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FreeHistorical Comment

“Her Joyous Enthusiasm for Her Life-Work …”: Early Women Authors in The American Naturalist

Abstract

Women have long been underrepresented in the natural sciences, and although great progress has been made in recent decades, many subtle and not-so-subtle barriers persist. In this context, it is easy to get the impression that the early history of ecology and evolutionary biology was exclusively the domain of male researchers. In fact, a number of women made very substantial contributions to The American Naturalist in its first decades. In a follow-up to a series of retrospective essays celebrating 150 years of this journal, we highlight the scientific contributions of the women published in it during its first 50 years (1867–1916). We also discuss the diverse paths that their scientific careers took and the barriers they faced along the way.

Online enhancements:   appendix tables.

In the countdown to its recent 150th anniversary, The American Naturalist published a series of retrospective essays highlighting important but often overlooked articles from the journal’s history. With only two exceptions—LoPresti and Weber (2016) and Metcalf (2016)—the highlighted articles were all authored by men. It is thus easy to come away with the impression that the journal (like the natural sciences in general) was an all-male club in its early years. The vast majority of articles, including almost all the classics that are still actively discussed today, were indeed written by men.

However, women scientists made significant scientific contributions throughout The American Naturalist’s early years. The goal of this historical commentary is to recognize the women who contributed during the first half-century of the journal (1867–1916). We highlight their research contributions but also use this opportunity to explore their fascinating lives and what they reveal about gender disparities during the first 50 years of professional science in North America. How often did women publish in The American Naturalist during this early era? What were their stories? How did they enter science? Were they able to establish scientific careers? How did their backgrounds and fates change over the 50-year period that we cover? The answers to such questions both teach us about the history of our field and reveal familiar themes that echo in modern discussions of inequality in academia (see, e.g., Moss-Racusin et al. 2012; Larivière et al. 2013; Holman et al. 2018). Although here we focus exclusively on the first half-century of this journal, readers will readily identify parallels to the barriers and biases that persist for scientists who are women, minorities, first-generation college students, or from developing nations.

How Often Did Women Publish in The American Naturalist in Its First 50 Years?

To explore the early history of women authors in this journal, we scanned every issue published from 1867 to 1916 (580 issues, 2,889 articles). Of these, 76 (2.6%) had a female author (88% of these were sole-authored, typical of the era). Female representation among authors increased only incrementally over the period we consider (fig. 1).

Figure 1.
Figure 1.

Proportion of female-authored articles in The American Naturalist, 1866–2018, counting all authors (filled circles, thick black line) or first authors only (open circles, thin black line). Each filled circle from 1870 to 1955 summarizes all articles in cumulative 5-year increments. From 1955 onward, when issues contained more articles, we only counted articles every fifth year (>100 articles per volume). Part of the rise in female-authored articles could reflect increasingly long author lists, in addition to increasing proportions of female authors. To adjust for this, we also present the proportion of articles whose first authors were women (open circles, representing 5-year intervals using only articles in the first issue of each year). Prior to 1960, most articles were written by a single author, so the two lines are approximately the same. We did not count articles by authors whose gender we could not determine (e.g., when only initials were given). This may cause us to underestimate female authorship if women were more likely to omit their first name from their publication, although this does not seem to have been a common practice (see text).

We read each female-authored or coauthored article from this period, noting the topic, general approach, major conclusions, and institutional affiliation (if any) of the author. An annotated list of articles authored by women between 1867 and 1916 is provided in Table A1. We then searched books, journals, and websites for information about the authors’ lives. Brief biographical information and sources can be found in Table A2.

These 76 articles were authored by 57 different women; that is, some women published more than once—a few rather prolifically—in the journal. It is possible that we missed articles by women who published under a pseudonym or their first initials. However, the large majority of authors in this era of the journal published with their full names, with the exception of certain extremely well-known men (including the journal’s founders), so we doubt that we missed many female-authored works.

These women were almost exclusively American, although many studied in Europe (where some institutions granted PhDs to women before American institutions did). International authors appear periodically from the early years of the journal but became commonplace only toward the end of our focal era. It is difficult to identify other aspects of diversity.

Who Were the Earliest Women Authors?

To give these early scientific pioneers the credit they deserve, we start by presenting the first three women (shown in fig. 2) to publish in this journal. We describe their backgrounds, research contributions, and how these reflected the scientific themes of the time. These stories are similar in some respects but also highlight the diversity represented in this very small pool of extraordinary scientists.

Figure 2.
Figure 2.

First three women authors in The American Naturalist.

Lucie L. Hartt was the first woman to publish in The American Naturalist, in its third year, 1869. She was personally connected to the journal through her husband, Charles Frederick Hartt, a prominent Canadian geologist who studied alongside the journal’s founders at Louis Agassiz’s museum at Harvard. Lucie Hartt apparently visited her husband-to-be while he was the founding director of the Section of Geology at the National Museum of Rio de Janeiro (1866–1867); the two married in 1869 and then returned to Brazil in 1870. Lucie Hartt’s 1869 article describes general information about cuttlefish natural history and her personal observations of one individual cuttlefish. It includes an endearing passage about how the cuttlefish in question, an “uncivil individual,” seized her finger “with a pressure far too ardent to be agreeable” (p. 33) and would not let go. Her writing was typical of many articles from that time: entertaining, personal, and inquisitive but focused on describing observations of organisms. (Interestingly, she published an almost verbatim article in the October 2, 1869, issue of Scientific American under a different title. Clearly, copyright and plagiarism rules have changed.) Lucie Hartt apparently stopped contributing to science soon thereafter. She gave birth to twins and then became pregnant again while in Brazil, where her husband was looking for geological evidence of Agassiz’s theory that the biblical flood left glaciation-like signatures in both the tropic and temperate zones. She later apparently left her husband because he worked too hard: a brief genealogical essay on Charles states that “his decision to put geology first had cost him dearly.” He died of yellow fever a few years later, in 1878. Lucie Hartt had her estranged husband’s remains returned to New York, where she ran a girl’s school that emphasized college preparation.

Grace Anna Lewis (1821–1912) published two articles in the fourth and fifth volumes (1870 and 1871). In contrast to Lucie Hartt’s personal anecdotes, but emblematic of another writing style of the time, Lewis offered textbook-like descriptions. One article described the lyrebird in Australia, while the next documented microscopic structures in bird feathers. In contrast to Lucie Hartt’s, Lewis’s life is well documented. She was a pioneer on many fronts: as a female American scientist, an abolitionist, a suffragist, and a temperance activist. Her Quaker family placed her in a boarding school after her father died. The Quakers were early proponents of girls’ education, as well as abolition and women’s right to vote. Consequently, several early women scientists were also active in these social movements. Lewis, who never married, learned to be an accomplished naturalist and illustrator. After the Civil War, at age 39, she turned her attention to ornithology, supported by her patron and teacher John Cassin, curator of the Philadelphia Academy of Natural Sciences. She published a widely acclaimed book, The Natural History of Birds. It was intended to be the first of a several-volume magnum opus, but Cassin died in 1869 of arsenic poisoning from handling preserved bird skins, and Lewis, without his sponsorship, saw her access to the Academy’s museum curtailed. Yet her expertise was widely acknowledged: T. H. Huxley, visiting the United States in 1876, judged her work as “fully up to the latest and most advanced ideas in regard to systematic classification.” For a time, Lewis supported her own research by selling off her family’s land and by periodically giving private lectures on ornithology in her parlor. She applied to many university jobs but was unable to obtain a steady teaching position that would allow her to continue her research. It is hard to say how much this reflected the pervasive sexism of the time or her lack of formal education beyond high school, but the two factors certainly went hand in hand. She ultimately took a teaching job at the Foster School for Girls and never published again.

Mary Treat (1830–1923), the third woman to publish in The American Naturalist, had a long and prominent scientific career, unlike her predecessors. She published five articles in the journal between 1873 and 1881 on diverse topics, including carnivorous plants, sex determination in butterflies, tarantula behavior, and flycatchers. Like Grace Anna Lewis, Mary Treat dove into research relatively late in life: she published her first article (in The American Entomologist) when she was 39 years old. Her work began in collaboration with her husband, Joseph Treat, an abolitionist and professor, and then continued after they separated. She ultimately published 76 scientific articles and five books, one of which (Injurious Insects of the Farm and Field) was reprinted five times. She corresponded avidly with Asa Gray and Charles Darwin, who engaged in a spirited debate with “Mrs. Treat” (as she was generally called) about control of sex in insects. Although Darwin disagreed with her, his serious attention to her ideas legitimized her in the eyes of many other scientists of the day.

These first women authors were under immense societal pressures to conform to a culturally specific vision of womanhood that is hard for us to fathom today. The title of our article comes from a contemporary description of the spectacularly successful Mary Treat (Harshbarger 1899). There is supreme irony, then, in reading the full passage from which we took it:

Her most prominent characteristic is a modesty so shrinking as to make any public recognition of her services painful to her, while her joyous enthusiasm for her chosen life-work is so great and so contagious that her home is always a centre of attraction, where are welcomed all who care to learn even the alphabet of her beloved book of nature, and where she dispenses the bounty of her gifts and attainments with a modest lavishness and an unwearied patience, which appears to be to her their own reward. (pp. 300–301, italics added for emphasis)

How Did Early Women Authors in The American Naturalist Enter Science?

Hartt, Lewis, and Treat, our three pioneers, were soon followed by a small but steadily growing list of other female authors (Table A1). They offered novel observations on topics ranging from botany to zoology to anthropology, conducted experiments, and presented literature reviews; they worked both nationally and internationally, at prominent field stations, and in their own backyards. In these ways, they represented a typical cross-section of American Naturalist scientists of their time, and indeed of our own time. However, their experiences in science often had little in common with their male peers. By exploring the lives of this diverse set of women, we can glean some common themes about what it was like to be a female scientist in America in the mid- to late-1800s and how their experiences changed as science was increasingly professionalized in the early 1900s.

Then, as now, people are drawn into science by a combination of their own innate curiosity and by the example and guidance of mentors. Today, most people meet their academic mentors during their undergraduate and graduate studies. This was also true for men in the era we consider here. Most notably, when the Swiss naturalist Louis Agassiz moved to Harvard University, he mentored a vast number of students spanning all areas of the natural sciences. These students (all men) went on to become the founders of many academic disciplines (and started The American Naturalist, in revolt against their mentor’s strict rule; Conklin 1944). But women had very limited opportunities for higher education and hence little access to mentors. Some private colleges began to admit women in the early to mid-1800s (e.g., coeducational Oberlin College was founded in 1833 and all-women’s Mount Holyoke College in 1837). However, these institutions often lacked departments in which the natural sciences were taught. Graduate education opportunities lagged still farther behind. Emily Gregory (1840–1897) was the first American woman to receive a PhD in botany in the mid-1880s, though she had to go to Zurich to do so (she was also the first woman elected to the then invitation-only American Society of Naturalists in 1886). She received her undergraduate degree from Cornell University in 1881 (at the age of 41). Cornell is a common thread linking many of the women publishing in The American Naturalist. It began admitting undergraduate women in 1870 (though they were initially not welcomed by the male students), and its strong natural sciences faculty apparently drew many women into science.

Some women became authors in The American Naturalist with very little formal mentorship. Erminnie Smith (1836–1886), for example, has been called the first woman field ethnographer (Miller 2007). She was the first female member of the New York Academy of Sciences (1877) and the American Association for the Advancement of Science (AAAS). She received a degree in mining in Germany, and then, without formal training, became a self-taught expert on the culture and language of the Iroquois. She started the Women’s Anthropological Society of America after she was barred, as a woman, from membership in her major professional society; this policy was apparently a backlash to the exceptionally large numbers of women anthropologists working in the Washington vicinity in the 1880s (Rossiter 1982). Miller (2007) points out that “for the nation’s pioneer women anthropologists, membership in an organization of like-minded women reinforced their identity as professionals and serious scholars” (p. 75). Networking among women in academia clearly had an early origin.

With few undergraduate and no graduate opportunities, the early women authors found mentors in other ways. Erminnie Smith lacked a professional mentor, but her research was funded by John Wesley Powell, the first director of the Bureau of Ethnology at the Smithsonian Institution and director of the US Geological Survey. A handful of prominent evolutionary biologists mentored large numbers of women, including David Starr Jordan, Thomas Hunt Morgan, William Bateson, and Charles Davenport. These training opportunities, while essential, did not generally translate to professional success: indeed, women were marginalized in most of these labs even when hired into permanent positions (Richmond 2007b). Further, sometimes these mentors’ motives were not entirely admirable. Edward Cope (an early editor of this journal) taught for some time at the Quaker-founded, coeducational Haverford College, where he mentored many young women. For example, Helen Abbott (whom we discuss in greater depth below) went on a paleontological expedition with Cope to the Western United States, chaperoned by her father. She later wrote that Cope believed women required extra education to compensate for what he saw as their innate lack of ability (LoPresti and Weber 2016). An alternative explanation was proffered by a contemporary, Charles Knight, who once said of Edward Drinker Cope that “no woman was safe within five miles of him” (Davidson 1997, p. 109).

Many women were encouraged by husbands or other family members who were themselves studying natural history. These collaborations were an entry point into science but at the same time could be a constraint on advancement (Rossiter 1982). Susanna Phelps Gage (1857–1915) was an embryologist and comparative anatomist who later studied neural development and anatomy. She received her PhD in 1880 from Cornell, where she met her husband, a professor. Although she was productive, pursued independent research, and was highly respected by leading European anatomists (Williamson 1899), Gage never held a formal job commensurate with her abilities. Instead, she supported her husband’s career, for instance, illustrating his articles and editing his book. In 1886, she published a first-authored article with him in this journal on aquatic respiration in turtles. Despite these constraints, she was elected to the AAAS and belonged to several major professional societies. She helped found George Washington University.

Not all husband-wife teams were this asymmetric. Gertrude Davenport (1866–1946) was another frequent American Naturalist author who began by collaborating with her husband, Charles Davenport. Together and separately, they published many articles on heredity in humans. They were among the most prominent eugenicists of their time, using this journal to promulgate many of their more repellent views. Unlike many of her predecessors, Gertrude Davenport held her own academic position (instructor at the Brooklyn Institute of Arts and Sciences). Rosa Eigenmann (1858–1947) began as an independent ichthyologist under David Starr Jordan’s mentorship at Indiana University, where she met and married Carl Eigenmann. They moved together to Harvard, where she was the first woman to take graduate classes. They cofounded a biological station in San Diego, California. Both were curators at the California Academy of Sciences, where they copublished 15 articles from 1888 to 1893. Today, the descriptions of roughly 150 species of fish are credited to “Eigenmann & Eigenmann.” Elizabeth and George Peckham seem to have been a similarly egalitarian couple. They were known for their studies of animal behavior, particularly in jumping spiders and insects. Elizabeth Peckham (1854–1940) obtained her BA from Vassar College in 1876 as one of its first science graduates. Together, the Peckhams published early studies on sexual selection (Peckham and Peckham 1889, 1890), supporting Darwin over Wallace. From 1883 to 1909, they described 63 genera and 366 species of arthropods.

Toward the end of the nineteenth century, women’s access to higher education expanded. However, women still had to fight for formal training. For example, Ida H. Hyde (1857–1945) was a physiologist known for developing a powerful microelectrode. She studied at Cornell University and then was mentored by Jacques Loeb and Thomas Hunt Morgan at Woods Hole Biological Laboratory, a common training ground for women scientists in this era. Hyde then went to Heidelberg University in Germany to train for her PhD with Wilhelm Kühne studying jellyfish development. Heidelberg was her second choice; Strasburg denied her admission on the basis of her gender. Even at Heidelberg, the faculty barred her from entering the lectures and laboratories. Kühne disliked teaching a woman and demanded far more work from her than from male students. Despite these significant hurdles, Hyde succeeded. Her male colleagues shared their lecture notes with her, and she passed her doctoral exam with honors in 1896 (at age 39), becoming the first woman to obtain a doctorate there. She wrote wryly about her experiences in an article entitled, “Before Women Were Human Beings” (Hyde 1938).

By the late 1890s to early 1900s, the list of universities that were training and, to a lesser extent, employing women expanded considerably. We begin to see women authors from Bryn Mawr College, University of Cincinnati, University of Illinois, Lake Forest University, University of Texas, Smith College, University of Michigan, Columbia University, and the University of Chicago. Graduate programs in some subfields, in fact, were dominated by women to a level unmatched today. Langenheim (1996) notes that women received every PhD awarded in ecology prior to 1920 at the University of Illinois and half of those awarded at Cornell. She points out that these figures are particularly significant considering that women did not get suffrage in the United States until 1920. We stress, however, that receiving a PhD and finding a position in academia were two quite different things.

Under What Circumstances Were the Early Women Authors in The American Naturalist Able to Establish Careers in Science?

As we have pointed out, some women published only a single article in The American Naturalist and then dropped off the scientific map (e.g., our first woman author, Lucie Hartt). Others had long careers and multiple articles over many years. There is no doubt they all faced barriers, from the bias of their male colleagues to limited educational and employment opportunities. We illustrate these barriers by describing some of their career trajectories. We then ask how it was that, among the many who either dropped out of science or toiled for decades with little professional support, a few women managed to establish thriving scientific careers.

“Without Work, Life Is Not Worth Living”

One story of unfulfilled promise involves Helen Abbott (1857–1904), who published a visionary article in this journal in 1887 arguing how principles of evolutionary biology could be applied to explain variation in plant biochemistry (LoPresti and Weber 2016). After several years conducting research on that theme, she returned to medicine, lacking the advanced degrees that her male colleagues in biology expected. Eventually, Abbott did return to school to obtain her PhD from Tufts University in 1903. Tragically, she died of influenza a year later, never having obtained the research position she richly deserved.

An advanced degree, though, was not a guaranteed ticket to success. Julia B. Platt (1857–1935) had credentials to impress. She studied embryology at Harvard in 1887 and then conducted research at Woods Hole, Bryn Mawr, the University of Chicago, Radcliffe, Hopkins Marine Laboratory, and at several German universities. She received her PhD in developmental biology in 1898 and then published 12 articles in just 10 years, including one in The American Naturalist in 1899. Most notably, she showed that neural crest cells formed the jaw cartilage and tooth dentine in salamanders. This conclusion was rejected by her contemporaries, who believed that only mesoderm formed bones and cartilage. It took 50 years for her hypothesis to be confirmed. Despite her depth of training and her productivity, she landed none of the academic positions for which she applied. Platt then wrote, “Without work, life is not worth living. If I cannot obtain the work I wish, then I must take up the next best” (quoted in Zottoli and Seyfarth 1994). She moved to Pacific Grove, California (where the American Society of Naturalists has recently held its stand-alone conferences), and became its first female mayor. She is noted for initiating marine protected reserves that were crucial for the survival of the California sea otter (Palumbi and Sotka 2012). We can take some solace from the hindsight that her municipal leadership probably had a bigger impact than most of her male peers’ research ever did, though it is unclear how she herself eventually felt about her life’s path.

Nevertheless, She Persisted

There were women who did succeed in gaining a coveted research or teaching position. Some of their stories are phenomenal examples of persistence, perhaps none more than Lilian Vaughan Morgan (1870–1952). Morgan was an American experimental biologist who made seminal contributions to the genetics of Drosophila melanogaster that cemented its status as one of the most powerful model systems in biology. Yet she remained at the margins of science for decades because of her own husband. After graduating from Bryn Mawr College with honors in biology in 1891, she published a few articles from her graduate work on Anuran development. In 1904, at the age of 34, she married her graduate advisor, Thomas Hunt Morgan. She then went nearly two decades without conducting her own research while she supported her husband’s career and raised four children. Once their children were grown, she resumed research, now focusing on Drosophila genetics, even though her husband refused to collaborate with her. Her achievements include the discovery of the attached-X and ring chromosomes in Drosophila. Despite Thomas Morgan’s admirable record of mentoring women at Bryn Mawr, including American Naturalist authors Annah Hazen, Helen Dean King, Hannah Teresa Rowley, Henrietta Thatcher, and Eleanor Rathbun (see tables A1, A2), he was not comfortable with them in the “exclusive men’s club” that was his own lab (Keenan 1983). It was only when Thomas died (1945) that Lilian Morgan was granted the first paid job of her life, at the age of 76, more than a half-century after her start in research.

Mary Rathbun (1860–1943) was another long-marginalized but ultimately successful scientist. Like many of her generation, she lacked a formal college education. She started her scientific work in 1884, when she became an unpaid assistant to her brother studying crustacean taxonomy at the Smithsonian Institution. After 3 years as an unpaid assistant, she was granted a clerkship at the Smithsonian. After another 28 years of mostly solo work describing more than a thousand new species, during which she published four American Naturalist articles, she was finally promoted to be an assistant curator of Crustacea. Upon retirement in 1915, she was granted an honorary master’s degree by the University of Pittsburgh and a PhD from George Washington University in 1917.

Not all women had to wait so long for a formal job. One of the earliest women scientists to establish a career was the malacologist Mary Willcox (1856–1953). She first studied at the Massachusetts Institute of Technology, then went to Cambridge, and eventually obtained her PhD from the University of Zurich. Despite her mother’s concern that a scientific career meant spinsterhood, Willcox sought a position at the Museum of Comparative Zoology at Harvard. Louis Agassiz’s son denied her application, saying there were “three windows, and a man for each window” (Palmieri 1997, p. 119), but he sponsored her to start the zoology department at Wellesley College in 1883. She was a full professor by 1890 and continued to publish articles on gastropod taxonomy and anatomy and to mentor other women scientists. She published a new species description and two studies in comparative anatomy in this journal between 1901 and 1906. This is not to say that she had an easy life. Palmieri (1997) reports that Willcox resigned at age 54, ostensibly because of ill health but more likely due to guilt over her professional success compared to a sickly sister who never had such opportunities. After her retirement, she lived another 45 years, devoting herself to her sister’s care.

Like Mary Willcox, Ida Hyde held prestigious positions and, spurred by her own difficulties along the way, became a vigorous advocate for women in academia. After obtaining her PhD, she was chosen as a prestigious investigator of residence to study marine invertebrate physiology at the Naples Zoological Station. During this time, she published a comparative study of mammalian hearts in this journal. She was ultimately hired as an assistant professor at the University of Kansas in 1899, where she was the founding chair of the Department of Physiology. She organized a fund (the Naples Table Association) to provide financial aid and professional support to women pursuing scientific research, founded a scholarship at the University of Kansas, and endowed the Ida H. Hyde International Fellowship with the Association of American University Women (AAUW). She argued for everything from equal pay for women to installing women’s’ toilets in the science buildings at her university.

Edith “Becky” Saunders is probably the most dramatically successful early woman author in The American Naturalist. She is not well known among evolutionary biologists today, but she should be. In an obituary in Science, J. B. S. Haldane described her as the “mother” of plant genetics, saying, “It is clear that she and Bateson had independently rediscovered some at least of Mendel’s laws before his work was known to them” (p. 385). Her first article on plant genetics was published in 1897, before Mendel’s work was uncovered. She ran the Balfour Biological Laboratory for Women, where she was able to conduct and publish the most extensive and careful plant genetics crosses in the early days of Mendelian rediscovery. Her 1902 book with Bateson introduced the terms “allelomorph” (later shortened to “allele”) and “heterozygote.” In 1904, The American Naturalist published a review of her book, the first discussion of Mendelian genetics in this journal: “As this is the first extensive post-Mendelian account of hybridization experiments, it may be fairly called epoch-making” (American Naturalist 38:322). In a 1906 awards ceremony where she received the Banksian medal from the Third International Conference on Genetics, Bateson said, “Had it not been for the work that has been done by my friends and pupils—first of all by my colleague Miss Saunders, whose name has been so deservedly honoured to-night […] I could never have dared, without that force behind me, to have asserted that Mendelian research has been and is of the importance that we now know it must possess” (Wilks 1906, p. 75). For her central role in early genetics, she became one of the first female members elected to the Linnean Society, served as its vice president in 1912–1913, and was named president of the British Genetical Society in 1938. She left science to assist the war effort in Europe (exactly what she did is not clear) and was killed in a bicycle accident shortly after returning to England.

“I Have Been Bossed My Entire Life”

It is easy to assume, from our own biases, that the ultimate goal of these early women scientists was a position in academia and that working in a nonacademic setting was a tragic fate rather than a matter of choice. But when the option is available to us, it is important to let these women speak for themselves. Sarah Monks (1841–1926), an authority on many topics in natural history, published in The American Naturalist in 1881 on the behavior and life history of lizards. She obtained her MS at Vassar College in 1871. She never married and taught for 22 years at a variety of small colleges and the Los Angeles State Normal School (now the University of California at Los Angeles). However, Monks seems to have really come into her own after she quit academia and devoted herself wholeheartedly to natural history. She wrote a high school physiology textbook, discovered regeneration in starfish, studied diatoms, wrote poetry, and more generally seems to have had the best time of her life. She was widely known as “the consulting naturalist”: “From the four corners of the earth they come to her, seekers of scientific knowledge. Scholars and students have long made a beaten path to her door, for she is a recognized authority on zoology, botany, geology, and minerology” (Boeckmann 1920, p. 54). She wrote, with obvious satisfaction, “I am pleasing myself as I please. I have been bossed my entire life. First, I had to obey my parents, then the college professors, and later the president and faculty and social life at the normal school” (Boeckmann 1920, p. 54).

Conclusions

Examining the early days of a field serves to remind us of how far we have come but also of how much we owe those who came before us. We have highlighted here the lives and achievements of a diverse, previously overlooked set of our forebears: the women scientists who published in The American Naturalist during its first 50 years (1867–1916). These individuals led rich and complex lives, contributed to the cutting-edge science of their day, and received varying degrees of credit for their work. Sadly, with some notable exceptions highlighted in the text and in Table A2, they have left behind little or no written record, beyond their fascinating contributions to this journal. We hope that readers will be inspired to investigate their lives and scientific contributions further. There are other sets of overlooked authors that deserve attention as well. Not all of these, including racial minorities, are as easily identified as we found our women authors to be, but they too deserve recognition as pioneers who enriched our field in its earliest days.

The lives of women scientists changed markedly across the era we cover here. Our earliest women authors were mostly self-trained naturalists with the leisure, resources, and connections to be able to spend significant amounts of time observing nature and recording what they saw. They were unconnected with universities or museums, except insofar as their husbands, fathers, and brothers were. The late 1800s saw a clear shift, with surprisingly large numbers of women being trained in university labs; a few key institutions (Wellesley, Bryn Mawr, Cornell) and mentors (T. H. Morgan, David Starr Jordan) played outsized roles. However, very few of these women remained in science after publishing their American Naturalist articles, which were often the products of their MS or PhD research. Those that did establish scientific careers often occupied unpaid assistantships for many years; they appear in many cases to have been deeply appreciated by their institutions, but professional advancement did not come with those sentiments. We were struck by how the lives and career trajectories of the women we have explored, for both better and worse, were strongly tied to those of powerful men (for good or ill) but also by the fact that professional networking among women scientists had emerged as early as the 1880s.

The causes of these patterns are complex and reflect broadscale social trends of the era that go well beyond academia—and within academia, well beyond the biological sciences. We refer readers interested in a deeper historical and sociological consideration to several outstanding analyses of early women in science: Rossiter (1982), Abir-Am and Outram (1987), and Schiebinger (1989). Valuable works are available on women’s experiences in certain subfields of biology, including ecology (Langenheim 1996) and genetics (Richmond 2007b), and at certain key institutions, including Wellesley College (Palmieri 1997) and Radcliffe College (Tonn 2017). Finally, biographies and autobiographies have been written of a handful of our authors (Hyde 1938; Warner 1979; Keenan 1983; Miller 2007; Ogilvie 2007; Richmond 2007a; Palumbi and Sotka 2012; LoPresti and Weber 2016).

We are left with many questions. First, did the women of this (or any other) era do science in any fundamentally different way than did the men of the same era? Did they ask different questions, use different forms of reasoning, rely on different tools? We will only note here that the marked topical shift in the journal across this period—from natural history observations toward rigorous laboratory investigations in physiology and genetics—is evident in articles published by both female and male authors. Second, what was the publication experience like for these pioneering scientists: Did women have to struggle more to publish their work, and were they held to any different publication standards? Were certain journals (including The American Naturalist) more, or less, welcoming for them? Third, to what extent did the experience of women scientists in other countries parallel the American trends we have discussed here?

Finally, when did things begin to change?—those things, that is, that did change. At least with regard to publication frequency, the answer is clear (fig. 1). During the period we considered, less than 3% of articles were authored by women. This rate held fairly steady until the mid-1950s, when it began to rise and then accelerate toward its present state. In recent years, women have been lead authors on slightly more than 40% of American Naturalist articles and authors on nearly 70%. Of all 414 articles published from January 2016 through August 2018 (at the time this article was written), 40% had women as first authors, 27% had women among the coauthors, and 33% were authored by men only. It is difficult to know what null expectation to compare these numbers to, because genders are not equally represented at all career stages. But, clearly, more improvements can be made. We should note that during the time covered by these numbers, two of the three editors were women and the journal used mostly double-blind review (except when authors opted out). However, many questions remain. What cultural and political events drove this change, particularly its onset in the 1950s? How important a role did mentorship of women students by women faculty play in this increase?

We hope that this comment has sparked interest in our readers to further explore our past, with the goal of documenting how it led us to where we are today and where we might yet go. The women whose lives and contributions we have highlighted here made real and, in many cases, highly significant scientific contributions. But they did more for us than that: they set the stage for later women by breaking barriers to university admission, professional degrees, and financial resources and by creating networks to support and mentor other women. Sadly, barriers remain for women in science, as well as for other underrepresented groups (see, e.g., Moss-Racusin et al. 2012; Larivière et al. 2013). These historical women’s stories of perseverance (and, often, failure) remind us of how strongly social norms can limit who participates in science and how our science advances, or does not, as a result.

We thank Trish Morse and Betty Smocovitis for discussions about the history of The American Naturalist, as well as Editor Russell Bonduriansky and two anonymous reviewers. This essay grew out of a series of Twitter posts between D.I.B. and Daniel Weissmann about early diversity of authors in The American Naturalist. Weissmann provided pointers to many early women authors, forming the initial foundation for this historical comment. While working on this essay, D.I.B. was supported by National Institutes of Health grant 1R01AI123659-01A1.

Appendix. Supplementary Tables

Table A1.

Articles by women authors published in The American Naturalist during its first 50 years, 1867–1916

YearAuthorTitleCoauthors, if anyVolume (issue): pagesSummary
1869Hartt, Lucie L.A Chapter on Cuttle-Fishes 3(5):257–261Observations of one cuttlefish and general information about cuttlefish natural history
1870Lewis, Grace AnnaThe Lyre Bird 4(6):321–331Textbook-like descriptions of the lyrebird in Australia; no personal observations
1871Lewis, Grace AnnaSymmetrical Figures in Birds’ Feathers 5(11):675–678Observations of feathers under the microscope
1873Treat, Mrs. MaryControlling Sex in Butterflies 7(3):129–132Experiment to test whether how much butterflies ate could determine which sex they matured into; the prominent entomologist Riley later wrote that her article was much discussed and complimented her “thoroughness and exactness”
1873Treat, Mrs. MaryObservations on the Sundew 7(12):705–708Experimental studies of how sundews digest prey; spoken of favorably by other authors
1874Boyce, CarolineThe Robin 8(4):203–208Basic description of robin life history, with some notes on her own observations
1875Treat, Mrs. MaryPlants That Eat Animals 9(12):658–662Observations and experiments on bladderworts
1878Smith, Emily A.The Maple-Tree Bark-Louse 12(10):655–661Economic entomology contribution containing basic pest life history; it is stated that this work is abstracted from a longer entomological article
1878Smith, Emily A.Modes of Spreading and Means of Extinguishing the Maple-Tree Bark-Louse 12(12):808–809Economic entomology contribution; conclusion of her earlier article
1879Treat, Mrs. MaryThe Habits of a Tarantula 13(8):485–489Behavior and life-history observations of a single animal
1880Smith, ErminnieConcerning Amber 14(3):179–190Paper read at the annual American Association for the Advancement of Science meeting; reviews what was known about amber, including its presence in literature
1881Monks, Sarah P.A Partial Biography of the Green Lizard 15(2):96–99Behavior and life-history observations of animals kept in captivity
1881Furbish, K. (Kate or Catherine)Botany: A Botanist’s Trip to “The Aroostook” 15(6):469–475Observations on the 208 species found and illustrated during a 2-month field trip in Maine
1881Treat, Mrs. MaryThe Great Crested Flycatcher 15(8):601–604Observations of mating behavior of one individual
1882Hinkley, Mary H.The Development of the Tree-Toad 16(8):636–639Observations of life histories in nature
1885Martin, LillieA Botanical Study of the Mite Gall Found on the Black Walnut 19(2):136–140Observations of some galls, with fine drawings
1886Gage, Susanna PhelpsAquatic Respiration in Soft-Shelled Turtles: A Contribution to the Physiology of Respiration in VertebratesSimon Gage (husband), professor of anatomy at Cornell University20(3):233–236Experiments and observations combined to argue how turtles respire
1887Abbott, Helen C. de S.Comparative Chemistry of Higher and Lower Plants 21(8):719–730Path-breaking work in chemical ecology; see LoPresti and Weber (2016)
1887Abbott, Helen C. de S.Comparative Chemistry of Higher and Lower Plants (concluded) 21(9):800–810Conclusion of the previous article
1891Hyde, Ida H.Notes on the Hearts of Certain Mammals 25(298):861–863Comparative notes on the hearts of different mammal species
1892Bodington, AliceMental Evolution in Man and the Lower Animals 26(306):482–494Humans (and specific human races) are here argued to have more impressive intellectual capacities than other animals (first part)
1892Bodington, AliceMental Evolution in Man and the Lower Animals (continued) 26(307):593–606Second part of previous article
1893Bodington, AliceLegends of the Sumiro-Accadians of Chaldea 27(313):14–19Anthropology report on the earliest civilization of Western Asia (first part)
1893Bodington, AliceLegends of the Sumiro-Accadians of Chaldea 27(314):105–112Second part of previous article
1893Eigenmann, R. S. (Rosa)Preliminary Descriptions of New Fishes from the NorthwestCarl Eigenmann (husband), ichthyologist, University of Indiana professor, and longtime collaborator27(314):151–154New species descriptions
1894Bodington, AliceThe Parasitic Protozoa Found in Cancerous Diseases 28(328):307–315Offers evidence cancer could be caused by protozoa
1894Claypole, Edith J.The Action of Leucocytes toward Foreign Substances 28(328):316–325Experiments with salamanders to study their immune systems
1895Bodington, AliceInsanity in Royal Families: A Study in Heredity 29(338):118–129An early look at inherited insanity; argues for sterilization
1897Stone, Ellen A.Some Observations on the Physiological Function of the Pyloric Caeca of Asterias vulgaris 31(372):1035–1041Careful lab experiments exploring the physiology of starfish digestion
1898Davenport, Gertrude C.Agassiz’s Work on the Embryology of the Turtle 32(375):187–188Reviews Agassiz’s contributions to this field; published in a memorial issue highlighting Agassiz’s work
1899Platt, Julia B.On the Specific Gravity of Spirostomum, Paramæcium, and the Tadpole in Relation to the Problem of Geotaxis 33(385)31–38Experimental studies on negative geotaxis in several organisms, conducted as part of mentor Charles Davenport’s course in experimental morphology
1899Slater, Florence WellsThe Egg-Carrying Habit of Zaitha 33(396):931–933Observations showing that males carry the eggs in this insect
1900Rathbun, Mary J.Synopses of North-American Invertebrates. VII. The Cyclometopous or Cancroid Crabs of North America 34(398)131–143Taxonomic descriptions
1900Richardson, HarrietSynopses of North-American Invertebrates. VIII. The Isopoda. Part I. Chelifera, Flabellifera, Valvifera 34(399):207–230Taxonomic descriptions
1900Richardson, HarrietSynopses of North-American Invertebrates. VIII. The Isopoda. Part II. Asellota, Oniscoidea, Epicaridea 34(400):295–309Taxonomic descriptions
1900Rathbun, Mary J.Synopses of North-American Invertebrates. X. The Oxyrhynchous and Oxystomatous Crabs of North America 34(402):503–520Taxonomic descriptions
1900Rathbun, Mary J.Synopses of North-American Invertebrates. XI. The Catometopous or Grapsoid Crabs of North America 34(403):583–592Taxonomic descriptions
1900Sampson, Lilian V.Unusual Modes of Breeding and Development among Anura 34(405):687–715Review and observations of life history, some based on animals she reared herself
1900Peckham, Elizabeth G.Instinct or Reason?George Peckham (husband), entomologist, secondary school teacher, and long-term collaborator34(406):817–818Offers evidence that insect used reasoning in how it captured prey, contrary to arguments made by prominent entomologist Fabre
1900Enteman, Minnie MarieVariations in the Crest of Daphnia hyalina 34(407):879–890Describes variation in Daphnia spines and crests, today one of the most prominent examples of phenotypic plasticity; shows variation between lakes, mostly in summer
1901Chamberlain, Lucia SarahPlants Used by the Indians of Eastern North America 35(409):1–10List of plants used by the North American Indians east of the Mississippi River; compiled during a course given to students of Radcliffe College in 1899–1900
1901Rucker, AugustaThe Texan Kœnenia 35(416):615–630Detailed taxonomic, anatomical, and neuroanatomical description of a new scorpion species in Shoal Creek in Austin, Texas
1901Nichols, M. LouiseThe Spermatogenesis of Oniscus asellus Linn., with Especial Reference to the History of the Chromatin 35(419):919–926Two-year study of spermatogenesis in the common woodlouse
1901Willcox, Mary A.A Parasitic or Commensal Oligochaete in New England 35(419):905–909Describes species found in a stream near Wellesley
1902Hazen, Annah PutnamRegeneration in Hydractinia and Podocoryne 36(423):193–200Multiple experiments on regeneration, inspired by a published article of Peebles
1902Johnson, Sarah W.The Course of the Blood Flow in LumbricusJ. B. Johnson of West Virginia University, presumably her husband36(424):317–328Experiments on blood circulation in worms
1902King, Helen DeanThe Gastrulation of the Egg of Bufo lentiginosus 36(427):527–548Laboratory study of development
1902Rowley, Hannah TheresaHistological Changes in Hydra viridis during Regeneration 36(427):579–583Laboratory studies devoted to regeneration
1902Anthony, Maude H.The Metamorphosis of Sisyra 36(428):615–631Observations of insect development conducted at an entomological field station
1902Thacher, Henrietta F.The Regeneration of the Pharynx in Planaria maculata 36(428):633–641Observations on regeneration in Planaria, based on material collected at Woods Hole
1902Snow, Lætitia M.The Microcosm of the Drift Line 36(431):855–864Observations of insects washed up from lakes; early observations of succession and diversity in insect communities
1903Rathbun, Eleanor P.On the Shell of Littorina litorea as Material for the Study of VariationRobert Paine Bigelow, marine invertebrate zoologist and MIT professor37(435):171–184Statistical study of variation in an introduced species, conducted with a goal of testing whether there is ongoing evolutionary change by comparing new shells with older ones; they found too much erosion to get convincing results
1904Townsend, Anne B.The Histology of the Light Organs of Photinus marginellus 38(446):127–151Laboratory study of the structure of firefly light organs
1904Mitchell, Evelyn GroesbeeckOral Breathing Valves of Teleosts, Their Modifications and Relation to the Shape of the Mouth 38(446):153–164Describes teleost breathing valves
1904Pritchett, Annie H.Observations on Hearing and Smell in Spiders 38(455/6):859–867Experiments on two species; some of the earliest work on sensory ecology of spiders
1905Haynes, Julia AnnaThe Angle of Deviation from the Normal Vertical Position at Which Stems Show the Strongest Geotropic Response 39(458):77–85Experiments on multiple plant species to study geotropism and to resolve a controversy in the literature regarding the angle of response
1905Willcox, Mary A.Biology of Acmaea testudinalis Müller 39(461):325–333Excerpt from a monograph on this topic; comparative studies of a marine invertebrate
1905Worthington, JuliaContribution to Our Knowledge of the Myxinoids 39(465):625–663Observations of hagfish behavior and anatomy
1906Willcox, Mary A.Anatomy of Acmaea testudinalis Müller. Part I. Introductory Material—External Anatomy 40(471):171–187Excerpt from a monograph on this topic; comparative studies of a marine invertebrate
1907Williamson, Helen V.Observations on the Natural History of Diving BeetlesJames Needham, professor at Lake Forest College and later Cornell41(488):477–494Describes diving beetle behavior from a campus pond, observing how seven species partition habitats; they discuss swimming speeds, structural adaptations, body size distributions, and provide detailed anatomical descriptions
1908Davenport, Gertrude C.Heredity of Hair-Form in Man 42(498):341–349Examines data on hair morphology from 500 children from 230 families, as well as their parents and grandparents, and proposes a Mendelian interpretation for these data, concluding that straight hair (round cross-section) is recessive to spiral hair (elliptical cross-section); concludes with statements about the outcomes of various human marriages, hinting at strident eugenics views about intermarriage: “Two blue-eyed, straight-haired parents will have only blue-eyed, straight-haired children. Two wavy-haired parents may have straight, wavy- or curly-haired children but the chances for curly hair are slight.” (p. 345)
1908Hubbard, Marian E.Some Experiments on the Order of Succession of the Somites in the Chick 42(499):466–471Mostly inspired by an 1889 article by Julia Platt on the formation of somites during development; to verify past claims, Hubbard surgically damaged somites (using “Miss Peebles’ method”) to determine whether that stopped formation of new somites; the results challenged Platt’s interpretation
1909Davenport, Gertrude C.Heredity of Hair Color in Man 43(508):193–211Suggests that there are no discrete hair color types but rather a continuum of hair colors and then examines inheritance of hair color as if it were categorical; she tries to interpret it as a two-locus Mendelian trait
1909Buckingham, Edith NasonA Light-Weight, Portable Outfit for the Study and Transportation of Ants 43(514):611–614Methods paper offering an improved design for an artificial nest that can be used for field studies of ants
1909Dederer, P. (Pauline) H.Comparison of Cænolestes with Polyprotodonta and Diprotodonta 43(514):614–618Detailed discussion of the morphology of fossil shrewlike mammals (Caenolestidae) and their putative relatives, presenting very detailed skeletal anatomy drawings and description; she uses this to question previously proposed phyletic relationships
1910Davenport, Gertrude C.Heredity of Skin Pigment in Man. I 44(527):641–672Very detailed pedigree-based analysis of the inheritance and genetic architecture of human skin color, the main question being the extent to which human skin color matches Mendelian segregation rules
1910Davenport, Gertrude C.Heredity of Skin Pigment in Man. II 44(528):705–731Part 2 of the previous article
1911McPheters, Lottie E.A Note on Certain Biometrical ComputationsRaymond Pearl, leading mathematical population biologist of the era and a formative figure in ecology45(540):756–760One of the journal’s earliest mathematical/statistical articles; calculates the statistical properties of a linear regression on a log-transformed variable; Pearl published copiously in The American Naturalist in this era
1913Henchman, Annie P.Clonal Variation in Pectinatella 47(558):361–371Describes heritable and plastic phenotypic variation in bryozoans
1913Lacy, Mary G.A Discussion of the Results Obtained by Crossing Zea mais L. (Mais Djagoeng) (-Reana luxurians Dur. -teosinte) and Euchlaena mexicana Schrad 47(560):511–512Analyzes an F2 hybrid cross of maize and teosinte, in which the parental types never reappear; apparently a summary of an article published elsewhere; concludes that the phenotypes produced are not of much practical value
1913Thompson, CrystalThe Variations in the Number of Vertebrae and Ventral Scutes in Two Snakes of the Genus ReginaAlexander Ruthven, University of Michigan zoology professor and eventual university president47(562):625–636Establishes a correlation between the number of belly scales vs. vertebrae in a group of snakes; Thompson was the “junior” researcher who did the actual work
1914Hagedoorn, Ann C.Another Hypothesis to Account for Dr. Swingle’s Experiments with Citrus 48(571):446–448Poses a new hypothesis to describe the results of crosses among types of citrus and outlines an experiment (then still in progress) to test the hypothesis
1915Hodge, Mildred A.Another Gene in the Fourth Chromosome of Drosophila 49(577):47–49Describes the “eyeless” mutation in Drosophila and, based on crosses, suggests it is linked to the gene causing T. H. Morgan’s bent-wing mutation; she interprets this in the context of gene arrangement on chromosomes
1915Metz, Blanche StaffordMutations in Two Species of DrosophilaCharles Metz (husband), geneticist, member of T. H. Morgan’s Drosophila group49(579):187–189Describes mutants in lab-bred Drosophila (after six generations) with abnormal wing venation; they showed a curious 1∶2 ratio of phenotypes and attributed this to the inviability of one homozygote; the mutant arose independently in two separate Drosophila species in the lab; the authors propose that these independent mutations in different species with different chromosomal arrangements might make it possible to determine which chromosomes were homologous across species
1915Wheldale, MurielFlower Pigments 49(580):256Describes the biochemical steps to purify some floral color chemicals; proposes that red and magenta flower colors can be explained in terms of flavones and anthocyanins
1915Platt, Emilie LouisThe Population of the “Blanket-Algae” of Freshwater Pools 49(588):752–762Describes the algal and animal communities in algal mats from stagnant pools; descriptive natural history applied to discuss community diversity; explicitly discusses the concept of the food web (though not in those terms) and nutrient flows through the community: “This brief account of some of the feeding-habits will serve to show how much all the members of this society are dependent upon the others, and, at the same time, are in constant danger of extinction. Each form acts as a check upon too rapid multiplication of some other form.” (p. 762)
1916Saunders, Edith Rebecca (Becky)On Selective Partial Sterility as an Explanation of the Behavior of the Double-Throwing Stock and the Petunia 50(596):486–498Author had previously published a paper elsewhere on a rare phenotype of Petunia flowers (“double-throwing”), suggesting a “factorial scheme” to explain it (epistasis between two partially linked loci); a male critic published an article questioning her interpretation; here she responds to this “mansplaining” by vigorously pointing out the flaws in his logic
1916Saunders, Edith Rebecca (Becky)The Results of Further Breeding Experiments with Petunia 50(597):548–553Reports new data (5,000 plants from crosses of five lines) building on a 1910 article focused on describing the inheritance of “single” vs. “double” flower morphology in Petunia

Note. Articles are listed in chronological order.

View Table Image
Table A2.

Biographical information on women publishing in The American Naturalist during its first 50 years, 1867–1916

AuthorDatesBiographical informationSources
Hartt, Lucie L.*?Married Charles Frederick Hartt, a student of Agassiz’s who wrote frequently in the journal, in 1869. Husband was a geologist, paleontologist, and naturalist who worked extensively in Brazil and died there of yellow fever at 38. She had left him by then because he worked too much.https://en.wikipedia.org/wiki/Charles_Frederick_Hartt; http://www.myhartt.com/graves.php?ID=679&NAME=Charles%20Frederick%20Hartt
Lewis, Grace Anna*1821–1912Naturalist, illustrator, and social reformer. An expert in the field of ornithology, she is remembered as a pioneer female American scientist as well as an activist in the antislavery, temperance, and women’s suffrage movements. Her life is well documented.Warner 1979; https://en.wikipedia.org/wiki/Graceanna_Lewis
Treat, Mary*1830–1923Naturalist and correspondent with Charles Darwin. One of four major female US botanists pre-1880. Her life is well documented.https://en.wikipedia.org/wiki/Mary_Treat; Rossiter 1982
Boyce, Caroline?Said by Samuel Lockwood to have written “very pleasantly on the robin.” No biographical information found.Lockwood 1888
Smith, Emily A.?American entomologist; little biographical information found. Worked at the US Entomological Commission studying locust outbreaks. 
Smith, Erminnie*1836–1886Geologist and anthropologist at the Smithsonian Institution’s Bureau of American Ethnology; called the “first woman field ethnographer.” Elected the first female member of the New York Academy of Sciences in 1877. Her life is well documented.Rossiter 1982; Miller 2007; https://en.wikipedia.org/wiki/Erminnie_A._Smith
Monks, Sarah P.*1841–1926Referred to as “the consulting naturalist,” she was a recognized authority in many sciences. Discovered regeneration in starfish.Boeckmann 1920; https://en.wikipedia.org/wiki/Sarah_P._Monks
Furbish, Kate (Catherine)1834–1931Botanist, illustrator, and collector of more than 4,000 specimens of the flora, ferns, and mushrooms of Maine (now housed in the Gray Herbarium at Harvard University). An inheritance from her father, who had owned a hardware store, provided enough means for her to pursue her passion. Although she had no formal education, she was renowned for the accuracy of her illustrations and for her intrepid field trips alone to wilderness regions of Maine. Two of her discoveries were named for her: Pedicularis furbishiae (Furbish lousewort) and Aster cordifolius L. var. furbishiae.https://www.britannica.com/biography/Catherine-Furbish; https://en.wikipedia.org/wiki/Catherine_Furbish
Hinkley, Mary H.?Lived in Massachusetts and published natural history observations on frogs and toads. Limited information found on her background or education, but her brief biography makes mention of many scientific contributions. Her father was a well-known artist.Williamson 1899
Martin, Lillie1851–1943Widely known, pioneering woman psychologist who was the first head of a department at Stanford. Early in her career, however, she taught botany and chemistry in an Indianapolis high school, and her contribution to this journal dates from that era. Forced into retirement by Stanford at 65, she went on to become one of the first consulting psychologists. Her career in psychology—but not natural science—is well documented.http://faculty.webster.edu/woolflm/martin.html; https://www.feministvoices.com/lillien-jane-martin/.
Gage, Susanna Phelps*1857–1915Embryologist and comparative anatomist who worked in her husband’s shadow but was widely respected by European anatomists. In the second edition of American Men and Women of Science, she was one of only 25 women to be featured as highly significant in her field.Williamson 1899; https://en.wikipedia.org/wiki/Susanna_Phelps_Gage
Abbott, Helen C. de S.*1857–1904Privately educated chemist and physician. She was a member of many professional societies but never held any formal post.LoPresti and Weber 2016
Hyde, Ida H.*1857–1945American physiologist, mentored by T. H. Morgan. Received her PhD at Heidelberg University and became the first woman allowed to do research at Harvard Medical School. She founded the Department of Physiology at University of Kansas. A significant advocate for women’s opportunities in science. She wrote about her difficulties in a revealing account entitled “Before Women Were Human Beings.” Her life is well documented.Hyde 1938; Rossiter 1982; https://en.wikipedia.org/wiki/Ida_Henrietta_Hyde
Bodington, Alice1840–1897Wrote on a wide variety of topics, including religion, race, marriage, and evolution. She believed that many scientists lacked the ability to express their findings in a way clear and understandable to nonprofessionals and viewed herself as a promoter of science. Her single book was criticized for her comments questioning why writers on science were meant to perform experiments. Though English, she spent most of her life in Canada.https://en.wikipedia.org/wiki/Alice_Brooke_Bodington
Eigenmann, Rosa*1858–1947Ichthyologist trained by David Starr Jordan. Along with her husband, Carl S. Eigenmann, described many North American fish for the first time. After her husband’s death in 1927, she left science to take care of her children. Her life is well documented.Williamson 1899; https://en.wikipedia.org/wiki/Rosa_Smith_Eigenmann
Claypole, Edith J.1870–1915English-American physiologist and pathologist who was head of Wellesley College’s Zoology Department. She eventually left natural science, obtained her MD, and resumed research focusing on medical issues. Her twin sister was also a published scientist. Her life is well documented.https://www.revolvy.com/main/index.php?s=Edith%20Claypole; http://science.sciencemag.org/content/sci/41/1064/754.2.full.pdf; Williamson 1899
Stone, Ellen A.1870–1952Received her bachelor’s from Radcliffe College and her MD from Johns Hopkins. After starting a scientific career, she moved on to become superintendent of child hygiene in the City Department of Heath of Providence, Rhode Island, where she made inroads in improving children’s health. She never married and lived with her sister, who was an architect. She was a noted advocate for careers for women.Creese 2000
Davenport, Gertrude C.*1866–1946Received degrees from Kansas State University and Radcliffe and was an instructor at Brooklyn Institute of Arts and Science. Published extensively with her husband, Charles Davenport, a leading American eugenicist who was a professor at Harvard. Some of her work was on turtles, but mostly she focused on humans, such as her articles in this journal on the genetics of hair traits associated with race. She published two books with her husband on zoology and two solo books on embryology and genetics. She apparently strongly encouraged her husband to promote the idea of forced sterilization of people deemed undesirable.https://en.wikipedia.org/wiki/Gertrude_Crotty_Davenport; https://thenewinquiry.com/blog/white-feminism-and-eugenics-the-case-of-gertrude-davenport/
Platt, Julia B.*1857–1935Highly trained embryologist who did landmark research but failed to obtain an academic position. She eventually left science and became mayor of Pacific Grove, California, where she was instrumental in gaining protection for fragile marine habitats. Her life is well documented.Creese 2000; Palumbi and Sotka 2012; Tonn 2017; https://en.wikipedia.org/wiki/Julia_Platt
Slater, Florence Wells?Little biographical information found. Her work for the journal states that her observations were made at the entomological library at Cornell, from which she graduated in 1884.Creese 2000
Rathbun, Mary J.*1860–1943Did not attend college, worked first as her brother’s assistant and ultimately as a curator at the Smithsonian for decades. She was highly prolific, and her life is well documented; the Smithsonian Institution, in particular, has promoted her landmark achievements.https://en.wikipedia.org/wiki/Mary_J._Rathbun; http://nmnh.typepad.com/no_bones/2015/03/womens-history-month-highlight-mary-jane-rathbun.html
Richardson, Harriet1874–1958Published extensively on marine invertebrates and was known as “the first lady of isopods.” She received her PhD at George Washington University and held a curatorial position at the Smithsonian Institution for decades.https://en.wikipedia.org/wiki/Harriet_Richardson; http://www.jstor.org/stable/1549222?seq=1#page_scan_tab_contents
Sampson, Lilian V.*1870–1952American experimental biologist who made seminal contributions to the genetics of Drosophila melanogaster. In addition to her scientific career, she was involved in science education and was one of the founders of the Children’s School of Science in Woods Hole, Massachusetts. Married to T. H. Morgan, who both facilitated and impeded her career, she received her first research appointment at age 76, upon his death.Keenan 1983; https://en.wikipedia.org/wiki/Lilian_Vaughan_Morgan
Peckham, Elizabeth M. G.*1854–1940Evolutionary behaviorist who collaborated with her husband, George Peckham, her entire career. One of the first science graduates of Vassar College, she, along with her husband, was a secondary school teacher and notable for introducing Darwinian concepts into secondary education. The two were among the earliest taxonomists to emphasize the value of behavior in classification. In 1889–1890, they published among the first studies on sexual selection, supporting Darwin’s concept against Wallace’s alternative explanation of courtship behavior. They described 63 genera and 366 species.https://en.wikipedia.org/wiki/George_and_Elizabeth_Peckham
Enteman, Minnie Marie1872–1955Received her PhD from the University of Chicago in 1901 and published on Daphnia and vespid wasps. Little biographical information found.Creese 2000
Chamberlain, Lucia Sarah?Graduated from Radcliffe College in 1901. No biographical information found.https://books.google.com/books?id=dIYEAAAAYAAJ&q=Chamberlain,+Lucia+Sarah&dq=Chamberlain,+Lucia+Sarah&hl=en&sa=X&ved=0ahUKEwjcv-yUkPfaAhXHrFQKHaYgA4oQ6AEITjAH
Rucker, Augusta1873–1963Physician and invertebrate zoologist. Educated at University of Texas and taught biology there in 1899–1900. She received her MD from Johns Hopkins in 1911, entered private practice in pediatrics, and became known as a pioneer in that field.https://en.wikipedia.org/wiki/Augusta_Rucker
Nichols, Mary Louise1873–1953Received her PhD from University of Pennsylvania. She worked as a high school teacher in Philadelphia but spent summers doing research at Woods Hole on diverse organisms including bees, plants, and crustaceans. Her textbook Science for Boys and Girls was first published in 1924.Ogilvie and Harvey 2003
Willcox, Mary A.*1856–1953Prominent professor of zoology at Wellesley for 27 years, appointed in 1883. She is responsible for securing the presence of physiology in the college. She also wrote an anti-anti-vivisection position paper arguing that biology students sometimes need to kill animals in order to study them.Palmieri 1997; Creese 2000
Hazen, Annah Putnam?Affiliated with Bryn Mawr College and Smith College. Evidently mentored by T. H. Morgan, with whom she published. No other information found. 
Johnson, Sarah W.?Published several articles with J. B. Johnson at West Virginia University; acknowledgments in one article state that the reported observations were made in conjunction with coursework she was carrying out. 
King, Helen Dean1869–1955An expert on rats who developed inbred lines for laboratory genetics use (half of all currently used lab rats derive from her lines). Trained by T. H. Morgan, she was the only US woman to hold a full research professorship between 1910–1920. She was also noted for her writings on eugenics. Her life and contributions are heavily documented.Ogilvie 2007; https://en.wikipedia.org/wiki/Helen_Dean_King
Rowley, Hannah Theresa?Student at Bryn Mawr College with T. H. Morgan. Later wrote Principles of Chemistry Applied to the Household, a text that appears to have been in widespread use. No other information found. 
Anthony, Maude H.?Affiliated with the Biological Laboratory of Lake Forest College, she published in a few entomology journals, and her work was spoken of approvingly by others. She also contributed scientific illustrations to other authors’ works in aquatic entomology. No other information found. 
Thacher, Henrietta F.?Another Bryn Mawr student trained by T. H. Morgan who did research at Woods Hole but disappears from the record afterward. 
Snow, Lætitia M.?A botanist and charter member of the Ecological Society of America, she received her PhD at the University of Chicago and was a professor at Wellesley College.Kohler 2002
Rathbun, Eleanor P.?Associated with MIT, she published extensively on marine invertebrates, occasionally with T. H. Morgan. No other information found; there is no evidence that she was related to her contemporary Mary Rathbun. 
Townsend, Anne B.?Received her BA at Cornell University, where her work for this journal comes. Worked at Woods Hole between 1903 and 1906 and, while there, was listed as a teacher from Philadelphia. Married Charles Howard, who took at professorship at University of Minnesota. No other information found. 
Mitchell, Evelyn Groesbeeck1879–1964African American entomologist who worked extensively with mosquitoes and mosquito-borne diseases but was also known for her work in psychiatry. Received her BA from Cornell and her MD from Howard University. Was a histology and physiological chemistry instructor at Howard. Involved in educational efforts for African Americans. She may have been the first or one of the first African American authors to publish in The American Naturalist.Ogilvie and Harvey 2003
Pritchett, Annie H.?Her work for this journal was listed as a contribution of the Zoological Laboratory of the University of Texas and was submitted as a requirement for obtaining her MS degree. Limited information found. 
Haynes, Julia Anna?Her work was a contribution of the University of Michigan Botanical Laboratory. She was affiliated with University of Michigan and Wellesley College; limited information found. 
Willcox, Mary A.*1856–1953Malacologist and head of the Zoology Department at Wellesley. Well-documented advocate for women in science.Palmieri 1997; Ogilvie and Harvey 2003
Worthington, Julia?Received her BS and MS degrees at University of Cincinnati, where her work for this journal was conducted. No other information found. 
Williamson, Helen V.?Associated with Lake Forest University. No other information found. 
Hubbard, Marian E.1868–1956Professor at Wellesley College. Received her BS (her highest degree) at the University of Chicago in 1894. She also studied salamander behavior and insect genetics, as well as bird embryology. A prominent advocate for women’s suffrage, she wrote many papers to the school’s president about women scientists and their struggles.Palmieri 1997; https://en.wikipedia.org/wiki/Marian_E._Hubbard
Buckingham, Edith Nason?Authored the book Division of Labor in Ants in 1911. A genealogy website describes her this way: “Featherland Farm was owned by Edith Nason Buckingham, a very patrician single lady with a PhD in Entomology from Radcliffe College. Her family was Boston Blue Blood, and some of them were scandalized that she had turned lady farmer.”http://www.worldcat.org/title/division-of-labor-among-ants/oclc/2357693
Dederer, Pauline H.1878–?Zoology instructor at Barnard College, later professor of zoology at the Connecticut College for Women. No other information found. 
McPheters, Lottie E.?Worked at the Maine Agricultural Experimental Station. No other information found. 
Henchman, Annie P.?Worked with Charles Davenport at Cold Spring Harbor. No other information found. 
Lacy, Mary G.?Worked at the Bureau of Plant Industry at the US Department of Agriculture. No other information found. 
Thompson, Crystal1886–1996Earned BA and MA at University of Michigan, then worked in its new Museum of Zoology, first as scientific assistant in charge of fish and invertebrates and later in other positions, including education and extension. Her work forms the basis of the museum’s current public exhibits. She cowrote the first edition of The Herpetology of Michigan (1912).http://um2017.org/2017_Website/History_of_Zoology_Museum.html
Hagedoorn, Ann C.?With her husband, Arend Hagedoorn, formed a team that published a book and many articles together. In their 1921 book, they outlined what would later be called genetic drift. Fisher initially described it as the “Hagedoorn effect,” before Wright gave it its present-day name. No other biographical information found. 
Hodge, Mildred, A.?Published on Drosophila karyotypes and may have trained with T. H. Morgan. No biographical information found. 
Metz, Blanche Stafford1889–1963Geneticist, worked with husband Charles Metz, following him from Columbia University to the Carnegie Institute, to Johns Hopkins, University of Pennsylvania, and Woods Hole. Her husband was quite prominent, but her life in science remains apparently undocumented. 
Platt, Emilie Louise?Cornell graduate whose MS thesis in 1913 produced her article for this journal. No biographical information found. 
Saunders, Edith Rebecca (Becky)*1865–1945Prominent British geneticist who carried out some of the most extensive and careful plant genetics crosses in the early days of Mendelian rediscovery, publishing books and articles with Bateson, Punnett, and others. Her data gave rise to the idea of linkage and was the subject of the first use of a Punnett square analysis. She ran the Balfour Biological Laboratory for Women. She was one of the first female members elected to the Linnean Society and was its vice president in 1912–1913. She became president of the British Genetical Society in 1938.Richmond 2007b; https://www.britannica.com/biography/Edith-Rebecca-Saunders; https://www.nature.com/articles/156385b0.pdf
Wheldale, Muriel1880–1932One of the first women appointed as a lecturer at Cambridge, though the institution would not grant her a PhD (none for women until 1948). She joined Bateson’s lab in 1903 to study the inheritance of snapdragon color. In 1916, she published a book from her very extensive crosses and study of Mendelian segregation of color. In 1914, she changed her focus to flower color biochemistry. She published two other books on plant biochemistry. In 2010, the Royal Institution of Great Britain staged a play, Blooming Snapdragons, about her and three other early twentieth-century women biochemists.Richmond 2007a; https://en.wikipedia.org/wiki/Muriel_Wheldale_Onslow

Note. Asterisks indicate women discussed in the text. Select references are provided.

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Literature Cited

References Cited Only in the Online Appendixes

“Then raising himself on his long limsy arms, he stalked away towards the water, making such a comical figure, that in spite of my fright I indulged in a hearty laugh. ” From “A Chapter on Cuttle-Fishes” by Lucie L. Hartt (The American Naturalist, 1869, 3:257–261).

Editor: Russell Bonduriansky