Gonzalo Giribet - US grants

Cyphophthalmi constitutes a suborder of the arachnid order Opiliones (daddy-long-legs). Of ancient origin and widespread distribution, they live in the leaf litter of undisturbed temperate to tropical humid forests. Cyphophthalmids have been found in all continents, except Antarctica, and in most islands of continental origin, such as New Zealand, New Caledonia, Sumatra, and Madagascar. Many of the places they inhabit qualify as biodiversity hotspots, areas collectively comprising only 1.4% of the land surface area of the Earth and recommended for preferential conservation policies for having high diversity. Due to their localized distribution and scant dispersal ability, cyphophthalmids have been chosen as a model to study evolutionary radiations in islands of continental origin, biogeography, and potential issues of conservation biology. To do so, the taxonomy and evolutionary relationships among its species need to be worked out. We will use scanning electron microscopy to study the morphology of the cyphophthalmids and will generate lists of characters to be analyzed with standard phylogenetic algorithms. Other specimens will be used for genetic studies in order to generate DNA sequence data also to be analyzed phylogenetically. Both sets of information (morphology and DNA) will be used to propose the evolutionary relationships of the members of the group and to establish a classification. The evolutionary trees will also aid studies of biogeographic patterns and earth history. Evolutionary analyses will be aided with the use of software written to work in parallel on a large multiprocessor computer cluster. Extensive field work in New Zealand, South Africa, Madagascar, tropical West Africa and southeast Asia is required to obtain material suitable for laboratory work and to document undescribed diversity.

Systematics, biogeography, and the study of evolutionary radiations have contributed to a better understanding of the patterns observed in nature. Optimizing the knowledge of diversity across geographic areas, cladogenesis, and speciation may benefit from the use of model systems. The potential use of Cyphophthalmi as a new model in evolutionary biology has driven a dual study of the higher systematics of the group and the exploration of one of their most spectacular adaptive radiations, the one undergone in New Zealand. The research will develop a new model system to integrate phylogenetic and biogeographic patterns with evolutionary studies of explosive radiations, and will form the basis for future studies on biodiversity estimates and conservation biology. Global knowledge of the Cyphophthalmi, with a database containing geographical information and illustrations for all the estimated 180 species of the group, will be made available. During the process, one undergraduate and one graduate student will be fully trained in arachnid evolutionary biology. The value of the research project thus has both scientific and social interests.

A grant has been awarded to Dr. Gonzalo Giribet of Harvard University to assemble the basic backbone of the protostome tree of life, which includes all triploblastic animals except those directly related to vertebrates. Protostomes constitute more than one million of the 1.7 million named species of living organisms, including familiar animals such as mollusks, insects, flatworms, roundworms, and many others of medical and economic importance. However, relationships among these groups are still contentious, so the backbone of the animal tree of life cannot currently be proposed without a large degree of uncertainty. This proposal will integrate and disseminate the broadest possible collection of information on selected representative protostomes to address questions of their origin and evolution over more than 540 million years. In order to do this we will integrate anatomical data on extant and extinct fauna with developmental data using cell-lineage studies and broad-based genomic information. Those data will be collected for a large pool of protostome species which will be studied at a level never before attempted in non-model organisms.
The awarded grant will contribute to the training of students at different academic levels, emphasizing underrepresented groups in science. Results will be disseminated to society through partnership with five of the most prominent natural history museums in the world, including the Harvard Museum of Natural History and the American Museum of Natural History in the USA, and the most important museums in Australia and Denmark. It is also the intention of the PI to publish an article in non-specialist scientific magazines for a broad lay audience.

In a renewal of a highly successful program of taxonomic research and training in the PEET activity, Partnerships for Enhancing Expertise in Taxonomy, Prof Gustavo Hormiga at George Washington University and colleagues at Harvard University, the California Academy of Sciences, and the Smithsonian seek to train additional systematic biologists (two likely Ph.D. students and one postdoctoral scholar) in monographic and revisionary study of a megadiverse group, spiders. The trainees, working in close collaboration with the PIs and several expert collaborators, will study and describe spider species in the orb weaving families Tetragnathidae and Mysmenidae. The main goal is to train three taxonomic experts in several critically understudied groups of mostly tropical spiders. These trainees will receive state of the art education in systematic methods and will interact with a diversity of project collaborators and academic and research institutions. Their training will allow them in the future to carry out systematic research on any taxonomic group of spiders. The phylogenetic component of these projects will use character evidence from morphology, behavior and DNA sequences. The taxonomic aspects of this research will make extensive use of computerization and web-based dissemination. The monographs and systematic data resulting from this research will be published in peer reviewed journals.
In addition, the project web site (www.gwu.edu/~spiders/) will serve as an outlet to advertise research results and to disseminate these, including: Interactive identification keys, Digital Image Banks (of taxonomic images of the world genera of Tetragnathidae and Mysmenidae to assist with identification), Database of images of web architecture (with images hyperlinked to a file with the specimen voucher data), and Specimen
databases of geo-referenced material for the monographed taxa.

Like many isolated islands, New Zealand is home to evolutionary radiations in various animal groups such as velvet worms, cockroaches, land planarians, or the better-known ratite birds (kiwi birds, giant moas and allies). One of these radiations has occurred in the arachnid family Pettalidae, a group of tiny harvestmen with Southern Hemisphere distribution across the fragments of the former temperate zone of Gondwana. A remarkable 60% of the described diversity of this family is found in New Zealand. This study, focusing in one common species (Rakaia denticulata), will provide insight into the processes contributing to speciation in small organisms that live in the forest litter and that are essential for the functioning of soil ecosystems.

In addition, this grant will contribute to the training of a female graduate student, support of female and minority undergraduates, and the development of K-12 outreach programs.

A grant has been awarded to Dr. Gonzalo Giribet of Harvard University and collaborators to assemble the basic backbone of the protostome tree of life, which includes all triploblastic animals (animals with three body layers, as opposed to the simpler jellyfish, anemones, and sponges) except those directly related to vertebrates. Protostomes constitute more than one million of the 1.7 million named species of living organisms, including familiar animals such as mollusks, insects, earthworms, flatworms, roundworms, and many others of medical and economic importance. However, relationships among these groups are still contentious, so the backbone of the animal tree of life cannot currently be proposed without a large degree of uncertainty. This proposal will integrate and disseminate the broadest possible collection of information on selected representative animals to address questions of their origin and evolution over more than 540 million years. In order to do this we will integrate anatomical data on extant and extinct fauna with developmental data using cell-lineage studies and broad-based genomic information. Those data will be collected for a large pool of protostome species which will be studied at a level never before attempted in non-model organisms. The outcome of this research will contribute to rewriting what it is known about animal evolution.
The awarded grant will contribute to the training of students at different academic levels, emphasizing underrepresented groups in science. Results will be disseminated to society through partnership with five of the most prominent natural history museums in the world, including the Harvard Museum of Natural History and the American Museum of Natural History in the USA, and the most important museums in Australia and Denmark. It is also the intention of the AToL protostome team to publish an article in non-specialist scientific magazines for a broad lay audience.

In this project, a large team of investigators will resolve evolutionary relationships within the bivalves (clams, oysters, mussels, scallops, etc.). Bivalves are a diverse and familiar group of mollusks with an old and well-preserved fossil record. Bivalves have important ecological roles in marine and freshwater ecosystems, and economic roles including fisheries, the ornament industry, and health sciences. Bivalves are the second largest class of living mollusks, which in turn constitute the second largest animal phylum and the largest in the marine realm. The evolutionary history and relationships of bivalves will be investigated through a fresh look at bivalve anatomy in combination with study of selected DNA markers for the same species investigated morphologically. This international team of investigators will assemble morphological and molecular phylogenetic data at levels of detail never before attempted. Outreach activities will engage various audiences in the project's results and in evolutionary science as a whole.

This project presents an excellent opportunity for outreach to communicate the concepts of evolution, and this team of researchers is well-placed to accomplish this. Museum exhibits and instructional materials using bivalves will be developed for K-12 teachers and their students, post-secondary teachers and their students, and the general public, in the form of a traveling exhibit ("Evolution on the Half-Shell") and associated teaching materials.

Molluscs are megadiverse animals, and the roughly 200,000 species are economically and culturally important. Discerning the evolutionary relationships within this group has been particularly difficult using traditional approaches, but a detailed understanding of such relationships would facilitate studies of all aspects of their biology and provide insight into general evolutionary principles. This project will resolve molluscan evolutionary relationships with new gene sequence data collected with high-throughput DNA sequencing technology. The research will resolve important evolutionary questions, such as the selective factors leading to the evolution of wildly disparate body plans, ranging from worm-like mollusks to shelled forms to the highly modified jet-propulsion body plan of cephalopods.


In addition, new analytical tools will be developed that will improve the efficiency of research across the emerging field of phylogenomics (the use of genome-wide information for studying organismal evolutionary relationships). These tools will be made freely available to the scientific community. The project will also include outreach components that will integrate research and education through a series of lectures at the Harvard Museum of Natural History and a regular podcast series. The podcasts will be designed for use within high school biology classes, and understandable and informative to a general audience.

Arachnids comprise one of the most diverse groups of animals, with numerous representatives in all terrestrial ecosystems. Aside from Acari (mites and ticks), whose total estimated diversity might reach up to one million species, the orders Araneae (spiders) and Opiliones (harvestmen or daddy longlegs) are the most diverse arachnid lineages. The main goal of this project is to carry out taxonomic revisions (monographs) of several lineages across selected Neotropical groups in two arachnid orders (Araneae [spiders] and Opiliones [daddy long legs]). The groups that we have targeted in this proposal are mainly leaf litter inhabitants (except for mimetids), grossly undersampled and understudied, that lack modern revisionary work and include many undescribed taxa at the species level and above, and thus provide excellent research and training opportunities in modern monography and systematics. The largely overlapping Neotropical distribution of our study groups allows us to effectively collect these arachnids in joint field expeditions. Some of these study taxa also pose similar biogeographic and evolutionary questions that can be addressed with common methodology and expertise.

To carry out our research and training goals we have assembled a team of scientists with diverse backgrounds in systematics and arachnology and a demonstrated history of successful collaborative research efforts. Combining research efforts to tackle these taxonomic problems provides the empirical basis for our multi-ordinal and multidisciplinary approach for training doctoral students in the broadest possible way. Lab rotations and collaboration among the multiple investigators participating in the projects, in addition to the joint field expeditions, also contribute to this synergism. Research results will be disseminated in high impact portals such as Encyclopedia of Life and Tree of Life. Online interactive identification tools will also be produced.

The prorhynchid flatworms are a diverse and abundant group of predatory microscopic invertebrates living in humid soils and freshwaters on all regions of the globe. However, as is the case with most animals showing small body size, simple anatomy, and cryptic habitats, their biodiversity has gone almost entirely unstudied by modern biologists; unnamed species may outnumber the 29 species known to science by the hundreds, and their evolutionary relationships are essentially unknown. This thesis research aims to combine worldwide sampling of new and known prorhynchids with the collection of genome-wide DNA sequence data from many dozens of species to arrive for the first time at an unbiased picture of their global diversification.

The purpose of this dissertation improvement grant is to support the development of a novel method to cheaply sequence hundreds of genes from arbitrarily large numbers of organisms, which will contribute to an ongoing revolution in the way biologists analyze the evolutionary relationships of closely-related species, the leaves of the tree of life. Applying this method to prorhynchids, and showcasing these and other microscopic invertebrates to undergraduate students and community members (in collaboration with a local science education organization), will serve to further engage both scientific and public interest in such organisms, the largest and simultaneously least-understood components of global animal biodiversity.

This project will chronicle the evolution and diversification of the orb-weaving spiders to resolve a long-standing problem in animal evolution, that is, the origin and genealogical relationships of the main groups of orb-weavers. Ubiquitous, diverse, and exclusively predatory, spiders are one the most diverse and fascinating animal groups and the dominant arthropod predators in most terrestrial ecosystems. Orb-weavers are remarkable not only because of the high number of species described so far, but also because of the extraordinary architectural diversity of their foraging webs, ranging from highly geometric snare designs, often regarded as a pinnacle of animal engineering, to sheet and irregular cob webs. Understanding web evolution and diversification requires an empirically robust hypothesis about the underlying genealogical patterns - the spider tree of life. Traditional anatomical and genetic methods have not yet produced a compelling answer to this problem, and thus our understanding of the evolution of this group is far from satisfactory. The researchers of this project will use new methods to gather genetic data to answer this long-standing question. The results of this research will be of interest to many biologists working on any comparative aspects of orb-weavers because it will provide the genomic resources and the fundamental comparative basis for studies on a broad diversity of topics (silk biology, sexual selection, venom evolution, etc.). This project will serve as a training platform for postdoctoral, graduate and undergraduate researchers. In addition, the project will produce a public exhibit at the Harvard Museum of Natural History that will explore the evolution and biology of orb-weaving spiders.

A team of three investigators, experts in arachnid systematics and with a long and productive history of research collaboration, will generate novel data through the sequencing of 120 transcriptomes and target capture approaches using modern DNA sequencing techniques for phylogenetic inference for a taxonomic sample of orbicularians that would include representatives of all 21 known families of orb-weavers and a diverse array of outgroups. These genomic data will be used to infer the family-level phylogenetic relationships of orb-weaving spiders using a diversity of analytical techniques. The PIs will also analyze a morphological matrix of 85 selected orbicularians largely based on extensive data already at hand. This project will use the results of the phylogenetic analyses to carry out comparative analyses of orb-weaver diversification and of web architecture evolution.

Understanding what drives the evolution of diverse animal forms is a largely unanswered key question in evolutionary biology. This research project will investigate the tree of life of living and fossil groups of marine snails, and the changes that happened in the shape of their shells through geological time. Snail shells are an iconic example of an organisms' effective defense against predators. With more than 30,000 described living species and countless fossils, snails are the most diverse group of invertebrate animals in the oceans; the diversity of their shell forms being equally impressive. Their rich fossil record shows a transition of shell features starting in the middle of the Mesozoic Era, around 160 million years ago. At that time, while dinosaurs thrived on land, fishes and crabs with increasing capacity of crushing shells were diversifying in the oceans. Changes in shell shape are thus hypothesized to be an evolutionary response to increased predation. If predation pressure was indeed an important driver during the evolution of shell shape, a general dominance of better-defended forms would be expected after the mid-Mesozoic. The researchers will gather data from an ample set of shells from fossil and living species to infer how they are related to each other and to estimate the times of origin of snail groups with different shapes; these results will be used to investigate the trajectory of the shell shape during the Mesozoic. Results from this research will be published in peer-reviewed journals and science blogs broadly accessible to the public. A large number of shell images will be produced and made available online, including websites for science communication. In addition, high school and undergraduate students will be mentored in their first experience in science.
The members of Vetigastropoda, the specific group of marine snails under study (including abalone, key-hole limpets, top-shells and others), present great variation in shell height and level of coiling. Shorter and less coiled shells are particular features that could offer mechanical resistance against shell crushing. This study will test the hypotheses that: (1) the morphospace of shells became more restricted in response to predator selective pressure, and (2) the trajectory of shell shape trended towards shorter and less spired forms. Hypothesis testing will be accomplished by adding shell morphological data to a molecular phylogeny inferred from high-throughput sequence data. The fossil data will provide the basis for estimating divergence times in the phylogeny of vetigastropods, and for the comparison among distinct models of trait evolution.

The Eastern Seaboard of the United States (ESB, U.S. Exclusive Economic Zone) stretches from the Canadian border on the Atlantic along nearly 6,000 km of eastern coastline, around the Floridian Peninsula, and along the Gulf of Mexico to the south end of the Texan coast, including 18 U.S. states. The ESB region is densely populated, with 47% of the U.S. population expected to inhabit the counties adjacent to the shoreline by 2021. Habitat loss, pollution, overfishing, and climate change threaten commercially and ecologically important marine species all along the ESB. This project will make occurrence data with map coordinates available for over 3,000 species of mollusks that find their habitat along the ESB, including mussels, clams, conchs, snails, and squid. Data from these ecologically and commercially important species (over 4.5 million individual specimens) will be made available through public online data portals. While the geographic ranges for many species of mollusks are well-known, the extent of their distribution within the seafloor habitats they occupy is unknown. Adding map coordinates to occurrence records for live-collected mollusks in natural history collections will provide detailed knowledge of distributions. Because natural history collections have specimens collected from the mid-1800s to present, these occurrence records can help track distributional changes over time and lead to better fisheries and conservation management.

One hundred million mollusk specimens have been documented in natural history collections across North America, and the breadth, depth, and growth of these collections is exceptionally well-known compared to other invertebrate taxa. Mollusks are among the best sampled group of animals, with some species having over 2,000 digital records available in natural history collections making them extremely well-suited for environmental and biogeographical studies that track faunal change over time and space. However, already-digitized mollusk lots are missing essential data such as collecting date (30% of records) and reliable georeferences (85% of records). This project will generate reliable geo-coordinate data for all covered specimen lots using a collaborative georeferencing project in GeoLocate. GeoLocate will add layers for bathymetric data, benthic habitat, and marine conservation areas. Incorporating bathymetry into GeoLocate to determine the extent of locations will also provide that capability for complex elevational data for terrestrial species. Important trait data will also be incorporated. For the first time, molluscan occurrence data will distinguish between live- and dead-collected specimens, with a defined vocabulary for traits added to each record. Due to the long persistence of molluscan shells, the live/dead- collected distinction is crucial for all studies of biotic change using mollusks. Information on collecting dates will be refined where possible to increase resolution for detecting biotic changes. The data will be shared through public data repositories, including iDigBio, GBIF, OBIS, and the InvertEBase Symbiota portal.

This award reflects NSF's statutory mission and has been deemed worthy of support through evaluation using the Foundation's intellectual merit and broader impacts review criteria.