Evolution of Cenozoic Land Mammal Faunas and Ecosystems: 25 Years of the NOW Database of Fossil Mammals (Vertebrate Paleobiology and Paleoanthropology) 3031174909, 9783031174902

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Vertebrate Paleobiology and Paleoanthropology Series

Isaac Casanovas-Vilar Lars W. van den Hoek Ostende Christine M. Janis Juha Saarinen Editors

Evolution of Cenozoic Land Mammal Faunas and Ecosystems 25 Years of the NOW Database of Fossil Mammals

Evolution of Cenozoic Land Mammal Faunas and Ecosystems

Vertebrate Paleobiology and Paleoanthropology Series Edited by Eric Delson City University of New York and American Museum of Natural History NY, USA E-mail: [email protected] Home page: http://www.nycep.org/ed/

Eric J. Sargis Yale University, New Haven, USA E-mail: [email protected] Home page: http://www.yale.edu/anthro/people/esargis.html Focal topics for volumes in the series will include systematic paleontology of all vertebrates (from agnathans to humans), phylogeny reconstruction, functional morphology, Paleolithic archaeology, taphonomy, geochronology, historical biogeography, and biostratigraphy. Other fields (e.g., paleoclimatology, paleoecology, ancient DNA, total organismal community structure) may be considered if the volume theme emphasizes paleobiology (or archaeology). Volumes in the series may either be monographic treatments (including unpublished but fully revised dissertations) or edited collections, especially those focusing on problem-oriented issues, with multidisciplinary coverage where possible. The two Series Editors are assisted by an Editorial Advisory Board. All contributions in the series (whether monographs or chapters in edited volumes) will be peer-reviewed by at least three readers, at the level of a journal submission.

Editorial Advisory Board Ross D.E. MacPhee, American Museum of Natural History, New York, NY, USA Peter Makovicky, University of Minnesota, Minneapolis, MN, USA Sally McBrearty, University of Connecticut, Storrs, CT, USA Jin Meng, American Museum of Natural History, New York, NY, USA Tom Plummer, Queens College/CUNY, Queens, NY, USA

Evolution of Cenozoic Land Mammal Faunas and Ecosystems 25 Years of the NOW Database of Fossil Mammals

Edited by

Isaac Casanovas-Vilar Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Barcelona, Spain

Lars W. van den Hoek Ostende Naturalis Biodiversity Center, Leiden, Zuid-Holland, The Netherlands

Christine M. Janis Department of Earth Sciences, University of Bristol, Bristol, UK; Department of Ecology and Evolutionary Biology, Brown University, Providence, USA

Juha Saarinen Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland

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Editors Isaac Casanovas-Vilar Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA) Universitat Autònoma de Barcelona Barcelona, Spain Christine M. Janis Department of Earth Sciences University of Bristol Bristol, UK

Lars W. van den Hoek Ostende Naturalis Biodiversity Center Leiden, Zuid-Holland, The Netherlands Juha Saarinen Department of Geosciences and Geography University of Helsinki Helsinki, Finland

Department of Ecology and Evolutionary Biology Brown University Providence, USA

ISSN 1877-9077 ISSN 1877-9085 (electronic) Vertebrate Paleobiology and Paleoanthropology ISBN 978-3-031-17490-2 ISBN 978-3-031-17491-9 (eBook) https://doi.org/10.1007/978-3-031-17491-9 © The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 Chapters 2 and 3 are licensed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/). For further details see license information in the chapters. This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms 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 specific 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, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover image: Early Vallesian (Late Miocene) landscape and fauna at Höwenegg, Germany. The illustrated fauna includes the sabertoothed cat Machairodus giganteus; the cursorial hyaenid Thalassictis robusta that has hunted a pika of the genus Prolagus; the hornless rhino Aceratherium incisivum; the hipparion Hippotherium primigenium; the gomphothere Tetralophodon longirostris and the deinothere Deinotherium giganteum can be seen in the background. The vegetation consists of herbs and reeds around the lake and mesophytic forest farther from it. This reconstruction is part of the permanent exhibit of the Hessisches Landesmuseum Darmstadt, Hesse, Germany. Artwork by Óscar Sanisidro. NOW database of fossil mammals logo. © Noira Martiskainen This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Foreword

NOW—The Power of a Curious Community Curiosity. It makes life interesting and likely is why you opened this book. Collectively we are curious about our world over the long trajectory of past, present, and future. Paleontologists find new fossils and fit them into phylogenies and ecological roles, and paleoanthropologists search for faunal evidence for climate changes that shaped human evolution. We question, pursue ideas, and think up hypotheses to test; we figure things out. There is a rapid pulse of curiosity in the scientists I know and admire, and for many, it is not just about the thrill of personal discovery but the satisfaction of helping others ask new questions and learn more about where we have been and where we are going. These people are happy to inspire and facilitate curiosity that propels scientific enquiry for their peers and students as well as themselves, and they make up the New and Old World (NOW) Community. This flagship book provides an overview of where NOW has been from its earliest beginnings to the present, with a series of innovative research papers that reveal the potential for NOW data to tackle both new and enduring big questions in the future. The NOW Database has grown over three decades, building upon the cumulative knowledge of several centuries of paleontology and guided by the goal of discovering and communicating a new understanding of mammalian evolution during the Cenozoic Era. Three introductory chapters cover the history and structure of the database, including its scientific (and sociological) underpinnings. The remaining ten chapters take us on a tour of the research that NOW makes possible, focusing on the Neogene time interval (23–2.6 Ma), with data assembled from the fossils of Eurasia, North America, Africa, and South America. Authors include data tables and visually accessible graphics to summarize their results. Readers are treated to an array of research topics, from specific problems in taxonomy, functional morphology, and ecological change to broad comparisons of mammalian evolution and ecomorphic trends on different continents. All of these chapters celebrate what NOW can do to advance knowledge, and their various contributions reflect what is possible with open-source, trustworthy data “for the people” that make such advances possible. Any database seeking to assemble standardized information about the natural world must confront and solve a complex maze of technical and human-resource problems, and it must also be nimble enough to seize unexpected opportunities. NOW decisions had to be made about a standard chronology (e.g., MN Zones) and what to include or not include, such as capturing occurrences of taxa at localities rather than specimen-based data. The latter are undeniably valuable but would have vastly increased data entry effort. A strong vision for what the data should do is critical—many a promising databasing effort has fallen by the wayside by assuming that “if we build it, users will come.” To the credit of the NOW Community, its early leadership (particularly M. Fortelius) recognized the necessity of a strong scientific vision to sustain the time-consuming assembling and checking of the data.

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This vision included the practical benefits of having information about fossil taxa available in one easy-to-use reference system, but that was not enough. There also had to be the potential to address big questions about pattern and process in Cenozoic mammalian evolution, including continent-scale paleogeography, million-year trends in ecology, and competition versus environmental change as drivers of natural selection and community structure. The ten research chapters tackle various aspects of these big questions from a NOW Community perspective along with the benefits, limitations, and opportunities for the NOW database as it evolves into the future. NOW data depend on accurate information about fossil identity (taxonomy), place, and time. A great deal of paleontological research depends on taxonomy, and like many other descriptive scientific activities, this rarely gets the credit it deserves. Without knowing the identity of fossils, however, understanding of diversification, functional innovation, geographic range patterns, extinctions, and myriad other aspects of evolution and ecosystem history is seriously limited. Working paleontologists are often absorbed with questions such as “Do this jaw fragment and that partial cranium from two localities separated by a thousand kilometers represent the same species?” Their decisions about such matters are critical for data “consumers” engaged in synthetic research based on which species occur at different fossil localities. It often takes years of experience, collaboration, interaction with museum collections, and discoveries of new fossils to make trustworthy identifications. These are then formalized and communicated through peer-reviewed publications. Given this extended process, it is no surprise that much important taxonomic knowledge resides in the minds of paleontologists rather than “on paper.” In the early evolution of the NOW database, tentative identifications were offered with which some experts disagreed, motivating them to speed up their taxonomic assignments and avoid propagating incorrect information in the database. This strategy resulted in increased nomenclatural stability for the growing body of NOW data. Chapters on Siwalik rodents by Flynn and colleagues, giraffes by Solounias and Jukar, carnivorous mammals by Friscia and Borths, musk deer by Aiglstorfer and colleagues, and equids by Janis amply demonstrate why up-to-date taxonomy is so important for the continuing growth and utility of the NOW database. Even with excellent fossils and the latest taxonomy, however, there are many unsolvable problems in identifying fragmentary fossils. An alternative approach, advocated in the early years of the Evolution of Terrestrial Ecosystems Program (ETE) and embraced by NOW, is labeled “ataxonomic” or “taxon-free.” This means categorizing fossils according to functional (eco-) morphology, for instance using inferred diet, body size, locomotion, or habitat. The ecomorphic approach can be applied to teeth, limb, and other skeletal elements that may not be identifiable to taxon. Classifying mammals in a fossil assemblage or stratigraphic interval in terms of ecomorphology allows comparisons across time and space without requiring taxonomic identification to the species or genus level. Two decades of previous research in ecomorphology by NOW Community members provided exciting new perspectives on Neogene mammalian community change across Eurasia. In this volume, ecomorphic patterns are examined from several innovative viewpoints, including body size trends over time in carnivorous mammals (Friscia and colleagues), fossil mammals (Huang and colleagues), and geographic patterns in recent tropical mammals (Lintulaakso and Kovarovic). These studies use intercontinental comparisons to search for underlying global-scale controls on ecomorphological traits. Friscia and colleagues analyze NOW data for ecomorphic comparisons of distantly related carnivores isolated on Africa versus South America. Refined information on body mass estimates added to NOW data serves as the basis for Huang and colleagues’ discovery of divergent trends in body size increase over time on different continents. Lintulaakso and Kovarovic’s findings show the importance of continent-scale vegetation, climate, and history in shaping how modern tropical mammal ecomorphology correlates with different habitats. These chapters show both the potential for NOW-based research to reveal how ecomorphic patterns play out over deep time

Foreword

Foreword

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and also the need for more and better ecological trait information for extinct taxa in order to realize this potential. Applying NOW data to questions of diversification, species durations, and competitive exclusion is explored in chapters on Pakistan Siwalik rodents (Flynn and colleagues), small mammals (Van den Hoek Ostende and colleagues, Samuels and Schap), large mammals (Galbrun and colleagues), and horses (Janis). Two of these investigations are at a regional to continental scale while the others take an intercontinental approach to teasing out global-scale patterns. For the Siwalik record of southern Asia, Flynn and colleagues show how fine-scale (105 yr) temporal resolution and refined taxonomy can amplify patterns of rodent metacommunity change that support competitive replacement. Samuels and Schap use NOW and other North American data sources to reveal varying responses of rodents and lagomorphs to abiotic drivers, with some ecomorphic traits emerging earlier in different parts of the continent. Van den Hoek Ostende and colleagues compare evolutionary patterns in three distinct groups of small mammals on different continents and propose the hypothesis of “fast lane and slow lane evolution,” correlated with species longevities. They further suggest that competition was an important long-term driver of diversification in these groups. A different approach to testing for competition in community structure by Galbrun and colleagues uses large herbivores and carnivores and NOW data on co-occurrence of con-generic taxa at fossil localities in Europe and North America. Janis touches on many potential drivers of diversification in her analysis of the evolution and ecomorphology of Equinae in the New and Old World, including marked differences in species richness at single fossil localities in North America versus Eurasia. In these chapters as well as others, paleontological data from both small and large mammals is contributing new insights for a broad-scale understanding of evolutionary patterns. For any databasing effort, the bottom line for success is whether the data are used by the scientific community. NOW has proved its worth through the steady growth of on-line users and publications, and this book adds many more thought-provoking contributions to build an understanding of Cenozoic mammalian evolution. As a dynamic ecosystem of people, data, and ideas, the NOW Community is designed to be resilient and long-lived. Perhaps NOW’s most outstanding “character trait” is the dynamic relationship between substance, i.e., facts about past life, and the dedication of its contributors—the essential data coordinators. Remarkably, this dynamic has been sustained over four decades and has gained a permanent home at the Natural History Museum of Helsinki. This promises a bright future for database growth and more scientific discoveries. Other databases, perhaps similarly idealistic and well-conceived to begin with, have faltered and gone extinct. The success of NOW is a tribute to the leadership of Mikael Fortelius and the trust and support he has received from institutions, granting agencies, private benefactors, and far-flung colleagues willing to join the NOW Community. Their combined curiosity about the past, which powered a vision for what NOW could be and was sustained through good times and bad, has given us a global-scale “digital museum” and examples of what such data can do, enjoyably portrayed in the pages of this milestone volume. Anna K. Behrensmeyer National Museum of Natural History Smithsonian Institution Washington, DC, USA

Preface

Dear Reader, In front of you lies a compilation of studies brought together on the occasion of the 25th anniversary of the New and Old Worlds Database. It is a testimony of what NOW has become. What started as a small joint initiative has turned into a thriving community in which mammal paleontologists from all over the world join to reveal the secrets of the past using a collective database. Of course, an anniversary is just a date, but 25 years is a respectable age for a database. A milestone. And just as a milestone marks the distance traveled, so is this book intended to give an impression of where we are. But you cannot consider what you have reached without knowing where you came from or pondering the road on which you travel. It may be no surprise that chapters on the history of the NOW and on the database itself were provided by the beating heart of the NOW community, the Helsinki team that started the initiative and coordinated the building and maintenance of the database. For many years the name of Mikael Fortelius was all but synonymous with the NOW database. Certainly, the present volume would never have seen the light of day if it wasn’t for his dedication, and especially his continuous effort to seek collaboration and cooperation. In short, in creating a true community. The rest of the book is filled by the community itself. When the decision was made to create a celebratory volume, we made a call for contributions. The response was overwhelming, and we were very sorry that we had to turn down several very promising offers. What remains is an array of studies with vastly different subjects. Some with a clearly paleoecological focus, which was very much the original purpose of the database, but others deal with the evolutionary process. Some studies are regional in nature, others focus on a single group, but there are also studies that cover the entire planet. Looking at all these studies and their authors, it is also evident that NOW is a global community, with scientists from far and wide joining in this celebration. This is also reflected in our editorial team, consisting of a Catalan, a Dutchman, a Brit, and a Finn. Writing this preface, it is already hard to realize that the contributions were written in a difficult period when the COVID-19 pandemic forced academics from all over the world to work from home. We are grateful for all their perseverance that helped bring this project to an end. Whereas we know all about writing scientific papers, we have little to no experience in turning them into a book. We were very happy to find Springer interested in joining the venture. In particular the book series “Vertebrate Paleobiology and Paleoanthropology” seems a very apt place for the NOW anniversary volume. It demonstrates that the NOW community is part of a far larger global community, which shares the love for the mysteries of the past and the way in which the rich record of vertebrate fossils can help us understand the history of vertebrate biodiversity and indeed that of our planet itself. It turned into a very pleasant cooperation with what was soon called “The Erics”, both always more than willing to help with any issues that arose during the project. A good book needs a good cover, and we were

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Preface

delighted that our own Óscar Sanisidro, a longstanding and valued member of the NOW community, agreed to have his artwork used for this purpose. We are grateful to Kay Behrensmeyer for crafting a wonderful, insightful prologue to this volume. Although ours was a modest contribution alongside the hard work of the actual contributors, we are very proud of the result. It has become a book worthy of the community it represents. It is a work that those pioneers that started the database could probably only dream of. But their database turned into not only a useful and reliable tool, but also a versatile instrument for scientists. Some may access the NOW database (https://nowdatabase.org/) only to find a first occurrence to help in calibrating a molecular phylogeny. Others use the database as a starting point for their elaborate analyses. We hope that you will enjoy the examples given in this book and will get a good impression of what the possibilities of the NOW database are. But most of all, we hope that you will be inspired by your own studies. The NOW database is there, being continuously improved and updated. And we anticipate that it will be there for many years to come to the benefit of all who want to join in. Barcelona, Spain Leiden, The Netherlands Bristol, UK Helsinki, Finland

Isaac Casanovas-Vilar Lars W. van den Hoek Ostende Christine M. Janis Juha Saarinen

Contents

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Introduction: Then, NOW and Beyond . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isaac Casanovas-Vilar, Lars W. van den Hoek Ostende, Christine M. Janis, and Juha Saarinen

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The Origin and Early History of NOW as It Happened . . . . . . . . . . . . . . . . Mikael Fortelius, Jordi Agustí, Raymond L. Bernor, Hans de Bruijn, Jan A. van Dam, John Damuth, Jussi T. Eronen, Gudrun Evans, Lars W. van den Hoek Ostende, Christine M. Janis, Jukka Jernvall, Anu Kaakinen, Wighart von Koenigswald, Kari Lintulaakso, Liping Liu, Majid Mirzaie Ataabadi, Hans-Walter Mittmann, Diana Pushkina, Juha Saarinen, Sevket Sen, Susanna Sova, Laura K. Säilä, Alexey Tesakov, Jouni Vepsäläinen, Suvi Viranta, Innessa Vislobokova, Lars Werdelin, Zhaoqun Zhang, and Indrė Žliobaitė

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The NOW Database of Fossil Mammals . . . . . . . . . . . . . . . . . . . . . . . . . . . Indrė Žliobaitė, Mikael Fortelius, Raymond L. Bernor, Lars W. van den Hoek Ostende, Christine M. Janis, Kari Lintulaakso, Laura K. Säilä, Lars Werdelin, Isaac Casanovas-Vilar, Darin A. Croft, Lawrence J. Flynn, Samantha S. B. Hopkins, Anu Kaakinen, László Kordos, Dimitris S. Kostopoulos, Luca Pandolfi, John Rowan, Alexey Tesakov, Innessa Vislobokova, Zhaoqun Zhang, Manuela Aiglstorfer, David M. Alba, Michelle Arnal, Pierre-Olivier Antoine, Miriam Belmaker, Melike Bilgin, Jean-Renaud Boisserie, Matthew R. Borths, Siobhán B. Cooke, Jan A. van Dam, Eric Delson, Jussi T. Eronen, David Fox, Anthony R. Friscia, Marc Furió, Ioannis X. Giaourtsakis, Luke Holbrook, John Hunter, Sergi López-Torres, Joshua Ludtke, Raef Minwer-Barakat, Jan van der Made, Bastien Mennecart, Diana Pushkina, Lorenzo Rook, Juha Saarinen, Joshua X. Samuels, William Sanders, Mary T. Silcox, and Jouni Vepsäläinen

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Siwalik Rodent Assemblages for NOW: Biostratigraphic Resolution in the Neogene of South Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lawrence J. Flynn, Michèle E. Morgan, John C. Barry, S. Mahmood Raza, Iqbal Umer Cheema, and David Pilbeam

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Comparing the Evolution of the Extinct, Endemic Carnivorous Mammals of South America and Africa (Sparassodonts and Hyaenodonts). . . . . . . . . . Anthony R. Friscia, Matthew R. Borths, and Darin A. Croft

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Contents

Environmental Change and Body Size Evolution in Neogene Large Mammals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shan Huang, Alison Eyres, Susanne A. Fritz, Jussi T. Eronen, and Juha Saarinen

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Continent-Wide Patterns in Mammal Community Structure: Diet, Locomotion, and Body Mass. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kari Lintulaakso and Kris Kovarovic

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Live Long and Prosper? Assessing Longevity of Small Mammal Taxa Using the NOW Database . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lars W. van den Hoek Ostende, Melike Bilgin, Yanell Braumuller, Florentin Cailleux, and Panagiotis Skandalos Patterns of Competitive Exclusion in the Mammalian Fossil Record . . . . . . . Esther Galbrun, Jo Skeie Hermansen, and Indrė Žliobaitė

10 Asymmetry of Evolutionary Patterns Between New World and Old World Equids and Among New World Equine Tribes . . . . . . . . . . . . . . . . . Christine M. Janis 11 Musk Deer on the Run – Dispersal of Miocene Moschidae in the Context of Environmental Changes . . . . . . . . . . . . . . . . . . . . . . . . . . Manuela Aiglstorfer, Israel M. Sánchez, Shi-Qi Wang, Jorge Morales, Chunxiao Li, Serdar Mayda, Loïc Costeur, Elmar P. Heizmann, and Bastien Mennecart 12 A Reassessment of Some Giraffidae Specimens from the Late Miocene Faunas of Eurasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nikos Solounias and Advait M. Jukar 13 Regional Topography and Climate Influence the Nature and Timing of Changes in the Structure of Rodent and Lagomorph Faunas Through the Cenozoic of North America . . . . . . . . . . . . . . . . . . . . . . . . . . . Joshua X. Samuels and Julia A. Schap

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General Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Taxonomic Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Contributors

Note * indicates preferred address for correspondence Jordi Agustí, Institut Català de Paleoecologia Humana i Evolució Social, Universitat Rovira i Virgili, Tarragona, Spain Manuela Aiglstorfer, Naturhistorisches Museum Mainz, Landessammlung für Naturkunde Rheinland-Pfalz, Reichklarastr, Mainz, Germany David M. Alba, Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, c/ Columnes s/n, Campus de la UAB, Cerdanyola del Vallès, Barcelona, Spain Pierre-Olivier Antoine, Institut des Sciences de l’Evolution de Montpellier, CC64, Université de Montpellier, CNRS, IRD, EPHE, Montpellier, France Michelle Arnal, Vertebrate Palaeontology Department, La Plata Museum, National University of La Plata, Argentina Majid Mirzaie Ataabadi, Department of Geology, Faculty of Science, University of Zanjan, Zanjan, Iran John C. Barry, Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA Miriam Belmaker, Department of Anthropology, The University of Tulsa, 800 South Tucker Drive, Tulsa, OK, USA Raymond L. Bernor, *College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, Howard University, 520 W St. N.W, Washington DC, USA; Human Origins Program, Department of Anthropology, Smithsonian Institution, Washington, DC, USA Melike Bilgin, Naturalis Biodiversity Center, Leiden, The Netherlands; *Department of Geology and Palaeontology, Comenius University, Bratislava, Slovakia Jean-Renaud Boisserie, *Laboratory Paleontology Evolution Paleoecosystems Paleoprimatology (PalEvOPrim), CNRS & Université de Poitiers, Poitiers, France; French Centre for Ethiopian Studies (CFEE), CNRS & Ministère de l'Europe et des affaires étrangères, Addis Ababa, Ethiopia Matthew R. Borths, Division of Fossil Primates, Duke Lemur Center, Durham, NC, USA Yanell Braumuller, Naturalis Biodiversity Center, Leiden, The Netherlands Hans de Bruijn (deceased), Department of Earth Sciences, University of Utrecht, The Netherlands Florentin Cailleux, *Department of Geology and Palaeontology, Comenius University, Bratislava, Slovakia; Naturalis Biodiversity Center, Leiden, The Netherlands

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Isaac Casanovas-Vilar, Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, c/ Columnes s/n, Campus de la UAB, Cerdanyola del Vallès, Barcelona, Spain Iqbal Umer Cheema, Islamabad, Pakistan Siobhán B. Cooke, Center for Functional Anatomy and Evolution, Johns Hopkins University, School of Medicine, Baltimore, MD, USA Loïc Costeur, Naturhistorisches Museum Basel, Basel, Switzerland Darin A. Croft, Department of Anatomy, Case Western Reserve University School of Medicine, Cleveland, OH, USA Jan A. van Dam, *Department of Earth Sciences, Faculty of Geosciences, Utrecht University, Utrecht, The Netherlands; Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain John Damuth, Department of Ecology, Evolution & Marine Biology, University of California, Santa Barbara, CA, USA Eric Delson, *Division of Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY, USA; Anthropology, Lehman College and the Graduate Center, The City University of New York, New York, NY, USA; Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain; NYCEP (New York Consortium in Evolutionary Primatology), New York, NY, USA Jussi T. Eronen, *Ecosystems and Environment Research Programme & Helsinki Institute of Sustainability Science (HELSUS), Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki, Finland; BIOS Research Unit, Helsinki, Finland Gudrun Evans, Freelance, formerly Institute of Biotechnology, University of Helsinki, Helsinki, Finland Alison Eyres, Department of Zoology, University of Cambridge, Cambridge, UK Lawrence J. Flynn, Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA Mikael Fortelius, *Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland; Finnish Museum of Natural History LUOMUS, University of Helsinki, Helsinki, Finland David Fox, Department of Earth & Environmental Sciences, University of Minnesota, 116 Church St SE, Minneapolis, MN, USA Anthony R. Friscia, Department of Integrative Biology & Physiology, University of California, Los Angeles, CA, USA Susanne A. Fritz, *Senckenberg Biodiversity and Climate Research Centre (SBiK-F), Frankfurt Am Main, Germany; Institut für Geowissenschaften, Goethe-Universität Frankfurt, Frankfurt Am Main, Germany Marc Furió, *Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain; Department of Geology, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain Esther Galbrun, School of Computing, University of Eastern Finland, Technopolis, Kuopio, Finland

Contributors

Contributors

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Ioannis X. Giaourtsakis, Department of Earth and Environmental Sciences, Section of Paleontology & Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany Elmar P. Heizmann, Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany Jo Skeie Hermansen, Centre for Ecological and Evolutionary Synthesis, University of Oslo, Oslo, Norway Lars W. van den Hoek Ostende, Naturalis Biodiversity Center, Leiden, The Netherlands Luke Holbrook, Department of Biological Sciences and Biomedical Sciences, Rowan University, Glassboro, USA Samantha S. B. Hopkins, Department of Earth Sciences, University of Oregon, Eugene, OR, USA Shan Huang, School of Geography, Earth and Environmental Sciences, University of Birmingham, Birmingham, UK John Hunter, Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Columbus, OH, USA Christine M. Janis, *Bristol Palaeobiology Group, School of Earth Sciences, University of Bristol, Bristol, UK; Department of Ecology and Evolutionary Biology, Brown University, Providence, RI, USA Jukka Jernvall, *Institute of Biotechnology, University of Helsinki, Helsinki, Finland; Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Advait M. Jukar, *Department of Geosciences, University of Arizona, Tucson, AZ, USA; Division of Vertebrate Paleontology, Yale Peabody Museum, New Haven, CT, USA; Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA Anu Kaakinen, Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Wighart von Koenigswald, Steinmann Institut für Geologie, Mineralogie und Paläontologie, Bereich Paläontologie, Bonn, Germany László Kordos, Eötvös Loránd University, Szombathely, Hungary Dimitris S. Kostopoulos, Faculty of Sciences, School of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece Kris Kovarovic, Department of Anthropology, Durham University, Durham, UK Chunxiao Li, University of Chinese Academy of Sciences, Beijing, China Kari Lintulaakso, Finnish Museum of Natural History LUOMUS, University of Helsinki, Helsinki, Finland Liping Liu, *Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland; Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden Sergi López-Torres, *Institute of Evolutionary Biology, Faculty of Biology, University of Warsaw, Warsaw, Poland; Division of Paleontology, American Museum of Natural History, New York, NY, USA; NYCEP (New York Consortium in Evolutionary Primatology), New York, NY, USA Joshua Ludtke, Department of Biological Sciences, San Diego Miramar College, San Diego, CA, USA Jan van der Made, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (CSIC), Madrid, Spain

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Serdar Mayda, *Faculty of Science, Biology Department, Ege University, Bornova, Izmir, Turkey; Natural History Research & Application Center, Ege University, Bornova, Izmir, Turkey Bastien Mennecart, Natural History Museum Basel, Basel, Switzerland Raef Minwer-Barakat, *Departamento de Estratigrafía y Paleontología, Universidad de Granada, Granada, Spain; Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain Hans-Walter Mittmann, Retired, formerly Staatliches Museum für Naturkunde, Karlsruhe, Germany Jorge Morales, Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (CSIC), Madrid, Spain Michèle E. Morgan, Peabody Museum of Archaeology and Ethnology, Harvard University, Cambridge, MA, USA Luca Pandolfi, Dipartimento di Scienze, Università della Basilicata, Potenza, Italy David Pilbeam, Department of Human Evolutionary Biology, Harvard University, Cambridge, MA, USA Diana Pushkina, Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland S. Mahmood Raza, National University of Medical Sciences, The Mall, Rawalpindi, Pakistan Lorenzo Rook, Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Firenze, Italy John Rowan, Department of Anthropology, University at Albany, Washington Avenue, Albany, NY, USA Juha Saarinen, Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Laura K. Säilä, Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Joshua X. Samuels, *Department of Geosciences, East Tennessee State University, Johnson City, TN, USA; Don Sundquist Center of Excellence in Paleontology, East Tennessee State University, Johnson City, TN Israel M. Sánchez, Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain William Sanders, Museum of Paleontology, University of Michigan, Ann Arbor, MI, USA Julia A. Schap, Spatial Ecology and Paleontology Lab, School of Biological Sciences, North Avenue, Georgia Institute of Technology, Atlanta, GA Sevket Sen, Laboratoire de Paléontologie du Muséum, CR2P-CNRS-UPMC, 8 rue Buffon, Paris, France Mary T. Silcox, Department of Anthropology, University of Toronto Scarborough, Military Trail, Toronto ON, Canada Panagiotis Skandalos, Naturalis Biodiversity Center, Leiden, The Netherlands

Contributors

Contributors

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Nikos Solounias, *Department of Anatomy, New York Institute of Technology, College of Osteopathic Medicine, Old Westbury, NY, USA; Division of Paleontology, American Museum of Natural History, New York, NY, USA Susanna Sova, *Institute of Biotechnology, University of Helsinki, Helsinki, Finland; Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland Alexey Tesakov, Geological Institute, Russian Academy of Sciences, Moscow, Russia Jouni Vepsäläinen, Department of Computer Science, University of Helsinki, Helsinki, Finland Suvi Virantana, Department of Anatomy, Faculty of Medicine, University of Helsinki, Helsinki, Finland Innessa Vislobokova, Borissiak Paleontological Institute, Russian Academy of Sciences, Moscow, Russia Shi-Qi Wang, *Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China Lars Werdelin, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden Zhaoqun Zhang, Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China Indrė Žliobaitė, *Department of Computer Science, University of Helsinki, Helsinki, Finland; Finnish Museum of Natural History LUOMUS, University of Helsinki, Helsinki, Finland; Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland

Chapter 1

Introduction: Then, NOW and Beyond Isaac Casanovas-Vilar, Lars W. van den Hoek Ostende, Christine M. Janis, and Juha Saarinen

Abstract The NOW database of fossil mammals was released in 1996, slightly more than 25 years ago. Initially the acronym stood for ‘Neogene Old World’, but as the database later expanded to include data for Cenozoic mammals worldwide it was changed to ‘New and Old Worlds’. NOW is a comprehensive research database and its creation approximately coincided with that of several other similar databases, many of which have been discontinued. A clear policy for data preservation and management as well as to ensure data access has been key to the success of NOW database. This volume celebrates this enduring history with several contributions that are briefly introduced in this chapter. These include chapters on the development and history of NOW, on its contents and management, or that make use of NOW data to address particular research questions from paleoecology and paleobiogeography to macroevolution.











Keywords Paleontological databases History of paleontology Fossil mammals Fossil record Cenozoic

I. Casanovas-Vilar (&) Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, c/Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain e-mail: [email protected] L. W. van den Hoek Ostende Naturalis Biodiversity Center, P.O. Box 95172300 RA Leiden, The Netherlands C. M. Janis Bristol Palaeobiology Group, School of Earth Sciences, University of Bristol, Bristol, BS8 1RL, UK Department of Ecology and Evolutionary Biology, Brown University, Box G-W, Providence, RI 02912, USA J. Saarinen Department of Geosciences and Geography, University of Helsinki, P.O. Box 6400014 Helsinki, Finland

“Data! Data! Data! [...] I can’t make bricks without clay.” Sherlock Holmes in The Adventure of Copper Beeches (Arthur Conan Doyle, 1892)

It is 1996. Spice Girl’s “Wanabe”, Oasis’s “Wonderwall” and (yes) Los del Río’s “Macarena” hit the radio stations, the sitcom “Friends” becomes a popular phenomenon, and the ancestors of our mobile phones start spreading worldwide. There is even something called the ‘internet’, a global system of interconnected computer networks that is used by less than 1% of the world population (data from United Nations, 2022). There is no Wi-Fi so you connect to the internet using a modem, which sounds like a tortured droid and achieves really low connection speed. The latest science news is about Dolly the sheep, the first cloned mammal, and Deep Blue, the computer that beat chess grand-master Garry Kasparov. This is the world when the NOW database was born, more than 25 years ago. The NOW database on fossil mammals was released in December 1996, on the same day the book The Evolution of Eurasian Neogene Mammal Faunas (Bernor et al., 1996) was published, and the first web-based interface was launched in 1999. Initially, the acronym ‘NOW’ referred to the ‘Neogene of the Old World’ but since the database later expanded to include data from all Cenozoic mammals worldwide it was officially changed to ‘New and Old Worlds’ in 2012. The origin and colorful history of NOW is covered in first person by its protagonists in Chap. 2 (Fortelius et al., 2023) so we do not want to introduce spoilers here. Instead, our purpose is to stress what makes the NOW database special. When did paleontological databases start? We tend to associate them with computers, but they are in fact much older, as old as paleontology itself. Indeed, museums and other entities have collection databases, typically listing the specimens and associated information under their care, and many collection catalogs are as old as the museums

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 I. Casanovas-Vilar et al. (eds.), Evolution of Cenozoic Land Mammal Faunas and Ecosystems: 25 Years of the NOW Database of Fossil Mammals, Vertebrate Paleobiology and Paleoanthropology, https://doi.org/10.1007/978-3-031-17491-9_1

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themselves. However, these must be distinguished from research databases; that is, those created to address particular research questions (Uhen et al., 2013). Today, these are electronic databases but they were preceded during more than 100 years by regular printed compilations in the form of books, tables or card catalogs. These started in 1843 with the compilation of British fossils by Morris (1843) and include for example the global compendia by Harland et al. (1967) and Sepkoski (1982) (for a review of the early history of databases see Benton, 1999). Indeed, such exhaustive and authoritative compendia continue to be published and are often important data sources for electronic databases, see for example the two volumes of the Evolution of Tertiary Mammals of North America (Janis et al., 1998, 2008), the Cenozoic Mammals of Africa (Werdelin & Sanders, 2010) or Fossil Mammals of Asia (Wang et al., 2013). The switch of NOW from an Old World to a global database was marked by the incorporation of the data from Janis et al. (1998, 2008) in 2012. As personal computers became commonplace by the late 1970s and 1980s so did electronic databases: collection management databases came first and were followed in the late 1980s and early 1990s by the first electronic research databases, such as FAUNMAP (FAUNMAP Working Group, 1994). These first research databases were distributed on physical media using back up tapes or floppy disks (Uhen et al., 2013). The next major step would not come until the internet era, by the late 1990s and early 2000s, when several databases were launched online, including NOW. The list of paleontological databases (and acronyms) is long and includes databases that record occurrences of taxa in sites or regions, such as NOW and also the popular Paleobiology Database (https://paleobiodb.org/#/) and the Neotoma Paleoecology Database (https://www.neotomadb.org/), for example. Others, such as MorphoSource (https://www. morphosource.org/), are repositories of 3D and 2D data and images of fossil and extant specimens. There was an initial ‘boom’ in paleontological databases by the late 1990s and early 2000s, and while many of these are still active or have been integrated into larger databases, others have ultimately vanished and are now extinct (see Uhen et al., 2013). The NOW database is one of the survivors and could be considered a long-ranging taxon, but it is certainly not a living fossil. Uhen et al. (2013) remark that many defunct databases were victims of the lack of a long-term strategy for data preservation, so in many cases there were no well-defined policies to ensure the data would remain available, even less so interactive, in the future. Another cause may be that databases were led by principal investigators’ grant projects, and once those ended or the investigators retired, they were abandoned. This has not happened to NOW and is unlikely that it will happen in the future. It is true that the database is

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mostly the brainchild of Mikael Fortelius, but from the very beginning there has been a clear policy for data preservation and management as well as the means to ensure data access (see Chap. 2 by Fortelius et al., 2023, and Chap. 3 by Žliobaitė et al., 2023). Clearly, great care was put into creating and maintaining the database. Maybe this was partly because the NOW database was initially developed in Finland, a country which is basically devoid of fossils. Its geology is a mixture of very old Precambrian rocks of the Fennoscandian Shield and Quaternary glacial deposits. During the last ice age, the ice sheet that covered Finland was huge and indeed glaciers caused intense erosion, stripping clean all the mainland from sedimentary rocks down to the Precambrian basement (Johansson et al., 2011; Saarnisto & Salonen, 1995). The Dutch football guru Johan Cruyff famously remarked that every disadvantage has its advantage. Not having fossils in their own country provided Finnish paleontologists, who had to travel elsewhere to study fossils, with a wide international network and a cosmopolitan view of the fossil record. This was particularly useful when in the 1990s the realization dawned that anthropogenic influence was changing our planet and Global Change became a focal point in science. Paleontologists turned their attention to large scale environmental changes. Databases were the tool to deal with these issues and Finnish colleagues were in an ideal position to organize the international effort. The international character of NOW markedly contrasts with that of other databases of its time. Take for example FAUNMAP (FAUNMAP Working Group, 1994), which was developed by and aimed at researchers living in a fossil-rich country, the United States, and only considered ‘national’ fossils. But on the other side of the Atlantic, an international scope was needed right from the beginning. Therefore, the first step was assembling an international panel of contributors and advisors (see Fortelius et al. 2023, Appendix 2.1) who would take care of contributing, reviewing, and updating the data. Initially, paleoecology was very much at the forefront of the minds of the group taking the initiative for the new database. For them, it was clear that building a database not only involved getting more and more data, but also ensuring consistency so that these data could be used in large scale analyses. The NOW database quickly became a scientific community and a huge online ‘museum’ of data. The same as a physical museum, there were data curators, specialists (the members of the advisory board) undertaking research on the ‘collections’, and even strict protocols regarding the preservation and management of the data heritage. The foundations for a huge online data museum had now been established. More than 25 years after its creation we may say that what makes NOW enduring and special is this, perhaps unintended, museum-like approach. Those of us who contribute to NOW and also manage museum collections find

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that, except for the lack of physical fossils to work with, there is not much difference. However, the value of NOW is not only storing and curation of the data, but also making them available to everyone interested through the internet. Contemporary paleontology, as for other sciences, is no longer an individualistic activity but a group effort. Scientists rely on each other for results, peer-review, and many other activities, including data sharing. Indeed, most journals now require that all the data used in a study are immediately available, either by their inclusion in the publication or by posting them on online repositories. From the very beginning NOW was envisioned as a public access database and all data are freely available for browsing and downloading (except for those data that are being uploaded or reviewed). This data democratization, one of the core principles of NOW, is now commonplace, but was not so much during the late 1990s and early 2000s. The NOW database was ahead of its time in this crucial aspect. The time of the writing of this introduction is 2022. Both the Spice Girls and Oasis disbanded long ago. Traditional television is being replaced by subscription streaming platforms where you can still watch ‘Friends’, now regarded as a sitcom classic. More than half of the world population (more than 80% in Europe, the United States and Australia; data from United Nations, 2022) has high-speed internet connection at their homes, working places or smartphones. This has really transformed our lives and society. The world has changed a lot, and NOW has come of age and keeps growing, evolving with changing times. In 2018, Mikael Fortelius was eventually succeeded as general coordination of the NOW database by Indrė Žliobaitė, initially a computer scientist who took her interest in paleontology too far, and has already led a spectacular renovation of NOW considering the short time she has been in charge. You can now access to it from almost anywhere in the world if you have a smartphone and you will find much information that was not there in 1996. Yet, the spirit, illusion and ideals that promoted its creation are there and have engaged new generations of paleontologists forming a strong community. We shall have NOW for years to come. This volume This volume celebrates these slightly more than 25 years of history of the NOW database. As such, it contains various contributions that either focus on the database, are based on NOW data (or on data that will soon be uploaded), or discuss subjects directly related to the NOW community research interests. It can be divided into two parts. The first one comprises Chaps. 2 and 3 and deals with the history, purpose, content, and management of the NOW database. The much longer second part, comprising Chaps. 4 to 13, provides various case studies mostly based on NOW data

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covering a wide range of topics, from paleoecology and paleobiogeography to macroevolution. Chapter 2, by Fortelius et al. (2023), tells the story of the origin and early years of the NOW database firsthand. Early concepts date back to the 1980s, although the database did not take shape and was not publicly released until 1996. The early story of NOW follows, first as a node of the ETE (Evolution of Terrestrial Ecosystems) database of the Smithsonian Institution to the launch of the first standalone online version in 2005. Originally set to include data only from the Old World, it became a global database in 2012, encompassing data on mammals from all continents and all the Cenozoic. Also, research based on NOW data grew from the initial goals of paleoecology and paleoenvironmental reconstruction to give birth to ecometrics and include macroevolutionary studies as well. At the same time, the NOW community grew as well to include specialists from all over the world and even computer scientists. All this colorful story is explained in detail in Chap. 2, with reference to all the main characters, unsuspected dramatic turns, old pictures, and the right amount of humor. Chapter 3, by Žliobaitė and most of the NOW advisory board (2023), briefly explains what the NOW database is, what are the data included and how it works. It also pays special attention to the quality and biases inherent to the fossil record and clearly explains how data are entered into NOW, reviewed, curated, and constantly updated. Finally, there are also specific sections explaining the information that can be found in the database that goes beyond species-by-locality tables and includes data on age, lithology, taphonomy and paleoenvironment (among other things). Considering that NOW started at a time when one could not foresee current developments, the database is surprisingly well equipped to meet contemporary challenges. Chapter 4 by Flynn et al. (2023) deals with the rich mammal faunas from the Siwaliks of the Potwar Plateau, Pakistan, which are in the process of being uploaded to NOW. After decades of research, the Siwaliks stand out as one of the best-known Neogene terrestrial mammal successions worldwide, including thousands of fossils coming from more than 300 different localities, most of which have been tightly dated to a precision of 100,000 years. Rodents stand out as one of the most abundant and well-studied groups and are the subject of this chapter. Flynn et al. (2023) consider the occurrences and relative abundances of Siwaliks rodent species during the Middle and Late Miocene and recognize successive characteristic metacommunities dominated by different murine species. These communities temporally replace one another, metacommunity turnover being consistent with their hypothesis of competitive replacement among muroid rodents. In Chap. 5, Friscia et al. (2023) examine evolutionary patterns and constraints in two extinct clades of

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hypercarnivorous mammals, the African creodonts (Eutheria) and the South American sparassodonts (Metatheria). The comparison, which focusses on the carnassialized molars, offers insights into constraints on body size evolution (deduced from molar size) and ecological specializations (deduced from carnassial blade lengths). While both clades showed trends to increased carnivory over time, it was mainly the hyaenodontids that coupled this with an increase in body size. In both clades the carnassial complex occupies each molar position, in contrast with extant carnivorans, and this morphology may have resulted in greater specialization leading to greater risk of extinction. In Chap. 6, Huang et al. (2023) explore how climate and environmental conditions affected body size within the key orders of large terrestrial mammals, Proboscidea, Artiodactyla, Perissodactyla, Primates and Carnivora, during the Neogene. They show that mean, maximum and minimum size of large terrestrial mammals, with Proboscidea as a newly analyzed case, track changes in global mean temperature during the Neogene, with larger sizes occurring during times of global cooling and subsequently increased seasonality and mid-latitude drying (“harsh” conditions in general). Nonetheless, there are also differences in the body size patterns of large mammals on different continents that reflect different patterns of biome change on separate continents from the Miocene to the Pliocene. While average body size of browsing and grazing ungulates increased significantly from Early Miocene to Pliocene in the increasingly open and dry biomes in North America, mean body size remained more constant in Europe where warm-temperate forests remained the dominant biome type. In Chap. 7, Lintulaakso and Kovarovic (2023) use mammalian community properties (species diets, body size and locomotion type) to distinguish vegetation types in modern tropical ecosystems in America, Africa and Asia. They found that dietary categories are the best at differentiating between vegetation types, followed by locomotor categories. Body size also gives a moderate signal of vegetation in America but not in Africa. In general, the models are not able to significantly differentiate between the more limited range of vegetation types in tropical Asia, but for the wider range of tropical vegetation types in America and Africa such distinctions based on mammalian community structure are significant. In Chap. 8, Van den Hoek Ostende et al. (2023) wonder for how long various taxa are on Earth and what determines their duration. Their dataset comprises the rodents, insectivores and lagomorphs. There appears to be a relationship between the diversity of a group and the longevity of its included genera or species. The shortest longevities are, for instance, found in rodents, the most diverse mammal order and within that order in the murids, by far its most diverse family. The authors attribute this pattern to higher

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competition with a group leading to Red Queen dynamics, resulting in a shorter duration of (chrono-) species. Galbrun et al. (2023) also focus on competition-based patterns in Chap. 9. Related species are more likely to be in competition with one another and this would particularly hold true for congenerics. The authors try to find empirical evidence for the hypothesis of competitive congenerics exclusion by looking at various subsets in the NOW database. Indeed, the co-occurrence of congenerics within a mammal assemblage is lower than expected from a random distribution. This pattern is most clear in the large herbivores, but also present in the carnivorans and small mammals. In Chap. 10, Janis (2023) considers other aspects of equid evolution besides the “progression” to modern day Equus. In North America the three tribes within the Equinae – Equini (containing Equus), Hipparionini, and Protohippini – all increased in hypsodonty during the Neogene. The Equini tended to be the largest in body size, but were never highly species diverse, especially in comparison with the Hipparionini. Only in the Plio-Pleistocene did the Equini become the dominant, and later the sole, equid tribe. Comparison with patterns in the Old World shows that during the Late Miocene the number of equid species per fossil locality was two to three times greater in North America than in the Old World. In Chap. 11, Aiglstorfer et al. (2023) investigate the effects of climatic and environmental changes on the patterns of dispersal of moschids, small hornless ruminants that survive today as the Asian musk deer, Moschus. They first review the current knowledge about Miocene moschids, including their phylogenetic position within the Ruminantia, and show that many taxa previously considered to be moschids do not belong in the family. Secondly, they show that the spread of moschids across the Old World northern hemisphere, from Asia into Europe, can be correlated with the onset of cooler, more seasonal, and more arid climatic conditions; moschids were especially impacted by the Middle Miocene Climatic Transition. This contrasts with the common view that moschids in the fossil record are indicators of closed and humid environments. Chapter 12 by Solounias and Jukar (2023) puts into value systematic studies and the revision of old museum collections in these times of paleontological databases. The authors review various giraffid specimens from the Late Miocene of Pakistan (Siwaliks), Iran (Maragheh) and Greece (Pikermi and Samos). This allows reporting several new occurrences for various genera and refining some identifications. Finally, the implications of these results for Eurasian giraffid diversity and paleobiogeography are thoroughly discussed. Chapter 13 by Samuels and Schap (2023) deals with fossil small mammals, this time North American rodents and

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lagomorphs through the Cenozoic. The authors analyze the turnover in rodent and lagomorph faunas related to the spread of open and arid habitats at a regional scale which contrasts with previous studies that had considered a wider continental scale. Their results show asynchronous changes across North America, with specialized dietary (hypsodonty) and locomotor (jumping, burrowing) adaptations occurring earlier in the relatively cooler and more arid regions at higher latitudes. The authors further infer that topographic complexity and volcanic activity were key factors pushing ecomorphological evolution in rodents and lagomorphs in those areas. Later, as the climate became more arid through the Cenozoic, arid-adapted taxa would extend their range into other regions. As this volume illustrates, the NOW database provides abundant high-quality data to address various research questions. What new matters will emerge in the future? For now, the questions are open, but NOW data will be there to help answering (or perhaps generating) them. Acknowledgments This publication is part of R+D+I project PID2020-117289GB-I00, funded by MCIN/AEI/https://doi.org/10. 13039/501100011033/. I. C.-V. is also supported by CERCA Programme/Generalitat de Catalunya and is member of consolidated research group 2022 SGR 00620 of the Generalitat de Catalunya. J.S. worked at the University of Helsinki funded by the Academy of Finland during this work (AoF. projects nr. 315691 and 340775).

References Aiglstorfer, M., Sánchez, I. M., Wang, S.-Q., Morales, J., Li, C., Mayda, S., et al. (2023). Musk deer on the run—Dispersal of Miocene Moschidae in the context of environmental changes. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 165–186). Springer. Benton, M. J. (1999). The history of life: Large databases in palaeontology. In D. A. T. Harper (Ed.), Numerical palaeobiology (pp. 249–283). Wiley. Bernor, R. L., Fahlbusch, V., & Mittmann, H.-W. (Eds.). (1996). The evolution of Western Eurasian Neogene mammal faunas. Columbia University Press. Conan Doyle, A. (1892). The adventure of copper beeches. George Newness. FAUNMAP Working Group. (1994). FAUNMAP: A database documenting late Quaternary distributions of mammal species in the United States. Illinois State Museum Scientific Papers, 25, 1–690. Flynn, L. J., Morgan, M. E., Barry, J. C., Mahmood Raza, S., Cheema, I. U., & Pilbeam, D. (2023). Siwalik rodent assemblages for NOW: Biostratigraphic resolution in the Neogene of South Asia. In I. Casanovas-Vilar, L.W. Van den Hoek Ostende, C.M. Janis & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 43–58). Springer. Fortelius, M., Agustí, J., Bernor, R., de Bruijn, H., Croft, D., Damuth, J., et al. (2023). The origin and early history of NOW as it happened. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C.

5 M. Janis & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 7–32). Springer. Friscia, A. R., Borths, M. R., & Croft, D. A. (2023). Comparing the evolution of the extinct, endemic carnivorous mammals of South America and Africa (sparassodonts and hyaenodonts). In I. Casanovas-Vilar, L.W. Van den Hoek Ostende, C.M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 59–77). Springer. Galbrun, E., Hermansen, J. E., & Žliobaitė, I. (2023). Patterns of competitive exclusion in the mammalian fossil record. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 131–141). Springer. Harland, W. B. C., Holland, C. H., House, M. R., Reynolds, A. B., Rudwick, M. J. S., Satterthwaite, G. E., Tarlo, L. B. H., & Willey, E. C. (1967). The fossil record: A symposium with documentation. Geological Society of London. Huang, S., Eyres, A., Fritz, S. A., Eronen, J. T., & Saarinen, J. (2023). Environmental change and body size evolution in Neogene large mammals. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 79–93). Springer. Janis, C. M. (2023). Asymmetry of evolutionary patterns between New World and Old World equids and among New World equine tribes. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 143–164). Springer. Janis, C. M., Scott, K. M., & Jacobs, L. L. (Eds.). (1998). Evolution of Tertiary mammals of North America, Vol. 1: Terrestrial carnivores, ungulates, and ungulate-like mammals. Cambridge University Press. Janis, C. M., Gunnell, G. F., & Uhen, M. (Eds.). (2008). Evolution of Tertiary mammals of North America, Vol. 2: Small mammals, xenarthrans, and marine mammals. Cambridge University Press. Johansson, P., Lunkka, J. P., & Sarala, P. (2011). Chapter 9—The glaciation of Finland. Developments in Quaternary Sciences, 15, 105–116. Lintulaakso, K., & Kovarovic, K. (2023). Continent-wide patterns in mammal community structure: Diet, locomotion and body mass. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 95–110). Springer. Morris, J. (1843). A catalogue of British fossils: Comprising the genera and species hitherto described. John Van Voorst. Saarnisto, M., & Salonen, V. P. (1995). Glacial history of Finland. In J. Ehlers, S. Kozarski, & Ph. L. Gibbard (Eds.), Glacial deposits in Northeast Europe (pp. 3–10). Balkema. Samuels, J. X, & Schap, J. A. (2023). Regional topography and climate influence the nature and timing of changes in the structure of rodent and lagomorph faunas through the Cenozoic of North America. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 199–215). Springer Sepkoski, J. J. Jr. (1982). A compendium of fossil marine animal genera. Paleontology Research Institution. Solounias, N., & Jukar, A. M. (2023). A reassessment of some Giraffidae specimens from the late Miocene faunas of Eurasia. In I.

6 Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 187–198). Springer. Uhen, M., Barnosky, A., Bills, B., Blois, J., Carrano, M., Carrasco, M., et al. (2013). From card catalogs to computers: Databases in vertebrate paleontology. Journal of Vertebrate Paleontology, 33, 13–28. United Nations. (2022). World Telecommunication/ICT Indicators Database. Retrieved March 25, 2022, from http://data.un.org/ Data.aspx?d=ITU&f=ind1Code%3aI99H Van den Hoek Ostende, L. W., Bilgin, M., Braumuller, Y., Cailleux, F., Skandalos, P. (2023). Live long and prosper? Assessing longevity of small mammal taxa using the NOW database. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, &

I. Casanovas-Vilar et al. J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 111–129). Springer. Wang, X., Flynn, L. J., & Fortelius, M. (Eds.). (2013). Fossil mammals of Asia: Neogene biostratigraphy and chronology. Columbia University Press. Werdelin, L., & Sanders, W. J. (Eds.). (2010). Cenozoic mammals of Africa. University of California Press. Žliobaitė I., Fortelius, M., Bernor, R., Van den Hoek Ostende, L. W., Janis, C., Lintulaakso, K., et al. (2023). The NOW database of fossil mammals. In I. Casanovas-Vilar, L. W. Van den Hoek Ostende, C. M. Janis, & J. Saarinen (Eds.), Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (pp. 33–42). Springer.

Chapter 2

The Origin and Early History of NOW as It Happened Mikael Fortelius, Jordi Agustí, Raymond L. Bernor, Hans de Bruijn, Jan A. van Dam, John Damuth, Jussi T. Eronen, Gudrun Evans, Lars W. van den Hoek Ostende, Christine M. Janis, Jukka Jernvall, Anu Kaakinen, Wighart von Koenigswald, Kari Lintulaakso, Liping Liu, Majid Mirzaie Ataabadi, Hans-Walter Mittmann, Diana Pushkina, Juha Saarinen, Sevket Sen, Susanna Sova, Laura K. Säilä, Alexey Tesakov, Jouni Vepsäläinen, Suvi Viranta, Innessa Vislobokova, Lars Werdelin, Zhaoqun Zhang, and Indrė Žliobaitė

Abstract The NOW database of fossil mammals came to be through a confluence of several initiatives spanning multiple decades. The first public version of NOW database was released in 1996 and the first Advisory Board was established the year after. Originally, NOW stood for Neogene of the Old World but with the gradual expansion of the database the acronym was eventually reassigned to stand for New and Old Worlds. The structure of what would become NOW was originally cloned from the ETE database of the Smithsonian Institution and the first NOW version accessible over the internet was a node of the ETE database.

M. Fortelius (&)
J. Jernvall
A. Kaakinen
L. Liu
D. Pushkina
J. Saarinen
S. Sova
L. K. Säilä Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, 00014 Helsinki, Finland e-mail: mikael.fortelius@helsinki.fi M. Fortelius
K. Lintulaakso
I. Žliobaitė Finnish Museum of Natural History LUOMUS, University of Helsinki, P.O. Box 44, 00014 Helsinki, Finland J. Agustí Institut Català de Paleoecologia Humana i Evolució Social, Universitat Rovira i Virgili, 43002 Tarragona, Spain R. L. Bernor College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, Howard University, 520 W St. N.W, Washington DC, 20059, USA Human Origins Program, Department of Anthropology, Smithsonian Institution, PO Box 37012 Washington, DC 20013-7012, USA H. de Bruijn (deceased) Department of Earth Sciences, University of Utrecht, Utrecht, The Netherlands J. A. van Dam Department of Earth Sciences, University of Utrecht, Utrecht, The Netherlands Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Barcelona, Spain

The first standalone, online version of NOW was launched in 2005 and the first formal steering group was established in 2009. During its existence, NOW has been funded, directly or indirectly, by several organizations but fundamentally it has always been an unfunded community effort, dependent on voluntary work by the participants.












Keywords Fossil Mammal Species List Stratigraphic Age Functional Trait Ecometrics Database

J. Damuth Department of Ecology, Evolution & Marine Biology, University of California, Santa Barbara, CA 93106, USA J. T. Eronen Ecosystems and Environment Research Programme & Helsinki Institute of Sustainability Science (HELSUS), Faculty of Biological and Environmental Sciences, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland BIOS Research Unit, 00170 Helsinki, Finland G. Evans Freelance, formerly Institute of Biotechnology, University of Helsinki, Helsinki, Finland L. W. van den Hoek Ostende Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands C. M. Janis Bristol Palaeobiology Group, School of Earth Sciences, University of Bristol, Bristol, BS8 1RL, UK Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman Street, Box G-W, Providence, RI 02912, USA J. Jernvall
S. Sova Institute of Biotechnology, University of Helsinki, Biocenter 2, P.O. Box 56, 00014 Helsinki, Finland

© The Author(s) 2023 I. Casanovas-Vilar et al. (eds.), Evolution of Cenozoic Land Mammal Faunas and Ecosystems: 25 Years of the NOW Database of Fossil Mammals, Vertebrate Paleobiology and Paleoanthropology, https://doi.org/10.1007/978-3-031-17491-9_2

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Introduction The advent of computers accessible to ordinary academics very quickly generated widespread interest in systematic computational analysis of observational data that had been developing for a long time, in some disciplines for centuries. One such discipline was paleontology, where a vast archive of formal descriptions of fossils had accumulated since the late eighteenth century and even earlier, scattered over an enormous range of scientific journals and monographs from several disciplines and in many languages. One of the first attempts to harness this information for computational analysis, and arguably still the most widely known, was Jack Sepkoski’s “compendium” from the later 1970’s, of first and last occurrences of known marine metazoan taxa of the Phanerozoic (see Sepkoski, 2002). For terrestrial vertebrates an early manifestation of this trend was the Workshop on Computerization organized by the Society of Vertebrate Paleontology in 1989 and the resulting Guidelines and Standards for Fossil Vertebrate Databases (Blum, 1991).

W. von Koenigswald Steinmann Institut für Geologie, Mineralogie und Paläontologie, Bereich Paläontologie, Nussallee 8, 53115 Bonn, Germany M. M. Ataabadi Department of Geology, Faculty of Science, University of Zanjan, 45371-38791 Zanjan, Iran H.-W. Mittmann Retired, formerly Staatliches Museum für Naturkunde, Karlsruhe, Germany S. Sen Laboratoire de Paléontologie du Muséum, CR2P-CNRS-UPMC, 8 rue Buffon, 75005 Paris, France A. Tesakov Geological Institute, Russian Academy of Sciences, Pyzhevsky 7, 119017 Moscow, Russia J. Vepsäläinen
I. Žliobaitė Department of Computer Science, University of Helsinki, P.O. Box 68, 00014 Helsinki, Finland S. Viranta Department of Anatomy, Faculty of Medicine, University of Helsinki, Haartmaninkatu 8, 00014 Helsinki, Finland I. Vislobokova Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117647 Moscow, Russia L. Liu
L. Werdelin Department of Palaeobiology, Swedish Museum of Natural History, P.O. Box 50007, 104 05 Stockholm, Sweden Z. Zhang Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China

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The roots of the NOW database (originally Neogene of the Old World, subsequently New and Old Worlds) run deep in this fertile ground, or perhaps more accurately, it first developed as nothing but one branch of several of the Evolution of Terrestrial Ecosystems (ETE) database, developed by John Damuth and the ETE Consortium. Based at the Smithsonian, the ETE Program designed a novel kind of database for the terrestrial fossil record. Such a database would not include merely species-occurrence data for particular fossil localities, but in addition would include pertinent interpretations of sedimentary context, ecological environment and taphonomy (for localities) and biological and ecological trait characteristics (for the species). Such a database could be used to explore terrestrial paleoecology in a way not previously possible with large compilations of data. One could ask not only, “Where/when is such-and-such a taxon found,” but also questions such as, “Where during this time do we find localities with species of grazing mammals over 100 kg?” Such a database asks much of its compilers and needs frequent revision as ideas and techniques develop. But even a small set of relatively basic ecological variables opens up a wide array of avenues of research. This concept resonated with the small community of researchers involved in the beginnings of NOW. What follows here is a long and strange story, littered with coincidences and lucky breaks and not devoid of passion. One might think that databases are just neutral repositories of primary data, necessary but boring. Instead, they seem to call forth strong personal ambitions, emotions and impulses, including the darker urges related to possession and ownership. The collective history of fossil databases is accordingly dotted with drama – at least one field catalogue of fossils even ending up as an item in divorce proceedings. But, while seemingly unavoidable and prudent to keep in mind, this dark and gossipy side is largely irrelevant to the real purpose of databases and we shall not review it further here. At the end of the story there is a moral: a public database is fundamentally really a kind of museum collection. And, like all museum collections, it needs dedicated care and curation to deliver its potential for discovery, insight, beauty, and wonder. Just as the objects in a museum and data in a database tell a story of their own, we can only appreciate the collection as a whole if we know the ambitions and quirks of those who caringly brought them together. We have accordingly allowed a certain amount of local color and “fun facts” in the text that follows and trust that readers familiar with the events will not object. Such detail is rarely preserved and may help future generations to put things in context. For a timeline of the events and developments described, see Fig. 2.1 and for the growth in size and spatial coverage see Fig. 2.2. More technical and current aspects of NOW are described in Žliobaitė et al. (2023).

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Fig. 2.1 Timeline of the prehistory and history of the NOW database, showing the sequence and timing of key developments mentioned in the text. The division into stratigraphic eras is heuristic and purely informal. Note that most of the Cosmopolitan Era postdates the time frame of this chapter

The Kurténian Era and the Epi-Kurténian The Setting The NOW database was long in the making. It has deep roots in the legacy of Helsinki mammal paleontologist and writer Björn Kurtén, his emphasis on function and ecology and his strong will to systematically compile and organize information about fossil mammals and environments, as in his classic books Pleistocene Mammals of Europe (Kurtén, 1968), The Age of Mammals (Kurtén, 1971), and Pleistocene Mammals of North America (Kurtén & Anderson, 1980). Together with his then PhD students Lars Werdelin and Mikael Fortelius, Kurtén as early as the early 1980s developed a plan and a proposal for a basic database of Quaternary fossil mammals, based primarily on the raw data in the

appendices to the 1968 and 1980 books, which would have run on mainframe computers using punched cards for data input (Fig. 2.3). Thinking back to this heroic initiative one must admit that the Academy of Finland may have been right in turning the proposal down. Neither the hardware nor the software available then were quite up to the task, nor was the conceptual understanding of database structure. Nevertheless, it is undeniable that the idea was there at a very early stage. Nor can it be considered a coincidence that Kurtén’s former students were later to play an important role in furthering the idea. After Björn Kurtén’s death in 1988 there was a strong wish among his many colleagues worldwide to arrange a memorial meeting and publish a volume to honor his legacy. The meeting took place in Helsinki in the autumn of 1989, incidentally coinciding with the fall of the Berlin wall, and a resulting volume was eventually published as a special issue of Annales Zoologici Fennici (Forstén et al., 1992). The

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Fig. 2.2 All public NOW localities plotted at different times of the development of the database, showing the successive expansion of spatial coverage. Graph: Mikael Fortelius and Indrė Žliobaitė

original idea had been to publish a much more comprehensive book in the tradition of Kurtén’s own synthesis volumes, but these were the days of the first enthusiasm for computerized databases and the book initiative accordingly soon evolved into a project for a database. A proposal to this effect was submitted to the European Science Foundation, which eventually funded a Network on Quaternary Mammalian Faunas for 1991–93. One explicit goal of the Network was the establishment of a database of Quaternary Mammals of Europe, a parallel to the American initiative FAUNMAP (FAUNMAP Working Group, 1994), with a similar concept and structure. In September 1991, Lars Werdelin hosted a technical workshop in Stockholm (Fig. 2.4) where the properties of a Quaternary Mammal Faunas database were discussed in some detail, under the expert guidance of Russell Graham of

FAUNMAP, a Quaternary fossil mammal database currently part of the Neotoma Paleoecology Database (https://www. neotomadb.org/) and David Mayhew, a database professional with roots in mammal paleontology. An unpublished report with detailed plans was later circulated as “Report #1” of the Network (Werdelin et al., 1992). The Stockholm meeting fed directly into the first of three ESF-funded meetings under the Network, Mammalian Migration and Dispersal Events in the European Quaternary at Andernach on the Rhine, hosted by Wighart von Koenigswald in October of the same year. The Network was quite successful and arranged two more workshops, at Dijon in 1992 and at Sant Feliu de Guíxols, north of Barcelona, in 1993 (Agustí & Werdelin, 1995; Chaline & Werdelin, 1993; Von Koenigswald & Werdelin, 1992). Data on the occurrence of Quaternary mammals were compiled in the process

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but an actual database did not yet materialize. Those data were eventually made available through the Gaia data depository, latterly Pangaea, and later gradually made their way into NOW.

Making It Work

Fig. 2.3 Example of coding for Björn Kurtén’s Quaternary mammals database in the 1980s. A bold but precocious initiative that never quite materialized. Scanned document from the archives of Lars Werdelin. Inset: Björn Kurtén in 1986. Photo: Mikael Fortelius

Fig. 2.4 The Stockholm database meeting in 1991. A. Group photo with Mikael Fortelius, Lars Werdelin, Patrick Brunet-Lecomte, Albert van der Meulen, David Mayhew. Photo: Russell Graham. B. Database structure presented by David Mayhew (lower right). Photo: Lars Werdelin

Anyone who has been seriously involved with the development of scientific databases will recognize a number of universals. The first and also the most trivial of them is the early obsession with technical matters, such as the choice of software platform or programming language, on which hecatombs of time and effort have been and are still being sacrificed. Interoperability, industry standard query languages, open-source software, high-level object-oriented computer languages, and the internet itself were new ideas in the early 1990s. Accordingly, there was a fear (unreasonable, as it turned out) that if one did not choose wisely, one's data would become “stuck” in a particular system and format, and could not be recovered (anyone old enough to remember trying to extract large datasets from mainframe tapes to simple ASCII files recognizes the source of that fear and frustration). Hot on the heels of this comes the question of data access: who shall be permitted to use our incomparable database? In the present days of open access and open data it may not be obvious, but the instinctive and near-universal answer used to be some version of reciprocity: only if you contribute data will you gain access. This seemingly rational and fair choice of policy has spelled the doom of many a promising database. The population of potential contributors is simply far smaller than the population of potential users, and it is the latter that determines the visibility and perceived success of the database. The NOW database was somehow blessed in avoiding the worst of such standard mistakes. The key factor here may paradoxically have been the lack of funding and resources in general. For software we just took what happened to be available, making progress through trial and error. The list is long and includes at least the early platforms dbase, Kman, Clarion and Paradox. Once you have built a relational database several times in different environments, passionate arguments for any one of them appear rather futile. The far more important matter of database structure and standards was never an issue for NOW, having been built originally as a clone of the ETE database. As for data contributors, it soon became clear that it was far easier to compile data on behalf of potential contributors than to persuade them to do so themselves. When actually faced with a complication of data, purportedly from her own published work, a normal scientist will be utterly unable to resist the urge to correct the mistakes. And even the few who resist this urge cannot prevent the database from compiling

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their data unless they (suicidally) refuse to allow public use of said data. This is recognised among professionals as Damuth’s Vice. It was accordingly always obvious that NOW should have unrestricted access, right from the time that it was officially launched in 1996. But we are running ahead of our story. Let us step back and review the circumstances that surrounded the birth of NOW.

Building Blocks To understand the beginnings of NOW it is necessary to appreciate the special problems of European mammal stratigraphy. In much of North and South America, Asia and Africa, fossil localities often occur in long, continuous and well-exposed stratigraphic sequences that can be dated directly by radioisotopic and/or magnetostratigraphic methods. In contrast, European fossil localities tend to lack a clear lithostratigraphic context; they also frequently have no datable rocks and are dependent on biochronologic dating techniques. Especially in central Europe, the fossils often occur in isolated patches of sediment or in fissure fillings and often the only means of dating them is the taxonomy of the fossils themselves. This is why the MN-system (from Mammal Neogene; De Bruijn et al., 1992; Fahlbusch, 1991; Mein, 1975, 1979, 1989), a chronology based on the presence (and, in some implementations, the evolutionary stage) of mammal taxa has been such a fundamental framework for the Neogene mammal stratigraphy of Europe. Needless to say, fossil localities from long continental sequences, especially from Spain, were critical in the building of the MN system from the beginning and helped define its chronology and improve its resolution. These included the Early-Middle Miocene sequence of the Calatayud-Daroca Basin, the Middle to Late Miocene sequences of the Vallès-Penedès Basin and the Late Miocene sequences of the Teruel-Alfambra Basin. Thus, the important bipartition of the Vallesian into MN9 and MN10 was a direct result of the local biozonation proposed for the Vallès-Penedès Basin (Fahlbusch, 1976). Similarly, the tripartition of the Turolian into MN11, MN12 and MN13 directly followed the biozonation proposed by Anne van der Weerd for the Teruel-Alfambra Basin (Van de Weerd, 1976). Indeed, a more conventional alternative to the MN-system, applicable to such long sequences, was the definition of a number of European Land Mammal Ages (ELMA), such as the Vallesian, the Turolian and the Aragonian (Crusafont Pairó, 1950; Crusafont Pairó & Truyols Santonja, 1960; Daams et al., 1977). The inception of what later became known as The Reisensburg Concept, focused on central Europe and the eastern Mediterranean, must be seen against this background

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of challenging stratigraphic relationships. This idea underwent a slow gestation period during a one-year Alexander von Humboldt fellowship that Ray Bernor enjoyed during the 1989–1990 academic year. This fellowship ended with a five week stay at the University of Helsinki with Mikael Fortelius, working on the horses from the Late Miocene Sinap Formation in central Anatolia, where Fortelius was co-directing a field project at the time. Along with the systematics of the Sinap horses arose clear evidence of provinciality of Pikermian faunas in general and hipparionin horses in particular. Bernor expressed interest in dissecting the Pikermian phenomenon to derive a clearer history of the rise, dispersion and fall of Late Miocene savanna faunas. Upon returning to Howard University in July, Bernor formulated a plan for a workshop and study of Eurasian Miocene mammal faunas. Bernor originally approached Fritz Steininger in early September, 1990. Steininger showed interest but in fact was too occupied with the renovation of the Paleontological Institute in Vienna and rising interest in him becoming Director of the Senckenberg Museum, Frankfurt, Germany. Bernor then turned to Siegfried Rietschel, Director of the Staatliches Museum für Naturkunde, Karlsruhe and Volker Fahlbusch, Professor at the University of Munich. Both agreed to help Bernor undertake this project. Rietschel wished to host this meeting at the classic locality of Höwenegg, Hegau, Germany and to that extent opened a test trench across the site in the summer of 1991 to expose the geological section. Karlsruhe Biology Curator Hans-Walter Mittmann took over this organization at Rietschel’s direction. In the meantime, Volker Fahlbusch applied to the VW Stiftung for money to support the workshop. This request was successful and funded the 1992 workshop. The workshop began in Immendingen, near the Höwenegg site and the invitees visited the site to see a complete Miotragocerus cranium retrieved from the site and Carl Swisher sampled the section for paleomagnetics and argon/argon dating (which successfully gave a congruent age of 10.3 Ma, MN9; Swisher, 1996).

The Reisensburgian Era The Reisensburg Process The primary objective of the Reisensburg workshop (Fig. 2.5) was to update the systematic framework for the small and large mammal groups being reported. Workshop participants agreed that revising the taxonomy and acquiring a contemporaneous view of systematics and evolution would lead to the best resolved report for Eurasian Miocene mammals. De Bruijn et al. (1992) provided a revision of MN

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of Western Eurasian Neogene Mammal Faunas (Bernor et al., 1996). The Reisensburg Volume was the originally envisioned end product of the meeting, promised to the VW Stiftung and the participants, along with a compilation of up-to-date raw data to be shared among the group. But by the time of the workshop, Fortelius already had one foot firmly in the world of databases and he accordingly brought up the idea of making the newly compiled high-quality data the core of a more permanent database, to be updated and offered to the scientific community in general. This idea received some support, but by the end of the meeting the matter was still undecided and no discussion of practical details had yet taken place. At this point another NOW coincidence occurred, paradoxical as in a Tang Dynasty parable, where good bad fortune cannot be told from good except in retrospect. Fortelius returned to Helsinki to learn that the Academy of Finland had unexpectedly failed to continue his main research project on the evolution of occlusal relationships in suoids and he was therefore both in acute need of a more fundable research project and free to commit himself fully to it. Given the situation, the idea of a database project based on the Reisensburg process and data seemed an obvious choice. We quote from p. 6 of the original proposal to the Academy of Finland, submitted in January 1993 and approved in May the same year:

Fig. 2.5 A. Heinz Tobien and Ray Bernor at the Höweneg. Quarry in 1986. Photographer Unknown. B. Ray Bernor at the Schloss Reisensburg meeting in 1992, flanked by Carl Swisher III (left) and Jens Franzen (right). Dorsal view of Sevket Sen in the background. Photo: Mikael Fortelius

units, which subsequently was placed in the wider scope of circum-Mediterranean geochronology by Steininger et al. (1996). Swisher redated Höwenegg and Maragheh while large mammal and small mammal groups were revised by several invited specialists. Fossil plant occurrences were revised by Kovar-Eder and colleagues. Biogeographic and paleoecological interpretations were made by Bernor, Fortelius, and their collaborators. It took four and a half years to complete the project, from the time of the workshop in the summer of 1992 until the publication of the Reisensburg volume in 1996. It is true enough that there were some political challenges along the way, the pervasive one being “why is a young American vertebrate palaeontologist in such a prominent role for this initiative?” Volker Fahlbush and Siegfried Rietschel staunchly defended Bernor for leading this initiative and it should be told that Bernor’s principal detractor was distinctly proud of the outcome that fifty colleagues produced in the Reisensburg Volume Evolution

Fossil land mammals offer both 1) a source of independent data for investigating the nature and timing of these and other similar changes, and 2) a direct window on other, related and/or simultaneous changes in the mammal communities in different habitats and on several continents. Conversely, the evolution of land mammals in relation to such an increasingly detailed background of environmental change offers challenging opportunities for evolutionary research with an ecological perspective and a geological time scale. An important step towards a more global analysis of the European fossil land mammals was recently taken by Bernor, Fahlbusch and Rietschel, who organised a roundtable workshop at Schloss Reisensburg (Germany) on the Evolution of Continental Biotopes in Central Europe and the Eastern Mediterranean (15-5 Ma) in July 1992. For this workshop specialists were invited to revise the material for all relevant taxa and for a large number of fossil localities from Poland to Iran, and to analyse temporal and biogeographic patterns. This particular area was chosen because it can be related to the history of the Paratethys complex, and the interval because major physical and faunal changes are known to have taken place then. The Neogene land mammals are diverse enough and have a sufficiently rapid turnover to allow quantitative treatment of the data. This data will form the basis of the first scientific investigation undertaken using the NOW database, a collaborative project between Fortelius, Bernor & Mittmann (Letter 1).

The funding of this project instantly created the resources needed for going ahead with the database idea. It also raised some questions about the principles and practices of such an endeavor. In retrospect, one of them stands out as crucial:

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should it continue to be an informal coalition of friends and collaborators or should a more formal structure be created? In particular, should there be a formal advisory board? The fact that this question was eventually answered by affirming the latter position may have been one of the most decisive steps in the early history of the NOW database, providing it with both the scientific credibility and the sense of commitments and continuity that a fledgling database so badly needs. The original invitation to the Advisory Board members is included as Appendix 2.1.

NOW Begins Once the database was established other developments followed naturally. In 1995, the ESF approved a new network called “Hominoid Evolution and Environmental change in the Neogene of Europe” (HOMINET). The network was coordinated by Jordi Agustí and members of the Steering Committee included, besides Mikael Fortelius, also Peter Andrews, Lorenzo Rook, Louis de Bonis and George Koufos. Although originally centered in the climatic and environmental context of the hominoid species in Europe, one of the explicit main goals of the network was to produce a database of the hominoid sites, mainly those belonging to the Vallesian and Turolian European Land Mammal Ages. In this way, the HOMINET became an early and major contributor to NOW. The original NOW was intended to include Neogene data from Spain, Italy, Germany, Hungary, Greece, Turkey and Ukraine, among other countries. In order to achieve these goals, three workshops were organized. The first one took place in 1996 at Sant Feliu de Guíxols north of Barcelona, Spain, organized by Jordi Agustí and devoted to “The Vallesian”. A second workshop was organized in 1997 by Lorenzo Rook at Certosa de Pontignano, Siena, Italy, devoted to “Climatic and environmental change in the Neogene of Europe”. As a result of these two workshops, a volume of more than 500 pages was published in 1999 by Cambridge University Press, edited by Jordi Agustí, Lorenzo Rook and Peter Andrews, entitled The Evolution of Terrestrial Ecosystems in Europe (Agustí et al., 1999). This volume included significant contributions such as The paleoecology of the Pikermian Biome and the savanna myth (Solounias et al., 1999) or Vicariance biogeography and paleoecology of Eurasian Miocene hominoid primates (Andrews & Bernor, 1999). A third workshop took place at Nikiti, Greece, in 1998, organized by George Koufos, under the title Phylogeny of the Neogene Hominoid Primates of Eurasia. Contributions included sites from Spain, Italy, Hungary, Greece, Turkey, Georgia and Pakistan. As happened with the first two workshops, a second volume was published by Cambridge University Press in

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2001, edited by Louis de Bonis, George Koufos and Peter Andrews (De Bonis et al., 2001). Smaller initiatives also added to the NOW. The insectivorans had fallen through the cracks at the Reisensburg meeting, considered to be “insufficiently known at the point” (De Bruijn et al., 1992). This struck a nerve with some colleagues who had either published on the group or even dedicated most of their career to the study of insectivores. During a meeting at Senckenberg, Frankfurt, the Working Group on Insectivores of the Neogene of Eurasia (WINE) was formed and two years later a volume on the fossil record of the group in many countries was published (Van den Hoek Ostende et al., 2005). Fortelius quickly recognized this as a welcome addition to the NOW and the data were transferred into the database, turning one of the lesser documented groups into one of the best elaborated, as it were, overnight. This was also the start of Van den Hoek Ostende becoming more involved in the NOW and taking on a role as Associate Coordinator of small mammals. Which was a quite remarkable conversion, considering that at the Senckenberg meeting he had opposed the idea of publishing the insectivores as a database in the first place.

NOW as an ETE Node In 1986, the Evolution of Terrestrial Ecosystems Program (ETE) was established in the Department of Paleobiology at the National Museum of Natural History (Smithsonian). One of its signature early projects was the development of a computer database for the terrestrial fossil record. By 1992, ETE had developed a design and detailed schema for such a relational database (Damuth et al., 1997). A chance meeting in the fall of 1992 between Mikael Fortelius and John Damuth (then of ETE) started a conversation about coordination of databases. Fresh from the Reisensburg meeting, Fortelius knew that the kernel of a database of Old World Neogene faunas had taken shape, but the remaining question was the technical issue of how best to turn the data compilation into a database that would have potential for longevity and growth. ETE offered a general, public structure that could guarantee a degree of generality without sacrificing functionality. Fortunately for NOW, it was also organized as a system of nodes in order to allow participants to work from multiple locations, independently of institutional affiliation. The first version of what we now know as NOW was compiled in the autumn of 1993 by Mikael Fortelius and Suvi Viranta in Paradox for Windows, using the ETE structure. At this point Providence once more smiled upon NOW. In October the same year there was an infrastructure call by the Academy of Finland, to which Fortelius on short notice submitted an application for an HP-UNIX server to

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host the still embryonic database, with specifications to match those chosen by the ETE for its own database. Against all expectations, the Academy approved this application in December the same year. Moving NOW to the University of Helsinki's server platform meant that not only the structure, but the user interfaces that ETE had developed could be used seamlessly by NOW. Furthermore, the rise of the Internet made it possible to update and troubleshoot the software applications remotely. Significantly, in the days before the Web, it was nevertheless possible to work directly with the database for research remotely, using the X Windows system in any terminal application that was available at the time in some form or other on all computer platforms. NOW could be served. The user logged in directly to the NOW server in Helsinki, and the interactive graphical user interface appeared on their own computer. This was based on a custom-designed GIS application written for ETE and would later provide the architectural foundation for other databases as well, among them the original Paleobiology Database. The user was presented with a digital map of the world and could zoom in on any region, specify a timespan, and the localities would appear as points. Their contents could be seen via mouseover, and the map display could be queried for specific taxa or, for that matter, any properties recorded for the localities. Subsets of the data could be created this way and downloaded as delimited text files, or SAS datasets, or other useful formats. Even the map display with localities could be downloaded as a fairly basic PostScript file (this was before PDF). Additional features of the GIS interface included typical ones such as scale bars and measurement of distances. Another graphical interface could be used locally to manage the database. In the summer of 1994, the latest Paradox version of NOW was successfully ported to the new server and over the ensuing months various bugs and implementation issues were worked out. By 1995, NOW had become a peer-node database in the ETE Consortium, and NOW and ETE were in regular communication. It may be of interest here to record that the Helsinki NOW office was using an early Linux Debian installation on a PC as a NOW terminal during this time, starting from November 1995. The first machine was very nearly installed by Linux creator Linus Torvalds himself, but owing to a mismatch of schedules it was eventually done instead by another early Linux legend, Lars Wirzenius. Linux support in those critical days was most generously supplied by Jussi Sjöström, IT support of Hanken School of Economics, Helsinki (which may or may not have been aware of these goings-on). In time, the ETE Program reset its priorities, and by 2000 it was no longer building and hosting its own database. Significant subsequent American palaeontology database

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efforts such as PaleobioDB (https://paleobiodb.org/) and FossilWorks (http://fossilworks.org/) have focused on ambitious national projects to database the entire published fossil record, terrestrial and marine, at first primarily to study patterns of global Phanerozoic diversity. These projects necessarily have a different set of goals and a different relationship with the professional community than does NOW, which was a more focused but also more open-ended and actively maintained database effort. Under these circumstances NOW had little choice but become an independent entity with its own infrastructure and institutional support. This was a time of trial and error, with false starts and temporary fixes of various kinds. But crucially, the NOW community persisted through the crisis and the building of the database continued without any serious disruptions. In this phase, crucial support from the Finnish Society of Sciences and Letters (one of the two national science academies of Finland) meant that students could be employed for data entry and checking. Thus, the database continued to grow in scope as well as in usefulness, providing an increasingly fertile platform for the kind of research that would later be labeled as ecometrics (Eronen et al., 2010a).

An Excursion into the Land of Nod This chapter is the story of how the NOW database came to be and how it grew. There were, of course, all sorts of disappointments, setbacks and failures along the way but rehearsing them at length would serve no useful purpose here. But there is one spectacular failure that may be worth a brief section as it is of some historical interest as well as a great cautionary tale. This is the story of the EEDEN Programme. As the first results of NOW analyses were emerging in the mid-1990’s there was considerable enthusiasm and hope that a new and better understanding of deep time ecosystem change might be within reach. The multidisciplinary ambitions of the HOMINET workshops mentioned above also provided a fertile substrate for thinking along such lines. Thus, it came to be that a remarkably ambitious initiative was conceived and proposed to the European Science Foundation as a Scientific Programme. The idea, first formulated in an informal conversation between Nicholas Shackleton and Mikael Fortelius, was to extend the scope of HOMINET to include the marine realm, add a strong modeling component, and to shift the emphasis from hominoid evolution to the collapse and recovery of ecosystems. In short, this was to be what today would be called Earth system science. The programme was approved by the ESF for the five-year period 2000–2004. As a programme,

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Fig. 2.6 Mosaic of participants from several meetings of the EEDEN Programme. A. Sabadell, Spain, 2001: Jordi Agustí, Zlatko Kvacek, Louis Françoise, Jan A. van Dam. B. Frankfurt am Main, Germany, 2002: George Doukas, Silvia Iaccarino, Johann Meulenkamp, Jussi Eronen. C. Frankfurt am Main, Germany, 2002: Mikael Fortelius, John Damuth, Volker Mosbrugger. D. Stara Lesna, Slovakia, 2004: Hans de Bruijn, Gudrun Daxner-Höck, Madelaine Böhme. Photographers unknown

rather than a network like HOMINET, it had substantial resources for researcher mobility and research as well as for meetings. The EEDEN programme organised a series of inspiring meetings with memorable discussions between participants who were in many cases being exposed for the first time to completely different methodologies and viewpoints. Especially the world of modeling was new to many and not everybody was ready to embrace it without reservations, but over time there was genuine increase of understanding. Many transdisciplinary friendships were also formed under the EEDEN umbrella, especially among the young researchers who benefited from the laboratory visits that EEDEN made possible (Fig. 2.6). There is little doubt that EEDEN contributed significantly towards integration in a broad sector of earth and life sciences and was in that sense a real success. It nevertheless failed spectacularly to achieve its specific goal of providing a synthetic, system-level understanding of ecosystem collapse and recovery. Indeed, it even failed to overcome the technical obstacles it had identified at the outset and promised to clear away, such as mismatches between regional

stratigraphic schemes or a routine for easily plotting present-day coordinates on paleogeographic maps. Much incremental progress was made during the tenure of EEDEN in its individual participant groups but, apart from inspiring discussions at meetings, it was largely business as usual. Perhaps the time was not yet ripe. Years later concepts such as “critical transition”, “tipping point” and “system state shift” would enter into the global change debate and public vocabulary as a result of quite different efforts (e.g., Barnosky et al., 2012; Scheffer, 2009; Steffen et al., 2011). In the history of the NOW database, EEDEN passed almost without a trace.

The Ecometrian Era The Birth of Ecometrics In the late 1990s, there was a surge of interest in using fossil vertebrate faunas to quantitatively reconstruct aspects of Neogene and Paleogene habitats and paleoclimates. Other

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than in the province of Pleistocene and Holocene vertebrate researchers, who often can directly rely on the known properties of extant species, vertebrate-based paleoecology of even deeper time must usually employ traits that have a more general, functional relationship with the environment. An early attempt in this direction was E.C. Olson’s reconstruction of a Permian food web, which showed how different taxa occupied similar ecological roles over time (Olson, 1952). An ETE conference in 1987 on Terrestrial Ecosystems Through Time and the book resulting from it (Behrensmeyer et al., 1992) included discussion of what was then called ataxonomic characters, functional traits that would enable different taxa to be compared over long periods of time in terms of their ecological roles. The chapters on Ecological characterization of fossil plants (Wing & DiMichele, 1992) and Taxon-free characterization of animal communities (Damuth, 1992) are good examples of that novel direction. An early example of using a trait-based approach to reveal paleoenvironmental patterns was Jernvall et al. (1996), using a global data compilation published in book form (Savage & Russell, 1983). The obtained patterns were also shared with Helsinki researchers in meteorology and atmosphere physics at an early date and their enthusiastic support for interpreting them in relation to past climates definitely contributed to thinking about them as potential climate proxies. A more database-oriented attempt to address this was carried out by a working group at meetings between 1998 and 2001 at the National Center for Ecological Analysis and Synthesis in Santa Barbara (Fig. 2.7). The results were somewhat disappointing and in retrospect the effort was perhaps premature. But what did emerge was yet another potential paleoclimatic relationship between rainfall, on the one hand, and mean hypsodonty divided by species richness on the other (Damuth et al., 2002). A parallel exploration of patterns of hypsodonty and lophedness in fossil data was also in progress at this time (e.g., Fortelius & Hokkanen, 2001; Fortelius, 2003) but these early efforts were severely hampered by the crude options available for displaying the results. In this intellectual climate, then, it may not be surprising that Fortelius was alert to something that he discovered by chance in 2001 while trying out a new kind of plot offered by a statistics package (incidentally during the Golden Dragon New Year celebrations in a wintery Beijing): the summed pattern of the distribution of molar crown heights in the totality of NOW data showed a striking resemblance to the present-day distribution of rainfall on the Eurasian continent. This suggested that hypsodonty alone, combined with the readily available NOW data, might be able to reconstruct geographic patterns of past rainfall. The challenge of displaying these patterns was finally solved when Jussi Eronen joined Fortelius as a masters student and brought with him from his geography

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Fig. 2.7 Mikael Fortelius (left) and John Damuth (right) at the National Center for Ecological Analysis and Synthesis in Santa Barbara in 1998. Photo: Christine Janis

background cartographic skills including knowledge of the then still new-fangled GIS. Very soon after, the first mean hypsodonty maps were generated and presented to a broader audience at a meeting honoring the 70th birthday of Hans de Bruijn in May 2001 (Fortelius et al., 2003). Technically more advanced maps of a similar kind were published more accessibly the following year in Evolutionary Ecology Research, one of the first journals to make space for this emerging field (Fortelius et al., 2002; Fig. 2.8), significantly with Helsinki meteorologist Juhani Rinne as one co-author. The support of professional meteorologists was critical at this early stage, but eventually the need to anchor the colorful and suggestive maps in more than scientific intuition led to the development of the mean ordinated hypsodonty proxy for past rainfall (Eronen et al., 2010a, 2010b, 2010c; Fortelius et al., 2006a), the beginning of a vigorous line of research that continued to produce interesting results over the last decade. But we must leave the narrative at this point, ten years before the time of writing this text.

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Fig. 2.8 Example of inverse-distance interpolated maps of ordinated mean hypsodonty overlain on a map of Eurasia, interpreted as changing rainfall distribution over Neogene time. From Fortelius et al. (2002)

NOW Comes of Age Meanwhile, in parallel with the development of ecometric proxies for environmental conditions, a second line of NOW-based research was emerging. Here, the aim was not reconstructing environmental patterns of the past as such, but rather to understand the evolutionary dynamics generating them, through adaptive trait evolution driven ultimately by biotic interactions as well as changing physical conditions. The first to reach publication was Jernvall and Fortelius (2002), showing that the changes in spatial hypsodonty patterns in the European Neogene were due mainly to the evolutionary success of herbivores with increased crown height, which became progressively more common relative to herbivores lacking this key feature. The follow-up study of Jernvall and Fortelius (2004) delved deeper into the dynamics, looking at the history of the most common lineages in the same dataset. In retrospect, the most significant finding of the paper was undoubtedly the discovery of what is known today as the hat pattern: the unimodal occupancy history of taxa in the fossil record (Foote et al., 2007; Liow & Stenseth, 2007), currently another subject of vigorous research, still partly involving NOW data.

The first explicitly NOW-based publications are from 1996, all but one of them (Fortelius et al., 1996a) being contributions to the Reisensburg Volume (Bernor et al., 1996), described earlier in this narrative. These early papers were strongly focused on the analysis of quantitative patterns of diversity, turnover, biogeography and community structure, but following years saw a broadening of the scope, with a steady stream of papers citing NOW for their data. Some of these papers are cited on the NOW webpage (https:// nowdatabase.org/now/publications/) but the list is surely incomplete. It is striking in retrospect how many of these early papers were concerned with age and timing (among others, Agustí & Oms, 2001; Agustí et al., 2001; Garcés et al., 2001; Krijgsman et al., 2000). Paleoecological and paleobiogeographic work of course continued (Bernor & Rook, 2008; Bernor et al., 2009; Casanovas-Vilar et al., 2010; Furió et al., 2011; Koufos, 2003; Koufos et al., 2009; Madern & Van den Hoek Ostende, 2015; Nargolwalla, 2009; Solounias et al., 2010). Some of the earliest papers to computationally assess the completeness of the mammalian fossil record (Alba et al., 2001; Bingham et al., 2007; Saarinen et al., 2010) also used NOW data, as did some early

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paleontology gave 3,677 researchers in January 2021. A search at the Web of Science Core Collection provided 4,817 authors for the Web of Science Category “Paleontology” for publications during 2017–2021. Similarly, the Society of Vertebrate Paleontology reports its membership as “approximately 2,000” while the Palaeontological Association gives “well over 1,000” and even the Facebook groups of these two have only about 5,000 and 2,200 members, respectively. A more detailed search at the Web of Science Core Collection (https://clarivate.com/webof sciencegroup/solutions/web-of-science-core-collection/) provided 1,086 authors for the Web of Science Category “Paleontology” for publications during 2017–2021 with additional 60 keywords, including “Mammalia”, “Mammals”, “Mammalian” and 57 taxonomic order names listed in NOW database. In this perspective, hundreds of visitors per month is quite a respectable number.

Connecting People (and Disciplines)

Fig. 2.9 Towards resolving the ecometric biome space. Snapshot of the blackboard in Fortelius’ office in 2010, showing an early version of the work published by Liu et al., 2012. Photo: Jukka Jernvall

work on computational biostratigraphy (Alroy et al., 1998; Fortelius et al., 2006b; Puolamäki et al., 2006; Ukkonen et al., 2005). Macroevolutionary work also started to emerge at this time (Liow et al., 2008; Peláez-Campomanes & Van der Meulen, 2009) as did paleoclimatology (Eronen et al., 2009; Liu et al., 2009; Micheels et al., 2009). The research fed by NOW has continued to grow and diversify and many more publications have appeared over the last decade but a description of those developments is beyond our present scope. Figure 2.9 shows an example of work in progress from 2010, moving towards an ecometric concept of paleobiome space. It is important here to emphasize that NOW has always served two main functions: to provide datasets for quantitative analysis and publication, but equally, if not even more importantly, to function as an online source of information about fossil mammals and localities. This latter role is difficult to quantify (and records from the early years do not exist), but a rough idea is given by the number of mean monthly visitors (173 in 2012, 485 in 2020) and by the number of mean monthly pages visited (1933 in 2012, 3059 in 2020). These numbers should be compared to the size of the relevant research community, which, although numbers are not readily available, is probably on the order of a few thousand individuals worldwide. For example, ORCID (https://orcid.org/) queries matching variant spellings of

As is often the case in research, funding opportunities can inspire creative solutions by connecting previously separate disciplines and people. One such opportunity was the 2003 call of Academy of Finland on Systems Biology. Whereas systems biology and fossils did not exhibit any obvious links, Jukka Jernvall, Irma Thesleff, and Mikael Fortelius nevertheless proposed that studying mammalian teeth from evolutionary, ecological, and developmental biology perspectives could in fact be considered systems biology. A key component of the proposal, which ended up being fully funded, was the aim to examine ways to link NOW to datasets ranging from molecular data on developing mouse teeth to digitized, three-dimensional tooth shapes. Here, a key person hired in the project was Gudrun Evans, who developed the database structures to link the different kinds of data. One outcome of the collaboration was that it set NOW in motion towards incorporating specimen level data, something that continues to this day. Dental morphology of extant taxa posed its own challenges as the same specimen could be represented by multiple instances in the data. These kinds of cases include tooth wear data obtained from wild animals captured multiple times (King et al., 2005), or time-lapse images of growing teeth on the petri dish (Harjunmaa et al., 2012). For these vastly diverse kinds of phenotypic data, Gudrun and Alistair Evans constructed a MorphoBrowser database (http://pantodon.science.helsinki. fi/morphobrowser/) that houses many of the datasets collected in individual studies (e.g., Evans et al., 2007; King et al., 2005; Wilson et al., 2012). Serendipitously, MorphoBrowser also became the basis for a new database in the US, thus continuing the transatlantic exchange of ideas that had been part of NOW from the onset. In 2008, Doug Boyer

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was working at Stony Brook University with Jernvall when an opportunity arose to re-engineer MorphoBrowser, resulting in the widely used MorphoSource database (https:// www.morphosource.org/), currently coordinated at Duke University by Boyer (Boyer et al., 2017). One of the aims of the systems biology collaboration was to develop methods to compare very different, almost incomparable morphologies. To do this, teeth from morphologically diverse taxa were studied using new taxon-free approaches (Andrews & Hixson, 2014). Initial steps included molar crown types that provide a simple tabulation of number and position of tooth cusps and crests. These ecometrical descriptions were incorporated into NOW and used in some of the early analyses of environmental patterns (Jernvall & Fortelius, 2002, 2004). Crown types were soon followed by more quantitative methods that were developed with Alistair Evans and Gregory Wilson (Evans et al., 2007; Wilson et al., 2012). Unlike work on tooth morphology, tooth genes remained a more protracted challenge. Already by 1998, Irma Thesleff had established a database for genes expressed in developing teeth (Nieminen et al., 1998). Whereas this Bite-It database allowed the visual inspection of hundreds of genes expressed in developing teeth, the actual number of genes expressed dynamically in a developing tooth is in the thousands (Hallikas et al., 2021). Indeed, new, fast-throughput methods to acquire huge amounts of gene expression data, increasingly at single cell resolution, have transformed the field of developmental genetics to rely on large international data depositories. Combining these system-level data on genes with data on phenotypic detail remains a challenge, but NOW-inspired ‘ecometrics of genes’ have provided new ways to disentangle the principles of development (e.g., Hallikas et al., 2021; Morita et al., 2020).

The Cosmopolitan Era Opening the NOW Database up to the World The very first attempt of making NOW open to the world was made by Kari Lintulaakso using the development tools of a private software company where he was working at the time. During 2001–2002 Kari made a proof-of-concept solution of a regular, HTML-based user interface that allowed adding and editing the data. Eventually, the solution proved to be too costly for the essentially volunteer-based NOW, and it was sadly buried without further development. When Gudrun Evans took on the role of technical database coordinator in 2004, the existing UNIX version of NOW still only allowed users to view data remotely, but not

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to make any changes to it. Gudrun designed the structure for the new Systems Biology database, which linked the NOW database to the new MorphoBrowser and the Gene database (Fig. 2.10). She rebuilt the NOW database in MySQL, replicating the ETE-based table structure, and strengthening it by enforcing data rules restricting many fields to specific entries at the database level. Significant work was then undertaken to clean the data to meet these rules, and the data was subsequently transferred to the new MySQL database. Evans created a new, user-friendly web interface in Servoy (Java-based, cross-platform software, able to be used by both Mac and Windows users). By downloading the client software, users were then able to directly search and view the database remotely and were also able to log in and enter and edit data directly from anywhere in the world. To maintain the integrity of the data, individual users were granted access to edit only a restricted set of data, which related to their area of expertise, determined either by taxonomy or locality. Researchers were able to edit records that they themselves had contributed, and regional and taxonomic coordinators were able to edit records related to their field. All changes were logged and could be checked by coordinators. Servoy offered the benefit that the interface could be developed relatively quickly, allowing focus to be directed to how the data should be displayed, and what functionality was required. It also allowed further time to be spent cleaning the data to improve the integrity of the dataset – many issues came to light as the data were able to be searched and viewed in more detail, including adequately dealing with diacritical marks in names, locality and taxon synonyms, consistency in usage of ‘cf.’, and errors in calculations of fractions of age in time units, errors in latitude and longitude, and of course spelling errors and inconsistencies in taxa, localities and museums. This is a very time-consuming process, but an essential part of ensuring database integrity, so that a user conducting a search will be able to find all the matching records. Evans created synonym tables and functions for creating synonyms and merging duplicate records to assist with this process. With increasing numbers of users able to edit and enter data remotely, it was also essential to keep track of user permissions, and to create a log of changes that keeps track of who made what changes when and why (by linking these to a reference). It was also important to ensure that the database was being backed up regularly and adequately. As a result of the Systems Biology project described in the previous section, a new Systems Biology database, incorporating NOW, was hosted at the University of Helsinki on a new server called Pantodon. Evans was helped by database technician Joonas Kauhanen, who set up this server and created back-up routines, and also a website traffic monitoring feature so that we could keep track of the number of visitors to the site. Kauhanen continued his association

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Fig. 2.10 The structure of the “Systems Biology” database, integrating NOW with the MorphoBrowser. Graph: Gudrun Evans

with the project for a number of years, and his expertise was invaluable in trouble shooting many issues as the database and interface were developed. The Servoy user database interface went live in April 2005. Over time the limitations of using Servoy became apparent: many users experienced problems using the client software, the server software required updating frequently, which inevitably led to necessary fixes in the interface, and there was an annual license fee that was unsustainable in the long term. Therefore, in 2007, Evans commenced building a new user interface in PHP, replicating the look and functionality of the Servoy interface, still connecting to the MySQL database and using an Apache server (Fig. 2.11). This eliminated all the problems with Servoy, but it was more complicated to develop and took longer to create because everything had to be coded manually. Evans designed and created much of this interface herself, but was helped by a few database technicians, each of whom brought invaluable expertise to the project. Veli-Pekka Kestilä helped check the new interface for potential security flaws before it was released, and also helped move the database from the aging Pantodon server to a new server called Mutikka, hosted by the Finnish Museum of Natural History,

Fig. 2.11 Gudrun Evans in the office at Helsinki, coding the new PHP interface for NOW. Photo: Alistair Evans

in 2008. Jouni Vepsäläinen from Enporia Oy was contracted to write a log-in function and editing, saving, and logging functions for the PHP interface in March 2008, essential requirements before being able to open the interface to internet access. Jouni continued his association with NOW,

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with Jouni Vepsäläinen and Atro Tossavainen as the database continued to be developed, working with Susanna Sova who was preparing the Harvard Siwaliks dataset for entry, and, together with Alistair Evans, on preparing the North American dataset for upload: the latter were uploaded in 2011, as described below.

Across the Vastness of Asia

Fig. 2.12 From left to right, Gudrun Evans, Susanna Sova and Mikael Fortelius at the farewell party for Gudrun and Alistair Evans in 2008. The tiramisu cake by Susanna Sova shows the NOW logo by Noira Martiskainen. Photo: Alistair Evans

helping Evans on various aspects of the interface development, and finishing the implementation of the database after Evans moved back to Australia in June 2008, with Susanna Sova taking over the management of the database (Fig. 2.12). The new PHP interface finally went live in October 2009, and users were now able to view and edit the data directly over the internet, without the need to download any client software. This was a critical step in the development of what was by then fast becoming a serious, international resource maintained by an interconnected NOW community. Although NOW at the time of writing enjoys a comfortable and hopefully long-lasting relationship with the Finnish Museum of Natural History, the first attempt to place the database within the museum was not successful. In 2010, the database moved servers again, this time to a server called Mormyrus, hosted by the Institute of Biotechnology, part of the University of Helsinki. Atro Tossavainen provided a lot of assistance moving the database, and getting it functioning correctly on the new server. Evans continued her association with NOW from Australia for some years, communicating

Inevitably, and as has already been testified, the history of the NOW database was always intertwined with the individual histories of its participants. A case in point concerns its expansion from the original scope of central Europe versus the eastern Mediterranean to cover the entire Eurasian continent. While the NOW database was being born, Fortelius was shifting his field project from western to eastern Asia. This was a very deliberate move, motivated partly by the decline of the Sinap Project in Turkey (Sen, 2003:14) and partly by a (retrospectively remarkably naïve) hypothesis arising from the first analyses of the initial NOW data: that the faunal developments seen in western Asia (e.g., Fortelius et al., 1996b) might be mirrored on the eastern half of the continent. After a few false starts, a cordial collaboration was initiated with the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) in Beijing and a first visit and planning session took place in Beijing in May–June 1996, where the junior Chinese colleague Zhaoqun Zhang was given the task of minding the guests Mikael Fortelius and Juha Pekka Lunkka and keeping a record of the daily negotiations. The area selected for field work was the Miocene section at the Bahe River by Lantian town in Shaanxi Province near the city of Xi’an, known from previous work to sample strata from the Middle to Late Miocene and therefore an appropriate match for the previous work in the Sinap Formation in Turkey. The first field season was agreed for September 1997 and young IVPP staff members Zhaoqun Zhang and Liping Liu were given the task of preparing the project and surveying the field area. The first field season was a challenging one, not least because Fortelius broke his leg just before it and therefore arrived in China three weeks late and on crutches. He was met in pouring rain by a dispirited field party, which had endured exceptional heat and very meagre findings in the previous weeks. Despite further challenges of similar kind, the field season was not only completed but became the beginning of a successful collaboration that still continues at the time of writing. From the very beginning, the project was set up with the dual purpose of data compilation as well as field work, and it naturally became the task of Zhang and Liu to help with the practical aspects of the database part as well, joining the NOW the advisory board along with a few more senior colleagues. The data compilation progressed well in the

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early years of the project and had by the turn of the millennium reached a sufficient stage of completeness to provide a first inkling of continental-scale patterns, as related above. One startling detail in the hypsodonty patterns, which might well have been ignored without critical ground-truthing through the field work at Lantian, was the reversal of the temporal sequence familiar from Europe and North America. Instead of a gradual drying during the later Miocene a reversal to more humid conditions at about 8 million years ago was indicated by mean hypsodonty values. This reversal was interpreted by Fortelius et al. (2002) as showing an intensification of the east Asian summer monsoon. Later work has largely confirmed both that this regional reversal of the planetary climate was real and that it allowed the establishment of the characteristic and diverse Chinese Hipparion fauna of the latest Miocene, known since the earliest days of Chinese paleontology from the famous dragon bone mines in the Red Clay deposits of the Loess Plateau (Fortelius & Zhang, 2006; Jokela et al., 2005; Liu et al., 2009; Passey et al., 2009). These developments left the interior of the Asian continent still largely unsampled by NOW, but this was soon to be remedied. By the beginning of 1990s, the reference Neogene mammal faunas that occupied a mid-latitude Eurasian belt, from Moldova to the Transbaikalia and Mongolia, had been already correlated with the European Neogene Mammal Units (MN system), and those from Asia also with Chinese Land Mammal Ages due to their intensive investigations mainly by specialists from the Paleontological Institute of the Russian Academy of Sciences (PIN) and Geological Institute of the Russian Academy of Sciences (GIN), and due to input from researchers of academic institutions of the former Soviet Union republics (Ukraine, Georgia, Kazakhstan). The correlations were based on biochronological evidence and also on the paleomagnetic dating and correlations with marine stages for some of them. After the International Congress BiochroM’97 in Montpellier in 1997, Innessa A. Vislobokova (PIN) joined the NOW team as a Coordinator, and later Alexey Tesakov (GIN) also took part in this work. It was not an easy task to convert many faunas from this huge territory to NOW standards, because some old localities were tied only to the regional stratigraphical schemes, and their place in the NOW database was uncertain. Vislobokova and Tesakov subsequently continued in charge of these questions and with the updating of mammal lists from the localities of this territory. In 2000, Diana Pushkina joined the work on the NOW database, mainly searching for the interglacial Eemian faunas on the former USSR territories, while completing her PhD under Mikael Fortelius’ supervision. Yet another NOW-story merits brief mention here. One morning in April in 2000, Fortelius was traveling on the morning train to downtown Helsinki and happened to bump

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into a friend who was at the time employed at the Academy of Finland. In the course of the 11-min commute the friend urged Fortelius, known for his habit of “going to strange places”, to apply for funding for collaborative research with partners from the Islamic Republic of Iran. The Academy had opened a call for this but had received disappointingly few proposals and the deadline was fast approaching. Thus, it came about that a research project was constructed at lightning speed, possible only because of the network that the newly established EEDEN programme offered, and funding was received in due course for a visit by Fortelius and then PhD student Anu Kaakinen to prospect for terrestrial mammals in the poorly sampled but paleobiogeographically crucial Oligo-Miocene of Iran. The continental red beds bracketing the marine Qum Formation in the Central Desert were selected as a first target, with structural geologist Ali Hamedani of Isfahan as a most obliging and helpful host. While this first exploration in August 2001 turned out to be futile in the sense that no significant discoveries were made during the two weeks it took, it was the beginning of a collaboration that is still ongoing at the time of writing, twenty years later. The key person here has been Majid Mirzaie Ataabadi, who joined the excursion as an undergraduate student assistant and thereby later came to get his Ph.D in a very much NOW-based project in Helsinki (Mirzaie Ataabadi, 2010).

Into the New World In the mid-1980s, Christine Janis (Fig. 2.13) hatched the idea of compiling a book on North American Tertiary mammals (i.e., Paleogene and Neogene mammals, the term “Tertiary” is now obsolete): the Pleistocene was excluded because of the existing compilation of Pleistocene North American mammals by Kurtén and Anderson (1980), and additionally the fact that including this time period would have vastly expanded the nature of the project. The notion was to encompass a who’s who for each family that also included information on taxonomy, higher level systematics, ecomorphology (including a molar measurement for body size estimates), evolutionary patterns, standardized phylogenies and figures showing range times for each genus, and – perhaps most importantly in terms of future utility for databases such as NOW – a link for each species to a specific fossil locality. The locality list was compiled as a separate appendix with numbered localities ordered by geographic region (e.g., the prefix CP = Central Great Plains), and the locality numbers then assigned to each taxon. In this sense the Tertiary Mammals (TM) book concept was unlike faunal compilation books of the time (such as Maglio & Cooke, 1978, Evolution of African Mammals) and more in the spirit of Savage and Russell, Mammalian Paleofaunas of the

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Fig. 2.13 Christine Janis contemplating the ecomorphology of Hypohippus, a pony-sized browsing horse (Anchitheriinae) from the Middle Miocene of North America. Hypohippus plushie by Savannah Olroyd. Photo: Christine Janis

World, 1983, although with a more detailed locality compilation. Janis initially embarked on this project with the editorial help of Kathy Scott and Louis Jacobs, but the process of trying to organize the many contributors was a process akin to not just to herding cats, but with some of the cats having behavioral issues. A problem with a volume of this nature, unlike most edited volumes, is that it is simply not an option to leave out a recalcitrant contributor (and hence omit a taxon). A decision was eventually made to turn the book into two volumes and to go ahead with the taxonomic groups with all of the contributions in place, and in 1998 the first volume (TM1 Carnivores, Ungulates, and Ungulate-Like Mammals; Janis et al., 1998) was published by Cambridge University Press. The second volume (TM2 Small Mammals and Marine Mammals; Janis et al., 2008) was published a decade later, this time with new contributing editors, Gregg Gunnell and Mark Uhen. This second volume also had some updates of both the locality information (including additional localities and some new dates) and the taxa covered in the first volume. Although this book was not originally devised as an online database, the hope always was that somebody would make such use of the information. Starting in 2008–2009, in the aftermath of the publication in the Proceedings of the National Academy of Sciences of the United States of America of a paper entitled Distribution history and climatic

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controls of the Late Miocene Pikermian chronofauna (Eronen et al., 2009), Janis began corresponding with Fortelius and others about North American chronofaunas. This turned into an endeavor to include TM1 and TM2 into the NOW database. The work started in late 2009, and went through 2010 and 2011, producing the first synthesis papers soon after (Eronen et al., 2012; Figueirido et al., 2012). Laura Säilä-Corfe and Jussi Eronen were responsible for the correspondence and much of the work in addition to Mikael Fortelius and Susanna Sova. The locality-related work and issues (see below) took much time, correspondence and travel. For example, Eronen and Janis were working on the TM1 and TM2 data in Providence when hurricane Sandy hit the Eastern Seaboard of the United States in 2012. The process of incorporating the Tertiary mammals data into NOW concluded in 2016, when they were made public, and the North American data have been subject to normal updates since then. A potential problem with converting the TM locality data to the NOW format is that Janis had not separated all the localities to the level of individual sites as was done in NOW (and in other databases such as the ETE or the Paleobiology Database. So, for example, locality CP110 (the Olcott Formation in Nebraska) includes around three dozen separate quarries or sites of varying sizes and quality (which are named in the locality list but not separately numbered). At least 10% of the localities are of this nature. This ‘lumping’ approach was taken for several reasons: (1) contributors often did not provide the necessary detailed information; (2) it was considered that, for the purposes of the book, a degree of locality lumping would actually be more informative, rather than reporting on every little sample individually; (3) given that Janis was working essentially alone, and at least for the first volume without the type of computerized assistance (e.g., Excel files) that we now all take for granted, she likely would still be completing TM1 had she decided to include every locality incidence. However, this lack of faunal locality equivalence with the existing structure of NOW remains a problem to be rectified.

Running NOW From the beginning, the NOW interface was designed to be as intuitive as possible so that the users can concentrate on its essential reason for being – the data. The aim was that even a lazy but excited researcher can insert the data smoothly. Adding data was made as straightforward as possible, human mistakes were minimized, and collecting information in different ways was steadily improved. Rules were enforced at both the interface and the database levels, restricting the range of possible entries in many fields, and

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enforcing the integrity of relationships between the various fields. Gudrun Evans was the main person behind this thinking and Jouni Vepsäläinen continued on the same track. We were very lucky to get Jouni just before Gudrun and Alistair Evans left Helsinki for their native Australia. It has been a surprise how much abstract thinking in three dimensions database development requires and Jouni had that ability in spades. Jouni quickly took the technical aspects in hand and his meticulous work enabled the development of NOW without compromising the safety and robustness of the database. NOW officers all had database-related projects of their own, which made them alert to errors and inclined to think about improvements. Thus, the interface was continually improved in many small steps. Localities with unknown coordinates no longer ended up in the Gulf of Guinea, under the armpit of Africa and the Map- and Export-functions became easier to use and adapted for a range of different purposes. The biggest improvement to the user interface may have been the Import-function. Previously, large, curated datasets had required a great deal of manual labor in harmonizing the data exactly to the NOW format, but with the improved Import-function, automatic format checking and reformatting was introduced. Jussi Eronen was the first brave NOW participant to actually use it, successfully as it turned out, for importing the North American data described above. This import significantly increased the records in the database, which in turn made it important to further develop checks on the consistency of logic as well as the formal validity of entries. History has shown the value of these improvements again and again, as many datasets still come to NOW as Excel-files that may have grown over years or even decades, with changes in usage that typically occur over time. From 2009 onwards, the NOW office has worked together with the taxonomic coordinators, updating and unifying taxonomic information of all families, under the umbrella of ‘NOW synonymy project’ (ridding the database of synonymous entries). This was initiated by Laura Säilä during her post-doctoral period in the Academy of Finland funded research project Extinction dynamics of taxa in the fossil record. This has vastly improved the taxonomic accuracy of the NOW database and how it can be reliably used for large-scale evolutionary studies that rely on taxonomic data. Additionally, improvements in how changes are recorded, and can be viewed, in the references of each locality and taxon entry and how synonymies and other taxonomic issues are detected in the database were implemented. Simultaneously, enhancements took place in search functions for taxonomic entries and their geographic/temporal occurrences in both the main database and its Export and Maps interface, and the types of data that could be exported out of the NOW database were increased.

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One late but crucial development must be mentioned here, even though it falls outside the temporal scope of the main narrative. One of the main challenges of running NOW during its first two decades was the lack of an employed curator to provide stability and continuity. This was finally remedied when, in the autumn of 2016, the University of Helsinki received a major donation from the Ella & Georg Ehrnrooth Foundation and decided to use this, in accordance with the wish of the donor, for furthering the research into hominin paleoenvironments that had sprung up as a collaboration between Fortelius and Meave Leakey’s research network centered on the Kenyan Turkana Basin. This donation both allowed the university to establish a professorship of Hominin Environments and to finally establish, in the Finnish Museum of Natural History, a permanent technical data coordinator position that includes responsibility for the maintenance of the NOW database. For this position Kari Lintulaakso was fortunately available, a person already involved with NOW for many years and intimately familiar with the structure and purpose of the database. This way, NOW finally achieved an administrative position in accordance with its international role and weight. As is evident from this narrative, NOW came together gradually, through an unplanned and rather amorphous process of collaboration between friends and colleagues who for various reasons shared an interest in developing this common resource. In retrospect it seems clear that what gave NOW direction, coherence and credibility in the critical early phase was the establishment of an Advisory Board of respected scholars, many of whom had also been active during the gestation period that preceded the formal launching of the database (cf. Appendix 2.1). At the time this was by no means seen as a necessity by all and several early participants indeed felt that it would be an unnecessary complication or even a waste of time. Nevertheless, the Board was established and was immediately engaged in the daily running of the database, some members taking a more active part than others. Despite its name, the Advisory Board did not at any time collectively advise. Rather, it provided individual scientific expertise and opinion in specific cases and in this way acted as a filter and quality control, deciding on such matters as how errors should be handled and how new results and opinions should be considered. It was during this time that an important NOW principle came to be established: more than advisors, the Board members should be rulers, with dictatorial power to decide on such issues as taxonomic nomenclature and stratigraphic correlation. It doesn’t have to be right, consistency is everything! has always been a key slogan for NOW data development and in this NOW differs significantly from other leading databases, for example the Paleobiology Database (https://paleobiodb. org/), which instead favors recording of alternative scientific opinions. While both systems have their advantages, the

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dictatorially induced consistency of NOW definitely makes it easy for users to locate all occurrences of a species, but inevitably also creates extra work for users who disagree about synonymy. Needless to say, few if any decisions of this kind have ever been made without consultations among coordinators and with the approval, implicit or explicit, of the general coordinator. NOW was always very much a consensus-based community effort. Originally, NOW had no administrative organs, there was just the somewhat misnamed Advisory Board, an Overall Coordinator and an Associate Coordinator. When executive decisions had to be made, this was done by the signatories of the founding document (Appendix 2.1), ironically self-styled as the NOW Mafia. As NOW grew and new members joined the Advisory Board it was gradually realized that some more transparent administrative structure was desired. A new structure was accordingly put in place in December 2013 with a Steering Group appointed by the now renamed General Coordinator and approved by the Advisory Board. At this time a five-year cycle of membership in both the Steering Group and the Advisory Board was also introduced. The main task of the Steering Group has been to decide on major issues such as the appointment of new board members or updating the stratigraphic reference for the NOW Time Unit Table. Thus, the daily running of the NOW database gradually caused its administrative structure to stabilize and mature. Looking at the structure of the NOW and the people involved, it is clearly an international consortium serving a global community. At the same time, history leaves no doubt that the main office of this consortium is in Helsinki. As is clear from the narrative above, it was this firm ground that allowed the database to develop all of its technical and conceptual aspects, supported by the Finnish academic infrastructure. Many colleagues and students travelled to Helsinki over the years and these visits were the basis for fruitful discussions and collaborations. However, they were still individual visitors enjoying the hospitality of the Finnish team. A major step towards creating a stronger community feeling came, when in 2016 NOW switched to a creative commons license, expressing the role of the community in each reference made to the database. In addition, in 2018, the long-cherished wish of having a NOW meeting finally came to be, when Lars van den Hoek Ostende and Peter Joniak organized the workshop NOW and the future of the past in Bratislava (Fig. 2.14). The benefits of having users and developers at one table became quickly apparent and the meeting produced a long list of action points to be taken.

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Fig. 2.14 Pablo Peláez-Campomanes, Lars van den Hoek Ostende and Peter Joniak at the Bratislava NOW meeting in 2018. Photo: Laura Säilä

Cauda: Changing of the Guard Following the pioneering ecometric papers by Jernvall and Fortelius (2002, 2004), a new line of research was opened in Helsinki that relied on collaboration with computer scientists. This successful initiative actually came about by administrative fiat, a rare occurrence in academia. In October 2002, Fortelius was extolling the virtues and potential of the hypsodonty proxy for rainfall to then University of Helsinki Rector Kari Raivio, who asked the critical question “would you need money or people?”. Upon receiving the reply “People, or money only to acquire them”, Rector Raivio promised to give the matter some thought. Soon after, Fortelius was contacted by Academy Professor Heikki Mannila, a prominent data mining scientist, and a meeting was agreed. Discussions ensued and it soon appeared both that the complex and fragmentary nature of the NOW data offered a highly attractive object for algorithmic analysis and that realizing the potential would require serious and patient two-way exchange of information. Thus began an enjoyable and productive collaboration with Mannila and his group (Bingham et al., 2007; Eronen et al., 2010a, 2010b, 2010c, 2011; Fortelius et al., 2006a, 2006b, Heikinheimo et al., 2007, 2012; Kallio et al., 2011; Liu et al., 2012; Puolamäki et al., 2006; Saarinen et al., 2010; Ukkonen et al., 2005), which unexpectedly came to an abrupt near halt when Mannila was appointed Vice Rector of the newly founded Aalto University in 2009. When he went on to become President of the Academy of Finland in 2012 the need for a replacement became urgent.

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History of NOW

In this situation, with an abundantly proven concept and the opening prospect of a collaboration with Meave Leakey and her network of collaborators on the legendary fossil record of the Turkana Basin in northern Kenya (see Fortelius et al., 2016), one might have expected that it would be easy to find such a replacement. Instead, it gradually became clear that the long and patient discussions that preceded the actual collaboration could not be easily repeated and that the slow rate of progress in paleontological research was seen as a major deterrent by most hypothetical candidates for this post. Fortunately, one of the early collaborators from Mannila’s group, Aristides Gionis, came up with a long-shot suggestion that a Lithuanian postdoc at Aalto University, Indrė Žliobaitė, might be interested in trying her hand at fossils in the newly funded Turkana Basin project ECHOES (Academy of Finland, 2014–2018). An interview was accordingly agreed and, after the differences in expected salary levels between geoscience and computer science had been overcome by good will and creative solutions, Žliobaitė accepted an offer to become the ECHOES postdoc. This was the beginning of a second wave of computer science collaboration based on the NOW data that would eventually see Žliobaitė transformed into a paleontologist in her own right and the successor of Fortelius as the General Coordinator of NOW. But this is another story. Acknowledgments Although there has been some dedicated funding for NOW, including the Academy of Finland grant 1993-1996 that made the first implementation possible, NOW was always overwhelmingly an unfinanced community effort. Most of the work has been contributed by NOW participants in their own time, and much has no doubt been sponsored by a large number of academic employers worldwide, knowingly or otherwise. In addition to the Academy of Finland we would like to acknowledge generous support from the European Science Foundation, the Volkswagen Foundation, the Finnish Society of Sciences and Letters, the Finnish Museum of Natural History, the University of Helsinki, the Naturalis Biodiversity Center, the Hanken School of Economics and the CSC – IT Center for Science. Our special gratitude goes to the Ella and Georg Ehrnrooth Foundation and Mrs. Elsa Fromond for the generous endowment that enabled, among other things, the establishment of a position at the Finnish Museum of Natural History that includes technical database coordination of NOW. This is R.L. Bernor’s NSF FuTRES publication 35.

Appendix 2.1. The Original Invitation Letter to Join the First NOW Advisory Board NOW c/o Prof. Mikael Fortelius Department of Geology P.O. Box 11 FIN-00014 University of Helsinki Finland Vox: + 358-9-1912–3419 Fax: + 358–9-1912–3466

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Email: [email protected].fi Dear colleague, 15.10.1997 The first years of the NOW database have gone, and with them the years of highly coherent and organised data entry. NOW, such as it was used for the synthetic chapters of Bernor-Fahlbusch-Mittmann (and is available on request from [email protected].fi), is far from free of errors, but it is still a uniquely up-to-date source of information regarding the occurrence and attributes of the Miocene mammals of Europe on a continental scale. It was never complete, and it still isn’t. Several taxa (notably Proboscidea) are missing altogether, and most localities have incomplete faunal lists. The selection of localities is also somewhat random (the case of Austria was especially remarked upon in this regard). We are gradually filling in holes but there is no concerted effort comparable to the “Reisensburg Process”. In our opinion the database is already a valuable resource, and we would like to contribute to its being more extensively used than has been the case so far. In the long run, such databases have the potential of bringing fossil mammals into contact with neighbouring fields, for example palaeoclimate and biodiversity modeling and research. Meanwhile, we know from first-hand experience that the database is already a wonderful resource for many mundane, everyday tasks involving Eurasian Neogene mammals. We feel strongly the need to continue the process of building the database as well improve the quality of the data. We are writing to you in the hope that you might share our interests and be willing to participate in the next step of developing the NOW database. To begin with, there are currently four main sources of new data into NOW: 1. Specific research projects that need to augment the existing data in some way (especially “ecomorphological” attributes of the animals) 2. The ESF “HOMINET”, which is revising and adding (primarily) the hominoid-bearing localities of Europe 3. A collaboration project with the IVPP to database all Chinese later Cenozoic mammal localities 4. Data extracted from the literature by our students and ourselves. In addition to these and somewhat different in nature are plans to fuse NOW with other existing databases with a similar structure (EUQUAM, ETE, FAUNMAP). Such plans will proceed independently of our development of NOW, and will not change the basic principles outlined below. Not all the data arriving to the database are of equal quality, and not all specialists agree on the details of taxonomy or dating, to say nothing of interpretation that is more speculative. Only some relatively powerful verification mechanism can prevent the growing database from decaying

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into a chaotic state, with taxa represented by multiple nomina and perpetuation of various long-since- recognised errors. In establishing such a verification mechanism one must balance several factors. It is important that the database retains a broad base in the field, so that it can grow as a common resource. On no account must it be seen as the toy, or worse, the exclusive research tool, of some small clique! It is also important to minimise the workload put on single individuals. On the other hand, for a database more important even than correctness is consistency. This means that once a certain classification and nomenclature has been adopted, any competing scheme must be put aside until, perhaps, it is verified and a conscious decision is made to adopt it. A similar principle applies to stratigraphic correlation and related issues, and these factors argue against having too many individuals directly involved with the supervision of the data. What we propose here is the creation of an Advisory Board for NOW, consisting primarily of the core group from the original Reisensburg Process, augmented (or replaced) by colleagues who have since joined the project. The Board would feature specialists and taxa, on stratigraphy, and on particular regions, and would complement itself as required. The Board would rarely (if ever) be expected to act as a body, but would be more like the advisory board of a scientific journal, where individual members would advise on matters with their special sphere of expertise. The default duty would be to review (or distribute for review), perhaps once a year, data listings of various sorts, such as lists of localities and ages, lists of localities and their fauna, lists of taxa with ecological interpretation, and so forth. Beyond keeping an eye on the quality of the data in this manner, board members would be well placed to guide the direction of development and to initiate or facilitate research projects. The first step in this direction was already taken when Hans de Bruijn kindly agreed to coordinate a complete overhauling of the small mammal data, a project currently underway. At the moment we visualise a Board of somewhat the following composition and primary responsibility: ***

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General large mammal coordinator – Ray Bernor Hominoid Primates – Peter Andrews Cercopithecoid Primates – Eric Delson Feloid Carnivores – Lars Werdelin Arctoid Carnivores – Suvi Viranta Suoids – Jan van der Made Tragulids – Elmar Heizmann Cervids – Getrude Rössner Giraffids – Nikos Solounias Bovids – Alan Gentry Equids – Ray Bernor Chalicotheres – Louis de Bonis Tapirs – Jens Franzen Rhinoceroses – Kurt Heissig Proboscideans – Bill Sanders Hyracoids –Elmar Heizmann Tubulidentates – Sevket Sen Stratigraphy General stratigraphic coordinator – Sevket Sen Remmert Daams, Volker Fahlbusch, John Kappelman, Pierre Mein, F.F. Steininger, Qiu Zhuding Regional coordinators Western Europe – Jorge Agusti Eastern Europe – George Koufos, Adam Nadachowski Carpathian Basin – Kordos Laszlo Southwest Asia – Sevket Sen Russia – Innessa Vislobokova Indian subcontinent – David Pilbeam China – Qiu Zhuding Geology and taphonomy Sedimentology – J.P. Lunkka

Proposal for Advisory Board of the NOW database Taxonomy and ecomorphology General small mammal coordinator – Hans de Bruijn Mein, Höck, Bolliger, Römer… (Hans: please provide a list of your people!)

Taphonomy – Miranda Armour-Chelu Palaeoecology Peter Andrews, Kay Behrensmeyer, Mikael Fortelius, Peter Ungar

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Contacts with related projects The ETE Consortium – John Damuth The ESF Network on Fossil Insects – ? Terrestrial molluscs – ? Leaf floras – Johanna Kovar-Eder Pollen floras – Jean-Pierre Suc General coordination and development Overall coordination and data entry – Mikael Fortelius Database structure and technical development – John Damuth Associate NOW coordinator – Suvi Viranta *** A few words about the “data policy” that we have in mind. As agreed during the preparation of the Bernor-Fahlbusch-Mittman volume, all contributors are entitled to the full data set that was produced. It was also agreed that this data set would become public with the publication of the volume, and this is the basic principle that we intend to maintain. New data deposited in the NOW database can be kept under restricted access until a project is completed, but will eventually join the growing mass of public data available to all bona fide researchers. Another basic principle is that the NOW data should not be distributed directly to third parties. Everybody is required to get their data directly from NOW ([email protected]. fi), to ensure that the currently available version is used, and to allow us to keep track of how and where NOW data are being used. (Backup copies of older versions will, of course, be kept and made available for verification purposes.) In the future, a version of the public data will also be available on the world wide web, probably as a collaboration project with the ETE database. We hope that you will accept our proposal and join the NOW database Advisory Board. Whether you do join or not, we hope that you will let us know your opinion and any suggestions you may have, including suggestions for additional people you feel should be involved. With our highest hopes and best wishes, Mikael Fortelius, Ray Bernor, Hans de Bruijn, Sevket Sen, Lars Werdelin.

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2

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Chapter 3

The NOW Database of Fossil Mammals Indrė Žliobaitė, Mikael Fortelius, Raymond L. Bernor, Lars W. van den Hoek Ostende, Christine M. Janis, Kari Lintulaakso, Laura K. Säilä, Lars Werdelin, Isaac Casanovas-Vilar, Darin A. Croft, Lawrence J. Flynn, Samantha S. B. Hopkins, Anu Kaakinen, László Kordos, Dimitris S. Kostopoulos, Luca Pandolfi, John Rowan, Alexey Tesakov, Innessa Vislobokova, Zhaoqun Zhang, Manuela Aiglstorfer, David M. Alba, Michelle Arnal, Pierre-Olivier Antoine, Miriam Belmaker, Melike Bilgin, Jean-Renaud Boisserie, Matthew R. Borths, Siobhán B. Cooke, Jan A. van Dam, Eric Delson, Jussi T. Eronen, David Fox, Anthony R. Friscia, Marc Furió, Ioannis X. Giaourtsakis, Luke Holbrook, John Hunter, Sergi López-Torres, Joshua Ludtke, Raef Minwer-Barakat, Jan van der Made, Bastien Mennecart, Diana Pushkina, Lorenzo Rook, Juha Saarinen, Joshua X. Samuels, William Sanders, Mary T. Silcox, and Jouni Vepsäläinen Abstract NOW (New and Old Worlds) is a global database of fossil mammal occurrences, currently containing around 68,000 locality-species entries. The database spans the last 66 million years, with its primary focus on the last 23 million years. Whereas the database contains records from all continents, the main focus and coverage of the database historically has been on Eurasia. The database includes primarily, but not exclusively, terrestrial mammals. It covers a large part of the currently known mammalian fossil record, focusing on classical and actively researched fossil localities. The database is managed in collaboration with an international advisory board of experts. Rather than a static archive, it emphasizes the continuous integration of new knowledge of the community, data curation, and consistency of scientific interpretations. The database records species occurrences at localities worldwide, as well as ecological I. Žliobaitė (&)
M. Fortelius
K. Lintulaakso Finnish Museum of Natural History LUOMUS, University of Helsinki, P.O. Box 44, 00014 Helsinki, Finland e-mail: indre.zliobaite@helsinki.fi I. Žliobaitė
J. Vepsäläinen Department of Computer Science, University of Helsinki, P.O. Box 68, 00014 Helsinki, Finland I. Žliobaitė
M. Fortelius
L. K. Säilä
A. Kaakinen
D. Pushkina
J. Saarinen Department of Geosciences and Geography, University of Helsinki, P.O. Box 64, 00014 Helsinki, Finland R. L. Bernor College of Medicine, Department of Anatomy, Laboratory of Evolutionary Biology, Howard University, 520 W St. N.W, Washington, DC 20059, USA Human Origins Program, Department of Anthropology, Smithsonian Institution, Washington DC, Washington, DC 20013-7012, USA L. W. van den Hoek Ostende
M. Bilgin Naturalis Biodiversity Center, P.O. Box 9517 2300 RA Leiden, The Netherlands

characteristics of fossil species, geological contexts of localities and more. The NOW database is primarily used for two purposes: (1) queries about occurrences of particular taxa, their characteristics and properties of localities in the spirit of an encyclopedia; and (2) large scale research and quantitative analyses of evolutionary processes, patterns, reconstructing past environments, as well as interpreting evolutionary contexts. The data are fully open, no logging in or community membership is necessary for using the data for any purpose.













Keywords Paleontological databases Mammals Fossil Paleoecology Paleobiology Neogene record Cenozoic

C. M. Janis School of Earth Sciences, University of Bristol, Bristol, BS8 1RL, UK Department of Ecology and Evolutionary Biology, Brown University, Box G-W, 80 Waterman Street, Providence, RI 02912, USA L. Werdelin Department of Palaeobiology, Swedish Museum of Natural History, Box 50007104 05 Stockholm, Sweden I. Casanovas-Vilar
D. M. Alba
J. A. van Dam
E. Delson
M. Furió
R. Minwer-Barakat Institut Català de Paleontologia Miquel Crusafont (ICP-CERCA), Universitat Autònoma de Barcelona, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain D. A. Croft Department of Anatomy, Case Western Reserve University School of Medicine, 10900 Euclid Ave., Cleveland, OH 44106-4930, USA L. J. Flynn Department of Human Evolutionary Biology, Harvard University, Cambridge, MA 02138, USA

© The Author(s) 2023 I. Casanovas-Vilar et al. (eds.), Evolution of Cenozoic Land Mammal Faunas and Ecosystems: 25 Years of the NOW Database of Fossil Mammals, Vertebrate Paleobiology and Paleoanthropology, https://doi.org/10.1007/978-3-031-17491-9_3

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Introduction The NOW database just turned 25 years old. Its colorful history is described in Chap. 2 (Fortelius et al. 2023). Started as a compilation of the European Neogene fossil record, the NOW database is now fully global, covering all continents, as well as the Paleogene and Quaternary in addition to the Neogene, although not equally densely. NOW stands for New and Old Worlds. The name was changed in 2012 from Neogene of the Old World to reflect expanding geographic and temporal coverage. The database collects, curates, and presents the global fossil record of mammals from roughly 66 million years ago to the very S. S. B. Hopkins Department of Earth Sciences, 1272 University of Oregon, Eugene, OR 97403, USA L. Kordos Eötvös Loránd University, Savaria Campus, Károlyi Gáspár tér 4, Szombathely, 9700, Hungary D. S. Kostopoulos Faculty of Sciences, School of Geology, Aristotle University of Thessaloniki, 541 24 Thessaloniki, Greece L. Pandolfi Dipartimento di Scienze, Università della Basilicata, Via dell’Ateneo Lucano, 10, 85100 Potenza, Italy

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recent past. The primary source of the data in NOW is the published scientific literature. Harmonization of data from different sources and synchronizing diverging interpretations takes most of the curatorial effort. For integrity and consistency, NOW strictly follows the scientific interpretations of dedicated experts – the Advisory Board members. While NOW is not the only database covering fossil mammals (Uhen et al., 2013), it is the only global fossil database dedicated solely to mammals. It aims to include all the classical and well-studied fossil mammal localities known to research. Currently, we believe it roughly encompasses about two-thirds of the entire Cenozoic M. Bilgin Department of Geology and Palaeontology, Comenius University, 84215 Bratislava, Slovakia J.-R. Boisserie Laboratory Paleontology Evolution Paleoecosystems Paleoprimatology (PalEvOPrim), CNRS & Université de Poitiers, 6 Rue Michel Brunet, 86000 Poitiers, France French Centre for Ethiopian Studies (CFEE), CNRS & Ministère de l’Europe et des affaires étrangères, P.O. 5554 Addis Ababa, Ethiopia M. R. Borths Division of Fossil Primates, Duke Lemur Center, Duke University, 1013 Broad St., Durham, NC 27705, USA

L. Rook Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121 Firenze, Italy

S. B. Cooke Center for Functional Anatomy and Evolution, Johns Hopkins University School of Medicine, 1830 East Monument Street, Baltimore, MD 21205, USA

J. Rowan Department of Anthropology, University at Albany, 1400 Washington Avenue, Albany, NY 12222, USA

J. A. van Dam Department of Earth Sciences, Faculty of Geosciences, Utrecht University, Princetonlaan 8, 3584 CB Utrecht, The Netherlands

A. Tesakov Geological Institute, Russian Academy of Sciences, Pyzhevsky 7, 119017 Moscow, Russia

E. Delson Anthropology, Lehman College and the Graduate Center, The City University of New York, 365 5th Avenue, New York, NY 10016, USA

I. Vislobokova Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117647 Moscow, Russia Z. Zhang Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, China M. Aiglstorfer Naturhistorisches Museum Mainz, Landessammlung für Naturkunde Rheinland-Pfalz, Reichklarastr. 10, 55116 Mainz, Germany M. Arnal Vertebrate Palaeontology Department, La Plata Museum, National University of La Plata, La Plata, Argentina P.-O. Antoine Institut des Sciences de l’Evolution de Montpellier, CC64, Université de Montpellier, CNRS, IRD, EPHE, 34095 Montpellier, France M. Belmaker Department of Anthropology, The University of Tulsa, 800 South Tucker Drive, Tulsa, OK 74104, USA

E. Delson
S. López-Torres Division of Paleontology, American Museum of Natural History, 200 Central Park West, New York, NY 10024, USA NYCEP (New York Consortium in Evolutionary Primatology), New York, NY, USA J. T. Eronen Ecosystems and Environment Research Programme & Helsinki Institute of Sustainability Science (HELSUS), Faculty of Biological and Environmental Sciences, University of Helsinki, P.O. Box 65, 00014 Helsinki, Finland BIOS Research Unit, Meritullintori 6, 00170 Helsinki, Finland D. Fox Department of Earth & Environmental Sciences, University of Minnesota, 116 Church St SE, Minneapolis, MN 55455, USA A. R. Friscia Department of Integrative Biology & Physiology, University of California, Los Angeles, Box 957246 Los Angeles, CA 90095-7246, USA

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mammalian fossil record in terms of species occurrences known to research. It does not, of course, record all the individual specimens ever found, as that would be an impossible task, but focuses on the temporal and geographic distribution of taxa and their ecological characteristics. The database does not include mammals or mammalian ancestors from the Mesozoic. From the start of operation, NOW has put a very strong emphasis on the Advisory Board and the curation of data. Each taxonomic group, geographic area or geological time has dedicated experts, whose roles are similar to those of associate editors in scientific journals. This system allows us to incorporate new scientific insights into the database while ensuring consistency throughout. Rather than being an archival repository of static data snapshots, NOW aims at integration and continuous incorporation of the evolving and expanding knowledge of the research community. In that sense, NOW is not only a database of fossil data, but equally a database of current scientific interpretations. The consistency of curatorial treatment is important not only in taxonomic assignments, but equally in the level of conservatism when compiling faunal lists, age estimation, and selection of localities for reporting. NOW puts a major emphasis on data curation and global integrity of the fossil record. All records will never be at the same level in terms of accuracy and reliability, but NOW aims to apply the same curatorial principles in handling uncertainties, especially taxonomic uncertainties. “Why study the fossil record?” is an eternal question that has many answers, which may be obvious to members of the paleontological and related research communities. For

M. Furió Department of Geology, Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès, Barcelona, Spain I. X. Giaourtsakis Department of Earth and Environmental Sciences, Section of Paleontology & Geobiology, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, 80333 Munich, Germany L. Holbrook Department of Biological and Biomedical Sciences, Rowan University, 201 Mullica Hill Rd., Glassboro, NJ 08028, USA J. Hunter Department of Evolution, Ecology and Organismal Biology, The Ohio State University, 318 W. 12th Avenue, 300 Aronoff Laboratory, Columbus, OH 43210, USA S. López-Torres Institute of Evolutionary Biology, Faculty of Biology, University of Warsaw, Warsaw, Poland J. Ludtke Department of Biological Sciences, San Diego Miramar College, San Diego, CA 92056, USA

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outsiders, the main take-away message is that the fossil record allows us to see alternative scenarios of how living worlds could be and to infer principles for how the living world works in general. The fossil record allows understanding the patterns of evolution and ecosystem structure, as these cannot be inferred from the study of living organisms alone. It also provides evidence of ancient climate and geography, and helps to estimate the age of geological formations. Last but not least, the fossil record is the key to understanding how current global biodiversity and ecology developed over geological time. The greatly impoverished biodiversity of today provides only a very limited view of what was typical of most of the Cenozoic. Paleontological research is thus essential for studying the principles of how the living world works and how continental ecosystems once supported higher mammalian biodiversity. For example, tropical African savannas and forests today include one or two species of proboscideans and rhinocerotids, while over the last 25 million years it was not unusual for many species of equids (Janis, 2023), proboscideans (Huang et al., 2023), and rhinocerotids to occur together across Eurasia, Africa, and North America. A wide spectrum of scientific questions can be addressed using fossil databases, including inquiries about global and local environmental dynamics in the past, mass and background extinctions, ancestral relationships of species and their diversity and the response of life to major changes in our planet’s geological past. Global fossil data compilations can be used for analyzing evolutionary patterns and evolutionary contexts, studying ways of life and reconstructing evolutionary contexts of individual species, including humans. They also have potential applications in conservation, the latter defining the relatively new field of conservation paleobiology (Barnosky et al., 2017; Dietl & Flessa, R. Minwer-Barakat Departamento de Estratigrafía y Paleontología, Universidad de Granada, Avenida de Fuente Nueva s/n, 18071 Granada, Spain J. van der Made Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (CSIC), Madrid, Spain B. Mennecart Natural History Museum Basel, Augustinergasse 2, 4051 Basel, Switzerland J. X. Samuels Department of Geosciences, East Tennessee State University, P.O. Box 70357 Johnson City, TN 37614-1709, USA W. Sanders Museum of Paleontology, University of Michigan, 1105 North University Avenue, Ann Arbor, MI 48109, USA M. T. Silcox Department of Anthropology, University of Toronto Scarborough, 1265 Military Trail, Toronto, ON M1C 1A4, Canada

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2011; Kiessling et al., 2019). Certainly, many new research questions are yet to be formulated. The main goal of the NOW database is to summarize and represent the current scientific knowledge of the global research community about the mammalian fossil record in a way that researchers, students, journalists, policy makers and the public all over the world can use. In this way, NOW contributes to recording global geoheritage. It also gives coordinates and information that can help policy makers to locate and delimit important fossiliferous areas for protection.

When many data points are available, regional- to global-scale studies become possible only if scientific interpretations of, e.g., taxonomic assignments, classification of taxa or age estimation of a locality, are reliable and consistent. The NOW database collects, integrates, and harmonizes the mammalian fossil record, aiming for consistency of interpretations. The data in NOW are a precious collection of evidence and knowledge generated over many years by research communities worldwide. It is not just a sample dataset, but to a large extent embraces the known record of mammals in the past.

The Nature of NOW Data

Is the Fossil Record Biased?

The fossil record is as much about the contexts of remains of past organisms found in sedimentary rocks as it is about the fossils themselves. The NOW database compiles and harmonizes secondary data – data from publications identifying what has been found, where and in what geological contexts. Specimens that we see on display in museums are usually exceptionally well preserved or well restored and often include nearly complete skeletons. However, the majority of mammalian fossil specimens are only fragments of organisms that once lived. Identifying them to taxonomic level and placing them into relevant geological contexts requires extensive specialist training and expertise. Collection of primary data in paleontology is extremely labor intensive. Finding fossils in the first place requires skill acquired through training and practice, as well as a certain amount of luck. Fossil finds primarily come from targeted searches and expertise is needed to know where and how to search. Often the target is an area of known exposures, but within that, paleontologists normally range over the outcrops looking for fossils or concentrations of them. Sometimes they target particular taxonomic groups. Collecting methods also differ, for example most small mammal collections come from screen-washing fossiliferous sediments in field campaigns rather than systematic excavation. Not uncommonly though, fossil finds can result accidentally from roadworks and other building activities, probably an increasing trend as human land use intensifies. The excavation and preparation of fossil specimens that may be broken or encased in rock is another labor intensive activity that requires special training. Next, fossils and their geological contexts are studied by experts, identified, interpreted, and described in publications. It may take years or decades for a fossil specimen to become a quality data point with accurate temporal, spatial and taxonomic information. Such data points can be analyzed on their own or as part of larger scale analyses covering extensive geographic areas and time spans.

Yes, of course it is. Fossils do not represent all times and all places of the past equally well. In On the Origin of Species, Charles Darwin dedicated more than a full chapter to the imperfection of the geological record (Darwin, 1859): “I look at the natural geological record, as a history of the world imperfectly kept, and written in a changing dialect; of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved; and of each page, only here and there a few lines.” (pp. 310–311). The fossil record known in Darwin’s day represented a very small part of potentially available material, and he understood that these fossils reflected a very incomplete and fragmentary record of past life (Newell, 1959). With over a century of active collection and documentation of fossil evidence, the global fossil record is now broader and more abundant, and includes increasingly rich contextual information, but still, most certainly, we have only a glimpse of a subset of the former life on Earth that was preserved and is available for study. Even though concerns about the quality of the fossil record are voiced frequently (Benton et al., 2000; Kalmar & Currie, 2010; Kidwell & Flessa, 1996; Saarinen et al., 2010; Seddon et al., 2014; Turner, 2007; Valentine et al., 2006), others argue that as knowledge is cumulative and sampling biases are gradually corrected, gaps in the fossil record are better understood (Benton & Storrs, 1994; Currie, 2019; Newell, 1959; Plotnick, 1993), and major patterns stabilize. Preservational patterns in vertebrate fossils are to some extent consistent over time within known preservation modes (Behrensmeyer et al., 1992), which brings good and bad news. The good news is that potential biases can be quantified and accounted for. The bad news is that what rarely or never gets preserved will remain virtually absent from the record. Understanding that the fossil record will never be complete raises a broader question: how representative is the knowledge that we derive from this record and how robust

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are our conclusions about how the living world works in general? Surely, the fossil record does not cover the complete history of life. Even within the mammalian fossil record, which has received focused attention over centuries, not only new species, but new genera of mammals are still being discovered by fieldwork (e.g., Madden et al., 2010; Ríos et al., 2017; Turvey et al., 2018) and even on the shelves of museum collections (e.g., Borths & Stevens, 2019). Yet, the existing fossil record contains pieces of evidence of different kinds – momentary disaster snapshots and long-term attritional accumulations, fissure fillings and lagerstätten (sites with exceptionally rich accumulation or preservation), carnivore traps or cave deposits, anthropogenic base camp and kill site accumulations and even past environments that are otherwise unfavorable to fossilization. Given the variety and vast number of such fragments, even if the fossil record is incomplete in a historical sense, the record is expected to be relatively more complete in representing the past diversity of function and forms. Thus, if many different pieces of evidence are available and those pieces represent different environmental circumstances and preservation contexts, we can hope to reconstruct the functional ecosystems of the past by drawing on overlaps among those pieces.

Data Curation and the NOW Community At the time of writing this chapter the NOW database contains records on around 16,200 mammalian taxa (not all of which are identified to the species rank), about 6,400 localities, and approximately 68,000 species by locality occurrences as publicly available open data, licensed under the Creative Commons attribution 4.0 license (CC BY 4.0 by The NOW Community). Species information in NOW is always public. New localities may be initially entered in private mode and released as public data after the faunal lists and the contextual information are curated. Inevitably, some entries in the database will contain errors. Existing data are curated either via targeted revisions of the database or following notifications from users about potential errors. New data chunks are added following larger research projects or data releases in the community, or on an ad hoc basis when new publications come out, typically under initiatives or following pointers flagged by the Advisory Board members. An active community of users helps to keep the database up to date as much as possible. The database does not strive to react immediately to major taxonomic updates or other revisions, but rather takes a conservative long-term approach and allows some time before major revisions within the database, as new perspectives settle in the research community.

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The primary authority for any data interpretation and treatment is the NOW Advisory Board, listed on https:// nowdatabase.org/now/board/. The Advisory Board consists of the General Coordination and the Management team, the Steering Group, the Coordinators and the Emeritus Board. The General Coordination and the Management team runs NOW on a daily basis. The NOW management team is headed by the General Coordinator and includes the Steering Group, Associate Coordinators, specialists for database infrastructure, and junior data curators. The Steering Group makes strategic decisions and appoints Coordinators. Coordinators have individual dedicated areas of responsibility. The Emeritus Board Members have no dedicated tasks but advise when needed on a range of issues. NOW strictly follows scientific interpretations and perspectives of the coordinators, responsible for taxonomic groups, regions, times, geological and ecological contexts. Currently NOW has about 100 dedicated coordinator roles. Coordinators are appointed by the Steering Group based on experience, expertise and reputation in the research community for a minimum of five years. The work of coordinators is similar to that of editors of scientific journals. The main responsibilities of coordinators are twofold – answering queries from data curators when interpretations or edits for small snapshots of data are needed, and monitoring their own areas of responsibility in NOW, flagging potential issues with data and editing data when necessary. The NOW database is run as a community service on a voluntary basis. The database has no dedicated funding. Over the years, most of the funding used to build and maintain the technical infrastructure, user interface, and carry out large scale data preparation comes from regular research projects of the NOW Community members. Appendix 3.1 provides more details on the working procedures via Frequently Asked Questions.

The Database and Data Infrastructure The database is physically hosted in Helsinki, Finland. The history chapter by Fortelius et al. (2023) provides details on how and why the database came to be hosted there, where it has operated for the last 25 years. The infrastructure consists of the relational database itself and the user interface via which users can access and query the database. The NOW website acts as a gateway to the database, as well as providing news, lists of advisory personnel, background information, instructions for usage and pointers to the user interface (https://nowdatabase.org/). Currently, the user interface of the database runs on a container platform at the University of Helsinki. The user interface code is written in PHP. The database itself is currently hosted by the Finnish

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Fig. 3.1 A simplified database scheme, a map of public NOW localities over all ages and NOW logo

Museum of Natural History. For many years, the database used to be MySQL but now it runs on MariaDB. At the center of the database are the species table, the locality table and the relational table between the two. There are over twenty accompanying tables, of which the most important are shown in Fig. 3.1. Treatment of time in NOW is modular. Only in rare cases when absolute time in years is available (such as from radiometric dating) is the age of a locality entered as a number. In most cases, localities are assigned to time units. Time units can be continental or regional mammal biozones, magnetochrons, geochronological units and more. When the boundaries of time units are updated globally, locality ages in the database adjust automatically. Each locality has a minimum and a maximum age estimate, which do not have to come from the same basis of age: for example, the upper age bound of a locality can be derived from magnetostratigraphy and the lower bound from an assignment to a mammalian biozone. All entries or edits to the database are referenced and most of the references are scientific publications. However, NOW allows referencing personal communications or

unpublished sources if necessary and if those sources give the most reliable information according to the opinion of the responsible coordinator. The opinion of a coordinator within his or her jurisdiction in NOW can be used as a reference. Strictly following the authority of individual coordinators allows consistent treatment and data harmonization within NOW, which, in case of inconsistencies, is prioritized over information derived verbatim from the publications. The policy of NOW is to provide a conservative species list per locality. This means that in a case where two occurrences of the same genus are reported in the literature, one of which is identified to the species level and the other is species indeterminate, NOW assumes that unless there is strong evidence otherwise, both represent the same taxon. If there is strong evidence that the second occurrence represents a different species, this can be recorded in NOW with the second occurrence highlighted in the species list. Similar conservative treatment applies at higher taxonomic ranks. NOW allows the recording of yet unnamed species at localities. NOW also has an extensive infrastructure for handling synonyms of taxa and localities. Data entry

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Table 3.1 Major information available in NOW database in addition to species occurrences at localities. Stars indicate mandatory fields, which means this information is available for all the entries. Information in the remaining (optional) fields may be sparse Attributes of speciesa

Attributes of locality

Attributes of locality-speciesb

Taxonomic affiliation* Diet, selectivity, digestion Habitat, locomotion Body mass, body length Brain mass, sexual dimorphism Tooth shape Hypsodonty Mesowear Microwear Crown types (2 schemes) Localities

Name* Country, region* Geographic coordinates* Minimum and maximum age* Basis for age* Lithostratigraphy Sample unit Lithology Depositional context Sedimentary environment Taphonomy Climate, vegetation, seasonality Pollen record Ecometric estimates of climate Museums Species

Species* Locality* Body mass Mesowear Oxygen isotopes Carbon isotopes

*

The attribute is mandatory for entry Attributes of species mean that the values are the same for a particular species no matter in which locality it occurs b Attributes of locality-species mean that the values for a particular species may vary from locality to locality a

conventions and treatment instructions are available on the NOW website (https://nowdatabase.org/now/conventions/). Users can export selected data as a flat table, wherein each row is one occurrence of a species at a locality, and columns represent attributes of localities, attributes of species in general and attributes of species by locality. Selected data can be viewed via integrated mapping services. Table 3.1 outlines what major information in addition to species occurrences is available in the database. Users can also get information on the references associated with each entry in the database as well as the history of updates for each entry.

The Future The future remains to be seen. It is clear that we are entering an era in which the study of large and integrative data sets is becoming an increasingly common practice in science. In paleontology, the treatment of large collections of data has become as important as the careful excavation and study of individual fossil assemblages. Considering that the NOW initiative started at a time in which we could not foresee current developments, the database is surprisingly well-equipped to meet contemporary challenges. The focus on integrity and consistency of data, rather than attempting to catch each new development, has proven to be a

successful formula. Because of this down-to-earth approach, the database continues to support research and inquiries into the world of fossil mammals. The NOW database not only provides data but acts as an authority for scientific interpretations of the mammalian fossil record. From the curatorial perspective NOW is fundamentally like a physical fossil collection at a museum, growing and developing in many ways, some of them goal-oriented and others more random. And like a museum collection, it is indispensable as a reference for scientists. NOW summarizes the knowledge of a research community and the database is curated and refined as knowledge of the community accumulates. In that way NOW is like a “living monograph” of information. More than a simple collection of data, NOW is a community. Acknowledgments This chapter was completed and accepted after revision in August 2021. NOW does not have dedicated institutional funding. The database and data development are funded from regular research projects of the NOW Community members. Current and recent (last 5 years) funding sources include: The Ella and Georg Ehrnrooth Foundation and The Academy of Finland. ICP researchers acknowledge funding from the “Generalitat de Catalunya (CERCA Programme)”, R+D+I projects “PID2020-117289GB-I00” and “PID2020-116908GB-I00” (MCIN/AEI/10.13039/501100011033/) and consolidated research group from the Generalitat de Catalunya “2022 SGR 00620”. This is Bernor’s NSF FuTRES publication 35. L. K. Säilä acknowledges Academy of Finland Postdoctoral grant (275551). We thank three reviewers for helpful suggestions regarding the manuscript text. Contributions from the Valio Armas Korvenkontio Unit of Dental Anatomy in Relation to Evolutionary Theory are acknowledged.

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Appendix 3.1. NOW Frequently Asked Questions What is NOW used for? Many users simply look up information about individual localities and species, using NOW as an encyclopedia. You can also use NOW as a time machine to look at how the distribution of species has changed over time, or to check the oldest and youngest record of a given species or taxon. NOW is also widely used as a source for professional publications all around the world. Why are there only mammals in NOW? Because NOW is a community working on mammals. The database is not a stand-alone data repository, it summarizes knowledge of the research community, and is curated accordingly as new knowledge accumulates. NOW is compiled from many data sources of different quality; is it accurate? Just like any individual dataset compiled by any researcher, NOW most certainly contains inaccuracies and uncertainties. The data in the NOW database are coordinated with emphasis on consistency, meaning that the final say on presence or absence of species or other characteristics is up to the coordinator responsible for that particular group or region. Rather than presenting a collection of many available opinions, now presents interpretations of experts. In some ways the treatment is similar to that of scientific committees such as the International Commission on Stratigraphy overseeing the International Chronostratigraphic Chart. Why should I use NOW? You are most welcome to use any source on the internet. What is special about NOW is that it contains information about the traits and properties of the taxa, their occurrences and the localities at which they occur. NOW aims at being consistent in this. That is why we have coordinators with full rights to decide issues of taxonomy and stratigraphy, for example. Of course, and even with complete formal consistency, a residual of underlying disorder and inconsistency will always remain. Will NOW ever be complete? No. Although NOW has very good coverage of many areas and intervals, it will never be complete. NOW is

fundamentally similar to a physical fossil collection at a museum. The collection grows and develops in many ways, some of them goal-oriented and others due to chance. NOW summarizes the knowledge of a research community, the database is curated and refined as knowledge of the community accumulates. Completeness is not the aim. More than a simple repository of data, NOW is a community of researchers, scholars, and specialists. I found an error in the database, what can I do? Wonderful! This is one important way in which the database can grow and develop. The first step is to discuss it with the coordinator responsible for the respective group or region. Please get in touch with the database curator Kari Lintulaakso (kari.lintulaakso@helsinki.fi) who will guide you to the right person or simply look at the NOW Advisory Board listing on the NOW web page and contact the appropriate coordinator directly. If the responsible coordinator agrees, you can make the update yourself or leave it for the responsible coordinator to make the update. To make updates yourself you will need an account, which will be provided for any updates cleared by the coordinator. I found/published a new paper describing the fauna of locality X, I would like the database to reflect that, what can I do? You are most welcome to add data. Please get in touch with the database curator Kari Lintulaakso who will guide you to the right person and through the process. Can I report data based on my own observations of museum specimens? Yes, please do! The database is referenced, and each update must have an associated reference. The primary practice is to reference publications, but professional personal communications and observations are also acceptable as references. All of that must be properly referenced and tractable, for example by DOI. What is a locality? It is a unit referring to a place (and time) where fossils come from. Some places have a finer resolution/ finer division into localities than others. A locality may be a single layer of rocks or sediment or refer to a broader area, such as a whole rock series, for example. The latter situation occurs especially with older collections. NOW follows locality designations of the original publications from which data come.

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What is a General locality?

Can I get a user account to the database?

General localities have been established to record data that has no fine-grained locality information or comes from historical collections where only limited information is recorded.

To browse and download data you do not need an account. If you would like to add or edit data an account can be provided. The Steering group makes the decision. To apply for an account please get in touch with the database curator Kari Lintulaakso.

What is an MN unit? MN is a biochronological system that gives a relative age for European Neogene localities based on mammalian faunal assemblages. The system was introduced by French paleontologist Pierre Mein in 1975 and has undergone multiple revisions and updates. Individual MN units are characterized either by first or last occurrence of selected taxa or by similarity to reference localities (reference faunas). The current age boundaries for MN units in NOW are set according to Hilgen et al. (2012). Does NOW use the MN system as the time framework for the Neogene? NOW does not use the MN system as a framework, but MN happens to be the most frequently encountered time reference unit for European localities and historically has received a lot of attention in NOW. Globally NOW uses many different time frameworks. I have heard that North American localities in NOW are not comparable to Eurasian localities, is that so? Yes, North American localities have been aggregated during the data development process; the raw data have not been preserved. Therefore, North American localities may seem to have richer faunal lists than European ones. In addition, relative ages for North American and European data have been determined using slightly different methods. Users should be mindful of these differences when doing faunal richness comparisons across continents. NOW is a public database, what are log-ins for? Just as in physical fossil collections, a small portion of data might remain private while the researcher or team that collected it is working on it, or a portion of data compiled from public sources needs extensive curatorial attention and harmonization before releasing it for public use. NOW does not accommodate permanently private user data; all data initially entered as private are meant to become public in a few years.

I am very interested in fossils and paleontology, how can I contribute to NOW? Great! Please contact us and let us know how you think you could contribute. You may contact any NOW member you might know or else database curator Kari Lintulaakso.

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Chapter 4

Siwalik Rodent Assemblages for NOW: Biostratigraphic Resolution in the Neogene of South Asia Lawrence J. Flynn, Michèle E. Morgan, John C. Barry, S. Mahmood Raza, Iqbal Umer Cheema, and David Pilbeam

Abstract The NOW database makes knowledge of extinct mammals available broadly, especially for temperate-latitude Eurasia. We add dated biostratigraphic information for the subtropical Siwaliks of the Potwar Plateau, Pakistan. Careful stratigraphic control yields dating precision to 105 yr for most Miocene age Siwalik localities. This improves correlation of local biostratigraphic ranges and turnover with abiotic events and change in other biogeographic regions. As a complement to occurrences, relative abundances of fossil rodent species are tracked for well-represented Middle and Late Miocene localities sorted in 100 kyr time bins. Siwalik rodent associations represent a succession of widespread metacommunities that are distinctive for South Asia. The Middle Miocene Antemus chinjiensis metacommunity (13.8–12.7 Ma) is replaced by crown murine (12.4– 10.1 Ma) and later Karnimata darwini (9.7–8.0 Ma) metacommunities dominated by abundant murine species. The replacement corresponds to fundamental innovations in dentition and mastication that gave early murines a competitive advantage over other muroids.

L. J. Flynn (&)
J. C. Barry
D. Pilbeam Department of Human Evolutionary Biology, Harvard University, 11 Divinity Avenue, Cambridge, MA 02138, USA e-mail: [email protected] J. C. Barry e-mail: [email protected] D. Pilbeam e-mail: [email protected] M. E. Morgan Peabody Museum of Archaeology and Ethnology, Harvard University, 11 Divinity Avenue, Cambridge, MA 02138, USA e-mail: [email protected] S. M. Raza National University of Medical Sciences, The Mall, Rawalpindi, 46000, Pakistan I. U. Cheema House 309, Gali 53, G-10/4, Islamabad, Pakistan










Keywords Rodents Biochronology Pakistan Metacommunity




Competition




India

Introduction The New and Old Worlds (NOW) database of fossil mammals began as a compilation of taxonomic and contextual information on Eurasian Neogene localities, initially emphasizing Europe (Fortelius et al., 2023). It is increasingly global in scope, and one current goal of the Harvard-based project on the Siwaliks of Pakistan is to integrate an extensive biostratigraphic record from South Asia into the NOW database. This record expands the NOW database in a number of domains. The Siwalik sequence of the Potwar Plateau is well-calibrated, and its team of investigators applies updated and uniform taxonomy, allowing fine-grained analyses of taxon histories, biogeography, and dispersal events across the region. Our dataset for South Asia is based on constrained sets of continental deposits in Pakistan, and differs from simple compilations of fossil occurrences spanning the area of the entire Indian Subcontinent. Our best fossil record is from the Potwar Plateau of northern Pakistan, a region of some 20,000 km2, well-defined geologically and geographically, and comprising several thousand meters of primarily fluvial deposits in the stratotype areas of the Miocene Siwalik Kamlial, Chinji, Nagri and Dhok Pathan formations. These deposits entomb multiple, superposed mammal assemblages that, during the Miocene, accumulated at subtropical latitude (