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E-raamat: Phylogenomics: A Primer 2nd edition [Taylor & Francis e-raamat]

(American Museum of Natural History, USA), ,
  • Formaat: 380 pages, 55 Tables, black and white; 165 Illustrations, black and white
  • Ilmumisaeg: 19-Aug-2020
  • Kirjastus: CRC Press
  • ISBN-13: 9780429397547
Teised raamatud teemal:
  • Taylor & Francis e-raamat
  • Hind: 350,83 €*
  • * hind, mis tagab piiramatu üheaegsete kasutajate arvuga ligipääsu piiramatuks ajaks
  • Tavahind: 501,18 €
  • Säästad 30%
  • Formaat: 380 pages, 55 Tables, black and white; 165 Illustrations, black and white
  • Ilmumisaeg: 19-Aug-2020
  • Kirjastus: CRC Press
  • ISBN-13: 9780429397547
Teised raamatud teemal:

Phylogenomics: A Primer, Second Edition

is for advanced undergraduate and graduate biology students studying molecular biology, comparative biology, evolution, genomics, and biodiversity. This book explains the essential concepts underlying the storage and manipulation of genomics level data, construction of phylogenetic trees, population genetics, natural selection, the tree of life, DNA barcoding, and metagenomics. The inclusion of problem-solving exercises in each chapter provides students with a solid grasp of the important molecular and evolutionary questions facing modern biologists as well as the tools needed to answer them.

Section I Foundations of Phylogenomics
Chapter 1 What is Phylogenomics?
3(12)
Phylogenomics and Bioinformatics
3(2)
Bioinformatics Tools for Finding Patterns in Biological Experiments
5(2)
The rise of phylogenomics
5(2)
Sub-Branches of Phylogenomics
7(1)
The Phylogenomic Toolbox
8(1)
Basic Computational Tools in Phylogenomics
8(1)
Statistics Help Compare Genetic Sequences and Generate Phylogenetic Trees
8(2)
Parametric Statistics Are Derived from Distributions
10(1)
Nonparametric statistical Analyses Are useful in Many Situations
10(1)
Maximum Likelihood and Bayesian Analysis Are Standard Statistical Methods Used in Phylogenomics
11(1)
Key Attributes of Phylogenomicists
11(2)
Summary
13(1)
Recommendations for students
13(1)
Discussion Questions
13(1)
Further Reading
13(2)
Chapter 2 The Biology and Sequencing of Genetic Information: DNA, RNA, and Proteins
15(18)
Nucleic Acids
15(2)
DNA molecules efficiently transmit information
15(1)
DNA is synthesized by specific pairing
15(2)
DNA can mutate and lead to descent with modification
17(1)
Polymerase chain reaction (PCR) is a milestone development
17(1)
Proteins
17(6)
Proteins are linear polymers of amino acids
17(1)
Proteins have multiple levels of structure
18(3)
Translation of DNA to amino acids is accomplished by the genetic code
21(2)
Reading frame in nucleic acid sequences
23(1)
The DNA Data Explosion
23(4)
Nucleic acid sequencing methods are increasingly powerful
23(1)
Next-generation sequencing allows for rapid analysis of genomes
24(2)
Other applications of next-generation sequencing
26(1)
Alternatives to Whole Genome Sequencing
27(1)
Single-nucleotide polymorphisms (SNPs) differ at one position in a designated DNA sequence
27(1)
Microarrays
27(1)
Genome reduction methods
27(1)
Analyzing Gene Expression
28(2)
RNA-Seq is a method for obtaining transcriptomic data
29(1)
Summary
30(1)
Recommendations for students
30(1)
Discussion Questions
30(1)
Further Reading
31(2)
Chapter 3 Evolutionary Principles: Populations and Trees
33(16)
Darwin, Wallace, and Evolutionary Theory
33(2)
Four early contributions
33(1)
Darwin's ideas lacked a valid genetic mechanism
34(1)
The study of evolution can be divided into microevolution and macroevolution
34(1)
Microevolution
35(3)
Population genetics focuses on microevolution
35(1)
Advances in molecular techniques led to new thinking in evolutionary biology
35(1)
Codon changes and usage can provide insights into natural selection
36(1)
Microevolutionary studies often rely on computational modeling
36(2)
Macroevolution
38(3)
Macroevolution studies rely heavily on systematics and phylogenetics
38(1)
Relationships and systematics
38(1)
There are several approaches to tree building
38(1)
Tree thinking
38(2)
Phylogenetics can help establish homology
40(1)
Species
41(2)
The definition of species is heavily debated
41(1)
Defining species phylogenetically
42(1)
Updates on Darwinian Evolution
43(2)
Punctuated equilibrium suggests that not all evolution is gradual
43(1)
Epigenetic changes are caused by influences outside of the genetic system
44(1)
Summary
45(1)
Suggestions for Students
45(1)
Evolution Websites
45(1)
Discussion Questions
45(1)
Further Reading
46(3)
Section II Data
Chapter 4 Data Storage---The Basics
49(14)
Databases and Phylogenomics
49(2)
DNA sequences are stored in large international databases
50(1)
Specific data sets may be held in special repositories
50(1)
These databases offer free access and availability for scientific inquiry
51(1)
Information Retrieval from the NCBI Database
51(10)
Publications are archived in the PubMed database
52(1)
Working with molecular sequences stored in GenBank
52(6)
Whole genomes are accessible on the Genome Page
58(3)
Summary
61(1)
Recommendations for Students
61(1)
Discussion Questions
61(1)
Further Reading
61(2)
Chapter 5 Sequence Alignment and Searching Sequence Databases
63(18)
Homology of Genes, Genomic Regions, and Proteins
63(5)
Genomes can diverge by speciation and by duplication
63(1)
Sequence alignment is an important procedure in phylogenomics
64(1)
Basic, paired nucleic acid sequence alignment
65(1)
Basic, paired protein sequence alignment
66(2)
Dynamic programming and sequence alignment
68(1)
Database Searching via Pairwise Alignments: The Basic Local Alignment Search Tool
68(9)
BLAST identifies highly similar sequences
69(1)
BLAST is optimized for searching large databases
69(2)
There are multiple types of BLAST for nucleotide and amino acid sequences
71(1)
BLAST searches are easy to do
72(3)
Whole genome alignments can also be performed
75(2)
Summary
77(1)
Recommendations for Students
78(1)
Discussion Questions
78(1)
Further Reading
79(2)
Chapter 6 Multiple Alignments
81(10)
Multiple Sequence Alignment
81(4)
Changing Alignment Parameters
85(1)
Multiple optimal alignments may exist
85(1)
Specialized Alignment Programs
86(1)
Choosing an Alignment Program
86(2)
Automated alignment results are frequently adjusted "by eye"
87(1)
Alignment programs can be compared by use of benchmark data sets
88(1)
Dynamic versus Static Alignment
88(1)
Summary
89(1)
Recommendations for Students
89(1)
Discussion Questions
89(1)
Further Reading
90(1)
Chapter 7 Genome Sequencing and Annotation
91(10)
Whole Genome Sequencing (WGS)
91(4)
Next-generation sequencing
91(1)
The nuts and bolts of assembly
92(3)
Gene Finding and Annotation
95(3)
Gene finding can be accomplished via extrinsic, ab Initio, and comparative approaches
96(1)
Gene functional annotation
97(1)
Genome completeness
97(1)
Summary
98(1)
Recommendations for Students
98(1)
Discussion Questions
98(1)
Further Reading
99(2)
Chapter 8 Genomics Databases: Genomes and Transcriptomes
101(14)
Genome information is Stored in Multiple Locations
101(7)
BioSample/BioProject/Short Read Archives (SRA) store archival information for projects used in broader genomics research archived in INSDC
101(7)
Data Archiving and Databases Outside of the INSCD System
108(6)
Organismal-focused genome and transcriptome databases
110(4)
Summary
114(1)
Recommendations for students
114(1)
Problems and Discussion
114(1)
Further Reading
114(1)
Chapter 9 Amplicon Databases: Bold and Bacterial 16S rDNA Databases
115(18)
DNA Barcoding and the BoLD Database
115(6)
DNA barcoding
115(1)
Taxonomy and speciation studies involve the species delimitation
116(1)
DNA taxonomy and DNA barcoding
116(1)
Character-based or distance-based approaches to DNA barcoding result in identification of species
117(1)
Is there enough information in a single gene to do DNA barcoding?
118(1)
Potential new species are flagged by DNA barcoding
119(1)
The BOLD Repository
120(1)
Ribosomal RNA Databases
121(7)
Amplicon sequencing, microbiomes, metagenomics, and eDNA
121(3)
Databases are used to identify the species in a microbiome, metagenome, and eDNA sample
124(1)
Classifiers for identifying microbial species in eDNA, microbiome studies, and metagenomics
125(3)
Summary
128(1)
Recommendations for Students
128(1)
Discussion Questions
128(1)
Further Reading
129(4)
Section III Phylogenetic/Phylogenomic Analysis
Chapter 10 Introduction to Tree Building
133(14)
Phylogenetic Tree Building Overview
133(1)
Which phylogenetic method should be used?
134(3)
The number of trees grows with each additional taxon
134(1)
Trees can be rooted by several methods
135(2)
Characters and Weighting
137(1)
Character states in molecular data may include the presence of genes and the sequence of nucleotides or amino acids
137(3)
Some discrete and numerical character states are ordered
137(1)
Characters can be weighted relative to one another
137(1)
Which characters should be used?
138(1)
A matrix for demonstrating phylogenetic analysis
139(1)
Basics of Parsimony Analysis
140(4)
Fitch's algorithm uses set theory
141(3)
Rescoring characters
144(1)
Summary
144(1)
Recommendations for Students
145(1)
Discussion Questions
145(1)
Further Reading
145(2)
Chapter 11 Distance and Clustering
147(10)
Corrections for Multiple Hits May Be Introduced
149(1)
Corrections Using Evolutionary Models
149(5)
Neighbor joining is a stepwise-based approach to tree-building
151(3)
Minimum Evolution Uses Minimal Distance as a Criterion to Choose the Best Solution among Multiple Trees
154(1)
Summary
155(1)
Recommendations for students
156(1)
Discussion Questions
156(1)
Further Reading
156(1)
Chapter 12 Maximum Likelihood
157(10)
Transformation and Probability Matrices
161(3)
Character weighting schemes
161(1)
Likelihood analysis incorporate probability matrices
162(2)
Summary
164(1)
Recommendations for students
164(1)
Discussion Questions
164(1)
Further Reading
165(2)
Chapter 13 Search Strategies and Robustness
167(10)
So Many Trees, so Little Time
167(4)
Tree space basics
167(2)
Selection of a starting tree
169(1)
Peaks in tree space can be reached by branch swapping
169(1)
Moving from local optimality peaks to peaks with higher optimality
170(1)
Robustness of Phylogenetic Trees
171(1)
Bremer Support Estimates Robustness of a Node
171(3)
Resampling to Determine Node Robustness
172(1)
Bootstrapping assesses node robustness by resampling with replacement
172(2)
Jackknifing assesses node robustness by resampling without replacement
174(1)
Parametric bootstrapping applies a distribution model to the data
174(1)
Resampling Gene Partitions
174(1)
Summary
175(1)
Recommendations for Students
176(1)
Discussion Questions
176(1)
Further Reading
176(1)
Chapter 14 Rate Heterogeneity, Long Branch Attraction, and Likelihood Models
177(10)
Long Branch Attraction
177(1)
Rate Heterogeneity
178(5)
Rate heterogeneity and invariant sites (I)
178(1)
Rate heterogeneity and the gamma distribution (rorG)
179(1)
Combining the invariant-sites parameter and a gamma distribution
180(1)
Other methods accommodating rate heterogeneity
180(3)
Comparing Likelihood Models
183(1)
Programs can compare models
183(1)
Summary
184(1)
Recommendations for Students
185(1)
Discussion Questions
185(1)
Further Reading
185(2)
Chapter 15 Bayesian Approaches in Phylogenetics
187(12)
Bayesian inference
187(6)
Generating a distribution of trees is an important application of the Bayesian approach
189(2)
What do we need from a Bayesian phylogenetic analysis?
191(1)
MCMC is critical to the success of Bayesian analysis
192(1)
Bayesian Parameters in a Phylogenetic Context
193(4)
Model selection can be utilized on any biologically meaningful partition
194(1)
Selection of priors
194(1)
More MCMC generations improves results at an increased computational cost
195(1)
Assessing the efficiency of a Bayesian phylogenetic analysis
196(1)
Interpreting posterior probabilities of clades
196(1)
Summary
197(1)
Recommendations for Students
197(1)
Discussion Questions
198(1)
Further Reading
198(1)
Chapter 16 Incongruence of Gene Trees
199(14)
Incongruence of Trees
199(5)
Taxonomic congruence via supertrees
200(1)
Character congruence via total evidence supermatrices
201(1)
Assessments of incongruence can help decide what to concatenate
202(1)
The incongruence length difference test
202(2)
Likelihood tests for incongruence
204(1)
Fork indices provide measures of tree similarity
204(1)
Robinson-Foulds Metric and Subtree Prune-and-Regraft Distance (SPR distance)
204(2)
The Gene Tree/Species Tree Problem
206(2)
Examples of incomplete lineage sorting in closely related taxa
207(1)
Coalescence and the gene tree/species tree problem
208(1)
Horizontal Transfer
208(1)
Programs That Consider Nonvertical Evolution and Incomplete Lineage Sorting to Infer Phytogeny
209(2)
Coalescence programs use both gene trees and species trees as input
209(1)
Programs that consider horizontal gene transfer generate nets and webs
210(1)
Summary
211(1)
Recommendations for students
211(1)
Discussion Questions
211(1)
Further Reading
212(1)
Chapter 17 Phylogenetic Programs and Websites
213(12)
Website Summaries of Programs
213(1)
The Classics
214(4)
Likelihood programs
216(1)
Bayesian phylogenetic inference programs
217(1)
Parsimony programs
217(1)
Networks
218(1)
The Comparative Method
218(1)
Tree Visualization Programs
218(1)
All-Purpose Websites and Software Companies
218(3)
Programming Languages and Packages
221(1)
Summary
221(1)
Recommendations for Students
221(1)
Discussion Questions
221(1)
Further Reading
221(4)
Section IV Population Genomics
Chapter 18 Population Genetics and Genomes
225(20)
High-Throughput Methods and Population Genetics
225(1)
Kimura and Lewontin contributed important new ways to think about genes in nature
225(1)
The Hardy-Weinberg theorem has been extended in modern population genetics
226(1)
DNA Variation among individuals
226(5)
Single-nucleotide polymorphisms (SNPs)
227(1)
Microsatellites provide another analytical tool for species where SNPs are less abundant
228(2)
RAD markers are a source of data for modern population genomics
230(1)
Extending Basic Population Genetics to DNA Sequences
231(4)
Tajima's D distinguishes between sequences evolving neutrally and those evolving non-neutrally using allele frequencies
231(1)
F statistics measure the degree of isolation of entities
232(1)
There are two approaches to estimating population-level statistics
233(2)
FST and related measures have four major uses in evolutionary biology
235(1)
Imputation
235(1)
Population-Level Techniques: Mismatch Distribution Analysis, STRUCTURE Analysis, Principle Components Analysis, and Analysis Platforms
236(7)
Mismatch distribution analysis compares haplotype data of populations
237(1)
Structure analysis reveals substructure and genetic cross talk
237(1)
Principle components and genomic data
238(3)
Population genomics analysis platforms
241(2)
Summary
243(1)
Discussion Questions
243(1)
Further Reading
244(1)
Chapter 19 Population Genomics Approaches
245(18)
Genome-Wide Association Studies
245(2)
A simple example illustrates the association technique
246(1)
The National Human Genomics Research Institute maintains a database of genome-wide association studies
247(1)
Programs That Can Perform GWAS Analyses
247(1)
Role of the Coalescent in Population Genetics
247(4)
The coalescent addresses the time for an allele to coalesce and the variation in populations under drift
248(1)
The coalescent in practice explores a large number and a broadly representative sample of plausible genealogical scenarios
248(1)
High-quality DNA sequence data from a random sample constitute the best input for a coalescence analysis
249(1)
Importance sampling and correlated sampling are used to generate a collection of simulated genealogies
249(1)
Programs for coalescence analysis include BEAST and Lamarc
250(1)
Genetic Hitchhiking and Selective Sweeps
251(9)
Selective sweeps are detected in four basic ways
252(2)
Empirical examples of selective sweeps include boxers, flies, and humans
254(1)
Hard and soft sweeps produce different effects in the genome
255(1)
Genome-wide scans to address population genetic and evolutionary questions
256(2)
Phylogenetic shadowing identifies regulatory elements in DNA sequences
258(1)
Regions of the human genome experience accelerated evolution
258(1)
Regions that are both strongly conserved and rapidly deleted are of interest
259(1)
Summary
260(1)
Discussion Questions
260(1)
Further Reading
261(2)
Chapter 20 Detecting Natural Selection: The Basics
263(14)
Analyzing DNA Sequences for Natural Selection
263(12)
DNA sequences can be examined for silent and replacement changes
263(2)
Several variables affect the detection of natural selection at the genomic level
265(2)
Approximate methods of determining dN/dS
267(1)
Basic dN and dS calculations begin with counting the observed number of changes
268(1)
Scaling for redundancy and getting the number of potential substitutions is necessary for determining dN/dS
268(3)
Pathways of codon change are an important element in calculating dN/dS
271(2)
Codon change pathways can be used to account for redundancy
273(2)
Summary
275(1)
Recommendations for Students
275(1)
Discussion Questions
275(1)
Further Reading
276(1)
Chapter 21 Refining the Approach to Natural Selection at the Molecular Level
277(18)
Accounting for Multiple Hits in DNA Sequences for dN/ds Measures
277(1)
The Jukes-Cantor conversion corrects for multiple hits
277(1)
Estimating Natural Selection Requires Adjusting the Calculation of Sequence Changes
278(1)
Expanding the Search for Natural Selection at the Molecular Level
278(8)
Statistical tests of significance are required at various levels
278(1)
Species 1
279(1)
Species 2
279(1)
Natural selection is variable across protein components and across time
280(1)
Examples of nonuniformity are seen in Drosophila and in the BRCA1 gene
280(2)
Maximum likelihood approaches are implemented in selection studies at the molecular level
282(1)
Statistical tests using dN and dS
283(1)
There are caveats when detecting selection at the molecular level
284(1)
Transcriptomics and whole genome sequencing has opened the way for searches for natural selection at an unprecedented level
285(1)
Codon Selection Bias
286(4)
Codon selection bias can be calculated manually or by various analytical methods
286(2)
Codon usage bias usually occurs in cellular housekeeping genes and varies among species
288(2)
Summary
290(1)
Recommendations for Students
290(1)
Discussion Questions
290(1)
Further Reading
291(4)
Section V Phylogenomics in Action
Chapter 22 Constructing Phylogenomic Matrices
295(16)
Why Choose the Programs We Focus On?
295(2)
Formatting matrices for population genomics analysis
295(2)
Formatting Arlequin Files
297(1)
Formatting STRUCTURE Files
297(1)
Formatting HYPHY Files
297(2)
Formatting PAML Files
299(1)
Formatting PLINK Files
300(1)
Constructing phylogenomic matrices
300(1)
Determining Orthology and Constructing Individual Gene Matrices
301(1)
Concatenating individual Gene Alignments
302(1)
Partitions and Partitioning
302(1)
Formatting Partitions in PAUP* and MrBayes (NEXUS)
302(3)
Formatting Partitions in PHYLIP
305(1)
Formatting Partitions in RaxML and IQtree
305(1)
Formatting Partitions in TNT
305(1)
Summary
306(1)
Web-Based Programs for Formatting Phylogenomic Matrices
306(1)
Recommendations for Students
307(1)
Discussion Questions
307(1)
Further Reading
308(3)
Chapter 23 Phylogenomics and the Tree of Life
311(20)
Problems with Phylogenomic Studies
312(1)
Supertrees or Supermatrices
313(3)
Grafting supertree approach
314(1)
Matrix representation approach
315(1)
Divide-and-conquer approach
316(1)
Examples of Phylogenomic Studies
316(10)
Shallow targeted sequencing of over 70,000 eukaryotes recovers major eukaryotic groups
316(1)
Whole genome microbial phylogenomics
317(1)
Specific problems in bacterial phylogenomics
317(2)
Does a tree of life really exist for bacteria?
319(1)
Microbial Trees of Life
319(1)
The deep relationships of Metazoa
320(2)
Green phylogenies
322(4)
Yeast and Drosophila Represent Examples of Concatenation and Lineage Sorting Problems in Phylogenomics
326(2)
Coalescence Can Partially Solve the Problem of Incongruence
328(1)
Summary
328(1)
Recommendations for Students
328(1)
Discussion Questions
329(1)
Further Reading
329(2)
Chapter 24 Comparative Genomics
331(16)
Characterizing Genomes by Orthology
331(6)
Clusters of orthologous groups is a method that enables identification of orthologs of genes across multiple species
332(1)
Single linkage clustering compares genes in a cross-species context based on sequence
332(1)
A presence/absence matrix is constructed via single linkage clustering
333(4)
Comparative Genomics Approaches
337(6)
Venn diagrams, EDGAR, and Sungear visualize the overlap of genes from two or more genomes
337(2)
The pangenome
339(1)
Genome content analysis was first accomplished for bacterial genomes
339(2)
Caveats with genome content analysis in phylogenetic analysis
341(1)
Using genome content in evolutionary studies
342(1)
Summary
343(1)
Recommendations for students
343(1)
Discussion Questions
344(1)
Further Reading
344(3)
Chapter 25 Environmental DNA (eDNA)
347(12)
Any Environment Can Be Examined for Its Microbial Makeup
347(1)
Amplicon Sequencing, Microbiomes, Metagenomics, and eDNA
348(8)
The next-generation approach
349(1)
Data management---format
350(1)
Data management---processing
350(1)
Data management storage
350(2)
Shotgun sequencing
352(1)
Software
352(1)
Making ecological/environmental inferences
352(3)
Caveats and recommendations
355(1)
Summary
356(1)
Recommendations for Students
356(1)
Discussion Questions
357(1)
Further Reading
357(2)
Chapter 26 Phylogenomic Approaches to Understanding Gene Function and Evolution
359(14)
Transcription-Based Approaches
360(4)
Transcriptomics is used for class comparison, prediction, and discovery
360(1)
Data are transformed for use in dendrograms and other clustering techniques
361(1)
Specific next-generation sequencing approaches applied to transcriptome analysis
362(1)
Transcriptomic approaches are useful in evolutionary and phylogenomic studies
363(1)
Protein--Protein Interactions
364(7)
Generating data for protein--protein interaction research
364(1)
2H screening
364(1)
PCA screening
365(1)
Computational methods for examining protein--protein interactions
365(1)
Model organism gene and protein function can be studied by Web-based approaches like ENCODE
365(1)
Functional phylogenomics employs common ancestry to infer protein function
366(1)
Phylogenomic gene partitioning can be used to explore function
367(1)
A gene presence/absence matrix was employed to examine evolution in the major metazoan lineages
367(1)
Transcript sequences and phylogeny can be used to study plant function
368(1)
Gene function clustering in Caenorhabditis elegans from RNA interference phenotypes
368(1)
Gene ontology facilitates the comparison of genes
369(2)
Summary
371(1)
Recommendations for Students
372(1)
Discussion Questions
372(1)
Further Reading
372(1)
Index 373
Rob DeSalle is Curator at the Sackler Institute of Comparative Genomics in the Division of invertebrate Zoology at the American Museum of Natural History. DeSalle works in molecular systematics, microbial evolution, and genomics. His current research concerns the development of bioinformatic tools to handle large-scale genomics problems using phylogenetic systematic approaches. Dr. DeSalle has worked closely with colleagues from Cold Spring Harbor Labs, New York University, and the New York Botanical Garden on seed plant genomics and development of tools to establish gene family membership on a genome- wide scale. His group also focuses on microbial genomics, taxonomy, and systematics. In particular, they approach tree-of-life questions concerning microbial life using whole genome information.



Jeffrey Rosenfeld is Assistant Professor for Pathology and Laboratory Medicine and the Manager of the Biomedical Informatics Shared Resource at the Rutgers Cancer institute. His research focuses on the use of new genomics technologies to investigate previously unsolvable problems. He is currently working with long-read and single-cell sequencing. Dr. Rosenfeld also has an appointment as a Research Associate at the American Museum of Natural History where he works on whole-genome phylogenetics. With collaborators at the Museum, he has sequenced and assembled the genomes of non-model insects.



Michael Tessler is Adjunct Faculty in Ecology at Sterling College. He received his PhD from the Richard Gilder Graduate School, American Museum of Natural History. His research explores the evolution and ecology of overlooked organisms and includes phylogenetic research on terrestrial leeches, combining his collections from China and Cambodia with AMNHs legacy collections to produce a phylogenetic revision of all terrestrial leech groups. His dissertation focused on the evolution of leech anticoagulants and on how leeches process difficult to digest blood such as urea-packed shark blood, and the ways anticoagulants evolved in leech lineages that no longer drink blood and instead eat invertebrates.