Records of the Western AlIstralian MlIsellm Supplement No. 58: 321-334 (2000).

Palaeobiogeographic affinities of Late brachiopods from Iran

Glenn A. Brockl and Mehdi YazdF

1 Centre for Ecostratigraphy and Palaeobiology, Department of Earth and Planetary Sciences, Macquarie University, Australia 2109 2 Department of Geology, Faculty of Science, University of Esfahan, Esfahan 81744, Islamic Republic of Iran

Abstract -A total of 34 Late Devonian brachiopod genera from north (Alborz Mountains), east (Tabas region) and central-southeast Iran were compared, using the Jaccard similarity index, with synchronous faunas from 28 localities spread across Gondwana, Euramerica, Siberia, Kazakhstan, and South China. The highest levels of similarity occur between brachiopod taxa from the three Iranian localities and terranes traditionally regarded as part of the northern margin of Gondwana (especially Afghanistan, Armenia, NW Pakistan, Libya, Cantabrian Mountains and Iberia). The distribution of benthic shelf faunas was probably strongly influenced by climatic (latitudinal) continuity, geographic proximity and open dispersal routes along the "North Gondwana Current". Jaccard analysis also reveals some biogeographic links between IranlAfghanistan and Euramerica throughout the Late Devonian, supporting the hypothesis that a narrow « 1000 km) Prototethys Ocean separated the two super continents during this interval.

INTRODUCTION and northern Gondwana. This is based on the fact There is general, but not universal agreement (cf. that modem oceans act as formidable barriers to Soffel and Forster 1984), that the Iranian dispersal by shallow marine shelf benthic microcontinent(s), comprising central-east Iran and invertebrates, especially those organisms (the vast the Alborz Mountains was located near Saudi majority) that produce nonplanktotrophic larvae Arabia along the norLl-tern margin of Gondwana with relatively short lifespans « 4 weeks), such as with numerous other 'Asian' and 'European' articulate brachiopods (Talent 1985; Talent et al. terranes (possibly forming an extensive 1986; Gratsianova et al. 1988; Richardson 1997). "Cimmerian belt" of terranes according to Sengor The aim of this study is to probe the and Hsu 1984) during the Devonian (Berberian and biogeographic affinities of Late Devonian King 1981; Wensink 1983; Lensch et al. 1984; Talent brachiopods from Iran by separately comparing et al. 1986; Belov 1986; Husseini 1991; Dastanpour and faunas with a selection of 1996). Marine transgressions during the Late synchronous faunas from Gondwana, Euramerica, Devonian-Early deposited thick Siberia, Kazakhstan and South China in an attempt carbonate dominated sequences, replete with to; 1) ascertain whether the faunas conform to diverse shallow marine faunaI assemblages, in patterns of 'cosmopolitanism' previously identified north, east and centraliSE Iran, consistent with the for the Late Devonian and, 2) constrain the position view that Iran (Figure 1), along with much of of the Iranian plate with respect to other northern Gondwana, was positioned within the Gondwanan and Euramerican terranes and test the carbonate development zone, between 0° and 30° validity of the claim that these two supercontinents (Witzke and Heckel 1988; Dickens 1993) in the were widely separated during the Frasnian­ southern hemisphere (Eldridge et al. 1997; Figures Famennian. 4-5). Some palaeogeographic reconstructions of the Late Devonian, especially those based on palaeomagnetic data, indicate that the northern BIOSTRATIGRAPHY OF LATE DEVONIAN margin of Gondwana was separated from the BRACHIOPODS FROM IRAN supercontinent of Euramerica, to the north, by a Biostratigraphy of Devonian faunas from Iran has 4000+ km ocean basin, variously referred to as been recently reviewed by Dastanpour (1996), with Prototethys or the Prehercynian Ocean (Morel and additional refinements presented by various hving 1978; Van der Voo 1988; Bachtadase et al. authors at the recent IGCP421 meeting held in 1995; Metcalfe 1996, figure 13). If this were true, Esfahan, Iran (e.g. Jafarian 1998; Brice et al. 1999). one could expect sharp provincial boundaries Late Devonian brachiopods from Iran are known between coeval benthic faunas from Euramerica from thick carbonate dominated sequences from 322 G.A. Brock, M. Yazdi

Khoshyielagh Formation and these have been ....=_=...5OOkm documented by Vandercammen (1967), Brice (1971), Brice et al. (1974, 1978) and Sartenaer and Plodowski (1975). The faunas are relatively diverse (Table 1) and broadly correlate with brachiopod 35' ; zones 6-12 (Frasnian-Famennian) of Brice (1977). r' ( The brachiopod fauna from the Geirud Formation \ '-., (sensu Stepanov 1967) in the central Alborz Range " '\ (Figure 1) was documented by Gaetani (1965) with IRAQ i revisions of some taxa provided by Sartenaer (1966a). The Geirud Formation is up to 340 m thick and consists of fossiliferous calcarenite, marly limestone, shales and lithic sandstones. Diagnostic brachiopods from limestones near the base of the sequence include Ripidiorhynchus elburzensis and Cyrtospirifer cf. syringothyriformis equivalent to biozone 8 of Brice (1977) and indicative of an upper Frasnian age. The overlying Centrorhynchus Figure 1 Map showing location of the three principal deltidialis zone of Gaetani (1965) with C. deltidialis, Late Devonian brachiopod bearing regions in Gastrodetoechia iranica and Cyrtospirifer verneuili has Iran (modified after Hamedi et al. 1997). no direct biozone equivalent using Brice's (1977) scheme, but Brice et al. (1978: 22) indicate the C. deltidialis zone approximates a lower Famennian three main regions (Figure 1). These faunas can be age. Centrorhynchus is a characteristic Famennian directly correlated with brachiopod biozones 6-12 taxon (Sartenaer 1970). erected by Brice (1977) for Late Devonian Dastanpour (1996) reported the presence of Frasnian brachiopod faunas from Afghanistan. According to (Ripidiorhynchus) and unnamed Famennian Brice's (1977) scheme brachiopod zones 6-8 are brachiopods from limestones outcropping in the assigned a Frasnian age and zones 9-12 (part) are vicinity of Zanjan in the western Alborz Range (Figure Famennian in age. 1), but these have not been formally described and are not included in the current analysis. Northern Iran (Alborz Mountains and Robat-e­ Gharabil) Eastern Iran The Khoshyielagh Formation consists of up to The Ozbak-Kuh Group, a 1500-1700 m thick 1800 m of interbedded limestones, shales and sequence of Devonian-Carboniferous sediments, sandstones outcropping extensively in the eastern outcrops extensively in the Tabas area in the Ozbak­ Alborz Range (Dastanpour 1996) and the Robat-e­ Kuh Mountains (Ruttner et al. 1991), the Shotori Gharabil region in northeastern Iran (Brice et al. Range (Stocklin et al. 1965), and at Shirgesht 1974). The age of the base of the formation is in (Ruttner et al. 1991). The Ozbak-Kuh Group is dispute. Brice et al. (1978) subdivided the formation subdivided into four formations, with the principal into seven members and indicated the basal 560 m Upper Devonian sequences being the Bahram and (members 1-4) could be assigned an ­ Shishtu formations. Detailed conodont age based on brachiopods, bryozoans, biostratigraphy presented by Weddige (1984) for conodonts and corals. The presence of conodont the Ozbak-Kuh area and by Yazdi (1996, 1997,1998, species such as Icriodus aff. obliquimarginatus, 1999) for exposures in the Shotori Range indicate Polygnathus pseudofoliatus, P. cf. timorensis, P. aff. the Bahram Formation ranges in age from Givetian­ xylus and P. linguiformis linguiformis provides strong Frasnian and, according to the latest work of Yazdi evidence for a Middle Devonian age (Bultynck in (1999), the conformably overlying Shishtu Brice et al. 1978). A possible age for Formation spans an interval from Frasnian (Early lowermost strata of the Khoshyielagh Formation hassi Zone or older) to latest (anchoralis­ has been suggested by Hamdi and Janvier (1981) latus Zone). Brachiopods are known to occur in the based on conodont and vertebrate occurrences. Bahram and Shishtu formations from all three However, Ghavidel-Syooki (1994, 1998) argues that regions, but only a few taxa have been formally the entire Khoshyielagh Formation is Late Devonian described. in age based on acritarch and spore evidence, and Sartenaer (1966b) described Cyphoterorhynchus that "Lower and Middle Devonian strata are not arpaensis, C. koraghensis and C. k. interpositus from present on the Iranian platform" (Ghavidel-Syooki the Bahram and Shishtu formations at Ozbak-Kuh. 1998: 12). Late Devonian brachiopods have been C. arpaensis and C. k. interpositus also occur in the recovered from strata 600 m above the base of the basal Shishtu Formation at Howz-e-Dorah (Yazdi Palaeobiogeography of Late Devonian brachiopods from Iran 323

Gaatani 1965; sonooo", 1966b; Brloo ot aI 1973, 1974, Von

Sarten.aef 1966a, 1968b; Legrand-Blain 1998

Ojafartan & BrI"" 1973; L99ramHllaln 1998, Brloo 1998; Brloo et al. in pro..

Durl

Abromlan 1957 & 1974; Mamb«OY& Rlhon.1985

sartana9l1964; Morante 1989

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Sartenaor 1975; Legrand-Blaln 1995

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Cooll EO III 1986

Sanonaor 19660; Brlco & MealS 1971; Brlco et 011976; Bnoo 1988

Sartenaer 1966a, 1968b; Vandercan"V'rl8n 1956,1969

x & Krath 1974; Plo

Balnskl 1979, 1995; B!omat 1971

Kala 1900; Malyla & Zbiko_a1974

Reed 1943, Butler 1981

Martyoova 1988; Lltv1no~ch OIa11975; Nasikanova 1975; Sldiachenko 1961

Numerous sources

Choo Yuan..f\en 1984: Sartenaer 1985: Tan 1987: Hanca et al. 1994

Tachlbana 1981

Oay 1988. 1995

Carter & Kam""" 1990

sartena'" & Sandberg 1974

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Sartenaor 1969,1975, Raasch 1988

V99vers 1959: Roberts 1971

Maxwelt 1953, 1954; McKolIar 1970 324 G.A. Brock, M. Yazdi

1996; pers. observ.). Conodonts indicate a Frasnian Frasnian taxa include the rhynchonellides age (older than Early hassi Zone) for this horizon. Ripidiorhynchus and Ladogilina and the spiriferids The presence of C. arpaensis and C. koraghensis Cyrtospirifer and Rigauxia, whilst Famennian faunas equate with brachiopod zone 6 of Brice (1977) include Sulcatospirifer, Cyrtiorina and the productids indicative of an early-mid Frasnian age. Sartenaer Sinoproductella (providing a link with South China) (1966b) also described Coeloterorhynchus tabasensis and Nigerinoplica. from the Shishtu Formation in the Kale Sardar area. Brice (1977) indicated C. tabasensis was diagnostic of the succeeding Frasnian brachiopod zone 7. MATERIALS AND METHODS Description of Famennian brachiopod taxa from Talent et al. (1986) and Gratsianova et al. (1988) eastern Iran are limited to Stenaulacorhynchus highlighted some of the problems associated with cheshmehshirensis and Pampoecilorhynchus arianus attaining a consistent taxonomic database for from lower Famennian exposures (= zone 9 of Brice palaeobiogeographic studies using Devonian 1977) of the Shishtu Formation in the Ozbak-Kuh brachiopods. Some of the problems include area (Sartenaer 1968a, b) and poorly preserved differences in taxonomic style (lumping vs. splitting Sentosia sp. from late Famennian horizons of the Shishtu Formation in the Shotori Range (Legrand­ Blain 1998). The presence of Pampoecilorhynchus s... arianus in the Khoshyielagh Formation, northern j!3 Iran (Brice et al. 1978) and Robat-e-Pal, Afghanistan ::E FRASNIAN t:! (Brice 1971) provides a level of regional correlation 0 BRACHIOPOD DATA .0 for lower Famennian sequences (Brice 1977). ~ ~ z Jaccard Coefficient z w < Sartenaer (1985: Figure 1) indicated Pampoecilo­ < l- (% similarity) Q; z ~ en rhynchus and Stenaulacorhynchus are restricted to the < Z :x:: Q;

• Ripidiorhynchus + Cyphoterorhynchus o Coeloterorhynchus b. Uchtospirifer @ Spinulicosta @ Whidbomella - \s _._. (

(> V ANTARCTICA () AFRICA~~

S.AMERICA

Figure 4 Global distribution of selected 'core' brachiopod genera during the Frasnian. SIB = Siberia, KAZ = Kazakhstan, SA = Saudi Arabia, SC = South China, NC = North China, ADS = Australia, NGC = "North Gondwana Current". Mercator base map modified from Eldridge et al. (1997).

(Dickins 1993). The majority of faunas included in situated on crustal blocks associated with North the analysis also display comparable levels of Gondwana, Euramerica, Siberia, Kazakhstan, preservation (mainly original shell and/or casts or South China and Australia (Table 1, Figures 2-5) moulds). using the Jaccard Similarity Coefficient. As noted Described Frasnian faunas from Iran are by Talent and Mawson (1979: 90) discussions "of characterised by the rhynchonellid genera the deficiencies of similarity/ dissimilarity Coeloterorhynchus, Cyphoterorhynchus and coefficients used in palaeobiogeography have Ripidiorhynchus, the productids Spinulicosta, become almost routine". Detailed review of the Thomasaria and Whidbornella, and the spiriferid advantages and disadvantages of the numerous Uchtospirijer (Figures 2, 4; Table 1). Famennian statistical equations used in palaeobiogeographic faunas are moderately more diverse and are studies is beyond the scope of this paper. For characterised by taxa such as rhynchonellid genera review and discussion of the various quantitative Centrorhynchus, Megalopterorhynchus, Leptocaryo­ methods employed in palaeobiogeographic rhynchus, Pampoecilorhyncus, Stenaulacorhynchus, the investigations see Raup and Crick (1979), Talent spiriferids Dichospirijer, Enchondrospirijer, and Mawson (1979), Shi (1993, 1995). The Jaccard Sulcatospirifer, Eobrachythyris, and the productids Coefficient, one of the more robust binary Ericiatia, and Sentosia (Figures 3, 5; Table 1). measures of similarity (Shi 1993), has been chosen The 34 "core" Frasnian and Famennian genera as the main statistical measure of similarity for identified from the three main brachiopod bearing palaeobiogeographical studies associated with regions in Iran (Figure 1) were compared to IGCP 421. To facilitate future comparison between described faunas from 28 separate localities different invertebrate groups the brachiopod Palaeobiogeography of Late Devonian brachiopods from Iran 327 • • t::. SIB @ + @

(> LJ () AFRICA~~

S.AMERICA

Figure 5 Global distribution of selected 'core' brachiopod genera during the Famennian. For abbreviations see Figure 4. Mercator base map modified from Eldridge et al. (1997).

dataset was analysed using the Jaccard Coefficient Alborz Mountains display greatest similarity with (Je), those from East Iran (40%) and central-southeast JC =c I a + b - ex 100 Iran (45%) (Figure 2). The degree of similarity is not unexpected given that a widespread transgressive Where a is the number of genera from locality A, episode at the beginning of the Frasnian resulted in b is the number of genera from locality B, and c is the development of an extensive shallow water the number of genera shared by localities A and B. carbonate platform that covered much of central The equation is multiplied by 100 to provide a Iran from the northeast Alborz to the Faraghan percentage similarity. Future analyses will focus on Mountains, south of Kerman (Dastanpour 1996: species comparisons (once faunas are adequately figure 6). Despite the fact that each of the Iranian described) using binary and probabilistic methods. localities has a relatively small total number of Cosmopolitan taxa, such as Cyrtospirifer, Productella, genera for analysis (Figure 2) the Jaccard Coefficient Schizophoria and many of the Frasnian atrypids with still reflects a moderate level of generic overlap. little or no biogeographic signature were omitted Indeed, each of the localities has a significant from the analysis. number of species in common (Gaetani 1965; Djafarian and Brice 1973; Brice et al. 1978). Moderate RESULTS levels of similarity occur with regions known to have been associated with the northern margin of Frasnian brachiopod indicies (Figure 2) Gondwana, including Afghanistan (26'i"o), Armenia (25%), and NW Pakistan (24%), based mainly on Alborz Range the co-occurrence of Coeloterorhynchus, Cyphotero­ During the Frasnian, brachiopod faunas from the rhynchus and the more cosmopolitan Spinatnjpina 328 G.A. Brock, M. Yazdi

(Table 1). Though the levels of similarity are Range also show linkages to regions such as Poland relatively low, there is also some evidence of links (up to 19%), Japan (13%) Idaho (10%) and Australia with Euramerica, including England (18%), (up to 19%). Boulonais, France (15%), Belgium (14%), and New Mexico (15%). Links with Kazakhstan (14%) are also East Iran recorded. Brachiopod diversity is relatively low in Famennian sequences from East Iran, with only East Iran Pampoecilorhynchus, Stenaulacorhynchus, and Sentosia Taxa from East Iran show greatest similarity with formally described (Sartenaer 1966b, 1968a, b; central/SE Iran (67%), northern Iran (40%), Legrand-Blain 1998). Stenaulacorhynchus is only Afghanistan (29%) and Armenia (40%). There is also known from East Iran (Sartenaer 1985) and Seville, some weak links with other regions from north Spain (Weyant et al. 1988). Other cosmopolitan taxa Gondwana, including NW Pakistan (15%) Libya such as Cyrtospirifer, "Productella", and Cyrtiopsis (16%), Morocco (14%) and southern Spain (18%). have been reported from East Iran (see Ruttner et al. Links with Euramerica include Boulonnais, France 1991), but these require formal systematic treatment (14%), England (17%), New Mexico (10%) and (Brock et al. in prep.). The lack of formal taxonomy western Canada (11%). from this region precludes meaningful comparison with other regions. Central-southeast Iran Apart from moderate similarity with the Alborz Central-southeast Iran Range and east Iran, the greatest similarity for Strongest links are with Afghanistan (48%), the central/SE Iran occurs with Afghanistan (43%), Cantabrian Mountains (29%), and Iberian Peninsula Armenia (23%), Morocco (20%), and with the (22%) along the periphery of Gondwana. Compared Euramerican localities Boulonnais, France (23%), with the Alborz faunas, faunal links between England (29%), South China (20%), New Mexico central/SE Iran and Euramerica are not as strong. (13%) and western Canada (11%). The similarity The highest levels of similarity are with Poland index for England is artificially inflated because (16%), Belgium (15%) and western Canada (19%). there are two shared genera from a total of only 6 genera. Description of diverse faunas from the Late Devonian of Kerman (Brice, 1999) not included in DISCUSSION the present analysis will undoubtedly decrease this The Late Devonian is acknowledged as a time index. Unlike the faunas from the Alborz Range and interval represented by high levels of faunal and east Iran, the faunas from central/SE Iran show a floral cosmopolitanism, in both the marine and higher level of similarity with South China (20%), terrestrial realms (Boucot et al. 1969; Oliver 1977; but this is mainly based on the shared occurrence of Talent 1984; Young 1987; Boucot 1988). The the widely distributed taxa Ripidiorhynchus, relatively low levels of faunal and floral Aulacella and Spinatrypina. provinciality during the Late Devonian is generally considered to be the result of continental or terrane Famennian brachiopod indicies (Figure 3) amalgamation, favourable oceanic circulation patterns, and warm climatic conditions (Dickins Alborz Range 1993) that permitted interchange of once isolated Famennian faunas from the Alborz Range display faunas. On a global scale, the distribution of Late significant levels of similarity with central-southeast Devonian brachiopods (at least at generic level) Iran (56%), but only one genus (Pampoecilorhynchus) tends to fit this pattern of weak provinciality is shared with East Iran. This is mainly due to the (Boucot 1988). However, comparison of Frasnian fact that Famennian faunas from East Iran, though (Figure 2) and Famennian (Figure 3) data indicates not diverse, have not been formally described. The that Frasnian brachiopods from Iran appear to be presence of Sentosia (Legrand-Blain 1998) and other more provincial than Famennian faunas. This is in undescribed genera from sections in the Shotori accord with more detailed comparisons, such as Range (Brock et al. in prep.) indicates that future those reported by Talent et al. (1986: 106), for Late investigations will probably boost the degree of Devonian brachiopod species from Western similarity between these regions. Within Australia, central and southwestern Kazakhstan Gondwana, the Alborz faunas display highest and South China which reveals such low levels of similarity with Cantabrian Mountains (50%), Libya species similarity, that by modern standards, these (21 %), Bergischen Landes (24%), and Iberia (20%). regions would be placed in separate zoogeographic Again, though the biogeographic signal is weak, provinces and regions. Talent and Mawson (1979: there are some links with Euramerica, especially 4) also documented strong "prOVincial with the Franco-Belgium Basin (21%) and Western heterogeneity" of Frasnian brachiopod faunas from Canada (23%). Famennian taxa from the Alborz Pakistan, Siberia, Kazakhstan, Vietnam and r _ Palaeobiogeography of Late Devonian brachiopods from Iran 329

Afghanistan consistent with reproductive isolation strongly suggestive that, at least for some taxa, associated with the existence of oceanic waters dispersal was occuring between these regions. between the various blocks. Soja (1992) has also If the same range of biological (short larval life recorded high levels of endemism in Famennian span, substrate selection) and physical faunas from eastern North America. These records (temperature, substrate, salinity) factors known to of unusually high levels of provinciality for the Late limit the distribution of modern articulate Devonian, imply significant geographic separation brachiopods (Richardson 1997) also prevailed and isolation of faunas, at least for these regions during the Palaeozoic, then this distribution pattern (Figures 4 and 5). Whilst it is true that Late can only be adequately explained if the shallow Devonian faunas are not as markedly provincial as marine shelves of Euramerica and North Early Devonian faunas, it is clear that regional Gondwana were close enough to allow larval differences were apparent during the Late dispersal and some degree of faunal mixing (Brice Devonian and the suggestion that all Frasnian­ et al. 1994; 1999). Support for this interpretation is Famennian brachiopod faunas were uniformly provided by Galle et al. (1995: 233) who presented cosmopolitan is a gross oversimplification. detailed biogeographic data on Middle Devonian In terms of Iranian faunas, previous authors such (Eifelian-Givetian) brachiopods and corals from the as Gaetani (1965, 1968), Brice (1971) and Brice et al. Czech Republic that indicated the northern (1973; 1999) have commented on the remarkable Gondwanan margin "approached closely to similarity between Late Devonian brachiopod southern Laurussia" (=Euramerica) during the faunas from Iran and synchronous faunas from Devonian. Afghanistan, Annenia and Pakistan, with Brice et The suggestion that the Iran-Afghan al. (1978) suggesting the probable existence of an microcontinent may have been situated midway Irano-Afghan platform during this interval. The between Euramerica and Gondwana in a 2000 km + quantitative data presented herein strongly wide ocean and acted as a staging post for trans­ supports these conclusions (Figures 2, 3). Frasnian oceanic dispersal of brachiopods and other marine and Famennian genera from Iran display strongest invertebrates cannot be totally ruled out, especially links with faunas on terranes situated at a similar if the subtropical latitude for the entire region is latitude along the northern margin of Gondwana accepted. Jaccard Coefficents presented here show (Figures 4, 5). that Late Devonian brachiopods from Iran and Though the palaeobiogeographic signal is not as Afghanistan had much stronger ties to Gondwana strong, there is also clear evidence of some links than to Euramerica, though this may simply reflect with Euramerica (Figures 2-5; Table 1). Within prevailing oceanic current patterns. Euramerica, Brice et al. (1994) documented faunal Talent and Mawson (1979: figure 5) have links between Frasnian faunas from Boulonnais, presented results of a Jaccard analysis of Frasnian North France and eastern North America. Detailed brachiopod faunas from Pakistan. Though no direct evaluation of biostratigraphy and location of first comparison was made with Iranian brachiopod occurrences of corals, brachiopods, crinoids and faunas, links are recorded with western (29'10) and ostracods from these areas led Brice et al. (1994) to eastern (17'10) Afghanistan. Comparison of Frasnian suggest an easterly migration route along the faunas from NW Pakistan and Afghanistan southern margin of Euramerica from eastern North presented herein record a Jaccard Coefficient of 9%, America to Boulonnais. However, Oczlon (1990: (Figure 2), though this may be due to an increase in figure 1) used palaeocurrent evidence to suggested knowledge of faunas from both regions. that the southern margin of Euramerica was Young (1987, 1990, 1995, 1998) provided probably dominated by strong westerly directed comprehensive palaeobiogeographical analyses of "South Equatorial Current" during the Middle global mid Palaeozoic vertebrate faunas and Devonian that swept down to North Africa to concluded that there was gradual convergence eventually form the easterly directed "North between Euramerica and Gondwana throughout the Gondwana Current" along the southern margin of Devonian leading to eventual "collision and Prototethys. Heckel and Witzke (1979) suggested exchange of continental floras and faunas at or near similar current directions in their palaeoclimatic the Frasnian-Famennian boundary" (Young 1987: reconstructions for the Devonian. If Oczlon's (1990) 299). However, Talent (1984) has pointed out that if interpretation is accepted, it seems more likely that such a connection took place at the Frasnian­ the larvae of North American taxa would be more f;amennian boundary, the resultant isolation of shelf likely to have along the northern margin faunas either side of the collision zone should have of Gondwana following the dominant current generated increased provinciality in Famennian and direction (Figures 4, 5). The presence of congeneric early Carboniferous marine faunas. In terms of the (in some cases conspecific) brachiopods in Frasnian­ brachiopod data presented herein there is no Famennian sequences in eastern Euramerica evidence of such provinciality. Indeed the presence (western Europe) and Northern Gondwana is of taxa such as Centrorlzynclzlls, Megalopterorlzynclllls 330 G.A.Brock,~.Yazdi and Toryniferella in eastern North America and northern Gondwana during the Late Devonian (e.g. Northern Gondwana (Figure 5) during the Morel and Irving 1978; Van der Voo 1988; Famennian indicates that some sort of latitudinal Bachtadase et al. 1995; Metcalfe 1996, figure 13). seaway must have been open between the two supercontinents. Becker (1998: 5) has also presented evidence suggesting migration of Late Devonian ACKNOWLEDGEMENTS ammonoids took place along the northern margin GAB extends sincere thanks to colleagues and of Gondwana via the "Rhenohercynian" seaway. It students at the Department of Geology, University seems more likely, given the increase in of Esfahan for hospitality, access to collections and provincialism of marine faunas globally after the logistical support whilst in Iran. Funding for this Visean, that the collision of Euramerica and North research was provided by a 1998-9 Macquarie Africa coincided with the Variscan orogeny (Oczlon University New Staff (MUNS) Grant to GAB. Dr 1992), sometime during the Visean (Ross 1979; Denise Brice and Professor Neil Archbold provided Talent 1984). constructive reviews of the manuscript. Dr. Brice A few interesting examples of disjunct also kindly allowed access to brachiopod distribution are apparent in the Famennian data distribution data from unpublished manuscripts. presented (Figures 3, 5; Table 1), possibly reflecting Judy Davis, GeoGraphics Unit, Macquarie rising cosmopolitanism. The genus Sulcatospirifer University drafted the figures. 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