Museum of Comparative Zoology

Harvard University

Annual Report 2015. 2016

Director’s Message This past year we celebrated marine biology at the MCZ and the completion of several facilities renovation projects, some of which had been in the works for years.

Harvard’s collection of Blaschka glass bright, climate-controlled rooms with marine invertebrates made its literary modern presentation technology and debut in Sea Creatures in Glass. This book better functioning—and much more offers a brief history of the Blaschka comfortable—tables and chairs. family’s manufacture of remarkable, lifelike Sadly, we lost one of our MCZ Faculty glass models and details the decade-long members this year. Dr. John Constable conservation process that restored and was a longtime member of the board of stabilized MCZ’s priceless collection. directors, whose existence dates back Eighty-eight pages of exquisite, detailed to the founding of the museum in 1859. color photographs follow the text. A remarkable man in many respects, Marine Life in the Putnam Family Gallery, John came from a distinguished British a permanent exhibit in the Harvard family. His father, a distant relative of the Museum of Natural History, opened in English Romantic painter John Constable, Melissa Aja Melissa 2015. The exhibit features specimens from immigrated to the U.S. to become curator several MCZ collections and modern of the (then) Boston Museum of Fine James Hanken models, which both immerse visitors Arts. John began his association with the Director in an underwater world and feature MCZ as a teenager in the 1940s, during contemporary research and conservation World War II, when he began a taxonomic efforts. We are tremendously grateful to study of the reptiles of India under the the Putnam family for their generous gift, supervision of herpetology curator Arthur which supported this new installation. Loveridge and in response to a request from the U.S. defense department, which In the spring, we celebrated completion needed help in producing an identification of the MCZ’s state-of-the-art collections guide to poisonous snakes. facility in the adjacent Northwest Building with a dedication and open house where Finally, we initiated a formal search for a hundreds of guests from the Cambridge faculty member/Curator of Invertebrate campus and beyond toured the two Paleontology to fill a position that has floors of storage, teaching and lab spaces. been vacant for several years. At the time Collections staff spent weeks preparing for this report goes to press, I am optimistic the event, wowing visitors with a rainbow- that we will welcome a new colleague in like, 15-foot display of birds and offering a the 2018–2019 academic year. rare glimpse of the Blaschka glass marine As you read more about the MCZ’s invertebrates in their storage cases, among newsworthy research and projects, other crowd-pleasers. remember that none of it would be Collection rooms were not the only possible without the ongoing efforts of our spaces to be modernized and upgraded— many faculty-curators, staff, students and About the Cover: The newly discovered Conception the MCZ renovated two classrooms, a postdocs, as well as the critical support we Bank silver boa (Chilabothrus conference room, and a staff/student receive from the Harvard administration argentum). Photo by R. Graham lounge. The charm of our 19th-century and dedicated alumni. We look forward to Reynolds. Opposite page: One of the MCZ's main building was maintained in all reporting more next year. recently renovated classrooms. spaces, while transforming them into Photo by Melissa Aja.

Annual Report 2015 . 2016 1 Highlighting the Importance of Collections Harvard’s Northwest Building provides more than ordinary storage for many collections of the MCZ.

“MCZ’s new facility in the Northwest Zoology collections, plus the Blaschka glass Building is carefully designed to provide a animals and the historic Harvard Embryological secure, temperature- and humidity-controlled Collection. Specimens are housed in custom- space to preserve six million specimens,” designed steel cabinets that maximize use of available space, including those for oversized specimens such as elephant hides, kangaroos and ostriches. A wall-mounted pegboard system creatively stores hundreds of mounted bird specimens prepared for exhibition, and precise climate control suppresses insect infestations without the use of pesticides. Preparation spaces have specialized equipment that makes it easier and safer for staff to work with particular types of specimens. The Paleontology lab has rock saws, dust collectors and giant fume hoods, while the Vertebrate Zoology lab is equipped to process tissue samples and store them cryogenically. To date, the environmentally friendly, energy- efficient 50,000-square-foot build-out has received nine LEED certifications.

Melissa Aja Melissa In March 2016, the facility’s official grand opening attracted more than 200 MCZ says Director James Hanken. “It facilitates personnel, the MCZ Faculty, and members of the use of these collections for research and the Harvard administration and community. specimen-based teaching and offers faculty, Director James Hanken and Faculty of Arts and collections staff, visitors and students a pleasant Sciences Dean Michael D. Smith addressed the work environment.” visitors, who then enjoyed refreshments and Planning for this space began long collection tours. before construction started. Consultant Jeff Weatherston of Toronto-based WeatherstonBruer Associates spent years performing an extensive space-needs analysis to determine how to best accommodate collections destined for the facility. Now, the space is filled with all or part of seven collections accumulated during the MCZ’s 157-year history. The building houses portions of the Invertebrate Paleontology, Ornithology, Vertebrate Paleontology, Malacology, Mammalogy, Entomology and Invertebrate Melissa Aja Melissa

2 Museum of Comparative Zoology MCZ Faculty-Curators

Andrew A. Biewener Charles P. Lyman Professor of Biology Director, Concord Field Station Prof. Biewener’s research focuses on understanding the biomechanics, neuromuscular control and energetics 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 their movement in natural environments.

Scott V. Edwards Professor of Organismic & 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 utilize genomics technologies to study comparative genomics and the evolution of flightlessness in birds; phylogeography and speciation of Australian and North American birds; and the genomics of host–parasite co-evolution between house finches and a recently acquired bacterial pathogen,Mycoplasma . Tony Rinaldo Tony

Brian D. Farrell Gonzalo Giribet Professor of Biology Professor of Organismic & Curator of Entomology Evolutionary Biology Director, David Alexander Agassiz Rockefeller Center for Professor of Zoology Latin American Studies Curator of Invertebrate Zoology Prof. Farrell’s research is broadly concerned Prof. Giribet’s primary with the evolution of research focuses on the ecological interactions evolution, systematics between host plants and biogeography of and animals and their invertebrate animals, parasites, such as including the use insects and other tiny of morphology and Stephanie Mitchell consumers. His current next-generation Dunn © Casey projects include applying next-generation sequencing to sequencing techniques. speciation and phylogenetic studies of associated species, Current projects in the Giribet lab include the evolution documenting biodiversity in the Dominican Republic, of orb-weaving spiders and other arachnids, and and repatriating digital information from scientific systematics and biogeography of arthropods, mollusks and specimens of insects and fossils in museums to their onychophorans, among other groups. He is also interested countries of origin. in the use of genomic-level data for inferring phylogenies.

Annual Report 2015 . 2016 3 FACULTY-CURATORS

James Hanken Professor of Biology Alexander Agassiz Professor of Zoology Curator of Herpetology MCZ Director Prof. Hanken utilizes laboratory-based analyses and field surveys to examine morphological evolution, developmental biology

and systematics. ErtlGretchen Current areas of Hopi E. Hoekstra Catherine Weisel research include the evolution of craniofacial patterning; the developmental Professor of Organismic & Evolutionary Biology basis of morphological novelty; biodiversity informatics; Professor of Molecular & Cellular Biology and systematics and evolution of neotropical salamanders. Alexander Agassiz Professor of Zoology Prof. Hanken also serves on the Executive Committee of the Curator of Mammalogy Encyclopedia of Life (eol.org). Howard Hughes Medical Institute Investigator Harvard College Professor Prof. Hoekstra combines field and laboratory work to understand the evolution of mammalian diversity from morphology to behavior. Her research focuses on the genetic basis of adaptive variation—identifying both the ultimate causes and the proximate mechanisms responsible for traits that help organisms survive and reproduce in the wild. Research in the Hoekstra lab integrates ecological, behavioral, genetic and molecular approaches.

Jonathan B. Losos Monique & Philip Lehner Professor for the Study of

Stephanie Mitchell Latin America Professor of Organismic & George V. Lauder Evolutionary Biology Professor of Biology Curator of Herpetology Henry Bryant Bigelow Professor of Ichthyology Prof. Losos’ research Curator of Ichthyology focuses on the behavioral Prof. Lauder’s research focuses on the biomechanics of fishes and evolutionary ecology and the development of robotic models for studying aquatic of lizards, specifically locomotion. how lizards interact with His current studies focus on the function of shark skin and their environment and how lizard clades have

other surface structures, the role of flexibility in improving Lincoln Rose the efficiency of aquatic propulsion, and how fishes control diversified evolutionarily. body and fin position as they maneuver through obstacles. His laboratory integrates approaches from systematics, Additional broad interests include biological fluid mechanics ecology, behavior, genetics and functional morphology, and theoretical approaches to the analysis of form and taking both observational and experimental approaches in function in organisms. the field and in the laboratory.

4 Museum of Comparative Zoology FACULTY-CURATORS

James J. McCarthy Naomi E. Pierce Professor of Biological Sidney A. & John Hessel Oceanography Professor of Biology Alexander Agassiz Professor of Curator of Lepidoptera Biological Oceanography Acting Curator of Malacology Prof. Pierce’s research focuses on the behavioral Prof. McCarthy’s research ecology of species focuses on factors that interactions, particularly regulate the processes of insect/plant associations, primary production and and symbioses between nutrient supply in the ants and other organisms, ocean. Using field studies including bacteria, fungi, and modeling, Prof. plants and caterpillars of Kris SnibbeKris Jon Chase McCarthy and his group butterflies in the family examine the effects of Lycaenidae. Prof. Pierce is seasonal or interannual climate change on marine life interested in how parasitic and mutualistic life histories can from plankton to whales. influence the evolutionary trajectories of each partner.

Stephanie E. Pierce Assistant Professor of Organismic & Evolutionary Biology Curator of Vertebrate Paleontology Prof. Pierce’s research is focused on major morphological and ecological transitions

in vertebrate BertrandJean-Francois evolution through an examination of the Robert M. Woollacott fossil record. Professor of Biology Paul Whatmough Paul Curator of Marine Invertebrates Her work tends toward 3-D modeling and experimentation of the musculoskeletal system, with particular attention Prof. Woollacott’s research focuses on aspects of marine to the link between form and function. Current projects invertebrate life history, such as synchronization of reproductive include the fin-to-limb transition, the evolution of the events and ecology and physiology of larvae. Topics of particular mammalian backbone, and the origin of the avian neck. interest include larval dispersal and population connectivity, as well as human impacts on the distribution of marine organisms.

Mansi Srivastava Assistant Professor of Organismic & Evolutionary Biology Curator of Invertebrate Zoology Dr. Srivastava's research focuses on understanding the evolution of animal development and regeneration. Her group utilizes the three-banded panther worm, Hofstenia miamia, which Dr. Srivastava has developed as a new acoel model system. Acoels represent the earliest lineage of animals with bilateral symmetry, which allows the study of genetic mechanisms that span 550 million years of animal evolution. Current projects in the lab range from identifying gene regulatory networks for regeneration to

Isaac Orderberg determining the embryonic origins of pluripotent stem cells.

Annual Report 2015 . 2016 5 MCZ Emeriti

A. W. “Fuzz” Crompton Faculty-Curator, Emeritus Fisher Professor of Natural History, Emeritus Prof. Crompton, former Curator of Mammalogy, was the Director of the MCZ from 1970 to 1982 and the former Director of the Peabody Museum of Natural History, Yale University, and the South African Museum, Capetown. His primary research interests are the origin and evolution of mammals, functional anatomy, neural control and evolution of feeding in recent and fossil vertebrates. Prof. Crompton received two Guggenheim fellowships for his research on vertebrate paleontology and functional morphology and in 2011 received the Romer-Simpson Medal from the Society of Vertebrate Paleontology. Justin Ide Justin

Richard C. Lewontin Professor of Biology, Emeritus Alexander Agassiz Professor of Zoology, Emeritus An evolutionary geneticist, Prof. Lewontin pioneered the field of molecular population genetics by merging molecular biology and evolutionary theory, as well as the philosophical and social implications of genetics and evolutionary theory. Prof. Lewontin’s current research involves computer simulation and evaluation of statistical tests for selection. Among his many books are The Genetic Basis of Evolutionary Change; Biology as Ideology: The Doctrine of DNA; Human Diversity; and The Triple Helix: Gene Organism and Environment.

Edward O. Wilson Honorary Curator in Entomology Pellegrino University Professor, Emeritus Prof. Wilson is considered the founder of sociobiology and evolutionary psychology and has developed the basis of modern biodiversity conservation. He has received many of the world’s leading prizes in recognition of his research and environmental activism. He was awarded two Pulitzer Prizes for his books The Ants (1990, with Bert Hölldobler) and On Human Nature (1978). Prof. Wilson received the TED Prize in 2007, where he articulated the concept of the Encyclopedia of Life, and the National Geographic Society’s Hubbard Medal

Beth Maynor Young Beth in 2013.

6 Museum of Comparative Zoology Courses in 2015–2016 Led by MCZ Faculty-Curators Jonathan Losos Jonathan

OEB 101: Biology of Mammals Organismic and Evolutionary OEB 130: Biology of Fishes Biology George V. Lauder OEB 57: Animal Behavior Explores the unparalleled diversity of fish Naomi E. Pierce (and Bence P. Olveczky) across different aquatic environments, including deep seas, intertidal zones, coral A review of the behavior of animals under reefs, polar waters, the vast Amazonian basin natural conditions, with emphasis on both and great East African lakes. mechanistic and evolutionary approaches. OEB 130: Biology of Fishes

OEB 101: Biology of Mammals Jonathan Losos An introduction to the biology of mammals. Lectures and laboratories examine the morphology, systematics, natural history, behavior, ecology, evolutionary relationships and biogeography of all major taxa.

OEB 126: Vertebrate Evolution Stephanie Pierce A comprehensive survey of the origin and evolution of vertebrates through an examination of the fossil record, focusing on major events in Earth’s evolutionary history, with an emphasis on anatomical and physiological transformations in fish, amphibians, reptiles, birds and mammals. Morgan Furze

Annual Report 2015 . 2016 7 COURSES George LauderGeorge

OEB 130: Biology of Fishes OEB 155r: Biology of Insects OEB 173: Comparative Biomechanics Naomi E. Pierce (and Michael R. Canfield) Andrew A. Biewener Introduction to the major groups of Explores how animals and plants contend with insects—life history, morphology, physiology their physical environment, considering their and ecology—through a combination of biomaterial properties, structural form and lecture, lab and field exercises. mechanical interactions with the environment. OEB 217: What Makes a Cat? OEB 157: Global Change OEB 181: Systematics Biology Gonzalo Giribet James J. McCarthy Introduces theory and practice of Examines how natural and systematics, emphasizing issues associated anthropogenic changes with homology statements and alignments, in the earth system are methods of tree reconstruction and affecting the composition hypothesis evaluation. and the functioning of the world’s land and ocean OEB 190: Biology and Diversity of Birds ecosystems. Scott V. Edwards An introduction to the biology of birds. OEB 167: Herpetology Covers the fossil record and theories for avian James Hanken and origins, physiology and anatomy, systematics, Jonathan Losos speciation processes, behavior, vocalizations, An introduction to the demography and conservation. biology of amphibians and reptiles. Lectures and OEB 217: What Makes a Cat? laboratories examine the Stephanie Pierce morphology, systematics, Explores the origin and evolution of cats natural history, behavior, through a mixture of literature discussions, ecology, evolutionary cadaveric dissection and fossil observation. relationships and Discussions include evolutionary dynamics, biogeography of all ecomorphology, and feeding and locomotor major taxa. performance. Stephanie Pierce

8 Museum of Comparative Zoology COURSES

OEB 234: Topics in Marine Biology Robert M. Woollacott Explores the fragility and resilience of marine life and ecosystems in the face of perturbations such as habitat fragmentation, land use change, anthropogenic climate change, pollution, alien species and unsustainable fishing practices. General Education Science of Living Systems 22: Human Influence on Life in the Sea Robert M. Woollacott, James J. McCarthy Over-harvested fish stocks, pollution and anthropogenic climate change affect the stability and productivity of marine ecosystems. This course asks what we need Scott Edwards to know about the causes and effects of OEB 190: Biology and Diversity anthropogenic change to best protect OEB 325: Marine Biology of Birds marine ecosystems and ensure sustainable Robert M. Woollacott harvests from the sea. OEB 334: Behavioral Ecology Environmental Science and Naomi E. Pierce Public Policy OEB 355: Evolutionary Developmental Biology ESPP 90j: Environmental Crises, Climate James Hanken Change and Population Flight James J. McCarthy (and Jennifer Leaning) OEB 362: Research in Molecular Evolution Explores the consequences of population Scott V. Edwards flight due to war, drought and famine in OEB 367: Evolutionary and Ecological which climate change is a contributing factor, Diversity relating to the extent and permanence of Jonathan Losos environmental destruction wrought by these crises, people’s attachment to their homes and ecosystems, the circumstances of departure, the destinations of refuge and the possibilities for return. Graduate Courses Reading and Research

OEB 307: Biomechanics, Physiology and Musculoskeletal Biology Andrew A. Biewener OEB 310: Metazoan Systematics Gonzalo Giribet OEB 320: Biomechanics and Evolution of Vertebrates George V. Lauder OEB 323: Advanced Vertebrate Anatomy Stephanie Pierce Scott Edwards

Annual Report 2015 . 2016 9 COURSES

Freshman Seminar FRSEMR 21R: The Evolutionary Transition from Dinosaurs to Birds: Fossils, Genomes and Behavior Scott V. Edwards Explores the dinosaurian origins of modern birds through exploration of Harvard’s excellent collections of dinosaur fossils, skeletons and specimens of extant birds, and focused readings and discussions. The goal will be to gain a greater appreciation of dinosaur diversity and to better understand the deep origins of modern bird adaptations. FRSEMR 22T: Why We Animals Sing Brian D. Farrell Investigates the sounds and structures of different kinds of acoustic animals— including birds, mammals, frogs and insects—and the different kinds of habitats in which they produce their songs and calls. Life Sciences LIFESCI 2: Evolutionary Human Physiology

and Anatomy MCZ, Harvard University Andrew Biewener, George V. Lauder (and Daniel E. Lieberman) MCZ History Explores human anatomy and physiology from an integrated framework, combining In June 1938, MCZ paleontologist Alfred Sherwood Romer functional, comparative and evolutionary received a letter from a Boston psychiatrist. Was he willing to perspectives on how organisms work. meet to discuss possible employment at the MCZ for her old schoolmate, Tilly Edinger? BIOS S-158: Study Abroad: Biodiversity of the Dominican Republic Edinger was then a vertebrate paleontologist at the Senckenberg Brian D. Farrell Museum in Frankfurt, Germany. Starting in the early 1920s, she Explores the interplay of ecological niches and had documented the structure of cranial endocasts, using these evolutionary diversification in the organisms “fossil brains” to reconstruct the lifestyle of extinct vertebrates. and habitats of a tropical island as a microcosm Her seminal 1929 work, Die fossilen Gehirne (Fossil Brains), of the evolution of biodiversity on Earth. had established her international reputation and the field of paleoneurology. Now, restrictive Nazi “racial laws” threatened both her job and her life. OEB 130: Biology of Fishes Romer’s subsequent actions paved the way for Edinger’s safe departure from Germany and her eventual addition to the MCZ staff. Edinger worked happily at the MCZ for a quarter-century, writing widely on the evolution of the brain and the historical pattern of neural innovations. More than anyone else, she introduced the concepts of behavior, body size and phylogeny to the study of the brains of fossil vertebrates. —Emily A. Bucholtz, Wellesley College George LauderGeorge

10 Museum of Comparative Zoology MCZ Research Making Headlines

Precious Metal In this day and age, it is rare to discover large In all, 33 of the meter-long boas were identified vertebrate species. It is even rarer to have one during three separate expeditions funded by crawl across your face while sleeping. But that the MCZ. The researchers have kept the exact is exactly what happened to expedition leader location of this R. Graham Reynolds during a Putnam-funded snake confidential expedition to the remote Conception Island in order to protect Bank in the Bahamas. It is especially remarkable the species that, that the new boa was found in 2015 in the by using IUCN Bahamas, a region that has been well studied Red List criteria, since the 1860s, and by the MCZ since the they estimate 1880s. It is the first new species of West Indian to be critically boa to be discovered in 73 years. endangered. The home of the silver The boa was christened the Conception Bank boa is protected silver boa (Chilabothrus argentum), in large by its remote part because of its metallic silver scales, but location, lack of also because it was first found in a silver palm human habitation (Coccothrinax argentata). Unlike the other and status as a ground-dwelling boas in the region, the silver

national park. Reynolds R. Graham boa lives in trees, moving agilely among the However, natural limbs and hunting birds. disasters, a population of feral cats The team, which included postdoctoral fellow discovered by the researchers and poaching Anthony Geneva and graduate student Nick for the pet trade are serious concerns. Herrmann, who found the first boa, were Reynolds RG, Puente-Rolón AR, Geneva AJ, Aviles- able to hand-capture six live specimens for Rodriguez KJ, Herrmann NC (2016) Discovery of a study, taking measurements, photos and DNA remarkable new boa from the Conception Island samples prior to their release. Bank, Bahamas. Breviora 549:1-19

Arachnid Sex, Frozen in Time Arachnid genitalia are an excellent way to preserved in a rather unique condition. identify not only gender, but also species, This harvestman, frozen in time in the both ancient and extant. Most arachnids moments before copulation, provides transfer sperm indirectly, without genital the first example of a fully extended contact. Harvestmen (Arachnida: Opiliones), penis and offers valuable insights into an nicknamed daddy longlegs, are unusual extinct species, Halitherses grimaldii. in that they transfer sperm directly via The slender penis, with a spatulate, heart- copulation. The male’s penis is retractable

shaped tip, and the specimen’s large Gonzalo Giribet and spends most of the time inside the body, eyes, suggest that H. grimaldii represents but as in the case of the spider genitalia, a new, extinct family of harvestmen. it yields crucial characteristics for species Gonzalo Giribet, who together with identification. Jason Dunlop described this Cretaceous One of the biggest challenges in reconciling species back in 2005, co-authored the studies of fossil and modern harvestmen is new findings, presented inThe Science the lack of visible male genitalia in fossils. of Nature. However, a fossil harvestman from the Dunlop JA, Selden PA, Giribet G (2016) Penis Cretaceous Burmese amber of Myanmar, morphology in a Burmese amber harvestman.

around 99 million years old, has been found Sci Nat 103:11 Gonzalo Giribet

Annual Report 2015 . 2016 11 RESEARCH

Answers from Anoles in Amber An enduring question in ecology is whether researcher in the Losos lab—and others the structure of ecological communities— began scanning the amber with X-ray micro the group of interacting species living in the computed tomography (micro-CT). The same place—can be stable over millions of surprise was that, in some cases, the intact years. Anole communities of the Caribbean lizard was preserved in the amber, preserving have been a living laboratory for Jonathan a perfect impression of the lizard down to the Losos, but due to a dearth of anole fossils, minute toepad scales. These toepad scales answers to this question have remained allow anoles to cling to surfaces, and the elusive. Undeterred, Losos turned to different types of toepad scales correlate with specimens preserved in amber. the lizards’ ecological niche and help identify them as ground, bush or tree dwellers. Caribbean anole specimens in amber— fossilized tree resin—are not easy to come by. By examining fossilized scales as well Ten years ago, Losos heard of an anole specimen as overall morphology, the researchers for sale by a private collector—only the fourth were able to determine that the lizard known to scientists—for one million dollars. communities living on Hispaniola 20 million Years later, Losos finally obtained access to years ago were much the same as those that the specimen and began to expand his search, exist there today. leading to a collector in Italy and a museum in Lead author Sherratt was joined by co- Germany. Losos managed to obtain 38 amber authors Rosario Castañeda, Jonathan Losos anole specimens from the island of Hispaniola and four former members of the Losos lab. to study, with one caveat—half of them could not leave Europe.

Emma Sherratt Sherratt E, Castañeda MdR, Garwood R, Mahler DL, Sanger TJ, Herrel A, de Queiroz K, Losos JL (2015) Amber At the Natural History Museum in London, fossils demonstrate deep-time stability of Caribbean lizard Emma Sherratt—at that time a postdoctoral communities. Proc Natl Acad Sci USA 112:9961-9966

In the Belly of the Whale Five years ago, then-undergraduate Annabel Peter Girguis and others, the researchers Beichman and former MCZ Hrdy Fellow Joe explore how these microbial communities Roman collected northern allow whales to extract the most nutrition right whale feces in the Bay possible from their crustacean diet, including of Fundy to determine which the chitin-rich exoskeletons, giving them microbes were present. The access to about 10 percent more calories. great whales are carnivores, The ancestors of whales are the same as often feeding on minute those of cows, camels, and other ruminants, shrimp-like animals, and and it is from these ancestors that they their microbiome—the inherited the multichambered foregut microbes that live in their that digests chitin in the way terrestrial guts—in some respects herbivores digest cellulose. resembles that of other carnivores, such as lions and In the future, the researchers hope to sample tigers. DNA sequencing that the microbial community in whales’ stomach Beichman conducted as part chambers and extend the study to toothed © Carl Buell of her senior thesis revealed whales. Aquariums have shown interest in that the microbiome of right whales and the research, in the hope that it may yield other baleen species also shares traits with additional information about how to keep ruminants—plant eaters, like cows. whales healthy in captivity. In a paper in Nature Communications by OEB Sanders JG, Beichman AC, Roman J, Scott JJ, Emerson D, McCarthy JJ, Girguis PR (2015) Baleen whales host graduate student Jon Sanders, Beichman, unique gut microbiome with similarities to both Roman, James McCarthy, OEB professor carnivores and herbivores. Nat Commun 6:8285

12 Museum of Comparative Zoology RESEARCH

A Blooming Partnership Flowering plants—angiosperms—are found pollen clump from a in numerous ecosystems, from tundra to gymnosperm, most tropical rain forests. The estimated 352,000 likely a bennettitalean, species are the result of nearly 130 million attached to its body, years of evolution, and that success is due offering direct evidence of in part to their symbiotic relationship with of a pollination pollinators, especially insects such as bees, relationship between butterflies and long-proboscid flies. However, ancient insect and 105 million years ago during the Early plant species. Cretaceous, gymnosperms (such as pines, The researchers note cypresses and cycads) were still dominant, that if adaptations for and the wind was a more prevalent mode of pollination became pollination than it is today. highly specialized in

New research by Farrell lab NSF postdoctoral certain insect lineages (IGME, AMBERIA UB) project J.A. Peñas, fellow Ricardo Pérez-de la Fuente and as a result of their others has described new species from an interaction with gymnosperms over tens of extinct group of long-proboscid flies (family millions of years, this may have predisposed Zhangsolvidae) encased in Cretaceous these insects to develop similar symbiotic amber from Spain and Myanmar. The amber relationships with the emerging angiosperms, preserved the delicate tubular proboscides, contributing to the evolutionary success of which the scientists were able to examine both flowering plants and their insect partners. in minute detail through micro-CT imaging, Peñalver E, Arillo A, Pérez-de la Fuente R, Riccio ML, and they were found to have adapted for Delclòs X, Barrón E, Grimaldi DA (2015) Long-proboscid siphoning nectar and possibly feeding on flies as pollinators of Cretaceous gymnosperms.Curr pollen. One Spanish specimen even had a Biol 25:1917-1923

Energy for the Run For decades researchers thought that the the IT band changes length during a series of primary function of the iliotibial (IT) band, movements. Finally, they created a computer which runs along the outside of the thigh model to measure how much energy is from hip to knee, was to stabilize the hip stored in running and walking, finding that during walking. However, new research by the human IT band has the capacity to Carolyn Eng examines how the IT band acts store 15 to 20 percent more energy than the as a spring to aid in locomotion, storing and similar structure in chimpanzees. releasing elastic energy to make running and Eng and co-authors Andrew Biewener, walking more efficient. research associate Allison Arnold-Rife and The IT band is made up of fascia, an elastic Daniel Lieberman describe the studies in connective tissue found throughout the the Journal of Experimental Biology and the body. Fascia is a sheath that encloses Journal of Biomechanics. The researchers muscles, connects muscles to bone, and hope to expand their investigation to other groups muscles that serve a similar function. primates adapted for running, and explore The IT band is the largest piece of fascia in how understanding the IT band can help the human body and much more developed treat and avoid injury. than that of chimpanzees. Eng CM, Arnold AS, Biewener AA, Lieberman DE (2015) To investigate the energy storage capacities The human iliotibial band is specialized for elastic energy of the IT band in humans and chimpanzees, storage compared with the chimp fascia lata. J Exp Biol 218:2382-2393 Eng and colleagues first had to precisely Eng CM, Arnold AS, Lieberman DE, Biewener AA (2015) describe which muscles attach to the IT The capacity of the human iliotibial band to store elastic band and where, then measure how much energy during running. J Biomech 48:3341-3348

Annual Report 2015 . 2016 13 Highlights from the Collections Historical MCZ Collection Yields Evolutionary Clues Historical specimens are increasingly valuable as species face extinction. They can be used to compare modern members of a species with those hundreds of years old, revealing changes in everything from diet to environment. They are also proving to be vital when scientists cannot obtain modern tissues for research, as with the threatened tuatara. The tuatara Sphenodon( punctatus) is a lizard-like reptile found in New Zealand. Adult males have no penis, making it a useful species in which to study the evolution of the phallus in terrestrial vertebrates. Researchers have wondered whether the common ancestor of amniotes— reptiles, birds and mammals—had a penis, which was subsequently lost in some species, or if the penis evolved later and independently in different groups. Because of the enormous diversity in adult anatomy, answering this question required the study of genital development in tuatara embryos, but fresh samples are very difficult to obtain for this threatened species. Researchers Thomas Sanger, Marissa Gredler and Martin Cohn instead turned to the MCZ’s collection of Victorian-era tuatara embryos. “This collection goes back to Charles S. Minot (1852– 1914), curator of the Harvard Embryological Collection,” says Curatorial Associate José Rosado. Arthur Dendy, an Hannah Kim Frank English zoologist, collected the tuatara embryos between 1896 and 1897. In 1909 he transferred his collection to Minot, who prepared the specimens. “The collection contributed significantly to the study of embryology,” says Rosado, “only to be lost in obscurity for much of the 20th century, stored in various places at the Harvard Medical School. Their recent rediscovery has made this collection available to add new dimensions to our study of natural history and evolution, as exemplified by Dr. Sanger's recent work.” Sanger photographed 82 of the fragile microscope slides containing thin sections of embryos more than a century old. He and colleagues then digitally removed tissues that were deteriorating and combined the remaining data into one 3D image. The image revealed swellings consistent with the development of paired genitals in the tuatara’s closest living relatives, snakes and lizards, and the beginnings of a single phallus in mammals, turtles, crocodilians and some birds. As the tuatara embryo matures, the swellings fail to develop into a penis, but their existence supports

the theory that the penis evolved just once in MCZ, Harvard University mammals and reptiles and that some animals have lost theirs over time. The findings were published inBiology Letters. The original photos and the digitally restored images used for the research are archived at the MCZ.

14 Museum of Comparative Zoology COLLECTIONS

Innovative Storage for Bird Specimens Jeremiah Trimble

The Ornithology collection has a new display scheme designed specifically for the needs of bird specimens prepared for exhibit. These specimens are often difficult to store, but the new pegboard system allows the birds to be spaced in a way that can be customized for all the different shapes and sizes of specimens, better utilizing the space. “This system allowed us both the room and ability to store a large collection of Ornithology specimens that were rotated off exhibit during a major renovation of the HMNH public space,” says Curatorial Associate Jeremiah Trimble. “By designing this innovative and novel storage solution, we are easily able to accommodate and properly store all of these valuable specimens.” “The specialized but simply designed equipment provides secure storage that is both flexible and efficient for these individually unique preparations of bird specimens,” says Linda S. Ford, MCZ Director of Collections Operations. LEED Recognition for MCZ The MCZ’s Northwest Building facilities recently received LEED platinum certification— the highest rating—from the U.S. Green Building Council. This is the ninth LEED certification and the fourth platinum certification for the building. The 50,000-square- foot build-out of state-of-the-art laboratory facilities and stable, climate-controlled storage accommodates six million zoological specimens from all or part of seven different collections: Invertebrate Paleontology, Ornithology, Vertebrate Paleontology, Malacology, Mammalogy, Entomology and Invertebrate Zoology. Thirteen percent of the materials in the project space are made of recycled content, and regional materials make up 36 percent of the new construction materials. Energy- efficiency measures include an integrated laboratory pressurization control system and occupancy sensors to control lighting and automate temperature settings. LED lighting will use 31 percent less power, and efficient plumbing fixtures will reduce water use by 32 percent. Aja Melissa

Annual Report 2015 . 2016 15 COLLECTIONS

MCZ First Floor Renovations An institution that opened in 1859 would naturally evolve over the years. The latest renovations to the MCZ building’s first floor, completed in fall 2016, were made possible when the staff, collection and prep spaces for Vertebrate Paleontology relocated to the Northwest Building. Photos by Melissa Aja by Melissa Photos

Two outdated, uncomfortable classrooms are now modernized and side-by-side, offering more functionality—MCZ 101 (left) and the Oceanography seminar room (right). These classrooms were updated with state-of-the art technology and mobile chairs and desks. Off a stairwell and impossible to find, the former Oceanography classroom will become a faculty office.

The Agassiz Room (left), a much-used and beloved conference space, was moved from the basement level of the OEB administration offices to a temporary home on the first floor of the MCZ. (It will move again in a later stage of renovations.) The former Agassiz Room was revamped and turned into greatly needed office space for the Harvard Museums of Science and Culture. At right is the new employee lounge.

16 Museum of Comparative Zoology Projects & Initiatives Encyclopedia of Life Learning + Education Group The Encyclopedia of Life eol.org( ) is a global effort to bring together species information in a free, trusted online resource. Content on EOL is provided by hundreds of partners, including the MCZ. The EOL Learning + Education Group (education.eol.org), based at the MCZ, encourages the development of innovative and effective uses of EOL content and tools in educational settings. Species Cards Species Cards (eol.org/info/species_cards) are built from EOL Collections and trait data from TraitBank and other sources. Traits may include species interactions, size categories, colors of organisms, body measurements, plant growth habits, phenology data and more. Learn more about TraitBank at eol.org/info/traitbank, including the breadth of data available and how to contribute or download data. Card decks can be viewed online or PDFs can be downloaded for printing. Species Cards are being used in lessons and activities for students in elementary and middle schools (ages 7–13). eol.org/info/ed_resources Okaloosa SCIENCE Grant EOL is now in the third and final year of the Okaloosa SCIENCE project, supported by a Department of Defense Education Activity grant to improve STEM education through outdoor activities and community partnerships. This year, EOL L+E staff are running professional development workshops for teachers and schoolyard BioBlitzes—species inventories—to bring its biodiversity learning tools and resources into classrooms. education.eol.org/ecosystems/ ecoproj.php?proj_id=4 Broader Impacts for the Kurator Project Harvard University and the University of Illinois Urbana–Champaign were awarded a National Science Foundation grant to develop software tools for natural history collection data digitization, sharing, integration and use. This grant will develop Kurator, an open-source toolkit for providing data quality information for natural history collections. EOL is collaborating with Integrated Digitized Biocollections (idigbio.org) and Advancing Integration of Museums into Undergraduate Programs (aimup.unm.edu) to develop an educational module for undergraduate students focusing on using natural history collections to answer ecological questions and to improve student data literacy. U.S. National Parks Centennial The EOL L+E group has been participating in the U.S. National Park Service and National Geographic Society annual BioBlitz species inventories since 2009. EOL has supported these events through the development of educational activities, EOL Collections and field guides. There were hundreds of BioBlitzes around the country in 2016 to commemorate the centennial of the U.S. National Park Service, and EOL staff participated in events at Gulf Islands National Seashore and the National Mall in Washington, D.C. Amy Lorenz

Annual Report 2015 . 2016 17 PROJECTS & INITIATIVES

William Brewster’s Field Notes The digitization and transcription of ornithologist William Brewster’s diaries (1851–1919), field journals, photographs and correspondence has been an ongoing project at the Ernst Mayr Library. Supported by various grants from the Institute of Museum and Library Services and the Council on Library and Information Resources, the goal of the project is to make all Brewster’s notes and correspondence digitally accessible for study. Currently, the CLIR Field Notebook Project grant is supporting the digitization of 9,821 remaining items in the William Brewster collection. Betsy Meyer, who has been scanning and transcribing Brewster’s notes, began highlighting interesting tidbits from these documents in a series of Ernst Mayr Library blog posts as a way to bring the library’s community into the ongoing project and showcase the value of texts where the scientific and the personal overlap. So far, the blog series has touched on a range of scenes from Brewster’s life, from wild animal encounters to Boston courtrooms in early debates

Kris SnibbeKris over wildlife protection legislation. library.mcz.harvard.edu/blog Laura Bush 21st Century Librarian Award The Ernst Mayr Library was awarded an IMLS grant, Foundations to Actions: Extending Innovations in Digital Libraries in Partnership with National Digital Stewardship Learners. Five residents across the United States will work on different Biodiversity Heritage Library projects. The resident based at the Ernst Mayr Library will develop a mechanism to incorporate transcription files from manuscripts into the BHL infrastructure, eventually allowing searching and indexing of handwritten documents such as field notes and correspondence. Science without Borders Intern Bruno Costelini joined the Ernst Mayr Library for three months to process and organize the papers of Ruth Turner, one of the foremost marine scientists of the 20th century, and delve into the Hassler Archives. He scanned 166 photographs of the Hassler expedition and organized metadata, transcribed James H. Blake’s expedition journal and built a spreadsheet of his roughly 100 watercolors. He explored materials from the Hassler expedition at several libraries across Harvard and plans to write a paper about his work. Costelini presented a poster on the Hassler Archives at the summer intern poster session and actively contributed to the

MCZ, Ernst Mayr Library MCZ, Ernst library blog. library.mcz.harvard.edu/blog

Expanding Access to Biodiversity Literature Awarded in October 2015 to the New York Botanical Garden with partners Ernst Mayr Library and Missouri Botanical Garden, the goal of this grant is to increase the availability of biodiversity literature by seeking out content providers who may need assistance in digitization or deposit and negotiating with copyright holders for more current publications. EML will also work with the Digital Public Library of America to ensure metadata is available for ingest, thus broadening the prospective audience and enhancing accessibility to the literature of biodiversity. Through this grant, the library was able to hire Patrick Randall to negotiate copyright agreements, provide social media outreach for the Ernst Mayr Library and respond to internal and external digitization, thus improving access to the Ernst Mayr Library collections, particularly those that are offsite or fragile.

18 Museum of Comparative Zoology PROJECTS & INITIATIVES

Marine Life On November 21, 2015, the new Marine Life exhibition in the Putnam Family Gallery opened at the Harvard Museum of Natural History. Marine Life is made possible by a generous gift from George Putnam III, AB 1973, JD 1977, MBA 1977, and Kathy Putnam. The centerpiece ofMarine Life is an immersive, floor-to-ceiling recreation of life in New England’s coastal waters, highlighting the diversity and dynamic interplay of local marine communities. As models of glowing jellies, a giant sea turtle and other sea animals swim overhead, visitors explore new research and displays of real fishes, mollusks, crustaceans, corals and other marine organisms. These “wet” specimens— demonstrating the breadth of the collections and the science behind them—were provided by the MCZ Invertebrate Zoology, Malacology and Ichthyology collections. Bird specimen mounts were selected from the Ornithology collection. Aja Melissa A special interactive display introduces the world of jellyfish, and the ocean exploration theater offers a multimedia journey of discovery from the surface to the deep oceans. MCZ faculty-curators James McCarthy and Gonzalo Giribet are featured in a video about the history of oceanography at Harvard, and they are among the four Harvard faculty members that provided the foundation for the exhibition. HMSC exhibit designers, Ichthyology staff and graduate students collaborated on the digital components. Sea Creatures in Glass Sea Creatures in Glass: The Blaschka Marine Animals at Harvard, by Elizabeth R. Brill and Florian Huber, was published to celebrate the restoration and permanent display of these extraordinary glass representations of marine organisms. It is the first publication featuring the MCZ’s exceptional collection of models created by father and son artists Leopold and Rudolf Blaschka in the late nineteenth century. Delicate jellyfish and anemones, octopus, squid and soft-bodied sea creatures—commissioned by universities and museums throughout the world as teaching models—are renowned for their beauty and exacting detail and are still relevant to marine study today. Sea Creatures in Glass features stunning photography of 60 of the most exquisite

David Brown models from this superb collection.

Annual Report 2015 . 2016 19 Awards & Recognition

George V. Lauder was elected a Fellow of the American Physical Society for fundamental contributions to understanding aquatic propulsion in fishes through experimental hydrodynamics. Naomi Pierce received the E. O. Wilson Naturalist Award from the American Society of Naturalists for significant contributions to the knowledge of a particular ecosystem or group of organisms. Mansi Srivastava was named a Searle Foundation Scholar for her potential to make significant contributions to chemical and biological research. Staff Bear Cieri Photography th Hopi Hoekstra Penny Benson marked her 5 -year service Emeritus anniversary at the MCZ. Edward O. Wilson received the Harper Jessica Cundiff and Breda Zimkus became Lee Award for Southern Literature from Co-Chairs of the Best Practices Committee the Alabama Literary Association. The PBS for the Society for the Preservation of special E. O. Wilson: Of Ants and Men was Natural History Collections. released; it won awards in two categories at Linda S. Ford became President of the the Jackson Hole Wildlife Film Festival and Society for the Preservation of Natural has received Emmy nominations for best History Collections. television program, also in two categories. Keleigh Quinn received a Dean’s Distinction Faculty Award from the Harvard Faculty of Arts and Brian D. Farrell was awarded funding from Sciences. the Harvard University Climate Change Postdoctoral Researchers Solutions Fund. Gonzalo Giribet was awarded a 2016 John Simon Guggenheim Fellowship in the field of Organismic Biology and Ecology. Guggenheim fellows are appointed on the basis of impressive achievement in the past and exceptional promise for future accomplishment. Hopi Hoekstra was elected to the U.S. National Academy of Charlotte Jandér Sciences, in recognition of her Charlotte Jandér received a distinguished and continuing research grant from Stiftelsen Extensus, achievements in original Sweden, and a travel award from the research. Jenn Thomson Jenn Smithsonian Tropical Research Institute. Mansi Srivastava 20 Museum of Comparative Zoology AWARDS Awards & Recognition

Patrick Gorring, Alyssa Hernandez and Sang Il Kim received travel grants from the David Rockefeller Center for Latin American Studies. Gorring also received a Huron Mountain Wildlife Foundation Research Grant. Kim also received a Systematics Research Fund grant from the Brent Hawkins Systematics Association and the Graduate Students Linnean Society of Felix Baier was awarded a Howard Hughes London. Medical Institute International Research Fellowship to support three years of his Emily Hager Sang Il Kim PhD research. was awarded an American Society of Mammalogists Caitlin Baker, Julia Cosgrove and Vanessa Grant-in-Aid of research for her work on Knutson received an NSF Graduate Research morphological and behavioral adaptation. Fellowship Program award. Talia Moore received a Robert A. Chapman John H. Boyle, Brianna McHorse, Zachary Memorial Scholarship. Morris and Elizabeth Sefton were awarded the Derek Bok Certificate of Distinction in Shayla Salzman was a recipient of the Grady Teaching. McHorse also received a Sigma L. Webster Award, American Society of Plant Xi Grant-in-Aid of Research and Chapman Taxonomists, for the most outstanding paper Memorial Scholarship for Locomotion published in Systematic Botany or Systematic Research funding. Morris also received a Botany Monographs in the field of plant Wood Student Research Award from the systematics for the years Society of Paleontology. 2014 and 2015. Brent Hawkins and Bruno de Medeiros received an NSF Doctoral Dissertation Improvement Grant. Medeiros also received a Jorge Paulo Lemann Fellowship. Undergraduates Andreé Franco-Vasquez and Silvia Golumbeanu were awarded Harvard College Program for Brianna McHorse Research in Science and James Crall was awarded a research grant Engineering fellowships. from the Winslow Foundation. Bruno de Medeiros Annual Report 2015 . 2016 21 MCZ Grant Recipients Academic Year 2015–2016 Grants in Aid of Undergraduate Research (GUR) These grants support research by Harvard College undergraduates under faculty supervision. Priority is given to projects that utilize MCZ, Harvard University Herbaria (HUH) and Arnold Arboretum (AA) research collections, laboratories and facilities. Support for these grants comes from the MCZ’s Myvanwy M. and George M. Dick Scholarship for Students, HUH and AA.

Recipient Faculty Sponsor/ Project Title Amount Academic Dept. Madeleine V. Ankhelyi George Lauder/ Morphological differences in placoid scales on $2,430 Organismic and different species and body regions of sharks Evolutionary Biology Lorena M. Benitez Stephanie Pierce/ Vertebral morphology of a basal pelycosaur $2,500 Organismic and and its implications for locomotor evolution in Evolutionary Biology synapsids

Elizabeth Benson Elizabeth Elizabeth L. Benson Naomi Pierce/ Testing convergent interactions and community $2,500 Organismic and succession with Australian carnivorous pitcher Evolutionary Biology plants Christopher H. Chen Donald Pfister/ Evolution of hyperparasitism in a system of $1,800 Organismic and Laboulbeniales, bat flies and bats Evolutionary Biology Eamon C. Corbett Scott Edwards/ Thesis research on biogeography and genetics $970 Organismic and of birds of northeastern Brazil in Recife, Evolutionary Biology Pernambuco Andree M. Franco Mansi Srivastava/ Gene expression profile of stem cells in acoels $2,500 Vasquez Organismic and characterized by in situ hybridization Evolutionary Biology Kaitlyn A. Gibson Stephanie Pierce/ Digital vertebral modeling of Chiniquodon and $1,770 Organismic and Ctenorhachis Evolutionary Biology Silvia Golumbeanu Mansi Srivastava/ Piwi-1 presence and cell pluripotency in early $2,500 Organismic and embryonic development of Hofstenia miamia Evolutionary Biology Ann H. Opel Colleen Cavanaugh/ The effect of coral outplant sites on local fish $900 Organismic and communities Evolutionary Biology Christian A. Perez Jonathan Losos/ Habitat partitioning and locomotor patterns in $2,500 Emily Venable Organismic and three species of Costa Rican anoles Evolutionary Biology Elizabeth Benson Elizabeth Silvia Golumbeanu Jones Katrina Lorena Benitez 22 Museum of Comparative Zoology GRANTS Polat DiyarPolat Annie Opel

Recipient Faculty Sponsor/ Project Title Amount Academic Dept. Andres Vega Andres Kristin M. Tsuo Terence Capellini/ Genome-wide studies of genetic variants $2,500 Christian Perez Human Evolutionary affecting body proportion and height Biology Emily M. Venable Richard Wrangham/ Investigation of the observed sexual $2,000 Human Evolutionary dimorphism in chimpanzee wood eating Biology Anthony W. Wohns Terence Capellini/ Why the long tendon? The role of muscle force $2,500 Human Evolutionary on gene expression and morphology in the Biology developing Achilles tendon Total Awards $27,370

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 Laslo Mara Mara Laslo Herpetology Thyroid hormone-independent limb $1,407 development in a direct-developing frog, Eleutherodactylus coqui Patrick Gorring Entomology Determining host plant influence on speciation $2,500 in sky island beetles Total Awards $3,907 Patrick GorringPatrick Patrick Gorring Annual Report 2015 . 2016 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.

Recipient Institutional Project Title Amount Affiliation Consuelo N. Alarcon San Antonio Abad An integrative taxonomic revision of genus $863 Rodriguez National University of Oxyrhopus Cusco Michael G. Branstetter University of Utah The leaf litter ants of Colombia $1,420 David A. Bullis State University of Taxonomy and shell morphology of a nearly $1,424 New York extinct Pacific endemic land snail group: The endodontoids of Drs. Yoshio Kondo Lilian Macedo (BPBM: 1934–1980) and Alan Solem (FMNH: 1957–1990) Sarah Z. Gibson University of Kansas Examining the evolutionary relationships $1,500 of fishes of the order Redfieldiiformes (Actinopterygii: Palaeoniscimorpha) from the Early Mesozoic María Rebecca Granja- Universidad Taxonomy of the Ophiuroidea (Echinodermata) $1,500 Fernández Autónoma from the tropical eastern Pacific Metropolitana Maike Hernández Institute of Ecology Review family Cepolidae of the Caribbean, with $1,500 Renata Stopiglia Renata Quinta and Systematics, emphasis in the Jeanneretia genus at MCZ Havana Paul J. Johnson South Dakota State Otto Schwarz primary types of Andean and $1,420 University Amazonian Elateridae Gustavo S. Libardi Instituto de Diversidad Taxonomic revision of the genus Necromys $1,500 y Evolución Austral, Ameghino, 1889 (Rodentia: Cricetidae) CONICET Lilian Cristina Macedo Federal University of Revision of the Physaloptera spp. Rudolphi, 1819 $1,500 Para from South American reptiles Shirley Daniella Universidad Nacional Taxonomic review and geographical $1,500 Martínez-Torres de Colombia distribution of the neotropical millipede genus Psammodesmus Cook, 1896 (Diplopoda: Polydesmida: Platyrhacidae) Robert K. McAfee Ohio Northern Expanded systematics of the Caribbean $1,260

Italo Salvatore de Castro Pecci-Maddalena de Castro Salvatore Italo University fossil sloths Robert McAfee Robert HernándezMaike Oleksander Varga 24 Museum of Comparative Zoology GRANTS Yongying Ruan (2) Ruan Yongying Recipient Institutional Project Title Amount Affiliation Katherine Nazario University of Revision of the cicada genus Tettigades $1,500 Connecticut (Auchenorrhyncha, Cicadidae) Brittany E. Owens Louisiana State Monography of the New Zealand Pselaphini $1,500 University (Coleoptera: Staphylinidae: Pselaphinae) Italo Salvatore de Castro Federal University of Taxonomic revision of Mycotretus Lacordaire $1,500 Pecci-Maddalena Viçosa (Coleoptera: Erotylidae: Tritomini): The type María Rebecca Granja-Fernández specimens from MNHN Abel Perez-Gonzalez Museo Argentino de Taxonomy and fine morphological $1,500 Ciencias Naturales, characterization of poorly known key species of CONICET Malagasy harvestmen (Opiliones: Laniatores) Gareth S. Powell Purdue University Revision of Nearctic Carpophilus Stephens $1,080 Matthew M. Prebus University of The ant genusTemnothorax Mayr 1861: $1,330 California, Davis Redefinition, subgeneric classification and Hispañola species revision ndez

Yongying Ruan Chinese Academy of Revision of Oriental Chaetocnema (Coleoptera, $1,500 María Rebecca Granja- á Fern Sciences Chrysomelidae) species David Salazar- Ohio State University Clarifying the systematics of the “ultimate $1,500 Valenzuela pitvipers” (Bothrops asper species complex) a medically important group of Neotropical snakes Renata Stopiglia Universidad de São Taxonomic revision of the species of the $1,500 Paulo subtribe Synallaxina (Synallaxini, Furnariidae, Passeriformes, Aves) Oleksandr Varga Schmalhausen An investigation of Ichneumonidae types (Insecta, $1,100 Institute of Zoology Hymenoptera, Ichneumonidae) deposited in the Bavarian State Collection of Zoology Hessarghatta Murthy University of Examination and study of Miridae types at the $1,500 Yeshwanth Agricultural Natural History Museum Sciences, Bangalore

Total Awards $30,897 Camila Baptista Folly Italo Salvatore de Castro Pecci-Maddalena Daniela Martinez Torres Martinez Daniela Renata Stopiglia Renata Abel Perez-Gonzalez Annual Report 2015 . 2016 25 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 Nicole L. Bedford Mammalogy Quantifying burrow usage patterns in a near- $7,875 threatened beach mouse species (Peromyscus polionotus leucocephalus) Ligia R. Benavides Silva Invertebrate Zoology Discovering the soil arthropods of the “path to $3,010 the panther” Leonora S. Bittleston Entomology The third convergent host: Insect communities $8,060

Claire DufourClaire from Cephalotus follicularis in Australia Tauana Cunha Invertebrate Zoology Resolving the deep relationships of the $8,475 Gastropoda Claire Marie-Soleil Herpetology Coexistence mechanisms between the native $9,432 Dufour species Anolis oculatus and a new intruder Anolis cristatellus in Dominica Rosa M. Fernández Invertebrate Zoology Exploring terra incognita: Soil fauna of New $12,876 Caledonia Kadeem J. Gilbert Entomology Phylogeography of Nepenthes-associated $9,198 dipterans of the Philippines Emily Hager Mammalogy Behavioral and morphological change across a $11,348 transect in habitat type Oriol Lapiedra Gonzalez Herpetology Does individual variation in behavior determine $10,154 Tauana Cunha Tauana natural selection in new environments? Gerard Talavera Entomology Phylogeny of the most cosmopolitan animal $8,000 migrator: the butterflyVanessa cardui Total Awards $88,428 Ligia R. Benavides Silva Nate Dappen / DaysNate Edge Productions Tauana Cunha Tauana Vanessa Knutson Vanessa Tauana Cunha Oriol Lapiedra Gonzalez 26 Museum of Comparative Zoology MCZ Publications in Calendar Year 2015

• Andrade SCS, Novo M, Kawauchi GY, Worsaae • Corbett-Detig R, Jacobs-Palmer E, Hartl DL, K, Pleijel F, Giribet G, Rouse GW (2015) Articulating Hoekstra HE (2015) Direct gamete sequencing “archiannelids”: Phylogenomics and annelid reveals no evidence for segregation distortion in relationships, with emphasis on meiofaunal taxa. house mouse hybrids. PLoS One 10:e0131933 Mol Biol Evol 32:2860-2875 • Crall JD, Gravish N, Mountcastle AM, Combes SA • André T, Specht C, Salzman S, Palma-Silva C, (2015) BEEtag: A low-cost, image-based tracking Wendt T (2015) Evolution of species diversity in system for the study of animal behavior and the genus Chamaecostus (Costaceae): Molecular locomotion. PLoS ONE 10:e0136487–13 phylogenetics and morphometric approaches. Phytotaxa 204:265-276 • Crall JD, Ravi S, Mountcastle AM, Combes SA (2015) Bumblebee flight performance in cluttered • Badenhorst D, Hillier LW, Literman R, Montiel EE, environments: Effects of obstacle orientation, body Radhakrishnan S, Shen Y, Minx P, Janes DE, Warren size and acceleration. J Exp Biol 218:2728-2737 WC, Edwards SV, Valenzuela N (2015) Physical mapping and refinement of the painted turtle • Crompton AW, Musinsky C, Owerkowicz T (2015) The evolution of the mammalian nose. InGreat Cover story “Dual embryonic origin genome (Chrysemys picta) inform amniote genome and patterning of the pharyngeal evolution and challenge turtle-bird chromosomal Transformations in Vertebrate Evolution (Dial K, Shubin NH, Brainerd EL, eds) University of Chicago Press: Chicago skeleton in the axolotl (Ambystoma conservation. Genome Biol Evol 7:2038-2050 mexicanum)” is authored by Elizabeth Sefton, Nadine Piekarski • Balenger SL, Bonneaud C, Sefick SA,Edwards • Cuff AR, Randau M, Head J, Hutchinson JR, and James Hanken. SV, Hill, GE (2015) Plumage color and pathogen- Pierce SE, Goswami A (2015) Big cat, small cat: induced gene expression in a wild bird. Behav Ecol Reconstructing body size evolution in living and 26:1100-1110 extinct Felidae. J Evol Biol 28:1516-1525 • Bedford NL, Hoekstra HE (2015) Peromyscus • Dierickx EG, Shultz AJ, Sato F, Hiraoka T, Edwards mice as a model for studying natural variation. eLife SV (2015) Morphological and genomic comparisons 4:eO6813 of Hawaiian and Japanese black-footed albatrosses Phoebastria nigripes using double digest RADseq: • Bhullar BAS, Morris ZS, Sefton EM, Tok A, Tokita Implications for conservation. Evol Appl 8:662-678 M, Namkoong B, Camacho J, Burnham DA, and Abzhanov A (2015) A molecular mechanism for • Dinca V, Backström N, Dapporto L, Friberg M, the origin of a key evolutionary innovation, the bird García-Barros E, Hebert PDN, Hernández-Roldán beak and palate, revealed by an integrative approach J, Hornett E, Lukhtanov V, Marec F, Montagud S, to major transitions in vertebrate history. Evolution Munguira ML, Olofsson M, Sichova J, Talavera G, 69: 1665-1677 Vicente-Arranz V, Vila R, Wiklund C (2015) DNA barcodes highlight unique research models in • Biewener AA (2015) Going from small to large: European butterflies.Genome 58:391 VOLUME 218 (8) APRIL 2015 Mechanical implications of body size diversity in terrestrial mammals. In Great Transformations in • Dinca V, Montagud S, Talavera G, Hernández- Research by Glenna Clifton, Vertebrate Evolution (Dial K, Shubin NH, Brainerd EL, Roldán J, Munguira M, García-Barros E, Hebert P, Andrew Biewener and eds) University of Chicago Press: Chicago Vila R (2015) DNA barcode reference for Iberian colleague, “Western and Clark's butterflies enables a continental-scale preview of grebes use novel strategies for • Bittleston LS, Baker CCM, Strominger LB, Pringle potential cryptic diversity. Sci Rep 5:12395 running on water,” was featured A, Pierce NE (2015) Metabarcoding as a tool for on the cover. investigating arthropod diversity in Nepenthes • Dreszer TB, Rađa T, Giribet G (2015) pitcher plants. Austral Ecol 41:120-132 Cyphophthalmus solentiensis sp. nov. (Cyphophthalmi, Sironidae) a new endogean • Boyle JH, Kaliszewska ZA, Espeland M, Suderman mite harvestman species from Croatia, with an TR, Fleming J, Heath A, Pierce NE (2015) Phylogeny application of confocal laser microscopy to illustrate of the Aphnaeinae: Myrmecophilous African genitalia in Opiliones. Breviora 543:1-15 butterflies with carnivorous and herbivorous life histories. Syst Entomol 40:169-182 • Dunn DW, Jandér KC, Lamas AG, Pereira RAS (2015) Mortal combat and competition for • Clifton GT, Hedrick TL, Biewener AA (2015) oviposition sites in female pollinating fig wasps. Western and Clark's grebes use novel strategies for Behav Ecol 26:262-268 running on water. J Exp Biol 218:1235-1243 • Economo EP, Sarnat EM, Janda M, Clouse R, Klimov • Clouse RM, Janda M, Blanchard B, Sharma P, PB, Fischer G, Blanchard BD, Ramirez LN, Andersen Hoffmann BD, Andersen AN, Czekanski-Moir JE, AN, Berman M, Guénard B, Lucky A, Rabeling C, Krushelnycky P, Rabeling C, Wilson EO, Economo Wilson EO, Knowles LL (2015) Breaking out of EP, Sarnat EM, General DM, Alpert GD, Wheeler WC biogeographical modules: Range expansion and (2015) Molecular phylogeny of Indo-Pacific carpenter taxon cycles in the hyperdiverse ant genus Pheidole. ants (Hymenoptera: Formicidae, Camponotus) reveals J Biogeogr 42:2289-2301 Four chapters were contributed waves of dispersal and colonization from diverse by Andrew Biewener, George , and source areas. Cladistics 31:424-437 • Edgecombe GD, Vahtera V, Giribet G, Kaunisto Lauder A.W. Crompton P (2015) Species limits and phylogeography of Catherine Musinsky.

Annual Report 2015 . 2016 27 MCZ PUBLICATIONS

Newportia (Scolopendromorpha) and essential sperm–egg binding partners. R Soc Miletinae (Lepidoptera: Lycaenidae). Evolution implications for widespread morphospecies. Open Sci 2:150296 69:571-588 ZooKeys 510:65-77 • Groh S, Giribet G (2015) Polyphyly of • Kambic RE, Roberts TJ, Gatesy SM (2015) • Edwards SV, Shultz A, Campbell-Staton Caddoidea, reinstatement of the family Guineafowl with a twist: Asymmetric limb SC (2015) Next-generation sequencing and Acropsopilionidae in Dyspnoi, and a revised control in steady bipedal locomotion. J Exp the expanding domain of phylogeography. classification system of Palpatores (Arachnida, Biol 218:3836-3844 Folia Zool 64:187-206 Opiliones). Cladistics 31:277-290 • Kim SI, Farrell BD (2015) Phylogeny of • Eng CM, Arnold AS, Biewener AA, • Hanken J (2015) Is heterochrony still an world stag beetles (Coleoptera: Lucanidae) Lieberman DE (2015) The human iliotibial effective paradigm for contemporary studies reveals a Gondwanan origin of Darwin’s stag band is specialized for elastic energy storage of evo-devo? In Conceptual Change in Biology: beetle. Mol Phylogenet Evol 86:35-48 compared with the chimp fascia lata. J Exp Scientific and Philosophical Perspectives on Biol 218:2382-2393 Evolution and Development (Love AC, ed) • Kvist S, Chernyshev AV, Giribet G (2015) Boston Studies in the Philosophy and History of Phylogeny of Nemertea with special interest • Eng CM, Arnold AS, Lieberman DE, Science, Vol. 307. Springer-Verlag: Berlin in the placement of diversity from Far East Biewener AA (2015) The capacity of the Russia and northeast Asia. Hydrobiologia human iliotibial band to store elastic energy • Harrison AS, Revell LJ, Losos JB (2015) 760:105-119 during running. J Biomech 48:3341-3348 Correlated evolution of microhabitat, morphology, and behavior in West Indian • Lauder GV (2015) Fish locomotion: Recent • Espeland MM, Hall JPW, DeVries PJ, Lees Anolis lizards: A test of the habitat matrix advances and new directions. Ann Rev Mar DC, Cornwall M, Hsu Y-F, Wu L-W, Campbell model. Behaviour 152:1187–1207 Sci 7:521-545 D, Talavera G, Vila R, Salzman S, Ruehr S, Lohman DJ, Pierce NE (2015) Ancient • Herrera-Bachiller A, Kvist S, Giribet G, Junoy • Lauder GV (2015) Flexible fins and fin rays Neotropical origin and recent recolonisation: J (2015) Redescription of Micrura dellechiajei as key transformations in ray-finned fishes. In Phylogeny, biogeography and diversification of (Hubrecht, 1879) (Nemertea, Pilidiophora, Great Transformations in Vertebrate Evolution the Riodinidae (Lepidoptera: Papilionoidea). Lineidae), a rare Mediterranean species. J Mar (Dial KP, Shubin N, Brainerd E, eds) University Mol Phylogenet Evol 93:296-306 Biol Assoc UK 95:1091-1100 of Chicago Press: Chicago • Evans AM, McKenna DD, Bellamy CL, • Hojo MK, Pierce NE, Tsuji K (2015) Lycaenid • Lauder GV, DiSanto V (2015) Swimming Farrell BD (2015) Large-scale molecular caterpillar secretions manipulate attendant mechanics and energetics of elasmobranch phylogeny of metallic wood-boring beetles ant behavior. Curr Biol 25:2260-2264 fishes. InPhysiology Vol. 34A, Physiology of (Coleoptera: Buprestoidea) provides new Elasmobranch Fishes: Structure and Interaction insights into relationships and reveals • Jandér KC (2015) Indirect mutualism: Ants with Environment (Shadwick RE, Farrell AP, multiple evolutionary origins of the larval protect fig seeds and pollen dispersers from Brauner CJ, eds) Academic Press: New York parasites. Ecol Entomol 40:500-510 leaf-mining habit. Syst Entomol 40:385-400 • Lauder GV, Tangorra JL (2015) Fish • Feilich K, Lauder GV (2015) Passive • Jennings DH, Evans B, Hanken J (2015) locomotion: Biology and robotics of body mechanical models of fish caudal fins: Effects Development of neuroendocrine and fin-based movements. InRobot Fish: Bio- of shape and stiffness on self-propulsion. components of the thyroid axis in the direct- inspired Fishlike Underwater Robots (Du R, Li Bioinspir Biomim 10:036002 developing frog Eleutherodactylus coqui: Z, Youcef-Toumi K, Valdivia Y Alvarado P, eds) formation of the median eminence and onset Springer-Verlag: Berlin • Fernández R, Giribet G (2015) Unnoticed of pituitary TSH production. Gen Comp in the tropics: phylogenomic resolution of Endocrinol 214:62-67 • Laumer CE, Bekkouche N, Kerbl A, Goetz the poorly known arachnid order Ricinulei F, Neves RC, Sørensen MV, Kristensen RM, (Arachnida). R Soc Open Sci 2:150065 • Johnson CH, Woollacott RM (2015) Hejnol A, Dunn, CW, Giribet G, Worsaae Analyses with newly developed microsatellite K (2015) Spiralian phylogeny informs the • Fernández R, Lemer S, McIntyre E, Giribet markers elucidate the spread dynamics of evolution of microscopic lineages. Curr Biol G (2015) Comparative phylogeography Tricellaria inopinata d'Hondt and Occhipinti- 25:2000-2006 and population genetic structure of Ambrogi, 1985—a recently established three widespread mollusc species in the bryozoan along the New England seashore. • Laumer CE, Hejnol A, Giribet G Mediterranean and near Atlantic. Mar Ecol Aquat Invasions 10:135-145 (2015) Nuclear genomic signals of the 36:701-715 "microturbellarian" roots of platyhelminth • Jones KE (2015) Evolutionary allometry of evolutionary innovation. eLife 4:e05503 • Giribet G (2015) Morphology should lumbar shape in Felidae and Bovidae. Biol J not be forgotten in the era of genomics—a Linn Soc 116:721-740 • Laurindo da Silva F, Trivinho-Strixino phylogenetic perspective. Zool Anz 256:96-103 S, Silva Neubern de Oliveira C (2015) • Jones KE, Goswami A, Smaers J (2015) Chironomidae types in the reference • Giribet G, Sharma PP (2015) Evolutionary Impact of the terrestrial-aquatic transition collection of the Laboratory of Ecology biology of harvestmen (Arachnida, on disparity and rates of evolution in the of Aquatic Insects, São Carlos, Brazil. Opiliones). Annu Rev Entomol 60:157-175 carnivoran skull. BMC Evol Biol 15:8 Chironomus 28:20-29 • González VL, Andrade SCS, Bieler R, Collins • Jones KE (2015) Evolutionary allometry • Laurindo da Silva F, Wiedenbrug S, Farrell TM, Dunn CW, Mikkelsen PM, Taylor JD, of the thoracolumbar centra in felids and BD (2015) A preliminary survey of the non- Giribet G (2015) A phylogenetic backbone bovids. J Morphol 267:818-831 biting midges (Diptera: Chironomidae) of the for Bivalvia: An RNA-seq approach. Proc R Soc Dominican Republic. Chironomus 28:12-19 Lond B Biol Sci 282:20142332 • Kaliszewska ZA, Lohman DL, Sommer K, Adelson GS, Rand DB, Mathew J, • Leal M, Losos JB (2015) A naturalist’s insight • Grayson P (2015) Izumo1 and Juno: the Pierce NE (2015) When caterpillars attack: into the evolution of signal redundancy. Am evolutionary origins and coevolution of Biogeography and life history evolution of the Nat 186:ii-iv

28 Museum of Comparative Zoology MCZ PUBLICATIONS

• Lemer S, Kawauchi GY, Andrade SCS, chromosomes in an evolutionarily ancient • Ros IG, Badger MA, Pierson AN, Bassman González VL, Boyle MJ, Giribet G (2015) Re- reptile, the tuatara Sphenodon punctatus. G3 LC, Biewener AA (2015) Pigeons produce evaluating the phylogeny of Sipuncula through (Bethesda) 5:1439-51 aerodynamic torques through changes in transcriptomics. Mol Phylogenet Evol 83:174-183 wing trajectory during low speed aerial turns. • Molnar JL, Pierce SE, Bhullar BAS, Turner J Exp Biol 218:480-490 • Lemer S, Saulnier D, Gueguen Y, Planes S AH, Hutchinson JR (2015) Morphological (2015) Identification of genes associated with and functional changes in the vertebral • Rykken JJ, Farrell BD (2015) Developing a shell color in the black-lipped pearl oyster, column with increasing aquatic adaptation in pollinator monitoring program for Boston Pinctada magaritifera. BMC Genomics 16:568 crocodylomorphs. R Soc Open Sci 2:150439 Harbor Islands National Recreation Area: A pilot study. Natural Resource Report NPS/BOHA/NRR • Liu L, Xi Z, Wu S, Davis CC, Edwards SV (2015) • Moore TY, Biewener AA (2015) Outrun or 950. National Park Service: Fort Collins, CO Estimating phylogenetic trees from genome- outmaneuver: How ecological context can scale data. Ann N Y Acad Sci 1360:36-53 make biomechanical studies more broadly • Salzman S, Driscoll HE, Renner T, André T, relevant. Integr Comp Biol 55:1188-1197 Shen S, Specht C (2015) Spiraling into history: • Lowe CB, Clarke JA, Baker AJ, Haussler D, A molecular phylogeny and investigation of Edwards SV (2015) Feather development genes • Moore TY, Organ CL, Edwards SV, biogeographic origins and floral evolution for and associated regulatory innovation predate Biewener AA, Tabin CJ, Jenkins Jr FA, Cooper the genus Costus. Syst Bot 40:104-115 the origin of Dinosauria. Mol Biol Evol 32:23-28 KL (2015) Multiple phylogenetically distinct events shaped the evolution of limb skeletal • Sanders JG, Beichman AC, Roman J, Scott • Lucas KN, Thornycroft PJM, Gemmell BJ, morphologies associated with bipedalism in JJ, Emerson D, McCarthy JJ, Girguis PR (2015) Colin SP, Costello JH, Lauder GV (2015) Effects the jerboas. Curr Biol 25:2785-2794 Baleen whales host unique gut microbiome of non-uniform stiffness on the swimming with similarities to both carnivores and performance of a passively-flexing, fish-like foil • Novo M, Fernández R, Fernández Marchán herbivores. Nat Commun 6:8285 model. Bioinspir Biomim 10:056019 D, Trigo D, Díaz Cosín DJ, Giribet G (2015) Unearthing the historical biogeography • Scarpetta S, Gray L, Nieto-Montes de Oca A, • McCullough EL, Ledger KJ, Moore TY (2015) of Mediterranean earthworms (Annelida: Castañeda MdR, Herrel A, Losos JB, Luna-Reyes Variation in cross-sectional horn shape within Hormogastridae). J Biogeogr 42:751-762 R, Jiménez Lang N, Poe S (2015) Morphology and among rhinoceros beetle species. Biol J and ecology of the Mexican cave anole Anolis Linn Soc 115:810-817 • Owerkowicz T, Musinsky C, Middleton KM, alvarezdeltoroi. Mesoam Herpetol 2:260-268 Crompton AW (2015) Respiratory turbinates • McKenna DD, Farrell BD, Caterino MS, and the evolution of endothermy in mammals • Sefton EM, Piekarski N, Hanken J (2015) Farnum CW, Hawks DC, Maddison DR, Seago and birds. In Great Transformations in Vertebrate Dual embryonic origin and patterning of the AE, Short AEZ, Newton AF, Thayer MK (2015) Evolution (Dial K, Shubin NH, Brainerd EL, eds) pharyngeal skeleton in the axolotl (Ambystoma Phylogeny and evolution of Staphyliniformia University of Chicago Press: Chicago mexicanum). Evol Dev 17:175-184 and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous • Öztürk B, Recevik M, Geyran K (2015) New • Shapiro LR, Scully ED, Roberts D, Straub beetles. Syst Entomol 40:35-60 alien molluscs in the Mediterranean Sea. Cah TJ, Geib SM, Park J, Stephenson AG, Rojas Biol Mar 56:205-212 ES, Liu Q, Beattie G, Gleason M, De Moraes • McKenna DD, Wild AL, Kanda K, Bellamy CM, Mescher MC, Fleischer SG, Kolter R, CL, Beutel RG, Caterino MS, Farnum CW, • Peñalver E, Arillo A, Pérez-de la Fuente R, Riccio Pierce NE, Zhaxybayeva O (2015) Draft Hawks DC, Ivie MA, Jameson ML, Leschen ML, Delclòs X, Barrón E, Grimaldi DA (2015) genome sequence of Erwinia tracheiphila, an RAB, Marvaldi AE, McHugh JV, Newton Long-proboscid flies as pollinators of Cretaceous economically important bacterial pathogen AF, Robertson JA, Thayer MK, Whiting MF, gymnosperms. Curr Biol 25:1917–1923 of cucurbits. Genome Announc 3:e00482-15 Lawrence JF, Ślipiński A, Maddison DR, Farrell BD (2015) The beetle tree of life reveals that • Quinn DB, Lauder GV, Smits AJ (2015) • Sherratt E, Castañeda MdR, Garwood R, Coleoptera survived end-Permian mass Maximizing the efficiency of a flexible Mahler DL, Sanger TJ, Herrel A, de Queiroz K, extinction to diversify during the Cretaceous propulsor using experimental optimization. Losos JL (2015) Amber fossils demonstrate deep- terrestrial revolution. Syst Entomol 40:835-880 J Fluid Mech 767:430-448 time stability of Caribbean lizard communities. Proc Natl Acad Sci USA 112:9961-9966 • McPhillips T, Song T, Kolisnik T, Aulenbach • Ravi S, Crall JD, McNeilly L, Gagliardi SF, S, Belhajjame K, Bocinsky RK, Cao Y, Cheney Biewener AA, Combes SA (2015) Hummingbird • Talavera G, Kaminski LA, Freitas AVL, Vila J, Chirigati F, Dey S, Freire J, Jones C, Hanken flight stability and control in freestream R (2015) One note samba: Biogeographical J, Kintigh KW, Kohler TA, Koop D, Macklin JA, turbulent winds. J Exp Biol 218:1444-1452 history of the relict Brazilian butterflyElkalyce Missier P, Schildhauer M, Schwalm C, Wei Y, • Reid CR, Lutz MJ, Powell S, Kao AB, Couzin cogina. J Biogeogr 43:727-737 Bieda M, Ludäscher B (2015) YesWorkflow: ID (2015) Army ants dynamically adjust living A user-oriented, language-independent tool • Wen L, Weaver jC, Thornycroft PJM, bridges in response to a cost-benefit trade-off. Lauder GV (2015) Hydrodynamic function for recovering workflow information from Proc Natl Acad Sci USA 112:15113-15118 scripts. IJDC 10:298-313 of biomimetic shark skin: Effect of denticle • Riesgo A, Maldonado M, López-Legentil S, pattern and spacing. Bioinspir Biomim 10:1-13 • Meegaskumbura M, Senevirathne G, Biju Giribet G (2015) A proposal for the evolution SD, Garg S, Meegaskumbura S, Pethiyagoda • Williams CD, Biewener AA (2015). Pigeons of cathepsin and silicatein in sponges. J Mol trade efficiency for stability in response to R, Hanken J, Schneider CJ (2015) Patterns of Evol 80:278-291 reproductive-mode evolution in Old World level of challenge during confined flight.Proc tree frogs (Anura, Rhacophoridae). Zool Scr • Roček Z, Boistel R, Lenoir N, Mazurier A, Nat Acad Sci USA112:3392-3396 44:509-522 Pierce SE, Rage JC, Smirnov SV, Schwermann • Witt WC, Wen L, Lauder GV (2015) AH, Valentin X, Venczel M, Wuttke M, Hydrodynamics of C-start escape responses • Miller HC, O’Meally D, Ezaz T, Amemiya C, Zikmund T (2015) Frontoparietal bone in Marshall-Graves JA, Edwards S (2015) Major of fish as studied with simple physical models. extinct Palaeobatrachidae (Anura): Its variation Integ Comp Biol 55:728-739 histocompatibility complex genes map to two and taxonomic value. Anat Rec 298:1848-1863

Annual Report 2015 . 2016 29 Financial Data

These charts describe the income and expenses of the Museum of Comparative Zoology in fiscal year 2016.

Endowment income funds much of the Museum’s activities, (Overhead Charged). Capital Projects includes deployment including acquisition and maintenance of collections, of collections to the newly constructed space in the faculty and staff salaries, capital projects, facilities Northwest Building. Building expenses such as maintenance, renovation and maintenance. Included in Endowment facility improvements and utilities are captured in the income is the annual distribution, revenue generated from Space & Occupancy category. Operating Expenses assets purchased through endowments and endowed funds consist of equipment purchases, supplies, and consultant decapitalized per donor request. Transfers include Harvard and conference fees, as well as annual subventions to the University-funded faculty research, financial support for Department of Organismic and Evolutionary Biology (OEB) the Ernst Mayr Library and other Harvard-funded projects. for administrative services. Support for MCZ-affiliated Other Income comprises miscellaneous income from graduate students in OEB is included in Scholarships, publication subscriptions, royalties, sales and fees, and cost Awards & Travel. Institutional Expenses are support for recovery from other MCZ-sponsored activities. Overhead is other University activities outside the MCZ, including FAS funding paid from MCZ-based sponsored projects to cover and University initiatives and general operating support to facilities and administrative costs for those projects. It is the Harvard Museum of Natural History. shown as both income (Overhead Earned) and expenses

Income Expenses & Non-Operating Funds

Endowment 74% Salaries & Fringe Benefits 45%

Capitalized Balances < 1% Capital Projects 1% Scholarships, Awards & Gifts Travel 4% 1% Operating Expenses Other Federal Overhead 22% Income Sponsored Charged 2% Transfers Nonfederal Overhead Revenue (Sponsored) Space & Institutional 3% Sponsored Earned 4% 12% 5% Occupancy 10% Expenses 13% Revenue 4%

Income Expenses Endowment $15,448,974 Salaries & Fringe Benefits $9,082,957 Federal Sponsored Revenue $2,606,436 Operating Expenses * $4,497,959 Overhead Earned $912,655 Institutional Expenses * $2,732,858 Nonfederal Sponsored Revenue $830,192 Space & Occupancy $2,130,409 Transfers $582,441 Overhead Charged (Sponsored) $912,636 Other Income $307,645 Scholarships, Awards & Travel $770,104 Gifts $265,646 Capital Projects $158,494 Total $20,953,989 Capitalized Balances $21,252 Total $20,306,669

* These two expense categories are calculated differently than in past years.

30 Museum of Comparative Zoology MCZ PERSONNEL

Faculty-Curators Postdoctoral Fellows, Research Zachary Lewis Mark Cornwall Herpetology, Hanken Lab Entomology, N. Pierce Lab Andrew A. Biewener Associates & Visiting Scholars Charles P. Lyman Professor of Biology; Allison Arnold-Rife Michael Logan Julia Cosgrove Director, Concord Field Station Concord Field Station, Biewener Lab Herpetology, Losos Lab Invertebrate Zoology, Giribet Lab Scott V. Edwards Ligia Benavides Silva David Lubertazzi James Crall Professor of Organismic & Evolutionary Invertebrate Zoology, Giribet Lab Global Ant Project, Wilson Lab Entomology, N. Pierce Lab Biology; Alexander Agassiz Professor of Ricardo Mallarino Tauana Cunha Zoology; Curator of Ornithology Andrés Bendesky Mammalogy, Hoekstra Lab Mammalogy, Hoekstra Lab Invertebrate Zoology, Giribet Lab Brian D. Farrell Ricardo Pérez-de la Fuente Blake Dickson Professor of Biology; Curator of Partha Bhagavatula Entomology, Farrell Lab Vertebrate Zoology, S. Pierce Lab Entomology; Director, David Rockefeller Concord Field Station, Biewener Lab Center for Latin American Studies Alison Cloutier Dennis Persson Amanda Evans Invertebrate Zoology, Giribet Lab Entomology, Farrell Lab Gonzalo Giribet Ornithology, Edwards Lab Professor of Organismic & Evolutionary David Combosch Sebastian Pohl Kara Feilich Biology; Alexander Agassiz Professor of Invertebrate Zoology, Giribet Lab Entomology, N. Pierce Lab Ichthyology, Lauder Lab Zoology; Curator of Invertebrate Zoology Valentina Di Santo Robert Graham Reynolds Jacob Gable James Hanken Ichthyology, Lauder Lab Herpetology, Losos Lab Mammalogy, Hoekstra Lab Professor of Biology; Alexander Agassiz Colin Donihue Lorenzo Ricci Kadeem Gilbert Professor of Zoology; Curator of Herpetology, Losos Lab Invertebrate Zoology, Srivastava Lab Entomology, N. Pierce Lab Herpetology; Director, MCZ Claire Dufour Ivo Ros Patrick Gorring Hopi E. Hoekstra Herpetology, Losos Lab Concord Field Station, Biewener Lab Entomology, Farrell Lab Professor of Organismic & Evolutionary Biology; Professor of Molecular & Pierre-Henri Fabre Seyed Mohammad Saadat Philip Grayson Cellular Biology; Alexander Agassiz Herpetology, Losos Lab Ichthyology, Lauder Lab Ornithology, Edwards Lab Professor of Zoology; Curator of Stacy Farina Tim Sackton Michael Brent Hawkins Mammalogy; Howard Hughes Medical Ichthyology, Lauder Lab Ornithology, Edwards Lab Herpetology, Hanken Lab Institute Investigator; Harvard College Professor Rosa Fernández García Martin Schwentner Ambika Kamath Invertebrate Zoology, Giribet Lab Invertebrate Zoology, Giribet Lab Herpetology, Losos Lab George V. Lauder Professor of Biology; Henry Bryant Heidi Fisher Elizabeth Sefton Mara Laslo Bigelow Professor of Ichthyology; Curator Mammalogy, Hoekstra Lab Herpetology, Hanken Lab Herpetology, Hanken Lab of Ichthyology Hunter Fraser Lori Shapiro Kelsey Lucas Jonathan B. Losos Mammalogy, Hoekstra Lab Entomology, N. Pierce Lab Ichthyology, Lauder Lab Professor of Organismic & Evolutionary Anthony Geneva Yung Wa (Simon) Sin Brianna McHorse Biology; Monique & Philip Lehner Herpetology, Losos Lab Ornithology, Edwards Lab Concord Field Station, Biewener Lab Professor for the Study of Latin America; Curator of Herpetology Andrew Gehrke Mary Stoddard Bruno Souza de Medeiros Invertebrate Zoology, Srivastava Lab Ornithology, Edwards Lab Entomology, Farrell Lab James J. McCarthy Professor of Biological Oceanography; Alexis Harrison Katherine Stryjewski Talia Moore Alexander Agassiz Professor of Biological Herpetology, Losos Lab Ornithology, Edwards Lab Concord Field Station, Biewener Lab Oceanography; Acting Curator of Caroline Hu Gerard Talavera Pavitra Muralidhar Malacology Mammalogy, Hoekstra Lab Entomology, N. Pierce Lab Herpetology, Losos Lab Naomi E. Pierce Charlotte Jandér Chris Tijs Shayla Salzman Sidney A. & John H. Hessel Professor of Entomology, N. Pierce Lab Concord Field Station, Biewener Lab Entomology, N. Pierce Lab Biology; Curator of Lepidoptera Katrina Jones Ana Lúcia Miranda Tourinho Allison Shultz Stephanie E. Pierce Vertebrate Paleontology, S. Pierce Lab Invertebrate Zoology, Giribet Lab Ornithology, Edwards Lab Assistant Professor of Organismic & Evolutionary Biology; Curator of Robert Kambic Tracy Washington Kari Taylor-Burt Vertebrate Paleontology Vertebrate Paleontology, S. Pierce Lab Ornithology, Edwards Lab Concord Field Station, Biewener Lab Mansi Srivastava Betul Kacar Melissa Whitaker Kira Treibergs Assistant Professor of Organismic Ornithology, Edwards Lab Entomology, N. Pierce Lab Marine Invertebrates, Woollacott Lab & Evolutionary Biology; Curator of Albert Kao Dylan Wainwright Invertebrate Zoology Entomology, N. Pierce Lab Graduate Students Ichthyology, Lauder Lab Robert M. Woollacott Christopher Kenaley Caitlin Baker Xuemai Zhai Professor of Biology; Curator of Marine Ichthyology, Lauder Lab Invertebrate Zoology, Giribet Lab Biological Oceanography, McCarthy Lab Invertebrates Nicolai Konow Maude Baldwin Emeritus Faculty Concord Field Station, Biewener Lab Ornithology, Edwards Lab Associates A.W. “Fuzz” Crompton Sebastian Kvist Nicole Bedford Gary Alpert Faculty-Curator, Emeritus; Fisher Invertebrate Zoology, Giribet Lab Mammalogy, Hoekstra Lab Associate of Entomology Professor of Natural History, Emeritus Harvard University Oriol Lapiedra John Boyle Richard C. Lewontin Herpetology, Losos Lab Entomology, N. Pierce Lab Brian S. Arbogast Professor of Biology, Emeritus; Alexander Associate of Mammalogy Jean-Marc Lassance Rebecca Buckman Agassiz Professor of Zoology, Emeritus University of North Carolina Wilmington Mammalogy, Hoekstra Lab Invertebrate Zoology, Giribet Lab Edward O. Wilson Bruce Archibald Fabio Laurindo da Silva Richard Childers Honorary Curator in Entomology; Associate of Entomology Entomology, Farrell Lab Entomology, N. Pierce Lab Pellegrino University Professor, Emeritus Simon Fraser University Sarah Lemer Glenna Clifton Invertebrate Zoology, Giribet Lab Concord Field Station, Biewener Lab

Annual Report 2015 . 2016 31 MCZ PERSONNEL

Aaron Bauer Leslie S. Kaufman Stephen Tilley Katherine Eldridge Associate of Herpetology Associate of Ichthyology Associate of Herpetology Curatorial Assistant, Ornithology Smith College Villanova University Boston University Charles Farnum Andrew Berry Gisele Y. Kawauchi James Traniello Curatorial Assistant, Entomology Associate of Entomology Associate of Population Genetics Associate of Invertebrate Zoology Helene Ferranti Boston University Harvard University University of São Paulo Faculty/Collection Assistant, Biological Elizabeth Brainerd Timothy Laman David Wagner Oceanography & Marine Biology Associate of Entomology Associate of Ichthyology Associate of Ornithology Dana Fisher University of Connecticut Brown University National Geographic Assistant to the Librarian/Special Jae Choe Ruth Hortencia Bastardo Landrau David Wake Collections, Ernst Mayr Library Associate of Herpetology Associate of Entomology Associate of Entomology Linda S. Ford University of California, Berkeley Ewha Womans University Universidad Autónoma de Santo Director, Collections Operations Domingo Marvalee Wake Janet Collett Matthew Gage Associate of Herpetology Associate of Population Genetics Phillip Lobel Herpetology University of Sussex Associate of Ichthyology University of California, Berkeley Brendan Haley Boston University Philip S. Ward Bruce Collette Senior Database Manager Associate of Ichthyology David Lohman Associate of Entomology National Marine Fisheries Service Associate of Entomology University of California, Davis Karsten Hartel The City College of New York Curatorial Associate, Ichthyology David Bruce Conn Jacqueline Webb Associate of Invertebrate Zoology Vladimir A. Lukhtanov Associate of Ichthyology Rachel Hawkins Berry College Associate of Entomology University of Rhode Island Curatorial Assistant, Entomology Russian Academy of Sciences James Costa R. Haven Wiley Andra Hollis Associate of Entomology James Mallet Associate of Ornithology Staff Assistant, Concord Field Station Western Carolina University Associate of Population Genetics University of North Carolina Kathleen Horton Harvard University Catherine Craig Cheryl Wilga Assistant with Professor Wilson, Associate of Invertebrate Zoology Russell Mittermeier Associate of Ichthyology Entomology Conservation Through Poverty Associate of Herpetology University of Rhode Island Jared Hughes Alleviation, International Conservation International Judith Winston Faculty/Collection Assistant, Herpetology Harlan Dean Piotr Naskrecki Associate of Marine Biology & Vertebrate Paleontology Associate of Invertebrate Zoology Associate of Entomology Virginia Museum of Natural History Nikki Hughes Harvard University Conservation International Faculty/Collection Assistant, Lloyd Demetrius Diane B. Paul Staff Mammalogy & Ornithology Associate of Population Genetics Associate of Population Genetics Melissa Aja Amie Jones Harvard University Harvard University Museum Projects Coordinator Faculty/Collection Assistant, Philip DeVries David L. Pawson Adam Baldinger Entomology Associate of Entomology Associate of Marine Biology Curatorial Associate, Invertebrate Marcia Kazmierczak University of New Orleans Smithsonian National Museum of Zoology, Malacology & Marine Staff Assistant, Collections Operations Natural History Invertebrates Gregory D. Edgecombe Michelle Kennedy Associate of Invertebrate Zoology Stewart Peck Dorothy Barr Collections Information & Database Natural History Museum, England Associate of Entomology Public Services/MCB Liaison Librarian, Specialist Carleton University Ernst Mayr Library Ben Evans Laura Leibensperger Associate of Herpetology Paulo Petry Penny Benson Curatorial Assistant, Invertebrate McMaster University Associate of Ichthyology Curatorial Assistant, Malacology Zoology The Nature Conservancy Brooke E. Flammang Ronnie Broadfoot Jennifer Lenihan Associate of Ichthyology Steve Poe Circulation/Reference, Ernst Mayr Library Curatorial Assistant, Invertebrate New Jersey Institute of Technology Associate of Herpetology Zoology University of New Mexico Linnaea Burt Kelvin A. Guerrero Cryogenic Collections Lisa Litchfield Associate of Entomology Michael Rex Administrator, Concord Field Station Systematic Entomologist/Environmental Associate of Malacology Christopher Capobianco Consultant University of Massachusetts, Boston Vertebrate Paleontology Technician & David Lowery Preparator Project Programmer, Biodiversity Michael Hadfield Jessica Rykken Informatics Associate of Marine Biology Associate of Entomology Judith Chupasko University of Hawaii Harvard University Curatorial Associate, Mammalogy Joseph Martinez Curatorial Assistant, Herpetology Berthold HÖlldobler Çağan H. Şekercioğlu April Collins Associate of Entomology Associate of Ornithology Acquisitions and Technology Specialist, Patrick McCormack Arizona State University University of Utah Ernst Mayr Library Curatorial Assistant, Entomology Gustavo Hormiga Andrea Sequeira Stefan Cover John Mewherter Associate of Invertebrate Zoology Associate of Entomology Curatorial Assistant, Entomology Curatorial Assistant The George Washington University Wellesley College Jessica Cundiff Paul J. Morris Helen F. James Steven O. Shattuck Curatorial Associate, Invertebrate & Biodiversity Informatics Manager Vertebrate Paleontology Associate of Ornithology Associate of Entomology Catherine Musinsky National Museum of Natural History, Commonwealth Scientific and Industrial Tatiana De Souza Varges Faculty/Collection Assistant, Smithsonian Research Organization (CSIRO) Curatorial Assistant , Collections Operations Mammalogy Alan Kabat Scott R. Shaw Joseph DeVeer John Nevins Associate of Malacology Associate of Entomology Head of Technical Services, Ernst Mayr Laboratory Systems Manager for Biological Attorney, Bernabei & Wachtel University of Wyoming Library Oceanography & Marine Biology

32 Museum of Comparative Zoology MCZ PERSONNEL

Mark Omura Temporary Staff Miguel San Kristin Pennarun Curatorial Assistant, Mammalogy Anthony Buda Ernst Mayr Library Manager of OEB Research Administration Services Julianne Pelaez Ernst Mayr Library Carly Sanker Bridget Power Curatorial Assistant Kimberly Carlin Encyclopedia of Life Administrative Coordinator Philip Perkins Mammalogy Elizabeth Schatzkin Christopher Preheim Curatorial Associate, Entomology Anne Everly Mammalogy Senior Academic Administrator Pedro Ramirez Ichthyology Khan Shairani Research Assistant, Concord Field Ernst Mayr Library Keleigh Quinn Kenneth Fu Station Senior Research Administrator Ernst Mayr Library Lauren Shear Patrick Randall Collections Operations Peg Richard Michael George Ernst Mayr Library Financial Assistant Cryogenic Collection Kaitlin Sheridan Murat Recevik Invertebrate Zoology Damari Rosado Morgan Groom Curatorial Assistant, Malacology Associate Director of Administration Ernst Mayr Library Emily Smith Mark Renczkowski Ernst Mayr Library Anna Salvato Sarah Guth Curatorial Assistant, Invertebrate Manager of Financial Operations Entomology Mingyuan Song Paleontology Ernst Mayr Library Geoff Tierney Emily Hamblet Constance Rinaldo Associate Director of Finance and Ernst Mayr Library Ashley Steinkraus Librarian, Ernst Mayr Library Research Administration Encyclopedia of Life Jane Harrison Alana Rivera Ornithology Michael Vitale Curatorial Assistant, Collections The MCZ deeply appreciates Encyclopedia of Life Operations Gwendolyn Fougy Henry the additional support and Ernst Mayr Library Yuchen Xiao José Rosado contributions of numerous interns Marine Invertebrates Curatorial Associate, Herpetology Madeleine Higgins and undergraduate students during Mammalogy Mary Sears Encyclopedia of Life, Learning the 2015–2016 academic year. Head of Public Services, Ernst Mayr Yiyuan Hu Library Ernst Mayr Library + Education Group Tracy Barbaro Diane Sheridan Jyhjong Hwang Project Coordinator Faculty/Collection Assistant, Collections Operations Invertebrate Zoology Jeffrey T. Holmes Kalvin Janik Digital Learning Editor Deborah Smiley Ernst Mayr Library MCZ Faculty Managing Editor, MCZ and HUH Amy Lorenz Kaitlyn Justus Project Coordinator The MCZ’s charter, signed in Meaghan Sorce Ernst Mayr Library 1859, mandates that the Curatorial Assistant Marie M. Studer Museum’s activities will be Sang Il Kim Learning + Education Director Ernst Mayr Library overseen by a governing Margaret Starvish board, the Faculty of the Faculty/Collection Assistant, Ichthyology Lacey Klingensmith Administration for the Museum of Comparative & Entomology Herpetology Zoology. Tsuyoshi Takahashi Department of Organismic Adam Kowalczyk & Evolutionary Biology Dr. John D. Constable Curatorial Assistant, Herpetology & Invertebrate Paleontology Collections Operations Krista Carmichael Mr. Robert G. Goelet Alina Lazar Senior Research Administrator Jennifer Thomson Ernst Mayr Library Mr. George Putnam, Jr. Faculty/Collection Assistant, Populations Rebecca Chetham Genetics Maria Lindquist Executive Director Mr. George Putnam III Ernst Mayr Library Jeremiah Trimble Irv Dumay Dr. Barbara Jil Wu Curatorial Associate, Ornithology Lily Lu Building Manager Mr. Paul J. Zofnass Ornithology Diana Turmenne Paul Dwyer President Drew Gilpin Faust Curatorial Assistant, Collections Liam McGill Mailroom Staff Assistant Operations Mammalogy Donna Gadbois Kenneth Wilcox Elizabeth Meyer Financial Associate Building Services Coordinator Ernst Mayr Library Jason Green Victoria Wilke Rachel Moon Financial Associate Acknowledgements Curatorial Assistant, Collections Ernst Mayr Library Alexander Hernandez-Siegel This annual report was Operations Robert Morris Senior Academic Programs produced by the Office of the Andrew Williston Biodiversity Informatics Administrator Director of the Museum of Curatorial Assistant, Ichthyology Comparative Zoology. Jessica Mullen Wendy Heywood Jonathan Woodward Malacology Communications & Events Coordinator Editors Curatorial Assistant, Collections Keiko Nishimoto Megan McHugh James Hanken, Director Operations Botany Library Human Resources Coordinator Melissa Aja, Museum Projects Robert Young Robert Oppenheimer Philip Norton Coordinator Special Collections Librarian, Ernst Concord Field Station Assistant Building Manager Mayr Library Copy, Design & Production Valentine Poumadere Jeremiah O’Connor Breda Zimkus Cyndi Wood Ernst Mayr Library Financial Analyst Cryogenics Collections Manager for Creative Project Genetic Resources Jessica Roth Monica Oyama Management, Inc. Collections Operations Financial Associate creativeprojectmgmt.com

Annual Report 2015 . 2016 33 26 Oxford Street Cambridge, MA 02138 617.495.2460 mcz.harvard.edu