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DEVONA BRUŅUZIVJU UN AKANTOŽU PALEOEKOLOĢIJA Un to MAZUĻI LODES IEGULĀ

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DEVONA BRUŅUZIVJU UN AKANTOŽU PALEOEKOLOĢIJA Un to MAZUĻI LODES IEGULĀ DISERTATIONES GEOLOGICAE UNIVERSITAS LATVIENSIS Nr. 21 IEVA UPENIECE DEVONA BRUŅUZIVJU UN AKANTOŽU PALEOEKOLOĢIJA un to MAZUĻI LODES IEGULĀ PROMOCIJAS DARBS Doktora grāda iegūšanai ģeoloģijas nozarē Apakšnozare: pamatiežu ģeoloģija LATVIJAS UNIVERSITĀTE Promocijas darbs izstrādāts Latvijas Universitātes Ģeoloģijas nodaļas Pamatiežu ģeoloģijas katedrā no 1995. gada līdz 2011. gadam. Šis darbs izstrādāts arī ar Eiropas Sociālā Fonda atbalstu projektā “Atbalsts doktora studijām Latvijas Universitātē” (2010./2011.māc.gads). Promocijas darba vadītājs: Ervīns Lukševičs, LU profesors, Dr. geol. (Latvijas Universitāte) Recenzenti: Ivars Zupiņš, Dr. geol., Latvijas Dabas Muzejs Elga Kurika, PhD, Tallinas Tehnoloģiju universitātes Ģeoloģijas institūts Aleksandrs Ivanovs, Cand. geol.-min. sci, Sankt-Pēterburgas Universitātes docents Promocijas padomes sastāvs: Vitālijs Zelčs, profesors, Dr. geol. – padomes priekšsēdētājs Ervīns Lukševičs, profesors, Dr. geol. – padomes priekšsēdētāja vietnieks Aija Dēliņa, docente, Dr. geol. Laimdota Kalniņa, asoc. profesore, Dr. geogr. Valdis Segliņš, profesors, Dr. geol. Ivars Zupiņš, Dr. geol. Padomes sekretārs: Ģirts Stinkulis, asoc. profesors, Dr. geol. Promocijas darbs pieņemts aizstāvēšanai ar LU Ģeoloģijas promocijas padomes 2011. gada 19. maija sēdes lēmumu Nr. 02/2011. Promocijas darba atklāta aizstāvēšana notiks LU Ģeoloģijas promocijas padomes sēdē 2011. gada 2. septembrī, Rīgā, Alberta ielā 10, Jāņa un Elfrīdas Rutku auditorijā (313. telpa). Ar promocijas darbu ir iespējams iepazīties Latvijas Universitātes Zinātniskajā bibliotēkā Rīgā, Kalpaka bulvārī 4 un Latvijas Akadēmiskajā bibliotēkā Rīgā, Lielvārdes ielā 4. Atsauksmes sūtīt: Dr. Ģirts Stinkulis, Latvijas Universitātes Ģeoloģijas nodaļa, Raiņa bulvāris 19, LV-1586, Rīga. Fakss: +371 6733 2704, e-pasts: [email protected] © Ieva Upeniece, 2011 Latvijas Universitāte www.lu.lv DISERTATIONES GEOLOGICAE UNIVERSITAS LATVIENSIS Nr. 21 IEVA UPENIECE PALAEOECOLOGY and JUVENILE INDIVIDUALS of the DEVONIAN PLACODERM and ACANTHODIAN FISHES from LODE site, LATVIA DOCTORAL THESIS In Partial Fulfillment of the Requirements of the Doctor Degree in Geology Subdiscipline of Bedrock Geology UNIVERSITY OF LATVIA The doctoral thesis was carried out: Chair of Bedrock Geology, Department of Geology, University of Latvia, 1995-2011. This work has been supported by the European Social Fund within the project “Support for Doctoral Studies at University of Latvia” (2010/2011 acad.year). Supervisor: Ervīns Lukševičs, Professor, Dr. geol. (University of Latvia) Reviewers: Ivars Zupiņš, Dr. geol., Natural History Museum of Latvia Elga Kurika, PhD, Institute of Geology, Tallinn University of Technology, Aleksandrs Ivanovs, Cand. geol.-min. sci, University of St.-Petersburg Doctoral Committee: Vitālijs Zelčs, Professor, Dr. geol. – chairman Ervīns Lukševičs, Professor, Dr. geol. – deputy chairman Aija Dēliņa, Dr. geol. Laimdota Kalniņa, Associate Professor, Dr. geogr. Valdis Segliņš, Professor, Dr. geol. Ivars Zupiņš, Dr. geol. Secretary: Ģirts Stinkulis, Associate Professor, Dr. geol. This thesis is accepted for the commencement of the degree of Doctor of Geology (in Bedrock Geology) on May, 19 by the Doctoral Commitee of Gelogy, University of Latvia. The thesis will be defended at the public session of the Doctoral Committee of Geology University of Latvia, on September, 2, 2011, Alberta Street 10, Jāņa un Elfrīdas Rutku auditorium (Room 313). The thesis is available at the Scientific Library of the University of Latvia Kalpaka Blvd. 4, Rīga, and Academic Library of Latvia, Lielvārdes Street 4, Rīga. Address for submitting of comments: Dr. Ģirts Stinkulis, Department of Geology, University of Latvia, Rainis bulvāris 19, LV- 1586, Rīga. Fax: +371 733 2704, e-mail: [email protected] © Ieva Upeniece, 2011 University of Latvia www.lu.lv CONTENTS Anotācija …………………………………………………………………………….. 4 Abstract ……………………………………………………………………………… 5 Introduction ………………………………………………………………………….. 6 1. Geological background and biostratigraphy of the Lode Formation ………. 11 1.1. Geological setting ……………………………………………………………. 11 1.2. Faunal characteristic ………………………………………………………….. 14 1.3. Comparison with similar ichthyofaunas of other areas ………………………. 17 2. Material and methods …………………………………………………………… 21 2.1. Examined fish and invertebrate material …………………………………….. 21 2.2. Field study methods ………………………………………………………….. 22 2.3. Methods of morphological research ………………………………………….. 23 2.4. Methods of taphonomical research and palaeoecological reconstructions …… 24 3. Palaeoecology and fish taphonomy of the Lode fossil site ……………………. 25 3.1. Distribution of fossils in the “juvenile fish” lens ……………………………. 25 3.2. Fish taphonomy ……………………………………………………………… 35 3.2.1. Disintegration process of placoderm Asterolepis ornata body ……….. 35 3.2.2. Disintegration process in other antiarchan fishes ……………………... 40 3.2.3. Taphonomy of acanthodian Lodeacanthus gaujicus ………………….. 41 3.2.4. Discussion on fish taphonomy ………………………………………… 41 3.3. Types of fossil fish taphocoenoses …………………………………………… 42 3.4. Trophic relations of Lode faunal and floral assemblage ……………………... 47 4. Morphology and ontogeny of juveniles of placoderm Asterolepis ornata Eichwald sensu Agassiz …………………………………………………………. 52 4.1. Morphology of juveniles of Asterolepis ornata ……………………………… 53 4.1.1. General remarks ………………………………………………………. 53 4.1.2. Size and general proportions …………………………………………. 55 4.1.3. Ridge system …………………………………………………………. 59 4.1.4. Head shield …………………………………………………………... 61 4.1.5. Visceral skeleton . …………………………………………………….. 75 4.1.6. Trunk-armour …………………………………………………………. 76 4.1.7. Pectoral fins …………………………………………………………… 97 4.1.8. Types of sutural connections ………………………………………… 100 4.1.9. Squamous part of the body …………………………………………… 105 4.1.10. Sensory canal system ………………………………………………… 109 4.1.11. Ornamentation ……………………………………………………….. 110 4.1.12. Internal structures ……………………………………………………. 114 4.2. Changes of characters in Asterolepis ornata during the early growth ………. 117 4.2.1. Morphological changes ………………………………………………. 117 4.2.2. Ontogenetic development of exoskeleton ……………………………. 119 4.3. Discussion …………………………………………………………………… 129 5. Morphology and ontogeny of acanthodian Lodeacanthus gaujicus Upeniece … 133 5.1. Morphology of Lodeacanthus gaujicus ………………………………………. 133 5.1.1. General remarks ………………………………………………………. 133 5.1.2. Systematic palaeontology ……………………………………………… 136 2 5.1.3. Histology of scales …………………………………………………….. 158 5.1.4. Comparison of Lodeacanthus with related genera ……………………. 159 5.2. Ontogeny of Lodeacanthus gaujicus ………………………………………… 160 5.2.1. Development of squamation ………………………………………….. 160 5.2.2. Sequence of ossification of skeletal elements ………………………… 165 5.2.3. Changes of proportions of body ………………………………………. 169 Conclusions ………………………………………………………………………… 171 Appendices ………………………………………………………………………… 174 1. Abbreviations used in text and figures Index of anatomical abbreviations used for placoderms ……………………… 174 Apzīmējumu saraksts bruņuzivīm …………………………………………….. 177 Index of anatomical abbreviations used for acanthodians ……………………. 180 Apzīmējumu saraksts akantodēm …………………………………………….. 180 2. Measurements and indicated distances of the head-shield and trunk armour plates of juveniles and adults of placoderm Asterolepis ornata ……………… 181 3. List of tables and figures ……………………………………………………… 193 Tabulu un attēlu un saraksts………………………………………………….. 201 References ………………………………………………………………………… 210 3 Anotācija Promocijas darbs veltīts unikālu, izcili labi saglabājušos bruņuzivju mazuļu, žokļžaunzivju un dažādu bezmugurkaulnieku izpētei, kurus autore ievākusi ilglaicīgu izrakumu laikā Liepas (Lodes) māla karjerā Latvijā. Materiāla ģeoloģiskais vecums attiecas uz augšdevona Lodes svītu. Veikta abu izmirušo zivju klašu pārstāvju - bruņuzivju Placodermi un žokļžaunzivju Acanthodii dažādās attīstības pakāpēs esošu mazuļu skeletu izpēte, izveidotas ķermeņu rekonstrukcijas. Izstrādāta to agrīnās ontoģenēzes un attīstības stadiju pētījumu metodoloģija. Katras agrīnās augšanas stadijas izdalīšanai tika noteikts kvalitatīvo un kvantitatīvo īpašību izmaiņu kopums. Detalizēti aprakstīta devona bruņuzivju mazuļu Asterolepis ornata Eichwald sensu Agassiz morfoloģija un bruņu plātņu veidošanās process. Bruņuzivju mazuļiem izdalītas un aprakstītas piecas secīgas agrīnās augšanas attīstības stadijas. Veikta dažādu augšanas stadiju salīdzināšana, lai atšķirīga vecuma vienas sugas bruņuzivis netiktu aprakstītas kā piederošas jauniem, dažādiem zivju taksoniem. Aprakstīta devona akantožu (žokļžaunzivju) Lodeacanthus gaujicus Upeniece morfoloģija un piecas secīgas augšanas stadijas, kas pamatojas uz dažādu izmēru veselu eksemplāru izpēti. Izpētīts zvīņojuma veidošanās process un veikts tā attīstības salīdzinājums ar citiem Acanthodiformes kārtas pārstāvjiem. Izdalītas sešas zvīņojuma zonas, kas ļauj morfoloģiski atšķirīgās zvīņas kļūdaini neaprakstīt kā piederošus jauniem, dažādiem zivju taksoniem. Izpētīts Lodes laikposma deltu nogulumu ihtiofaunas un bezmugurkaulnieku komplekss. Lodes svītas organismu saraksts papildināts ar jauniem zivju, bezmugurkaulnieku un augu taksoniem. Bruņuzivju mazuļos un akantodēs atrastas un aprakstītas unikālas parazītisko tārpu fosilās atliekas, kas ir senākie līdz šim zināmie zivju parazītu atradumi paleontoloģiskajā hronikā. Tie pārstāv arī senāko mugurkaulnieku- parazītorganismu asociāciju. Tiek sniegts parazītisko tārpu fosilo atlieku salīdzinājums ar mūsdienu parazītiskajiem tārpiem. Tika apstiprināta parazitologa Llewellyn
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