Molluscan immigrations via biogeographical ecotone of the Middle Russian Sea during the Jurassic 143
Molluscan immigrations via biogeographical ecotone of the Middle Russian Sea during the Jurassic
Mikhail ROGOV 1, Viktor ZAKHAROV 1 and Dmitry KISELEV 2
1 Geological Institute of Russian Academy of Sciences, Pyzhevskii Lane 7, Moscow – 109017, Russia; e-mail: [email protected]; [email protected] 2 Yaroslavl State Pedagogical University, Kotoroslnaya nab. 46, Yaroslavl - 150000, Russia; e-mail: [email protected]
Key-words: paleobiogeography, paleobiology, ecotone, Middle-Late Jurassic, Middle Russian Sea, stratigraphy, immigrational paterns.
ABSTRACT: Patterns of molluscan immigration via the biogeographical ecotone zone of the Middle Russian Sea during the Middle – Late Jurassic have been studied in detail. Firstly, the migration routes and ranges of recent molluscs are briefly reviewed, with some comments about their applicability to the Jurassic. Secondly, the migrational events have been classified by their direction, duration and intensivity. We recognize two main types of immigration by their direction (unidirectional and multidirectional) and by their intensity (mass immigrations and isolated strayings). Recent progress in infrazonal ammonite biostratigraphy leads us to the more precise recognition of paleobiogeographical events and to the understanding of the immigration patterns that can produce more precise correlations.
INTRODUCTION The so-called Middle Russian Sea flooded the East-European Platform during Bajocian-Batho- Jurassic molluscs, especially ammonoids, are nian time. After that for ca. 30 Myr this sea the only group of fossils which has been repeatedly changing its configuration, but always traditionally used to elaborate new approaches in remaining connected with Boreal Basin via the biostratigraphy such as the concept of zones by Timan-Pechora Sea to the north. The southern Oppel (1856-58) and the hemerae of Buckman connection with Peri-Tethyan (Caucasian) seas was (1893) as well as in paleobiogeography (Neumayr not permanent and sometimes disappeared. 1878; Uhlig 1911) and, recently, Sachs et al. (1971), Westermann (2000). This paper summarises the results of recent MIGRATION PATTERNS IN MODERN CEPHALO- detailed investigations of the biostratigraphy and PODS AND BIVALVES AS KEY TO THE PAST paleobiogeography of the Middle Russian Sea (Fig. 1) during Middle-Late Jurassic time. This Modern cephalopods are represented by water body was chosen mostly because of its different living forms which inhabit all seas and position as an ecotone between two main oceans from tidal areas to the hadal zone. The taxa Superrealms during the Middle-Late Jurassic include both neritic and oceanic species, and both (Zakharov and Rogov 2004). assemblages include pelagic and bottom dwellers, 144 Volumina Jurassica, Volumen VI
(Nesis 2003). Post-mortal drift is common for Sepiidae and Spirulidae shells. Cephalopods are very prone to expand and to reduce their species ranges quickly. After the opening of Suez Channel more than 200 different hydrobionts penetrated into the Mediterranean Sea, including BALTIC SHIELD the Indo-Pacific Octopus kagoshimensis (Salman et al . 2005) Recent warming of MIDDLE RUSSIAN SEA the climate (the so-called “Global war- ming”) has provoked a gradual disappeara- Moscow nce of the squid Loligo forbesi from Portugal (Chen et al. 2006) and, presuma- bly, well before this, in Morocco, where it was recorded in the past but never met during intensive investigations in 1995-1998 (Laptikhovsky, pers. comm). The influence of interspecific competition on the ranges of extant cephalopods is still insufficiently known. Inverse patterns of abundance of two commercial squids Illex TETHYS argentinus and Loligo gahi has been suggested as the possible result of inter- specific competition (Arkhipkin and 1234 Middleton 2002), and the same causal hypothesis was invoked to explain Fig. 1. Middle Russian Sea and adjacent areas during the Late Jurassic (example of Late Kimmeridgian – Early Volgian): 1 – land areas; 2 – oceanic basins; the strong depth segregation in some North 3 – shallow epicontinental seas; 4 – coastal areas, sometimes flooded by sea. African coleoids (Arkhipkin and Laptikho- Palaeogeography and paleolatitudes are from Thierry and Barrier (2000), corrected. vski 2005). Nevertheless, numerous occasions of the coexistence of morpho- with the maximum diversity in the tropics and logically close species are known, amongst comme- subtropics (Nesis 2003). They occur in water rcial taxa (Bonaud et al. 1998; Herke and Foltz of normal marine salinity, between 27 and 37‰ 2002). (Jereb et al. 2005) with some rare exclusions. With respect to cephalopods, bivalves are slow- The bulk of modern cephalopods is characterized moving when adult, (except when attached to algae by very fast growth and short life-cycles with or wood) and their migratory ranges are restricted terminal spawning. Cephalopods are voracious, up to several kilometers, such as in Macoma active predators (Nesis 1985; Jereb et al. 2005). baltica (Hiddink 2002). However, many species The major factors which influence cephalopod have planktonic larvae that can be widely dispersed distribution are temperature, food and currents. by oceanic currents. The major factor limiting their Oceanic species usually undergo daily vertical spreading is water temperature during spawning. migrations, occurring at depths of about 200 to 700 Temperature limits for gonad maturation and m during the day, and ascending into the uppermost hatching could be much narrower in comparison 200 m for the night (Jereb et al . 2005). Some with those of adult animals (Scarlato 1981). cephalopods are characterized by ontogenetic Another important factor is the presence/absence and/or seasonal migrations, either active (Omma- of a suitable substrate for juvenile settling. strephidae) or passive, through areas with closed Gastropod taxa with pelagic larvae are widely circulation ( Spirula ) (Nesis 1985). Some neritic distributed along the west coast of South America, cephalopods have small natural habitats and some where the necessary rocky substrates are of them have very wide distribution, for example abundant. In contrast to this, the east coast of Octopus vulgaris (Nesis 1987). Oceanic coleoids the continent is represented almost exclusively usually have latitudinal distribution, but also pan- by sandy substrates, and species with direct boreal, bi-subtropical amphi-atlantic ranges occur development predominate from the subtropics Molluscan immigrations via biogeographical ecotone of the Middle Russian Sea during the Jurassic 145 to Antarctica (Gallardo and Penchaszadeh 2001). during the last decades (Kennedy and Cobban 1976; Bivalves probably could suffer the same problem. Olóriz 1990; Cecca 1992; Westermann 1996; The critical temperatures differ between species Westermann and Tsujita 1999, among others). and populations depending on latitude or seasonal In a similar manner to modern cephalopods the temperature acclimatization. Peck and Conway ammonoids were possibly animals with different (after Pörtner 2001) suggested that tropical and tem- modes of life, from planktonic to quasi-nektonic perate species are more eurythermic than polar ones. and nekto-benthic ones. The ammonoids inhabited The polarbound limit of the species range different biomes, and the oceanic groups of in summer is determined in bivalves by the ammonites are usually different from those minimum temperature suitable for reproduction, of neritic areas. Perhaps their penetration from and in winter by the minimum survival temperatu- shelf areas to the oceans, in contrast to coleoids, re. The equator-ward limit of the species distribu- happened to planktonic, non nektonic, forms. tion in winter is determined by the maximum tem- It is very possible, that many ammonoids perature of reproduction whereas the maximum (especially heteromorphs) were vertical migrants survival temperature restricts this boundary in (Westermann and Tsujita 1999), while others summer (Scarlato 1981). These adaptive patterns mostly swam above the bottom. The bulk of permit the bivalves of high latitudes to inhabit the ammonoids was restricted to waters with normal deep waters of the temperate zone, whereas tro- oceanic salinity, with some exceptions like pical or subtropical species are able to penetrate Ceratites which lived in restricted carbonate far into high latitudes in summer. For example, platforms surrounded by evaporate basins (Wester- a Boreal species Astarte crenata inhabits shallow mann and Tsujita 1999). Ontogenetic migrations areas in both the Barents Sea and the North Sea, leading to spatial segregation of juvenile and adult and occurs in deep biotopes of the Gulf of Biscay. growth stages are also known in ammonoids Some tropical-subtropical (for example, Solen cor- (Westermann 1996). neus ) and numerous subtro-pical relatively deep- Migrations which are related to long-time water bivalves occur in the shallow habitats of changes in the natural habitat, sometimes Possjet Bay (42° N, Japan Sea) (Golikov and accompanied by diversification or sharp changes in Scarlato 1967; Scarlato 1981). fossil (ammonite) communities, should be These biological features controlling the separated from cyclic and repeated vertical, geographical ranges of bivalves in modern seas, ontogenetic and seasonal migrations. Some of these were important for bivalve spreading in the past. types of migration in ammonites were recently That is why, in our opinion, the southward reviewed by Reyment (2005) with some additions spreading of natural habitats of Boreal bivalves (for concerning necroplanktonic drift and “spurious example, belonging to the genera Buchia and migrations”, caused by tectonics (continental drift, Retroceramus ) and the northward penetration of terrane movement, etc ). Long-time and non-cyclic Tethyan Trigoniidae were caused by movement of migrations we could call “immigrations”, following larvae by currents. Another explanation of these the earlier usage of this term in ammonite events is the movement of water masses caused by biogeography (Westermann 1996; Fernandez-Lopez transgressive-regressive events. In comparison to and Melendez 1996; Schweigert 2000; Navarro et al. the situation with ammonites and other active 2005) and study of recent coleoids (Neige 2003), or poikilotherm animals, we should suppose more “invasions” (Sei and Kalacheva 1983). The term developed eurythermy within bivalves (Scarlato “invasion” usually means a strong event, leading to 1981). In spite of the planktonic mode of life of the alteration of ecological relationships ( i.e . mass bivalve larvae, sometimes “ecological opportunity” migrations), while “immigration” could be used for rather than current direction influenced their all the characters of ammonite range movement. dispersion (Amano 2005). The analysis of ammonite immigrations involves difficulties due to problems in the precise outlining of their natural habitat (Bengtson and Kakabadze DISCUSSION ABOUT THE AMMONITE MODE OF 1999). Ammonoids could be assigned to biogeogra- LIFE AND THEIR MIGRATIONAL PATTERNS phic categories (eudemic, miodemic and parade- mic, carrying by currents, and so on (Fernandez- The biogeography, life habitat and migrational Lopez and Melendez 1996)), but even analyzing patterns of ammonites were intensively studied modern cephalopod areas we sometimes could 4 5 2 5 0 5 8 4 6
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o i n n f e : ; Molluscan immigrations via biogeographical ecotone of the Middle Russian Sea during the Jurassic 147 confuse these categories. Examples of post-mortem the following major factors: climate, geographical shell distribution published in the last decades barriers and currents. Climate and geographical were peculiar to the some groups of ammonoids barriers often were interrelated, because an only (Westermann 1996), especially from the appearance or a disappearance of a barrier usually oceanic biome. lead to a change in the water mass circulation and In comparison to most of juvenile coleoids, very hence to climate change. The effect of currents was small ammonite hatchlings with a spherical shell expressed mostly in the differences between fossil filled by gas perhaps were easily scattered assemblages within different parts of a single basin by currents. After growing and maturation, some with constant paleolatitude and sedimentation. ammonites remained planktonic while other began A good example of sign of a cold paleocurrent via near-bottom or demersal life styles. There are the Middle Russian Sea along the Urals during the different ways to reconstruct the mode of life Middle-Late Jurassic has been suggested by Kiselev in ammonites, from morphofunctional studies (2004) due to the strong prevalence of Boreal to facies and stable isotope analysis and to the stu- ammonites. The same current also persisted during dying of palaeopathologies and stable isotope the Volgian (Rogov 2004). signals (Ziegler 1967; Keupp and Ilg 1992; Westermann 1996; Anderson et al . 1994; Zakharov et al. 2006). We have to admit the high diversity in MOLLUSCAN IMMIGRATIONS THROUGH ecology of different ammonites. THE MIDDLE RUSSIAN SEA Following Rawson (1973) we have recognized two different types of immigration according to Any paleobiogeographical reconstruction, intensivity: 1) mass immigrations (or expansions) in contrast to modern biogeography, needs which lead to the movement of high-ranked preliminary correlation and the revealing of the bio- biogeographic boundaries and alter the characti- events. It could be carried out accurately through stics of ammonite communities, and 2) isolate the faunal horizons and their temporal equivalents, straying, when immigrants are scarce and didn’t so-called hemerae (~ca. 0.08-0.2 My, after Page give rise to new stocks. These types sometimes 1995). Only sometimes important changes in could be similar both spatially and temporally. relative abundance of taxa were widely traced One of the most unusual examples of the conside- within the smaller part of faunal horizons. rable distance travelled by small ammonite The Middle Russian Sea existed on the populations is shown by the aspidoceratids of latest boundary zone of two climatic belts, Boreal and Kimmeridgian age. Already living in the Subboreal Tethyan. Analysis of the biogeographical distribu- seas of Poland and central areas of the Russian tion of mollusks shows a permanent location of Platform, aspidoceratids of the late Autissiodo- the biogeographical ecotone in this region rensis Chron are extremely rare (Kutek and Zeiss (Zakharov and Rogov 2004). This is shown by 1997; Rogov 2004). But some representatives of this the contemporaneous existence of the mixed group travelled as far as the Pechora River Basin Boreal, Subboreal and Submediterranean taxa in and even the Subpolar Ural (Mesezhnikov 1984; the bulk of the Middle-Upper Jurassic zonal Zakharov et al . 2005). assemblages (Hantzpergue et al. 1998; Mitta and There are also two main types of immigrations Seltzer 2002; Rogov 2004). The Middle Russian Sea which differ in their direction. was influenced by Arctic and Peri-Tethyan water Unidirectional immigrations that usually masses. This is demonstrated by the temporal coincide with movements of the biochore bounda- dynamics of the Boreal and Tethyan elements in ries and drastically change ammonite communities. molluscan faunas (Figs 2-3). Multidirectional (bidirectional) immigrations Boreal taxa were dominant and widely reflecting the simultaneous southward penetration distributed at the end of the Early (?) and the Late of the Boreal taxa and the northward of Tethyan Bathonian, the earliest Callovian (when multidire- taxa. Perhaps, bidirectional immigrations occur ctional migrations were important), in the latest rarely and reflect mainly a high sea-level stand. Callovian and beginning of the Oxfordian, in the The biogeographical ecotone is usually enlarged at Late Oxfordian, in the beginning of the Middle this time (Zakharov and Rogov 2004). Volgian and near to the Jurassic-Cretaceous The ranges of ammonite species were boundary. Prolonged Tethyan influences occurred influenced (with the exception of evolution) by in the earliest Bathonian, in the beginning of 148 Volumina Jurassica, Volumen VI F i g .
3 O x f o r di n K i m m e r id g ia n Age
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a in Timan-Pechora region and West Siberia. abundance of Buchia. n a t i o n ) . Molluscan immigrations via biogeographical ecotone of the Middle Russian Sea during the Jurassic 149 the Late Callovian and during the Late Kimme- by the vast Boreal transgression connected with ridgian – Early Volgian. Also two short but strong the southward migration of Boreal ammonites, Tethyan events were restricted to the Oxfordian: in especially cardioceratids (Boreal Spread of Arkell, the Early Oxfordian (Cordatum Chron, baccatum 1956). These ammonoids mostly predominated hemera) and the Middle Oxfordian (Densiplicatum- during the Oxfordian, but during short warming Tenuiserratum chrons). The Late Kimmeridgian – episodes Tethyan cephalopods also inhabited Early Volgian penetration of the Peri-Tethyan the Middle Russian Sea (Fig. 3). The Tenuiserratum ammonoids, such as the Oppeliidae and the Aspido- Chron could be listed among the strongest Tethyan ceratidae were mostly long termed in comparison faunistic influence intervals, when oppeliids with other Middle – Late Jurassic invasions. (Richeiceras, Creniceras, Glochiceras, Neoprio- But even during this time the predominance of Peri- noceras ) have penetrated to the Unzha River. Tethyan taxa was combined with the presence of The Early Kimmeridgian also was characterized numerous Boreal genera (Fig. 3). In summary, by a few short events of the penetration of Submedi- the analysis of the fossil assemblages shows strong terranean ammonites (Zakharov and Rogov 2004), Boreal influence during the Middle-Late Jurassic. while the Late Kimmeridgian shows even a predo- In spite of the mixed character of the faunas, Boreal minance of these ammonites (aspidoceratids are taxa prevailed during the Callovian, Oxfordian, common in the Eudoxus and earliest Autissiodo- Early Kimmeridgian and Middle-Late Volgian. rensis Chron, whereas oppeliids became numerous Nevertheless, by detailed investigation of the during the Autissiodorensis Chron and persist till outcrops situated in the East-European Platform, the Early Volgian). Nevertheless, some strata in the representatives of both Peri-Tethyan and Boreal Upper Kimmeridgian and Lower Volgian are taxa are encountered within nearly all the ammo- characterized still by numerous Boreal fossils nite zones. (volgae horizon of the basal Autissiodorensis Zone; Among the most important discoveries of the pavida horizon of the Sokolovi Zone). At the last few years influencing our paleobiogeographic beginning of the Middle Volgian numerous Boreal knowledge, were the finding of true Boreal molluscs Pavlovia penetrated to the Orenburg area (~36° N of Early Bathonian age in the Saratov Volga area in paleolatitudes) and possibly even to the Northern (~37° N in paleolatitudes). These are the cardioce- Caucasus (Rogov 2004). The Regularis hemera ratid ammonites Arcticoceras and Arctocephalites of the Middle Volgian shows a wide spreading of and also the bivalve Retroceramus (Mitta et al . Zaraiskites , from the Pechora area to the Peri- 2004). During the Late Bathonian the first signs Caspian region. During the Middle-Late Volgian of Boreal influence appeared in West Europe and, (Virgatus Chron and later) Submediterranean at the same time, in the Northern Caucasus, where mollusks were very rare; only the bivalve family recently some Bathonian Kepplerites have been Trigoniidae is known. At that time Boreal bivalves discovered (Mitta 2003). The beginning of the Callo- (Buchia ) penetrated very far south. Recently some vian coincides with the opening of the long-term Middle-Upper Volgian Buchia have been recorded Boreal-Tethyan connection of the Arctic and Tethys from the Minor Caucasus (~28° N in paleolatitudes; seas through the Middle Russian Sea perhaps Zakharov and Kasumzadeh 2005). Latitudinal caused by sea-level rise. Simultaneously Boreal immigrations of Boreal ammonites also were Paracadoceras penetrated to the Northern Cauca- common, especially around the Middle-Late Volgian sus (and even its south slope, in Azerbaijan), transition. The latest Volgian deposits are while the Tethyan Macrocephalitidae ( M. jaquoti ) characterized by the appearance of the Arctic travelled as far as the Pechora region (Gulyaev Volgidiscus in the Yaroslavl region (Kiselev 2003). 2005). During the Callovian some oscillations in Significant Boreal influence also persisted through the predominance of the Boreal, Subboreal and to the earliest Ryazanian, where ammonites of the (scarcely) Submediterranean ammonites occured. genus Praetollia are known (Mitta 2004a). Major events of Tethyan ammonite immigration were observed in the late Early Callovian, when Oxycerites and Parapatoceras penetrated north- CONCLUSION wards to the Unzha River Basin (~46° N in paleo- latitudes; Gulyaev 2002; Mitta 2004b), and in The precise study of molluscan immigration the beginning of the Late Callovian. The turn of leads to the more accurate recognition of paleo- the Callovian and Oxfordian has been marked geography, while contemporaneous records of 150 Volumina Jurassica, Volumen VI fossils with different paleogeographic affinities Bonnaud L., Rodhouse P. G. and Boucher-Rodoni R. permits the correlation of different biostratigraphic 1998. A phylogenetic study of the squid family schemes. The European part of Russia seems to Onychoteuthidae (Cephalopoda: Oegopsida). be one of the most suitable areas for the recognition Proceedings of the Royal Society of London , of both the Boreal and Tethyan influences in Series B , 265 : 1761-1770. the Middle-Late Jurassic due to its unique position Buckman S. S. 1893. The Bajocian of the Sherborne between the Arctic Basin, Tethys and the West- District: its Relation to Subjacent and European shallow seas. Superjacent Strata. Quarterly Journal of the Geological Society , 9: 479-522. Cecca F. 1992. Ammonite habitats in the Early Acknowledgements Tithonian of Western Tethys. Lethaia , 25 : 257-267. This study has been supported by RFBR grants Chen C. S., Pierce G. J., Wang J., Robin J.-P., Poulard no.06-05-64284 and 09-05-00456, Russian Science J.-C., Pereira J., Zuur A. F., Boyle P. R., Bailey N., Support Foundation and grants MK-3235.2006.5 Beare D. J., Jereb P., Ragonese S., Mannini A. and MK-856.2008.5 of the President of Russian and Orsi-Relini L. 2006. The apparent Federation. We warmly thank Dr. V. Laptikhovsky disappearance of Loligo forbesi from the south for his valuable comments about the migrational of its range in the 1990s: trends in Loligo spp. patterns of modern molluscs and their ecology and abundance in the northeast Atlantic and some references provided, and reviewers (G. Pavia, possible environmental influences. Fisheries A. Wierzbowski and anonymous reviewer) for their Research , 78 : 44-54. comments. Fernández-López S. and Meléndez G. 1996. 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