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Andrew V Z Brower

Andrew V Z Brower
USDA; AMNH; USNM · National Identification Services

Ph.D. Cornell University, 1994

About

265
Publications
77,700
Reads
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8,383
Citations
Citations since 2017
30 Research Items
2238 Citations
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Introduction
Assistant Director, National Identification Service, USDA-APHIS. I supervise the national entomological taxonomists, who provide authoritative identifications of insects intercepted on cargo and goods at U.S. ports of entry. I am a molecular systematist of butterflies, with an interest in theory of biological systematics and speciation. I formerly taught at Oregon State University and Middle Tennessee State University. I also have substantial experience in insect collection curation.
Additional affiliations
October 2018 - present
United States Department of Agriculture
Position
  • Managing Director
Description
  • I work with a team of systematists to provide prompt, authoritative identifications and make regulatory decisions about insect pests imported into the U.S. We keep the country's food and natural resources safe from exotic pests.
October 2018 - May 2021
USDA; AMNH; USNM
Position
  • Managing Director
August 2006 - October 2018
Middle Tennessee State University
Position
  • Professor
Description
  • Molecular systematics of nymphalid butterflies
Education
September 1987 - October 1993
Cornell University
Field of study
  • Ecology and Systematics
September 1985 - May 1987
Yale School of Forestry and Environmental Studies
Field of study
  • conservation biology and museum science
September 1981 - May 1985
Yale College
Field of study
  • Biology

Publications

Publications (265)
Article
Full-text available
The claim that parsimony can be statistically inconsistent remains the chief criticism of the cladistic approach, and also the main justification for alternative model-based approaches such as maximum likelihood and Bayesian inference. Despite its refutation in the 1980s, this persistent myth of parsimony's Achilles’ heel is entrenched in the prima...
Article
Full-text available
This paper reviews the ontogeny of pattern cladistics from the 1970s and 1980s, rebuts criticisms by contemporary anti‐cladists, and endeavours to clarify persistent misunderstandings about the philosophical foundations of the approach.
Article
This is a review of the version 6.0 of the "PhyloCode", with comments on the historical development of phylogenetic nomenclature over the past 30 years.
Article
This essay provides an alternative perspective on the history of classification, species and subspecies concepts to that presented in a recent review by Padial and de la Riva. Central points of difference relate to Darwin’s role in revolutionizing classification, Willi Hennig’s role in the development of the phylogenetic species concept, the metaph...
Conference Paper
Full-text available
A lecture I presented to the Systematics Association, 27 Nov. 2019. The essay is published online at https://systass.org/meetings/past-events/systematics-and-parsimony/
Article
This is a review of four recent philosophy of science books. A main point is the centrality of peer review ("intersubjective corroboration") for the legitimization of scientific knowledge.
Book
This is a graduate-level textbook on systematics, with an emphasis on the cladistic approach. However, most of the book is about character coding, classification, nomenclature and other aspects that are not method-specific, so it should be useful for anyone who wants to be well-grounded in systematic theory (from a biological perspective, rather th...
Article
a book review of "Phylogenetic Ecology" by Nathan Swenson.
Chapter
The exuberant diversity present in the Neotropics naturally leads to questions about its age of origin. The butterfly fossil record is so depauperate that most of what we know today about the age of butterflies is based on relaxed clock estimates. Here we compile relaxed clock-based crown age estimates for clades of Neotropical butterflies from 6 f...
Article
Full-text available
A book review of Tandy Warnow's "Computational Phylogenetics", critically assessing the schism between mathematical and empirical approaches to phylogenetic inference.
Article
Full-text available
This is a short biographical essay written for the online journal Inference, International Review of Science. It addresses the life and ongoing influence of Willi Hennig, founder of phylogenetic systematics and originator of words such as synapomorphy. The PDF posted here was downloaded from https://inference-review.com/article/a-quiet-revolution...
Article
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A book review of Michael Arnold's most recent. The epistemological framework of systematics is defended against the reticulistic world view.
Article
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a book review of two recent compendia of previously published historical lectures and publications by A. M. Marshall and A. W. F. Edwards, respectively . The latter is of particular interest as a developer of likelihood methods in phylogenetic inference.
Article
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A book review of David Quammen's "Tangled Trees". The main point is that phylogenetic trees are metaphors that allow a clear hierarchical representation of relationships among taxa and a framework for classification, not pictures of the process of evolution.
Article
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Background: Butterflies (Papilionoidea) are perhaps the most charismatic insect lineage, yet phylogenetic relationships among them remain incompletely studied and controversial. This is especially true for skippers (Hesperiidae), one of the most species-rich and poorly studied butterfly families. Methods: To infer a robust phylogenomic hypothesi...
Article
Full-text available
Mallet et al. (2007 BMC Evolutionary Biology, 7, 28) employed a database of putative interspecific hybrid specimens of the genus Heliconius to advance a hypothesis of "the species boundary as a continuum." Here, each of those specimens, as well as subsequently documented specimens, is individually reassessed regarding its phenotype, potential paren...
Article
Full-text available
Bootstrap and other percentage‐based measures of nodal “support” cannot distinguish between strong support and very strong support. Clearly, if there is no upper bound to the number of potential characters, there should be no upper bound to measures of support.
Article
A book review that addresses strengths and weaknesses of this 2017 book.
Article
Quinn (Biol Philos 32:581–598, 2017) offered seven definitions of “cladist” and discussed the context in which they are used in relation to historical and current debates in systematics. As a member of her study taxon, I offer some contextual color commentary, clarifications on the views of “pattern cladists” regarding monophyly, ancestors, synapom...
Article
Hamadryas februa Hübner and close relatives H. amphichloe Boisduval, H. glauconome Bates and H. julitta Fruhstorfer are a homogeneous group of taxa loosely defined as different species mainly by a combination of wing color characters. Here we explored the utility of the mtDNA COI barcode region in delimiting these taxa and examined its overall perf...
Article
Full-text available
This is a book review of Olivier Rieppel's "Phylogenetic Systematics from Haeckel to Hennig." It discusses the development of systematics in the German-speaking part of Europe during the century 1870-1970 (more or less), including the Nazi impact on science.
Article
Full-text available
Heliconius butterflies have become a model for the study of speciation with gene flow. For adaptive introgression to take place, there must be incomplete barriers to gene exchange that allow interspecific hybridization and multiple generations of backcrossing. The recent publication of estimates of individual components of reproductive isolation be...
Article
Full-text available
The problem of ‘rogue taxa’ explored by Buenaventura et al. (2017, Cladistics 33: 109–133) is due to complementary patterns of missing data in their matrix, which are evident from perusal of their supplementary table S1.
Article
Full-text available
Kozak et al. (2015) portrayed the inference of evolutionary history among Heliconius and allied butterfly genera as a particularly difficult problem for systematics due to prevalent gene conflict caused by interspecific reticulation. To control for this, Kozak et al. conducted a series of multispecies coalescent phylogenetic analyses that they clai...
Article
Full-text available
A book review of Elliot Sober's "Ockham's Razors: a user's manual" - a book about the principle of parsimony. Among other things, the review questions the relevance of the Smith-Quackdoodle theorem for empirical systematics.
Article
Full-text available
Despite multiple attempts to infer the higher-level phylogenetic relationships of skipper butterflies (Family Hesperiidae), uncertainties in the deep clade relationships persist. The most recent phylogenetic analysis included fewer than 30% of known genera and data from three gene markers. Here we reconstruct the higher-level relationships with a r...
Data
Saturation curve of 3rd codon positions on F84 sequence distance. Transitions: s; blue cross and transversions: v; green triangle. For all genes, the curve saturate at higher sequence divergence or when the genetic distances between sequences increase. (The saturation curves for CAD and GAPDH are not shown because of consistent technical error duri...
Data
Newly designed primer pairs for an IDH (Isocitrate dehydrogenase) amplicon in Hesperiidae. This primer pair, along with attached universal tail (T7 promoter and T3) were tested at annealing temperature 55 °C.
Data
ML trees from concatenated and combined nuclear dataset under different partition scheme or model of evolution. ML trees from the analyses of concatenated dataset with the partitions from PartitionFinder (A) and with TIGER partitions (B). ML trees from the analyses of combined nuclear dataset with the partitions from PartitionFinder (C) and with TI...
Data
The maximum likelihood tree from mitogenome analysis. The maximum likelihood tree from the analyses of 13 protein-coding mitochondrial genes of 6 hesperiids using codon-based partitions. The node supports were derived from 100 bootstrap analysis.
Data
Analyses of gene clusters. Trees from the ML analyses of different gene combinations (A) ArgKin, EF1a, GAPDH, RpS2, RpS5, wingless, (B) CAD, EF1a, IDH, wingless, (C) ArgKin, GAPDH, MDH, RpS2, RpS5 and (D) CAD, MDH using gene partitions. The node supports were derived from 1,000 bootstrap trees. The gene combinations were made based on the relations...
Data
Summarization of partitioned bremer support. (A) The node specific partitioned bremer support for ten-gene partitions. (B) The positions of the nodes are shown on the consensus tree reconstructed using concatenated dataset.
Data
Cross representation of bootstrap support. (A) Summarization of the ML-bootstrap trees from the concatenated dataset on the ML-best tree from the combined nuclear dataset. (B) Summarization of the ML-bootstrap trees from the combined nuclear dataset on the ML-best tree from the concatenated dataset.
Data
Summarization of tree properties at different proportion of missing data. The bootstrap support (BS) and branch length for the deep nodes from the ML trees reconstructed using gene partitions for different datasets. The dataset1 is same as the concatenated dataset (> 80% missing data per taxa allowed) and the other datasets were generated after rem...
Data
New taxonomic positions of few genera. List of genera for which we observed new taxonomic positions or ambiguity in taxonomic position. The taxonomic positions of rest of the sampled genera follow the list supplied with Warren, Ogawa & Brower (2009).
Data
Bayesian tree from concatenated dataset. The analysis was performed on the concatenated dataset with TIGER partitions under reversible jump MCMC. The values at nodes represent posterior probabilities. The topology in this tree is similar to that of Fig. 1B except that Euschemoninae and Eudaminae are sisters.
Data
Proportion of contrasting tree topologies across datasets. The proportion of trees, that show either topology 1 (as in Fig. 1A) or topology 2 (as in Fig. 1B), in the set of 105 or 108 independent ML trees from the analyses of different datasets as mentioned in Table S2. The number on the top of the bar represents the number of ingroup taxa in the r...
Data
Concatenated aligned sequence file. The gene partitions in this concatenated aligned sequence file is: ArgKin = 1-596, CAD = 597-1446, COI = 1447-2921, EF1A = 2922-416, GAPDH = 4162-4852, IDH = 4853-5562, MDH = 5563-6298, RPS2 = 6299-6709, RPS5 = 6710-7326, WGL = 7327-7726.
Data
List of sequences of length less than 200 bp in fasta format. The definition of each sequence include the name of organism, the specimen voucher, gene name, sequence length and the starting position of codon.
Chapter
Willi Hennig (1913–76), founder of phylogenetic systematics, revolutionised our understanding of the relationships among species and their natural classification. An expert on Diptera and fossil insects, Hennig's ideas were applicable to all organisms. He wrote about the science of taxonomy or systematics, refining and promoting discussion of the p...
Article
Phylogenetic analysis of the south-temperate members of Pronophilina (Nymphalidae: Satyrinae) using mitochondrial and nuclear gene data corroborated monophyly of the clade and provided a framework for its systematic revision based on morphology. Of the 19 genera, 8 have been synonymized with 16 new combinations: Cosmosatyrus stelligera n. comb.,C....
Article
I offer comments on two recent articles in The American Biology Teacher by Davenport and colleagues addressing the interpretation and construction of phylogenetic trees. The "tree-thinking" literature suggests that students need to acquire a clear understanding of the meaning of phylogenetic tree diagrams. To this end, I provide clarifications of t...
Article
Full-text available
A book review of Gunther Wagner's treatise on developmental homology. The main point is that like systematics, developmental biology involves epistemological issues that are not evident from a reductionist perspective.
Article
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The origins and meanings of "cladogram" are reviewed. Traditionally, "cladogram" has been defined as a graphical representation of an empirical hypothesis of relationships among taxa, based on evidence from synapomorphies alone. Disturbingly, numerous recent authors treat "cladogram" as synonymous with "dendrogram" and do not appreciate the particu...
Article
Full-text available
The recent publication of a time-tree for the plant family Solanaceae (nightshades) provides the opportunity to use independent calibrations to test divergence times previously inferred for the diverse Neotropical butterfly tribe Ithomiini. Ithomiini includes clades that are obligate herbivores of Solanaceae, with some genera feeding on only one ge...
Article
Full-text available
Article
Evidence from butterflies and birds suggests that most extant Amazonian species arose during the Pleistocene ( Keywords: Amazonia; Neogene; Neotropics; Pleistocene; Quaternary; crown ages; refuge hypothesis; tropical diversity Document Type: Correspondence DOI: http://dx.doi.org/10.1111/jbi.12539 Publication date: July 1, 2015 $(document).ready(fun...
Article
Full-text available
This is a book review of Richard Zander's "A framework for post-phylogenetic systematics"
Article
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This is a book review of "the Nature of Classification" by John S. Wilkins and Malte Ebach.
Article
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Morphological and molecular data from two prior publications are combined with new data resulting in a mostly well-supported cladistic hypothesis of relationships among subtribes and genera of Ithomiini, a diverse clade of Neotropical butterflies. A revised classification for relationships among the nine tribes and 46 genera comprising the group is...
Article
Rejection of the Pleistocene refugium hypothesis (PRH) as an explanation for the high biodiversity of Neotropical forest is based in part on the assertion that biotic elements of these forests evolved during the Neogene. That argument is justified, in turn, by the ages of crown groups (the age of the most recent common ancestor of extant species of...
Article
Full-text available
Usages and meanings of the terms “taxic” and “transformational homology” are reviewed from 1982 to the present. While “taxic homology” has been relatively invariant in its connotation, “transformational homology” as employed by different authors refers to at least three different concepts. This has resulted in confusion.
Article
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Article
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This is a book review of a Willi Hennig biography by Michael Schmitt.
Article
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In light of recent terminological controversy, this article reviews cladistic conceptions of character states coded as absences, symplesiomorphies, and secondary losses. The first section addresses absence as a question of ontology vs. epistemology. The second and third sections address the evidentiary status of symplesiomorphy in cladistics, the f...
Article
Full-text available
The diverse Müllerian mimetic wing patterns of neotropical Heliconius (Nymphalidae) have been proposed to be not only aposematic signals to potential predators, but also intra- and interspecific recognition signals that allow the butterflies to maintain their specific identities, and which perhaps drive the process of speciation, as well. Adaptive...
Article
Review of Pietsch, "Trees of Life"
Article
A recent review of the homology concept in cladistics is critiqued in light of the historical literature. Homology as a notion relevant to the recognition of clades remains equivalent to synapomorphy. Some symplesiomorphies are “homologies” inasmuch as they represent synapomorphies of more inclusive taxa; others are complementary character states t...
Article
This paper provides a history of the use of the term "phenetic" to clarify current usage. Phenetic analyses convert character data into a matrix of pairwise distances among the study taxa that is used to infer a hypothesis of relatedness based on overall similarity.
Article
To investigate the utility of a region of wingless (wg) as a marker for molecular systematics, we compared wg sequences to mitochondrial COII sequences from twenty-two nymphalid butter¯y taxa and one outgroup. Compositional characteristics of the two gene regions are compared, and their contributions to a cladogram inferred from the combined data s...
Article
Full-text available
Ten species of the Astraptes fulgerator complex are named and diagnosed based on their mitochondrial ‘DNA barcode’ sequences. A rationale for this rather unorthodox approach to species description is offered, and some philosophical questions about whether or not such descriptions are desirable are addressed. It is emphasized that while these descri...
Article
The use of model-based methods to infer a phylogenetic tree from a given data set is frequently motivated by the truism that under certain circumstances the parsimony approach (MP) may produce incorrect topologies, while explicit model-based approaches are believed to avoid this problem. In the realm of empirical data from actual taxa, it is not kn...
Article
Full-text available
The evidence supporting the recent hypothesis of a homoploid hybrid origin for the butterfly species Heliconius heurippa is evaluated. Data from selective breeding experiments, mate-choice studies, and a wide variety of DNA markers are reviewed, and an alternative hypothesis for the origin of the species and its close relatives is proposed. A scena...
Article
Full-text available
Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index s...
Article
Full-text available
Case 3492 Heliconius tristero Brower, 1996 and Heliconius melpomene mocoa Brower, 1996 (Lepidoptera, NYMPHALIDAE): proposed conservation by suppression of Heliconius melpomene bellula Brown, 1979 Abstract. The purpose of this application, under Articles 23.9.3 and 81.2.3 of the Code, is to conserve the species-group names Heliconius tristero Brower...
Article
Full-text available
The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars...
Article
Abstract We propose a revised higher classification for the genera of Hesperiidae (skipper butterflies) of the world. We have augmented our published DNA data matrix with 49 morphological characters in order to infer relationships for taxa not sampled in the molecular study. We use the results of a combined analysis to identify morphological synapo...
Article
Full-text available
Global biodiversity peaks in the tropical forests of the Andes, a striking geological feature that has likely been instrumental in generating biodiversity by providing opportunities for both vicariant and ecological speciation. However, the role of these mountains in the diversification of insects, which dominate biodiversity, has been poorly explo...
Article
Full-text available
To test prior results with a more sensitive technique and larger sample sizes, we assessed genetic diversity among far-flung monarch butterfly, Danaus plexippus (L.) (Lepidoptera: Nymphalidae), populations from North and South America by using mitochondrial DNA sequences. Our new data corroborate the previously documented close genetic similarity a...
Article
Full-text available
Rapidly evolving nuclear coding sequences are highly desirable for phylogenetic studies of closely related species. Here, we investigated an 807-bp region, homologous to the testis-specific Tektin gene from Bombyx mori (L.), in 34 nymphalid butterfly taxa in the subfamilies Ithomiinae, Danainae, and Heliconiinae. Within Ithomiinae, relationships in...
Book
This is a textbook, published by Cornell University Press. As such, I am not at liberty to distribute electronic copies. Note that a third edition is in preparation as of 2019.
Article
Primer pairs were designed and protocols developed to selectively amplify segments of vertebrate mitochondrial cytochrome oxidase subunit 1 (COI) and cytochrome b (Cyt b) mtDNA from the bloodmeals of mosquitoes (Diptera: Culicidae). The protocols use two pairs of nested COI primers and one pair of Cyt b primers to amplify short segments of DNA. Res...
Article
A comprehensive tribal-level classification for the world’s subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I-subunit II, elongation factor-1α and wingless). Monophyl...
Article
Full-text available
DNA 'barcoding' relies on a short fragment of mitochondrial DNA to infer identification of specimens. The method depends on genetic diversity being markedly lower within than between species. Closely related species are most likely to share genetic variation in communities where speciation rates are rapid and effective population sizes are large, s...
Article
Full-text available
Butterfly long-wavelength (L) photopigments are interesting for comparative studies of adaptive evolution because of the tremendous phenotypic variation that exists in their wavelength of peak absorbance (lambda(max) value). Here we present a comprehensive survey of L photopigment variation by measuring lambda(max) in 12 nymphalid and 1 riodinid sp...
Article
Full-text available
Phylogenetic information is useful in understanding the evolutionary history of adaptive traits. Here, we present a well-resolved phylogenetic hypothesis for Heliconius butterflies and related genera. We use this tree to investigate the evolution of three traits, pollen feeding, pupal-mating behaviour and larval gregariousness. Phylogenetic relatio...

Questions

Questions (7)
Question
I have been examining some UCE datasets deposited in Treebase, and the alignments, as far as I can see, are very poor, with floating small fragments that are obviously "aligned" at random, and clear mismatches in gaps to which more parsimonious alignments are readily apparent. This seems unacceptable to me, but these datasets are published, so I guess editors and reviewers think this is not a problem. Is it? Should it be?

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Projects

Projects (11)
Archived project
The object of this work was to test the hypothesis that eastern and western North American populations of monarch butterflies are phylogenetically distinct from one another, based on mtDNA. We did not find any evidence to support that. However, this does not mean that the populations are not ecologically distinct, and thus our comments about the dangers of monarch transfer.
Project
Work mostly done in collaboration with former student Andy Warren and colleagues. The goal is to generate a stable tree to support the higher-level classification of Hesperiidae.