Systematics, bionomy, and metamorphosis of Coleoptera (Insecta): Papers celebrating the 80 th birthday of Cleide Costa

. We present a brief biography of Dr. Cleide Costa, eminent entomologist from Museu de Zoologia da Universidade de São Paulo (MZUSP). She has been dedicating the last six decades to the study of adults and immatures of Coleoptera. Dr. Costa is the pioneer in collecting and rearing immature beetles in Brazil, being responsible for establishing the most extensive Latin American collection of reared immatures Coleoptera. We discuss central aspects of her personal history, as well as career landmarks and achievements. A compilation of taxa introduced to science by her, taxa named in her honor, and a full list of her scientific, educational and cultural production are provided in chronological order. More than a biographical account, this publication is an acknowledgment of Dr. Cleide Costa’s legacy to entomology.

This special volume of Papéis Avulsos de Zoologia (PAZ) is dedicated to Dr. Cleide Costa, a highly prolific beetle taxonomist from the Museu de Zoologia da Universidade de São Paulo (MZUSP). Her list of contributions comprises over 166 publications, including peer-reviewed manuscripts, books, book chapters, and science outreach articles. Dr. Costa's comprehensive work encompasses diverse fields of biology such as natural history, comparative morphology of adults and immatures, phylogenetics, and taxonomy in several groups of insects, most notably Coleoptera. She pioneered the study of immature insects in Brazil. A highlight of her career is a volume on the beetle larvae from Brazil ("Larvas de Coleoptera do Brasil"), considered amongst the most relevant publications in Brazilian entomology.
Dr. Costa described over one hundred new taxa across different taxonomic ranks, from species to family-level. She contributed to her home institution in several academic and administrative roles, including student mentoring, teaching, and erecting a world-class collection of immature beetles.
This editorial provides an overview of Dr. Costa's academic trajectory, starting from her early career as an entomology student to some of her most significant achievements and discoveries as a researcher. We also emphasize the pivotal relationship with close colleagues whose collaborative ties contributed to her outstanding accomplishments.
On the occasion of her 80 th birthday, and in recognition of her exceptional career, we dedicate this volume to Dr. Costa, which introduces 35 original papers in 25 Coleoptera families. These manuscripts encompass most of the subjects and fields of study Dr. Costa has comprehensively explored in her career, featuring 53 new taxa, most of which are named in her honor. The distinguished team of experts who contributed with high-quality papers is a testimonial of her legacy and the esteem that her colleagues afford her.
This publication is part of the academic activities circumscribed to the "VI Simpósio de Coleoptera" occurring in the "XXXIII Congresso Brasileiro de Zoologia", which will be held in Águas de Lindóia, São Paulo, Brazil, March 2 nd -6 th , 2020. On this occasion, part of the Brazilian scientific community will acknowledge Dr. Costa in person for her contributions to our field. Her interest in entomology arose early as an undergraduate. Under the supervision of Zoology Professor Luiz Dino Vizotto, Cleide conducted a project on rearing gyrinid beetles in the laboratory. Such a study would re-quire the further assistance of a coleopterist with taxonomic knowledge. Seeking to learn more about her subject, Cleide started a close correspondence with eminent entomologist Dr. Ubirajara Ribeiro Martins, who advised her on the fundamentals of taxonomy and beetle identification (Fig. 2). The growing experience in Coleoptera and the new challenges stemming from these studies prompted her to seek closer guidance from a specialist, and thus Cleide would attend the DZ quite often, to work with Dr. Martins. These early visits were the beginning of a long and prolific career at MZUSP, which strongly influenced the field of entomology in Brazil.
One year after initiating the internship at DZ, Cleide's unique talent for beetle systematics earned her a position of curator of the entomology collection. After she became a staff member, Cleide started her graduate  Influenced by her early studies on the life cycle of insects, and concerned about the scarcity of data on beetle immatures, especially from the Neotropics, Cleide set out an ambitious goal: to study the morphology, bi-ology, and metamorphosis of Coleoptera. Under Cleide's leadership, in countless collecting trips and fieldwork, immatures samples were collected by herself and her students, colleagues, and collaborators. As a result of a long-lasting routine of collecting and rearing, Cleide assembled the most extensive Latin American collection of laboratory-reared immatures of Coleoptera, which is  also one of the world's most important holdings of its kind. Such endeavor generated an outstanding and innovative series of papers and books on immature beetles (series named "Larvas de Coleoptera Neotropicais", or "Larvae of Neotropical Coleoptera"). Early on this project, Costa advised on the importance of immature traits to systematics, therefore teaching how to collect and rear beetle larvae in the laboratory efficiently. Her efforts came to fruition, as most of her students continued to pursue this subject, which in turn influenced new generations of entomologists. In Brazil, the study of immature morphology and development is increasingly popular and appreciated among contemporary experts. Apart from research and curation, part of her professional attributions at MZUSP included advising graduate and undergraduate students, and teaching entomology classes for undergraduate and graduate courses at the Universidade de São Paulo (USP) and other Brazilian and international institutions. Cleide's professional activities extend beyond the real of Brazilian academia. She consistently traveled abroad to teach, participate in congresses, and scientific exchange events. Her tight relationship with foreign colleagues resulted in a vast network of external contributors, which helped introduce novel techniques and experimental methods to Brazilian academia, and draw the attention of the international community to Brazilian research.
Cleide was also very active in the administration and discussions about the future of her home institution. In 1969, the Museu de Zoologia da Universidade de São Paulo was then called "Departamento de Zoologia" ("Department of Zoology"), a branch of the State of São Paulo Secretariat of Agriculture, Industry, and Commerce. In 1969, this section of the Secretariat became an Institute within the State University and was renamed "Museu de Zoologia da Universidade de São Paulo" (MZUSP) (Nearns et al., 2019). Cleide was the head of the Entomology section from 1974 to 1977, and from 1993 to 1999. Notoriously determined, she obtained funds to implement simple changes that have drastically improved the section facilities as well as the conservation of the collection, like the transfer of all insect specimens from drawers with fixed cork bottoms to drawers with unit trays. She also brought innovation to the Coleoptera collections storage system by implementing sliding compact cabinets, which allowed the optimization of space and improved safety for the beetles. For several years Cleide was the head of the "Estação Biológica de Boracéia" ("Boracéia Biological station"), a conservation unit located in the Atlantic Forest managed by the MZUSP, to serve as a center for biological research and student training in fieldwork. In 1996, during the last period under her responsibility, she fully renovated all the buildings, rebuilding the accommodations for researchers and constructing a laboratory and classrooms.
Cleide was an active member in the discussions about the future of the MZUSP and the career path of the museum's members. Until 1986, the position of "Biologist" of the majority of the researchers of the MZUSP was assigned a salary and status that was largely incompatible with the activities developed by these professionals. Unlike researchers and professors from other University departments, apart from doing research and teaching, the biologists were also in charge of curatorial and other technical work activities. Despite the numerous attributions and requirements imposed on the biologists, the recognition of their importance and salaries were much lower than those of their colleagues from other departments. MZUSP researchers protested, then, for parity of career status and wage. Thanks to Cleide's scientific recognition and political influence, this issue was partially solved in 1986, when she convinced the influent USP Professor Fernando Henrique Cardoso (at that time a Senator of Republic who would later be elected President of Brazil), to transmit her appeals to the University President, Professor José Goldemberg. The salary disparity issue plaguing MZUSP researchers was finally solved. However, their career status situation was still not settled before Cleide's official retirement, when the MZUSP researchers achieved the recognition of equivalent career to those of other researchers and professors from USP. From 1988 to 1995, the University of São Paulo was undergoing a reorganization of their institutes, and the status of the University museums (including MZUSP, as well as history and art museums) was intricate. The University Rectory then appointed Cleide as a representative of the MZUSP community in the discussions about the integration of the museum to the Instituto de Biociências (IBUSP) (see Costa, 1987;Brandão & Costa, 2007).
Cleide was included in the active researcher team of MZUSP until 1999 when she officially retired. Even after retirement, she keeps working as a Senior Professor at the MZUSP, where she still conducts research, advises students, and teaches graduate-level classes.

Cleide as Coleoptera researcher
Cleide's scientific career has followed three overlapping lines of research: systematics, phylogeny and biology of Elateroidea, with an emphasis in Elateridae; study of Coleoptera larvae from Neotropical Region; and systematics, bionomy, and metamorphosis of Coleoptera (see publications below, chronologically listed). Early on, Cleide demonstrated a high amount of carefulness in her papers, which is evident from their high scientific quality and fantastic detail in her illustrations (Fig. 3). In her first published paper (Costa, 1966), she presented illustrations of the male genitalia, mouthparts, antennae, and tarsi, as line drawings covered with Nankin ink. Mr. Giro Pastore took most of the photographs of luminescent elaterids from live specimens. This high standard persisted during all her career, and the quality of the illustrations improved over time. Every time that a novice student arrived in her laboratory, their first task was to learn how to draw. It is a tradition in Cleide's laboratory, followed by her disciples, that every student should illustrate their papers adequately. Costa's early research efforts were related to luminescent elaterids, especially concerning Pyrophorus (e.g., Costa, 1968bCosta, , 1969Costa, , 1971aCosta, , b, c, 1972a) and related genera (Costa, 1975a(Costa, , b, c, 1976(Costa, , 1977a. Her doctoral thesis developed the project "Systematics and evolution of bioluminescent Pyrophorinae of New World (Coleoptera, Elateridae)", concluded in 1972. Many new species were described in the subsequent works, and a new classification for the then subfamily Pyrophorinae was established.
During this period of intense taxonomic works in luminescent elaterids, Cleide has caught attention due to her knowledge and descriptions of their immature forms and natural history (Costa, , 1971a(Costa, , 1972b. This particular interest gave rise to her second and successful line of research: the study of the immature forms of Neotropical Coleoptera. To accomplish this ambitious project, she missed no opportunity to collect beetle larvae and, in 1980, together with contributors, started intensive larvae collections and laboratory rearing of beetles. For over a decade, her works were primarily focused on beetle larvae. As a result of this inaugural project, the knowledge of Coleoptera larvae was greatly intensified, with the publication of dozens of papers, in a series called "Larvae of Neotropical Coleoptera" and one textbook with the state of art of immature beetles from Brazil . One essential aspect of Cleide's papers is that, whenever possible, she introduced the pupae in the descriptions and illustrations. It is common for papers dealing with immatures forms to omit or only briefly describe the pupae. On some occasions, she investigated the possible roles and evolution of pupae and prepupae in Endopterygota (Costa, 1984c;Costa & Vanin, 1985b). Cleide's pioneering in the study of immature beetles, always evidencing the importance of the associated knowledge to the study of systematics, has and still inspires many researchers and students around the world. The book "Larvas de Coleoptera do Brasil" , has been a fundamental reference for larval studies worldwide. Some of the descriptions and figures of larvae and pupae introduced therein or in her series of papers are the single references for many groups and are frequently reproduced in subsequent textbooks and manual of coleopteran larvae (e.g., Booth  The third line of research pursued by Costa essentially merges the precedent lines, comprehensively applying the immatures features sampled in broad systematic works, mostly focused on the superfamily Elateroidea. According to Costa (1971a), larval characters of species of Semiotus Eschscholtz, could be useful to establish a new subfamily for this genus. The subfamily Semiotinae was almost simultaneously erected by Golbach (1970), based on adult features (Costa, 1972b). Even though this subfamily is no longer valid, the monophyletic group that it represents is consistently recovered in different phylogenetic approaches (e.g., Kundrata & Bocak, 2011;Kundrata et al., 2018). If larval characters are robust enough to support generic and subfamilial ranks, they might be in-formative and useful in higher coleopteran ranks. Thus, starting in the 1990s, besides continuing the immatures studies, she also dedicated herself to the systematics of Elateriformia, whenever possible, combining data of immatures and adults. As a result, highly relevant systematics works and major taxonomic revisions were concluded in several groups, such as elaterid tribe Tetralobiini (Costa et al., 1994), the superfamily Byrrhoidea (Costa et al., 1999b), and the family Brachypsectridae (Costa et al., 2006i).
More recently, Cleide has been successfully transitioning to modern methodological approaches, combining detailed morphological studies with molecular phylogenetics. In her most recent paper, a comprehensive survey of immature forms and adults, combined with a molecular phylogeny, resulted in the discovery of a new family of Coleoptera within Elateroidea (Rosa et al., 2020a, b).
Cleide has repeatedly benefited from career grants and has recently been awarded renewals for her research grants to work on new projects and to conclude her upcoming book on the "Neotropical Immature Beetles".

Collecting and rearing beetles in the laboratory -the rise of the Immature Beetles Collection (CIC-MZUSP)
Cleide started collecting and rearing beetle larvae in 1965. At that time, the immature forms of Neotropical beetles were mostly unknown, and even primary literature was unavailable. Thus, rearing beetles in laboratory conditions was an experiential activity. The only proper process to have larvae determined to family level was to obtain adults that would then produce immatures, so that the developmental stages could be reliably associated. Considering that larvae collected without their adult counterparts were not yet identified to family, their behaviors and feeding habits had also to be discovered by trial and error. There was a great effort in keeping all the collected specimens alive, either to allow the larvae to reach full development to adults, or to obtain eggs and larvae from the adults. While rearing in the laboratory, every relevant information regarding their natural history, behavior, molting, and feeding habits was recorded (Fig. 4).
This project to understand and describe the immatures of Neotropical Coleoptera would require intensive fieldwork and daily devotion in the laboratory. In the beginning, Cleide frequently collected at the "Mata do Instituto Botânico", in São Paulo, a vegetation patch that is not far from the MZUSP, accompanied by the museum technician Lázaro Ribeiro Silva. In addition to the material collected by herself, she also received specimen donations, especially bioluminescent larvae and adults, from fellow zoologists such as F. Lane, L. Alonso, B.N. Dias, P. Biasi, P.E. Vanzolini, D.L. Tiemann, J. Bojard, and L. Otero.
She was accompanied in two collecting trips in 1969 by D.L. Tiemann, an amateur entomologist and engineering technician for the United States Navy, who contributed with his knowledge in finding luminescent larvae and females of Phengodidae (railroad worms). Tiemann was a lover of railroad worms and considered himself as "a compulsive rock kicker-over". To collect Phengodidae larvae and larviform females, he scattered tiles and concrete blocks in vacant lots and grassy fields, hoping railroad worms would crawl beneath them. He tried different techniques but concluded that the best way to collect his beloved worms was to search the grass and soil, night after night patiently. Cleide became obsessed with collecting phengodids, and in her fieldworks, every guest was invited to go along her at night, walking through roads and trails, in the dark, searching for bioluminescent insects. With plans of increasing her commitment to the study of larvae and developing a project on immature beetles, from 1977 on, she had an active collaboration with Dr. Sergio A. Vanin (IBUSP), to work in the collecting and also in the publication of data on immature forms. In 1980, she obtained funding from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) ("São Paulo Research Foundation") to initiate an intensive work of larvae collecting and laboratory rearing. This endeavor counted with the constant contribution of dedicated colleagues that were zoologists, botanists, entomologists, technicians, field assistants, and drivers, like Sônia A. Casari Along with her collaborators, Cleide participated in more than seventy expeditions to collect scientific material in various Brazilian ecosystems (Fig. 5).
In 1982 a partnership was commenced between Cleide and the chemist Prof. Dr. Etelvino Bechara (Instituto de Química, IQUSP), a joint project aiming to study the phenomenon of bioluminescence. Together they headed on several journeys to central Brazil, notably to Parque Nacional das Emas (Goiás), frequently accompanied by the driver and excellent field assistant Mr. José Sanches (IQUSP). Their main objective was to massively collect larvae occurring in termite mounds, primarily the luminescent specimens, which would then be used in chemical, biological, morphological, and systematic studies. The hundreds of bioluminescent larvae collected in those expeditions were kept in the laboratory housed in the MZUSP, under Cleide's supervision, and with the close assistance of three technicians, Maria Ines Migliaccio, in charge of the general Coleoptera, and Emília Arasaki and Adriana Pires, who cared for the bioluminescents.
With the growing relevance of her works, Cleide received exceptional support from world specialists in immature beetles, such as Roy A. Crowson and John F. Lawrence, and from specialists in several families, who joined her in her projects and became coauthors in her articles (Fig. 6).
The diaries and notes taken by Cleide in the laboratory and during fieldworks since 1965 are quite descriptive, detailed, and informative. In 1969, during fieldwork in Estação Biológica de Boracéia (EBB), she recorded in her field book the observation of a luminescent larva of Staphylinidae. This would be the first record of light emission in Staphylinidae and the first record in a non-Elateroidea beetle. She knew that this would represent an extremely crucial discovery, in particular for those interested in bioluminescence. Unfortunately, at that time, the larva died amidst molting, and the dissemination of that important record would have to be postponed. Countless other surveys were conducted in the EBB for 13 consecutive years. In 1983 and 1984, 14 years after the first observation, Cleide found that mysterious luminescent staphylinid species at the same spot where it had been before. This time, another four specimens were found and reared in the laboratory for several days, and the same light emission was observed. The staphylinid larva was tentatively identified as a species of Xantholinus Dejean, and this record was finally published, along with a morphological description of the larva (Costa et al., 1986b). The occurrence of luminescence in Xantholinus larva was later confirmed by Rosa (2010) from specimens from Mato Grosso state, Brazil.
Finding and collecting larvae can be easy if one knows their habits and natural history, although, for certain groups with completely unknown larvae, the chase may be a puzzle. Cleide's surveys for immatures collection were strictly planned, with the techniques and collecting methods carefully designed to catch target groups or to investigate the determined area, micro ambient, hosts, etc. However, occasionally some of the most pleasing surprises and important discoveries occur by chance when no one is expecting them.
During one call for inspection to investigate why the Casuarina sp. trees in São Paulo were dying, Édson P. Teixeira, an Agricultural Engineer from the Instituto Florestal de São Paulo, along with Cleide, found adults and larvae of Nosodendron angelus Reichardt, 1973 (Nosodendridae) in the exudate present on the tree's trunk, believed to be caused by the attack of Pantophthalmidae (Diptera) larvae. The immatures of the family Nosodendridae were still unknown. Still, after these observations, a myriad of larvae, pupae, and adults of this species was found in other locations in Casuarinas with exudate (Costa et al., 1986a) The same occurred with larvae of at least two other families that were still unknown: Lutrochidae  and Cneoglossidae (Costa et al., 1999b), collected inside wood underwater in lakes and streams, and those of Artematopodidae , found in ravine soil inside the forest, among others.
The vast collection of Neotropical Coleoptera (Coleção de Imaturos de Coleoptera (CIC-MZUSP)) ( Fig. 4) founded by Cleide and expanded by her and other for several years is deposited at the MZUSP and keeps receiving new contributions every year. The collection is currently com-posed of more than 50,617 specimens, including 20,364 larvae, 9,607 larval exuviae, 3,320 pupae, five pupal exuviae, and 17,317 adults. The collection comprises 103 beetle families, 471 genera, and 662 species, especially from the Neotropical region, most of them identified by specialists in each family (Barbosa et al., 2013).

Travels to study material and meetings abroad and in Brazil
Similar to what has been observed in different countries with colonial roots, the development of the knowledge on the Brazilian biodiversity was pro- foundly delayed in comparison with European countries. Taxonomy is a budding science in Brazil, dating only from the 20 th century. Although our diversity is relatively well represented in national collections, the main obstacles for research in systematics and taxonomy were the lack of reference material with reli-able identification and the absence of fundamental literature.
To compensate for these challenges, the solution found was to travel overseas for an extended period to examine the Old World collections, bringing back home a large amount of undetermined specimens for compar- ison, and to gather the most amount of information as possible, in hopes that it would be enough to enable future studies with Brazilian material. Traveling abroad was unusual for Brazilian scientists in the 1960's and 1970's, especially in light of the historically underfunded governmental institutions.
Thanks to Cleide's proactivity and prominence as a scientist, she was highly successful in obtaining funds for long travels abroad and nationally (Table 1). In her numerous trips and visits to institutions, Cleide was able to study thousands of essential materials, from type specimens to unidentified exemplars, enabling the present and future works for herself and her students' projects. Pursuing literature was an indispensable part of her travels. Whenever photocopies were unavailable, Cleide spent several days in libraries copying papers and books manually.
After collecting literature and analyzing the type material abroad, she visited Brazilian institutions trying to identify and gather the material deposited.
Cleide's good relationship with several curators and researchers abroad fostered the exchange of material and literature from other institutions. Due to these efforts, Cleide was able to build an impressive reference collection at the MZUSP, with numerous specimens identified from direct comparison with their types, and many more new types designated from her discoveries.
While traveling, Cleide always learned about best curatorial practices from other museums, updating her expertise with the most modern methodological and technical methods. Some were successfully applied in her research, as well as in improvements to the museum's organization and facilities.
One of her most prolonged periods abroad was during her Post-Doctoral research in the Biosystematics Research Institute, Ottawa, Canada, and the United States National Museum, Smithsonian Institution, Washington D.C, USA. She had, in 1989, the invaluable opportunity to be in Glasgow, Scotland, to work alongside the distinguished coleopterist Roy A. Crowson and discuss with him the morphology of immature beetles. Later, in 1991, she went to Australia to examine beetle classification with John F. Lawrence, another renowned coleopterist.
Apart from traveling with the specific purpose of studying beetle specimens, Cleide has also attended numerous international scientific meetings, congresses, courses, and conferences (Fig. 7).

Cleide as a University graduate and postgraduate professor and advisor
As a postgraduate professor and advisor, Dr. Costa taught classes in different areas of entomology, such as comparative morphology of adults and immatures, insect systematics, and insect physiology.
Cleide was part of postgraduate programs in several institutions. A few examples of the programs she was part of are "Zoologia" (IBUSP), "Sistemática, Taxonomia e Biodiversidade" (MZUSP), "Entomologia" (Universidade Federal do Paraná, Curitiba) and "Ciências Biológicas" (UNESP, Botucatu). During these courses, she always encouraged students to perform comprehensive investigations on many aspects of entomology, integrating natural history and morphology traits of adults and larvae. She also advised students at the master's, doc- toral, and post-doctoral levels and in several coleopteran families, such as Elateridae, Lycidae, Phengodidae, Passalidae, Scarabaeidae, Geotrupidae, Chrysomelidae, Curculionidae, and Carabidae (Fig. 8). Her former students include specialists in several Coleoptera families ( Table 2).
As part of her activities as professor and due to her expertise in insects, more specifically in beetles, she participated in dozens of graduate committees, including masters and doctoral degrees examining boards, as well as evaluation boards for public hiring committees for the role of tenure-track positions in different universities.

Dissemination of science, taxonomists training and the importance of collections
Most lines of research are only fully accessible and comprehended among a small circle of specialists. Unlike most of her fellows, Cleide was deeply concerned about disseminating science not only among her peers but also among the general public of biologists, early students, non-scientists, and children.
As part of her outreach program, Costa used her beloved bioluminescent beetles to catch people's attention to varying topics in entomology and natural history. She published various articles in magazines and journals directed at children on the chemistry of bioluminescence, insects' life-cycles and morphology, the importance of biodiversity, and others ( Besides formal classes for graduate and undergraduate students, Cleide was also worried about the amount of time necessary to train taxonomists adequately. This led her to organize and teach conferences and basic-level courses of entomology in Brazil and abroad, notably about Coleoptera identification and larval morphology. In one of these courses, held in the MZUSP, Dr. John F. Lawrence taught lessons on Coleoptera morphology and systematics for Brazilian and other South American students and professionals (Fig. 6F).
Another of Cleide's main interests is the importance and future of entomological collections. She authored a series of papers about the state of knowledge   ), a conference with the participation of many Ibero-American researchers dedicated to discussing and proposing the production of inventories and estimates of biodiversity in hyperdiverse insect orders (Fig. 7C).

Honors and awards
In recognition of the importance of her work, Cleide Costa was awarded with the following honors and distinctions: 1988 Award "Alexandre Rodrigues Ferreira" offered by the "Sociedade Brasileira de Zoologia", for the publication of the book "Larvas de Coleoptera do Brasil". 1994 Award "Bunka de Pesquisa" offered by the "Banco Sumitomo & Sociedade Brasileira de Cultura Japonesa" for the publication of the book "Larvas de Coleoptera do Brasil". 2002 Honorary Partner of "Sociedade Brasileira sobre Vetores e Pragas Sinantrópicas". 2015 Tribute from the MZUSP for her "50 years of outstanding performance, dedication, ethics and love for the research and teaching".
The entomological community also recognizes Cleide's immense contributions to the field in publishing high quality systematic and taxonomic works, through her intensive efforts of collecting, describing and archiving specimens, and by her unceasing fondness for forming professionals, teaching and disseminating science. The importance of her contributions can be veri-fied in the enormous repercussions of her scientific papers and textbooks, and the high attendance to her lectures and speeches. Furthermore, her legacy is cherished through all the taxa named in her honor (Table 3).
Once again, the coleopterists' community pays tribute to the outstanding entomologist and colleague Cleide Costa, by participating in this series of papers celebrating her career. This special volume of Papéis Avulsos de Zoologia features articles in numerous areas of Coleoptera research that were cherished by her, such as the description of new taxa, taxonomic revisions, studies in comparative morphology, phylogeny, natural history, and, especially, a several papers on the morphology and metamorphosis of immatures. The participants include experienced and novice coleopterists, old friends and young admirers, and authors from Brazil and from all over the world.
The wide coverage of taxonomic groups of Coleoptera, the diversity of subjects dealt with and the many nationalities of the authors assembled in this volume are a testament to the scope and importance of Cleide Costa's legacy. The following papers intend to demonstrate an appreciation of the legacy she has left to Coleopterology. Table 3. Taxa named in honor of Cleide Costa. Taxa marked with an asterisk are patronyms presented in this special volume.