Acarorum Catalogus I 1 PLANT – ARTHROPOD INTERACTIONS IN THE EARLY ANGIOSPERM HISTORY Evidence from the Cretaceous of Israel Editors Valentin Krassilov & Alexander Rasnitsyn 2 Plant – Arthropod Interactions in the Early Angiosperm History Acarorum Catalogus I 3 Plant – Arthropod Interactions in the Early Angiosperm History Evidence from the Cretaceous of Israel Editors Valentin Krassilov & Alexander Rasnitsyn Sofia – Moscow 2008 4 Plant – Arthropod Interactions in the Early Angiosperm History PLANT – ARTHROPOD INTERACTIONS IN THE EARLY ANGIOSPERM HISTORY Evidence from the Cretaceous of Israel Editors: Valentin Krassilov Institute of Evolution, University of Haifa and Paleontological Institute, Moscow E-mail: [email protected] Alexandr Rasnitsyn Paleontological Institute, Moscow and Natural History Museum, London UK, E-mail: [email protected] First published 2008 ISBN 978–954–642-315-3 © PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359–2-870–42–82 [email protected] www.pensoft.net Printed in Bulgaria, January 2008 Contents 5 Contents PART I Traumas on Fossil Leaves from the Cretaceous of Israel 1. Introduction 9 2. Acknowledgements 11 3. Material and methods 12 4. Stratigraphic overview of Cretaceous fossil plant/insect localities in Israel and adjacent countries 15 5. Plant communities 20 6. Taphonomy of phyllostigmas 25 7. Functional morphology of phyllostigmas 29 8. Morphological classification of phyllostigmas 28 9. Diversity of Cretaceous phyllostigmatic structures 41 10. Plant – arthropod interactions 50 11. Environmental variables 62 12. Systematic descriptions of phyllostigmas 65 References 124 Plates 133 6 Plant – Arthropod Interactions in the Early Angiosperm History PART II Fossil Insects in the Cretaceous Mangrove Facies of Southern Negev, Israel 1. Introduction 191 2. Systematic description 193 3. Conclusion 210 4. Acknowledgements 213 References 214 Plates 217 2. Acknowledgements 7 PART I Traumas on Fossil Leaves from the Cretaceous of Israel by Valentin Krassilov in cooperation with Natalia Silantieva & Zeev Lewy 8 PART I: Traumas on Fossil Leaves from the Cretaceous of Israel TRAUMAS ON FOSSIL LEAVES FROM THE CRETACEOUS OF ISRAEL by Valentin Krassilov Institute of Evolution, University of Haifa and Paleontological Institute, Russian Academy of Sciences Contributors: Natalia Silantieva* and Ze e v L e w y ** *Institute of Evolution, University of Haifa **Geological Survey of Israel 1. Introduction 9 1. Introduction Plant – arthropod interactions are generally recognized as a major force under the evolution- ary developments in both plants and arthropods, as well as of the terrestrial ecosystems as a whole. Yet surprisingly little was made for exploring the fossil record for documentation and understanding of these fruitful interactions over times. To enlist fossil plants and arthropods coming from one and the same locality or to depict insect traces on fossil leaves is the first step requiring an analytical procedure for further advancements. Since in Israel even the first step had been but timidly made, it was our task, after de- scribing the fossil flora from the richest Cretaceous locality (Krassilov et al., 2005), to provide an inventory of insect body fossils and of the mite and insect traces on fossil plants. Yet we attempted a step further into the realm of structural and functional relationships linking morphology to ecosystem evolution. The Cretaceous of Negev provides the richest regional material of arthropod traces on fossil leaves (phyllostigmas) at the time of a major restructuring stage in the history of terres- trial ecosystems brought about by the advent and rise of angiosperms. In the middle of Creta- ceous period already, these newcomers have surpassed all the other groups of higher plants in the diversity of their foliar and floral structures, as well as in the plasticity of their life forms. These developments were matched, and perhaps enhanced, by the burst of adaptive radia- tion in the leaf parasites providing ample evidence of plant – arthropod co-evolution. The material is favorably preserved for developing morphological classification of fossil traces, which is necessary for further accumulation and processing of the data. The insect body fossil fauna of Negev is not so rich, but of general interest on account of its paleogeographic situation and paleoecological settings, in particular, the early angiosperm mangrove habitats. Some insect taxa appear surprisingly modern in comparison with their contemporaries north of the Tethys having their closest relatives in the younger Cretaceous and Tertiary assemblage elsewhere. Fortunately, stratigraphic control is rigorous enough for all the insect localities in Israel, and the modern-looking faunistic elements are just evidence of high rate morphological evolution in the insect groups directly or indirectly linked to the explosive angiosperm radiation. Traces of arthropod feeding and habitation on fossil leaves may add to the diversity of contemporaneous fossil faunas, because the trace makers (both miners and gall inducers) are rarely if at all represented in the body fossil assemblages. Yet classification of traces primar- ily reflects the diversity of habits, being indirectly, and sometimes ambiguously, related to taxonomic diversity. In the trace studies, parasitic habits are inferred from the structures these habits induced or produced. It must be said from the very beginning that our comparison with phyllostigmas of modern arthropods do not imply taxonomic affinities, but indicate similarity of behavioral traits that has an evolutionary history of their own. 10 PART I: Traumas on Fossil Leaves from the Cretaceous of Israel Moreover, since parasitic behavior is regulated by the host responses on the one hand, and by the secondary parasites (parasitoids) and predators on the other, and insofar as we can infer the effectiveness of regulation from such structural features as selectivity of preda- tion marks, or the marks of temporary mining, as well as form the co-occurrence of various phyllostigmas and their on leaf diversity, the paleoecological dream of observing extinct ecosystems in action nearly comes true. 2. Acknowledgements 11 2. Acknowledgements This work is a contribution to the German Israeli Science Foundation project no. 1–888– 159.8/2005. We acknowledge the support by the University of Haifa where the main part of the research was conducted. The field works were on several occasions assisted by Israeli Geological Survey. Dr Sophia Barinova has participated both in the field studies and in pre- paring graphs for the mine and gall distribution analysis. Our thanks are due to Professor Eviatar Nevo for his encouragement and advice, to Alexander Rasnitsyn, Conrad Laban- deira and Carlos Herrera for their helpful comments on publications related to this study, to Meinolf Hellmund and Winfried Hellmund, who were involved in the study of Odonata egg sets from the Albian of Makhtesh Ramon, as well as to all the colleagues from the Insti- tute of Evolution, University of Haifa and Paleontological Institute, Russian Academy of Sciences for their interest in our research. 12 PART I: Traumas on Fossil Leaves from the Cretaceous of Israel 3. Material and methods Materials of this study came mainly from the lower – middle Albian Upper Hatira Formation of erosional depressions (‘craters”) Makhtesh Ramon and Makhtesh Hatira in the central – northern Negev and from the mid-Turonian Ora Formation of Arava Rift Valley (Fig. 1). The bulk of the material has been collected in 2001–2006 by Valentin Krassilov assisted by Natalia Silantieva and Sofia Barinova, partly in collaboration with Zeev Lewy and Shelomo Ashkenazi from Geological Survey of Israel, Jerusalem. Alexander Rasnitsyn took part in the field work in Arava in 2004. Additional materials were loaned from Yaakov Lorch’s col- lections deposited in the Hebrew University, Jerusalem. Fossil plants are collected from five localities at two fossiliferous horizons in the lower and upper parts of the Upper Hatira Formation, altogether about 500 fossil plant specimens preserved as the bedding plane impressions coated with pinkish to brick red ferruginous films showing microscopic details of leaf surface topography, including the traumatic structures, or phyllostigmas (on preservation of phyllostigmas see Chapter 5). The flora consists of ferns, ginkgophytes, conifers and angiosperms forming several plant communities (Silantieva & Krassilov, 2006). A number of gymnosperm leaves are stigmatized with marginal and apical cut-outs, and a few narrow-leaved angiosperms (or possibly pro-angiosperms) of Acaciaephyl- lum type bear sets of inserted eggs. No arthropod traces were found in the Cenomanian of Israel, but a few occurred in the material collected by Flavio Bacchia form the Nammoura locality in the Mont-Liban district of Lebanon (Krassilov & Bacchia, 2000; see also Dalla Vecchia et al., 2002 ). The flora is essentially angiospermous, with few ferns and gymnosperms. A leaf of gymnospermous Pseudotorrelia is mined and occasional angiosperm leaves show marginal cut-outs. The mid-Turonian plants and insects of Arava Valley, southern Negev are collected form two localities at the mouth of Gerofit