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Galvenā Devona Lauka Osteolepiformu DISERTATIONES GEOLOGICAE UNIVERSITAS LATVIENSIS Nr. 11 IVARS ZUPI ĥŠ GALVEN Ā DEVONA LAUKA OSTEOLEPIFORMU K ĀRTAS DAIVSPURZIVIS (SARCOPTERYGII, OSTEOLEPIFORMES) DISERT ĀCIJA RĪGA 2009 DISERTATIONES GEOLOGICAE UNIVERSITAS LATVIENSIS Nr. 11 IVARS ZUPI ĥŠ GALVEN Ā DEVONA LAUKA OSTEOLEPIFORMU K ĀRTAS DAIVSPURZIVIS (SARCOPTERYGII, OSTEOLEPIFORMES) DISERT ĀCIJA doktora gr āda ieg ūšanai ăeolo ăijas nozares pamatiežu ăeolo ăijas apakšnozar ē LATVIJAS UNIVERSIT ĀTE 2 Promocijas darbs izstr ādāts Latvijas Universit ātes Ăeolo ăijas noda Ĝas Pamatiežu ăeolo ăijas katedr ā no 2001. gada l īdz 2009. gadam Promocijas darba vad ītājs: Erv īns Lukševi čs, profesors, Dr. ăeol. (Latvijas Universit āte) Recenzenti: Promocijas padomes sast āvs: Vit ālijs Zel čs, profesors, Dr. ăeol. – padomes priekšs ēdētājs Erv īns Lukševi čs, profesors, Dr. ăeol. – padomes priekšs ēdētāja vietnieks Guntis Eberhards, emerit ētais profesors, Dr. h. ăeog. Laimdota Kalni Ħa, asoc. profesore, Dr. ăeog. Māris K Ĝavi Ħš, profesors, Dr. h. ėī m. Uldis Sedmalis, profesors, Dr. h. ėī m. Padomes sekret ārs: Ăirts Stinkulis, Dr. ăeol. Promocijas darbs pie Ħemts aizst āvēšanai ar LU Ăeolo ăijas promocijas padomes ……. gada …. ................................ s ēdes l ēmumu nr. .../........... Promocijas darba atkl āta aizst āvēšana notiks LU Ăeolo ăijas promocijas padomes s ēdē ……. gada …. ........................., R īgā, Alberta iel ā 10, J āĦ a un Elfr īdas Rutku auditorij ā (313. telpa). Promocijas darba kopsavilkuma izdošanu ir finans ējusi Latvijas Universit āte. Ar promocijas darbu ir iesp ējams iepaz īties Latvijas Universit ātes Zin ātniskaj ā bibliot ēkā Rīgā, Kalpaka bulv ārī 4 un Latvijas Akad ēmiskaj ā bibliot ēkā R īgā, Lielv ārdes iel ā 4. Atsauksmes s ūtīt: Dr. Ăirts Stinkulis, Latvijas Universit ātes Ăeolo ăijas noda Ĝa, Rai Ħa bulv āris 19, LV-1586, R īga. Fakss: +371 6733 2704, e-pasts: [email protected] © Ivars Zupi Ħš Latvijas Universit āte www.lu.lv 3 ANOT ĀCIJA Revid ēts osteolepiformu k ārtas daivspurzivju taksonomiskias sast āvs un stratigr āfisk ā izplat ība Galven ā devona lauka (Austrumeiropas platformas zieme Ĝrietumu da Ĝas) teritorij ā. Kritiski izv ērt ēts morfolo ăisko paz īmju kopums un sniegtas sugu preciz ētas diagnozes. Materi āla anal īze liecina, ka darb ā apskat āmaj ā teritorij ā k ārta p ārst āvēta ar 16 sug ām, kas pieder 9 ăint īm. Aprakst īta jauna tristihopter īdu suga Eusthenopteron kurshi sp. nov., izv ērt ējot osteolepiformu iesp ējamo pielietojam ību devona nogulumu biostratigr āfij ā, konstat ēts, ka grupai ir zin āma noz īme k ā vienam no mugurkaulnieku faunas kompleksu elementiem l īdz ās biež āk pieejamam bru Ħuzivju un bezžokle Ħu materi ālam, ko pier āda korel ācijas piem ēri, sal īdzinot Galven ā devona lauka, Skotijas un Centr ālā devona lauka griezumus. Analiz ēti daivspurzivju tafonomijas aspekti īpaši piev ēršoties Lodes m ālu karjer ā konstat ēto fosiliju sakopojumu veidošan ās apst ākĜiem. 4 ANNOTATION The diversity and stratigraphical distribution of osteolepiform sarcopterygians from the Main Devonian Field (north-western part of the East European Platform) are analysed. Morphological characters have been critically evaluated and amended diagnoses are given. It is concluded that the order is represented by 16 species, which belong to 9 genera. A new species of tristichopterids, Eusthenopteron kurshi sp. nov., is described. Evaluation of the potential application of the osteolepiforms in biostratigraphy shows that the group has certain value as elements in Devonian vertebrate assemblages along with the more frequently available placoderm and agnathan material as approved by the correlation examples between the sequences of the Main Devonian Field, Scotland and the Central Devonian Field. Certain aspects of sarcopterygian taphonomy have been discussed paying special attention to the fossil accumulations in the Lode clay quarry and their formation conditions. 5 SATURS IEVADS…………………………………………………………………………….…………….. 7 1. OSTEOLEPIFORMU K ĀRTAS RAKSTUROJUMS………………………………................. 14 1.1. P ētījumu v ēsture un teritorijas raksturojums……………………..………………… 14 1.2. Osteolepiformiem rakstur īgās īpaš ības…………………………………………….. 20 1.3. Sistem ātika………………………………………………………….………............ 22 2. MATERI ĀLS UN METODES………………………………………………………………….. 25 3. GALVEN Ā DEVONA LAUKA OSTEOLEPIFORMU SISTEM ĀTISKAIS APRAKSTS….. 30 TRISTICHOPTERIDAE …………………………………………………………………… 31 Eusthenopteron………………………………………………………………… …………..... 31 Jarvikina………………………………………………………………………… …………… 50 Platycephalichthys…………………………………………………… …………………… 52 OSTEOLEPIDIDAE…………………………………………………………………...… 61 Cryptolepis…………………………………………………………………………………….. 62 Latvius………………………………………………………………………………………..... 69 Osteolepis……………………………………………………………………… …………….. 73 Gyroptychius………………………………………………………………………………..…. 75 Thursius……………………………………………………………………………………..…. 83 Glyptopomus………………………………………………………………………………...… 86 Piez īmes par atseviš ėā m osteolepid īdu form ām………………………………. 89 4. OSTEOLEPIFORMU STRATIGR ĀFISK Ā IZPLAT ĪBA, TO PIELIETOJAM ĪBA BIOSTRATIGR ĀFIJ Ā UN PALEOBIO ĂEOGR ĀFISKIE ASPEKTI…………….…………… 91 5. DAIVSPURZIVJU TAFONOMIJAS ASPEKTI……………………………………………….. 100 SECIN ĀJUMI……………………………………….……………………………………………… 111 LITERAT ŪRA……………………………………………..……………………………………….. 113 6 CONTENTS INTRODUCTION…………………………………………………………………………………….. 7 1. PRINCIPAL ASPECTS OF THE OSTEOLEPIFORM SARCOPTERYGIANS………................ 14 1.1. History of research and characteristics of the study area…………………………….. 14 1.2. Characteristics of Osteolepiformes ….……………………………………………….. 20 1.3. Systematics……………………………………………………………………............ 22 2. MATERIAL AND METHODS………………………………………………………………….. 25 3. SYSTEMATIC DESCRIPTION OF OSTEOLEPIFORMS OF THE MAIN DEVONIAN FIELD…………………………………………………………………………………………… 30 TRISTICHOPTERIDAE ……………………………………………………………………. 31 Eusthenopteron………………………………………………………………… …….............. 31 Jarvikina………………………………………………………………………… …………….. 50 Platycephalichthys…………………………………………………… …………….……….. 52 OSTEOLEPIDIDAE……………………………………………………………….……… 61 Cryptolepis………………………………………………………………………...……………. 62 Latvius……………………………………………………………………………………………. 69 Osteolepis……………………………………………………………………… ……………… 73 Gyroptychius……………………………………………………………………………………. 75 Thursius…………………………………………………………………………………………. 83 Glyptopomus……………………………………………………………………………………. 86 Remarks on certain osteolepidid forms……………………………………………….. 89 4. STRATIGRAPHICAL DISTRIBUTION OF OSTEOLEPIFORMS, THEIR BIOSTRATIGRAPHICAL VALUE AND PALAEOBIOGEOGRAPHICAL ASPECTS …………….…………………………………………………………………………… 91 5. PECULIARITIES OF SARCOPTERYGIAN TAPHONOMY…………………….………..…… 100 CONCLUSIONS……………………………………….……………………………………………… 111 REFERENCES……………………………………………..………………………………………….. 113 7 IEVADS Veiktie p ētījumi velt īti Galven ā devona lauka (Austrumeiropas platformas zieme Ĝrietumu da Ĝas) osteolepiformu k ārtas daivspurzivju izp ētei. Apskat āmaj ā teritorij ā devona ieži daudzviet ir sastopami dab īgajos atsegumos, k ā ar ī der īgo izrakte Ħu ieguves viet ās gandr īz vis ā Latvijas teritorij ā, Igaunijas dienvidda Ĝā , da Ĝā Lietuvas un Baltkrievijas, Krievij ā – ěeĦingradas, Pleskavas, Novgorodas, Tveras un Vologdas apgabal ā. ĥemot v ērā devona sist ēmas iežu plašo izplat ību Latvij ā, tie ir noz īmīgs Latvijas ăeolo ăiskās v ēstures izp ētes objekts. Daivspurzivis, tai skait ā osteolepiformi, bija b ūtiski faunas elementi t ā laika biocenoz ēs. Kārtas raksturojums un noz īmība. Osteolepiformu k ārt ā ietilpst divas dzimtas – osteolepid īdi (Osteolepididae) un tristihopter īdi (Tristichopteridae). Kopum ā, atbilstoši pašreiz ējām zin āšan ām, osteolepiformu k ārta p ārst āvēta ar 28 ăint īm. Nov ērota ar ī plaša to ăeogr āfisk ā izplat ība. Pirmie š īs grupas p ārst āvji aprakst īti jau 19. gadsimta 30-os gados, kad public ēti pirmie dati par osteolepid īdiem no Skotijas. Šis laiks sakr īt ar ī ar devona sist ēmas izdal īšanu Anglij ā. Dr īz vien par ādās pirmie dati par osteolepiformiem piederošu atlieku atradumiem toreiz ējā Krievijas teritorij ā. 19. gadsimta otraj ā pus ē aizs ākas Skotijas un Kan ādas tristihopter īdu izp ēte (Egerton, 1861; Whiteaves, 1881; Hussakof, 1888). 20. gadsimta 30.-40. gados tiek public ēti dati par osteolepid īdiem no Baltijas (Gross, 1933, 1942; Paul, 1940). Attiec ībā uz citiem apgabaliem j āatz īmē vair āki zi Ħojumi par osteolepid īdiem un tristihopter īdiem no Turcijas (Janvier, Marcoux 1977; Janvier, 1983), Marokas (Janvier et al ., 1979; Leliévre, Janvier, 1986) un Indijas (Gupta, Janvier, 1981). Daudzveid īga devona osteolepiformu fauna zin āma no Austr ālijas – Gyroptychius ? australis (Young, Gorter, 1981); Canowindra grossi Thomson, 1973 (Thomson, 1973; Long, 1985b), Beelarongia patrichae (Long, 1987), Marsdenichthys longioccipitus (Long, 1985a), Eusthenodon cf. wängsjöi , Yambira thomsoni (Johanson, Ritchie, 2000; Johanson, 2004) un Antarktīdas – Gyroptychius ? antarcticus , Koharalepis jarviki Young, Long & Ritchie, 1992, Mahalalepis resima Young, Long & Ritchie, 1992, Platyethmoidia antarctica Young, Long & Ritchie, 1992, Vorobjevaia dolondon Young, Long & Ritchie, 1992 (Young, 1991; Young et al ., 1992). Pateicoties materi āla izcilai saglab ātībai, starp visdetaliz ētāk izp ētītajiem osteolepid īdiem j āmin Medoevia 8 lata Lebedev, 1995 no Krievijas teritorijas (Lebedev, 1995a) un Gogonasus andrewsi Rietumaustr ālij ā (Long, 1985c, 1987; Long et al ., 1997). L īdz ās daudzskaitl īgiem tristihopter īda Eusthenopteron foordi atradumiem Kan ādā, no Zieme Ĝamerikas aprakst ītas vair ākas devona osteolepid īdu sugas: Hyneria lindae (Thomson,
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