Monday, December 08, 2025

 

The Florida Museum of Natural History publishes “The Butterflies of California,” a book five decades in the making





Florida Museum of Natural History
Photograph of butterflies 

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The Florida Museum of Natural History has published “The Butterflies of California,” a free online book four decades in the making.

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Credit: Florida Museum photo by Kristen Grace





Key points

  • The Florida Museum of Natural History recently published “The Butterflies of California.” The book was co-written by the late Thomas Emmel (1941-2018), a founder and curator of the museum’s McGuire Center for Lepidoptera and Biodiversity, and two of the Center’s research associates: Tom’s brother, John Emmel (1944-2022) of Hemet, CA, and Sterling Mattoon (1932- ) of Chico, CA.
  • All three authors started collecting and studying California butterflies as children and continued to do so throughout their lives. Their work on the book began in 1974 and was nearly complete, but it remained unpublished following the recent deaths of Tom and John Emmel.   
  • The book draft is now available for free online and contains a history of butterfly collection and research in California, sections on the state’s geology, climate and plant communities, and detailed accounts of nearly 200 species and more than 400 subspecies.

GAINESVILLE, Fla. --- In 1973, brothers Thomas and John Emmel published “The Butterflies of Southern California.” The elder Emmel, Tom, had recently completed his doctoral degree at Stanford University and was working as a biology professor at the University of Florida. His brother John graduated from University of California, San Francisco, with a medical doctorate in 1971 and had just moved to Santa Monica with his wife to begin a residency at the Wadsworth Veterans Administration Hospital.

Both men were consummate naturalists, with a special fondness for butterflies that they’d developed early in their adolescence with support from their parents. From their home base in Los Angeles, the family made regular camping excursions to national parks, and their mother drove the boys to entomology conferences held at the Los Angeles County Museum of Natural History.

The brothers began publishing research articles together on the topic of butterfly ecology when John was still a teenager, and they didn’t stop. Between 1961 and 1973, they co-authored 21 manuscripts, with a primary focus on the diversity and life histories of butterflies in western North America.

After finishing their book on California butterflies, their diverging careers, the demands of John’s family life, and the geographical constraints of working on opposite ends of the country could have ended the brothers’ collaboration. But to assume such would be to underestimate their seemingly limitless passion and capacity for learning about the natural world.

“They both had a remarkable energy and drive,” said Andrei Sourakov, a collection coordinator at the McGuire Center for Lepidoptera and Biodiversity, which Thomas Emmel helped establish in 2004. “But their approaches were very different. While Tom chose Lepidoptera as a career, conducted field work around the world, wrote numerous popular books and engaged in a variety of projects, from population genetics work in Colorado to conservation-related work in Florida, John focused exclusively on the taxonomy, biogeography and ecology of Californian butterflies. John spent all his evening, weekends and vacations on his passion for butterflies, which he pursued as a hobby but with more dedication and methodology than most professionals would.”

“The Butterflies of Southern California” was well received, and a community of scientists and people who spent their free time chasing butterflies encouraged them to expand their focus to the entire state. The Emmel brothers needed little convincing, and they found a willing collaborator in Sterling Mattoon, who grew up near the northern terminus of California’s Central Valley and had nurtured a fascination for butterflies since he was seven. Both John and Sterling continued exploring the most remote corners of the state, including its islands, and frequently did so together. Both maintained close contact and extensive correspondence with other members of lepidopterological community of Western North America. Tom, while busy with numerous academic pursuits, often served as a point person to see their joint work results through the publication process.

Like John, who worked at a private dermatology practice for almost 40 years, Sterling was not a professional Lepidopterist. Most of his career was spent working for the Butte County Agricultural Commissioner’s Office, but his time outside of work was dedicated in great measure to the collection and study of butterflies, an avocation he often shared with his wife.

“My favorite memory is when we were collecting in the Ferndale, CA area where we caught a fantastic species called Speyeria cybele. This unnamed subspecies was later named S. cybele eileenae after my wife, Eileen, who grew up in the Ferndale area. Eileen accompanied me on most of my outings over our married lifetime and became an excellent field biologist,” Sterling wrote.

John also enjoyed the strong support of his wife, Phyllis, in his butterfly studies. “Much of our family life revolved around John’s scientific interest in butterflies, and I tried to make sure that he had the time for his studies,” she said. In 1981, John named a butterfly subspecies in her honor, calling it the Phyllis' Swallowtail. In the paper he wrote, "I take great pleasure in naming this subspecies after my wife…who has provided abundant support and encouragement for my studies of the P. indra complex."

The Emmel brothers and Sterling officially began their work on “The Butterflies of California” in 1974 and estimated it would take them five years to complete. This proved to be a slight miscalculation, as there was always another corner of the state to explore, another subspecies to describe and another life history to illustrate. Decades passed with no end in sight, and the project ultimately outlived two of its authors, Tom and John, who passed away in 2018 and 2022, respectively.

“Yet, short of the illustrations and a few other minor details, the manuscript was practically completed, and many lepidopterists awaited its publication with anticipation,” Sourakov said. “It would have been a shame if it had never seen the light of day.”

Now, more than 50 years after they began, the book draft has officially been published online and is freely available to the public. In its current form, the book clocks in at 854 pages, though it remains incomplete.

The manuscript contains a detailed history of butterfly collections made by Europeans in the region, which originated not with its longtime Spanish colonists but instead with Russian explorers who sailed down out of Alaska in the 18th and 19th centuries. Aspiring prospectors the world over descended on the region not long after. Most of them never found enough gold to cover even their meager living expenses, but among these, at least one — Pierre Joseph Michael Lorquin — made a fruitful career as a butterfly catcher out of his dashed hopes for fabulously incommensurate wealth.

The list contains names and biographies for dozens of additional naturalists, with a few left blank at the time of publication. The book also has detailed notes on the geology, climate and plant communities of California — these being essential for the study of butterflies — and a stark review of the effects humans have had on butterfly populations.

The rest of the book provides detailed accounts for some 200 California butterfly species and more than twice that number of subspecies. During their work on the book, the authors discovered and named several of the former and more than 100 of the latter. The accounts include firsthand observations of development and behavior made by John Emmel and Mattoon, which they acquired by rearing most of the species, from eggs through the caterpillar and pupal stages and on to their metamorphoses into butterflies. This required obtaining live specimens of butterflies and their host plants from across the state and figuring out their growth requirements through a painstaking process of trial and error. This step is one of the primary reasons for what the authors refer to as the project’s long “gestation.”

It should be noted that taxonomic field guides often become outdated soon after they’re published. Scientists are constantly finding new species and deriving an increasingly more accurate picture of how they are related to each other. The distribution of species changes over time, and knowledge gaps are filled in by younger generations of naturalists who are handed the baton by their predecessors in the endless scientific relay race of curiosity and personal discovery.

All the while, the value of books such as these only increases. They are faithful records of our planet’s past and a solid foundation of knowledge for all the work that comes after.

The authors embraced this spirit of continuity. In the last paragraph of the book’s preface, they wrote: “We also hope the appearance of this book will generate a great many independent publications on life histories, foodplant records, and all the basic ecology, behavior, genetics, and evolutionary biosystematic work that a basic knowledge of the California fauna should stimulate and encourage.”

The book is available on the Florida Museum’s website.

 

Parasitic fungus may have emerged 18 million years before the ants with which it lives today




Analysis of 309 strains indicates that the genus Escovopsis emerged 56.9 million years ago, but only began interacting with today’s mutualistic ants 38 million years ago, challenging the theory that they all emerged at the same time.



Fundação de Amparo à Pesquisa do Estado de São Paulo

Parasitic fungus may have emerged 18 million years before the ants with which it lives today 

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An ant (genus Atta) removes fragments of a fungus garden used to feed the colony. The insects’ cleaning behavior functions as a social immune system

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Credit: Quimi Vidaurre Montoya/IB-UNESP





A genus of fungi previously considered a parasite of fungi associated with ants may actually have much more complex ecological functions. According to a study published in the journal Communications Biology and supported by FAPESP, one piece of evidence is that they appeared 18 million years before the ants with which they are associated today.

The results are based on the analysis of 309 strains of the genus Escovopsis collected in eight countries in the Americas. By observing fragments of the genomes, morphological characteristics, geographic distribution, and phylogeny of the species – something like their evolutionary kinship – the researchers were able to determine the relationship with leafcutter ants over 38 million years. Escovopsis emerged 56.9 million years ago.

“Our main hypothesis is that they emerged associated with ancestral groups of fungus-growing ants and then began to coexist with the current leafcutter ants 38 million years ago. Another possibility is that they lived in a different context during those initial 18 million years, outside of their association with ants, as leaf colonizers or degrading organic matter, for example,” explains Quimi Vidaurre Montoya, the first author of the study. Vidaurre Montoya conducted the study as part of his postdoctoral research with a fellowship from FAPESP at the Institute of Biosciences of São Paulo State University (IB-UNESP) in Rio Claro, Brazil.

Leafcutter ants (subtribe Attina) cultivate fungi for food. According to a recent study by the group published in the journal Science in 2024, this mutualistic relationship is thought to have emerged 66 million years ago (read more at agencia.fapesp.br/52936). 

“Our current work is on Escovopsis, a genus of fungi that isn’t cultivated by ants, but is present in the colonies of some species of farming ants and can kill some of their cultivars. Because of this, it’s portrayed as a ‘parasite,’ when in fact only one of 24 species is known to cause infection in the fungus cultivated by ants,” Montoya explains.

The study is part of a project supported by FAPESP through the FAPESP Research Program on Biodiversity Characterization, Conservation, Restoration, and Sustainable Use (BIOTA-FAPESP), coordinated by André Rodrigues, a professor at IB-UNESP and a researcher at the Center for Research on Biodiversity Dynamics and Climate Change (CBioClima), which is one of the Research, Innovation, and Dissemination Centers (RIDCs) supported by FAPESP.

Adaptations

The authors note that Escovopsis underwent morphological and physiological adaptations over evolutionary time, apparently to increase reproductive efficiency and adapt to life inside anthills. These changes mainly occurred in the vesicles that produce conidia, which are structures that perform asexual reproduction.

“The vesicles change from a globular shape in species closer to the common ancestor to a cylindrical shape in more recent species. These changes may have been responses to barriers imposed by the ants or their symbiotic fungi,” Montoya says.

Physiological data indicate that the growth rate, number of vesicles, and production and viability of conidia gradually increased as the genus diversified. Species with cylindrical vesicles may grow faster than those with globular structures. The thinner, elongated vesicles of more recent species produce considerably more viable conidia than older groups with globular vesicles.

“Apparently, there’s a coevolution between ants, symbiotic fungi, and Escovopsis. We don’t know if they evolved to become parasites or if they're opportunists that feed on debris and can eat what remains when the system as a whole collapses. But if it were a specialized virulent host, as part of the literature assumes, it would destroy the system regardless of whether it was in equilibrium or not,” Montoya says.

Little-known fungi

This study is an offshoot of broader work on the genus Escovopsis that began during Montoya’s doctoral studies. He received a scholarship from FAPESP to conduct research under the guidance of Rodrigues. Montoya also completed an internship at Emory University in the United States.

At the time, Montoya analyzed the two largest existing collections of these fungi: the IB-UNESP collection, which was collected and maintained by Rodrigues’s group, and the Emory collection, which was maintained by Professor Nicole Marie Gerardo, Montoya’s supervisor abroad.

One of the first results of that effort was the description of two new genera previously classified as Escovopsis. Two others discovered in that work are still in the process of being described. Montoya and Rodrigues also led the description of 13 new species of Escovopsis, with ten more in the process of being described.

“These fungi are still poorly understood from physiological and ecological points of view. Therefore, it’s premature to treat them all as parasites. Our studies suggest that they have other functions and can live in relative harmony in colonies,” Montoya believes.

According to the researcher, only the Escovopsis weberi species has been proven to cause infections in fungi cultivated by ants. Experiments by his group and others show that several strains did not kill the fungi in the presence of these insects.

In vitro experiments that support the claim of parasitism disregard the fact that the affected fungi rely on ants for protection. Therefore, evaluating the effect of Escovopsis on mutualistic fungi without ants and their hygiene behaviors, which act as a social immune system, would not make sense (read more at agencia.fapesp.br/54199). 

“Some more virulent fungi are immediately removed by ants when inoculated into the colony. In experiments with Escovopsis, however, they don’t give it much importance,” the researcher says.

About São Paulo Research Foundation (FAPESP)
The São Paulo Research Foundation (FAPESP) is a public institution with the mission of supporting scientific research in all fields of knowledge by awarding scholarships, fellowships and grants to investigators linked with higher education and research institutions in the State of São Paulo, Brazil. FAPESP is aware that the very best research can only be done by working with the best researchers internationally. Therefore, it has established partnerships with funding agencies, higher education, private companies, and research organizations in other countries known for the quality of their research and has been encouraging scientists funded by its grants to further develop their international collaboration. You can learn more about FAPESP at www.fapesp.br/en and visit FAPESP news agency at www.agencia.fapesp.br/en to keep updated with the latest scientific breakthroughs FAPESP helps achieve through its many programs, awards and research centers. You may also subscribe to FAPESP news agency at http://agencia.fapesp.br/subscribe.

 

 

GREEN  CAPITALI$M

Illinois research uncovers harvest and nutrient strategies to boost bioenergy profits



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University of Illinois College of Agricultural, Consumer and Environmental Sciences

Harvesting biomass crops 

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New University of Illinois Urbana-Champaign research investigates harvest and nutrient management strategies to attain maximum profitability for biomass growers.

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Credit: University of Illinois Urbana-Champaign





URBANA, Ill. -- To meet ambitious U.S. Department of Energy targets for sustainable aviation fuel (SAF), production of purpose-grown energy crops must ramp up significantly. Although researchers have made substantial progress in understanding the management and conversion of these crops, key knowledge gaps hold the industry back. Now, two new studies from the University of Illinois Urbana-Champaign help fill in the blanks for Miscanthus and switchgrass management. 

“We have come a long way in our understanding of purpose-grown energy crops for SAF, but we still need to optimize agronomic management practices, like harvesting and nutrient management, to reduce production costs and incentivize growers,” said D.K. Lee, senior author of both studies and professor in the Department of Crop Sciences, part of the College of Agricultural, Consumer and Environmental Sciences at U. of I.

Previous studies have been limited in spatial scale or focused on the first years after establishment, early in the perennial life cycle. But Lee’s group is working to provide more realistic, long-term solutions to maximize biomass productivity and grow a more sustainable fuel industry. 

In the first of two recent studies, Lee’s team conducted an economic and environmental analysis of two harvest methods for switchgrass. 

“Harvesting operations account for 60-80% of the total production costs for switchgrass,” said Muhammad Umer Arshad, postdoctoral researcher in Lee’s group and first author on the Bioresource Technology paper. “We wanted to understand why the harvesting cost is so high and how each operation contributes to cost, energy use, and greenhouse gas emissions, as well as identify where reductions are possible.”

Arshad explains that switchgrass harvesting can happen via the stepwise method, in which tasks like mowing, raking, baling, and roadsiding are separated into individual operations; or the integrated method, which uses different equipment to consolidate mowing and raking into one pass. Hypothetically, an integrated approach could reduce effort, energy consumption, and costs. But, after analyzing data from 125 Virginia commercial-scale sites varying in field size and biomass yield, Arshad found a more nuanced answer.

“We found that the integrated method makes more sense for smaller fields (less than 3 hectares, or 10 acres) and low-yield (less than 3.2 tons per acre) conditions, reducing GHG emissions by 9% and energy use by 5%,” Arshad said. “The stepwise method was better for large fields with high biomass yield, reducing harvesting costs to $37.70 per ton and achieving the lowest GHG emissions.”

The costs were estimated assuming the farmers are using their own machinery, tractors, and harvesting equipment.

Lee adds that the results reveal the importance of tailoring harvesting strategies to site-specific conditions and provide the first evidence-based guidance that harvest methods can improve both economic and environmental outcomes.

In a separate study published in Biomass & Bioenergy, the team tackled age-related declines in Miscanthus biomass yield, a function of tiller (stem) mass and density. These long-lived perennial grasses follow a predictable growth trajectory, including a juvenile stage that builds over several years to reach peak biomass yield, followed by a slow decline after about 10 years. Until now, it wasn’t clear which components of yield change over time and how nutrient management might help.

Postdoctoral fellow Nictor Namoi analyzed data from a long-term Miscanthus trial with nitrogen fertilization treatments that varied in amount and timing. First, he looked at how tiller mass and density changed over time and with various nitrogen treatments. Then, he asked what other soil fertility factors may influence the decline in biomass yield in older stands. 

“We found that both tiller mass and density increase from the first year of establishment to the fourth year, and if you apply nitrogen, you get an increase in both factors,” Namoi said. “But over time, as you withdraw nitrogen by harvesting biomass, the first component to be impacted is tiller mass. So tiller mass is very sensitive to nitrogen management.”

Namoi adds that tiller density continues to increase until individual stands become saturated, with no more space for further expansion. After that point, biomass yield is determined by tiller mass. 

He notes that the decline in biomass yield over time may not be entirely tied to nitrogen. With every harvest, soil nutrients tied up in plant biomass are removed from the system, depleting elements that play a role in photosynthetic efficiency. When Namoi analyzed soil phosphorus and potassium in mature Miscanthus stands, he found significant deficits in both essential nutrients.  

“Our findings identify tiller mass as a key determinant of biomass yield in aging Miscanthus and highlight the need for nitrogen, phosphorus, and potassium management for long-term productivity,” he said.

Both studies provide practical guidance to increase profitability, a key factor for any producer looking to explore biomass crops.

The first study, “Optimizing bioenergy biofuel harvest: a comparative analysis of stepwise and integrated methods for economic and environmental sustainability,” is published in Bioresource Technology [DOI: 10.1016/j.biortech.2025.133288]. 

The second study, “Soil fertility management for sustainable Miscanthus × giganteus production: Increased tiller weight from nitrogen management explains yield gains in aged miscanthus,” is published in Biomass & Bioenergy [DOI: 10.1016/j.biombioe.2025.108394].

Research in the College of ACES is made possible in part by Hatch funding from USDA’s National Institute of Food and Agriculture. This work was also funded by the U.S. Department of Energy, Bioenergy Technologies Office (DOE-BETO) under Award Number (DE-EE0008521) and the DOE Center for Advanced Bioenergy and Bioproducts Innovation (U.S. Department of Energy, Office of Science, Biological and Environmental Research Program under Award Number DE-SC0018420).