Ichnoentomology Insect Traces in Soils and Paleosols Topics in Geobiology

Topics in Geobiology 37

Jorge Fernando Genise Ichnoentomology Insect Traces in Soils and Paleosols Topics in Geobiology

Series Editors Neil Landman Peter J. Harries

More information about this series at http://www.springer.com/series/6623 Jorge Fernando Genise

Ichnoentomology Insect Traces in Soils and Paleosols Jorge Fernando Genise Principal Researcher National Research Council of Argentina Founder, National Ichnological Collection and Division Icnología of the Museo Argentino de Ciencias Naturales President of the First International Congress on Ichnology (Ichnia 2004)

ISSN 0275-0120 Topics in Geobiology ISBN 978-3-319-28208-4 ISBN 978-3-319-28210-7 (eBook) DOI 10.1007/978-3-319-28210-7

Library of Congress Control Number: 2016946484

© Springer International Publishing Switzerland 2017 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made.

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This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG Switzerland Dedicated to (in order of appearance): Diego, Nahuel, Sofía, Federico, Ivan, and Santiago Foreword

In 1993, a mere year after fi nishing our Ph.D. theses, we traveled from Northwest Argentina to the city of Santa Rosa in the central part of the country to attend the First Argentinean Meeting of Ichnology. During the conference, we met an entomologist from the Natural Sciences Museum in Buenos Aires by the name of Jorge Genise. This person seemed to be quite an interesting character; somebody told us that he had spent days inside a mammal-produced cave near the coastal city of Mar del Plata, looking at its scratch-ornamented walls. Jorge was essentially an entomologist by training, but his interests were clearly far and beyond insect systematics. At that time, one of our research lines was freshwater ichnology with a focus on lake systems from a geologic perspective, and Jorge was working on terrestrial ich- nofaunas from a biologic standpoint. The two research lines were in perfect coordi- nation, but the interesting point is that it was by pure chance; neither Jorge knew about our existence, nor did any of us know about him. And, to make things even more curious, it happened in a remote area of the South American continent, a place away from the leading centers of ichnology in the Northern Hemisphere. Since then, the three of us have been lucky to be part of a group of researchers who started to revitalize the fi eld of continental ichnology. In fact, Jorge has been fundamental in “injecting some fresh blood” into research on insect trace fossils, and in reminding us that these structures are close relatives to modern ones; not just ancient objects but they provide real and reliable testimony of the activities of animals. During the last two decades, Jorge and his group of students accomplished a huge and highly signifi cant task, i.e., providing ichnologists, paleobiologists, sedimentologists, and stratigraphers with a conceptual and taxonomic framework to understand and classify continental trace fossils. Please note that for a scientist, it is critical to fi rst improve the understanding (i.e., the careful examination of data, detailed analysis, and fi nal synthesis based on solid observations) and only then focus on the classifi cation that is essential to provide tools of comparison and com- munication among scientists.

vii viii Foreword

Jorge and his team have carefully crafted a patchwork of pieces of information that ranges from a description of individual elements of trace fossils themselves as well as the larger scale ichnofacies model. Before Jorge started his ichnological research, general knowledge of paleosol ichnology was rather patchy and poorly articulated. A fair number of ichnotaxa had been defi ned, but there was limited understanding of their paleobiologic and paleoethologic meaning. The ichnofacies model was also heavily biased towards interpreting marine environments. Twenty years later, the picture of the whole fi eld is radically different. It is rather commend- able that a single working group has accomplished so much and been at the fore- front and center of such an impressive transformation. This book is the result of Jorge’s many years of intense and fruitful work. It is a book with personality, and discusses ichnological concepts in detail without losing sight of the big picture. The reader will discover details about the micromorphology of nest walls as well as the broader applications of ichnology to understand the evolution of insect behavior. Only seldom does one have the opportunity of reading a book that opens a whole new fi eld. This is one of those rare cases because the book represents the foundation of a novel area of research—ichnoentomology—connecting with blood the living to the fossil. Initiating such a fi eld requires the toolbox of an entomologist and the vision of a paleobiologist. Jorge has successfully combined these two facets. Readers will not only discover Jorge’s rigorous research practices, but also his unique sense of humor. We invite the readers to pay special attention to the epigraphs of each chapter which are delightful to explore, and help us connect science to other fi elds of intellectual endeavors. Who knows? While trying to understand how to classify enigmatic structures found during the last fi eld season, the reader may just end up discovering Borges or Cortazar—it is not just about science.

Luis A. Buatois M. Gabriela Mángano Pref ace

In a preface, an author generally briefl y explains what inspired him to write a book. I cannot do this without providing a brief history of discoveries and ideas that have fuelled the fi re of the South American tradition of research on insect fossils, and how I came across these traditions and some of its pioneering representatives. Whether or not it has been recognized, there is a South American school on continental ichnology, that is responsible for the development of pioneer work on paleosols and their trace fossils. The origins of this school can be traced back to the nineteenth century. Contemporary representatives include Eduardo Bellosi, Luis Buatois, Gabriela Mángano, Ricardo Melchor, Renata Netto, Mariano Verde, and myself among others, who organized the First International Congress on Ichnology (Ichnia 2004) in Trelew, Argentina. Surely, this book and my embracing of insect trace fossils research, either consciously or unconsciously, has something to do with that tradition. It is a tradition that in the past lacked any direct connection between masters and disciples. This fi eld has developed under a “pick up the gauntlet” style. We are not sure if the Uruguayan Lucas Roselli had read Serafi n Rivas’s fi rst mention of insect fossil nests in 1884 when he wrote his fi rst paper on insect paleoichnology in 1939. Serafín Rivas (1838–1913) was a Spanish physician who lived in Uruguay and mentioned for the fi rst time (to my knowledge) insect (hymenopteran) nests, from paleosols worldwide. They were from the red sandstones of the Asencio Formation of Uruguay. His work had been signed on February 1881, but it was actually published by 1884. It seems like a very simple discovery or interpretation today, but in the nineteenth century to even think that insect nests could be fossilized and found preserved in rocks was an impressive intellectual advance. Even when Asencio’s nests look very much as extant ones just transformed in rock, their complexity leaves few doubts on their origin. It is also unclear whether the Italian- Argentinian geologist and paleontologist Joaquin Frenguelli (1883–1958) knew of Rivas’s note when he described fossil bee cells from the limestone formations of Uruguay for the fi rst time in 1930 (Chap. 10 ). Frenguelli went on to publish pioneer works on insect fossil nests from hymenopterans, coleopterans, rhizoliths, and paleosols from Patagonia and other localities of Argentina (Chap. 20 ). While these works might seem simple, they were groundbreaking and inspirational achievements

ix x Preface of their time. The phrases nidos de véspidos solitarios meaning solitary wasps nests, and nidos de escarabeidos meaning scarab nests or dung beetle nests have been used in the South American geological literature until now. We do know, however, that Frenguelli was aware of Roselli’s work because of an article published in 1946, in which the former argued against the use of naming trace fossils as Roselli did, and also criticized his interpretations of some insect nests. Francisco Lucas Roselli (1902–1987), a self-taught Uruguayan paleontologist, naturalist, blacksmith, and professor of natural sciences in Nueva Palmira fi rst described and named what is still the largest number of insect trace fossils from paleosols. In 1939, Roselli named four ichnogenera and fi ve ichnospecies of insect nests and pupation chambers from the Asencio Formation of Uruguay and almost half a century later described 13 ichnogenera and 15 ichnospecies of trace fossils attributed to bees, dung beetles, cicadas, and chafers right before he passed away. His work is housed presently at the Museo Municipal Lucas Roselli in Nueva Palmira (Chap. 20). By the 1930–1940s, a North American researcher—William Brown—had concurrently described and named fossil bee cells and a wasp nest from the USA (Chap. 10 ). Between the years of 1971–1972, other geologists picked up the gauntlet and demonstrated that the tradition was still alive. Mario Teruggi (1919–2002) and Renato “Tito” Andreis (1935–2009) started their studies on paleosols. They published several papers including one at the “Symposium on the Age of Parent Materials and Soils”, held in Amsterdam in 1970, which set the foundation for the present state of research on paleosols (Chap. 20 ). Andreis, who recognized Frenguelli’s pioneer work, became an expert on paleosols and included dung beetle brood balls in some of his contributions. Other contemporary geologists, however, disregard the concept of fossil soils and nests. I shared several fi eld trips with Tito during the 1990s (Fig. 1) where he taught me, along with Tom Bown, many aspects of paleosols. I fi nally arrive at the part where I can talk about my meeting with one of the pioneers that infl uenced my own work—Jose “Pepe” Laza. Pepe began working at the Museo de La Plata as a technician in the Division of Paleovertebrates around 1964 where he went on to become one of the most recognized and fi nest Argentinean paleontological technicians. He acquired an interest in insect fossil nests when he recognized dung beetle brood balls in the museum’s exhibition, and when by sheer coincidence some paleontologists collected ant fossil nests from La Pampa and showed them to him. At that time, Pepe had not read Frenguelli’s work, and paleosol studies were still in their infancy. Pepe had a clear idea that the paleontologists have collected fossil nests preserved in ancient soils, and that the big balls of La Pampa were in fact, ant nests. Pepe soon began to collect insect fossil nests, particularly dung beetle ones, during several trips to Patagonia in the 1970s (Fig. 2 ). He published groundbreaking work on insect paleoichnology in 1982 where he described, named, and interpreted fossil ant nests from La Pampa, and also extracted paleoclimatological and paleogeographic inferences from them; an important mile- stone in insect paleoichnology, which involved not only the description, naming, and interpretation of insect fossil nests, but also the extraction of paleoecological Preface xi

Fig. 1 Tito Andreis and the author at Bajo Santa Rosa, Río Negro, Argentina, in 1990 inferences from them. A task that very simple to accomplish nowadays but was very innovative at that time (Chap. 12 ). Here is an excerpt from a textbook on ichnology authored by Ekdale, Bromley, and Pemberton (1984) that gives you a better idea of the knowledge prevalent in the ﬁ eld on this topic at the time. Continental subaerial (i.e. non.-aquatic) deposits can be conveniently grouped into two main categories: eolianites and soil zones. A common misconception regarding such deposits is that they are devoid of fossils . Although their paleontologic record is indeed meager, distinct suites of trace fossils are present that ultimately may prove useful as diag- nostic tools in identifying these environments in the rock record. xii Preface

Fig. 2 Pepe Laza at Sierra de Talquino, Chubut, Argentina, in 1982 (picture courtesy of Pepe Laza) Preface xiii

Fig. 3 From left to right , Mirta González, the author and Pepe Laza at Cañadon Puerta del Diablo, Chubut, Argentina, in 1999

Pepe Laza worked with me for about 10 years before his retirement, but we are still in contact. In fact, he shared the story of his beginnings with me personally. He taught me a lot of things; we both developed a lot of ideas (some published and some not), and shared fi eld trips, one of which, resulted in the discovery of the Cretaceous bee nests and coleopteran pupation chambers in Patagonia (Fig. 3 ). This is one of the oldest evidence of bees, and one of the few Cretaceous nests recorded worldwide. I believe that the preparation of these Cretaceous bee nests is one of Pepe’s fi nest works in his career as a technician (Fig. 9.15 ). Incredibly, when Pepe published his fi rst paper in Argentina, Thomas Bown (Tom) in North America also published one on Egyptian termite and ant nests collected from Egypt. Why incredibly? Because, like Pepe, Tom had also extracted paleoecological inferences from nests for the very fi rst time. It was a completely novel achievement at the time. The same year and independently from Pepe, Tom did the same thing but with Egyptian termite and ant nests. It was as if scientifi c ideas developed, grew, and exploded independently of scientists; as if ideas had their own life, and took their own time to shape up. How were Tom’s beginnings? Tom told me that he became interested in trace fossils in 1968 when he was shown impressive specimens of the coiled beaver burrows called Daemonelix in Nebraska. He soon began his own collection of trace fossils in Wyoming and then in Egypt in 1980, where he saw insect fossil nests for the fi rst time. He had not read about insect fossil nests before, and most sedimentologists xiv Preface spoke about concretions and worm burrows when referring to continental trace fossils. Tom read through more than 300 papers on termite and insect architecture after he discovered the nests in 1980, before writing his Egyptian contribution (Fig. 4). Even without much expertise in the subject, he interpreted Egyptian trace fossils to be nests of social insects just when he discovered them because he had read Edward Wilson’s masterful work on ants. This is when I enter the picture. In 1983, I graduated with a Doctorate in Biological Sciences from the University of Buenos Aires and began working at the

Fig. 4 Tom Bown at Fayum, Egypt, in 1984 digging the specimen of Krausichnus altus (picture courtesy of Tom Bown) Preface xv

División Entomología of the Museo Argentino de Ciencias Naturales. My doctoral thesis and expertise was in soil nests and behavior of wasps. I had always liked paleontology, but had no idea how to combine insect behavior with fossils. I was also clueless about ichnology. I used to walk along the coastal cliffs of Buenos Aires looking for fossils. In 1983, I discovered by myself (I wasn’t aware that this had already been published in the literature) that many of the small fossil mammals that I saw were preserved inside their own burrows. I was really excited by that fi nding because I fi nally had the opportunity to link animal behavior to fossil burrows in spite of the fact that they were not really made by insects. I collected data for 3 years and completed writing up the results generated from my work on rodent fossil burrows by 1986. This work was published in 1989 after being strongly critiqued about the comparison between fossil and extant burrows (Chap. 16). I was introduced to fossil mammal burrows, but was completely ignorant about the existence of ichnology and insect fossil nests. This was when Tom Bown appeared in my professional and personal life. In the late 1980s, Tom had been working on fossil primates in Argentina with John Fleagle. He was interested in insect fossil nests due to his prior work in Egypt, and his experience collecting specimens of different nests in Patagonia. He had collected hundreds of specimens of an enigmatic fossil, named Chubutolithes (by Ihering) (Chap. 18) which had never before been interpreted correctly. Tom published this new interpretation of Chubutolithes as a mud wasp nest with Brett Ratcliffe in 1989. The late paleontologist—Miguel Soria—who worked with Tom in the fi eld and whom I knew from university, showed me Tom’s paper on Chubutolithes that had been published because he knew about my interest and experience working with wasp nests. It was in that precise moment, I had discovered ichnology and the existence of insect fossil nests: the perfect combination of behavior, nests, and insects that I had studied before I was at the university, and fossils, which I have always loved. I had noticed that Chubutolithes was being compared to the wrong extant wasp mud nest. I immediately decided to write a letter to Tom with a picture of what I believed was a more similar extant analog. By then Tom was a recognized expert in the subject, but he welcomed the information in letter and to my surprise, invited me to publish this new idea with him. This story clearly tells us what kind of a person and scientist Tom is. Since then, I have learned a lot from him on ichnology and paleosols. Around this time, I was also introduced to José Laza’s work. Tom and Laza had published a paper together on a fossil termite nests from Patagonia (Chap. 12 ). Tom was generous enough to invite and support me on several fi eld trips including those to Patagonia (1991–1992), Uruguay (1994), and a wonderful one to Egypt (1992) Egypt (Figs. 5 and 6 ). Since then, my research has focused on insect fossil nests, and my work has involved fi eld and laboratory work with colleagues and graduate students, some of them who had already been awarded PhD’s. These researchers have been acknowledged in another section of this book. Years ago, when insect trace fossils were mostly unknown, a geologist told me: “You study things that we piss over”. Now, 25 years later, the same trace fossils that were rare objects in a single locality have begun to appear in other localities, countries, and even continents. xvi Preface

Fig. 5 Tom Bown and the author at Pan de Azúcar, Chubut, Argentina (1991) where most of the specimens of Chubutolithes were found

Fig. 6 Tom Bown and the author in a camp at Qattara Depression, Egypt (1992) Preface xvii

You might wonder what the need is for a book on ichnoentomology? Two quotes stated by Julio Cortazar and Jorge Luis Borges aptly describe my reasons for writing this book. Cortazar aptly summed up his feelings about his book Rayuela when he said: “In a way, it is the experience of a whole life and the attempt to bring it to writing”. Borges stated that “We publish our books to free ourselves from them, to avoid spending the rest of our lives correcting drafts”. I would add to Cortazar’s quote that although this book is the experience of my whole life, I am well aware that there is a lot more to do in this fi eld and I plan to continue doing so with my team; perhaps my team will continue advancing this research after I’m gone. I believe that scientifi c research involves not only fi nding the evidence that supports our ideas, but also fi nding the ones that test and reject them. The subject of this book does not require further explanation, but as the Cuban poet Silvio Rodríguez wrote: “I chose to write about impossible things because about possible ones it has been written too much”, or as Dr. Frankenstein explained: “I pursued nature to her hiding-places ”. What are my expectations from the publication of this book? Perhaps only one: that some students, perhaps from other countries and in the future, fi nd a copy of it on a dirty library shelf and decide to “pick up the gauntlet”. Speaking of dirt, let me conclude with the same words I said at the opening ceremony of Ichnia 2004 adapted from a song by an Argentinean rock band (Aquelarre, 1973): “They are no dirty rocks, they are trace fossils…true diamonds, touch them with your mind, and you will see … how they shine”. Antoine de Saint-Exupéry’s famous phrase from his book Pilote de Guerre (1942) was based on a similar idea: Un tas de pierres cesse d’être un tas de pierres, des qu’un seul homme le contemple avec, en lui, l’image d’une cathédrale (a rock pile ceases to be a rock pile the moment a single man contemplates it, bearing with him the image of a cathedral).

Buenos Aires, Argentina Jorge Fernando Genise Acknowledgements

Chapter 4 of this book deals with classifi cations. It shows (I hope) how diffi cult it is to classify trace fossils and to create groups. I face the same diffi culty when writing these acknowledgements. How does one group all these valuable and dear people into paragraphs? Who do I mention fi rst? Family? Inspiring colleagues? My research team? Friends? Direct contributors? It is impossible to split people into separate groups because they often took on different roles in my life. Members of my research team are my friends, inspiring colleagues, as well as direct collaborators of this book. Some of my friends and particularly my family outside the lab were also my inspiration and direct contributors; without them I would not be here. Everybody on the list below contributed to this book in one way or another. So, let me list the names and give less importance to the order in which they are presented. Tito Andreis, Dolf Seilacher, Jordi de Gibert, Miguel Soria, Manfredo Fritz, and Marcelo Grangeiro are no longer with us. I am grateful for conversations with them and their advice. I miss them. I recognize Tom Bown and Pepe Laza as my guiding light on the pathway of learning about insect paleoichnology. Their detailed stories are included in the preface. Axel Bachmann was, is, and will be always, my master. He is such a good person. He knows insects so deeply that he is capable to teach how they think, but also how to avoid anthropomorphic interpretations in ethology. His lessons, scientifi c rigor, enthusiasm, and generosity have been inherited by my graduate students. It is impossible to thank him for all he did for me; the list will be too long, and the appro- priate words to use to express my gratitude have not been invented yet. Mateo Zelich is another mentor who guided me in my former fi eld research and taught me a lot about insects, plants, vertebrates, fossils, and rocks and how to search for them in the fi eld. A scientifi c monograph may have a single author, but it is the result of a coopera- tive effort among many. The author is simply the person who puts the last brick in a wall that has been constructed by everybody else. The author makes the wall visible, but its construction has been progressing for years. The trace makers of this wall,

xix xx Acknowledgements those who share the facilities of the División Icnología of the Museo Argentino de Ciencias Naturales, day after day, and to whom I am deeply grateful, are (in order of appearance) Mirta González, Eduardo Bellosi, Viky Sánchez, Laura Sarzetti, Liliana Cantil, and Karina Cherñajovsky. However, the team that has worked with me in the fi eld all these years discussing ideas is not only composed of them, but also of colleagues (friends and in some cases former graduate students) from other institutions of Argentina and foreign countries: Mariano Verde, Marcelo Krause, Emilio Bedatou, Ricardo Melchor, Juan Farina, Alejandra Molina, Ana María Alonso-Zarza, Pablo Puerta, Guillermo Roland, Gabriela Cilla, Marcela Cosarinsky, and Roberto Straneck. I am also grateful to others with whom I have shared part of my research: Juan Carlos Sciutto, Gerardo Cladera, Martin Umazano, Julian Petrulevicius, Pablo Dinghi, Javier Muzón, Fabiana Cuezzo, Paco La Roche, Radek Mikuláš, Alfonso Meléndez, Markus Bertling, Nick Edwards, Daniel Poiré, Oscar Gallego, Luis Bala, Philippe Duringer, Frank Krell, Richard Bromley, Patricia Hazeldine, Jerry Rozen, Mariana Morando, Luciano Avila, Conrad Labandeira, Alfred Uchman, Andy Rindsberg, Andre Nel, Arturo Roig Alsina, Duncan McIlroy, Michael Engel, and Jorge Frana. Daniel Speranza, my long time friend, like my brother and an artist, my son Nahuel, Laura Sarzetti, and Mirta González have provided the drawings and paint- ings. Juan Farina (provided data), Viky Sánchez, Alejandra Molina, Laura Sarzetti, Liliana Cantil, Jo Darlington (provided data), Philippe Duringer (provided data), Frank Krell, Pepe Laza, Tom Bown, Conrad Labandeira, Finnegan Marsh, Eduardo Ruigómez, Marcela Cosarinsky, Mariano Verde, Mirta González, Emilio Bedatou, José Luis Román Carrion, Luis Buatois, Gabriela Mángano, Shirley Espert, Silvia Gnaedinger, Patricia Hazeldine, Franz Fürsich, Markus Wilmsen, Arpad David, Paco La Roche, Ana María Alonso-Zarza, Ariel Roth, Diego Genise, and Verónica Krapovickas took and/or gave pictures used in this book. André Nel, Dmitri Ponomarenko, Philippe Duringer, and Jo Darlington kindly provided some articles that I could not fi nd anywhere else. Nadia Garrido reviewed the language and Mirta González the bibliographic lists. The fl ame of South American continental ichnology has been kept alive for more than 20 years by Luis Buatois, Gabriela Mángano, Ricardo Melchor, Renata Netto, Mariano Verde, and me. These researchers supported my crazy idea of organizing the fi rst international congress on ichnology (Ichnia 2004) and bring together ichnologists from all continents and subdisciplines. Eduardo Bellosi and Mirta González provided the paleopedological support for my ichnological research during the last 20 years. Luis and Gabriela, who kindly wrote the foreword, were always available to answer my questions and doubts on theoretical ichnology. Ricardo Melchor provided sedimentological background to a lot of research accomplished during these years too. I wish to thank Duncan McIlroy, Denis Brothers, Xavier Delclós, Renata Netto, Alfred Uchman, and Radek Mikuláš for inviting me to give talks in their countries, which at that time represented important accolades for keeping me on the road. I am grateful to CONICET (Argentinean Research Council) and the ANPCYT (Argentinean Agency of Scientifi c and Technological Promotion) for the permanent Acknowledgements xxi

ﬁ nancial support to my research along these years and also to Alejandro Andreini, who has patiently managed those funds. Thanks to the crews of the Museo Argentino de Ciencias Naturales and the Museo Paleontológico Egidio Feruglio, where I have worked during these years. Wilfredo Fernández and Jorge Frogoni have kindly assisted me along these years every time I visited the el Museo Municipal Profesor Roselli at Nueva Palmira, Uruguay. Sher Saini, from Springer, constantly encouraged me while writing the book and Judith Terpos solved countless doubts on editorial matters. Thank you to all who improved my research with positive or negative criticism. Both made me progress forward. Betty Roberto encouraged me during the last stages of book production. Thank you for being there! Finally thank you papá Fernando, mamá Gladys, abuelo German, abuela Enriqueta, nono Pedro, nona Rosa, tío padrino, tía Emma, and Yaya, who departed early and could not even imagine that I would write this book. Contents

1 Introduction ...... 1 2 The Wall: Where Every Trace Begins ...... 7 2.1 Introduction ...... 7 2.2 Wall Types ...... 8 2.3 Surface Morphology ...... 32 3 Other Characters: Shape, Fillings and Further Micromorphological Characters ...... 35 3.1 Introduction ...... 35 3.2 Shape ...... 35 3.3 Fillings ...... 43 3.4 Other Micromorphological Characters ...... 50 3.4.1 Phytoliths ...... 50 3.4.2 Micromorphological Evidence of Termite Activity in Laterites ...... 52 4 Classifications: The Utopia of Classifying the Unclassifiable ...... 55 4.1 Introduction ...... 55 4.2 Classiﬁ cations with More Biological Taste ...... 57 4.3 Ethological Classiﬁ cations ...... 60 4.4 Ichnotaxonomy ...... 65 4.5 When Ichnotaxonomy Becomes Problematic ...... 67 5 The Keys I: Celliformidae and Coprinisphaeridae...... 71 5.1 Brief Introduction for Two Chapters ...... 71 5.1.1 Those Left Behind...... 72 5.2 Key to Separate Ichnofamilies of Chambered Trace Fossils Occurring in Paleosols ...... 73 5.3 Ichnofamily Celliformidae Genise (2000) ...... 74 5.3.1 Diagnosis and Comments ...... 74 5.3.2 Key to Separate the Ichnogenera of Celliformidae ...... 74

xxiii xxiv Contents

5.3.3 Cellicalichnus Genise 2000 ...... 74 5.3.4 Celliforma Brown 1934 ...... 78 5.3.5 Corimbatichnus Genise and Verde 2000 ...... 81 5.3.6 Elipsoideichnus Roselli 1987...... 81 5.3.7 Palmiraichnus Roselli 1987 ...... 84 5.3.8 Rosellichnus Genise and Bown 1996...... 86 5.3.9 Uruguay Roselli 1939 ...... 87 5.4 Ichnofamily Coprinisphaeridae Genise 2004 ...... 88 5.4.1 Diagnosis and Comments ...... 88 5.4.2 Key to Separate the Ichnogenera of Coprinisphaeridae .... 91 5.4.3 Coprinisphaera Sauer 1955 ...... 91 5.4.4 Chubutolithes Von Ihering 1922 ...... 97 5.4.5 Eatonichnus Bown et al. 1997 ...... 97 5.4.6 Feoichnus Krause et al. 2008 ...... 98 5.4.7 Monesichnus Roselli 1987 ...... 100 5.4.8 Quirogaichnus Laza 2006 ...... 100 5.4.9 Rebuffoichnus Roselli 1987 ...... 101 5.4.10 Teisseirei Roselli 1939 ...... 104 6 The Keys II: Krausichnidae and Pallichnidae ...... 107 6.1 Ichnofamily Krausichnidae Genise 2004 ...... 107 6.1.1 Diagnosis and Comments ...... 107 6.1.2 Key to Separate the Ichnogenera of Krausichnidae ...... 108 6.1.3 Archeoentomichnus Hasiotis and Dubiel 1995 ...... 109 6.1.4 Attaichnus Laza 1982...... 109 6.1.5 Barberichnus Laza 2006 ...... 110 6.1.6 Coatonichnus Duringer et al. 2007 ...... 112 6.1.7 Daimoniobarax Smith et al. 2011 ...... 112 6.1.8 Fleaglellius Genise and Bown 1994 ...... 114 6.1.9 Krausichnus Genise and Bown 1994 ...... 116 6.1.10 Masrichnus Bown 1982 ...... 116 6.1.11 Microfavichnus Duringer et al. 2007 ...... 118 6.1.12 Parowanichnus Bown et al. 1997 ...... 119 6.1.13 Socialites Roberts and Tapanila 2006 ...... 120 6.1.14 Syntermesichnus Bown and Laza 1990 ...... 122 6.1.15 Tacuruichnus Genise 1997 ...... 122 6.1.16 Termitichnus Bown 1982 ...... 123 6.1.17 Vondrichnus Genise and Bown 1994 ...... 127 6.2 Ichnofamily Pallichnidae Genise 2004 ...... 129 6.2.1 Diagnosis and Comments ...... 129 6.2.2 Key to Separate the Ichnogenera of Pallichnidae ...... 129 6.2.3 Fictovichnus Johnston et al. 1996 ...... 129 6.2.4 Pallichnus Retallack 1984...... 132 6.2.5 Scaphichnium Bown and Kraus 1983 ...... 132 Contents xxv

7 Dung Beetle Masonry ...... 135 7.1 Historical Perspective of Dung Beetle Behavior and Traces: From the Ancient Egyptians to Fabre ...... 135 7.2 The Modern Classiﬁ cations of Nesting Patterns and Behavioral Characters ...... 152 7.3 Some Ecological Preferences of Dung Beetles ...... 169 8 Trace Fossils of Dung Beetles ...... 173 8.1 Introduction ...... 173 8.2 Coprinisphaera ...... 173 8.3 Quirogaichnus ...... 183 8.4 Eatonichnus ...... 184 8.5 Chubutolithes ...... 187 8.6 Pallichnus ...... 189 8.7 Scaphichnium and Other Meniscate Burrows ...... 190 8.8 Brief History of Scarabaeinae Coprophagy and Necrophagy as Reﬂ ected by Trace Fossils ...... 191 9 Basic Architecture of Soil Nesting Wasps and Bees ...... 193 9.1 Introduction ...... 193 9.2 Basic Architecture of Solitary Hymenopterans ...... 196 9.3 Malyshev’s Phases and Wasp Behavior ...... 198 9.4 Bee Architecture ...... 206 10 Wasp and Bee Trace Fossils ...... 219 10.1 Introduction ...... 219 10.2 Wasp Trace Fossils in Paleosols...... 220 10.2.1 Fictovichnus sciuttoi and Fictovichnus aragon ...... 220 10.3 Bee Trace Fossils in Paleosols ...... 225 10.3.1 Celliforma ...... 226 10.3.2 Palmiraichnus ...... 229 10.3.3 Corimbatichnus ...... 233 10.3.4 Rosellichnus ...... 235 10.3.5 Uruguay ...... 236 10.3.6 Elipsoideichnus ...... 241 10.3.7 Cellicalichnus ...... 242 11 Blueprints of Termite and Ant Nests ...... 247 11.1 Introduction ...... 247 11.2 Termite Nests ...... 249 11.3 Ant Nests ...... 265 12 The Trace Fossil Record of Eusociality in Ants and Termites ...... 285 12.1 Introduction ...... 285 12.2 Reliability of Named Krausichnidae ...... 288 12.3 Archeoentomichnus ...... 291 12.4 Attaichnus ...... 291 xxvi Contents

12.5 Barberichnus ...... 293 12.6 Coatonichnus ...... 295 12.7 Daimoniobarax ...... 296 12.8 Fleaglellius ...... 297 12.9 Krausichnus ...... 299 12.10 Masrichnus ...... 301 12.11 Microfavichnus ...... 301 12.12 Parowanichnus ...... 302 12.13 Socialites ...... 304 12.14 Syntermesichnus ...... 304 12.15 Tacuruichnus ...... 305 12.16 Termitichnus ...... 306 12.17 Vondrichnus ...... 310 13 Other Insect Trace Fossils in Paleosols: Cicadas, Chafers, Weevils, and Sphinx Moths ...... 313 13.1 Introduction ...... 313 13.2 Cicadas ...... 314 13.2.1 Trace Fossils of Cicadas ...... 319 13.3 Chafers and Weevils ...... 330 13.3.1 Trace Fossils of Chafers and Weevils ...... 337 13.4 Sphinx Moths and its Trace Fossil: Teisseirei barattinia ...... 345 14 Traces from Nest Invaders ...... 353 14.1 Introduction ...... 353 14.2 The Fellowship of the Smoking Dung ...... 355 14.3 Bees and Wasps in the Neighborhood ...... 358 14.4 The Trace Fossil Record of Parasitoidism, Cleptoparasitism and Detritivory Reﬂ ected By Insect Trace Fossils in Paleosols ...... 369 14.4.1 Key to Separate the Ichnogenera of Lazaichnidae ...... 369 14.4.2 Tombownichnus Mikuláš and Genise ...... 369 14.4.3 Lazaichnus Mikuláš and Genise ...... 376 14.4.4 Unnamed Records of Trace Fossils in Insect Trace Fossils in Paleosols ...... 379 15 Soil Neighbors I: Traces of Other Organisms in Paleosols. Crustaceans and Earthworms ...... 383 15.1 Crustaceans ...... 383 15.2 Earthworms ...... 404 16 Soil Neighbors II. Traces of Other Organisms in Paleosols. Vertebrates and Roots ...... 417 16.1 Vertebrates ...... 417 16.2 Roots ...... 431 Contents xxvii

17 Insect Trace Fossils in Other Substrates than Paleosols I. Plant Remains ...... 447 17.1 Introduction ...... 447 17.2 Insect Trace Fossils in Wood ...... 447 17.3 Insect Trace Fossils in Leaves ...... 458 17.3.1 Excisions ...... 460 17.3.2 Incisions ...... 465 17.3.3 Galls ...... 467 17.3.4 Mines...... 471 17.4 Insect Trace Fossils in Other Plant Remains...... 474 18 Insect Trace Fossils in Substrates Other than Paleosols II. Bones, Caddisfly Cases, Trackways, Imprints and Aerial Nests ...... 477 18.1 Insect Trace Fossils in Bones ...... 477 18.2 Fossil Caddisﬂ y Cases ...... 482 18.3 Insect Trackways ...... 490 18.4 Insect Imprints ...... 501 18.5 Aerial Insect Nests ...... 502 19 Trace Fossils as the Physical Evidence of Evolution of Insect Behavior ...... 507 19.1 Introduction ...... 507 19.2 A Brief and Biased Early History of How Ideas on Evolution of Behavior Evolved ...... 508 19.3 Ethological Studies in Hymenoptera ...... 517 19.4 Some Hits in the Most Recent History of the Evolution of Behavior up to the Crossroad with Cladistics ...... 521 19.5 The Crossroad Between Cladistics and Evolution of Behavior ..... 522 19.6 The Crossroad Between Cladistics and Paleontology ...... 524 19.7 The Ichnological Approach of Evolution of Behavior ...... 525 19.8 Brief History of the Meeting Point among Ichnology, Cladistics, and Evolution of Behavior in Insects ...... 532 20 (The Most Remarkable Insect) Ichnofabrics in Paleosols ...... 537 20.1 Methodological Introduction ...... 537 20.2 The Paleogene Asencio Formation of Uruguay and Other Study Cases of Paleosol Tiering and Cross- Cuttings Related to Environmental Changes ...... 549 20.3 The Middle Eocene–Early Miocene Sarmiento Formation of Patagonia. Cross-Cuttings Related to Cleptoparasitism or Detritivory ...... 554 20.4 The Quaternary of Canary Islands. Self Cross-Cuttings Related to Density of Trace Fossils ...... 556 xxviii Contents

21 Paleosol Ichnofacies ...... 559 21.1 Introduction ...... 559 21.2 The Paleozoic Beginning of Paleosol Ichnofacies ...... 560 21.3 The Mesozoic Born Paleosol Ichnofacies ...... 565 21.4 The Cenozoic Born Paleosol Ichnofacies ...... 572 22 Paleoenvironmental Analysis and Ichnoentomological Synthesis ...... 579 22.1 Introduction ...... 579 22.2 Facing the Ghost of Waterlogging ...... 583 22.3 Increasing the Complexity of Interpretations. Gran Barranca and One of the Oldest Grassy Open Habitats ...... 587 22.4 Tracking Anthropoid Evolution in Africa and its Controversial Paleoenvironments with Insect Trace Fossils ...... 593 22.5 The Ichnoentomological Synthesis ...... 601

Appendix ...... 607

References ...... 623 Chapter 1 Introduction

“How between the various layers of stone are still to be found the tracks of worms that crawled about upon them when it was not yet dry” (Leonardo da Vinci, 1510, The Leicester Codex)

As attested by Gould (1998) in his book Leonardo’s Mountain of Clams and the Diet of Worms from where the quote has been taken, da Vinci was probably the first ichnologist. The first to have observed that rocks may preserve traces of extinct animals. What is ichnology? It is the science that studies traces, mostly trace fossils, which are defined as morphologically recurrent structures (nests, burrows, trackways, etc.) resulting from the life activity of an individual organism modifying the substrate (Bertling et al. 2006). Rocks with trace fossils coming from the ancient past reflect how animals behaved. They are Rosetta stones to be interpreted by ichnologists (Genise 2006a). I prefer to define ichnology as a time machine that allows us to observe as directly as possible extinct insects nesting in fantastic landscapes with huge animals, either ground sloths or dinosaurs. Humans have been looking for alternative realities since the antiquity, creating the magic worlds of fairies, elves, dragons, phantoms, and aliens. Cartoons, science-fiction movies and computers gave us the tools to live all these alternative worlds. However, those worlds are imaginary. In contrast, extinct plants and animals compose a paleontological world or worlds along time, which are the only alternate realities that truly existed. Ichnology is the time machine that turns on the movie projector and gives movement to those worlds. Ichnology, a discipline that should be biological in its conception, born, and early development, was paradoxically a geological born discipline. There is nothing more biological that the work of animals or the morphology of roots, and still, geologists were the first to recognize that the traces deserve its own universe. In contrast to organisms themselves, firstly studied by biologists and then discovered

© Springer International Publishing Switzerland 2017 1 J.F. Genise, Ichnoentomology, Topics in Geobiology 37, DOI 10.1007/978-3-319-28210-7_1 2 1 Introduction as fossils in the rocks, traces called attention first in rocks and then in extant environments. Probably biologists are more programed to follow systematics. An entomologist working on a group rarely considers other groups, to a point that there is almost no literature comparing nests or traces produced by representatives of different orders of insects (Chap. 11). This is more than understandable when it is necessary to deal with a million species in combination with an entomologist’s pas- sion for particular groups of insects. The approach of this book particularly contemplates this separation of worlds in an attempt to achieve some kind of confluence: the ichnoentomology. Entomologists, who are usually specialized in particular taxa of insects, will find a readily database on comparable behaviors and traces of other taxa, including their trace fossil record. This may stimulate new ideas, or may simply give a broader readily panorama of others entomological topics, or may introduce the trace fossils produced by the taxa of their own interest. Ichnologists will find the most relevant neo and paleoichnologi- cal information (probably with involuntary omissions) on insect traces in soils and paleosols in a single volume to configure a readily available and unique source database. This will avoid time consuming bibliographic search along innumerable papers spread in journals of very different disciplines and dating from the very beginnings of the scientific research in entomology. In addition, ichnologists devoted to paleoenvironments other than terrestrial ones will find the paleosol counterpart of the ichnological information that they deal with every day. Paleoentomologists will also find the complementary information on how extinct insects behaved, along with the physical evidence of their behavior. Paleopedologists will find some aid for interpreting several of the biogenic structures that they find in paleosols. With some exceptions, it is common in the literature on ichnoentomology that the traces described by former naturalists contain much more details than the more recently described ones. In our present science, time consuming research is only justified by important publishable achievements in specialized disciplines or subdisciplines. Probably the description of linings, surface morphology, fillings and other important ichnological characters is many times considered not worthwhile to include in entomological/behavioral papers. The consolidation of ichnoentomology as a subdiscipline between both, ichnology and entomology, should encourage and favor the return of detailed descriptions of traces and the behaviors involved in their construction to the scientific literature. Ichnologists may read the book without any further introduction. What do entomologists need to know before reading the book? Here are some tips: (1) Trace fossils are grouped in ichnofamilies, ichnogenera and ichnospecies and named following a binomial nomenclature ruled by the ICZN. Without ichnotaxonomy ichnology is not possible. (2) Most insect trace fossils in paleosols are composed of or are chambers, which are built for pupating or nesting. (3) The taphonomic processes are those involved in the preservation of traces, which start once the trace or the whole soil becomes buried and isolated from atmosphere. (4) One of the classical principles of ichnology is that the trace makers cannot be identified because the same organism may produce more than one ichnotaxon and the same ichnotaxon may be produced by more than one organism (Bromley 1996; Buatois and Mángano 2011). 1 Introduction 3

Bromley (1996) claimed that asking the question on which animal produced that trace fossil “is not the right way of addressing trace fossils”. By that time, ichnology was mostly marine ichnology and the one-to-one relationships between insects and their traces were only peeping in the ichnological scenario. Even when insects are rarely preserved inside their traces (Chaps. 8, 13 and 16) and some doubts will always persist on the actual trace makers, some nests are complex enough to allow hypotheses on behavioral homologies along time (i.e. extinct unknown producers are homologues of known extant ones) (Chap. 19). Ichnology should be taken as a dynamic discipline that provides interpretations on which are the producers of trace fossils. These interpretations are (or should be) considered working hypotheses to test and reject as soon as new data indicates it. Don’t be ashamed to change your mind (Chap. 13). The only condition for doing good ichnology is to base hypotheses on sound data: accurate descriptions and supporting documentation. (5) Trace fossil or not? The definition of a trace fossil is clear in the books: they are morphologically recurrent structures (Bertling et al. 2006). However, recognition in the field is not quite the same when they do not show an outstanding complex shape. Some inor- ganic structures are also morphological recurrent structures and several true trace fossils may be confused with other ones if they are not well preserved. In Chaps. 3 and 4 several of these points are addressed. Along this book and perhaps with involuntary omissions, special care was taken to mention the oldest literature on each subject, as an acknowledgement to those people who took the risk of describing and interpreting things that by the time were not considered, known or understood. Clearly, an exhaustive review would repre- sent a whole book in itself and would be far beyond the scope of this one. Actually, the published information that should be included, (and it is not), in the chapters on insect trace fossils in other substrates (Chaps. 17 and 18), is enough to deserve whole books written by experts in those substrates. The aim of including such chapters in this book is to provide a very concentrated overview on all insect trace fossils. By no means are those chapters intended to be comprehensive. Thus, each chapter is written with the aim of giving the tools and literature to further deepen in particular items if necessary. Weiss (2006) reviewing defecation behavior of insects proposed that her review was “meant as an attempt at collection and synthesis of disparate papers that can inform and perhaps nucleate the development of a literature…..and to stimulate further observation, research, and development of theory”. This is the aim of this book too. In contrast to that review on defecating behavior in insects, which had no antecedents, some authors have reviewed with uneven accu- racy and detail insect trace fossils in paleosols before (e.g. Ratcliffe and Fagerstrom 1980; Donovan 1994b; Genise 1999; Zherikhin 2002a, 2003; Hasiotis 2003). Some of these reviews are commented along this book. This book begins with a chapter devoted to the different types of wall that an insect trace may show (Chap. 2). However, to prove that every chapter is linked in different ways with several others and as part of a literary exercise, the book was thought as a circular tale that readers can start in any chapter without losing the thread. The last chapter that refers to the paleoenvironmental significance of insect trace fossils (Chap. 22) is strongly linked with the first one on walls. The last chapter, 4 1 Introduction which also deals with the function of walls, can be read first. In this last chapter, on paleoenvironments and the ichnoentomological synthesis, it is possible to appreciate why it was necessary to devote so many pages and chapters to the recognition of insect trace fossils, their walls (Chap. 2), shapes and fillings (Chap.3 ), their classifications and ichnotaxonomy (Chaps.4 , 5 and 6), and their detailed descriptions and interpretations for the different groups of insects (Chaps. 7–14). It is impossible to play chess if we cannot recognize chess pieces first. The big edifice of paleoenvironmental interpretations becomes a house of cards if it is not grounded in the sound identifications of trace fossils. If we confuse insect trace fossils, for instance, with those of water breather invertebrates the whole paleoenvironmental interpretation changes completely. In some cases interpretations may even range among different kingdoms of trace makers: Moran et al. (2010) interpreted ellipsoidal trace fossils in wood as done by mayfly larvae in subaquatic conditions, whereas the same trace fossils are interpreted as produced by fungi in subaerial conditions (Genise 2004b; Genise et al. 2012; Chap. 17). If we were to take the Argentine writer Julio Cortázar’s idea from his famous book “Rayuela”, chapters like the ones on trace fossils of other organisms in paleosols (Chaps. 15 and 16), or insect trace fossils in substrates other than paleosols (Chaps. 17 and 18), should have no chapter number. They should be included before or after the group of chapters that do follow the main thread of the story, that is 1-2- 3-4-5-6-7-8-9-10-11-12-13-18-19-20-21. However, as Chap. 22 ends with the wall and Chap. 2 starts with the wall, they will inevitably interrupt the circular story. They can be read then, maybe as separate articles, may be, sometime after reading the first book. The purpose of including two chapters about trace fossils in paleosols that are not produced by insects (Chaps. 15 and 16) is not reduced to giving a general overview of the ichnofauna that can be found with insect trace fossils in paleosols. Instead, these chapters have been included because in order to identify the producer of a trace fossil of any type it is necessary to fulfill a two folded procedure (Chap. 19). In the first step it is necessary to prove as thoroughly as possible that the trace fossil under study is similar to the purported extant one; and in the second step, in many cases neglected, we should prove that such trace is not produced by other organisms. A simple fossil burrow may be similar to extant ones produced by beetles, cicadas, wasps, or bees, but it may also be similar to those of crayfishes, earthworms, vertebrates, or it may be a root trace. Why is this structure an insect trace fossil and not an earthworm one? It is necessary to overfly ichnological literature on vertebrates, crayfishes, earthworms and root traces to recognize insect trace fossils. It is necessary to set the limits of neighbor universes to recognize the limits of the one studied. It is impossible to know insect trace fossils without knowing ichnology of other groups as well. Cases of false termite nests are one of the clearest examples of misinterpretations with earthworm and root traces. In the last past years, almost any boxwork recognized in terrestrial environments has been considered as a termite nest, despite other possible and more sound interpretations (Chap. 12). The second step is lacking in all these cases. 1 Introduction 5

The historical approach has been prioritized along this book. How have ideas on insect, earthworm, crayfish, root and vertebrate traces evolved? As a biologist involved in evolutionary studies, I feel it necessary to reconstruct the evolution of ideas, why do I or most people think this way now? Why are we paying attention to some characters and overlooking others? Why are root traces considered trace fossils and why aren’t they named? Why are most vertebrate burrows not named? Why have earthworm burrows been so misinterpreted and understudied? (Chaps. 15 and 16). We are cutting the historical timeline in a place that as a result of evolution of ideas tells us how trace fossils should be described, named, compared with, used for and identified. This will obviously change. Descriptions of trace fossils will remain as the core of ichnology, while present ideas may disappear because they may be improved or dismissed. In some cases old ideas may return and become usable again. It is necessary to understand these historical processes to know how we think or research in the way we do currently. Please, enjoy the book!