Museum of Comparative Zoology

Harvard University

Annual Report 2019. 2020 About the MCZ

The Museum of Comparative Zoology at Harvard University is a global center for research and education focused on the comparative relationships and evolution of animal life. The MCZ collections comprise approximately 21 million extant and fossil invertebrate and vertebrate specimens, which are a focus of research and teaching for the MCZ, Harvard, and outside students and researchers.

Contents Feature Story 2 Faculty-Curator Profiles 3 In Memoriam: James J. McCarthy 6 Emeritus Profiles 7 Courses Led by Faculty-Curators 8 Research Making Headlines 12 Highlights from the Collections 15 Projects & Initiatives 18 Awards & Recognition 20 Grant Recipients 22 Publications in 2019 26 Financial Data 29 Personnel 30 Director’s Message Put on your mask, wash your hands, stay six feet apart, and what on Earth is Zoom?

As spring semester began last has come at great cost. Yet, we had January, none of us had any idea of already experienced sadness and how dramatically—and, for some, loss last fall following the passing tragically—our world was about to of two key members of the MCZ change. Just two months later, as we community, Prof. James J. McCarthy were administering midterm exams, and Mr. Robert G. Goelet. Jim finalizing plans for spring break McCarthy was an internationally course field trips, awarding summer recognized biological oceanographer travel grants, and dozens of us were and the MCZ’s eighth director planning field and conference trips, but, as described elsewhere in this Harvard responded to the COVID-19 report, he was perhaps best known pandemic by closing the Cambridge for his accomplishments in the area campus and cancelling all domestic of public policy and for bringing and international travel. With only a the world’s attention to the drastic few days to prepare, curatorial staff consequences of climate change. were instructed to work from home Bobby Goelet was a longtime Jennifer Berglund and all courses—lectures, labs and member of the Museum’s governing James Hanken final exams—were moved online. I board, the MCZ Faculty. To say that Director am intensely proud of the many ways Bobby led a very full life doesn’t come the MCZ community responded close to capturing all that he did and to these challenges—developing experienced. After graduating from innovative software to adapt to online Harvard in 1945 with an AB in history, teaching, transitioning undergraduate Bobby successfully managed his family’s Contents summer projects to remote research investment firm. At the same time, opportunities, and seizing the he was a passionate believer in the Feature Story 2 opportunity to complete collection value of natural history collections and management tasks compatible with collections-based research, and in that Faculty-Curator Profiles 3 home offices. Whether it was embarking capacity he held many distinguished on that cross-country bike ride they’d appointments, including Chair of the In Memoriam: James J. McCarthy 6 always dreamed of, developing skills Board of Trustees of the American Emeritus Profiles 7 as a birder and nature photographer, Museum of Natural History. At MCZ, or creating learning opportunities for Bobby contributed a series of gifts that Courses Led by Faculty-Curators 8 students living thousands of miles enriched our programs, from faculty- Research Making Headlines 12 from campus, MCZ’s faculty-curators curator salaries to the purchase of found ways to fill the void left when specimen cabinets to grants for student Highlights from the Collections 15 research field trips weren’t possible. projects. The Harvard Museums of Science & Projects & Initiatives 18 More so than in any of my previous Culture also successfully migrated 19 years as director, MCZ’s About the cover: Osprey, Pandion Awards & Recognition 20 its public programs online and, in so accomplishments over the past year haliaetus, landing in nest in Cape Cod. doing, expanded its global reach and Photo by Gonzalo Giribet. Grant Recipients 22 are the result of the tremendous “visitations” well beyond what it had Opposite page: Insect eggs come in energy, commitment, hard work and Publications in 2019 26 experienced previously. all shapes, sizes and ornamentation. resilience of our dedicated students, Clockwise from top: Hawaiian fly, Drosophila mimica; two-spotted cricket, Financial Data 29 Responding to the pandemic has staff and faculty-curators. To them I Gryllus bimaculatus; milkweed bug, been MCZ’s preoccupation for most say, thanks for a job well done. I hope Oncopeltus fasciatus; and imperial Personnel 30 hairstreak butterfly,Jalmenus evagoras. of 2020, and while we have been able you enjoy reading about them on the Images by Samuel Church and Seth to manage it successfully, our success following pages. Donoughe.

Annual Report 2019 . 2020 1 Significant Progress and Unexpected Benefits

During early 2020, the world faced a pandemic that was rapidly evolving. When the decision was made to close Harvard's campus in mid-March, the MCZ had only a week’s notice before the shutdown.

According to Linda Ford, director for research that seeks to analyze species of collection operations, “The distribution over time and space. mood was urgent due to the While much of this work has been seriousness of the situation, and all voluminous but routine, some collections staff dedicated their limited time efforts required more creative solutions. The on campus to getting what they majority of specimen data in the Entomology needed so they could effectively collection, for example, is recorded on transition to working from home.” slips of paper pinned beneath the insects. Many curatorial associates quickly These tiny bits of information—many more gathered files and scanned than a century old, handwritten in myriad materials that would be useful languages—often provide the only data for for remote projects for their an insect specimen. During remote work, staff. The way forward was clear: Entomology staff transcribed thousands focus on ongoing data entry and of photographed specimen labels for cleanup efforts forMCZbase , Lepidoptera (butterflies and moths). But the museum-wide collections when labels had challenging handwriting, database containing more than 2.2 million records for over 21 Mark Omura million specimens in 9 research Thanks to focused database efforts and a Japanese visiting collections. These records include species researcher's translation assistance, information, images, geographical data and this record for Lutra lutra whiteleyi in MCZbase is enriched with 3D models. valuable information. Data enhancement and standardization have been tasks for collections staff for many years, but the shutdown provided the unexpected benefit of time to concentrate on the effort. “Typically, curatorial staff only spend a portion of their days were written in non-Latin alphabets or attending to the database,” says James had translations of long-outdated names, Hanken, director of the MCZ, “but curatorial staff enlisted the help of the because of the campus closure, they have global Twitter community, posting the labels been able to dedicate an unprecedented for interpretation by entomologists and amount of time to it.” transcription enthusiasts, often with success. Significant progress has been made during “On one hand, remote work has been difficult the extended period of remote work. Staff because we can’t physically access the across the various collections undertook specimens we use for research and teaching,” numerous projects to improve data Hanken says. “On the other hand, because so At right, challenging labels for Papilio phorcas, above, were quality, with a special emphasis on many of our past activities have emphasized deciphered via an appeal on georeferencing, which assigns GPS coordinates digital data and digital representations Twitter. to locality information described in words in of these specimens, we’ve been able to the database, information especially critical successfully make this transition.”

2 Museum of Comparative Zoology Faculty-Curator Profiles

Andrew A. Biewener Charles P. Lyman Professor of Biology Director, Concord Field Station Prof. Biewener studies the biomechanics and neuromuscular control of animal movement on land and in the air. His goal is to understand general principles that govern the biomechanical and physiological design of vertebrate animals related to movement in natural environments, work with engineers to develop bio-inspired robotic designs and develop improved neuromuscular models for treating human movement disorders.

Scott V. Edwards Professor of Organismic and Evolutionary Biology Alexander Agassiz Professor of Zoology Curator of Ornithology Prof. Edwards' research focuses on the evolutionary biology of birds and related species, combining field, museum and genomics approaches to understand the basis of avian diversity, evolution and behavior. Current projects use genomics technologies to study the evolution of flightlessness and other traits in birds; phylogeography and speciation in Neotropical and Australasian birds; and the genomics of host–parasite coevolution between house finches and a recently acquired bacterial

pathogen, Mycoplasma. Snibbe/HarvardKris University

Brian D. Farrell Monique & Philip Lehner Professor for the Study of Latin America Professor of Organismic & Evolutionary Biology Curator of Entomology Director, David Rockefeller Center for Latin American Studies Faculty Dean, Leverett House Prof. Farrell's research is broadly concerned with the evolution of ecological interactions between host plants and animals and their parasites, such as insects and other tiny consumers. His current projects include applying next-generation sequencing to speciation and phylogenetic studies of associated species, documenting biodiversity in the Dominican Republic, and repatriating digital information from scientific specimens of

Stephanie Mitchell insects and fossils in museums to their countries of origin.

Annual Report 2019 . 2020 3 FACULTY-CURATOR PROFILES

Gonzalo Giribet James Hanken Alexander Agassiz Professor of Biology Professor of Zoology Alexander Agassiz Professor of Professor of Zoology Organismic & Curator of Herpetology Evolutionary Biology Acting Curator of Curator of Malacology Invertebrate Zoology MCZ Director Harvard College Professor Prof. Hanken utilizes Prof. Giribet’s laboratory-based primary research analyses and field focuses on surveys to examine the evolution, morphological evolution, systematics and developmental biology biogeography and systematics. Current © Casey Dunn © Casey of invertebrate areas of research include the evolution of animals, including the use of morphology and next- Catherine Weisel generation sequencing techniques. craniofacial patterning, the developmental basis of morphological novelty and life- Current projects in the Giribet lab include a comprehensive history evolution, biodiversity informatics, and systematics and study of the harvestmen of New Zealand, their systematics evolution of neotropical frogs and salamanders. and biogeography; mollusk phylogenomic projects; and exploring techniques to use degraded DNA from old museum specimens in phylogenomics and population genomics. The lab also works on other projects on systematics and biogeography of arthropods and onychophorans, among other groups.

Hopi E. Hoekstra Professor of Organismic & Evolutionary Biology Professor of Molecular & Cellular Biology Alexander Agassiz Professor of Zoology

Curator of Stephanie Mitchell Mammalogy Howard Hughes George V. Lauder Medical Institute Henry Bryant Bigelow Professor of Ichthyology Investigator Curator of Ichthyology Harvard College Professor Kris Snibbe/HarvardKris University Prof. Lauder's research focuses on the biomechanics of fishes Prof. Hoekstra combines field and laboratory work to and the development of robotic models for studying aquatic understand the evolution of mammalian diversity. Her locomotion. research focuses on the genetic basis of morphological and behavioral variation, primarily in rodents, identifying both His current studies focus on the structure and function of the evolutionary processes and the molecular mechanisms shark skin and other fish surface structures and research responsible for traits that help organisms survive and with various robotic fish models, including a tuna robot. reproduce in the wild. Research in the Hoekstra lab Additional broad interests include biological fluid mechanics integrates ecological, behavioral, genetic, developmental and theoretical approaches to the analysis of form and and neurobiological approaches. function in organisms.

4 Museum of Comparative Zoology FACULTY-CURATOR PROFILES

Javier Ortega-Hernández Assistant Professor of Organismic & Evolutionary Biology Curator of Invertebrate Paleontology Prof. Ortega-Hernández's research focuses on the evolution of metazoans that first appeared and rapidly diversified during the Paleozoic Era (ca. 541 to 251 million years ago). His group specializes in the study of exceptionally preserved Cambrian and Ordovician fossil biotas around the world, with Isabella Kirkland a strong interest in the morphology, phylogeny and development of panarthropods and their relatives. The Naomi E. Pierce lab combines traditional paleontology with cutting- Sidney A. & John H. Hessel Professor of Biology edge techniques to investigate exceptional fossils, test Curator of Lepidoptera macroevolutionary hypotheses through deep time, and better understand the origin of the major animal Prof. Pierce’s research focuses on the behavioral ecology of groups that have shaped the biosphere for more than species interactions, particularly the coevolution between 500 million years. plants, pathogens and herbivores, and symbioses between ants and other organisms.

Her laboratory integrates approaches from phylogenetics, ecology, behavior, genomics and comparative methods to investigate patterns of reciprocal adaptation and diversification exhibited by organisms that live in close association with each other.

Mansi Srivastava John L. Loeb Associate Professor of the Natural Sciences Curator of Invertebrate Zoology Prof. Srivastava’s research focuses on understanding

Paul Whatmough Paul the evolution of animal development and Stephanie E. Pierce regeneration. Her group Thomas D. Cabot Associate Professor of Organismic & utilizes the three-banded Evolutionary Biology panther worm, Hofstenia Curator of Vertebrate Paleontology miamia, which she has Prof. Pierce’s research is focused on major morphological developed as a new acoel BradburyTim and ecological transitions in vertebrate evolution through model system. Acoels represent the sister-group to all an examination of the fossil record. Her work tends toward animals with bilateral symmetry, which allows the study 3D modeling and experimentation of the musculoskeletal of genetic mechanisms that span 550 million years of system, with particular attention to the link between form animal evolution. Current projects in the lab range from and function. Current projects focus on two key events identifying gene regulatory networks for regeneration to in the fossil record, the fish-to-tetrapod and “reptile”-to- determining the embryonic origins of pluripotent stem cells mammal transitions. to understanding the origins of bilaterian nervous systems.

Annual Report 2019 . 2020 5 IN MEMORIAM

James J. McCarthy A mid-winter’s day grew even colder on December 11, 2019, when Prof. James J. McCarthy died following a years-long battle with pulmonary fibrosis. So ended a remarkable career of scholarship and service to the Museum of Comparative Zoology, where Jim had been the Alexander Agassiz Professor of Biological Oceanography since 1980 and the Museum’s director from 1982 to 2002. Yet, his professional accomplishments had profound impacts well beyond MCZ, Harvard and even the United States. Among many distinctions, Jim led a working group of the third Intergovernmental Panel on Climate Change, whose report identified the global scale of climate impacts already evident in ecological systems, as well as the vulnerability of people in both developed and developing countries who are without sufficient adaptive measures to deal with the impacts of impending extreme events and sea level rise linked to climate change. As a result of this report and related efforts, the IPCC

shared the 2007 Nobel Peace Prize with Al Gore. C. Cambon Jim was a master at working behind the scenes to effect substantive change in the status quo. Indeed, he may be the one person over the last 40 years most responsible for the current configuration of environmental biology at Harvard, in all its many manifestations. He was, for example, the main driving force, together with a small number of faculty colleagues, responsible for the founding of the Harvard Museum of Natural History in the late 1990s. Today, the HMNH manages the public programs of the MCZ, the Harvard University Herbaria, the Botanical Museum, and the Mineralogical and Geological Museum. As MCZ director, Jim championed MCZ’s public outreach, including educational programs for Boston and Cambridge schoolchildren, evening science lectures, and the overhaul of many of MCZ’s outdated exhibit galleries. These efforts reflected his ardent belief that scientists have an obligation to share their discoveries with the public—both young and old—and that museums, in particular, should play a dominant role in public education. He was particularly fond of MCZ’s natural history travel program, which literally was the first of its kind in North America and possibly the world. Beginning with a single whale-watching trip to the Gulf of California in 1975, the program grew under Jim’s encouragement. By 1990 the program was sponsoring trips to Central and South America, the Galapagos, Africa, Antarctica, New Zealand, French Polynesia, Bali and the North Sea. Many of the trips were led by Jim and his wife Sue. To many of us in the MCZ, however, what we will remember most about Jim McCarthy was his humanity, kindness and basic decency. He always sought to bring out the best in others, be they friends, relatives, students or colleagues, and could always be counted on for sage advice and thoughtful words of encouragement. We will miss him. Our planet will miss him.

Peter Wiebe Peter —­James Hanken

6 Museum of Comparative Zoology Emeritus Profiles

A. W. “Fuzz” Crompton Fisher Professor of Natural History, Emeritus Prof. Crompton, former curator of Mammalogy, was the director of the MCZ from 1970 to 1982, having served as director of both the Peabody Museum of Natural History at Yale University and the South African Museum in Cape Town.

His primary research interests include the Richard C. Lewontin origin and evolution of Professor of Biology, Emeritus

Justin Ide Justin mammals, functional Alexander Agassiz Professor of Zoology, Emeritus anatomy, and neural An evolutionary geneticist, Prof. Lewontin pioneered control and evolution of feeding in recent and fossil vertebrates. the field of molecular population genetics by merging Prof. Crompton received two Guggenheim fellowships for his molecular biology and evolutionary theory, as well as research on vertebrate paleontology and functional morphology, the philosophical and social implications of genetics and in 2011 received the Romer-Simpson Medal from the Society and evolutionary theory. of Vertebrate Paleontology. Among his many books are The Genetic Basis of Evolutionary Change; Biology as Ideology: The Doctrine Edward O. Wilson of DNA; Human Diversity; and The Triple Helix: Gene, Honorary Curator in Organism and Environment. Entomology Pellegrino University Professor, Emeritus Robert M. Prof. Wilson is Woollacott considered the founder Professor of Biology, of sociobiology and has Emeritus developed the basis of Prof. Woollacott modern biodiversity joined the faculty conservation. He has in 1972 and received many of retired in 2018. the world’s leading His teaching prizes in recognition and research of his research, Beth Maynor Finch focus is on the creative literature and reproduction environmental activism and holds 40 honorary doctoral degrees of marine from universities and colleges in America and Europe. Prof. invertebrates, Wilson was awarded two Pulitzer Prizes for his books The Ants especially larval (1990, with Bert Hölldobler) and On Human Nature (1978). He biology, as well received the TED Prize in 2007, where he articulated the concept as human impacts of the Encyclopedia of Life, and the Hubbard Medal in 2013, the on life in the sea. rarely given highest award of the National Geographic Society.

Annual Report 2019 . 2020 7 Courses Led by Faculty-Curators Academic Year 2019–2020 Naomi Pierce Naomi

OEB 155R: Biology of Insects Organismic and Evolutionary OEB 57: Animal Behavior Biology Naomi E. Pierce (and Bence P. Olveczky) A review of the behavior of animals under OEB 10: Foundations of Biological Diversity natural conditions, with emphasis on both Brian D. Farrell (and Elena Kramer, Peter Girguis) mechanistic and evolutionary approaches. An integrated approach to the diversity of life, emphasizing how chemical, physical, OEB 126: Vertebrate Evolution OEB 167: Herpetology genetic, ecological and geologic processes Stephanie E. Pierce contribute to the origin and maintenance of A comprehensive survey of the origin biological diversity. and evolution of vertebrates through an examination of the fossil record. OEB 11: Introduction to Tropical Biology The primary focus is on major events Gonzalo Giribet (and David Haig) in Earth’s evolutionary history, with an Introduction to concepts of tropical emphasis on anatomical and physiological biology and tropical biodiversity with transformations in fish, amphibians, reptiles, a focus on the ecology, physiology and birds and mammals. diversity of rainforest and tropical coral reef ecosystems. OEB 150: Exceptional Paleobiological Insights into Animal Evolution OEB 51: Biology and Evolution of Javier Ortega-Hernández Invertebrate Animals Explores the importance of soft-tissue Gonzalo Giribet (and Cassandra Extavour) preservation in the rock record through an Introduction to invertebrate diversity, covering overview of major exceptional fossiliferous the development, adult anatomy, biology and sites around the world and throughout evolutionary relationships of the main animal the Precambrian to Mid-Phanerozoic, with phyla including sponges, mollusks, annelids particular emphasis on the evolutionary

Inbar Maayan and arthropods, among others. history of invertebrate animals.

8 Museum of Comparative Zoology COURSES

OEB 155R: Biology of Insects OEB 307: Biomechanics, Naomi E. Pierce Physiology and Musculoskeletal An introduction to the major groups Biology of insects. The life history, morphology, Andrew A. Biewener physiology and ecology of the main taxa are OEB 310: Metazoan Systematics examined. Topics include the phylogeny of Gonzalo Giribet terrestrial arthropods, an analysis of abiotic OEB 320: Biomechanics and and biotic factors regulating populations, Evolution of Vertebrates and the use of insects in biological control. George V. Lauder OEB 167: Herpetology OEB 321: Evolution of James Hanken Regeneration and Development An introduction to the biology of Mansi Srivastava amphibians and reptiles. Lectures and OEB 323: Advanced Vertebrate laboratories examine the morphology, Anatomy systematics, natural history, behavior, Stephanie E. Pierce ecology, evolutionary relationships and biogeography of all major taxa. OEB 334: Behavioral Ecology Naomi E. Pierce OEB 173: Comparative Biomechanics OEB 341: Coevolution Andrew A. Biewener Brian D. Farrell An exploration of how animals and plants contend with their physical environment, OEB 355: Evolutionary Ortega-Hernández Javier Developmental Biology considering their biomaterial properties, OEB 150: Exceptional structural form and mechanical interaction James Hanken Paleobiological Insights into Animal Evolution with the environment. OEB 362: Research in Molecular Evolution Scott V. Edwards OEB 190: Biology and Diversity of Birds Scott V. Edwards OEB 370: Mammalian Evolutionary Genetics An introduction to the biology of birds. Hopi E. Hoekstra Covers the fossil record and theories for avian origins, physiology and anatomy, OEB 399: Topics in Organismic and systematics, speciation processes, behavior, Evolutionary Biology Scott V. Edwards OEB 11: Introduction to vocalizations, demography and conservation. Tropical Biology OEB 213: Evolutionary Convergence, Mass Extinctions and the Shape of Life Javier Ortega-Hernández An examination of how processes acting through deep time affect fundamental biodiversity patterns, including topics such as the origin of animals, the rapid diversification of major clades and the impact of extinction. Graduate Courses of Reading and Research OEB 275R: Phylogenetics and Phylogeography in the Era of Genomics Scott V. Edwards OEB 306: Invertebrate Paleobiology and Evolution

Javier Ortega-Hernández Gonzalo Giribet

Annual Report 2019 . 2020 9 COURSES Gonzalo Giribet

OEB 11: Introduction to Tropical Biology Freshman Seminar FRSEMR 41U: Museums James Hanken FRSEMR 22T: Why We Animals Sing Traces the history of museums from their Brian D. Farrell beginnings centuries ago as personal Investigates the sounds and structures collections maintained by private individuals of different kinds of acoustic animals— to the modern institutions of today. including birds, mammals, frogs and insects—and the different kinds of habitats Life Sciences in which they produce their songs and calls. FRSEMR 41U: Museums LIFESCI 1B: An Integrated Introduction to the Life Sciences: Genetics, Genomics and Evolution Hopi E. Hoekstra (and Andrew Berry, Pardis Sabeti) An integrated approach showing how genetics and evolution are intimately related, together explaining the patterns of genetic variation we see in nature and how genomics can be used to analyze variation.

LIFESCI 2: Evolutionary Human Physiology and Anatomy George V. Lauder (and Daniel E. Lieberman, Ian Wallace) Explores human anatomy and physiology from an integrated framework, combining functional, comparative and evolutionary

Melissa Aja Melissa perspectives on how organisms work.

10 Museum of Comparative Zoology COURSES Embracing Remote Learning

When it became clear that courses would be moving online for the remainder of Spring 2020, MCZ faculty-curators and their teaching fellows quickly transitioned their course content and the entire learning experience to the virtual space. In-person lab instruction, which relies heavily on fieldwork, scientific methods and interacting with physical specimens, required particularly creative solutions. Pre-pandemic, OEB 126: Vertebrate Evolution, taught by Stephanie E. Pierce, included lab visits to the MCZ for students to view and interact with fossil specimens from the Vertebrate Paleontology collection, reinforcing information learned in lectures Lab 3D, a website created by through hands-on observation of specimens Phil Fahn-Lai for viewing 3D lecture for students scattered across the and student presentations. specimens and related materials globe. They tried to keep the flow of in the lab of OEB 126: Vertebrate Evolution In an online environment, Prof. Pierce and the course similar to how it was before her team had to ensure that students could spring break, but without student access observe hundreds of specimens remotely. Phil to labs, lab modules had to be modified. Fahn-Lai, the teaching fellow for the course, is One assignment required students to an OEB graduate student and graduate fellow collect biological samples from their at Harvard’s Derek Bok Center for Teaching & environments, extract the DNA and Learning. In collaboration with fellow Pierce analyze the resulting data. For the lab members, Fahn-Lai spent his last days on online space, Prof. Hoekstra and her campus using photogrammetry to scan and team collected samples from a variety create 3D models of as many of the course’s of familiar objects, including a cell remaining fossil specimens as possible, and phone, the sole of a boot and a dog toy, then used his web and graphic design skills to and they recorded a teaching fellow develop a tool to make them viewable. modeling the DNA extraction process. The result is Lab 3D, OEB 126’s online These samples were sent to an external platform for conducting labs. The website lab for sequencing and the resulting data consists of three major components: a were provided to students for analysis. specimen viewer, where students interact Students then matched the DNA profiles Hopi Hoekstra of the samples to those in a database. Sample from Hopi Hoekstra's with high-quality 3D models hosted on the cell phone for remote labwork Sketchfab website; a text pane that contains Even though the students weren’t able in LIFESCI 1B: An Integrated streamlined versions of lab handouts curated to do the sampling and extraction, they Introduction to the Life Sciences: Genetics, Genomics and for the online environment and with links to still participated in the scientific process, Evolution relevant 3D models; and a phylogenetic tree providing foundational knowledge and pop-up that allows students to locate their experience for future courses. specimens in an evolutionary context. Creative solutions like these demonstrate In the course taught by Hopi E. Hoekstra, instructional innovation and a LIFESCI 1B: An Integrated Introduction to dedication to educational excellence the Life Sciences: Genetics, Genomics and that will support high-quality learning Evolution, Prof. Hoekstra and colleagues experiences for students during recorded some elements of discussion and exceptional times.

Annual Report 2019 . 2020 11 Research Making Headlines

Becoming a Snake The elongate and limbless micro-computed tomography (microCT) body plan of snakes is one of scan of the skull revealed that it still had the most extreme examples of some primitive lizard-like features, like the the evolutionary versatility of jugal bone—which is analogous to the cheek vertebrates. It is also very old, bone—but that it also had characteristics beginning in the upper Middle of a modern snake skull. The lack of a jugal Jurassic almost 170 million years bone had been considered a key defining ago, but few early fossils have been feature of all snakes, both ancient and living, found that can shed light on the an assumption disproved by the findings. timing and physical changes that Evolutionary analyses including Najash marked their divergence from and other snake fossils demonstrated that ancient lizards. early snakes retained robust rear limbs for at least the first 100 million years of their Based on new fossils of the extinct evolutionary history, signifying that this legged snake Najash found in was a successful and stable, rather than a Patagonia, Argentina, including transitional, body plan before absence of an exceptionally preserved 3D limbs became predominant in snakes. skull—the oldest articulated fossil snake skull in the world—Tiago R. Simões Garberoglio FF, Apesteguía S, Simões TR, Palci A, Gómez and colleagues clarify several long-standing RO, Nydam RL, Larsson HCE, Lee MSY, Caldwell MW (2019) New skulls and skeletons of the Cretaceous Ú l G ó mez problems on the origins of key features of

Ra legged snake Najash, and the evolution of the modern the modern snake skull. Their analysis of a snake body plan. Sci Adv 5:eaax5833

Developing Flexibility Mammals have specialized different regions occurred in the mammal spine. They have of the spine such that each section takes on a developed a tool to measure the relationship variety of shapes and functions independent between form and function by evaluating the of the other spinal biomechanics of two modern species with regions. This results very different spines—the tegu lizard and in amazing functional the domestic cat—that bracket the ancient diversity, such as the transition to mammals. giraffe’s elongated They then applied this method to CT scans of neck that allows it to synapsid fossils, the forerunners of mammals, nibble on treetops finding that even though spinal regions or the flexible back evolved earlier, an evolutionary trigger was of the cheetah that required in order to exploit this adaptation. allows it to run like For example, a burst in functional diversity the wind. But what was observed in advanced cynodonts, the prompted the early closest synapsids to mammals, and was linked evolution of this with the evolution of extreme spinal twisting. complexity—the This twisting is used in living mammals while physical development self-grooming fur, potentially indicating the of the regions or the evolution of an insulating pelage before the need to adapt to new Rose Lincoln/HarvardRose University origin of mammals. behaviors? Katrina Jones & Stephanie Pierce Jones KE, Gonzalez S, Angielczyk KD, Pierce SE (2020) Stephanie E. Pierce and Katrina Jones are Regionalization of the axial skeleton predates functional investigating these questions to determine adaptation in the forerunners of mammals. Nat Ecol Evol how and when—and why—changes 4:470-478

12 Museum of Comparative Zoology RESEARCH

Wondrous Wings Wings are useful for far more than flight. are especially susceptible to overheating. Wing patterns and colors help butterflies Specialized scales on the surface of the wing attract mates, warn off or hide from have intricate nanostructures that function predators, or mimic other animals. In as black body absorbers to modulate addition, recent research has discovered that temperatures of the tissues beneath them. butterfly wings have sophisticated physical Thus the wings as a whole act as sensitive adaptations to sense and control their temperature sensors, signaling to the temperature, triggering behaviors designed to butterfly to bask in the sun if the wings are cool or warm the wings as needed. too cool, orient as needed to catch the rays, and avoid overheating by flapping or closing Butterfly wings contain a matrix of living their wings, turning around or moving tissue that is supplied by circulatory, neural away. They also discovered a novel “wing and tracheal systems—indicating that they heart” that beats rhythmically to pump are dynamic, living structures—and these hemolymph (insect blood) directionally tissues require appropriate temperatures to through the scent pad organs of male function. To study the thermodynamic and hairstreak butterflies. Together these results thermoregulatory properties of the wings, show that to fully appreciate the patterns Naomi E. Pierce and colleagues developed produced on the wings of butterflies, it's a novel noninvasive infrared hyperspectral important not only to focus on colors that are imaging technique to map the temperature of visible to us, but also to analyze wavelengths in the thin and delicate wing structures in various Nanfang Yu the non-visible part of the spectrum, including natural environmental conditions, finding the UV and near infrared. that the living areas of the wing have a higher thermal emissivity, passively releasing heat to Tsai C-C, Childers RA, Nan Shi N, Ren C, Pelaez JN, Bernard GD, Pierce NE, Yu N (2020) Physical and keep them much cooler than the non-living behavioral adaptations to prevent overheating of the membranes in the rest of the wing, which living wings of butterflies.Nat Commun 11:551

The Tortoise and the Hare Bio-inspired robots use naturally occurring Then, Prof. Lauder,Dylan Wainright and properties of organisms to improve Valentina Di Santo, working with collaborators performance, and in turn, can help researchers at the University of Virginia, evaluated the better understand those same organisms. This tunabot’s performance in a custom-built flow fortuitous feedback occurs in the experimental tank in the lab. Using new study of an underwater robot design based data from their research on on the physical properties of tuna, remarkable live, free-swimming yellowfin endurance swimmers that are also capable tuna for comparison, they of bursts of high speed when escaping from found that the tunabot has predators or feeding. similar performance and power consumption, flapping The majority of fish-like robots have operated its tail up to 15 times per at lower speeds, so the challenge was to create second to reach higher speeds a robot with the capability of effectively and, if fitted with a 10-Wh propelling itself through the water at both battery pack, could swim as lower and higher speeds. To develop the far as 5.5 miles. Ultimately, LauderGeorge tunabot, a 10-inch-long robot based on a the tunabot could help shed simplified body plan of the yellowfin tuna, light on how open-ocean fish swim and inform Thunnus albacares, George V. Lauder and the design of high-performance bio-inspired colleagues first identified major anatomical underwater vehicles. structures that affect propulsion and made a 3D model of the body. A motor in the robot’s Zhu J, White C, Wainwright DK, Di Santo V, Lauder GV, Bart-Smith H (2019) Tuna robotics: A high-frequency head connects to a drive shaft to power the tail, experimental platform exploring the performance which is stabilized for fish-like lateral bending. space of swimming fishes.Sci Robot 4:eaax4615

Annual Report 2019 . 2020 13 RESEARCH

Head Games The Chengjiang site in China’s Yunnan of Chengjiang arthropods using microCT to Province is a richly fossiliferous create virtual 3D models of the iron-rich fossils. deposit from the early Cambrian, This imaging method reveals features that are dating to around 518 million years concealed within the rock, yielding high-quality ago. The animals preserved in morphological data along with information on Chengjiang lived during the so-called the subject’s affinities, ecology and evolutionary Cambrian Explosion, and include significance. Their recent paper inCurrent several extinct arthropod groups that Biology focuses on the head of the species have an unfamiliar morphology, are Leanchoilia illecebrosa, a member of Megacheira, extremely rare, or are incompletely an extinct group characterized by a pair of large preserved. These combined biases raptorial limbs known as “great appendages.” By hinder our understanding of the early studying small Leanchoilia juveniles, they found evolutionary history of this major the presence of a reduced labrum—a flap-like phylum of invertebrates, whose modern structure overlaying the mouth in most living members include insects, arachnids arthropods—a valuable anatomical feature and crustaceans. Cambrian arthropods suggesting that megacheirians are related to are particularly important as they modern chelicerates like horseshoe crabs, play a key role in understanding the scorpions and spiders. The use of microCT to origin of the arthropod head, whose study Chengjiang fossils holds great potential complex evolution has been scrutinized to revolutionize our understanding of these

Yu Liu through anatomical, developmental and important organisms during a critical time in paleontological means. the history of life. Javier Ortega-Hernández and Yu Liu of Yunnan Liu Y, Ortega-Hernández J, Zhai D, Hou X (2020) A University have been collaborating on the study reduced labrum in a Cambrian great-appendage euarthropod. Curr Biol 30:3057-3061.e2 Taking the Plunge For decades, from the African continent. The fossils—found paleontologists have alongside evidence of a freshwater ecosystem thought that dinosaurs dominated by ancient fish—reveal a flexible lived on land or flew tail, more than 16 feet long with 3-foot vertical in the air but did not spines that form a vertically extended, fin-like live in the water like structure unlike that of any other dinosaur. To some ancient reptiles. evaluate if the tail could be used for propulsion Spinosaurus, a large- in the water, Stephanie E. Pierce collaborated bodied theropod like with George V. Lauder to model the tail— tyrannosaurs, was along with tails of two land-based dinosaurs, thought to have fed a modern crocodile, and a newt—to compare on fish by wading swimming efficiency. They created small two-

Stephanie Pierce (2) Pierce Stephanie along the water’s dimensional plastic tails, attached them to a George Lauder edge. Intriguingly, robot arm and measured the amount of thrust Spinosaurus had other adaptations like reduced produced in water. They found theSpinosaurus hindlimbs, wide feet and a center of gravity model produced eight times the thrust of the more suited to water than land—but without land-based dinosaur tails, comparable to that a known method of propulsion, it was unclear of today’s crocodiles, pointing to Spinosaurus whether it was fully aquatic or not. having been an active and highly specialized large aquatic predator that hunted its prey in In 2018 extensive remains of Spinosaurus the water column. aegyptiacus, the largest known spinosaurid, were excavated from Morocco’s Cretaceous Ibrahim N, Maganuco S, Dal Sasso C, Fabbri M, 13 more including Lauder GV, Pierce SE (2020) Tail-propelled KemKem beds, including a young Spinosaurus aquatic locomotion in a therapod dinosaur. Nature thought to be the most complete dinosaur 581:67-70

14 Museum of Comparative Zoology Highlights from the Collections

Documenting Delicate Sea Creatures The MCZ’s spectacular 19th-century collection of glass teaching models by Leopold and Ralph Blaschka, the second largest in the world, has undergone significant transformations in the last two decades, including a major, multiyear restoration project. More recently, a similarly substantial effort has been undertaken to render the delicate glass sculptures of sea creatures in 3D to make them available to researchers and the public around the world. Traditional 3D photography, called photogrammetry, involves taking multiple images of the subject from various angles and stitching them together. While this method is effective for some models, the reflective and transparent surfaces of other models make them less suitable, so new techniques were needed. A paper by James Hanken, lead author Peter Fried and 3D rendering of the Blaschka model of a greater argonaut, colleagues details how traditional photogrammetry, Argonauta argo. View on combined with a mesh generated through X-ray Sketchfab at skfb.ly/6G7M9. computed tomography (CT) scanning, can render glossy, At left: 3D rendering of the Blaschka model of the blue translucent and intricate objects in 3D while preserving button jellyfish,Porpita porpita. the delicate colors of the models. Fried has utilized this View on Sketchfab at method to render more than a dozen of the Blaschka skfb.ly/6Nrx7. models of marine invertebrates in 3D and make them viewable on the website Sketchfab. "This procedure has the potential to render other reflective and translucent— and delicate—museum objects for archival purposes and make them widely available," says Prof. Hanken.

Exceptional Eggs & Notable Nests The Ornithology department has undertaken the task of imaging and transcribing data from specimen cards (egg cards) associated with the MCZ's collection of around 40,000 bird eggs and nests. In their final days on campus last spring, curatorial associate Jeremiah Trimble and curatorial assistants Katherine Eldridge and John Mewherter diligently scanned about 800 egg cards associated with 500 specimens. Working remotely, they then were able to translate, transcribe, vet and digitize the data contained on the cards. “Traditionally, egg cards contain a vast wealth of information related to the specimens, including habitat, incubation time, nest location, behavior, plant species associated with the nest, and other important data,” says Trimble. Egg cards in the Ornithology collection date from the 1850s, and they provide a snapshot of the birds’ environments as well as insight into their breeding seasons and nesting ranges. The new information will provide significant supplemental data

to enhance existing records for these specimens in MCZbase. Jeremiah Trimble

Annual Report 2019 . 2020 15 COLLECTIONS Digging Into the Data MCZ has been awarded three National Science Foundation grants to enhance and mobilize specimen records, which will benefit both science and society.

Cryo-preservation of Type Specimens The three-year grantCollections in Support of Biological Research: Preserving the Genomes of the Type Specimens in the Museum of Comparative Zoology will preserve the genetic identity of a large number of animal species represented in the MCZ collections, while also serving as a backup for the tissue of these specimens. The PI for the grant is Gonzalo Giribet, with co-PIs Scott Edwards and Breda Zimkus, assistant director of collections operations. “The main objective is to subsample and cryo-preserve tissues of nearly 17,000 primary types hosted at the MCZ, excluding insects,” says Prof. Giribet. “It will benefit from the recently created Cryogenic collection at the MCZ, where the subsampled tissues will be housed and made available to other researchers.” This initiative could then act as a trigger for similar initiatives at other museums and further global goals across disciplines that rely on

Jennifer Berglund genetic resources, including zoology, genomics and conservation biology. Breda Zimkus Documenting Marine Biodiversity About 75% of the 250,000 described species of marine organisms are invertebrates. The four-year grant Documenting Marine Biodiversity through Digitization of Invertebrate Collections will facilitate digitization of approximately 31,564 lots and 4,631 primary type specimens of MCZ's recent marine invertebrates, including cnidarians, echinoderms, crustaceans, brachiopods, polychaete worms and bryozoans. James Hanken is PI for the MCZ component. Adam J. Baldinger, curatorial associate, Invertebrate Zoology and Malacology, is co-PI. Aja Melissa “The MCZ collections of marine invertebrates are worldwide in scope, many resulting from expeditions in the 19th and early 20th centuries,” says Baldinger. Digitization efforts will consist of entering specimen metadata into the database, georeferencing localities, imaging specimens, and capturing data for specimens that are attached to or growing on each other, like a barnacle on a mussel.

Mobilizing Mollusks The MCZ Malacology collection includes 82,775 lots from the Eastern Seaboard that equal 710,600 non-fossil specimens of biologically and economically important species, including oysters, clams, mussels,

scallops, whelks and squids. The four-year grantMobilizing Millions of Trimble Jennifer W. Marine Mollusks of the Eastern Seaboard will database 7,500 uncataloged lots from the Eastern Seaboard in the MCZ collection, clean the georeferences of 16,000 unique localities in MCZbase for specimens from this area, and prepare high-quality digital images of nearly 14,000 primary types for MCZbase. PI Gonzalo Giribet is joined by Adam J. Baldinger and Vanessa Knutson Vanessa curatorial assistants Alana Rivera and Murat Recevik. “MCZ is part of a collaborative consortium that will collectively mobilize 53,000 fully georeferenced records representing 4.2 million specimens from the shore to the edge of the Exclusive Economic Zone, from the Canadian border to the Texas coast of the Gulf of Mexico, around 3,800 miles of coastline,” says Baldinger. Specimen data will eventually be accessible through portals like InvertEbase and MolluscaBase and various aggregators such as WoRMS, iDigBio and GBIF.

16 Museum of Comparative Zoology COLLECTIONS

Staff Highlight Adam J. Baldinger In 1997, Adam Baldinger became the curatorial associate in Malacology and, in 2008, he began to manage the Invertebrate Zoology collection as well. Both collections are extensive in number and diverse in scope—the Malacology collection has an estimated 10 million shells—with some groups within Invertebrate Zoology, such as spiders, being among the most important in the world. Many specimens date to MCZ expeditions and other collections activities in the late 19th century. “It’s intriguing to see a specimen collected in 1849 and reflect on the people that have touched it before me,” says Baldinger. “Hopefully, 150 years from now, my efforts will mean that others can continue to work with

Catherine Weisel these specimens and collections.” Adam works closely with Gonzalo Giribet and Mansi Srivastava, curators of Invertebrate Zoology, and James Hanken, acting curator of Malacology. Along with five staff members, they are managing four significant grants to further digitize the collections and cryo-preserve tissue samples. “After working with Adam for two decades, I could not have wished for a more thoughtful, reliable and efficient colleague,” says Prof. Giribet. In his spare time, Baldinger’s research interests are on amphipods, a group of ecologically important shrimp-like crustaceans on which he published his first paper as an undergraduate and which he continues to study. Several years ago, a park ranger in Death Valley National Park contacted him about a subterranean freshwater amphipod and sent samples. It turned out to be a new species, Hyalella sandra, which he described and named after his wife. A group of researchers also named a species of a small marine mollusk from Martinique Teinostoma baldingeri in recognition of his vital contribution to their work.

MCZ History Many people associated with the Museum of Comparative Zoology have achieved great fame and notoriety since its founding in 1859. Yet, other individuals who made highly significant contributions to the Museum, and comparative zoology more generally, have for a variety of reasons been denied due recognition of their talents and accomplishments. Perhaps foremost among the latter is Robert A. Gilbert, who as an African American was denied opportunities for education, training and professional advancement that many of his contemporaries took for granted. Mass Audubon Collection, Museum of American Bird Art Bird of American Museum Collection, Audubon Mass Born in South Carolina in 1869, Gilbert didn’t begin working Robert Gilbert (far right) during a field excursion for the MCZ until 1919. His association with zoology, however, near Bethel, Maine had begun in the mid-1890s when, at the age of 27, Gilbert was hired by Cambridge-based ornithologist William Brewster to assist with Brewster's studies of the birds of New England, and especially Massachusetts. This included the then-novel idea of bringing along a camera and glass- plate negatives in order to photograph birds and other animals and plants in their natural habitats. Brewster was long credited as principal photographer, with little more than passing mention of Gilbert’s contribution. That all changed in 2005, when biographer John Hanson Mitchell provided compelling evidence that most if not all of the remarkable images were actually created by Gilbert. Today, Gilbert is recognized as one of the earliest natural history photographers in North America, if not the world.

Annual Report 2019 . 2020 17 Projects & Initiatives The Importance of Primary Sources TheErnst Mayr Library holds a wealth of to the project, published by museums, biological resources and makes them available universities and scholarly societies from to researchers at Harvard and around the around the world.” world. Much like the MCZ’s specimens, the Creating custom software to search the EML’s diverse primary literature sources can be digital texts for egg morphological data, the used to disprove old hypotheses or develop researchers narrowed the results to 1,756 new ones. When the team of Samuel H. publications. The software analyzed images Church, Seth Donoughe and Bruno A.S. de and extracted text, resulting in about 10,000 Medeiros—led by OEB professor Cassandra egg measurements for more than 6,700 G. Extavour—needed help with references to insect species. Using the completed and create a historical dataset on insect eggs, they verified dataset, the researchers disproved turned to the EML. some popular hypotheses about egg size, To build this massive data set, the team finding instead that ecological considerations compiled more than 3,000 references from seem to be more important in determining the past 250 years, finding many online or on egg morphology, pointing the way for future the shelves of the EML. However, still missing research and highlighting the importance of 350 tough-to-find sources, they approached primary historical sources. Mary Sears, EML’s head of public services, Church SH, Donoughe S, de Medeiros BAS, Extavour CG for support. “It was like detective work,” (2019) A dataset of egg size and shape from more than says Sears, “because the citations were in 6,700 insect species. Sci Data 6:104 relatively obscure journals and in several Church SH, Donoughe S, de Medeiros BAS, Extavour CG different languages. Ultimately I was able to (2019) Insect egg size and shape evolve with ecology but

Samuel Church and Seth Donoughe (2) Donoughe Seth and Church Samuel contribute 265 books and journal articles not developmental rate. Nature 571:58-62

Ernst Mayr Library After campus closed in March, staff worked remotely to support MCZ research and teaching, making extensive use of digital repositories like the Biodiversity Heritage Library to provide materials to faculty, graduate students and curatorial staff. The EML website, library guides, blogs and other social media connected the MCZ and larger natural history community with library pages of William Brewster’s journals and diaries resources. this year. Only 6% of the Brewster material Biodiversity Heritage Library housed in the archives remains to be digitized, EML staff, working with partner institutions, and 20% of it has been transcribed and have enriched BHL content while focusing on validated, including 15 journals (2,416 pages). unique MCZ collections, adding more than

200 books (34,955 pages) in the last year. Digitizing MCZ Collection Documents EML has digitized the Paul Morris field notes, On average, EML content in BHL is viewed 470 cards for the Northcutt collection of by 9,513 users each month, a total of 29,108 microscope slides, two accession ledgers views per month. and 104 pages of the Catalogue of Fishes for William Brewster Collections Digitized the Boston Society of Natural History, and Using the Atlas of Living Australia’s DigiVol correspondence from the Herpetology and platform, volunteers and staff transcribed 2,647 Malacology collections.

18 Museum of Comparative Zoology PROJECTS & INITIATIVES Harvard Museums of Science & Culture While Harvard’s public-facing museums have been active in the online space for years, the COVID-19 pandemic places new emphasis on virtual tours, talks and gallery displays. MCZ personnel regularly contribute to such outreach efforts, and they continue to do so via Harvard’s web presence while campus is closed to visitors.

HMSC Connects!, a web initiative to expand online educational activities, launched in April. HMSC Connects! Exhibit Spotlight features Sharks: Streamlined Swimmers, which provides an orientation to shark physiology and highlights the Lauder lab’s research into the effectiveness of shark locomotion. In a video, George V. Lauder uses a blue shark from the MCZ collection to explain how the external aspects of shark anatomy—the fins, tail and scales—help sharks move efficiently through the water. In a second video, lab member Molly Gabler-Smith explains how these scales, or denticles, improve shark hydrodynamics. In HMSC Connects! Extraordinary Things, the website shares the fossilized skeleton of Dimetrodon milleri, a George Lauder mammal-like carnivore that lived before the emergence of dinosaurs almost 300 million years ago. It features Stephanie E. Pierce and explains how she uses digital scans of this fossil to create 3D models to understand how Dimetrodon may have moved. TheHMSC Connects! Podcasts take listeners behind the scenes of the museums, and in one episode, curatorial associate Andrew Williston talks about the MCZ Ichthyology collection and why collections are important for science and education as well as public viewing. Another podcast for Women’s Suffrage Month goes into the contributions of women working in MCZ collections in the late 19th and early 20th centuries and Reed Gochberg’s

Melissa Aja Melissa efforts to tell their stories in an exhibit at HMNH. Andrew Williston

A Personal Quest with a Larger Purpose This summer,Scott V. Edwards fulfilled his lifelong dream of cycling across the US, and, along the way, contributed to the quest for social justice. In early June he departed Massachusetts after dipping his wheels in the Atlantic Ocean. He reached Oregon and the Pacific Ocean in August, many days and adventures later. He traveled unassisted, with 60 pounds of gear on a bike festooned with printed signs: #BlackLivesMatter, #BlackBirdersWeek and #ShutDownSTEM, a one-day work stoppage to highlight systemic racism in the scientific and academic communities. Prof. Edwards, MCZ faculty member and curator of Ornithology, experienced natural beauty, physical and logistical challenges, random acts of kindness and occasional racist comments, and copious birdsong. “It’s important for folks to see that African Americans do enjoy nature,”

he says. “It’s important to showcase that we like camping and show it’s Ellie Steinberg not just the domain of others.” Scott Edwards

Annual Report 2019 . 2020 19 Awards & Recognition Javier Ortega-Hernández and Research Associate Rudy Lerosey-Aubril were jointly awarded a grant from Harvard’s William F. Milton Fund. Ortega-Hernández received an award from the Harvard China Fund to support his work on South Chinese fossils. Naomi E. Pierce was awarded the 2019 International Prize for Biology from the Japan Society for the Promotion of Science for her outstanding contribution to the advancement of research in fundamental biology. Stephanie E. Pierce received the 2019 Fannie Cox Prize for Excellence in Science Teaching from Harvard's Faculty of Arts and Sciences. A new genus of terrestrial lizard, Wilsonosaura, and a new bat species, Miniopterus wilsoni, were named in honor of Edward O. Wilson. Jennifer Berglund Edward O. Wilson and James Postdoctoral Researchers Hanken Faculty-Curators Andrew Gehrke received a Charles King Andrew Biewener was elected a fellow of the Trust Postdoctoral Research Fellowship. American Association for the Advancement Aaron Hartmann received a Certificate of of Science. Excellence in Teaching from the Derek Bok Center. Katrina Jones was awarded the Royal Society of London University Research Fellowship. Nicholas Jourjine received a Ruth L. Kirschstein Postdoctoral Individual National Research Service Award and a Harvard Brain Initiative Young Scientist Travel Award. Andreas Kautt received the Cameron Award Scott Edwards was elected to the from the American Society of Mammalogists American Philosophical Society and, along and a postdoctoral fellowship from the with a colleague, received the inaugural German Research Foundation. Inclusiveness, Diversity, Equity and Access (IDEA) Award from the American Society of Naturalists, the Society for the Study of Evolution and the Society of Systematic Biologists. Two new species were named in honor of Gonzalo Giribet, Panzosus giribeti, an arachnid, and Galathenemertes giribeti, a ribbon worm. Hopi E. Hoekstra received the 2019 C. Hart Merriam Award from the American Society Snibbe/HarvardKris University Sarah Losso of Mammalogists. Andrew Gehrke and Mansi Srivastava

20 Museum of Comparative Zoology AWARDS & RECOGNITION Awards & Recognition Elizabeth Sibert was an International Ocean Pavitra Muralidhar received a Discovery Program participant. National Science Foundation Postdoctoral Research Fellowship Tiago Simões received a postdoctoral in Biology and was a finalist for fellowship from the Natural Sciences and the James F. Crow Early Career Engineering Research Council of Canada. Researcher Award from the Genetics Kelsey Tyssowski received a Life Sciences Society of America. Research Foundation postdoctoral fellowship. Kari Taylor-Burt received a Dean Graduate Students Shinagel Exceptional Teaching Assistant Award from Harvard Jennifer Austiff, Nicole Bedford, Richard Extension School. Childers, Blake Dickson, Nathaniel Edelman, Kadeem Gilbert, Alyssa Mark Wright was awarded a Robert Hernandez, Sang Il Kim, Vanessa Knutson, A. Chapman Memorial Scholarship Sarah Losso, Olivia Meyerson, Zachary for Vertebrate Locomotion from the Morris, Shoyo Sato, Carl Schmitt, Harvard Faculty of Arts and Sciences. Zhengyang Wang and Brock Wooldridge each received a Certificate of Distinction in Undergraduate Students Teaching from the Derek Bok Center. Dylan Ryals and Allison Shu Ting Law each received a Hoopes Prize for their thesis Andrew Berry and Naomi Pierce projects. Adele Woodmansee received the Taliesin Prize for Distinction in the Art of Learning. Staff Christian Flynn, OEB administrative coordinator, and Breda Zimkus, assistant director, collections operations, each

Sloane Jackson Sloane received a Dean’s Distinction Award from the Faculty of Arts and Sciences. Inbar Maayan Ryan Hulett received a Ruth L. Kirschstein Breda Zimkus received a Society for National Research Service Award fellowship the Preservation of Natural History

from the National Institutes of Health. Collections Special Service Award, Pierce Naomi presented to the organizing committee Richard Childers Sang Il Kim received a Graduate Student for the SPNHC 2020 meeting. Research Enhancement Award from The Coleopterists Society. Special Commendation Julian Kimura received a Qbio Student Extraordinary Teaching in Extraordinary Times Fellowship supported by the National This spring, undergraduates named instructors for special Science Foundation–Simons Center for recognition, praising their flexibility and creativity in adapting the Mathematical & Statistical Analysis of to online teaching and their compassionate responses to the Biology at Harvard University. challenges faced by students during the pandemic. Inbar Maayan received a Society of The Harvard College special commendation was awarded to Scott Systematic Biologists Graduate Student Edwards and Teaching Fellow Gustavo Bravo (OEB 190); Teaching Research Award. Fellow Philip Fahn-Lai (OEB 126); Teaching Fellow Dave Matthews Zachary Morris received a National (OEB 53); and James Hanken and Teaching Fellows Inbar Maayan Science Foundation Earth Sciences and Jennifer Austiff (OEB 167). Postdoctoral Fellowship. Annual Report 2019 . 2020 21 Grant Recipients

Grants-in-Aid of Undergraduate Research L.André Martins These grants support research by Harvard College undergraduates under faculty supervision. Priority is given to projects that utilize MCZ research collections, laboratories and facilities. Support for these grants comes from the MCZ’s Myvanwy M. and George M. Dick Scholarship for Students.

Recipient Academic Dept./ Project Title Amount Faculty Sponsor Kim Anyeji Boerrigter OEB/Stephanie Pierce These fins are made for walkin’: A $4,000 Amy Li comparative musculoskeletal analysis of anglerfish pectoral fins Justin Duffy OEB/Mansi Srivastava Whole-body regeneration of Nematostella $4,000 vectensis Kate Gonzalez OEB/George Lauder The effects of fish abundance on coral disease in $4,000 Tela Bay, Honduras Jackson Kehoe OEB/Javier Ortega- Testing the evolution of body segmentation in $4,000 Hernández trilobites Anne Kennedy-Yoon OEB/Naomi Pierce The effect of ant species on the morphology $3,809 and fungal communities of their host, V. drepanolobium Keza Levine OEB/Hopi Hoekstra Deer mouse vocalization analysis $4,000

Inbar Maayan Amy Li OEB/Peter Girguis Classifying marine benthos via hyperspectral $4,000 imaging Chamberlain Mathis OEB/James Hanken Documentation and analysis of $1,000 Lepidobatrachus laevis stomach development Logan Qualls OEB/Javier Ortega- Immediate family matters: Understanding $4,000 Hernández interspecific differences in the genusIsotelus , and how members are related Maya Rayle Harvard Medical The eco-evolutionary dynamics of microbes $4,000 School/Ali R. Zomorrodi Oliver Riskin-Kutz OEB/Scott Edwards Determining ecological influences on sea $1,000 urchin grazing behavior in Northern California kelp forests Arianna Romero OEB/Stephanie Pierce Investigating locomotor adaptations in $4,000 Procynosuchus delaharpeae (Synapsida,

Naomi Pierce Naomi Cynodontia) with implications for ecological diversity in the mammalian stem Anne Kennedy-Yoon Dylan Ryals OEB/Brian Farrell Transmission of honeybee parasites due to $2,677 migratory beekeeping practice John Emory Sabatini OEB/Hopi Hoekstra Computational analysis of pup vocal recordings $4,000 from four Peromyscus species John Schaefer History of Science/ Carnivorous plants: History, conservation and $4,000 Janet Brown education Chidambaram OEB/Scott Edwards Convergence and coevolution between $4,000 Thillairajah plumage coloration and light-sensitive visual pigments in antbirds (Aves, Thamnophilidae) Madeleine Waskom OEB/Javier Ortega- Modeling of Walcott-Rust thin sections $4,000 Hernández

Total Awards $60,486 John Schaefer John

22 Museum of Comparative Zoology GRANTS

Putnam Expedition Grants Putnam Expedition Grants are intended to support MCZ faculty-curators, postdoctoral fellows and graduate students in collecting specimens and data relating to the study of comparative zoology. Priority is given to projects that collect living specimens in regions where habitats are threatened or fossil specimens in regions most likely to hold important clues for unraveling evolutionary strategies. These grants are made possible by a gift from Mr. George Putnam Jr., AB 1949 and MBA 1951, and Mrs. Nancy Putnam.

Recipient MCZ Department/ Project Title Amount Faculty Sponsor

Shahan Derkarabetian Invertebrate Zoology/ The unknown triaenonychid harvestmen of $7,785 Gonzalo Giribet Gonzalo Giribet Australia. Expedition II: Queensland Reinhardt Kristensen and Katrine Worsaae Gonzalo Giribet Invertebrate Zoology Following the leads of eDNA: In search of the $4,700 Pyrenees' new species of Micrognathozoa Nicholas Herrmann Herpetology/David Determinants of success in a novel $11,200 Haig environment: Testing how morphology and habitat use affect individual fitness in Anolis lizards Andreas F. Kautt Mammalogy/Hopi The genetic basis of predator aversion $6,997 Hoekstra behavior in deer mice in the Channel Islands Archipelago, California Kelsey M. Tyssowski Mammalogy/Hopi Natural variation in morphology and behavior $10,271 Hoekstra in arboreal and non-arboreal deer mice

Total Awards $40,953 Jennifer Austiff

Ken Miyata Grants The Ken Miyata Fund in Herpetology and Ken Miyata Fund for Field Research support students who share Ken’s interests as a naturalist, biogeographer and writer/photographer by defraying the costs of research by graduate students in herpetology. The funds were established by generous gifts from Barbara Wu, PhD 1981, and Eric Larson, AB 1977, and other close friends of Ken.

Recipient MCZ Department/ Project Title Amount Faculty Sponsor

Inbar Maayan Herpetology/David Island-wide variation and lineage $12,110 Javawney Crooks Haig diversification in Jamaican Anolis lizards Inbar Maayan Total Awards $12,110

Robert G. Goelet Research Awards Goelet Awards support MCZ graduate student research projects. These grants are made possible through a gift from Mr. Robert G. Goelet.

Recipient MCZ Department/ Project Title Amount Faculty Sponsor Jennifer Austiff Herpetology/James Developmental mechanisms that underlie $6,797 Hanken the evolution of the stomach of the frog, Lepidobatrachus laevis

Total Awards $6,797 Yale Peabody Museum specimen by Jackson Kehoe by Jackson specimen Museum Peabody Yale

Annual Report 2019 . 2020 23 GRANTS

Ernst Mayr Travel Grants in Animal Systematics Ernst Mayr Grants support travel for research in animal systematics and are open to the scientific community worldwide. The principal objective of these grants is to stimulate taxonomic work on neglected taxa and/or poorly described species. Ernst Mayr Grants typically facilitate visits to institutional collections, with preference given to research that uses MCZ’s collections. These grants are made possible by a gift from professor and former MCZ Director Ernst Mayr.

Shahan Derkarabetian Recipient Institutional Project Title Amount Affiliation Uttaran Bandyopadhyay Zoological Survey of Identifying and resolving higher taxonomy of $1,500 India Indian Himalayan Phlogophorini (Lepidoptera: Noctuidae: Xyleninae) with special reference to the Euplexia genus group Russell D. C. Bicknell University of New Redescription of Mesolimulus based on $1,500 England, Armidale, exceptional fossils with three-dimensional Australia preservation Damian Luis Castellini Instituto de Systematics and taxonomic review of the flatfish $1,500 Investigaciones Paralichthys genus in Southwestern Atlantic Marinas Y Costeras Ocean (Actinopterygii: Pleuronectiformes) CONICET, Argentina Patrick J. Ciccotto Warren Wilson College Resolving the taxonomy of carps of the $650 subfamily Labeoninae (Teleostei: Cyprinidae) of mainland Southeast Asia Brittany N. Damron Universidade de São Taxonomic review and revision of the family $1,500 Paulo, Brazil Cosmetidae (Opiliones, Arachnida) in the MCZ collection Matt Green Matt Vinicius C. Espindola Smithsonian National A taxonomic review of the Western South $1,460 Alexander B. Orfinger Museum of Natural Atlantic Chimaera (Chondrichthyes: History Holocephali), with the description of a new species and three new records Edgar Gamero Mora Universidade de São From species discovery to species description: $1,400 Paulo, Brazil The description of two new species ofCassiopea (Cnidaria: Scyphozoa) Olivia M. Gearner Purdue University Systematics and taxonomic revisions of the $1,500 tok-tokkie beetles of Africa (Coleoptera: Tenebrionidae: Sepidiini) Lucas Roberto Pereira Universidade Federal Revision of Neotropical Anthomyiidae $1,500 Gomes do Paraná, Brazil (Diptera) and phylogenetic analysis of the family Matthew W. Green Clemson University Inferring association and diagnosis of $1,500

Pamela Yesenia Sánchez Vendizú Yesenia Pamela Pycnopsyche (Trichoptera: Limnephilidae) larvae and adults Zachary H. Griebenow University of Resolving parallel taxonomy in the Leptanillinae $1,120 California, Davis (Hymenoptera: Formicidae), an enigmatic group of ants Karla Janet Humara Gil Centro Universitario Revision of Ophioderma teres complex $1,500 de la Costa, (Echinodermata: Ophiuroidea) from the Eastern Universidad de Pacific Guadalajara, Mexico Ilgoo Kang Louisiana State Important newly collected cardiochiline $1,242 University braconids housed in the Naturalis Biodiversity Center in Leiden (Hymenoptera: Braconidae) José Ricardo Assmann Universidade Federal Taxonomic revision and phylogenetic $1,500 Lemes do Paraná, Brazil analysis of Staphyus Godman & Salvin, 1896, Incisus Grishin, 2019 and Perus Grishin, 2019 (Lepidoptera: Hesperiidae) Pamela Yesenia Sánchez Vendizú

24 Museum of Comparative Zoology GRANTS Patrick J. Ciccotto J. Patrick

Recipient Institutional Project Title Amount Affiliation Abigail P. Martens South Dakota State The Petr Starý Collection of Aphidiinae $1,500 University (Hymenoptera: Braconidae) André L. Martins Universidade Federal Taxonomic revision of Dryinus Latreille species $1,500

do Paraná, Brazil (Hymenoptera, Dryinidae, Dryininae) from Emily L. Sandall Neotropical region: With a new proposal of classification for species groups Bruno A. S. de Medeiros Smithsonian Tropical New species of Anchylorhynchus Schoenherr, $1,000 Research Institute, 1836 (Curculionidae: Derelomini) Panama Jairo Andrés Moreno- Universidade de São Revision of the unidentified material of the $1,500 González Paulo, Brazil enigmatic order Schizomida (Arachnida) housed in the MCZ Alexander B. Orfinger University of Florida Taxonomic revision of Polycentropus $1,000 (Trichoptera: Polycentropodidae) from the Eastern Nearctic Lívia Pires do Prado Universidade Federal Taxonomic revision and phylogenetic $1,000 do Pará and Museu relationships of the Megalomyrmex Forel, 1885 Paraense Emílio (Hymenoptera: Formicidae: Solenopsidini) Goeldi, Brazil Willians Lourenço Porto Museo Argentino de Revision of the Andean-Patagonian $1,500 Ciencias Naturales Triaenonychidae (Opiliones, Laniatores, MACN-CONICET, Triaenonychidae) Emily L. Sandall Argentina M. Guadalupe del Río CONICET, División Systematics on the weevil tribe Leptopiini $1,500 Entomología, Museo (Insecta: Coleoptera: Curculionidae) de La Plata, Argentina Sebastian Salata California Academy of Taxonomic revision of Pheidole Westwood, 1839 $1,500 Sciences (Hymenoptera, Formicidae) in Madagascar Pamela Yesenia Sánchez Universidade Federal Systematic revision of the spiny tree rat $1,500 Vendizú do Pará, Brazil Mesomys (Rodentia: Echimyidae)

Emily L. Sandall Pennsylvania State Phylogeny of Ophiogomphus (Odonata: $882 Orfinger B. Alexander University Gomphidae) Aluska Tavares dos Universidade Federal Taxonomic revision of Acanthoscelides, the $1,500 Santos do Paraná, Brazil largest Neotropical genus of seed-beetles (Chrysomelidae: Bruchinae) Josenilson R. Santos Universidade Federal A taxonomic review of the New World $1,500 do Rio de Janeiro, genus Microcerella Macquart, 1851 (Diptera, Brazil Sarcophagidae) Wildio Ikaro da Graça Universidade Federal Taxonomic review of Eurybia [Illiger], 1807 $1,500 Santos do Paraná, Brazil and phylogenetic analysis of the subtribe Eurybiina Reuter, 1896 (Lepidoptera: Riodinidae) José Eduardo Serrano- Universidade de São Taxonomic revision of the genus Pithecia $1,500 Villavicencio Paulo, Brazil Demarest, 1804 in the eastern Peruvian Amazon

Total Awards $39,754 André L. Martins

Annual Report 2019 . 2020 25 Publications in 2019

• Alegre A, Gainett G, López Iborra G, Giribet • Campbell Mekarski M, Pierce SE, Caldwell MW G (2019) Two new species of Manahunca, (2019) Spatiotemporal distributions of non- redescription of its type species, current ophidian ophidiomorphs, with implications for their conservation status of the genus and a survey origin, radiation, and extinction. Front Earth Sci 7:245 of male glands in Stenostygninae (Opiliones: Laniatores: Biantidae). Zootaxa 4686:83-111 • Chen X, Ortega-Hernández J, Wolfe JM, Zhai D, Huijian M, Hou XG, Chen A, Yu L (2019) The • Álvarez-Campos P, Kenny NJ, Verdes A, appendicular morphology of Sinoburius lunaris and Fernández R, Novo M, Giribet G, Riesgo A (2019) the evolution of the artiopodan clade Xandarellida Delegating sex: Differential gene expression in (Euarthropoda, early Cambrian) from South China. stolonizing syllids uncovers the hormonal control BMC Evol Biol 19:1-20 of reproduction. Genome Biol Evol 11:295-318 • Cheng D, Chen S, Huang Y, Pierce NE, Riegler • Arellano C, Konow N, Gidmark N, Roberts TJ M, Yang F, Zeng L, Lu Y, Liang G, Xu Y (2019) (2019) Evidence of a tunable biological spring: Symbiotic microbiota may reflect host adaptation Elastic energy storage in aponeuroses varies with by resident to invasive ant species. PLoS Pathog transverse strain in vivo. Proc R Soc B 286:20182764 15:e1007942 • Baier F, Hoekstra HE (2019) The genetics of • Church SH, Donoughe S, de Medeiros BAS,

Dayou Zhai morphological and behavioral island traits in deer Extavour CG (2019) A dataset of egg size and shape mice. Proc R Soc B 286:20191697 from more than 6,700 insect species. Sci Data 6:104 • Barrett RDH, Laurent S, Mallarino R, Pfeifer SP, Xu • Church SH, Donoughe S, de Medeiros BAS, CCY, Foll M, Wakamatsu K, Duke-Cohan JS, Jensen Extavour CG (2019) Insect egg size and shape JD, Hoekstra HE (2019) Linking a mutation to evolve with ecology but not developmental rate. survival in wild mice. Science 363:499-504 Nature 571:58-62 • Benavides LR, Cosgrove JG, Harvey MS, Giribet • Clites TR, Arnold AS, Singh NM, Kline E, Chen G (2019) Phylogenomic interrogation resolves the H, Tugman C, Billadeau B, Biewener AA, Herr backbone of the Pseudoscorpiones tree of life. HM (2019) Goats decrease hindlimb stiffness Mol Phylogenet Evol 139:106509 when walking over compliant surfaces. J Exp Biol 222:jeb198325 • Benavides LR, Hormiga G, Giribet G (2019) Phylogeny, evolution and systematic revision of the • Cloutier A, Sackton TB, Grayson P, Clamp M, mite harvestman family Neogoveidae (Opiliones, Baker AJ, Edwards SV (2019) Whole-genome Cyphophthalmi). Invertebr Syst 33:101-180 analyses resolve the phylogeny of flightless birds (Palaeognathae) in the presence of an empirical Samuel Church and Seth Donoughe Donoughe Seth and Church Samuel • Bhullar BS, Manafzadeh AR, Miyamae JA, anomaly zone. Syst Biol 68:937-955 Hoffman EA, Brainerd EL,Musinsky C, Crompton AW (2019) Rolling of the jaw is essential for • Cordeiro IR, Kabashima K, Ochi H, Munakata mammalian chewing and tribosphenic molar K, Nishimori C, Laslo M, Hanken J, Tanaka M function. Nature 566:528-532 (2019) Environmental oxygen exposure allows for the evolution of interdigital cell death in limb • Bicknell RDC, Amati L, Ortega-Hernández J patterning. Dev Cell 50:155-166 (2019) New insights into the evolution of lateral compound eyes in Palaeozoic horseshoe crabs. • Cunha TJ, Giribet G (2019) A congruent Zool J Linnean Soc 187:1061-1077 topology for deep gastropod relationships. Proc R Soc B 286:21082776 • Biewener AA (2019) Getting to grips with how birds land stably on complex surfaces. News & • Cunha TJ, Lemer S, Bouchet P, Kano Y, Views. Nature 574:180-181 Giribet G (2019) Putting keyhole limpets on the map: Phylogeny and biogeography of the • Blackburn DC, Giribet G, Soltis DE, Stanley EL (2019) globally distributed marine family Fissurellidae Predicting the impact of describing new species on (Vetigastropoda, Mollusca). Mol Phylogenet Evol phylogenetic patterns. Integr Comp Biol 1:1-12 135:249-269 • Boyle JH, Martens D, Musili PM, Pierce NE (2019) • Delaney EK, Hoekstra HE (2019) Diet-based Population genomics and demographic sampling assortative mating through sexual imprinting. Ecol of the ant-plant Vachellia drepanolobium and its Evol 9:12045-12050 symbiotic ants from sites across its range in East Africa. Front Ecol Evol 7:206 • Derkarabetian S, Benavides LR, Giribet G (2019) Sequence capture phylogenomics of historical • Bravo GA, Antonelli A, Bacon CD, Bartoszek ethanol-preserved museum specimens: Unlocking K, Blom MPK, Huynh S, Jones G, Knowles LL, the rest of the vault. Mol Ecol Resour 19:1531-1544 Lamichhaney S, 11 more including Edwards SV (2019) Embracing heterogeneity: Coalescing the Tree • Dickson BV, Pierce SE (2019) Functional of Life and the future of phylogenomics. PeerJ 7:e6399 performance of turtle humerus shape across an Marjorie Liénard

26 Museum of Comparative Zoology MCZ PUBLICATIONS Publications in 2019

ecological adaptive landscape. Evolution • Giribet G, Baker CM (2019) Further seymouriamorphs (Tetrapoda) uncovered 76:1265-1277 discussion on the Eocene drowning of New by high-resolution x-ray microcomputed Caledonia: Discordances from the point of tomography, and the origin of the amniote • Dickson BV, Pierce SE (2019) How (and view of zoology. J Biogeogr 46:1912-1918 crown group. Palaeontology 63:131-154 why) fins turn into limbs: Insights from anglerfish.Earth Environ Sci Trans R Soc • Giribet G, Edgecombe GD (2019) • Lai A, Biewener AA, Wakeling JM (2019) Edinb 109:87-103 Perspectives in animal phylogeny and Muscle-specific indices to characterise the evolution: A decade later. In Perspectives functional behavior of human lower-limb • Du KS, Ortega-Hernández J, Yang J, Zhang on evolutionary and developmental biology: muscles during locomotion. J Biomech XG (2019) A soft-bodied euarthropod from Essays for Alessandro Minelli (Fusco G ed). 89:134-138 the early Cambrian Xiaoshiba Lagerstätte Padova University Press: Padova of China supports a new clade of basal • Lamichhaney S, Card DC, Grayson artiopodans with dorsal ecdysial sutures. • Giribet G, Edgecombe GD (2019) The P, Tonini JFR, Bravo GA, Näpflin K, Cladistics 35:269-281 phylogeny and evolutionary history of Termignoni-Garcia F, Torres C, Burbrink F, arthropods. Curr Biol 29:R592-R602 Clarke JA, Sackton TB, Edwards SV (2019) • Edgecombe GD, Giribet G (2019) The Integrating natural history collections and molecularization of centipede systematics. In • Hedrick BP, Schachner ER, Rivera G, comparative genomics to study the genetic Perspectives on evolutionary and developmental Dodson P, Pierce SE (2019) The effects of architecture of convergent evolution. Phil biology: Essays for Alessandro Minelli (Fusco G skeletal asymmetry on interpreting biologic Trans R Soc B 374:20180248 ed). Padova University Press: Padova variation and taphonomy in the fossil record. Paleobiology 45:154-166 • Laslo M, Denver RJ, Hanken J (2019) • Eng CM, Konow N, Tijs C, Holt NC, Evolutionary conservation of thyroid Biewener AA (2019) In vivo force–length • Heiss E, Schwartz D, Konow N (2019) hormone receptor and deiodinase and activation dynamics of two distal rat Chewing or not? Intraoral food processing in expression dynamics in ovo in a direct- hindlimb muscles in relation to gait and a salamandrid newt. J Exp Biol 222:jeb189886 developing frog, Eleutherodactylus coqui. grade. J Exp Biol 222:jeb205559 • Hsiou AS, Nydam RL, Simões TR, Pretto FA, Fronts Endocrinol 10:307 • Esteve-Altava B, Pierce SE, Molnar JL, Onary S, Martinelli AG, Liparini A, Martínez • Laumer CE, Fernández R, Lemer S, Johnston P, Diogo R, Hutchinson JR (2019) PRRdV, Soares MB, Schultz CL, Caldwell Combosch DJ, Kocot K, Andrade SCS, Evolutionary parallelisms of pectoral and MW (2019) A new clevosaurid from the Sterrer W, Sørensen MV, Giribet G (2019) pelvic network-anatomy from fins to limbs. Triassic (Carnian) of Brazil and the rise of Revisiting metazoan phylogeny with Sci Adv 5:eaau7459 sphenodontians in Gondwana. Sci Rep 9:11821 genomic sampling of all phyla. Proc R Soc B • Fernández E, Ferreras I, Farrell BD, de • Hu Z, Sackton TB, Edwards SV, Liu JS 286:20190831 Medeiros BAS, Romero-González GA (2019) (2019) Bayesian detection of convergent rate • Leiva C, Taboada S, Kenny NJ, Combosch Studies in Aristolochia (Aristolochiaceae) of changes of conserved noncoding elements on D, Giribet G, Jombart T, Riesgo A (2019) Hispaniola. Phytotaxa 420:1-20 phylogenetic trees. Mol Biol Evol 36:1086-1100 Population substructure and signals of • Friedman M, Pierce SE, Coates M, Giles S • Iwama R, Oceguera-Figueroa A, Giribet G, divergent adaptive selection despite (2019) Feeding structures in the ray-finned Kvist S (2019) The salivary transcriptome of admixture in the sponge Dendrilla fishEurynotus crenatus (Actinopterygii: Limnobdella mexicana (Annelida: Clitellata: antarctica from shallow waters surrounding Eurynotiformes): Implications for trophic Praobdellidae) and orthology determination the Antarctic Peninsula. Mol Ecol 28:3151- diversification among Carboniferous of major leech anticoagulants. Parasitology 3170 actinopterygians. Earth Environ Sci Trans R 146:1338-1346 • Lemer S, Bieler R, Giribet G (2019) Soc Edinb 109:33-47 • Jones KE, Angielczyk KD, Pierce SE (2019) Resolving the relationships of clams and • Gainett G, Sharma PP, Fernandes N, Pinto- Stepwise shifts underlie evolutionary cockles: Dense transcriptome sampling da-Rocha R, Giribet G, Willemart RH (2019) trends in morphological complexity of drastically improves the bivalve tree of life. Evolution of a sensory cluster on the legs of the mammalian vertebral column. Nat Proc R Soc B 286:20182684 Opiliones (Arachnida) informs multi-level Commun 10:1-13 • Lendemer J, Thiers B, Monfils AK, 16 more phylogenetic relationships. Zool J Linnean including Ford LS (2019) The Extended Soc 187:143-165 • Kenaley CP, Marecki MC, Lauder GV (2019) The role of an overlooked adductor Specimen Network: A strategy to enhance • Garberoglio FF, Apesteguía S, Simões TR, muscle in the feeding mechanism of ray- US biodiversity collection, promote research Palci A, Gómez RO, Nydam RL, Larsson HCE, finned fishes: Predictions from simulations and education. Bioscience 70:23-30 Lee MSY, Caldwell MW (2019) New skulls of a deep-sea viperfish.Zoology 135:125678 • Lerosey-Aubril R, Ortega-Hernández and skeletons of the Cretaceous legged • Kessler SE, Rainbow MJ, Lichtwark GA, J (2019) Appendicular anatomy of the snake Najash, and the evolution of the artiopod Emeraldella brutoni from the modern snake body plan. Sci Adv 5:eaax5833 Cresswell AG, D'Andrea SE, Konow N, Kelly LA (2019) A direct comparison of biplanar middle Cambrian (Drumian) of western • Gehrke AR, Neverett E, Luo Y-J, Brandt videoradiography and optical motion Utah. PeerJ 7:e7945 A, Ricci L, Hulett RE, Gompers A, Ruby capture for foot and ankle kinematics. Front • Lin Y, Konow N, Dumont ER (2019) How JG, Rokhsar DS, Reddien PW, Srivastava M Bioeng Biotechnol 7:1-10 moles destroy your lawn: The forelimb (2019) Acoel genome reveals the regulatory • Klembara J, Hain M, Ruta M, Berman kinematics of Eastern moles in loose and landscape of whole-body regeneration. compact substrates. J Exp Biol 222:jeb182436 Science 363:1191 DS, Pierce SE, Henrici AC (2019) Inner ear morphology of diadectomorphs and Annual Report 2019 . 2020 27 MCZ PUBLICATIONS

• Lindsay WR, Andersson S, Bererhi B, • Näpflin K, O’Connor EA, Becks L, Bensch • Wang Z, Huang Y, Pierce NE (2019) Höglund J, Johnsen A, Kvarnemo C, Leder S, Ellis VA, Hafer-Hahmann N, Harding KC, Radio telemetry helps record the dispersal EH, Lifjel JT, Ninnes CE, Olsson M, Parker GA, Lindén SK, Olsen MT, Roved J, Sackton patterns of birdwing butterflies in Pizzari T, Qvarnström A, Safran RJ, Svensson TB, Shultz AJ, Venkatakrishnan V, Videvall mountainous habitats: Golden Birdwing O, Edwards SV (2019) Endless forms of E, Westerdahl H, Winternitz JC, Edwards (Troides aeacus) as an example. J Insect sexual selection. PeerJ 7:e7988 SV (2019) Genomics of hosts-pathogen Conserv 23:729-738 interactions: Challenges and opportunities • Liu G, Geng B, Zheng X, Xue Z, Dong across ecological and spatiotemporal • Wang Z, Zhuang H, Wand M, Pierce NE H, Lauder GV (2019) An image-guided scales. PeerJ 7:e8013 (2019) Thitarodes shambalaensis sp.nov. computational approach to inversely (Lepidoptera: Hepialidae): A new host of determine in vivo material properties and • O'Connor EA, Westerdahl H, Burri R, the “caterpillar fungus” Ophiocordyceps model flow-structure interactions of fish Edwards SV (2019) Avian MHC evolution in sinensis supported by genome-wide SNP fins.J Comput Phys 392:578-593 the era of genomics: Phase 1.0. Cells 10:1152 data, with comments on effort towards a robust phylogeny of the genus Thitarodes. • Liu L, Anderson C, Pearl D, Edwards SV • Ortega-Hernández J (2019) Exceptionally Zookeys 885:89-113 (2019) Modern phylogenomics: Building preserved Cambrian fossils in the genomic phylogenetic trees using the multispecies era. In Old Questions and Young Approaches • Whitaker MRL, Baker CMM, Salzman coalescent model. In Evolutionary Genomics. to Animal Evolution. Springer Nature: SM, Martins DJ, Pierce NE (2019) Humana: New York Switzerland Combining stable isotope analysis with DNA metabarcoding improves inferences of • Machado FM, Dos Passos F, Giribet • Ortega-Hernández J, Janssen R, Budd trophic ecology. PLoS One 14:e0219070 G (2019) The use of micro-computed GE (2019) The last common ancestor of tomography as a minimally invasive tool Ecdysozoa had an adult terminal mouth. • Witzmann F, Brainerd EL, Konow N (2019) for anatomical study of bivalves (Mollusca: Arthropod Struct Dev 49:155-158 Eye movements in frogs and salamanders— Bivalvia). Zool J Linnean Soc 186:46-75 Testing the palatal buccal pump hypothesis. • Ortega-Hernández J, Lerosey-Aubril R, Integr Comp Biol 1:1-13 • McHorse BK, Biewener AA, Pierce SE Pates S (2019) Proclivity of nervous system (2019) The evolution of a single toe in preservation in Cambrian Burgess Shale-type • Xu L, Sin SYW, Grayson P, Edwards SV, horses: Causes, consequences, and the way deposits. Proc R Soc B 286:20192370 Sackton TB (2019) Evolutionary dynamics of forward. Integr Comp Biol 59:638-655 sex chromosomes of paleognathous birds. • Pérez-de la Fuente R, Gehler A, Farnum Genome Biol Evol 11:2376-2390 • McLean DJ, Cassis G, Kikuchi DW, Giribet CW, Farrell BD (2019) Digitisation as a G, Herberstein ME (2019) Insincere flattery? tool to promote transparency between • Yang J, Ortega-Hernández J, Drage HB, Du Understanding the evolution of imperfect collections: The case of the Baltic amber KS, Zhang ZG (2019) Ecdysis in a stem-group mimicry. Q Rev Biol 94:395-415 from the Königsberg collection at the euarthropod from the early Cambrian of Museum of Comparative Zoology. Span J South China. Sci Rep 9:5709 • de Medeiros BAS, Núñez-Avellaneda LA, Palaeont 34:145-151 Hernandez AM, Farrell BD (2019) Flower • Young JJ, Grayson P, Edwards SV, Tabin CJ visitors of the licuri palm (Syagrus coronata): • Sackton TB, Grayson P, Cloutier A, Hu (2019) Attenuated Fgf signaling underlies Brood pollinators coexist with a diverse Z, Liu JS, Wheeler NE, Gardner PP, Clarke the forelimb heterochrony in the emu community of antagonists and mutualists. JA, Baker AJ, Clamp M, Edwards SV (2019) Dromaius novaehollandiae. Curr Biol Biol J Linnean Soc 126:666-687 Convergent regulatory evolution and loss 29:3681-3691.e5 of flight in paleognathous birds.Science • Meegaskumbura M, Senevirathne 364:74-78 • Zazzi-Boulou AG, Zimkus BM, G, Manamendra-Arachchi K, Goma-Tchimbakala J, Boudzoumou S, Pethiyagoda R, Hanken J, Schneider CJ • Strong CRC, Simões TR, Caldwell MW, Mboungou-Louiki A, Jackson K (2019) (2019) Diversification of shrub frogs Doschak MR (2019) Cranial ontogeny of Herpetological survey of the proposed (Rhacophoridae, Pseudophilautus) in Sri Thamnophis radix (Serpentes: Colubroidea) Bambama-Sanaga Mine Site, Department Lanka­—Timing and geographic context. Mol with a re-evaluation of current paradigms of of Lékoumo, Republic of Congo. Herpetol Phylogenet Evol 132:14-24 snake skull evolution. R Soc open sci 6:182228 Conserv Biol 14:470–480 • Michalik P, Kallal R, Dederichs TM, • Valente RM, Da Silva PAL, de Medeiros • Zhai D, Ortega-Hernández J, Wolfe Labarque FM, Hormiga G, Giribet G, BAS (2019) The first species ofCotithene JM, Hou XG, Cao CJ, Liu Y (2019) Three- Ramírez MJ (2019) Phylogenomics and Voss (Coleoptera: Curculionidae: dimensionally preserved appendages in an genital morphology of cave raptor spiders Curculioninae) from Amazonian Brazil, early Cambrian stem-group pancrustacean. (Araneae, Trogloraptoridae) reveal an with notes on its role as a pollinator Curr Biol 29:171-177 independent origin of a flow-through female of Evodianthus funifer (Poit.) Lindm. genital system. J Zool Syst Evol Res 57:737-747 (Cyclanthaceae). Zootaxa 4576:461-482 • Zhu J, White C, Wainwright DK, Di Santo V, Lauder GV, Bart-Smith H • Mignano AP, Kadapa S, Tangorra JL, • Wainwright DK, Fish FE, Ingersoll S, (2019) Tuna robotics: A high-frequency Lauder GV (2019) Passing the wake: Using Williams TM, Leger JS, Smits AJ, Lauder GV experimental platform exploring the multiple fins to shape forces for swimming. (2019) How smooth is a dolphin? The ridged performance space of swimming fishes.Sci Biomimetics 4:23 skin of odontocetes. Biol Lett 15:20190103 Robot 4:eaax4615 • Morris ZS, Vliet KA, Abzhanov A, Pierce SE • Walker NS, Fernández R, Sneed JM, • Zhu X-J, Lerosey-Aubril R, Ortega- (2019) Heterochronic shifts and conserved Paul VJ, Giribet G, Combosch DJ (2019) Hernández J (2019) Occurrence of the embryonic shape underlie crocodylian Differential gene expression during substrate aglaspidid arthropod Beckwithia in the craniofacial disparity and convergence. Proc probing in larvae of the Caribbean coral Furongian Guole Konservat-Lagerstätte of R Soc B 286:20182389 Porites astreoides. Mol Ecol 28:4899-4913 South China. Palaeoworld 28:73-79

28 Museum of Comparative Zoology Financial Data These charts describe the income and expenses of the Museum of Comparative Zoology in fiscal year 2020. Endowment income funds much of the Museum’s activities, Reserves indicates net growth of balances in unrestricted gifts such as acquisition and maintenance of collections, faculty and endowments from, for example, interest payments and and staff salaries, capital projects, facilities renovation and unspent portions of the current year’s endowment payouts. maintenance. It includes the annual distribution (payout) and Accumulation of Restricted Reserves indicates net growth of endowed funds decapitalized per donor request. Nonfederal balances in highly restricted gifts and endowments. Building Sponsored Revenue does not include HHMI funds awarded expenses such as maintenance, facility improvements and to Prof. Hoekstra. Gifts are donations received in support utilities are captured in the Space & Occupancy category. of Museum activities that are available for current use; it Operating Expenses consist of equipment purchases, does not include donations for endowed funds. Transfers supplies, and consultant and conference fees, as well as include financial support for the Ernst Mayr Library and annual subventions to the Department of Organismic and other Harvard-funded activities. Other Income comprises Evolutionary Biology (OEB) for administrative services miscellaneous income from publication subscriptions, and MCZ support for faculty-curator research. Support royalties, sales and fees, and cost recovery from other MCZ- for MCZ-affiliated graduate students in OEB is included in sponsored activities. Overhead is funds paid from sponsored Scholarships, Awards & Travel. Institutional Expenses are projects to cover associated facilities and administrative costs. support for other University activities outside the MCZ, It is shown as both income (Overhead Earned) and expenses including FAS and University initiatives and general operating (Overhead Charged). Accumulation of Unrestricted support to the Harvard Museums of Science and Culture.

INCOME EXPENSES & NON-OPERATING FUNDS

Endowment 85% Capitalized Salaries & Fringe Balances 0.2% Benefits 40%

Overhead Charged (Sponsored) 3%

Scholarships, Awards & Travel 4% Gifts 0.6% Space & Operating Occupancy Expenses Nonfederal Federal 15% 21% Sponsored Sponsored Revenue 1% Other Revenue 8% Income Transfers Overhead Institutional 1.2% 2% Earned 3% Expenses 17%

Income Expenses Endowment $16,975,583 Salaries & Fringe Benefits $7,695,344 Federal Sponsored Revenue $1,525,650 Operating Expenses $4,040,458 Overhead Earned $579,254 Institutional Expenses $3,272,435 Transfers $403,783 Space & Occupancy $3,005,982 Other Income $242,974 Scholarships, Awards & Travel $731,341 Nonfederal Sponsored Revenue $143,532 Overhead Charged (Sponsored) $579,254 Gifts $111,067 Capitalized Balances $40,144 Accumulation of Unrestricted Reserves ($362,813) Total $19,364,958 Accumulation of Restricted Reserves ($254,072) Total $19,364,958

Annual Report 2019 . 2020 29 Personnel

Faculty-Curators Emeritus Faculty Robert Kambic Andrew A. Biewener A.W. “Fuzz” Crompton Vertebrate Paleontology, S. Pierce Lab Charles P. Lyman Professor of Fisher Professor of Natural History, Andreas Kautt Biology; Director, Concord Field Emeritus Mammalogy, Hoekstra Lab Station Richard C. Lewontin Michael Kelly Scott V. Edwards Professor of Biology, Emeritus; Invertebrate Zoology, Giribet Lab Professor of Organismic & Alexander Agassiz Professor of Evolutionary Biology; Alexander Zoology, Emeritus Sangeet Lamichhaney Ornithology, Edwards Lab Agassiz Professor of Zoology; Edward O. Wilson Curator of Ornithology Honorary Curator in Entomology; Rudy Lerosey Brian D. Farrell University Research Professor, Invertebrate Paleontology, Ortega- Monique & Philip Lehner Professor Emeritus Hernández Lab for the Study of Latin America, Robert M. Woollacott Marjorie Liénard Professor of Organismic & Professor of Biology, Emeritus Entomology, N. Pierce Lab Evolutionary Biology; Curator of Entomology; Director, David Cong Liu Rockefeller Center for Latin Postdoctoral Fellows, Research Entomology, N. Pierce Lab American Studies Associates & Visiting Scholars Gang Liu Gonzalo Giribet Nicole Bedford Ornithology, Edwards Lab Professor of Organismic & Mammalogy, Hoekstra Lab Yu Liu Evolutionary Biology; Alexander Ligia Benavides Silva Invertebrate Paleontology, Ortega- Agassiz Professor of Zoology; Invertebrate Zoology, Giribet Lab Hernández Lab Harvard College Professor; Curator of Invertebrate Zoology D. Marcela Bolaños Yi-Jyun Luo Invertebrate Zoology, Srivastava Lab Invertebrate Zoology, Srivastava Lab James Hanken Professor of Biology; Alexander Gustavo Bravo Juan Moles Agassiz Professor of Zoology; Curator Ornithology, Edwards Lab Invertebrate Zoology, Giribet Lab of Herpetology; Acting Curator of Robert Brocklehurst Andres Moya Simarro Malacology; Director, MCZ Vertebrate Paleontology, S. Pierce Lab Invertebrate Zoology, Giribet Lab Hopi E. Hoekstra Daren Card Luciano Naka Professor of Organismic & Ornithology, Edwards Lab Ornithology, Edwards Lab Evolutionary Biology; Professor Jennifer Chen Kathrin Näpflin of Molecular & Cellular Biology; Ornithology, Edwards Lab Alexander Agassiz Professor of Mammalogy, Hoekstra Lab Zoology; Curator of Mammalogy, Marina Cheng Stephen Pates Howard Hughes Medical Institute Invertebrate Zoology, Giribet Lab Invertebrate Paleontology, Ortega- Investigator Hernández Lab Shahan Derkarabetian George V. Lauder Invertebrate Zoology, Giribet Lab Altai Carlos Pavon Paneque Henry Bryant Bigelow Professor of Invertebrate Zoology, Giribet Lab Ichthyology; Curator of Ichthyology; Valentina Di Santo Ichthyology, Lauder Lab Lorenzo Ricci Harvard College Professor Invertebrate Zoology, Srivastava Lab James J. McCarthy Terry Dial Ichthyology, Lauder Lab Mehdi Saadat Professor of Biological Ichthyology, Lauder Lab Oceanography; Alexander Lu Dong Agassiz Professor of Biological Ornithology, Edwards Lab Nacho Sanguinetti-Scheck Oceanography; Acting Curator of Mammalogy, Hoekstra Lab Malacology Miquéias Ferrão Herpetology, Hanken Lab Elizabeth Sibert Javier Ortega-Hernández Ichthyology, Lauder Lab Assistant Professor of Organismic Molly Gabler Ichthyology, Lauder Lab Tiago Simões and Evolutionary Biology; Curator Vertebrate Paleontology, S. Pierce Lab of Invertebrate Paleontology Andrew Gehrke Invertebrate Zoology, Srivastava Lab Bruno Souza de Medeiros Naomi E. Pierce Entomology, Farrell Lab Sidney A. and John H. Hessel Elsa Goerig Professor of Biology; Curator of Ichthyology, Lauder Lab Flavia Termignoni Garcia Lepidoptera Ornithology, Edwards Lab Carlos Guerra Schrago Stephanie E. Pierce Ornithology, Edwards Lab Robin Thandiackal Thomas D. Cabot Associate Ichthyology, Lauder Lab Professor of Organismic & Aaron Hartmann Hrdy Fellow, Giribet Lab João Tonini Evolutionary Biology; Curator of Entomology, N. Pierce Lab & Vertebrate Paleontology Caroline Hu Ornithology, Edwards Lab Mammalogy, Hoekstra Lab Mansi Srivastava Kelsey Tyssowski John L. Loeb Associate Professor Katrina Jones Mammalogy, Hoekstra Lab of the Natural Sciences; Curator of Vertebrate Paleontology, S. Pierce Lab Invertebrate Zoology Dylan Wainwright Nicholas Jourjine Ichthyology, Lauder Lab Mammalogy, Hoekstra Lab

30 Museum of Comparative Zoology PERSONNEL Personnel

Pengcheng Wang Philip Fahn-Lai Ruth Hortencia Bastardo Michael Huben Ornithology, Edwards Lab Concord Field Station, Biewener Associate of Entomology Associate of Entomology Lab & Vertebrate Paleontology, S. Universidad Autónoma de Santo Independent Researcher Fernanda de Pinho Werneck Domingo Ornithology, Edwards Lab Pierce Lab Samuel Jaffe Mara Laslo Aaron Bauer Associate of Entomology Megan Whitney Associate of Herpetology Independent Researcher Vertebrate Paleontology, S. Pierce Lab Herpetology, Hanken Lab Villanova University Alan Kabat Joanna Wolfe Sarah Losso Invertebrate Paleontology, Ortega- Reinier Beeuwkes III Associate of Malacology Invertebrate Paleontology, Ortega- Associate of the Concord Field Attorney, Bernabei & Wachtel Hernández Lab Hernández Lab Station Leslie S. Kaufman Donghao Wu Inbar Maayan Ischemix, LLC (MA) Herpetology Associate of Ichthyology Entomology, N. Pierce Lab Andrew Berry Boston University Tao Xu Sophia MacRae Orzechowski Associate of Population Genetics Ornithology, Edwards Lab Anthony E. Kiszewski Ornithology, Edwards Lab Harvard University Associate of Entomology Wenqing Ye Marc Mapalo Elizabeth Brainerd Bentley University Invertebrate Paleontology, Ortega- Entomology, N. Pierce Lab Associate of Ichthyology Nicolai Konow Hernández Lab Brown University Chuang Zhou Associate of the Concord Field Ornithology, Edwards Lab David Matthews Jae Choe Station Ichthyology, Lauder Lab Associate of Entomology University of Massachusetts, Lowell Olivia Meyerson Ewha Womans University Timothy Laman Graduate Students Mammalogy, Hoekstra Lab Rockwell Anyoha Janet Collett Associate of Ornithology Mammalogy, Hoekstra Lab Zachary Morris Associate of Population Genetics National Geographic Vertebrate Paleontology, S. Pierce Lab University of Sussex Jennifer Austiff Phillip Lobel Herpetology, Hanken Lab Sofia Prado-Irwin Bruce Collette Associate of Ichthyology Ornithology, Edwards Lab Associate of Ichthyology Boston University Felix Baier National Marine Fisheries Service Mammalogy, Hoekstra Lab Amber Rock David Lohman Invertebrate Zoology, Srivastava Lab David Bruce Conn Associate of Entomology Caitlin Baker Associate of Invertebrate Zoology The City College of New York Invertebrate Zoology, Giribet Lab Samantha Royle Berry College Herpetology, Hanken Lab Vladimir A. Lukhtanov Wei-Ping Chan James Costa Associate of Entomology Entomology, N. Pierce Lab Shayla Salzman Associate of Entomology Russian Academy of Sciences Entomology, N. Pierce Lab Richard Childers Western Carolina University D. Luke Mahler Entomology, N. Pierce Lab Shoyo Sato Catherine Craig Associate of Herpetology Invertebrate Zoology, Giribet Lab Tauana Cunha Associate of Invertebrate Zoology University of Toronto Invertebrate Zoology, Giribet Lab Carl Jonathan Schmitt Conservation through Poverty James Mallet Ornithology, Edwards Lab Alleviation, International Blake Dickson Associate of Population Genetics Vertebrate Paleontology, S. Pierce Lab Kari Taylor-Burt Harlan Dean Harvard University Concord Field Station, Biewener Lab Associate of Invertebrate Zoology Russell Mittermeier Alexandria DiGiacomo Harvard University Ornithology, Edwards Lab Wendy Valencia Montoya Associate of Herpetology Entomology, N. Pierce Lab Philip DeVries Conservation International Jake Gable Zhengyang Wang Associate of Entomology Piotr Naskrecki Mammalogy, Hoekstra Lab University of New Orleans Entomology, N. Pierce Lab Associate of Entomology Emily Hager Gregory D. Edgecombe Connor White Conservation International Mammalogy, Hoekstra Lab Associate of Invertebrate Zoology Ichthyology, Lauder Lab Diane B. Paul Michael Brent Hawkins Natural History Museum, England Rebecca Zane Wolf Associate of Population Genetics Herpetology, Hanken Lab Ben Evans Harvard University Ichthyology, Lauder Lab Alyssa Hernandez Associate of Herpetology David L. Pawson Tyler Brock Wooldridge Entomology, Farrell Lab McMaster University Associate of Marine Biology Mammalogy, Hoekstra Lab Evan Hoki Brooke E. Flammang Smithsonian National Museum of Entomology, N. Pierce Lab Mark Wright Associate of Ichthyology Natural History Vertebrate Paleontology, S. Pierce Lab New Jersey Institute of Paulo Petry Ryan Hulett Technology Invertebrate Zoology, Srivastava Lab Xuemei Zhai Associate of Ichthyology Biological Oceanography, Michael Hadfield The Nature Conservancy Sang Il Kim McCarthy Lab Associate of Marine Biology Entomology, Farrell Lab Ricardo Pinto da Rocha University of Hawaii Associate of Invertebrate Zoology Julian Kimura Associates Berthold Hölldobler University of São Paulo Invertebrate Zoology, Srivastava Lab Gary Alpert Associate of Entomology Michael Reed Richard Knecht Associate of Entomology Arizona State University Associate of Ornithology Invertebrate Paleontology, Ortega- Harvard University Gustavo Hormiga Tufts University Hernández Lab Bruce Archibald Associate of Invertebrate Zoology Michael Rex Vanessa Knutson Associate of Entomology The George Washington Associate of Malacology Invertebrate Zoology, Giribet Lab Simon Fraser University University University of Massachusetts, Boston

Annual Report 2019 . 2020 31 PERSONNEL

R. Graham Reynolds Eva Biedron Nikki Hughes Associate of Herpetology Curatorial Assistant, Invertebrate & Faculty/Collection Assistant, University of North Carolina, Vertebrate Paleontology Mammalogy & Invertebrate Asheville Emily Blank Zoology Jessica Rykken Curatorial Assistant, Collections Amie Jones Associate of Entomology Operations Faculty/Collection Assistant, Harvard University Ronnie Broadfoot Entomology Carl Schmitt Coordinator of Access Services, Marcia Kazmierczak Associate of Ornithology Ernst Mayr Library Staff Assistant, Collections Independent Researcher Christina Byrd Operations Donna Schmitt Curatorial Associate, Vertebrate Michelle Kennedy Associate of Ornithology Paleontology Collections Information & Database Independent Researcher Christopher Capobianco Specialist Andrea Sequeira Technician & Preparator, Vertebrate Christopher Kirby Associate of Entomology Paleontology Research Assistant, Hoekstra Lab Wellesley College April Collins Jeremy Kisala Steven O. Shattuck Acquisitions & Technology Curatorial Assistant, Collections Associate of Entomology Specialist, Ernst Mayr Library Operations Commonwealth Scientific & Stefan Cover Laura Leibensperger Industrial Research Organization Curatorial Assistant, Entomology Curatorial Assistant, Invertebrate (CSIRO) Jessica Cundiff Zoology Scott R. Shaw Curatorial Associate, Invertebrate & Lisa Litchfield Associate of Entomology Vertebrate Paleontology Administrator, Concord Field University of Wyoming Joseph deVeer Station Stephen Tilley Head of Technical Services, Ernst David Lubertazzi Associate of Herpetology Mayr Library Research Scientist, Entomology Smith College Katherine Eldridge Crystal Maier James Traniello Curatorial Assistant, Ornithology Curatorial Associate, Entomology Associate of Entomology Boston University Charles Farnum Joseph Martinez Curatorial Assistant, Entomology Curatorial Assistant, Herpetology David Wake Associate of Herpetology Helene Ferranti Patrick McCormack University of California, Berkeley Faculty/Collection Assistant, Curatorial Assistant, Entomology Biological Oceanography & Marine Sade McFadden Marvalee Wake Biology Associate of Herpetology Research Assistant, Hoekstra Lab University of California, Berkeley Linda S. Ford John Mewherter Director, Collections Operations Philip S. Ward Curatorial Assistant, Collections Associate of Entomology Matthew Gage Operations University of California, Davis Curatorial Assistant & Lab Paul J. Morris Manager, Hanken Lab Jacqueline Webb Biodiversity Informatics Manager Associate of Ichthyology Emily Graham Madeleine Mullon University of Rhode Island Curatorial Assistant, Collections Curatorial Associate, Mammalogy Operations Haven Wiley Catherine Musinsky Associate of Ornithology Cyrus Green Faculty/Collection Assistant, University of North Carolina, Curatorial Assistant, Invertebrate & Mammalogy Chapel Hill Vertebrate Paleontology Mark Omura Cheryl Wilga Cory Hahn Curatorial Associate, Mammalogy Associate of Ichthyology Animal Technician, Herpetology University of Rhode Island Melinda Peterson Brendan Haley Research Lab Coordinator, Edwards Lab Judith Winston Senior Database Manager Associate of Invertebrate Zoology Bridget Power Rachel Hawkins Faculty/Collection Assistant, Virginia Museum of Natural Curatorial Assistant, Entomology History Invertebrate Zoology Andra Hollis Pedro Ramirez Staff Assistant, Concord Field Research Assistant, Concord Field Staff Station Melissa Aja Station Museum Projects Coordinator & Kathleen Horton Rona Razon Interim Managing Editor Assistant with Professor Wilson, Archivist, Ernst Mayr Library Entomology Adam Baldinger Murat Recevik Curatorial Associate, Invertebrate Jared Hughes Curatorial Assistant, Malacology Zoology & Malacology Faculty/Collection Assistant, Herpetology, Invertebrate & Mark Renczkowski Penny Benson Vertebrate Paleontology Curatorial Assistant, Invertebrate Curatorial Assistant, Malacology Paleontology

32 Museum of Comparative Zoology PERSONNEL

Constance Rinaldo Chloe-Rose Colombero Daniel Stevens Librarian, Ernst Mayr Library Ichthyology Collection Operations Alana Rivera Brendan Cramphorn Dexter Summers Curatorial Assistant, Collections Entomology Ichthyology Operations Sheridan Cunningham Gary Taylor José Rosado Ernst Mayr Library Concord Field Station Curatorial Associate, Herpetology Caroline Elson Chidambaram Thillairajah The MCZ deeply appreciates Mary Sears Entomology Ornithology the additional support and Head of Public Services, Ernst Mayr contributions of numerous Anne Everly Amy Wu interns and undergraduate Library Ichthyology Entomology Kaitlin Sheridan students during the 2019–2020 Cleo Falvey academic year. Laboratory Assistant, Invertebrate Herpetology Zoology Administration for the Caroline Fleming Department of Organismic Meaghan Sorce Invertebrate Zoology & Evolutionary Biology Curatorial Assistant, Ichthyology Jose Flores Lydia Carmosino Margaret Starvish Ichthyology Senior Academics Programs Faculty/Collection Assistant, Administrator MCZ Faculty Entomology & Ichthyology Zoe Flores Entomology Rebecca Chetham The MCZ’s charter, signed in Tsuyoshi Takahashi Executive Director Curatorial Assistant, Herpetology & Julia Foster 1859, mandates that the Collections Operations Ornithology Sarine Der Kaloustian Museum’s activities will be Associate Director of overseen by a governing Jennifer Thomson Ella Frigyik Administration board, the Faculty of the Lab Administrator, Srivastava Lab Entomology Paul Dwyer Museum of Comparative Jennifer Trimble Andrea Garza Erdmann Mailroom Staff Assistant Zoology. Curatorial Assistant, Invertebrate Ernst Mayr Library Zoology Jason Fleming Mr. Robert G. Goelet Jennifer Goldstein Senior Research Administrator Jeremiah Trimble Invertebrate Zoology Mr. George Putnam III Curatorial Associate, Ornithology Christian Flynn Jane Harrison Administrative Coordinator Mr. Jeff Tarr Diana Turmenne Ornithology Curatorial Assistant, Collections Donna Gadbois Dr. Barbara Jil Wu Evan Hoki Financial Associate Operations Entomology Mr. Paul J. Zofnass Kenneth Wilcox Diana Gjino President Lawrence S. Bacow Anna Holden Senior Research Administrator Building Services Coordinator Malacology Andrew Williston Jason Green Kalvin Janik Financial Associate Curatorial Associate, Ichthyology Ernst Mayr Library Wendy Heywood Greta Wong Jackson Kehoe Laboratory Assistant, Ichthyology Communications & Events Entomology Coordinator Jonathan Woodward Acknowledgements Adam Kowalczyk Julie Knippa Colby Curatorial Assistant, Collections Collections Operations This annual report was Operations Associate Director of Finance and produced by the Office of the Research Administration Philip Fahn-Lai Director of the Museum of Mark Wright Vertebrate Paleontology Research Lab Assistant, Vertebrate Megan McHugh Comparative Zoology. Kevin Liu Human Resources Administrator Paleontology Editors Invertebrate Zoology Jeremiah (JJ) O’Connor Robert Young James Hanken, Director Special Collections Librarian, Ernst Ian Magill Temporary Financial Analyst Mayr Library Concord Field Station Melissa Aja, Museum Projects Kristin Pennarun Coordinator Clayton Ziemke Francisco Matos Assistant Director of Research Curatorial Assistant, Entomology Entomology Administration Copy, Design & Production Breda Zimkus John Nevins Christopher Preheim Cyndi Wood Assistant Director, Collections Biological Oceanography Temporary Academic Program Creative Project Coordinator Operations Uzochi Nwoko Management, Inc. Ernst Mayr Library Keleigh Quinn creativeprojectmgmt.com Senior Research Administrator Temporary Staff Avalon Owens Griffin Andres Entomology Emily Reynolds Ichthyology Senior Research Administrator Carolina Sepulveda Adham Bedir Ernst Mayr Library Peg Richards Ernst Mayr Library Financial Assistant Toby Shaya Portia Berry-Kilby Entomology Anna Salvato Ernst Mayr Library Manager of Financial Operations Lauren Shear William Clark Collection Operations Ernst Mayr Library

Annual Report 2019 . 2020 33 26 Oxford Street Cambridge, MA 02138 617.495.2460 mcz.harvard.edu

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