FULL DESCRIPTION OF THE PROPOSED PROJECT

Circum-Arctic Lower-Middle Palaeozoic

Vertebrate Palaeontology and Biostratigraphy

BACKGROUND
Remote northern parts of the world have been studied relatively little with respect to their Ordovician-Devonian vertebrate palaeontology. However, the scientific expeditions and studies completed to date (Appendix 1) have yielded remarkably rich assemblages of early vertebrates and several examples of exceptionally preserved material. Based on these studies, what is now the remote Circum-Arctic region is thought to include important centres of origin and diversification for vertebrate higher taxa. Importantly, the vertebrate remains can be found in rocks of different lithology (both carbonates and terrigenous marine sediments) and thus more diverse paleoenvironments than some other fossil groups more commonly used for biostratigraphic correlation. Recent plate tectonic and biogeographic reconstructions have suggested that these presently northern regions were distributed around subtropical areas of the globe during early-mid Palaeozoic time. Therefore, stimulating the study of Ordovician, Silurian, and Devonian vertebrates and improving international cooperation in their interpretation will help to clarify the correlations, paleoenvironments, global reconstructions, and bioevents of that time, while yielding improved understanding of the structure and evolution of one of the most important and scientifically interesting groups of animals.

The territory to be studied includes a large number of islands and adjacent northern mainlands remote from each other and from more accessible parts of the mainland. The study areas also have geologically different structure. Research expeditions to these remote lands, which belong to four different countries (Canada, Denmark, Norway, Russia), are expensive. For this reason studies should be coordinated. Cooperation during expeditions supported by several international organizations and institutions is highly desirable (good examples are: 1939 English-Norwegian-Swedish Expedition to Spitsbergen; and 1994-95 Australian-Canadian-French-Estonian work in the Canadian Arctic).

The earliest work on Circum-Arctic vertebrates resulted from expeditions carried out nearly a century ago and descriptive paleontology published in the years since (Nathorst 1900; Woodward 1902). Most expeditions to the region were for geographical or geological surveying purposes, returning vertebrate fossils only as a side benefit. Nevertheless, some of the collections proved to be important. Examples of such expeditions are listed in Appendix 1.


RECENT DEVELOPMENTS

PALAEONTOLOGY. Discoveries of articulated fossil agnathans and fishes are relatively rare, yet they are essential for elucidation of early vertebrate morphology, taxonomy, and evolution. Some regions of the Arctic are extremely important as the localities of articulated agnathan and fish species. Some examples follow.

Articulated and more or less complete Upper Silurian and Lower Devonian thelodonts come from northern Canada and Spitsbergen (Heintz 1972; Dineley and Loeffler 1976; Wilson and Caldwell 1993). Abundant heterostracan shields have been found in many parts of the Arctic; osteostracan localities are less numerous, but exceptionally good specimens of these groups with both the whole head shield and tail region preserved are known from northern Canada (Dineley and Loeffler 1976; Soehn and Wilson 1990; Adrain and Wilson 1994; yet unpublished material of R.Thorsteinsson). Articulated anaspids from the Canadian Arctic and Severnaya Zemlya are still to be described. Of jawed fishes, articulated acanthodians have been described from the Lower Devonian of northern Canada (Bernacsek and Dineley 1977; Gagnier and Wilson, in press) and Severnaya Zemlya (Valiukevicius 1992). Many of these discoveries were made recently and many more could be made with an increase in focused field work.

Recent Canadian Arctic field work sponsored in part by IGCP 328 (Palaeozoic Microvertebrates) and supported by Canadas Polar Continental Shelf Project and other agencies, concentrated on collecting Silurian and Early Devonian rock samples for their contained microvertebrate remains. At the same time, macrovertebrate and invertebrate specimens were collected so that the microvertebrates could be compared with more complete, often articulated specimens. In 1994, several Silurian - Lower Devonian sections on Baillie Hamilton and Cornwallis Islands were sampled by scientists from Canada, France, and Estonia. Rocks being processed for microvertebrates are in the Institute of Geology, Tallinn, Estonia. Macrovertebrate collections are at the University of Alberta, Canada, and a representative collection is in the National Museum of Natural History, Paris. The 1995 expedition to Prince of Wales Island sampled five classic and newly reported localities known to yield acanthodian, placoderm and sarcopterygian fishes of Lower Devonian age. Institutions in Australia, Canada, and France collaborated to collect and now to process the rocks. At one camp on Drake Bay three sections were sampled, some 200 kg were taken, 60 m of section measured, and collections made of mainly heterostracans, placoderms and sarcopterygians. On the west peninsula of Prince of Wales Island articulated heterostracans and placoderms from the quarry of Dr. R. Thorsteinsson, Geological Survey of Canada, were collected, some 18 m of section measured and another 200 kg of samples collected. These samples are in Paris for processing; a reference collection is at the University of Alberta in Canada.

Also in 1995, the Canadian Museum of Nature in Ottawa and the Natural History Museum of the Humboldt University in Berlin collected Lower Devonian vertebrates from the Anderson River area, Northwest Territories, Canada. These materials are in Berlin being prepared for study.

An important collection from the Cape Storm Formation, Canadian Arctic Archipelago, is currently in Bristol where it is under study by D. Dineley, E. Loeffler, and D. Elliott. Additional field work is necessary to carry this work further.

BIOSTRATIGRAPHY. In biostratigraphical research superior results have been obtained by vertebrate microremains, especially thelodont scales. At present the standard left hand sides of national and international correlation charts are under discussion. In 1993 Silurian vertebrates were proposed for inclusion in the standard correlation chart (the so-called Global Standard). In addition to graptolites and conodonts, the vertebrate, chitinozoan, and spore biozones are now shown. Later, palaeoichthyologists and conodontologists (Mä rss et al. 1995) presented an improved vertebrate biozonal scheme. This will help to define time-slices for paleogeographic studies. The scheme cannot be considered as Global yet because it does not consider all areas of distribution of Silurian deposits, including some Arctic regions.

Biostratigraphy of Silurian and Devonian sequences is most thoroughly studied in the Russian Arctic. A regional vertebrate biozonal scheme has been published for the Silurian of Timan-Pechora Region (Karatajute-Talimaa, Melnikov 1987); characteristic vertebrates for the Silurian of the Severnaya Zemlya Archipelago (Mä rss 1982) and the Devonian of the same region (Karatajute-Talimaa 1985) have been defined. Acanthodian assemblages for the Russian Arctic were established by Valiukevicius (1988).

The Lower Devonian of Spitsbergen (Blieck 1983) and the Upper Silurian-Lower Devonian of the Canadian Arctic (Elliott 1984) have preliminary biozonal schemes based on vertebrates. Though these schemes have been tentatively accepted, they have not been extensively tested or checked against microvertebrate and invertebrate evidence. The most early attempt at microvertebrate work in the Canadian Arctic was that of Vieth (1980). Silurian and Lower Devonian samples to be processed for vertebrates were brought from Baillie Hamilton and Cornwallis Islands by the IGCP 328 international field party in 1994-1995. Their study is in progress. Also recently a book on the stratigraphy and fauna of the Lower Devonian deposits on the River Tareya, Taimyr, was published (Cherkesova, Karatajute-Talimaa and Matukhin 1994). In the next few years it will be followed by a monograph on the fauna and biostratigraphy of the Severnaya Zemlya Archipelago.

An urgent remaining task is comparison of the Silurian graptolite-conodont-chitinozoan-spore-vertebrate biozonal schemes. For example, in the Devonian some discrepancies exist between the zonations at the Silurian/Devonian boundary. Our project hopes to obtain the earliest independently dated occurrence of the thelodont vertebrate Turinia .

PALAEOGEOGRAPHY. S.V.Cherkesova and L.V.Nekhorosheva, St. Petersburgh are finishing work on Silurian and Devonian Russian Arctic paleozoogeography, based on bryozoans, brachiopods, and other fauna. Most other regions have been little studied.

MICROVERTEBRATE DATABASE. Though numerous geological expeditions have returned rock samples from the Arctic regions, many of them have not been properly recorded in databases or thoroughly studied. For example, in Russia there are 8000 samples with microremains from the Arctic for which data remain on cards because of lack of finances for computers in Eastern Europe. By encouraging study of such samples, we hope to make such data available to the international community.


COMMUNICATION:

We propose to publicize the project and report its activities in the newsletter Ichthyolith Issues that has until now been used to coordinate the activities of IGCP 328. We will also supply news to other circulars such as Silurian Times . Ichthyolith Issues has a mailing list of more than 400 specialists on Palaeozoic vertebrates, conodonts, and some other groups. By these means the activities of the group will be brought to the attention of a wider audience.

In addition, we have recently established a World-Wide-Web home page for Palaeozoic Microvertebrates (http://www.biology.ualberta.ca/wilson.hp/Palaeozoic.html) which will help advertise the project and disseminate information about it. Many of the participants have electronic mail addresses (see attached list), which we will use where possible to keep communication costs as low as possible.

National representatives for many countries have agreed to help organize colleagues in their respective countries and to submit regular reports and make funding applications for project activities.
Australia - S. Turner Canada - M. Wilson Estonia - T. Märss
France - A. Blieck Japan - M. Goto Latvia - E. Luksevics
Lithuania - V. Talimaa Netherlands - J.M.J. Vergoossen Russia - A. Ivanov
Sweden - H. Blom UK - V.T. Young USA - D. Elliott


MAIN OBJECTIVES OF NEW PROJECT

The Arctic regions are well known as a source of abundant early vertebrate fossils. Taxonomic studies of agnathans and fishes of this region have greatly contributed to our knowledge of these animals. Biostratigraphic data have been used to some extent in mapping of different areas, and in exploration for oil and gas. Still, studies of many regions are hampered by lack of funding and lack of international collaboration; studies of articulated skeletons from exceptional deposits have not been extensive; studies of microvertebrates are still in their early stages; studies of vertebrate paleoecology, paleogeography, and taphonomy have rarely been attempted; and studies of vertebrate biostratigraphy have seldom been closely integrated with those based on other biostratigraphically important fossils.

The new project aims to coordinate and stimulate research into Lower and Middle Palaeozoic vertebrates from remote northern areas. We plan to encourage detailed studies of the taxonomy, functional anatomy, ecology, phylogeny of vertebrates, using carefully documented collections from measured stratigraphic sections. These studies may include, where possible, reference to taphonomy, community structure and evolution, and identification of bioevents. Intra-provincial faunal variations and sub-provinces, especially with respect to facies distribution patterns, palaeoclimate and barriers, and apparent earlier evolution of groups in certain areas will be dealt with.

A major objective will be to record details of the stratigraphy and sedimentology of vertebrate-yielding sections (especially stratotype sections and those containing also invertebrates/conodonts, ostracodes, brachiopods, palynomorphs and plants etc.), for the purpose of correlation, ecostratigraphy and palaeoenvironmental reconstruction. As a primary goal we will promote refinement of correlations, where possible, through integrated regional biostratigraphical schemes, including comparison of microvertebrate with macrovertebrate records, and of vertebrate with non-vertebrate ones.

The project will emphasize material from the following areas (palaeocontinents after McKerrow and Scotese 1990; see also Li et al. 1993). Northern Laurentian palaeocontinent (studies will be focused in the Canadian Arctic Islands, the northernmost remote areas of the North American mainland, and Greenland). Barentsian microcontinent (Spitsbergen). Baltican palaeocontinent (the project will cover the Timan-Pechora region, the Polar Urals and the islands north of them). Siberian palaeocontinent (Arctic islands north of the Asian part of Russia and Taimyr Peninsula).


WORKING GROUP STRUCTURE AND OUTLINE OF SUBPROJECTS

We propose to establish working groups for each of the major paleocontinental or microcontinental areas represented in the Circum-Arctic region in order to promote field and laboratory studies and meetings in all of these areas. The groups have overlapping membership and will regularly consult and meet jointly to promote collaboration and correlation. The following structure is proposed:


Northern Laurentia Paleocontinent
Canadian Arctic Working Group will deal with Canadian Arctic Archipelagos (Devon, Melville, Somerset, Ellesmere, Cornwallis, Baillie Hamilton, Prinze of Wales Islands) and northernmost North America (e.g., Boothia Peninsula, Anderson River, Mackenzie Mountains, northern Alaska).
Leaders: M.V.H. Wilson, D. Elliott, D. Dineley
Other Vertebrate researchers: P. Ahlberg, H. Blom, C. Burrow, M. Caldwell, Chang Mee-mann, R. Cloutier, S. Cumbaa, P.-Y. Gagnier, D. Goujet, G. Hanke, Z. Johanson, E. Loeffler, T. Märss, M. Purnell, J. Repetski, H.-P. Schultze, K. Soehn, R. Thorsteinsson, S. Turner, J.M.J. Vergoossen
Conodonts & trilobites: C. R. Barnes, B. Chatterton, P. Mannik
Chitinozoans: S. Sutherland
Sponges, tectonics, stratigraphy: T. De Freitas
Spores: J. Richardson
Subproject Outline
The early work of D. Dineley, E. Loeffler, D. Elliott, R. Thorsteinsson, and D. Broad has stimulated renewed interest in study of the Canadian Arctic. Research will continue in:

(a) Palaeontology. Field work to collected important Lower Devonian vertebrates is planned for the Mackenzie Mountains and Anderson River areas in 1996, with materials to be studied in Edmonton and Ottawa, Canada and Berlin, Germany. The group will try to arrange to revisit Frasnian vertebrate localities, discovered at the turn of the century by the Norwegian Expedition, in the Okse Bay Formation, southern Ellesmere Island. At the University of Bristol, UK, D. Broad's collection of traquairaspidids will be published by D. Dineley and L. Loeffler. Extensive Wenlockian vertebrate fauna comes from the Cape Storm Formation. The large collection of D. Elliott, Flagstaff, USA, will be described as well. In Calgary, R. Thorsteinsson's work on Canadian heterostracans is nearly ready for publication. Collections made by the IGCP 328 groups in 1994 and 1995 will also be readied for publication in Edmonton, Paris, and Tallinn. C. Barnes has Ordovician microvertebrates being studied in collaboration with Sansom et al. and Llandovery thelodonts from the Arctic. As a first step, all of these descriptive and taxonomic works will be completed and then necessary additional field work and integrative studies planned.

(b) Biostratigraphy. The first attempt at a correlation between the Canadian Arctic, Spitsbergen and the Welsh Borderland was made by D. Elliott in 1984, using macrovertebrates. The correlations were mostly based on overall faunal composition and assessments of the evolutionary level reached. It remains necessary to work on the micropalaeontological aspects as the vertebrate microremains appear to be more widely distributed and serve both for regional as well as interregional correlations (at least one student is involved in this project). An urgent need is to compare the vertebrate-conodont-graptolite-spore zones. IGCP 328 samples from measured sections in 1994-95 will be important for correlating microvertebrate, macrovertebrate, and other paleontological data.

(c) Evolutionary and environmental aspects. The Canadian Arctic has been considered to be an area in which a number of early vertebrate groups originated (based on evidence from e.g. Cape Storm Formation and Delorme Formation vertebrate fossils). An important task is to investigate the apparent early development of heterostracans here (later evolution in this group was certainly linked to environments, either distal or proximal).


Greenland Working Group will study materials from eastern and northwestern coastal areas of Greenland.
Leaders: H. Blom, J. Peel
Other Vertebrate researchers: Chang Mee-mann, J. Clack, M. Coates, T. Märss
Ostracodes & conodonts: G. Miller
Palynology: J. Marshall
Sedimentology: T. Astin
Subproject Outline
The work has three aspects:

(a) To promote studies on Silurian vertebrate palaeontology and biostratigraphy. Silurian vertebrates were recently discovered for the first time in North Greenland (Bendix-Almgreen 1976, 1986; Turner, Peel 1986). A PhD student has started the study of agnathans and fishes collected from there. The initial objectives of this work are to increase taxonomic and biostratigraphical information. This study is based on unique and rich collections from Hall Land, containing thelodont scales, acanthodian scales and teeth, and heterostracan fragments, but more material is required. Field work, where lithology of the sections will be described and samples for vertebrate-conodont-spore-chitinozoan studies collected bed-by-bed, is needed.

(b) To collect the vertebrate remains from the localities of Eastern Greenland, and to re-examine Devonian fishes and early tetrapods.

(c) To revise Cambrian-Ordovician problematic taxon Anatolepis (lately suggested to belong to a taxon of Arthropoda).


Barentsia Microcontinent
Barentsia Working Group will study materials from Spitsbergen.
Leaders: D. Goujet, A. Blieck
Other Vertebrate researchers: P. Ahlberg, C. Bexiga, U. Borgen, Chang Mee-mann, D. Dineley, N. Heintz, R. Ilyes, Ph. Janvier, A. Ivanov, H. Lelièvre, V. Talimaa, J. Valiukevicius, S. Young
Subproject Outline
Vertebrates from Spitsbergen are better investigated than those from other areas under study. The first report on fragments of fish from Palaeozoic strata of Spitsbergen is by Lankester, 1884. Later, several expeditions visited Spitsbergen (see Appendix 1) and large collections of vertebrate fossils were obtained.

Our project will contribute to the knowledge about morphology, biology, interrelationships and environmental impact of different vertebrate groups (placoderms, rhipidistians, etc.). Succession of faunas will be studied and correlation of vertebrate and palynological biozonal schemes is planned. Work will also emphasize integrative studies of paleoenvironment and biogeography.

Of particular interest for clarifying the evolution of the fishes closely related to tetrapods (panderichthyids, osteolepiforms and rhizodonts) and the base of the lungfish-porolepiform radiation (Youngolepis , Powichthys , Porolepis etc.) and their biogeography is field work on Lower Devonian sites that might yield porolepiforms and lungfishes, and Middle-Upper Devonian sites that could produce osteolepiforms.

Extensive collections of vertebrates are in the Paleontologisk Museum, Oslo, including undescribed heterostracan material. There is also a need for bed-by-bed biostratigraphic work in the Spitsbergen sections.


Baltica Paleocontinent
Baltica Working Group will deal with material from the Baltican paleocontinent, including Novaya Zemlya, Vaigatch, Dolgii and Kolguev Islands, the Polar Urals and northern Timan-Pechora Region.
Leaders: V. Talimaa, V. Menner
Other Vertebrate researchers: O. Afanassieva, A. Ivanov, E. Kurik, E. Luksevics, T. Marss, J. Valiukevicius
Conodontologists: P. Mannik, S. Melnikov, V. Viira, A.V. Zhuravlev
Ostracode workers: A. Abushik, I. Evdokimova
Brachiopods: S. Cherkesova, T. Modzalevskaja
Bryozoa: L. Nekhorosheva
Others being asked (no reply yet): Vertebrates: L. Novitskaya; Bryozoa + stratigraphy: L.Nekhorosheva; stratigraphers: Z.P. Jurjeva, V.S. Tsyganko
Subproject Outline
The collections of agnathans and fishes, especially those from the dissolved rock samples of northeastern Europe, are housed in the institutions of the Baltic countries. They have been studied during the geological mapping of the region for the stratigraphical purposes (Karatajute-Talimaa 1970, 1981; Valiukevicius et al. 1983), and prospecting for oil. A large number of samples were taken both from drill cores and outcrops from Ordovician, Silurian and Devonian strata. Most fossil agnathan and fish remains are still to be described. For example, E. Luksevics has a collection in Latvia from the Upper Devonian of the Timan region, including North Timan, collected by L. Lyarskaya. New material can also be obtained by dissolving the samples from Dolgii Island; other vertebrate remains are to be found in samples taken for conodont research from Novaya Zemlya Island sections.

Co-operative study on taxonomy, succession, and biostratigraphy of conodonts, ostracodes and brachiopods will allow us to find reliable criteria for subdivision and correlation of sections. We recognize the need to find a researcher to study Ordovician to Devonian spores of this region.


Siberia Paleocontinent
Siberia Working Group will study Taimyr and Arctic Islands (Severnaya Zemlya, Novosibirskije = New Siberian Islands, and formally we include here also Wrangel Island from northeastern Russia).
Leaders: J. Valiukevicius, R.Matukhin, V. Menner
Other Vertebrate researchers: O. Afanassieva, A. Ivanov, E. Kurik, E. Luksevics, T. Marss, V. Talimaa
Conodontologists: P. Mannik, N. Sobolev
Ostracode workers: A. Abushik, I. Evdokimova
Brachiopod workers: S. Cherkesova, T. Modzalevskaya
Others being asked (no reply yet): Vertebrate researcher: L. Novitskaya
Subproject Outline
Here as everywhere in the Russian Arctic, Lower and Middle Palaeozoic vertebrates have been discovered during geological mapping. The main collections comprise vertebrate microremains obtained during dissolution of samples with acid. Data on microvertebrates and shelly faunas have been used for subdivision and correlation of the sections. A small part of research results has been published. Most of the publications are devoted to the stratigraphical problems of Severnaya Zemlya Archipelago. Vertebrate morphology and taxonomy is treated only in a few papers (Mark-Kurik 1975, 1976; Karatajute-Talimaa 1983; Novitskaya, Karatajute-Talimaa 1989; Valiukevicius 1992, 1993).

The working group will mainly deal with Early-Middle Palaeozoic agnathans and fishes from the Severnaya Zemlya region. Apart from the partly studied material, there are also many samples, collected during different expeditions to the Severnaya Zemlya islands, Kotelny Island, New Siberian Archipelago, stored in VNIIO and VSEGEI, St.Petersburg. All of these samples need to be processed and included in the work. V. Talimaa, V. Menner, and R. Matukhin are assembling a team to describe Severnaya Zemlya material. S. Cherkesova and L. Nekhorosheva will have data on Silurian and Devonian Arctic paleozoogeography, but many Devonian brachiopods cannot be described without assistance with photographic and publication expenses. Such problems may be solved by international research partnerships. A. Abushik, her student I. Evdokimova, and T. Modzalevskaya can help with ostracode biostratigraphy and have rock samples from Novosibirskiye (New Siberian Islands) available for processing. Samples from Dolgi Island are available, but quality of specimens may not be sufficient. Llandovery conodonts have also recently been found together with vertebrates in Novaya Zemlya. They are similar to those of the Baltic region, but preservation is different. Paleoichthyologists will also get at their disposal material from the more eastern Wrangel island. As the Upper Devonian vertebrates of Severnaya Zemlya are poorly known, field work to the sections is needed.

Possibly in the Silurian, particularly the Llandovery, many northern Russian basins were connected. The Severnaya Zemlya Archipelago causes some problems in palaeogeographic reconstructions, as it has fauna common with the Novaya Zemlya and Urals regions but not with the closely situated Taimyr Peninsula (e.g. Early Ludlow brachiopods); some Llandovery conodont taxa resemble those from Australia; the Silurian vertebrates are in some beds the same as in the Baltica palaeocontinent, but partly they comprise unknown taxa. Therefore there is a need to revise palaeotectonic and palaeogeographic reconstructions of these palaeocontinents using vertebrate data. Publication of results is planned for 1996-1997; they would contain the description of fossils, a bed-by-bed study of their distribution, and the lithology, subdivision and correlation of the sections on islands. Attention will be paid to correlation of sections of archipelagos and northern Siberia (Taimyr Peninsula), basin development, and plate-tectonic reconstructions.


NAMES AND ADDRESSES OF PROJECT ORGANIZERS:

Leader:
Prof. Mark V.H. Wilson
Department of Biological Sciences and
Laboratory for Vertebrate Paleontology
University of Alberta
Edmonton, Alberta
T6G 2E9, CANADA
tel: 1 403 492 5408
fax: 1 403 492 9234;
e-mail: mark.wilson@ualberta.ca

Co-leader:
Dr. T. Märss
Institute of Geology
Estonia Ave. 7
Tallinn EE-0001
ESTONIA
tel: 372 2 454652
fax: 372 6 312074
e-mail: marss@gi.ee


Mailing List of Participants and Prospective Participants

See Appendix 2