherti:E

G2 Maeneceras latilobatumet f--

u. G1 Maeneceras subvaricatum Lower marginifera "Enkeberg"f---

UD F2 Acrimeroceras n.sp.

11 F1 Paratornoceras polonicum

E2 Praemeroceras primaevum Upper rhomboidea

E1 Praemeroceras petterae

D Oxytornoceras n.sp.Lower rhomboidea _r::= Low. CondrozUppermost crepida

C "Falcitornoceras" n.sp. Upper crepida

A-B no record triangularis· M. crepida

L2 Manticoceras sp. lingui- Upper

L1b Manticoceras guppyi formis Kellwasser13 f---

L1a Crickites lindneri Upper

K (Aulatornoceras n.sp.) rhenana f-- LowerKellwasser

J2 Virginoceras erraticum12 Lower

J1 Maternoceras retorquatum rhenana

12 Playfordites tripartitus11

f--- semichatovae11 Serramanticoceras serratum jamieae

ZH Beloceras tenuistriatum 9 - 10 Upp. hassi

et G2 Mesobeloceras thomasi 8 Lower- 7 hassiZ G1 Naplesites housei RhinestreetfI) UD -et I F3 Sphaeromanticoceras affine

£t: F2 Gogoceras nicolliu. 6

F1 Prochorites alveolatus

E3 Manticoceras n.sp. punctata

E2 Ponticeras discoidale

E1 Probeloceras lutheri lutheri 5

D ?no record

C2 Manticoceras Sp.transitans -4

C1 Timanites angustus Timan

B3 Koenenites n.sp. 3falsi-

B2 "Protimanites" pons 2 ovalis ?Genundewac--

B1 Chutoceras n.sp.

Table 1 Regional ammonoid zonation of the Canning Basin and its correlation with conodont zones andchronostratigraphic 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 "totalspecies species similarity" species species similarity"

NW Canada 2 2 3 2 5W. United States 1 1 3 3E. United States 18 9 27 2 0 2SW England 12 6 18 3 3Armorican Massif 1 1 3 3Ardennes 16 1 17 5 5Germany 32 19 51 27 14 41Montagne Noire 15 5 20 11 2 13Kantabrian Mts. 6 6 5 1 6North Africa 14 8 24 22 5 27Poland 4 3 7 21 9 30Ural 9 3 12 18 7 25Timan 11 7 18 1 1Novaya Zemlya 4 5 9 6 3 9Chios, Greece 6 1 7Kazakhstan 6 6Turkestan 2 2Rudnyi Altai 7 1 8North China 8 1 9South China 5 3 8 1 1

Canning Basin 99 69

and Kirchgasser 1993, Becker et al. 1997, 2000,unpublished work in Morocco, North America,Germany and other regions).

Faunal comparisons have been made with twentyother regions (Table 2) which include some blockswith rather limited scientific investigation (e.g.,Chios, North China, Turkestan, Rudnyi Altai,Novaya Zemlya, Urals: Bashkiria) or with ratherpoor faunas in terms of abundance and diversity(e.g., Frasnian of Poland and western United States;Canada, South China; early to middle Famennian ofthe Ardennes Shelf, of England and of Kazakhstan).No distinction has been made between Germanregions south (Saxothuringia) and north (RhenishMassif/Harz Mountains) of a supposed "RheicOcean", since no intra-German biogeographicdifferences have been recognized at all in the UpperDevonian. The restricted Kazakhstan data baseexcludes the prolongation of the Urals into thewestern part (Aktyubinsk region/MugodzharMountains) of the country and leaves theKaraganda Basin and Semipalatinsk regions as oneentity. Records from other regions are so incompleteand sporadic that they were omitted. These includethe Frasnian of Kolyma (NE Siberia), of theKuznetsk Basin, and of Kirgisia as well as theFamennian records from Bohemia, Moravia, Bolivia,the Caucasus, and Taimyr. Faunas from the CarnicAlps and Iran are under current detailed study andinterpretations have to await the forthcomingresults.

Qualitative analysis: "Spot Taxa" and otherindicators

FrasnianTypical Frasnian cosmopolitan species are

Manticoceras cordatum (Sandberger and Sandberger),M. lamed (Sandberger and Sandberger),Aulatornoceras auris (Quenstedt), and Tornocerastypum (Sandberger and Sandberger). The latter iscommonly but wrongly quoted in the literature asT. simplex (v. Buch), which represents a MiddleDevonian nomen dubium, probably aholzapfeloceratid (Becker 1993b). The listed taxaseem to be both very widespread and ratherbradytelic but future taxonomic revision may showthat populations from various regions and stratacan also be separated. Species of Manticoceras (in awider sense), especially M. intumescens (Beyrich),have been frequently misidentified.

Significant Frasnian "spot taxa" of the CanningBasin are Prochorites alveolatus (Glenister) andProbeloceras lutheri (Clarke) (suggesting a link toNew York State), the Costamanticoceras koeneni(Roemer) and "Maternoceras" prumiense (Steininger)Groups (link to the Rhenish Massif), Manticocerasevollltllm Petter (link to North Africa), and rare M.latisellatum Yanishewsky and M. solnzeviBogoslovskiy (link to the Timan). Ponticerasdiscoidale described by Glenister (1958) fromWestern Australia was reported by Gatley (1983) tooccur also in the Ardennes (Belgium). Much richer

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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 distributionpatterns. Timanites is lacking in the western Prototethys region, Beloceras along the equatorial Transarctic Route.

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Western Australian ammonoid palaeobiogeography

middle Frasnian Ponticeras assemblages occur in theTiman. Southern Chinese species of Mesobeloceras,Beloceras, and Splweromanticoceras were partlydescribed under different generic names but arevery close, if not conspecific, with members of thegenera from the Canning Basin, Germany, andNorth Africa.

Conclusive is the distribution of early Frasnian(UD I-C) Timanites (Figure 2), which was spreadalong the equator from the Canning Basin via theUrals (Bashkiria, Tartar Republic, western slope ofMiddle Ural), Timan, Polar Ural and NovayaZemlya to Western Canada (Transarctic Route ofHouse 1973 and Becker 1993a, b). For unknownreasons, and neglecting false records ofhomoeomorphic late Givetian Eobeloceratidae (seeBecker and House 1993), the genus did not inhabitthe Western European, North African, and easternNorth American regions. The late middle to upperFrasnian Beloceras shows a partly reversedistribution excluding the Transarctic Route andNew York but with wide distribution on NorthGondwanan crustal blocks such as North Africa(South Morocco, Southwest Algeria and easternMoroccan Meseta), the Montagne Noire, Pyrenees,Cantabrian Mountains, West Iberia, Carnic Alps,Eastern Iran, and South China (Guangxi, Yunnan)as well as in the surrounding of the "Rheic Ocean"including SW England, Belgium/Northern France,and Germany (Rhenish Massif, Harz Mountains,Thuringia). Another branch of distribution ledaround Kazakhstan to the Rudnyi Altai and to theKusnetsk Basin. The westerly restricted Belocerasdistribution is even more remarkable in the light ofthe fact that ancestral members of its lineage(Naplesites, Mesobeloceras) occur in New York Stateand on Novaya Zemlya (also in Poland). Beloceraswilliamsi Wells of New York was re-assigned byHouse and Kirchgasser (1993) to the triainoceratidgenus Wellsites. An indication that Beloceras rangedinto the eastern part of the Ural seaway is based onan unconfirmed note in Tiasheva (1961: 99) that thegenus co-occurs with Manticoceras faunas inBashkiria.

FamennianEarly Famennian (UD II) goniatites have been

intensively revised by Becker (1993b), which allowsa detailed evaluation of Canning Basincheiloceratid, tornoceratid, dimeroceratid andsporadoceratid faunas. Praemeroceras petterae,originally described by Petersen (1975) as anendemic form of Western Australia, turned out tobe globally widespread (Becker 1993b). It waspreviously illustrated under different names or wassubsequently reported from Poland, Germany,Morocco, Chios (eastern Greek Agais), theCanadian Rocky Mountains and (new recordherein) from Eastern Iran (Figure 1). This example

391

confirms the necessity for a rigorous and consistenttaxonomy and the suspicion that true endemismwas perhaps somewhat lower than is apparent fromcurrent data.

Famennian examples of "spot taxa" are Tornian.sp. and the Cheiloceras postinversllm-semiinversllmGroup (indicating links to Poland), Protactoclymeniakrasnopolski (Tschernyscheff) (link to the Urals),Karaclymenia n.sp. juv. (link to Polar Russia), andMaeneceras milleri (Famennian link to New York;Figure 3). The last, however, is related to M.aClltolaterale (Sandberger and Sandberger) and M.sedgwicki Wedekind from Germany and Iran. Thelate middle Famennian (UD IV) Raymondiceras israre but occurs as widely separated as in theCanning Basin (R. inceptllm Petersen), SouthernUrals (R. aktllbense Bogoslovskiy), and in Montana(R. simplex (Raymond». Pseudoclymeniidae can bevery common in lower Hembergian (UD Ill-B) beds,but show a patchy distribution in Germany, theUrals and Western Australia. Similar irregularpatterns were recognized in the Prolobitidae butnew records of endemic genera from Morocco andthe Canning Basin expand ourpalaeobiogeographical knowledge of the familyconsiderably.

Simple quantitative analysisThere are strong biogeographic differences

between taxonomic levels. Australia has no endemicDevonian ammonoid family and at the generic levelthe relative rate of endemism is also low (around5% of all genera). At the species level, endemism issignificant and lies around 50% and over, both inthe Frasnian and in the Famennian. Analyzed zoneby zone, the average rate of endemism is around40% in both stages. This rather high rate seems toreflect the wide spatial distance between easternand western parts of the Prototethys (Figures 1-2).Since investigations of Canning Basin faunas haveproceded much farther than in other regions, it can,however, be expected that some Australiansupposed endemics will eventually be discoveredelsewhere, as has happened in the past (see above:Praemeroceras petterae and Pan ticeras discoidale).Despite the high number of endemic species-leveltaxa it is not suggested that Canning Basin faunasbelong to a clearly defined eastern Gondwanaprovince but subprovince status (see Westermann2000; chorotype Canning Basin, chronotype =Famennian) may be granted. The low-leveldistinction is based on the fact that there are noendemic genera in the studied faunas which arecommon as well as very distinctive.

The comparison of the Canning Basin with twentyother regions (Table 2) gives a clear picture offaunal similarities that bears no easy correlationwith estimated distances between crustal blocks orwith their postulated spatial configuration. There is

392

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R.T. Beeker

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Figure 3 Maeneceras milleri (Flower and Caster), a typical "spot taxon" known from a single specimen in eastern NorthAmerica (3A-C, natural size, from Miller 1938: PI. 37, Figures 5-7) and abundantly from the late UD 11 of theCanning Basin (D-E; GSWA 109145/AI, "Syncline Gully", Bed K, Mt. Pierre area, natural size).

---------

Western Australian ammonoid palaeobiogeography

also no fundamental palaeobiogeographicdifference between the two Upper Devonian stages.The global Kellwasser Crisis at the end of theFrasnian had no lasting effect on principle fauna1exchange although there are some interesting trendsin detail.

The greatest faunal similarities during all of theinvestigated time-span are with the classical faunasfrom Germany. In the Frasnian there are 32common species (33%) and further 19 similar taxa,giving a "total similarity" of more than 50(10. In theFamennian there are 27 common species (39%) andthe wider similarity is close to 60% (41 taxa).Frasnian ammonoids show also clear relationships("total similarity" between 20 and 30%) with theTiman, North Africa, the Montagne Noire, theArdennes Shelf, SW England, and with EasternNorth America (mostly New York State). Theseregions all formed one larger realm within oradjacent to the western Prototethys. Close affinitieswith the Canning Basin emphasize that there wasno general biogeographic barrier, neither too deepoceanic nor too shallow, between eastern andwestern Prototethys regions plus the Uralian­Transarctic seaway. This important conclusion is insome conflict with the plate tectonic reconstructionsof Scotese and McKerrow (1991) which shows anextensive central Prototethys ocean that should atleast have produced a clearer separation of Russianfaunas. Migrations from and to Europe and NorthAfrica could have taken place along the equatorialnorthern margin of Gondwana but this iscontradicted by the absence of Frasnianintermediate faunas from Tibet, Pakistan,Afghanistan, Iran or Turkey and by the lack ofTimanites in the western Prototethys. In the Scoteseand McKerrow reconstruction, Beloeeras faunasfrom the Rudnyi Altai and Kusnetsk Basin alsowould be completely isolated. The Devonianreconstruction of Kent and Van der Voo (1990)shows an even wider ocean without intermediateplates between Australia and Europe and is anywaydismissed here because of the highly unlikely wideocean separating North Africa and Europe.

Crustal blocks positioned in the neighbourhoodof northwest Australia, especially former parts ofWestern Indonesia-Malaysia-Thailand, Vietnam­Laos-Cambodia, Burma, and Tibet have norecorded Upper Devonian ammonoid faunas at all;ammonoid facies never developed there or did notoccupy significant areas. Southeast Asian blocksseem to have had a different shelf-facies history. Ifplaced just west of Australia, as in thereconstruction of Metcalfe (1996) and to a lesserextent (W jNW) also in Scotese and McKerrow(1990), these plates (Figure 4) would have restrictedthe faunal exchange to the west more than evidentfrom the data. The reconstruction of Heckel andWitzke (1979) leaves more space between South

393

East Asian plates to account both for pantropicalgenepools of pelagic forms and for diverging outershelf facies differences.

Assemblages from South China (Hunan, Guangxi,Guizhou, and the Baoshan region of Yunnan) arealso rather rare and of low-diversity althoughtypical cephalopod facies were present in somedepressions between wide reefal platforms. Thepoor affinities between the Canning Basin andSoutheast Asian regions is again thought to reflectmostly differences in shelf facies rather than thepresence of biogeographical barriers. It is possiblethat detailed future work will produce richerFrasnian faunas from South China. A fewunpublished goniatites have been seen by theauthor in regional museum collections at Guilin andNanning.

The Famennian picture differs insofar thatammonoid faunal similarities with North Americaand the Timan were lost. This is not based ondifferences in faunal composition but, again, on thesimple fact that early to middle Famennianammonoids are almost lacking in the two regions.After the Upper Kellwasser Event, typicalammonoid biofacies did not re-develop for a longperiod. The influence of regional structural andfacies evolution was the dominant factor controllinga restricted ammonoid distribution, not increasedspatial distance or the establishment of migrationalbarriers. A reverse process can explain the increasedFamennian faunal similarity between the CanningBasin, the Urals (18 common species) and the HolyCross Mountains (21 common species). At the sametime poorly studied faunas appeared in regionslacking Frasnian goniatites completely: GreekAgais, North China (Great Khingan), Caucasus,Turkestan, and Kazakhstan. Transgressionproduced new regions hospitable for ammonoidsbut spreading events, such as the one associatedwith the global Annulata Event (Becker andKullmann 1996: Figure 4), did not reach all basins.

The ammonoid analyses seems to contradict tosome extent palaeogeographic results obtained fromthe study of Asian and Australian vertebrates. Richand Young (1996) observed that fish faunas of bothregions became more similar at some stage in theUpper Devonian and suggested a narrowing of themarine barriere between both crustal units, fittingthe Metcalfe (1996) reconstruction. Palaeomagneticdata (Zhao et al. 1996), however, indicate thatChinese blocks were pulling away from the westernpart of Australia as early as between the EarlySilurian and Middle Devonian. Furthermore,Johanson and Ritchie (1999) recently pointed outthat latest Devonian fish faunal similarities betweenSouth China and eastern Australia were not as closeas originally thought and during most of the UpperDevonian there were close affinities withEuramerica. Stratigraphical data in Rich and Young

394

Late Devonian (Famenrnan)

R.T. Becker

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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 CanningBasin (black dots) by an array of Southeast Asian crustaI blocks. WB = West Burma, S =Sibumasu (parts ofIndonesia, 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 theTransarctic and Kazakhstan distribution of Timanites and Be/oeeras (Figure 2).

(1996: Figure 2) show that faunal links betweeneastern Australia and China appeared late in theFamennian when they were no more ammonoids inWestern Australia.

"Intra-Australian" relationshipsIn Famennian II and IV, the Canning Basin and

New South Wales have not a single species incommon. Eastern Australian ammonoid sourceareas are part of terranes which were attached laterto the craton. There was certainly again a faciesinfluence on different faunal composition since

Canning Basin DD IV-assemblages (Piker Hills)were found in a rather unusual hemi-pelagicshallow black shale facies with many gastropods,brachiopods, and plant remains. New South Walesfaunas (Jenkins 1966, 1968) contain both endemicspecies (e.g., "Genuclymenia" keepitensis) and formsknown from Western Europe and North Africa(Erfoudites, Pia tyclymenia). Generally, western andeastern Australian Devonian ammonoid faunasrepresent completely disjunct basins andsubprovinces that both had some permanent faunalexchange with western parts of the Prototethys.

Western Australian ammonoid palaeobiogeography

Migration from Eastern Australia via theProtopacific (Panthalassia) is ruled out because ofthe lack of similar faunas in western NorthAmerica.

Level of endemism in taxonomic groupsAll Canning Basin species-level taxa were sorted

according to their family affinity. Representativesof the Anarcestidae (Archoceras),Praeglyphioceratidae (Lagowites),Kosmoclymeniidae (Protoxyclymenia) andCarinoclymeniidae (Karaclymenia) are so rare thatthey were not considered. The Phenacoceratidae(Cycloclymenia), Prolobitidae (with a new genus)and Posttornoceratidae are only represented bysingle endemic species. The high endemism withinthe Prolobitidae was probably related to theirdemersal (suprabenthonic) lifestyle in conjunctionwith a global sealevel lowstand (Becker 1993a,Becker and Kullmann 1996) in the VD III-C(Prolobites delphinus Zone). The same may apply tothe "spot genus" Cycloclymenia which waspreviously only known in VD II and VD V ofGermany. Posttornoceras glenisteri (Petersen) is veryclose to P. contiguum (Miinster) from Germany. Allsix species of the Koenenitidae are restricted tonorthwestern Australia. These forms compose earlyFrasnian very low diverse assemblages which wereobviously more isolated in shallow hypoxicinterreefal basins than faunas from later in theFrasnian when the sealevel was much higher.Timanites angustus (Glenister) is very close to theRussian T. keyserlingi (Miller) and to the northwestCanadian T. occidentalis (Miller and Warren).

High numbers of endemic species (between 50and 70%) can be found in the Tornoceratidae,Pseudoclymeniidae, Dimeroceratidae,Acanthoclymeniidae and in the Beloceratidae.These groups represent a wide range ofmorphotypes that cannot be assigned to a singletype of lifestyle. The Famennian tornoceratidendemism is rather special since the majority ofspecies is regarded as typical pelagic forms of theopen sea (Becker 1995).

Lower numbers (between 30 and 50%) of endemicspecies are found in early FamennianCheiloceratidae, in the Frasnian Gephuroceratidae,and in the regionally low-diverse middleFamennian Platyclymeniidae. The dominance of theCheiloceras (Puncticeras) postinversum-semiinversumGroup, however, gives a unique structure tocheiloceratid assemblages. The highest degree ofcosmopolitanism IS apparent In theSporadoceratidae (see Teichert 1943 and Petersen1975) and in the Cyrtoclymeniidae. Again, there isno easy correlation between morphology, assumedlifestyle and wide distribution. Dispersal or geneticisolation was more influenced by episodes of facieschanges and eustatic movements than by general

395

types of morphology and paleoecology. Only amore general study might clarify whetherrepresentatives of certain morphotypes tend to bemore endemic or cosmopolitan than others.

Outer shelf synecology and palaeobiogeographyIn Canning Basin marginal slope and intereefal

environments, ammonoids are frequently associatedwith a range of other fossil groups, especially withconodonts, nautiloids, rhynchonellids, trilobites,crinoids, bivalves, gastropods, solitary rugosecorals, sponges, and fish remains. Only some ofthese groups have been studied sufficiently in detailto allow palaeobiogeographic comparisons. Itremains an interesting question which parts ofburied assemblages display similar or differentpatterns of endemism. Almost nothing is known atpresent about changes in Devonian "pelagiccommunities" with distance or latitude. Forexample, the complete lack of trilobites in pelagicassemblages of the Timan and of Eastern NorthAmerica is well-known but not understood.Klapper (1995) compared Frasnian conodonts andfound high levels of endemism (up to 1/3) at thespecies level; this group was previously regardedas cosmopolitan at that time. Even higherendemism seems to apply to many Canning Basinbenthic groups of the Gogo and Virgin HillsFormations: Frasnian and Famennian crinoids Gelland Jell 1999), sponges (Rigby 1986) andrhynchonellids. By contrast, buchiolinid bivalves(Grimm 1998) show close relationships withEuropean taxa. This suggests an interestingdifference in the spatial gene flow between faunalgroups living in the same environment. But,anyway, a range of faunal groups can be used todefine a biogeographic Canning Basin Subprovincein the Upper Devonian.

Regional diversity analysis

Canning Basin extinction and diversification phasesThe Canning Basin taxonomic diversity is

higher in the Frasnian (36 general subgenera with99 species/subspecies) than in the Famennian II­IV (32 genera with 69 species/subspecies). Theaverage diversity of Frasnian zones lies at ca. 9.5species but only at ca. 7 species in Famennianzones. These figures are based on realoccurrences and do not include regional "Lazarusepisodes" (record gaps) of taxa. These often donot reflect sampling gaps of rare taxa but episodicand facies-controlled absence of taxa from thestudied sections. Bactritids have been omittedsince their record is strongly biased bypreservational aspects (easy species identificationis only possible in haematitic faunas). Generally,faunas from individual levels of the CanningBasin inter-reefal deposits are not as diverse as

396

species

15

10

5

BUGLE GAPEVENT

LKWEVENT

R.T. Becker

UKWEVENT

ANNULATAEVENT

CONDROZEVENTS

B c E F G

UPPER DEVONIAN I

FRASNIAN

K GAB

UPPER DEVONIAN 11 UO III

FAMENNIAN

CAB

UDIV

Figure 5 Species level total extinctions in the Upper Devonian of the Canning Basin showing three major Frasnianextinctions (Bugle Gap, Lower Kellwasser and Upper Kellwasser Events) and some Famennian phases ofhigh faunal overturn (Condroz Events, UD II-F IG, extinction subsequent to the Annulata Event).

those from German or Polish submarineseamount sections positioned in deeper parts ofshelf margin basins. Several diversity peaksclearly correlate with transgressive episodes, butnot all eustatic rises (see Becker and House 1997)allowed diversification or the spread of morespecies into the Canning Basin. This indicates asignificant influence of regional structuralevolution. The maximum diversity with 31species/subspecies was reached in theVirginoceras erraticum Zone (VD I-J

2). Plots of total

species level diversity and of total extinctions(Figures 5-6) give the following stages ofdiversity development (for event sequence andterminology see Becker et al. 1993 and Becker andHouse 1997):a) Early Frasnian (VD I-B/C) immigration and

speciation of low-diverse faunas, probablycorresponding with the international

transgressive episodes associated with theGenundewa and Timan Events.

b) Small scale extinction at the end of VD I-Ccorresponding with the global Koenenitidaedecline near the end of MN Zone 4 (= transitansZone).

c) Important regional diversification, with a peakin VD I-F

2(Gogoceras nicolli Zone; ca. MN Zone

6 = upper part of punctata Zone), correlatingwith the regional main development of hypoxicgoniatite shales.

d) Bugle Gap Event (named herewith): Significantregional extinction associated with a retreat ofhypoxic facies and regression at the end of VDI-F

2(within MN Zone 6) in the Bugle Gap area.

This event may be an important andcharacteristic regional signature of the CanningBasin.

Western Australian ammonoid palaeobiogeography 397

cosmopolitan/

I endemiC

~

openes

BUGLE GAPEVENT

SEMICH.EVENT

LKW• EVENT

PRESERVED SPECIES LEVELTOTAL DIVERSITYCANNING BASIN, W. AUSTRALIA

NEHDENRADIATION

CONDROZEVENTS

UKWEVENT

ANN.EVENT

B C o El G H I KI

AIBCDE FIG HI AB CAB

UPPER DEVONIAN I

FRASNIANUPPER DEVONIAN 11 UD III

FAMENNIANUDIV

Figure 6 Preserved species level total diversity in the Upper Devonian of the Canning Basin and the total number ofendemic species. Diversifications occurred in the early middle Frasnian (UD I-ElF), with the semichatovaeTransgression, in the upper Frasnian, lower Famennian (Nehden Radiation), early in UD 1II and with theglobal Annlllata Event.

e) Stable and moderately low diversity in UD I-C/H (MN Zones 7-10, = ca. hassi Zones).

f) Important wave of immigration and speciationat the base of UD I-I corresponding with theinternational "semichatovae Transgression"(early MN Zone 11, jamieae Zone to lowestpart of Early rhenana Zone).

g) Stable high diversity in UD 1-12

and I-J l (MNZones 11-12, = Early rhellalla Zone), inagreement with the globally high sealevel.

h) Maximum regional diversity (31 species/subspecies) in the Virgilloceras erraticllm Zone(UD I-J

2, upper part of Zone 12, upper part of

Early rhellalla Zone), followed by a massiveextinction (27 species) during a significantregressive phase (top of MN Zone 1 Itcorresponds in time with the global extinctionjust prior to the Lower Kellwasser level.

i) Very low diversity and very rare faunas in thelower part of MN Zone 13 (UD I-K, = lowerpart of Late rhellalla Zone). The regional lack offaunas from this interval is intriguing but maypartly reflect undersampling.

j) Diversification in the latest Frasnian Crickiteslindneri Zone (UD I-Lln, upper part of MN Zone13, = upper part of Late rhenana Zone) which ischaracterized by several regional transgressivepulses.

k) Upper Kellwasser Event: A regional decline ofammonoid faunas started at the base of thelingllifomlis Zone (= topmost part of MN Zone13, UD I-Llb)' characterized by a short-termedregression with a spread of neritic faunas (e.g.,diverse gastropods). The main extinction liesabove a subsequent transgressive level(Manticoceras gllppyi Bed) within the linglliformisZone (see Becker et a/. 1991, Becker and House1997) which correlates with the UpperKellwasser level but which is not hypoxic. Thelast Mallticoceras range up to the last centimeterof the Frasnian. The regional extinction patternis more gradual than in Europe and NorthAfrica and this may be related to the completelack of hypoxic Kellwasser facies. The finalsudden extinction of manticoceratids must havebeen caused by factors such as temperature and

%endemics

100

R.T. Becker

.'

NEHDENRADIATION

marginifera Zone) and in the lower part of VDIll-A. The low diversity of VD-G

2and VD III­

A3

reflects inadequate sampling. The globaland regional transgression at the base of VDII-G (classical do liB, Maeneceras subvaricatumZone) is marked by the immigration of theoldest sporadoceratids but not by adiversification.

p) The immigration of pseudoclymenids took placeduring a minor diversification phase (VD Ill-B)and their sudden extinction (within the Lowertrachytera Zone) represents a highly interestingsmall-scale global extinction event that is notreflected by a change in lithofacies.

q) The immigration of the oldest clymenids (baseof VD III-C, ca. base of Vpper trachytera Zone)and prolobitids produced another minordiversity increase but faunas remained of verylow diversity in comparison with those from theVrals, Poland, and Germany. At the end of theregional Protactoclymenia euryomphala Zonethere was a small-scale extinction, correlating

:"

LKWEVENT

SEMICH.EVENT

BUGLE GAPEVENT

B

10

30 •

20

numberat species

Figure 7 Canning Basin relationships between preserved total species-level diversity and the relative (percentage) rateof endemism (interrupted lines), Low-diverse/highly endemic faunas alternate with species-rich/morecosmopolitan faunas, However, there is also a smaller-scale positive correlation between changes in diversityand endemism.

salinity that are not very obvious in thesedimentary record.

1) Earliest Famennian beds (VD II-A/B, Pa.triangularis to Middle crepida Zones), spanningca. 2 ma, have no ammonoids at all. Regionally,there were no ammonoid survivors at all.

m) The rapid Nehden Adaptive Radiation (VD 1I­CID, Vpper crepida Zone to lowermostrhomboidea Zone) produced the second regionaldiversity peak, which correlates well with aglobal and regional sealevel high.

n) The Lower Condroz Event at the end of VD lI­D (Oxytornoceras n.sp. Zone) led to a significantextinction which, however, was balanced bynew forms entering in VD I-E (Praemeroceraspetterae Zone, middle part of rhomboidea Zone).

0) Diversity remained moderately high duringthe higher parts of VD 11 and in the lowerpart of VD III (upper part of rhomboidea Zoneto Vppermost marginifera Zone), but therewere several faunal overturns, for examplebetween VD I-F and I-G (within basal

398

Western Australian ammonoid palaeobiogeography

with a basin-wide regression and discontinuityin the ammonoid record.

r) Annulata-Event: The global hypoxic andtransgressive Annulata Event (Upper trachyteraZone) allowed only a minor diversification inthe haematitic Piker Hills Formation butenabled the spread of ammonoids to New SouthWales (see Jenkins 1968). Regionally, the post­event major regression caused a drastic declineof ammonoid faunas from which the group didnot recover in the Middle Paleozoic.

s) The final extinction of the last and very rareCanning Basin clymenids (Protoxyclymenia,Falciclymenia) took place in the upper part ofUD IV, probably still within the postera Zonebut precise conodont data are not available fromthe shallow-water carbonates.

Palaeobiogeography and regional diversityLow diversity faunas have a relative high

percentage (50% or more) of endemic forms andseem to contain specialists that were adapted tolocal/regional environmental conditions whichwere not favourable for many other species.This applies to early Frasnian (UD I-B/C) andsome Famennian (UD II/IlI) faunas. Relativelylow rates of endemism « 20% of taxa) werefound during some but not all transgressiveepisodes. Examples are the basal Rhinestreeteustatic pulse (Naplesites housei Zone, UD I-G l ),

the "semichatovae Transgression" (UD I-J l ), thelatest Frasnian diversification (Crickites lindneriZone, UD I-L

l.) and the two eustatic rises

during the marginifera Zone (UD II-F2 and II­G

l). It is suggested that differences in

ammonoid diversity and endemism duringindividual transgressive phases reflect varyingoverprints of the basin subsidence history andby associated regional ecological developments.

The relationships between total (preserved)diversity and the relative rate of endemism (%of endemic species in each zone) are depicted inFigure 7. The alternation of low-diverse/highly­endemic and diverse/weakly-endemicassemblages can be seen with more endemicphases in the early Frasnian and in the late lowerto middle Famennian. In addition, however, thesecond order peaks and lows show paralleltrends suggesting that during eachdiversification there was also an incoming ofendemic taxa, probably from elsewhere. This issupported by the fact that not only cosmopolitanbut also endemic forms show record gaps(regional "Lazarus phases"). A good proportionof endemic species is also of cryptic origin;direct ancestors are not known from the region.Many of the Canning Basin forms must havelived outside the studied sections, at leastepisodically.

399

ConclusionsThe analysis of Canning Basin Upper Devonian

ammonoid faunas allows a range of conclusionsthat are of importance for the generalunderstanding of palaeobiogeographic relationshipsin Middle Palaeozoic pelagic environments and fora better placing of northwest Australia in thecomplex framework of Upper Devonian crustalplates:1) Canning Basin ammonoid faunas consist to a

varying degree of (semi-)cosmopolitan(pantropical), endemic and "spot taxa". Thelatter emphasize the incompleteness of theknown fossil record and are important markersfor biogeographic links.

2) Endemism was low at the generic but verysignificant at the species level. This seems toreflect the wide spatial distance betweenpopulations of the eastern and western parts ofthe tropical Prototethys. Pantropical genepoolsexisted contemporaneously only in some butnot in all ammonoid groups. Canning Basinfaunas belong to a small-sized subprovince ofthe Prototethys Province.

3) Closest similarities existed with Germany, andto a lesser degree with other European-NorthAfrican-Appalachian regions. The TransarcticRoute was only episodically significant for thedispersal of some taxa such as Timanites andPraemeroceras.

4) The individual structural evolution of basins(crustal blocks) determining the spread of outershelf ammonoid biofacies and not theestablishment or dismiss of biogeographicbarriers was the major controlling factor ofUpper Devonian ammonoid distributionpatterns. Asian plates adjacent to northwesternAustralia mostly lacked suitable facies. Anequal or greater importance of biofacies ratherthan biogeographic factors was also suggestedfor conodonts by Klapper (1995).

5) The lack of a general biogeographic barrierbetween northwest Australia and the westernPrototethys and Urals regions contradicts theplate tectonic reconstructions of Scotese andMcKerrow (1990) and Metcalfe (1996) but ismostly in agreement with that of Heckel andWitzke (1979).

6) There is no clear correlation between the rate ofendemism and morphology or higher leveltaxonomy (families). The different spreadingsuccess of various groups has to be interpretedin the light of the changing global sealevel. Itcould be argued that a Canning Basin faunalprovince was established and destructed in afine sequence of episodes with fluctuatingisolation.

7) The Upper Kellwasser event near the Frasnian-

400

Famennian boundary interrupted the CanningBasin ammonoid record totally for ca. 2 ma, butcaused no fundamental biogeographical change.

8) All known global extinctions of the Frasnian tomiddle Famennian, even small-scale eventssuch as the koenenitid and pseudoclymenidextinctions, can be recognized in the CanningBasin. Magnitudes and phases, however, werestrongly influenced by regional factors. TheBugle Gap Event at the end of VD I-E

2(upper

part of punctata Zone) was a significant regionalextinction.

9) Analyzed zone by zone, there was analternation of low diverse/endemic and highlydiverse/cosmopolitan faunas. "Lazarus Phases"of endemic forms and a small-scale positivecorrelation between diversity trends and therelative rate of endemism suggest that manyforms may episodically have lived outside thestudied sections. True endemism was probablysomewhat lower than is evident from currentdata.

ACKNOWLEDGEMENTS

This study would have been impossible withoutall the support and co-operation of M.R House(Southampton). He as well as G. Klapper and K.McNamara (Perth) read and corrected themanuscript. Field work in the Canning Basin wasfinanced by grants of NERC (to M.R. House) and bythe DFG. It was conducted jointly with W.T.Kirchgasser (Potsdam, New York), G. Klapper(Iowa City), P.E. Playford (Perth), RT. Brown(Canberra), R Feist (Montpellier), R Crick(Arlington) and others. The Geological Survey ofWestern Australia generously assisted in providingvehicles, equipment and other support that enabledextensive fieldwork.

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Manuscript received March 1999; accepted October 1999.