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Fabio Prezoto Fabio Santos Nascimento Bruno Corrêa Barbosa Alexandre Somavilla  Editors Neotropical Social Wasps Basic and Applied Aspects Neotropical Social Wasps Fabio Prezoto Fabio Santos Nascimento Bruno Corrêa Barbosa • Alexandre Somavilla Editors Neotropical Social Wasps Basic and Applied Aspects Editors Fabio Prezoto Fabio Santos Nascimento Universidade Federal de Juiz de Fora Universidade De Sao Paulo Juiz de Fora, Minas Gerais, Brazil Ribeirão Preto, São Paulo, Brazil Bruno Corrêa Barbosa Alexandre Somavilla Universidade Federal de Juiz de Fora Instituto Nacional de Pesquisas da Juiz de Fora, Minas Gerais, Brazil Amazônia Manaus, Amazonas, Brazil ISBN 978-3-030-53509-4 ISBN 978-3-030-53510-0 (eBook) https://doi.org/10.1007/978-3-030-53510-0 © Springer Nature Switzerland AG 2021 This work is subject to copyright. All rights are reserved 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. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Foreword Vespidology in Brazil: An Appreciation This is a landmark volume for anyone interested in social wasps and social insects in general. It contains authoritative chapters on a very wide range of topics, from keys for the identification of genera to specific aspects of morphology, ecology, behavior, and evolution. Most of the chapters feature the social wasps of Brazil through the eyes of Brazilian researchers and others who have worked in Brazil or jointly with Brazilian colleagues. The first chapter of the volume describes the lives of some important figures in the history of Brazilian studies of social wasps. Some of them are mentioned in this preface, where I take advantage of the chance to write a personal appreciation of that history. It would be difficult to overstate the importance of research in Brazil for the his- tory of social-wasp science, especially for studies of Neotropical species. I first began to appreciate this when preparing for fieldwork in Colombia on tropical wasps in the middle of the twentieth century (1964–65). Among the few publica- tions I took along as guides were copies of monographs by Adolpho Ducke (1876–1959) (Fig. 1), the Austro-Hungarian Trieste-born Brazilian biologist, who, before turning to botany (see Egler 1963), was an entomologist at the Museu Paraense (now Goeldi Museum) in Belem. Even though Ducke’s 21 papers with information on social wasps were published many years earlier (during the period 1904–1918), and in three different languages (German, French and Portuguese), they were clearly written and well illustrated. Some contained photographs and drawings of nests, for Ducke considered nests important evidence for studies of vespid phylogeny. His monographs also contained information on natural history and behavior not available elsewhere. Richards and Richards (1951, p. 2) paid trib- ute to “the great Adolfo Ducke, whose abilities as a naturalist, collector and taxono- mist were alike unrivalled” and whose 1910 paper “is still the foundation of all work on S. American POLYBIINAE.” Largely because of Ducke’s papers I still feel a sentimental attachment to now obsolete names of Neotropical social wasp genera such as Nectarina, Tatua, and Gymnopolybia (now Brachygastra, Epipona, and v vi Foreword Fig. 1 Adolpho Ducke (1876– 1959). (From Egler 1963) Agelaia, respectively). And I regret the loss of the sonorous name Marimbonda (now Leipomeles) assigned by Richards (1978) to two Brazilian species, one dis- covered by Richards in the Mato Grosso and the other named by Ducke. In addition to Ducke’s publications, I used early publications by other Brazilians on Brazilian social wasps. Some of those with information on castes and reproduc- tion were by Hermann von Ihering, a German-born naturalized Brazilian, (1850–1930) and his son Rodolpho (1883–1939). (For references to their work on social wasps see Richards and Richards (1951) and Chap. 1 of the present volume.) Like Ducke, the von Iherings published on many biological topics other than ento- mology and were primarily based in museums, not in teaching institutions. Similarly, R. L. Araujo, who published several important taxonomic papers on Brazilian social wasps, including notes on nests in the 1940s, was based at the Institute of Biology of São Paulo, dedicated primarily to agricultural research and outreach rather than teaching. Jose Francisco Zikán (1881–1949), who published monographs on the taxonomy of Mischocyttarus and other social wasps beginning in 1935 (see refer- ences in Ross and Matthews 1991), was employed as a naturalist of the Parque Nacional do Itatiaia (from introduction to Zikan 1949 by W. Duarte de Barros). In a paper published posthumously (Zikan 1949) he described several species of Mischocyttarus that he believed to be parasitic. To my knowledge this idea has not subsequently been investigated. The Brazilian authors of the present volume are in large part the second and third intellectual descendants of a later generation of university-based social-wasp researchers who began their work in the 1960s, including Professors Ronaldo Zucchi, Vilma Maule Rodriques, Vera Lígia Letízio-Machado, and Carminda do Cruz-Landim. At about the same time there was an influx of important social-wasp researchers from outside. At least some of this surge of activity was due to the pro- motion of interest in social insects by the charismatic and energetic Brazilian hon- eybee geneticist Warwick Kerr. It was Kerr who encouraged and advised V.M. Rodriquez, for example, beginning when she was an undergraduate and also during her doctoral work; Kerr was also host to a 1964 visit to Brazil by William D. Hamilton, who then met Rodriquez while he was beginning work on social wasps. Hamilton shared information on the natural history of wasps and supplied Foreword vii Fig. 2 O. W. Richards (1901–1984) at the campo-cerrado boundary near the Mato Grosso base camp of the Xavantina- Cachimbo expedition of the Royal Society of London and the Royal Geographical Society, March 1968. Richards, a noted naturalist and ecologist, is contemplating a plant, which he identified as Paepalanthus (Eriocaulaceae). (Photograph by Maude J. Richards; sent by O.W. Richards to the author in 1971) difficult-to-obtain reprints of articles (Rodriquez, 1968). Hamilton was in Brazil when he did the final revisions on his influential 1964 papers (Hamilton 1964a, b) on the evolution of social behavior by what was later called “kin selection.” While in Brazil he divided the original manuscript into two parts and added his own obser- vations on Brazilian social wasps (see Hamilton 1996, p.29), including especially Polistes. Hamilton’s papers appeared in print while he was still in Brazil and, coin- cidentally, while I was in Colombia studying Polistes. Hamilton’s mention of Brazilian Polistes attracted my attention and enabled me to immediately attempt the first field tests of Hamilton’s ideas (see West, 1966, 1967). O. W. Richards, a leader in taxonomic and biological studies of social wasps (see especially Richards and Richards 1951, Richards 1978) made three trips to Brazil (Richards, 1978, p. 1). The first and longest was in 1968 when he spent 14 weeks at a base camp in the central Mato Grosso maintained by the Royal Society of London- Royal Geographical Society Xavantina-Cachimbo expedition (Fig.  2). That research, and collections made by W.D. Hamilton at the same site, led to the deci- sion by Richards to write a monumental treatise on the social wasps of the Americas (Richards, 1978). He returned to Brazil in 1970 to study type specimens of Mischocyttarus species named by Zikán at the Universidade Federal do Paraná, where his host was Padre J. S. Moure. Then, in 1971, Richards travelled to São Paulo to study the historic collections at the Museu de Zoologia da Universidade de São Paulo, hosted by its director Paolo E. Vanzolini. Richards was 77 years old when the book was finally published. He had typed the entire manuscript himself, as I learned from conversations with Richards. The success of such visitors always depends on the experience and hospitality of their hosts, another notable contribution of Brazilian scientists. The hospitality of rural landowners has also facilitated fieldwork on the social wasps. Robert Jeanne, for example, spent more than a year in Brazil as a graduate student. His doctoral viii Foreword thesis is now well known as a classic study of social wasp Mischocyttarus drewseni. In his monograph on that species he acknowledged not only the help of Brazilian scientific colleagues but also the indispensable role of the Hagmann family at Fazenda Taperinha, located on the Rio Ayayá, a branch of the Amazon east of Santarém, Pará. Jeanne wrote that their “assistance, patience, understanding, sym- pathy, encouragement and hospitality were such as I have never experienced and shall not soon forget” (Jeanne 1972, p.141). This volume features the special contribution of modern research on social wasps in Brazil. It shows how Brazilian scientists have capitalized on the diversity of spe- cies available for study, the opportunity for in-depth studies by people actually liv- ing where the wasps live, their special open-ness to international collaborations, and their own long and rich tradition of research on social wasps. Mary Jane West-Eberhard Smithsonian Tropical Research Institute c/o Escuela de Biología, Universidad de Costa Rica San Pedro, Costa Rica e-mail: [email protected] References Egler W (1963) Adolpho Ducke – traços biográficos, viagens e trabalhos. Boletin do Museu Paraense Emílio Goeldi. Nova Série. Botánica No. 18:15–130 Hamilton WD (1964a, b) The genetical evolution of social behavior. I, II. J Theor Biol 7:1–16, 17–52 Hamilton WD (1996) Narrow roads of gene land, Volume 1. W.H. Freeman and Company, New York Jeanne RL (1972) Social biology of the Neotropical wasp Mischocyttarus drewseni. Bull Museum Comparative Zool 144(3):63–150 Richards OW (1978) The Social wasps of the Americas excluding the Vespinae. British Museum, London RichardsOW, Richards MJ (1951) Observations on social wasps of South America (Hymenoptera, Vespidae). Trans R Entomol Soc, London 102:1–107 Rodriguez VM (1968) Estudo sobre vespas sociais do Brasil (Hymenoptera, Vespidae). Doctoral Thesis, Universidade de Campinas, Rio Claro Ross KG, Matthews RW (eds) (1991) The social biology of wasps. Comstock Publishing Associates, London West MJ (1966) The nature and determination of castes in Polistes wasps. Am Zool 6(4):Abstract 346. West MJ (1967) Foundress associations in polistine wasps: dominance hierarchies and the evolu- tion of social behavior. Science 157:1584–1585 Zikán JF (1949) O genero Mischocyttarus Saussure (Hym. Vespidae), com a descricao de 82 espe- ies novas. Boletin Parque Nacional do Itatiaia 1:1–251 Prologue The Neotropical region has the greatest diversity of social wasps in the world, and this is the reason that, throughout the years, attracted the attention of specialist researchers, who investigated different biological, ecological, behavioral, and evo- lutionary aspects using the species of these social insects as a model. In recent decades, studies with Neotropical social wasps have increased signifi- cantly, as this group offers fertile ground for many researchers; however, the results of these works are fragmented in thousands of scientific articles, thus hindering an insight into the state of art of the taxon. In this way, the aim of this book is to share the knowledge obtained by several research groups around different countries (like Belgium, Brazil, Colombia, Costa Rica, Japan, New Zealand, Portugal, the UK, and the USA) whose efforts resulted in a huge amount of information on social wasps. This book provides an updated overview of different aspects of Neotropical social wasps and also pays a tribute to pioneering work, rescuing the natural history of these wasps and pointing out trends in different areas of research. We hope that the experience shared in this book gives readers an exciting glimpse into the fascinating history of Neotropical social wasps, thereby inspiring a new generation of motivated vespidologists. Best regards, The Editors ix Contents 1 A Brief Review of Studies on Social Wasps in Brazil . . . . . . . . . . . . . . 1 Sergio R. Andena, Luis Filipe Lopes, James M. Carpenter, and Orlando T. Silveira 2 The Evolution of Swarm Founding in the Wasps: Possible Scenarios . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Robert L. Jeanne 3 The Foraging Behaviour of Neotropical Social Wasps . . . . . . . . . . . . 47 Mateus Detoni and Fábio Prezoto 4 The Choice of Sexual Partner in Social Wasps . . . . . . . . . . . . . . . . . . . 71 André Rodrigues de Souza, Camila Folly Baptista, Fábio Santos Nascimento, and José Lino-Neto 5 Nesting Habits of Neotropical Social Wasps . . . . . . . . . . . . . . . . . . . . . 85 Bruno Corrêa Barbosa, Tatiane Tagliatti Maciel, and Fábio Prezoto 6 Castes and Polymorphisms in Neotropical Social Wasps . . . . . . . . . . 99 Marjorie da Silva, Sidnei Mateus, and Fernando Barbosa Noll 7 The Biology of Swarm-Founding Epiponine Wasp, Polybia paulista . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 Kazuyuki Kudô 8 Causes and Consequences of Reproductive Conflicts in Wasp Societies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 Cintia Akemi Oi, Tom Wenseleers, and Ricardo Caliari Oliveira 9 Post-embryonic Development in Brazilian Social Wasps . . . . . . . . . . 179 Edilberto Giannotti and Agda Alves da Rocha 10 Evolution and Adaptation of the Wings and Mandibles of Neotropical Social Wasps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195 Carlos E. Sarmiento and Orlando Tobias Silveira xi

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