REGIONAL AND NATIONAL TRENDS IN LEPIDOPTERA COLLECTING IN THE UNITED 

STATES SINCE 1800 

 

By 

 

Erica E. Fischer 

 
 
 
 
 
 
 
 
 
 
 
 
 

A THESIS 

 

Submitted to 

Michigan State University 

in partial fulfillment of the requirements  

for the degree of 

 

Entomology—Master of Science 

 
 
 
 
 

 
 
 
 
 
 
 
 
 
 
 
 
 
 

REGIONAL AND NATIONAL TRENDS IN LEPIDOPTERA COLLECTING IN THE UNITED 

ABSTRACT 

 

STATES SINCE 1800 

 

By 

 

Erica E. Fischer 

 

Though natural history collections began as private demonstrations of wealth and 

knowledge, they are now places of public learning and depositories of biodiversity.  Insects 

have been collected for their beauty and small size since the beginnings of natural history 

collections, making them ideal for studies of long-term collecting patterns. 

This thesis project characterizes collecting efforts focused on butterflies and moths 

within the United States, both at the institutional and national level.  The A. J. Cook 

Arthropod Research Collection (ARC) at Michigan State University contains 96,618 

databased Lepidoptera specimens, predominantly collected in the state of Michigan.  The 

ARC has a long history of involvement by avocational collectors, both in terms of specimen 

donation and curation.  Their contributions grew significantly in the 1950s.  However, the 

number of specimens added annually by both professionals and non-professionals has 

decreased greatly since 1970.  More than 1 million Lepidoptera specimens held in various 

US collections were also used to examine trends in Lepidoptera collecting nationally.  

Collecting has been inconsistent over time and markedly on the decline since the 1990s.  

States are not evenly represented in this dataset, mirroring the inconsistency seen in 

county representation among Michigan specimens in the ARC.  This uneven distribution is 

at least partially associated with a lack of funding for databasing and other areas of 

curatorial effort for insect collections, in addition to a decrease in natural history education 

in K-12 and undergraduate curriculums. 

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

 

For my best friend, Evan, for putting up with far too many  

panic texts at 3am and for always insisting that I could (and would!) do this. 

 
 

And for my dad, the best butterfly hunting partner,  

insect display case builder, and parent a person could ask for. 

 

iii 

ACKNOWLEDGEMENTS 

 
 
 

An immense amount of gratitude to my major professor, Dr. Anthony Cognato, for being 

unerringly supportive throughout the long process of me learning how coding works, 

getting things written, and just generally figuring out how to be a functioning student. 

 

Many additional thanks to the other members of my thesis committee: Dr. Amanda Lorenz-

Reaves, an amazing instructor to work with and for, and Dr. Barbara Lundrigan, whose 

commentary improved this document immensely. 

 

Thanks also to Jason Matlock, who provided a great number of explanations and coding 

basics in tidyverse and for illustrating advanced statistical concepts during overlapping TA 

office hours.  And thank you to Daniel Barnett for many hours of being a statistical 

sounding board. 

 

 

iv 

TABLE OF CONTENTS 

 

 
LIST OF TABLES …………………………………………………………………………………………………..….……vi 
 
LIST OF FIGURES ...……………………………………………………………………………..…………………….…..vii 
 
CHAPTER 1: INTRODUCTION TO THE HISTORY OF NATURAL HISTORY COLLECTING …......1 
“A View of the Whole”: Natural history collecting, 1500-2019 ..………………………....……......1 
 
1500-1600 …………………………………...………………………………………………………………….….1 
 
 
1600-1900………………………………………………...…………………………………………………….…..2 
 
 
 
 
1900-Present …...………………………………………………………………………………………..….…..10 
“Can learned skill on little specks display…”: Collecting and  
 
classifying insects, 1800-2019 ……………….…………………………………………………………….….13 
BIBLIOGRAPHY……………………...………………………………………………………………………………….….20 
 
CHAPTER 2: THE LEPIDOPTERA OF THE A.J. COOK ARTHROPOD RESEARCH COLLECTION 
AND THE IMPACT OF AVOCATIONAL COLLECTING ….……………………………...……………..……..26 
 
Introduction …………………………………………………………………………………………………….……26 
Methods ………………………………………………………………………………………………………….…….28 
 
Results …………………………………………………………………………………………………………….……29 
 
 
Discussion……………………………………………………………………………….……………………….……37 
BIBLIOGRAPHY ……………………………………………………………………………………………….……….….42 
 
CHAPTER 3: THE RISE AND FALL OF LEPIDOPTERA COLLECTING IN THE UNITED STATES 
SINCE 1800 ……………………………………………………………..………………………....…………………….….46 
Introduction ………………………………………………………………………………....……………...…….…46 
 
Methods ……………………………………………………………………….………………………………….……48 
Results ………………………………………………………………………………………………….………...….…50 
Discussion ………………………………………………………………………………………………………….…58 
BIBLIOGRAPHY ……………………………………………………………………………………….……….……....….65 
 
 
 

 

v 

LIST OF TABLES 

 
 
 

Table 2.1: Counts and percentages of specimens from represented Lepidoptera families ..30 
 
Table 2.2: Vocational vs. avocational collectors: comparison of specimen numbers by 
decade ..………………………………………………………………………………………………………..………………35 
 
Table 2.3: The top 10 avocational and vocational collectors represented in the ARC ...……..36 
 
Table 3.1: Counts and percentages of specimens from represented Lepidoptera families ..51 
 
Table 3.2: Avocational vs. vocational collectors: comparison of specimen numbers by 
decade ….………………………………………………………………………………………………………………….…..57 
 

 

vi 

LIST OF FIGURES 

 
 
 

Figure 2.1: Specimen collection events per year since 1800 in A. J. Cook Arthropod 
Research Collection.  The green line is a rolling decadal average, and the grey lines 
represent a single standard deviation from this average on a rolling decadal basis .………...30 
 
Figure 2.2: National distribution of Lepidoptera specimens in the A. J. Cook Arthropod 
Research Collection ………………………………………………………………………………………………….…..33 
 
Figure 2.3: Michigan distribution of Lepidoptera specimens in the A. J. Cook Arthropod 
Research Collection at the county level ………………………………………………………………..………..34 
 
Figure 2.4: Avocational and vocational specimen collection events ……………………….………..35 
 
Figure 3.1: Specimen collection events per year since 1800 in institutional collections 
across the United States.  The green line is a rolling decadal average, and the grey lines 
represent a single standard deviation from this average on a rolling decadal basis ...………51 
 
Figure 3.2: National distribution of Lepidoptera specimens held in institutional collections 
across the country normalized by state area in square kilometers.  States marked with a 
star have at least one institution participating in LepNet ……………………………………………….55 
 
Figure 3.3: Graph of Lepidoptera specimens held in institutional collections across the 
country normalized by state area in square kilometers ……………………………………………..…..56 
 
Figure 3.4: Avocational and vocational specimen collection events, 1963-2009 ………...……57

vii 

CHAPTER 1: INTRODUCTION TO THE HISTORY OF NATURAL HISTORY COLLECTING 

 
 
 

“A view of the whole”1: Natural history collecting, 1500-2019 

1500-1600 

 

Though they have undergone many extensive changes since their beginnings in the 

sixteenth century, natural history collections continue to fascinate the professional and the 

amateur alike.   In their influence on the professional scientist, collections of specimens 

from the natural world have informed and supported many major discoveries and theories 

in the biological sciences, from works in the field of anatomy to the development of 

Darwin’s theory of evolution (Darwin 2003, Lubar 2017).  Though early collections are a 

far cry from the professional institutions of today, these proto-collections would become 

one of the most important sources of scientific knowledge. 

In Medici-era Italy and in Prague, early groupings of objects—not yet called 

collections—were encyclopedic in nature and privately owned.  These collections were 

arranged according to some organizational scheme, largely based around ideas of 

differences between the items in the collection (Pearce 1995).  Collectors were not using 

their specimens and objects to answer questions about the natural world and its biological 

functioning; this development would come with the advent of the seventeenth century 

(Pearce 1995).   

The words “collection” and “collector” came to be associated with groupings of 

physical objects in the seventeenth century; up until this point these terms were only 

associated with written materials (Swann 2001).  The objects in these collections, like their 
                                                        
1 “A view of the whole” is the title given to the table of contents of the Catalogue of the Tradescant Collection, 
as published in 1656 by John Tradescant the Younger. 

 1 

predecessors in Italy and elsewhere, had been set aside by a single person, away from 

regular use but would come to hold a place in investigating the world (Swann 2001).  From 

early collections came the establishment of a new culture or form of culture.  Specimens 

that were kept and recorded by collectors represented a way to verify stories and previous 

writings, in addition to being a way for a collector to reevaluate works conducted in the 

past (Pearce 1995, Parrish 2006).  In this way, collections of natural history specimens in 

particular were involved in the reinvention of scientific endeavors by allowing for better 

description of details.  This, in turn, shaped writings on the exact appearance and other 

characters of objects in the natural world, though many such publications—like the catalog 

of the Tradescant collection—read like grocery lists (Tradescant 1656, Pearce 1995, Swann 

2001).   

 

1600-1900 

By the seventeenth century, natural history was an extension of the efforts of 

Classical era philosophers; it consisted of information about and explorations of the natural 

world.  However, Classical natural history also included a variety of fields of interest that 

would no longer be considered a part of natural history, such as astrology (Swann 2001).  

Collections revolutionized the study of natural history as a science because they allowed 

for the reevaluation of past efforts that looked into the nature of the world.   Observations 

of collected specimens, as made by discerning individuals, were to become the new way of 

discovering truths about the world.  This was a move away from prior efforts, which 

consisted of trying to boil knowledge gleaned from earlier writers down to some universal 

truth purely through philosophical thought.  By moving toward observation as a standard, 

 2 

collectors became able to question and revise the writings of the Classical world, though a 

deep understanding of classical writings was still greatly valued in determining the 

academic standing of an author (Pearce 1995, Swann 2001). 

Natural history was then a movement in search of truths that would ideally allow 

the observer to sort specimens and other objects according to a universal language (Pearce 

1995).  This effort to sort and classify was a manifestation of a desire to understand 

Creation through observation and collecting (Lubar 2017).  However, collections were not 

intended, at this time, to document all of life on Earth (Asma 2001, Parrish 2006)—instead, 

collections amassed things that were either from exotic locales or that were odd or rare for 

some other reason (Swann 2001, Parrish 2006).  These masses of oddities were the origins 

of ‘cabinets of curiosities’ or ‘wunderkammer’, which would eventually develop into more 

formal natural history museums and collections. 

Because of this European interest in things taken from exotic or foreign lands, 

America and other distant lands represented a veritable treasure-trove of specimens and 

stories.  For the American colonies in particular, this was due to the wild differences 

between the flora and fauna of the American and European continents. The existence of 

networks of correspondence between European countries and the New World aided in 

relating specimens and observations across the vast ocean (Parrish 2006).  By collecting 

specimens of living things, in addition to objects from the native peoples, European 

explorers and collectors subjugated both the natural world and the peoples in it.  In this 

way, collections of exotic objects became a symbolic representation of domination over 

newly discovered lands and a demonstration of prestige won overseas (Swann 2001, Lubar 

2017).  In a number of cases, living individuals from cultures perceived as ‘primitive’ were 

 3 

also collected and often displayed by people of European origin, both by early institutional 

efforts—such as the 1893 Columbian Exposition in Chicago, Ill. or the display of Ota Benga 

by the Bronx Zoo in 1906—and by private individuals like Carl Hagenbeck, a German man 

who dealt in animals (Domosh 2002, Purtschert 2015, Newkirk 2016).  Some of the shows 

put on by individuals were associated with educational institutions and a growing 

professional scientific community (Purtschert 2015). 

The prestige and social status won through collecting in foreign lands contributed to 

an individual’s status as an authority on a subject—though this status was not extended to 

those, such as colonists, with easy access to foreign materials (Parrish 2006).  For those in 

the British Empire, the expertise of a colonist—due to their geographic closeness to 

organisms and objects for observation—was sought after by higher-status naturalists in 

cities like London (Parrish 2006).   However, colonists were seen as becoming less and less 

English the longer they spent in the American colonies, and their position in the networks 

of communication between naturalists was variable (Parrish 2006).  A person’s social and 

economic status, assuming they were not a colonist, determined what they could collect; 

these seventeenth century collections were used to advance socially and to demonstrate 

social status (Pearce 1995, Swann 2001, Lubar 2017).  Only the fabulously wealthy could 

collect objects of fine art; middle class people collected natural specimens and cultural 

objects from around the world, both through their own travels and through 

correspondence with other collectors (Swann 2001, Mayhew 2005, Parrish 2006).  Those 

trying to be perceived as having a higher economic status also endeavored to collect such 

objects (Swann 2001).  

 4 

 

From this goal of being seen as high-status, private collections developed from 

emphasizing the odd and exotic to being a representative sampling of the objects a 

collector could acquire.  During the seventeenth and eighteenth centuries, the meaning of 

curiosity began to change from denoting ‘something that is out of the ordinary—be it 

exotic, disfigured, incomprehensible, or any number of other things’ to a word that signaled 

a grouping of objects taken from the natural world (Pearce 1995, Parrish 2006).  These 

groupings of objects were becoming more representative of the general state of things in 

nature due to a growing understanding that basic truths of the world could not be 

discerned by looking only at the rare and strange.  Unusual or irregular specimens could 

not be classified according to a logical scheme due to their differences from the nature of a 

typical representative (Pearce 1995).  When specimens that represented the population 

were chosen, there was a greater potential for the object to provide some understanding as 

to its nature and the nature of the place it came from.  In this way, biological classifications 

grew out of collecting efforts (Pearce 1995, Asma 2001).   

 

Unlike the private collecting that had been so concerned with proving wealth and 

status, collecting for some in the eighteenth century became an effort for the public good, 

for science and natural history.  However, the shift from specimens as curiosities of old to 

things of scientific value and merit was not without tension and resulted in the formation of 

a few different streams of collecting philosophy (Pearce 1995).  On the one hand were 

those who continued to collect exotic objects and aimed, often, for commercial success; the 

other main collecting philosophy was focused on formal scientific collecting with the desire 

to understand and explain (Pearce 1995).  Despite this divide between collectors and a 

middle-ground group existing with them, the growing trend toward organization and more 

 5 

thorough understandings of the world were reflected in the collections of the time and 

their increasing levels of order (Pearce 1995).   

 

Further organization of collections would come with the invention of a formal 

system of classification of living things.  Carolus Linnaeus assembled his first plant 

collection in the first half of the eighteenth century and from this would develop a 

systematic way to classify living things. This system would be published in Systema 

Naturae, Genera Plantarum, Species Plantarum, and Philosophia Botanica (Linné and Turton 

1806, Pearce 1995, Linnaeus 2003, Clark 2009, Reid 2009, Grande 2017).  Prior to 

Linnaeus’ system, there was not a universally accepted method of classifying living 

organisms in a scientific manner; though many had made attempts in one form or another 

none was implemented or accepted by all natural historians (Asma 2001, Clark 2009).  

There had also been a long-standing tendency among natural historians to keep their 

efforts in scientific thought close to their chest, without communicating their findings to 

their peers (Reid 2009), which likely limited the ability of natural historians to come to a 

consensus with regards to a system of naming and classification.  In his efforts, Linnaeus 

made use of contemporary methods of scientific thought, i.e. Linnaeus went back to the 

idea of description and trying to extract some fundamental truth about the thing he was 

describing through the action (Reid 2009, Grande 2017), while developing an ‘artificial 

system’ of boiling things down to essential characters, not using details to represent 

nature’s full complexity (Asma 2001, Clark 2009).  This was in keeping with the 

philosophies of the ancient world, especially Plato and Aristotle (Lloyd 1961, Morge 1973, 

Asma 2001).  However, Linnaeus’ system—importantly—required empirical evidence in 

the form of detailed observation or specimens in order for description and naming to be 

 6 

viable where many previous efforts at classification had been implicitly aimed at combining 

natural history with religious beliefs (Clark 2009, Reid 2009).  Linnaeus was also dedicated 

to communicating his ideas; by actively sharing his system with other natural historians, 

Linnaeus popularized his ideas in the community in a way others had not managed.   

 

In addition to Linnaeus’ scientific collecting endeavors, many others were engaged 

in the fervor of specimen collecting.  Natural history societies, social clubs, and field groups 

were formed in order to support collecting efforts; such collaborations enhanced the ability 

of the middle classes to contribute to scientific thought and developments to a degree  

(Pearce 1995).  These collecting groups arguably represented precursors for later 

professional groups and societies, and they mirror later occurrences to the amateur 

following the professionalization of the biological sciences in their relative exclusivity.    

 

For much of the late eighteenth and early nineteenth centuries, the focus in natural 

history studies was on collecting specimens, observing their morphologies, and developing 

an encyclopedic understanding of living things (Gates 2007).  The importance of singular 

characteristics in description and classification continued to define this period in the 

history of systematics.  Over the course of this period of time, many were fascinated by the 

natural world and wrote extensively on their observations (Gates 2007).  By the end of the 

nineteenth century, collecting of natural history specimens was focused towards the 

further development of a system of hierarchies for natural things.  Such a system of 

classification, building from Linnaeus but taking into account more than a single 

characteristic, would allow naturalists to better understand the biological world and the 

place of humans in this scheme (Pearce 1995, Asma 2001, Grand 2017, Lubar 2017).  The 

 7 

development of a biological hierarchy hinged on the perspective of objects and specimens 

in such a way that the observer could understand some part of a larger overall picture.   

 

Despite this larger trend of collecting for systematization, specimen collecting was 

never homogenous—there were always a small number of collectors who sought out the 

rarities and oddities, including things with supposed historical significance (Pearce 1995, 

Lubar 2017).  Within curiosities collecting, the idiosyncrasies of each individual collection 

were dependent on the collector, their interests, and their means (Guralnick and Van Cleve 

2005, Beck and Kitching 2007, Lubar 2017).   

 

This aspect of private collecting was not necessarily excluded completely from the 

encyclopedic and organized formal institutions, however, as curators and other 

professionals often built collections by engaging in expeditions.  Often these expeditions 

were intended for specimen collection; it was the purview of museum curators to acquire 

specimens themselves and then prepare them for addition to the museum collection.  In 

this way, the interests of the curator could be seen in the objects brought back and 

incorporated into the institutional collection, though curators were often required to be 

somewhat of a generalist in their collecting (Rader and Cain 2014, Lubar 2017).  What 

contributed to the separation of professional curators and the self-taught collectors was 

the level of collection information and description saved with natural history specimens; 

within institutions there was often a protocol for curators to document specimens, 

including a minimum requirement for the information to be saved (Lubar 2017).  Though 

museum professionals were often required to go on collecting trips, the public was also 

able to contribute specimens to the museum; museums would put out calls for well-

documented specimens and amateur collectors would answer (Lubar 2017).  Such donors 

 8 

saw their donation of specimens to museums as both a way to contribute to others’ 

learning and as a way to preserve their collections securely, among various other 

motivations in donating (Lubar 2017).  This system of call-and-respond would continue on 

in the twentieth century, extending the time that the amateur naturalist was able to 

contribute to professional science (Rader and Cain 2014).  In addition, the donation of 

large, private collections to public museums served both professional scientists and a 

visiting public (Lubar 2017). 

 

Calls to the public for specimen donations solved one of the more important issues 

facing professional natural history museums: the need for large numbers of specimens.  

Without a collection consisting of many specimens, both similar and dissimilar, 

professional scientists could not investigate areas of natural history such as geographic 

range and variations in species across regions and seasons.  Additionally, multiple 

specimens were necessary for comparison and description (Lubar 2017).  Detailed 

descriptions and the ability to compare one specimen to another were needed for the 

positive identification of newly-collected specimens and served a major role in the 

scientific community, making natural history museums important to scientists beyond 

their walls (Lubar 2017). 

 

The nineteenth century also saw the further development of the concept of a natural 

history museum as a public institution, rather than a private collection.  Through visits to 

an institution of natural history, a person could better both themselves and society at large, 

though museum professionals had mixed opinions on whether the public was an important 

audience to the museum (Rader and Cain 2014, Grande 2017, Lubar 2017).  Due to 

economic growth from both the First and the Second Industrial Revolutions and rapid, 

 9 

lucrative technological change in the United States and Western Europe, it was possible to 

more firmly establish formal institutions that served to educate the public (Pearce 1995, 

Levin et al. 2010, Rader and Cain 2014, Lubar 2017).  Once a handful of natural history 

museums had been formed, it became possible for staff to exchange or distribute 

specimens from one institution to another for the betterment of each collection (Lubar 

2017).   

 

1900-Present 

 

Over the course of the twentieth century, collections of natural history specimens in 

museums served a larger societal purpose with increasing frequency by serving an 

educational mission and by documenting biodiversity (Lubar 2017).  Though the public 

was welcome to the public portion of the museum, they were barred from the professional 

research division of museums’ scientific efforts, sometimes not even aware that biological 

research was happening behind-the-scenes (Lubar 2017).  In conducting these two 

separate functions, museums were often arranged with some specimens out for public 

viewing with many others saved just for professional researchers (Rader and Cain 2014).  

Public displays of collections were aimed toward presenting a complete picture of 

biodiversity with a goal of attaining an understanding in the public audience of biological 

concepts as they were portrayed in displays (Rader and Cain 2014).   

 

The effort to document all of life on earth was not just a goal for the public half of 

the museum.  Natural history museums attempted to be ever-growing collections of 

specimens that would serve as documentation of all forms of life on earth in preparations 

ranging from full specimens to frozen tissue samples (Rader and Cain 2014, Lubar 2017).  

 10

The professional scientists in a museum put their effort toward preserving type specimens, 

in addition to those that served to delineate geographic ranges.  In their documentation of 

geographic information, curators continued to rely on the well-documented donations of 

the amateur naturalist in addition to specimens coming out of the fieldwork of researchers 

(Rader and Cain 2014, Lubar 2017).  However, the professional researcher was the one to 

determine the worth of a collection or object, thus reinforcing a growing divide between 

the professional and the amateur in terms of having authority over such decisions (Lubar 

2017).  More and more, however, justification for the addition of an object to a collection 

would be needed; some perceived authority would no longer be adequate (Lubar 2017).  In 

addition, a movement to return unethically collected and held cultural objects has been 

started—particularly in the United States with the passage of the Native American Graves 

Protection and Repatriation Act (NAGPRA) in 1990 (Colwell 2014, Lubar 2017).  In doing 

so, authority over cultural heritage objects and human remains is returned to the peoples 

of origin (Colwell 2014).  However, returning authority over objects and stories is an 

ongoing process and source of debate. 

 

By the twenty-first century, museums in the United States would contain over 1 

billion objects, and 820 million of these would be natural history specimens. 

Internationally, institutions hold over 3 billion natural history specimens (Lubar 2017).  

Today, biological collections continue to be systematically arranged, now according to 

taxonomic relationships (Lubar 2017), though this sometimes poses a challenge due to the 

frequency of changes made to the classification of some organisms.  Digitization of 

collections and data is an ongoing struggle for museums, though biological collections lead 

this endeavor (Hill et al. 2014, Grande 2017, Lubar 2017, Short et al. 2018).  Increasingly, 

 11

digitization efforts are undertaken as collaborations among institutions (Grande 2017).  

For many institutions, the digitization of their collections serves not only a research 

purpose but an educational purpose as well.  Museums continue to acknowledge the 

educational value of their collections to the greater public (Winston 2007, Grande 2017, 

Lubar 2017). 

 

Museums also face the task of making their collections appear meaningful to the 

public and remaining useful to science; attitudes toward collecting have changed since the 

sixteenth century.  Collecting is no longer done willy-nilly (Rocha et al. 2014, Lubar 2017).  

These and other shifts in popular opinion toward collecting have contributed to an inability 

to ever potentially replace currently held specimens in institutional collections (Short et al. 

2018).  Specimens saved during the heyday of collecting a century or more in the past 

continue to play a role in biological research as they document changes over time and thus 

provide modern scientists with larger datasets for studies determining biological needs 

today (Brooks et al. 2014, Lubar 2017, Short et al. 2018).  Specimens collected over a large 

timeframe allow researchers to understand relationships between organisms and between 

organisms and their environment, including the influence of climate change (Suarez et al. 

2004, Winston 2007, Hill et al. 2012, Brooks et al. 2014, Grande 2017, Lubar 2017).  By 

allowing for a greater understanding of such influences, natural history museums assist in 

efforts to conserve biodiversity into the future (Grande 2017).   

 
 
 

 12

“Can learned skill on little specks display…”2: Collecting and classifying insects, 

1800-2019 

From the first cabinets of curiosity and beginning of natural history collecting, 

insects were sought after for the beauty they manage to exhibit even at their small size.  

This early inclusion of insects in collections made them of great interest to those 

developing systems of biological classification, and insect specimens were the subjects of 

many attempts at a system of classification (Swann 2001, Clark 2009, Roos 2012).  By 

examining the morphologies and natural histories of insects, eighteenth century insect 

collectors developed a number of different methods of classification that would inform 

later systems, though a single, accepted system took longer to develop for insects than it 

did for plants (Clark 2009).  All of the classification schemes that were developed centered 

on external morphologies, not internal.  The determination of which classification schemes 

would last the longest or be thrown out the fastest was linked to the idea of the status quo 

and authority (Clark 2009).   

 

This idea of the status quo in natural hierarchies was linked in many ways to 

attempts to justify a social hierarchy.  Through investigations into anatomy and physiology, 

naturalists assigned levels in the hierarchy, moving from things belonging to the ‘mineral 

kingdom’ that lacked many traits of living things up to mammals and other vertebrates that 

had some form of consciousness (Clark 2009). 

 

Insect classification—and the classification of other living things—in the nineteenth 

century was a reflection of a desire to better understand the theological concept of Creation 

through the collection of natural objects.  In a way, collecting and naming species was an 

                                                        
2 From “The Natural Philosopher” in “The Age of Frivolity” by Thomas Beck (“Timothy Touch’em”), 1806. 

 13

effort to not only own the specimen but also to exercise domination over nature, in 

addition to appreciating the diversity of divine Creation (Harris 2005, Clark 2009).  This 

idea of domination and ownership was a reflection of larger trends in the field of natural 

history collecting and more general social values. 

 

During the nineteenth century, the culture of collecting and naming insect 

specimens was not particularly disturbed by the publication of Charles Darwin’s theory of 

evolution, and entomologists were divided in their opinions on his ideas (Clark 2009).  

However, insects and their collection played a major role in the development of the theory 

of evolution, as both Darwin and Alfred Russel Wallace were coleopterists.  Wallace would 

come to attribute the agreement of these two men on the process of evolution to their 

shared interest in beetles.  In addition to contributing to Darwin’s work on the 

development of evolutionary theory, insects provided him with data reinforcing such 

concepts in further writings outside of Origin of Species, including investigations into 

pollination and other behaviors (Darwin 2003, Clark 2009).  Other naturalists found 

evidence supporting Darwin’s theory of evolution in the insect world.  Henry Walter Bates, 

for example, observed butterflies in the South American rainforest and described a form of 

mimicry.  Bates’ trip abroad was intended as a collecting trip to find specimens to put up 

for sale and justify the entire trip financially.  While collecting specimens, Bates discovered 

a form of mimicry in Heliconian butterflies that protected non-poisonous species from 

being eaten by appearing similar to species that were poisonous (Bates 1862).  The 

development of this pattern of similar coloration was easily explained using the idea of 

natural selection, thus supporting Darwin’s ideas (Clark 2009). 

 14

 

However, many members of the public entering museums were not fond of insect 

specimens.  Insects seemed to serve no purpose on their display shelves.  This opinion, for 

some, extended to the idea of the entire museum.  A general sense of the attitude 

museumgoers may have had is exemplified in the poem “The Age of Frivolity” written by 

Thomas Beck (under the pseudonym “Timothy Touch’em”) before the turn of the twentieth 

century (Touch’em 1806, Carty 2015, poem quoted in Lubar 2017):  

 

“Some spend a life in classing grubs, and try, 

New methods to impale a butterfly; 

…. 

This precious lumber, labell’d, shelv’d, and cas’d, 

And with a title of Museum grac’d 

Shews how a man may time and fortune waste, 

And die a mummy’d connoisseur of taste.”  

 

 

By the end of the nineteenth century, entomology—and biological science as a 

whole—had undergone and was continuing to undergo a process of professionalization.  

Entomology more than other fields of study had been accessible to the amateur naturalist 

or lay-collector due to the relative ease of acquiring and using the materials used to build 

an insect collection (Clark 2009).   This made insect collecting a potential hobby for many, 

assuming they had the economic means to purchase the tools necessary.  Some naturalists 

made an effort to interact with others who shared their interest in collecting insects.  To 

create a community of collectors, entomological societies and clubs were formed, often 

 15

specific to a particular state or region (Essig 1931).  Such groups began to arise in the mid-

nineteenth century and continued forming from there (Clark 2009, Kaplan 2009).  Many of 

the entomological societies formed from groups of amateurs would eventually become 

societies for the professional scientist and inaccessible to the amateur collector (Clark 

2009). 

The private insect collections of these individuals in the late nineteenth and early 

twentieth centuries were commonly donated to institutions, though they were occasionally 

sold to other private collectors (Essig 1931).  Donation of collections was not a common 

practice in natural history collecting in previous centuries.  In the twentieth century, these 

institutional collections would often come to hold specimens that had been named and 

designated as type specimens in the description of species (Essig 1931, Lubar 2017), often 

by amateur entomologists (Waters and MacKenzie 2018).  Though these specimens are 

extremely important, limitations of space would be a growing concern for natural history 

museums.  While insects often take up relatively little space, the single largest collection in 

all the museums in the world is Alfred Kinsey’s gall wasp collection in the American 

Museum of Natural History, New York and consists of over 7.5 million gall wasp specimens.  

This manages to outdo even Harvard’s collection of 1 million ants (Lubar 2017).   

Museums, by the end of the twentieth century, would involve insects in their 

mission of public education in a variety of ways.  Some used insects, along with specimens 

of other living things, in dioramas with the intention of teaching the public about various 

ecological concepts (Rader and Cain 2014).  However, many insect collections were saved 

for the purpose of professional scientific research—arguments in favor of maintaining 

insect collections harken back centuries to the motivations involved in founding public 

 16

museum collections in the first place.  For example, preserved specimens of insects which 

vector diseases were used to teach field medics and hospital workers in both World Wars 

(Winston 2007, Lubar 2017).  Specimens were lent out to these non-entomologist 

professionals in an effort to allow for the identification of such medically important species 

in the field (Lubar 2017).  Collections in natural history museums also supported 

agricultural aims and helped to spread Western-style science to other regions of the world 

(Suarez et al. 2004, Lubar 2017). 

Though amateur entomologists in some ways have been pushed from the 

entomological scene, they continue to make lasting contributions, particularly in the way of 

specimens.  For example, two amateur collectors recently donated their pooled 

Lepidoptera collection—representing over 70,000 specimens from a wide variety of taxa—

to the McGuire Center for Lepidoptera and Biodiversity (Watkinson 2018).  The sum of 

historical collections from a variety of amateur collectors that have been donated to 

institutions over time also represent a valuable resource to modern professional 

entomologists, e.g. the Lyman collection (Waters and MacKenzie 2018).  Groups specifically 

founded to support amateur entomologists and their contributions continue to exist 

despite growth in professional societies that cater to the vocational entomologist; the 

Amateur Entomologists’ Society in the United Kingdom continues to function both as a 

conservation-concerned Society and works to encourage potential future entomologists 

(Lonsdale 2012).  Groups of amateur entomologists have both noticed and investigated—

among other things—global insect decline, some likely being among the first to voice 

concern in the 1930s (Lonsdale 2012, Hallmann et al. 2017). 

 17

The debate over the value of keeping and adding to insect collections continues, and 

the pro-collecting side stresses the usefulness of collections to science today and in the 

future as a justification for continued collecting efforts.  Historical insect collections are 

almost entirely irreplaceable for many reasons—including habitat change, climate change, 

funding for collections staff and collecting trips, and decreases in global biodiversity—

making old collections invaluable to research (Suarez et al. 2004, Winston 2007, Brooks et 

al. 2014, Short et al. 2018).  The efforts of groups like iDigBio to digitize collections of 

insects and other natural history specimens make the information contained in these old 

collections available to researchers, in addition to preserving specimen data should the 

specimens ever be destroyed.  Researchers involved in such projects ask many diverse 

questions about the biological world that would otherwise be impossible to address.  

However, databasing and digitization is extremely time-consuming and often not well-

funded; less than 2% of the total number of institutionally-held insect specimens have been 

databased (Short et al. 2018).  For some digitization projects, the field of entomology has 

come full circle; volunteers and citizen science initiatives have been used to database 

specimens, once again involving the amateur in scientific pursuits (Hill et al. 2012).  Given 

that a relatively small percentage of the total number of museum specimens has been 

databased, there has historically been a limit on the variety and depth of studies using 

whole-collection data for various purposes.  The dearth of digitized specimens is not the 

only limitation to entomological research; though insect collections are irreplaceable, they 

will also never be complete due to various limitations on collectors over the past four 

centuries (i.e. there has not been consistent collecting everywhere in the world every single 

 18

year since collecting began) (Beck and Kitching 2007, Winston 2007).  Efforts to pool data 

from many institutions aim to minimize the effect of this limitation to future research.   

LepNet consists of 27 United States insect collections, which aim to collectively 

database and georeference the label information of 1.7 million Lepidoptera specimens 

(Seltmann et al. 2017).  Using the existing efforts of LepNet to database and pool data from 

Lepidoptera specimens housed in US institutional collections, this thesis project aims to 

characterize American efforts to collect butterflies and moths since 1800.  First, we 

examined the influence of avocational, or non-professional, collectors based in the state of 

Michigan on the A.J. Cook Arthropod Research Collection located at Michigan State 

University across time.  Using country-wide data generated through LepNet, changes across 

both time and space were characterized as trends associated with the growth and decline 

of natural history as a scientific study, in addition to examining the influence of non-

professional collectors at the national level.  This research was conducted with the 

intention of determining (a) the scope of the role of non-professional collectors in terms of 

specimens collected, (b) how the influence of non-professional collectors affects scientific 

research making use of institutionally-held Lepidoptera collections, (c) how evenly—or 

unevenly—distributed collection locations have been historically for Lepidoptera 

specimens collected in the United States, and (d) how collection efforts have changed over 

time, affecting research questions regarding time series, including studies around climate 

change. 

 

 

 19

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

 

BIBLIOGRAPHY 

 

 20

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 25

CHAPTER 2: THE LEPIDOPTERA OF THE A.J. COOK ARTHROPOD RESEARCH COLLECTION 

AND THE IMPACT OF AVOCATIONAL COLLECTING 

 

Introduction 

There has long been a history of the amateur naturalist or lay-person being engaged in 

scientific study to varying degrees—in the areas of collecting, observing, and 

communications with ‘professionals’, among other things (Vetter 2011).  However, in many 

cases, the line between the amateur and the professional, or lay-person and expert, has 

been blurred.  Despite a lack of consistent differentiation between the amateur naturalist 

and the professional scientist, there has also been a history of excluding the ‘stamp-

collecting naturalists’ from the realm of science as biology became more and more 

concerned with experimentalism (Johnson 2007, Vetter 2011).  In some cases, it has been 

said that amateur collecting efforts were simply a steppingstone to ‘proper’ science (Kohler 

2007).  However, the efforts of non-professional scientists continue to be valuable sources 

of data in many fields of biological science, including entomology.  For example, skilled 

amateurs working in Germany were the first to quantify a continuing decline in insect 

biomass based on decades of saved mixed-species alcohol samples from traps (Hallmann et 

al. 2017). However, the number of non-professionals working and publishing in natural 

history and insect taxonomy is on the decline, in spite of their importance in identifying 

and recording sightings of specific insect species (Hopkins and Freckleton 2002). For this 

reason, it is important to evaluate the overall impact of amateur collectors on existing 

specimen collections. 

The A.J. Cook Arthropod Research Collection (ARC) was founded by its namesake in 

1867 at Michigan State University (MSU, then the Agricultural College of the State of 

 26

Michigan).   The collection currently houses 1.1 million pinned insect specimens, in 

addition to others preserved on slides or in alcohol.  The collection holds specimens 

documenting Michigan insect biodiversity with areas outside of the state addressed to 

varying degrees dependent on the individual collecting efforts of its many donors.  The 

Lepidoptera in particular are well-represented, consisting of approximately 185,000 

specimens.  The ARC owes much to amateurs and volunteers in terms of both building the 

collection and its curation during the ARC’s 152-year history. 

Databasing of the ARC’s lepidopteran specimens is ongoing as part of a national 

collaborative databasing effort known as LepNet (lep-net.org, Seltmann et al. 2017). LepNet 

allows for a greater understanding of collecting patterns over time both nationally and 

locally.  Information about the specimens, as well as images, are available via the LepNet 

public website, providing information to both researchers and laypersons across the world 

(SCAN-bugs.org).  As such, this database represents an unparalleled opportunity to analyze 

the development of Lepidoptera collections, including the collection of North American 

Lepidoptera found in the ARC.  We focus on specimens in the ARC as a case study because 

there is a long history of extensive work done by non-professional entomologists building 

and curating this collection.  Examination of the influence non-professional and 

professional collectors have had on institutional collections is important due to the 

implications changes in collecting effort might have on the collections as a whole and thus 

their potential for supporting future research using historical specimens, particularly time 

series analyses.   

This study describes the pattern of collecting North American Lepidoptera found in 

the ARC.  Specifically, we quantify represented taxa; specimens collected per year; and 

 27

specimens collected by professional and non-professional (or avocational) collectors. In 

particular, we examine the relative impact of avocational collecting on the ARC.  

 

Methods 

Data associated with the 96,618 currently digitized specimens of the approximately 

185,000 total Lepidoptera specimens housed in the A.J. Cook Arthropod Research 

Collection at Michigan State University (ARC) were downloaded from the Symbiota 

Collections of Arthropods Network database (SCAN-bugs.org), accessed 23/03/2019.  

Using the ‘tidyverse’ R package (Wickham 2017), the collection data were revised to only 

include specimens that were collected in the United States between 1800 and 2018.  The 

remaining 83,682 specimens were assessed with respect to the number of collection events 

per year and the collection locality at state level.  A rolling decadal average and standard 

deviations for collecting events were calculated using the ‘TTR’ package (Ulrich 2018).  

Rolling decadal averages and standard deviations are the unweighted means and standard 

deviations of the values—specimen counts—for the previous ten years.  The package 

‘ggplot2’ from the ‘tidyverse’ suite of R packages was used to generate all graphs. The 

number of specimens per square kilometer collected per state was generated using the 

‘usmap’ R package.  A map of specimens collected per square kilometer in Michigan 

counties was also generated using the R packages: ‘ggmap’, ‘maps’, and ‘mapdata’.   

Specimen records were also used to determine the composition of Lepidoptera families 

represented in the collection and whether collectors were professionals or non-

professionals.  For the latter, the records were sorted by collector name and year, and 

tallied to generate a specimen count per year for each collector.  Collectors were identified 

 28

as professional or non-professional by reviewing ARC records (Houk 1954) and 1964 - 

2018 newsletters of the Michigan Entomological Society (michentsoc.org) written by and 

about people associated with the ARC and using Google Internet searches of names on the 

Internet.  Collectors were considered professional if they held positions as university 

professors or as collections staff in an institutionally-held natural history collection.  The 

collection efforts of professionals and non-professionals were then compared using 

repeated Mann-Whitney-Wilcoxon tests in the base R statistical package for each decade 

between 1950 and 1994 on a sliding window basis.   This window of time was used to 

analyze the collecting effort of different types of collector because it includes the top ten 

years in terms of total specimens collected.  During the period from 1950 to 1994, 68.7% of 

the specimens in the A. J. Arthropod Research Collection were collected, so the majority of 

the data was included in analyzing the differences between avocational and vocational 

collectors.  

 

Results 

Our initial review of the LepNet data removed 13.39% of the specimen records 

because of missing or erroneous collection year or country of origin, resulting in a dataset 

consisting of 83,682 records.   The maximum of the data, in terms of collection events per 

year, occurred in 1960 with a total of 3,753 specimens collected (Figure 2.1).  The second 

highest peak in collection events occurred in 1965 (3,224 specimens).  For almost 60 years, 

there has been, on average, a steady decline in annual collection events in Michigan.  This 

decline has become steeper in recent years, decreasing from 1,693 specimens in 1987 to 

fewer than 100 specimens collected annually since 2011.  The most frequently collected 

 29

family was the Erebidae (25.64% of total), the favorite taxon of a handful of Michigan 

collectors, while other commonly collected families included Geometridae (13.76%), 

Noctuidae (11.43%), and Tortricidae (8.23%) (Table 2.1).   

 

 

 
Figure 2.1: Specimen collection events per year since 1800 in A. J. Cook Arthropod Research 
Collection.  The green line is a rolling decadal average, and the grey lines represent a single 
standard deviation from this average on a rolling decadal basis.   
 
 
 
Table 2.1: Counts and percentages of specimens from represented Lepidoptera families.  
 
Family 
Acanthopteroctetidae 
Acrolepiidae 
Acrolophidae 
Adelidae 
Alucitidae 
Apatelodidae 
Arctiidae 
Argyresthiidae 

Percent 
0.002 
0.001 
0.002 
0.047 
0.013 
0.240 
0.002 
0.043 

Count 
2 
1 
2 
39 
11 
201 
2 
36 

 30

Table 2.1 (cont’d) 
 
Attevidae 
Autostichidae 
Batrachedridae 
Bedelliidae 
Blastobasidae 
Bombycidae 
Bucculatricidae 
Carposinidae 
Choreutidae 
Coleophoridae 
Copromorphidae 
Cosmopterigidae 
Cossidae 
Crambidae 
Dalceridae 
Depressariidae 
Doidae 
Drepanidae 
Dudgeoneidae 
Elachistidae 
Epermeniidae 
Epiplemidae 
Erebidae 
Eriocraniidae 
Euteliidae 
Galacticidae 
Gelechiidae 
Geometridae 
Glyphipterigidae 
Gracillariidae 
Heliodinidae 
Heliozelidae 
Hepialidae 
Hesperiidae 
Incurvariidae 
Lacturidae 
Lasiocampidae 
Lecithoceridae 
Limacodidae 
Lycaenidae 
Lymantriidae 
Lyonetiidae 
Megalopygidae 

126 
33 
2 
18 
76 
11 
49 
5 
55 
189 
1 
108 
178 
5,930 
3 
857 
2 
57 
3 
16 
125 
1 
21,460 
14 
34 
7 
951 
11,511 
8 
614 
19 
31 
54 
2,870 
7 
4 
1,814 
6 
612 
1,710 
32 
9 
139 

 31

0.151 
0.039 
0.002 
0.022 
0.091 
0.013 
0.059 
0.006 
0.066 
0.226 
0.001 
0.129 
0.213 
7.086 
0.004 
1.024 
0.002 
0.068 
0.004 
0.019 
0.149 
0.001 
25.645 
0.017 
0.041 
0.008 
1.136 
13.756 
0.010 
0.734 
0.023 
0.037 
0.065 
3.430 
0.008 
0.005 
2.168 
0.007 
0.731 
2.043 
0.038 
0.011 
0.166 

Table 2.1 (cont’d) 

Micropterigidae 
Mimallonidae 
Momphidae 
Nepticulidae 
Noctuidae 
Nolidae 
Notodontidae 
Nymphalidae 
Oecophoridae 
Opostegidae 
Papilionidae 
Pieridae 
Plutellidae 
Prodoxidae 
Psychidae 
Pterolonchidae 
Pterophoridae 
Pyralidae 
Riodinidae 
Saturniidae 
Scythrididae 
Sesiidae 
Sphingidae 
Stathmopodidae 
Thyrididae 
Tineidae 
Tischeriidae 
Tortricidae 
Uraniidae 
Yponomeutidae 
Ypsolophidae 
Zygaenidae 
N/A 
 
 

1 
69 
19 
62 
9,569 
61 
755 
2,569 
68 
2 
1,392 
616 
44 
86 
131 
1 
316 
1,689 
43 
1,703 
31 
2,499 
3,935 
2 
37 
585 
124 
6,883 
7 
206 
57 
73 
2 

0.001 
0.082 
0.023 
0.074 
11.435 
0.073 
0.902 
3.070 
0.081 
0.002 
1.663 
0.736 
0.053 
0.103 
0.157 
0.001 
0.378 
2.018 
0.051 
2.035 
0.037 
2.986 
4.702 
0.002 
0.044 
0.699 
0.148 
8.225 
0.008 
0.246 
0.068 
0.087 
0.002 

Given that the A. J. Cook Collection is a collection housed in Michigan and largely 

supported by collectors based in Michigan, it is unsurprising that the geographical spread 

of collection locales is heavily biased toward the state of Michigan (63,907 specimens).  

This is followed by New Mexico (2,393 specimens), Arizona (2,046 specimens), and Ohio 

 32

(1,775 specimens) (Figure 2.2).  Other states are represented to a lesser degree. Specimens 

collected within the state of Michigan are not evenly distributed across counties (Figure 

2.3).  The counties with the most specimens collected are Otsego (4,810 specimens), 

Kalamazoo (4,287 specimens), Oakland (3,503 specimens), and Clinton (3,332 specimens).  

 

Figure 2.2: National distribution of Lepidoptera specimens in the A. J. Cook Arthropod 
Research Collection. 
 
 
Using collection data sorted by collector name, we compared the collection contributions of 

vocational and avocational collectors to the Cook Collection over time (Figure 2.4). 

Avocational collectors contributed 56,323 Lepidoptera specimens while vocational 

collectors contributed 21,127 specimens.  Temporal patterns in collection event numbers 

are also inconsistent between avocational and vocational collectors; while vocational 

collectors were at their most active in the 1950s through the early 1970s, avocational 

collectors were in the midst of a gradual increase in collecting activity.  During a 20-year 

period of peak collecting (1950-1970), the collecting efforts of these two groups were 

 33

significantly different (Table 2.2).  However, after that time, neither group of collectors 

collected significantly more specimens than did the other, statistically (Table 2.2).  If the 

role of time is ignored completely, there is not a significant difference between the 

specimen numbers contributed by the two groups (W = 2243786, p = 0.7014). 

 

Figure 2.3: Michigan distribution of Lepidoptera specimens in the A. J. Cook Arthropod 
Research Collection at the county level. 
 
 

 

 34

 

Figure 2.4: Avocational and vocational specimen collection events, 1950-1993. 
 
 
 
Table 2.2: Vocational vs. avocational collectors: comparison of specimen numbers by decade. 
 
Year Range 
1950-1959 
1951-1960 
1952-1961 
1953-1962 
1954-1963 
1955-1964 
1956-1965 
1957-1966 
1958-1967 
1959-1968 
1960-1969 
1961-1970 
1962-1971 
1963-1972 
1964-1973 
1965-1974 
1966-1975 
1967-1976 

Interpretation 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 

P-value 
0.0137 
0.0002981 
1.381 e-5 
5.29 e-6 
5.381 e-6 
3.648 e-6 
2.107 e-6 
0.000115 
0.001311 
0.008534 
0.07216 
0.2681 
0.6377 
0.5317 
0.8308 
0.5104 
0.3096 
0.4148 

 

 35

Table 2.2 (cont’d) 
 
1968-1977 
1969-1978 
1970-1979 
1971-1980 
1972-1981 
1973-1982 
1974-1983 
1975-1984 
1976-1985 
1977-1986 
1978-1987 
1979-1988 
1980-1989 
1981-1990 
1982-1991 
1983-1992 
1984-1993 
 
 
The most prolific collector who contributed to the ARC is Mogens “Mo” Nielsen, an 

No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 

0.2713 
0.2717 
0.07194 
0.1682 
0.1528 
0.1766 
0.1078 
0.4201 
0.2665 
0.3383 
0.1664 
0.5256 
0.7449 
0.4907 
0.4828 
0.8075 
0.6029 

avocational collector active for 53 years (Table 2.3).  Mr. Nielsen contributed more than 

three times the number of Lepidoptera specimens that were donated by the top vocational 

collector and second most prolific collector (Julian Donahue). 

 
Table 2.3: The top 10 avocational and vocational collectors represented in the ARC 

Association 
Avocational 
Vocational 
Avocational 
Vocational 
Avocational 
Avocational 
Avocational 
Vocational 
Vocational 
Vocational 

Collector 
M. C. Nielsen 
J. P. Donahue 
E. H. Metzler 
R. L. Fischer 
W. C. Stinson 
J. H. Newman 
G. J. Balogh 
R. R. Dreisbach 
G. C. Eickwort 
G. L. Parsons 
 
 
 

Specimens 
19,326 
5,201 
5,188 
5,011 
2,902 
2,758 
2,040 
2,003 
988 
892 

 36

Years Active 
1957-2010 
1960-1993 
1962-2013 
1937-1991 
1930-1983 
1941-1986 
1944-1999 
1915-1963 
1959-1966 
1970-2018 

Discussion 
 

The A. J. Cook Arthropod Research Collection logically consists largely of specimens 

from Michigan (Figure 2.2), as this is the state the collection is located in, with Erebidae 

representing the majority of lepidopteran specimens (Table 2.1).  However, Michigan 

counties are not evenly represented in terms of Lepidoptera specimens collected per 

square kilometer. Most activity was near urban areas or vacation localities, and the 

Michigan “thumb” is the most under collected region (Figure 2.3).  The highest peak in 

annual collecting events occurred in 1960; since then there has been a general decline in 

annual collection events (Figure 2.1).  In total, avocational collectors have contributed 

more than twice the specimens of vocational collectors, and this contribution was 

significantly greater between 1950 and 1968 (Table 2.2).   

Clearly, avocational collectors are important to the growth of entomology 

collections; in the case of the A. J. Cook Arthropod Research Collection, they contributed the 

majority of specimens.  Historically, the vast majority of entomologists have been amateurs, 

working with insects out of enjoyment separate from their paying vocations (Wolfenbarger 

1954).  Many organized groups of avocational collectors arose from their shared 

enthusiasm for insects, especially in the nineteenth and twentieth centuries, even amidst a 

growing emphasis on professionals in the biological sciences (Essig 1931, Hunter and 

Jaros-Su 1997, Clark 2009, Kaplan 2009, Lonsdale 2012, Michigan Entomological Society 

2018).  These non-professional entomological groups include the Amateur Entomologists’ 

Society in the UK, the Florida Entomological Society, and the Michigan Entomological 

Society (MES), which are all still active (Florida Entomological Society 1917, Lonsdale 

2012, Michigan Entomological Society 2018).  These societies act as catalysts for citizen 

 37

participation in biodiversity science and support amateur collecting, along with helping to 

engage the next generation in entomological study.  For example, the members of the 

Michigan Entomological Society have organized group collecting trips in the past—

including the foundation of a formal annual spring collecting trip in 1965 (Michigan 

Entomological Society 1965)—which likely contributed to spikes in collecting activity 

following the founding of the Society in 1954.  The accumulated collections of insects and 

data from non-professional entomological societies continue to contribute, and likely 

enable, professional entomological research (Winston 2007, Brooks et al. 2014).   

In the case of the Michigan Entomological Society, there have been a number of 

highly dedicated Lepidopterists collecting both in Michigan and out of state. Of the many 

avocational collectors who contributed to the collection over the last century, Mr. Mo 

Nielsen was a particularly dedicated benefactor to Michigan Lepidopterology and to the 

development of the ARC.  Mr. Nielsen enrolled in Forest Management at Michigan State 

University after his service in World War II and was mentored by John H. Newman, who 

also contributed to the ARC through specimen identification, curation, and donation.  After 

graduating from MSU, Mr. Nielsen worked for the Michigan Department of Natural 

Resources and continued to develop a lifelong interest in butterflies and moths (Michigan 

Entomological Society 2014).  He was a founding member of the Michigan Entomological 

Society (est. 1954) and a mentor to young Lepidopterists.  He amassed a collection of over 

13,000 Lepidoptera specimens, and he concentrated on Erebidae and Noctuidae.  This 

collection was the basis for a field guide to the butterflies of Michigan (Nielsen 1999).  Mr. 

Nielsen assisted in the curation of the ARC’s butterfly and moth collection and donated 

 38

specimens throughout his adult life. He left the total of his personal collection to the ARC, 

which was incorporated into the research collection following his passing in 2014.  

As many of the collectors who have contributed to the ARC were living and working 

in the state of Michigan at or before the time they started donating specimens, it is 

unsurprising that the Lepidoptera collection is heavily skewed in favor of Michigan 

localities.  In particular, Mo Nielsen had what he referred to as his “hut” located in Otsego 

County.  Mo often did extensive collecting of Lepidoptera specimens while there, and he 

also hosted friends—many of them from MES— to collect as well (Michigan Entomological 

Society 2014).  In addition, Clinton County is home to portions of East Lansing, where MSU 

is located; it is likely that many collectors associated with the Michigan Entomological 

Society and Michigan State University collected frequently in the area.  Mo and others 

focused heavily on the Erebidae, making the family the most numerous in the ARC.  Other 

commonly collected families of moth—Geometridae, Noctudiae, and Tortricidae—are 

economically important crop pests; they were likely collected in large numbers as part of 

agricultural research.  Given the contributions to the ARC by Mo and his contemporaries, 

including nearly 5,000 specimens from Otsego county, it becomes clear that collecting 

efforts by non-professionals have been very important to the growth of the A. J. Cook 

Arthropod Research Collection, and likely other collections around the country.   

 

The difference in the magnitude of contributions made by avocational collectors as 

compared to their professional counterparts may be due to differences in the nature of 

their interest in entomology; avocational collectors tend to have greater enthusiasm for 

active collection of specimens relative to professional biologists (Strasser 2012).  Changes 

in the focus of biology, such as the rise of molecular studies and other laboratory-based 

 39

science in defining professional biology, have brought with them—in many ways—an 

exclusion of avocational collectors (Clark 2009, Strasser 2012), which likely contributed to 

the decline in specimen acquisition.  The availability of funding for particular biological 

sub-disciplines has greatly influenced the pursuits of professional scientists (Asma 2001, 

Tewksbury et al. 2014).  The decline of funding for natural history research has stymied 

dedicated professional collecting efforts (Tewksbury 2014).  Only the collections manager 

of the A. J. Cook Arthropod Collection (Gary Parsons) consistently contributes lepidopteran 

specimens (Table 2.3).   

 

However, the decline in Lepidoptera specimen collecting may not be as dire as the 

data suggests.  Institutional insect collections have an approximate 10-year lag between 

when donated specimens are physically added to public collections and when these 

specimens are catalogued in databases (ECN annual meeting attendees, personal 

communication, November 10, 2018).  In addition, collectors likely keep specimens in their 

private collection for years before donation, so specimens collected in 2015 may not 

become publicly available for 20-30 years (e.g. the Kenelm Philip collection at 

USNM/University of Alaska, the Finkelstein and Knudson/Bordelon collections at the 

McGuire Center at the University of Florida).  These two aspects of vocational and 

avocational specimen acquisition probably contribute to the observed steep decline in the 

last decade (Figure 2.1), making the number of recently collected specimens artificially low.  

In case of the A. J. Cook Arthropod Collection, there are only 4 or 5 substantial pending 

donations, which would increase the representation of specimens collected in the 21st 

century.  However, these collections and unknown future donations likely will not 

 40

approach the numbers of specimens added to the ARC in the height of collecting activity 

(Figure 2.1).  

Engaging new collectors is necessary for institutional entomological collections to 

continue to grow and represent insect diversity.  The existing networks of amateur 

collectors have encouraged new participation often through reduced membership dues for 

students (e.g. MES). In addition, the Amateur Entomologists’ Society conducts public 

outreach and has established a group for new, young collectors (Lonsdale 2012). Other 

opportunities for creating a future core of avocational collectors exist in K-12 schools and 

at undergraduate and graduate institutions. Extracurricular K-12 activities often include 

science clubs and summer camps, some with a focus on natural history (e.g., Science 

Olympiad, 4-H, Girl Scouts). General entomology and insect taxonomy courses are available 

at nearly all US state universities—often with associated insect-focused student clubs.  

Thus, the “infrastructure” is in place to grow and nurture amateur entomologist collectors, 

but the “metamorphosis” of these amateurs into lifelong avocational collectors is the 

challenge.  It is unknown how effective the “infrastructure” may prove to be into the future 

as far as inspiring lifelong collecting.   At the A. J. Cook Arthropod Research Collection, we 

know of only a few amateur and avocational Lepidoptera collectors that may donate 

specimens collected in the 21st century. We fear that the heyday of the avocational collector 

has passed.  

 

 

 41

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

 

BIBLIOGRAPHY 

 

 42

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Lonsdale, D. (2012) Insect conservation in the United Kingdom – The Amateur  

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 45

CHAPTER 3: THE RISE AND FALL OF LEPIDOPTERA COLLECTING IN THE UNITED STATES 

 
Introduction 

SINCE 1800 

 

Insects have been kept in natural history collections from their earliest days as 

Renaissance cabinets of curiosity (Alberti 2002, Harkness 2007).  Historical natural history 

collections, now often held by institutions, are unmatched in their ability to document 

information about species across time and space (Bartomeus et al. 2011, Polgar et al. 

2013).  These historical collections—combined with more recent collecting activity—

record changes in species range, seasonality, plasticity of characters, and allow for the 

identification of specimens in all fields of natural history (Prather et al. 2004b, Winston 

2007).  Museum collections of insects have been used in several studies regarding 

temperature shifts that have come with global climate change (Bartomeus et al. 2011, 

Polgar et al. 2013, Brooks et al. 2014).  For example, using the specimens in the Natural 

History Museum, London, Brooks et al. (2014) found a correlation between the collection 

date of early spring Lepidoptera and temperature over time.  Studies that use collections to 

test hypotheses or describe natural patterns tend to use specimens of a certain taxon, 

particularly those within one or a few collections.  The enormity of gathering specimen-

level data limits large-scale inferences.  Only the databasing of specimen-level data across 

many institutes enables testing of grand hypotheses, such as the effects of global climate 

change on the range of Lepidoptera native to southern states.  For example, a database of 

over 1-2 million historical and contemporary mammal specimens from multiple US 

institutes is available for such large-scale studies (Malaney and Cook 2018).   

 46

Specimens of butterflies and moths and their associated data offer detailed insight to 

environmental and species change over time and space.   Given their aesthetic appeal, they 

have been collected in earnest over time and are well-represented in collections 

throughout the United States and worldwide.  Approximately 17 million lepidopteran 

specimens exist in collections worldwide (Kawahara and Pyle 2012), which if databased, 

could provide the data needed to address large-scale hypotheses.  An effort to connect 

Lepidoptera collections in the US began in 2016 with LepNet: a group of 27 US collections, 

which aims, in part, to database and georeference label information for 1.7 million 

specimens, a third of the collections’ total holdings (Seltmann et al. 2017).  This sample 

represents most US states and the 86 lepidopteran families occurring in North America. 

However, details of the distribution and numbers of specimens collected in specific time 

periods and areas of the US are unknown, both because of inadequate sampling effort and 

incomplete label data associated with specimens.  As observed in other specimen-level 

datasets, the spatial and temporal distribution is likely uneven (Malaney and Cook 2018).  

This uneven spatial and temporal distribution has been noted in global inventories and 

observation records of Lepidoptera species (Girardello et al. 2019).  Declines in collecting 

efforts observed across many taxa in part explain contemporary gaps in collection records 

and specimen distributions (Gardner et al. 2014).  Also not widely documented is the 

profession of the collector; naturalists and non-professional systematists have made great 

contributions to science and collections (e.g., Alfred Russell Wallace) (Vetter 2011). 

However, their impact and significance as compared to professionals have not been 

quantified up until this point.  Anecdotal evidence, combined with analysis of the A.J. Cook 

Arthropod Research Collection in Chapter Two, suggests that their impact is significant 

 47

(e.g., Hallmann et al. 2017), so understanding their role in documenting lepidopteran 

diversity is pertinent.  This is especially true in the midst of an observed decline in amateur 

interest in the natural sciences (Hopkins and Freckleton 2002).  Because of the overall 

declining trend in specimen collecting, documented in numerous other studies, including 

Chapter Two of this thesis, we predict that Lepidoptera collecting as a whole has not been 

consistent over time, space, and collector profession.  Thus, the collections held in many 

institutions may not be truly representative of the natural world.  As such, this study 

describes the patterns of lepidopteran collecting in the US since the mid 1800s as reflected 

in one million LepNet databased specimens.  It also tests for correlations between patterns 

of collecting and collector profession and trends of lepidopteran interest in an effort to 

explain the observed patterns.  

 
 
Methods 
 

Data associated with 1,405,997 databased Lepidoptera specimens housed in 75 US 

insect collections were downloaded from the Symbiota Collections of Arthropods Network 

database (SCAN-bugs.org), accessed 16/05/2019.  Using the ‘tidyverse’ R package 

(Wickham 2018), the collection data were revised to include only specimens that were 

collected in the United States between 1800 and 2018.  These 1,031,401 specimens were 

used to determine the number of collection events per year and the collection locality at 

state level.  The package ‘ggplot2’ from the ‘tidyverse’ suite of R packages was used to 

generate all graphs.  A rolling decadal average and standard deviations (also on a rolling 

decadal basis) for annual collecting events were calculated using the ‘TTR’ package (Ulrich 

2018).   

 48

The base R statistics package and ggplot2 (Wickham 2018, R Core Team 2019) were 

also used to test for a statistical difference between the data from states with LepNet 

participants and states without institutions funded through LepNet.  We used a single 

data.frame of state names, specimens collected per square kilometer, a categorical variable 

to indicate if the state has a LepNet institution and the log10 of the specimens/km2 value 

for each state.  A one-sample Kolmogorov-Smirnov test was used to determine if the 

specimens/ km2 data represent a normal distribution (ks.test where x is specimens/km2 

and y is pnorm).  This was not the case and the data are heavily skewed, so the Central 

Limit Theorem does not apply.  The data for states with and without a LepNet institution 

were compared using a Kruskal-Wallis one-way analysis of variance. 

To understand potential reasons for the observed collecting activity, we compared 

membership information from the Lepidopterists Society with the Lepidoptera collection 

data using the base R statistics package and ggplot2.  This was conducted using two 

data.frames, one for LepSoc membership data (year joined, year paid through) and one for 

Lepidoptera collection events per year.  The Lepidoptera collection data were trimmed to 

fit the membership data from the Lepidopterists’ Society (LepSoc), which was founded 

1974 and whose data terminate at 2016 for new memberships.  LepSoc membership and 

Lepidoptera collection datasets were compared using a two-sample Kolmogorov-Smirnov 

test to determine if the number of members added per year was drawn from the same 

distribution as the number of collection events per year.  This would indicate a strong 

relationship between a major entomologists’ society and collecting activity.  

 

In addition, valid specimen records were sorted by collector name and year, and 

were tallied to generate a specimen count per year for each collector.  Collectors were 

 49

identified as professional or non-professional by reviewing institutional records of 

associated collectors and using Google searches of names on the Internet.  Collectors were 

considered professional if they held positions as university professors or as collections 

staff in an institutionally-held natural history collection.  The top 10% of collectors—both 

professionals and non-professionals—in terms of specimens collected were then compared 

using repeated Mann-Whitney-Wilcoxon tests for each decade in the between 1963 and 

2009 on a sliding window basis of highest total collection effort using the base statistical 

package in R.  This top 10% of collectors represented 80.89% of the total number of 

specimen records. 

We limit our inferences and conclusions of the analyses for collector data to the 

years prior to 2009 because of a potential 10-year lag for the curation of recently collected 

specimens (discussion with participants of Entomological Collections Network 2018).  That 

is, specimens collected in 2009 are mostly likely to be curated, identified, and available for 

databasing by 2019.  Percent lepidopteran family composition was also calculated for the 

data. 

 

Results 

 

Our initial review of the national LepNet data removed 26.64% of the specimen 

records because of erroneous collection years, non-US countries of origin, collections not in 

US states, or non-lepidopteran families.  The single year with most collected specimens 

(23,319) was 1996.  For the last couple decades, there has been a general decline in annual 

collection events of Lepidoptera in the United States.  This decline has become more 

prominent in recent years, decreasing from nearly 20,000 specimens in 2010 to less than 

 50

10,000 specimens collected annually since (Figure 3.1).  Nationally, the most collected 

family of Lepidoptera was the Noctuidae (16.29% of total) followed by Geometridae 

(13.47%), Erebidae (12.98%), and Nymphalidae (9.42%) (Table 3.1). 

 

 

Figure 3.1: Specimen collection events per year since 1800 in institutional collections across 
the United States.  The green line is a rolling decadal average, and the grey lines represent a 
single standard deviation from this average on a rolling decadal basis. 
 
 
Table 3.1: Counts and percentages of specimens from represented Lepidoptera families  
 
Family 
Acanthopteroctetidae 
Acrolepiidae 
Acrolophidae 
Acrolophilidae 
Adelidae 
Agonoxenidae 
Alucitidae 

Percentage 
0.002 
2.909 e -4 
0.010 
1.939 e -4 
0.041 
0.001 
0.012 

Count 
25 
3 
104 
2 
427 
13 
121 

 51

Table 3.1 (cont’d) 

Apatelodidae 
Arctiidae 
Argyresthiidae 
Attevidae 
Autostichidae 
Batrachedridae 
Bedelliidae 
Blastobasidae 
Bombycidae 
Bucculatricidae 
Carposinidae 
Choreutidae 
Coleophoridae 
Copromorphidae 
Cosmopterigidae 
Cossidae 
Crambidae 
Ctenuchidae 
Dalceridae 
Depressariidae 
Doidae 
Douglasiidae 
Drepanidae 
Dryadaulidae 
Dudgeoneidae 
Elachistidae 
Epermeniidae 
Epicopeiidae 
Epiplemidae 
Epipyropidae 
Erebidae 
Eriocraniidae 
Euteliidae 
Galacticidae 
Gelechiidae 
Geometridae 
Glyphipterigidae 
Gracillariidae 
Heliodinidae 
Heliozelidae 
Hepialidae 
Hesperiidae 
Hyblaeidae 

613 
235 
267 
598 
745 
98 
73 
1037 
14 
882 
102 
234 
803 
85 
2009 
874 
36734 
3 
36 
4476 
62 
7 
1507 
88 
78 
188 
264 
1 
2 
101 
133831 
251 
1417 
124 
19544 
138908 
255 
2951 
130 
196 
376 
95941 
1 

 52

0.059 
0.023 
0.026 
0.058 
0.072 
0.010 
0.007 
0.101 
0.001 
0.085 
0.010 
0.023 
0.078 
0.008 
0.195 
0.085 
3.562 
2.909 e -4 
0.003 
0.434 
0.006 
6.787 e -4 
0.146 
0.009 
0.008 
0.018 
0.024 
9.696 e -5 
1.939 e -4 
0.010 
12.976 
0.024 
0.137 
0.012 
1.895 
13.468 
0.025 
0.286 
0.013 
0.019 
0.036 
9.302 
9.696 e -5 

Table 3.1 (cont’d) 
 
Immidae 
Incurvariidae 
Lacturidae 
Lasiocampidae 
Lecithoceridae 
Libytheidae 
Limacodidae 
Lycaenidae 
Lymantriidae 
Lyonetiidae 
Meesiidae 
Megalopygidae 
Micropterigidae 
Mimallonidae 
Momphidae 
Nepticulidae 
Noctuidae 
Nolidae 
Notodontidae 
Nymphalidae 
Oecophoridae 
Opostegidae 
Papilionidae 
Phyllocnistidae 
Pieridae 
Plutellidae 
Praydidae 
Prodoxidae 
Psychidae 
Pterolonchidae 
Pterophoridae 
Pyralidae 
Riodinidae 
Saturniidae 
Satyridae 
Schreckensteiniidae 
Scythrididae 
Sesiidae 
Sphingidae 
Strathmopodidae 
Symmocidae 
Thyatiridae 
Thyrididae 

24 
31 
229 
4587 
39 
82 
4768 
59375 
61 
52 
23 
1231 
6 
220 
392 
356 
168016 
2947 
12908 
97016 
1578 
74 
31562 
1 
67394 
533 
26 
475 
706 
6 
2357 
13065 
2585 
17640 
30 
115 
335 
5885 
26468 
3 
2 
40 
377 

 53

0.002 
0.003 
0.022 
0.445 
0.004 
0.008 
0.462 
5.757 
0.006 
0.005 
0.002 
0.119 
5.817 e -4 
0.021 
0.038 
0.035 
16.290 
0.286 
1.252 
9.415 
0.153 
0.007 
3.060 
9.696 e -5 
6.534 
0.052 
0.003 
0.046 
0.068 
5.817 e -4 
0.229 
1.267 
0.251 
1.710 
0.003 
0.011 
0.032 
0.571 
2.566 
2.909 e -4 
1.939 e -4 
0.004 
0.037 

Table 3.1 (cont’d) 
 
Tineidae 
Tischeriidae 
Tortricidae 
Tridentaformidae 
Uraniidae 
Urodidae 
Xyloryctidae 
Yponomeutidae 
Ypsolophidae 
Zygaenidae 
N/A 
 
 
 

7510 
313 
50546 
10 
192 
252 
2 
943 
258 
628 
31 

0.728 
0.030 
4.901 
9.696 e -4 
0.019 
0.024 
1.939 e -4 
0.091 
0.025 
0.061 
0.003 

The number of specimens collected per square kilometer in states with a LepNet 

institution is drawn from a different distribution than the same metric in states without a 

LepNet institution (P = 0.005694).  In other words, the LepNet and non-LepNet states are 

statistically different populations.  Overall, there is a dearth of specimens collected 

throughout the middle of the United States, especially the states composing the Great Plains 

(Figure 3.2).  For the vast majority of the fifty states, less than one half of a specimen has 

been collected and databased per square kilometer (Figure 3.3).  Other states that are 

particularly lacking sampling of Lepidoptera include Alaska, Nevada, Kentucky, and 

Virginia.  The states with such low numbers of specimens per square kilometer are mostly 

those without an institution participating in LepNet; specimens may exist in collections, but 

they have not been added to the SCAN database.  If specimens from states that are 

underrepresented in the SCAN sample were in collections that are not databased, this 

would potentially change the pictured spatial distribution of US Lepidoptera in a significant 

way. 

 

 54

 

Figure 3.2: National distribution of Lepidoptera specimens held in institutional collections 
across the country normalized by state area in square kilometers.  States marked with a star 
have at least one institution participating in LepNet.  
 

 

The number of collectors who joined the Lepidopterists’ Society between 1947 and 

2016 is pulled from a different distribution than the number of specimens collected per 

year over the same time period (P < 2.2 e -16).  There is not a significant association 

between Lepidoptera Society membership numbers and the number of specimens collected 

for each year.  We also compared the collection efforts of avocational and vocational 

collectors over time.  The total number of specimens collected by avocational collectors 

was 487,152 in the top 10% of collectors, which is more than the 346,740 collected by 

vocational collectors (Figure 3.4).  The overall collection efforts of avocational collectors 

were significantly different from that of the vocational collectors (W = 74549054, P = 

0.01111).  

 55

 

                                                                                                                                                                    

Figure 3.3: Graph of Lepidoptera specimens held in institutional collections across the 

country normalized by state area in square kilometers. 

 

 56

 

Figure 3.4: Avocational and vocational specimen collection events, 1963-2008 
 
 
Between the years of 1963 and 1989, there was a significant difference between the 

numbers of specimens collected by avocational as compared to vocational collectors (Table 

3.2).  After 1989, there was not a statistical difference between the collecting effort of 

avocational and vocational collectors (Table 3.2). 

 
Table 3.2: Avocational vs. vocational collectors: comparison of specimen numbers by decade 
 
Year Range 
1963-1972 
1964-1973 
1965-1974 
1966-1975 
1967-1976 
1968-1977 
1969-1978 
1970-1979 

Interpretation 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 

P-value 
0.0002879* 
0.0002879* 
0.0002879* 
8.227 e-5* 
4.114 e-5* 
8.227e-5* 
4.114 e-5* 
4.114 e-5* 

 57

Table 3.2 (cont’d) 
 
1971-1980 
1972-1981 
1973-1982 
1974-1983 
1975-1984 
1976-1985 
1977-1986 
1978-1987 
1979-1988 
1980-1989 
1981-1990 
1982-1991 
1983-1992 
1984-1993 
1985-1994 
1986-1995 
1987-1996 
1988-1997 
1989-1998 
1990-1999 
1991-2000 
1992-2001 
1993-2002 
1994-2003 
1995-2004 
1996-2005 
1997-2006 
1998-2007 
1999-2008 
 
 
 
Discussion 

4.114 e-5* 
4.114 e-5* 
4.114 e-5* 
8.227e-5* 
0.0001645* 
0.0002879* 
8.227 e-5* 
8.227 e-5* 
0.0004936* 
0.00399* 
0.09391 
0.3401 
0.8633 
0.4894 
0.3865 
0.3865 
0.5457 
0.8633 
0.8633 
1 
0.6665 
0.8633 
1 
0.9314 
0.5457 
0.3865 
0.1359 
0.06253 
0.06253 

Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
Significant difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 
No difference 

In the past 150 years, there has been a significant increase in specimen acquisition 

with a peak in 1996 followed by an apparent decline in collecting events across the United 

States (Figure 3.1).  This decline in collection events is most prominent in the last 15 years.  

The more than 1 million specimens are not evenly distributed across the United States 

(Figure 3.2).  There is a significant difference in the number of databased specimens 

 58

between states that have at least one institution associated with LepNet and those that do 

not, suggesting that the database is geographically under-sampled.  Avocational collectors 

have contributed more specimens to institutional collections than vocational collectors 

(Figure 3.4).  However trends in their collecting are not significantly associated with 

membership in Lepidopterists’ Society.  Overall, there is a significant difference between 

the two groups but for some periods in the last 65 years, the efforts of avocational and 

vocational collectors varied. 

Given their aesthetic appeal, Lepidoptera are potentially the most collected insect 

group throughout history.  The appeal of Lepidoptera may inspire people to start 

collections, regardless of whether or not such collectors become professional 

entomologists or remain hobbyists.  Overall, the collection of 17 million US specimens 

(Seltmann et al. 2017) relied in part on the efforts of amateurs and those in the process of 

acquiring scientific training.  Collections of butterflies and moths may reflect the best 

dataset of ecologically important insects available to scientists, although their numbers in 

space and time are not evenly distributed.   

The geographical gaps in collecting are striking (Figure 3.2).  Most of the states are 

represented by < 0.5 specimens/km2.  In part, this maybe an artifact of state size—larger 

states require more specimens for a similar proportion of specimens as compared to 

smaller states.  For example, though the Kenelm Philip collection of over 83,000 arctic 

Lepidoptera specimens (Bakker 2014) is included in SCAN, Alaska is still represented by 

less than 0.05 specimens/km2 due to its sheer size.  In other cases, a lack of funding for 

databasing correlates with fewer databased specimens.  For example, no Texan institute 

was part of LepNet (Seltmann et al. 2017), and only 360 specimen records collected in 

 59

Texas from Texan institutes exist in SCAN database.  In contrast, the Texas A&M Insect 

Collection houses the Roy Kendall Lepidoptera collection, which contains approximately 

100,000 specimens from the southwestern United States (AgriLife Today 2004).  Funding 

for databasing this collection alone would have a significant positive impact on 

documenting the geographic occurrence of southwest Lepidoptera, though it would likely 

not have a significant effect on overall trends in collecting activity.  

The accumulation of US specimens greatly increased in the years following World 

War II.  Following the establishment of the GI Bill, the number of years men spent in 

postsecondary educational levels increased by an estimated 15-20%.  Nearly 70% of men 

who turned 21 between 1940 and 1955 attained a college education for little to no cost 

(Stanley 2003).  Many veterans who made use of the GI Bill studied the sciences (Bennett 

1996, Altschuler and Blumin 2009), and were likely inspired to continue collecting as a 

hobby and the addition of their student collections to institutional collections may have 

contributed to overall collection growth.  

According to the US Department of Education, 11% of undergraduate students 

starting in academic year 2003-2004 declared a biology major at some point between 2003 

and 2009 (Chen 2013).  College attendance is, in general, on the rise; enrollment in the fall 

semester was 12% higher in 2016 than it was in 2006 (Snyder et al. 2019).  The actual 

completion of bachelor’s degrees is also on the rise, including in the fields of agriculture 

and natural resource fields, with a 29% growth from 2010-2011 to 2015-2016, through 

growth in health-related fields is more than double that of agriculture and natural 

resources (Snyder et al. 2019).   

 60

Despite this growth, undergraduate biology majors are overwhelmingly choosing 

further education outside of biology.  In the academic year 2015-2016, 73,700 doctoral-

level degrees in health profession-related fields were granted in the United States as 

compared to 7,900 doctoral degrees in biology and biomedical sciences (Snyder et al. 

2019).  Given that relatively few students are pursuing advanced studies in biology and 

natural science, these ongoing educational shifts have serious implications for the growth 

of institutional collections.  Though college attendance is on the rise, comparatively few 

students are choosing fields where collections would be emphasized, decreasing potential 

levels of natural history training and eagerness to collect. 

The accumulation of specimens shows significant recent declines (Figure 3.1). 

Though Lepidoptera collecting increased dramatically from the late nineteenth century 

through the 1970s, collection events have plummeted since the late 1990s and this trend 

has become more pronounced from 1999 to 2009 (Figure 3.1).  Lepidoptera collecting is 

not alone in this decline; many herbaria have also hit a peak in annual specimen collection 

events (some of those peaks occurring in the 1990s) before declining dramatically (Prather 

et al. 2004a).  This decline may be associated with the number of students studying the 

natural sciences at the undergraduate college level; by 1990, interest in a number of 

scientific majors had greatly declined (Green 1989).  This trend in post-secondary 

educational choices, particularly in fields relating to natural history, continues today—both 

at the undergraduate and graduate level—and likely impacts the growth of natural history 

collections (Tewksbury et al. 2014).  

The decrease in Lepidoptera specimen collecting has the potential to greatly impact 

future research studies by limiting temporal data and geographic locations represented for 

 61

any given species of butterfly or moth.  Historical collections document environmental 

conditions and species distributions at a particular place and time.  Over time, such 

collections become more and more valuable sources of information to researchers, as the 

collections become increasingly irreplaceable and arguably more meaningful in studies 

dependent on a long timescale.  Malaney and Cook (2018) used such historical collections 

of mammals to investigate collection and accession efforts for mammalian specimens over 

time and across the United States; they reported a decline in such efforts over the last 

couple of decades.  Compared to mammals, insects are easier to collect, requiring relatively 

less preservation effort and few to no permits.  Despite the ease and appeal of collecting   

lepidopterans, declines in insect collecting follow those observed in many areas of biology 

that have historically added many biological specimens to museum collections (Prather et 

al. 2004a, Gardner et al. 2014).  The continued existence of specimens and collection 

records has many implications for the biological sciences; there are many areas of research 

that will depend on specimens into the future, including uses for these data that we cannot 

predict or foresee (Heberling et al., in press).   

Continued collection of biological specimens is critical for a greater understanding 

of biodiversity and environmental change (Prather et al. 2004b).  Through the efforts of 

avocational and professional entomologists, historical collections of biological specimens 

provide the data to track changes in patterns of biological diversity (Hill et al. 2014; Lubar 

2017; Short et al. 2018; Heberling et al., in press).  The availability of collection and 

specimen information in public databases allow researchers to a) observe morphological 

features in imaged specimens, preventing potential damage through the transfer of 

specimens among institutions and hastening the description of insect biodiversity before it 

 62

is lost, and b) to use large amounts of data to describe broad trends often with statistical 

significance (Short et al. 2018).  For example, skilled German amateurs were the first to 

quantify a continuing decline in insect biomass based on decades of saved samples from 

traps, and various scientific institutions in Europe are also making use of avocational 

entomologists to attempt further study into steep rates of insect decline (Hallmann et al. 

2017).  Professional scientists have also identified such trends over the course of their 

fieldwork (Lister and Garcia 2018).   

The importance of research into the natural history of organisms—long considered 

to be a hallmark of the amateur—has recently come to light despite a decades-long decline 

in such efforts (Tewksbury et al. 2014; Heberling et al., in press).  In some cases, this 

decline has been linked to a lack of funding for research into the natural history of 

organisms and a coinciding decline in the teaching of skills needed for natural history in 

university settings  (Tewksbury et al. 2014, Hiller et al. 2017).  Given that less than 2% of 

institutionally-held insect specimens have been databased (Short et al. 2018), there is a 

need for increased maintenance, growth, and accessibility of natural history collections.  In 

the United States, there is limited funding to support natural history collections, despite the 

need for curatorial staff, digitization of specimen records, and space (Dalton 2003, Prather 

et al. 2004b, Snow 2005).  This dearth of funding greatly impacts aspects of biological 

research that rely on geographical and time series data and often excludes additional and 

rare data from smaller collections (Snow 2005).   

Our results suggest urgency for continuation of funding for databasing of museum 

specimens and greater engagement of the public in collection-based sciences.  Without the 

availability of specimen data to researchers—particularly data pooled from multiple 

 63

sources and an increase in specimen acquisition to pre-1990 levels—investigations into 

many areas of critical concern to the biological sciences will, at best, be stalled for an 

unknowable amount of time. 

 

 

 

 64

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

 

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