Tuesday, June 17, 2025

 

War, trade and agriculture spread rice disease across Africa




PLOS

War, trade and agriculture spread rice disease across Africa 

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Dispersion of rice yellow mottle virus throughout East Africa visualized by Evolaps.

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Credit: François Chevenet





Since the mid-1800s, human activities have rapidly facilitated the spread of rice yellow mottle virus (RYMV), a pathogen that infects rice, far and wide across Africa, according to a new study led by Eugénie Hébrard, at the Institut de Recherche pour le Développement (IRD, France), published June 17, 2025 in the open-access journal PLOS Pathogens.

RYMV is a pathogen that infects rice and a few related grass species, and that poses a major threat to rice production in Africa. In the new study, researchers investigated how human history has shaped the spread of RYMV, looking at how distinct strains of RYMV evolved in different locations and times.

The research team compared the gene sequences that code for a viral protein or the full-length viral genomes, fromup to 335 virus samples collected across more than 770,000 square miles in East Africa between 1966 and 2020. Based on variations and similarities in the gene sequences, the researchers found evidence that RYMV emerged in the middle of the 1800s in the Eastern Arc Mountains, a biodiversity hotspot, located in what is now Tanzania, where people grew rice slash-and-burn agriculture. Several spillovers of RYMV from wild grasses into cultivated rice were identified, with the virus rapidly spreading to the nearby rice growing areas, including Kilombero valley and the Morogoro region in southern Tanzania.

The study also suggested that humans transported RYMV long distances in infected rice plant matter at multiple points in history. The virus spread along the caravan routes from the Indian Ocean Coast to Lake Victoria in the second half of the 1800s, from East Africa to West Africa at the end of the 1800s, from Lake Victoria to the north of Ethiopia in the second half of the 1900s, and then on to Madagascar at the end of 1970s. Unexpectedly, it moved from the Kilombero Valley to the southern end of Lake Malawi toward the end of the First World War, likely due to rice being a staple food for troops.

Altogether, these findings suggest that transporting contaminated rice seeds was a major factor in spreading RYMV across long distances, not only within East Africa, but also in bringing it from East Africa to West Africa and Madagascar. The researchers conclude that, due to human activities, RYMV can spread as efficiently as some highly mobile zoonotic viruses that humans have contracted from animals. The study also sheds light on the risk of transmitting RYMV and other plant viruses from Africa to other continents.

The authors add, “This paper highlights the role of human history in the transmission of plant pathogens and underscores the risks of intercontinental transmission. The paradoxical role of seeds in the spread of a major pest of rice - which is not seed transmitted, but seed associated – is explained in the light of rice biology and agronomy. This study is a major achievement of a long-term, multilateral and interdisciplinary partnership.”

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Press Preview: https://plos.io/44cjsAT

In your coverage, please use this URL to provide access to the freely available paper in PLOS Pathogenshttp://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1013168

Citation: Ndikumana I, Onaga G, Pinel-Galzi A, Rocu P, Hubert J, Wéré HK, et al. (2025) Grains, trade and war in the multimodal transmission of Rice yellow mottle virus: An historical and phylogeographical retrospective. PLoS Pathog 21(6): e1013168. https://doi.org/10.1371/journal.ppat.1013168

 

Movie caption: Spatio-temporal dispersion of rice yellow mottle virus throughout East Africa visualized by Evolaps. https://www.evolaps.org/

Movie credit: François Chevenet, © Mapbox © OpenStreetMap, CC-BY 4.0 (https://creativecommons.org/licenses/by/4.0/)

High-resolution movie link: https://plos.io/3Z30MRn

Author Countries: Belgium, Cote d’Ivoire, France, Kenya, Rwanda, Singapore, Tanzania, United States

Funding: This work was partly supported by the French National Research Agency as an “Investissements d’avenir” program (ANR-10-LABX-001-01 Labex Agro) coordinated by Agropolis Foundation (project no. 1504-004 E-SPACE to IN, EPG, NP, DF, EH) and by a bilateral project between Kenya and France (PHC PAMOJA no 36128PK to HKW, AA, MNW, EH) cofunded by National Commission for Science, Technology and Innovation (NACOSTI) and Ministère de l’Europe et des Affaires Etrangères (MEAE). PR’s internship at the University of Montpellier was founded by the I-SITE MUSE through the Key Initiative “Data and Life Sciences”. SD acknowledges support from the Fonds National de la Recherche Scientifique (F.R.S.-FNRS, Belgium; grant n°F.4515.22), from the Research Foundation - Flanders (Fonds voor Wetenschappelijk Onderzoek - Vlaanderen, FWO, Belgium; grant n°G098321N), from the European Union Horizon 2020 projects MOOD (grant agreement n°874850) and LEAPS (grant agreement n°101094685). GO acknowledges support from the Plant Health Initiative (PHI) funded by the CGIAR Trust Fund. The funders had no role in the study design, data collection and interpretation, or the decision to submit the work for publication.

 

Hyperspectral sensor pushes weed science a wave further



Spectroradiometer used to quantify plant response to herbicide




University of Arkansas System Division of Agriculture

Aurelie Poncet and Mario Soto 

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Aurelie Poncet, left, and Mario Soto conducted a study showing how a sensor and AI/Machine Learning can be used to assist weed scientists in rating herbicide effectiveness on plants. Poncet is an assistant professor of precision agriculture in the crop, soil and environmental sciences department for the Division of Agriculture and the Dale Bumpers College of Agricultural, Food and Life Sciences. Mario Soto is a master's student in the department.

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Credit: U of A System Division of Agriculture photo





FAYETTEVILLE, Ark. — By combining artificial intelligence and sensors that can see beyond visible light, Arkansas researchers have developed a system that exceeds human discernment when it comes to measuring herbicide-induced stress in plants.

Scientists with the Arkansas Agricultural Experiment Station, the research arm of the University of Arkansas System Division of Agriculture, recently published a study in Smart Agricultural Technology providing proof-of-concept that hyperspectral sensors like a spectroradiometer can help in quantifying herbicide effectiveness, a critical element of weed management that helps curb herbicide resistance.

While normal cameras use three visible light bands — red, green and blue — to create images in the spectral range of 380 to 750 nanometers, hyperspectral sensing captures bands ranging from 250 nanometers to 2,500 nanometers and thermal infrared.

The researchers used this technology to evaluate how common lambsquarters responded to glyphosate. They also turned up empirical evidence that photosynthesis in the plant actually increased when exposed to a sub-lethal dose of the herbicide. Common lambsquarters — Chenopodium album L. — is a weed in agricultural and garden settings.

“Plant response to herbicide application is measured using visual ratings, but accuracy varies with the quality of training and years of practice of the rater,” said principal investigator of the study Aurelie Poncet, assistant professor of precision agriculture in the crop, soil and environmental sciences department for the Division of Agriculture and the Dale Bumpers College of Agricultural, Food and Life Sciences. “We thought, if we could have a sensor that automates some of this decision, we might be able to implement it into applications down the road.”

Weed scientists are trained to rate herbicide efficacy within a 10 percent margin of error, plus or minus 5 percent. The researchers were able to use machine learning models on data collected with a spectroradiometer to reach a margin of error of 12.1 percent. Their goal is to get below 10 percent.

The researchers used a random forest machine learning algorithm to analyze thousands of vegetation index data points collected in the experiment. The algorithm combines the output of multiple decision trees to reach a single result.

“Our success using random forest to describe common lambsquarters response to glyphosate application opens the possibility of moving beyond the development of vegetation indices, another approach gaining traction in the published literature,” said Mario Soto, lead author of the study and a crop, soil and environmental sciences master’s student in Bumpers College.

Next steps

Once refined, hyperspectral sensing could be used to measure specific weed response to herbicide application and overcome limitations of a human’s visual assessment. Further development of the method and validation may also be used to create a platform for high-throughput categorization of weed response to herbicides and screening for herbicide resistance, the study’s authors noted.

While training can overcome lack of experience for evaluators, mental and physical fatigue from long workdays evaluating treatments in harsh environmental conditions can affect judgement for even the most experienced evaluator, said Nilda Roma-Burgos, professor of weed physiology and molecular biology for the experiment station and Bumpers College.

“This method, in principle, could remove the human factor in herbicide efficacy evaluations and will be an invaluable research tool for weed science,” said Burgos, a co-author of the study. “Meanwhile, much work still awaits to validate the method across key weed species, herbicide modes of action, time after herbicide application and environmental conditions.”

Co-authors of the study included Kristofor Brye, University Professor of applied soil physics and pedology; Wesley France, program associate, and Juan C. Velasquez, weed science graduate research assistant, of the crop, soil and environmental sciences department.

Cengiz Koparan, assistant professor of precision agriculture technology with the agricultural education, communication and technology department and the biological and agricultural engineering department, and Amanda Ashworth, research soil scientist with the U.S. Department of Agriculture’s Agricultural Research Service, were also co-authors.

The hyperspectral imaging study was supported in part by the National Science Foundation’s NSF-SBIR Phase II Award No. 2304528 and the USDA’s National Institute of Food and Agriculture, Hatch projects ARK0–2734 and ARK0–2852.

To learn more about the Division of Agriculture research, visit the Arkansas Agricultural Experiment Station website. Follow us on X at @ArkAgResearch, subscribe to the Food, Farms and Forests podcast and sign up for our monthly newsletter, the Arkansas Agricultural Research Report. To learn more about the Division of Agriculture, visit uada.edu. Follow us on X at @AgInArk. To learn about extension programs in Arkansas, contact your local Cooperative Extension Service agent or visit uaex.uada.edu.

About the Division of Agriculture

The University of Arkansas System Division of Agriculture’s mission is to strengthen agriculture, communities, and families by connecting trusted research to the adoption of best practices. Through the Agricultural Experiment Station and the Cooperative Extension Service, the Division of Agriculture conducts research and extension work within the nation’s historic land grant education system. 

The Division of Agriculture is one of 20 entities within the University of Arkansas System. It has offices in all 75 counties in Arkansas and faculty on three system campuses.  

Pursuant to 7 CFR § 15.3, the University of Arkansas System Division of Agriculture offers all its Extension and Research programs and services (including employment) without regard to race, color, sex, national origin, religion, age, disability, marital or veteran status, genetic information, sexual preference, pregnancy or any other legally protected status, and is an equal opportunity institution.

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‘Virtual ward’ bed uses 4 times less carbon than traditional inpatient bed



These wards have the potential to reduce need to build more high-carbon impact hospitals



BMJ Group





A virtual ward bed uses 4 times less carbon than a traditional inpatient bed, so helping the NHS achieve its net zero target by 2045, finds the first study of its kind, published in the open access journal BMJ Innovations.

And they are a promising way to care for more patients effectively, with the potential to reduce the need to build more high-carbon impact hospitals, suggest the researchers.

A virtual ward, also known as "hospital at home," provides hospital-level care to patients in the comfort of their own homes, leveraging digital technology and remote monitoring. 

In recent years, particularly in the wake of the COVID-19 pandemic, virtual wards have been widely adopted throughout England to ease pressures on hospital bed capacity and streamline patient flow, say the researchers.

But while existing research has primarily focused on the care outcomes and cost effectiveness of virtual wards, few studies have looked at their environmental impact and carbon footprint.

To plug this knowledge gap, the researchers compared the carbon cost of virtual ward care with that of traditional inpatient care between May 2022 and May 2023 in a large acute hospital trust. 

They quantified the avoided carbon emissions for 1260 virtual ward patients, 728 of whom were frail and 532 of whom had an acute respiratory infection.

The researchers used the Greener Care at Home toolkit to calculate the carbon emissions of a care pathway, including carbon activity points, such as accident and emergency (A&E) attendance, travel to hospital in an ambulance/car, and diagnostics. 

All inpatient bed days, virtual ward bed days, home energy and community/general practice (GP) call-outs were calculated initially for a random sample of 30 patients, using a manual audit and then for the entire group of 1260 patients, using an internal data collection system. 

The researchers used a previously created method to calculate ‘predicted stay’ in hospital for the virtual ward and traditional inpatient care pathways.

 

The results show that there was a significant difference between the carbon costs of a virtual ward and an inpatient stay, with virtual wards emitting significantly less carbon when evaluated across the entire episode of care.

On average, an inpatient bed emits 4 times more carbon  at 37.9 kg COthan a virtual ward bed day at 8.8 kg CO2. And avoided carbon emissions added up to 285 tonnes of CO2 between May 2022 and May 2023.

This doesn’t represent a carbon reduction for the hospital, emphasise the researchers, as hospital beds were still in use by other patients, but it does represent increased capacity. 

And the reduced carbon footprint of virtual wards is particularly important as the NHS aims to deliver 40–50 virtual ward beds per 100,000 of the population, say the researchers. 

“Having a [virtual ward] in place will not decrease overall carbon emissions for the hospital but enable more patients to be cared for in the most efficient and lowest carbon way possible, enabling the hospital's capacity to increase and for teams to manage more patients with the same number of inpatient beds,” they write.

Carbon costs were higher, using the manual audit, largely due to the addition of external factors that internal hospital data systems don’t capture, note the researchers.

The researchers acknowledge that they didn’t have any data on carbon emissions from typical home use, and relied instead on government calculations, which estimate average home carbon emissions of 7.4 kg CO2/day. Nor do patients always fit into a neat box of a mapped pathway, which highlights the complexity of carbon mapping care pathways, they point out.

Nevertheless, they conclude: “[Virtual wards] look like a promising way for hospitals to increase capacity in a model of sustainable healthcare that aligns with the triple bottom line analysis of high-quality care, value for money, and low associated carbon emissions.”

They add: “We know that a traditional inpatient bed is a very high carbon/resource intensive method of treating patients. As our population increases, we will need to create more effective, less resource-intensive ways to treat our local population, without having to build more hospitals as this is extremely high cost, high carbon, and will require additional workforce.” 

 

Father-daughter bonding helps female baboons live longer



University of Notre Dame
Infant baboon with adult male baboon 

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An adult male and infant baboon in the Amboseli ecosystem, Kenya.

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Credit: (Photo by Elizabeth Archie, professor at Notre Dame)





Besides humans, very few mammals receive care from their fathers. But when species do, it may benefit their children.

New research from the University of Notre Dame found that the strength of early-life father-daughter relationships predicts meaningful differences in the survival of female baboons.

Published in Proceedings of the Royal Society B, the study evaluated the impact father baboons may have when they choose to co-reside or interact with their daughters, even though baboon mothers provide all essential care. Until now, the consequences of early-life paternal relationships of offspring were mostly unknown.

“Male baboons tend to reach their peak reproductive success when they’re young adults,” said Elizabeth Archie, professor of biological sciences at Notre Dame and corresponding author of the study. “But once they’ve had a few kids and their condition declines, they sort of slide into ‘dad mode,’ where they don’t disperse as much and they don’t try as hard to mate. Then they have time to invest in and hang out with their kids.”

Looking at 216 female baboons and their fathers in the Amboseli ecosystem of East Africa, the researchers found about a third of the daughters lived in the same social group as their fathers for three years or more. The remaining two-thirds had fathers who either left the group or died within their daughter’s first three years of life.

Researchers also evaluated the grooming habits of juvenile females with their fathers and other adult males, which speaks to the potential strength of father-daughter and other relationships. Archie shared that grooming, which is used for hygiene and social bonding, could be considered the “human equivalent of sitting down, having a cup of coffee and a good chat.”

The study showed daughters who had a strong relationship with their fathers, who co-resided with their fathers for three years or more, or both, lived two to four years longer than females who had weak father-daughter relationships.

“Early life adversity has a powerful effect on lifespan, so this study suggests that having a dad allows females that have experienced other forms of adversity to recover some of those costs,” Archie said. “In a lot of mammals, dads have a reputation of not contributing very much to offering care, but we now know that even these seemingly minor contributions that males are making still have really important consequences, at least in baboons.”

Additionally, father-daughter pairs that lived together for longer had stronger grooming relationships. Meanwhile, strong relationships between juvenile females and other adult males did not predict adult survival. This could be because male baboons sometimes intervene on behalf of offspring in conflicts, protecting their daughters, and even the mothers, from other group members.

“Males seem to sort of expand a child’s social network, as they can be popular members of their social group. Lots of baboons are coming up and interacting with the male. So an infant who’s hanging out near a male has more diverse social interactions than if they’re only hanging out with mom,” Archie said. “And dads can create a sort of safety zone for their daughters.”

Although mammal fathers may not provide much, if at all, to their offspring, Archie believes this study may hold insight into the evolutionary roots of human parental care.

This study is part of the Amboseli Baboon Research Project, which began in 1971 and is among the longest-running primate studies in the world. Funded by the National Science Foundation (NSF) and the National Institutes of Health (NIH), the project is co-directed by Archie at Notre Dame, Susan Alberts from Duke University and Jenny Tung at the Max Planck Institute for Evolutionary Anthropology.

“We’re very grateful to the NSF and NIH for funding over the years that has allowed us to sustain this project. We could not have done over 50 years of this project without their support,” Archie said.

In addition to Archie, Alberts and Tung, study co-authors include David Jansen at the University of Wisconsin-Madison and J. Kinyua Warutere at the Amboseli National Park in Kenya. Archie is affiliated with Notre Dame’s Eck Institute for Global Health and Environmental Change Initiative.

Contact: Brandi Wampler, associate director of media relations, 574-631-2632, brandiwampler@nd.edu

 

New species of armored, monstersaur lizard that lived alongside dinosaurs identified by NHM paleontologists



Discovery of Bolg amondol, a name inspired by J.R.R. Tolkien’s Lord of the Rings universe, reveals, reveals the complex evolutionary history of giant Gila monster relatives that roamed Utah’s prehistoric tropical forests.


Natural History Museum of Los Angeles County

Artistic reconstruction of Bolg amondol, by Cullen Townsend 

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Artistic reconstruction of Bolg amondol, depicted raiding an oviraptorosaur dinosaur nest amidst the lush Kaiparowits Formation habitat. 

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Credit: Art by Cullen Townsend.





A newly discovered raccoon-sized armored monstersaurian lizard from Grand Staircase-Escalante National Monument in southern Utah reveals a surprising diversity of these very big lizards at the pinnacle of the Age of Dinosaurs. Named for the goblin prince from J.R.R. Tolkien’s The Hobbit, the new species Bolg amondol also illuminates the sometimes murky path that life traveled between ancient continents. Published in the open-access journal Royal Society Open Science, the collaborative research led by the Natural History Museum of Los Angeles County’s Dinosaur Institute reveals hidden treasures awaiting future paleontologists in the bowels of museum fossil collections, and the vast potential of paleontological heritage preserved in Grand Staircase-Escalante National Monument and other public lands. 

“I opened this jar of bones labeled ‘lizard’ at the Natural History Museum of Utah, and was like, oh wow, there's a fragmentary skeleton here,” says lead author Dr. Hank Woolley from the Dinosaur Institute. “We know very little about large-bodied lizards from the Kaiparowits Formation in Grand Staircase-Escalante National Monument in Utah, so I knew this was significant right away.”

A Moniker from Middle-earth
Bolg represents an evolutionary lineage that sprouted within a group of large-bodied lizards called monstersaurs, the most familiar example being the Gila monsters, which still roam the deserts where Bolg was recovered from. Woolley knew that a new species of monstersaur called for an appropriate name from an iconic monster creator: J.R.R. Tolkien. “Bolg is a great sounding name. It's a goblin prince from The Hobbit, and I think of these lizards as goblin-like, especially looking at their skulls,” says Woolley.  He used the fictional language Sindarin—created by Tolkien for his elves—to craft the species epithet.  “Amon” means “mound”, and “dol” means “head” in the Elvish language, a reference to the mound-like osteoderms found on Bolg’s and other monstersaur skulls. “Mound-Headed Bolg” would fit right in with the goblins—and it’s revealing quite a bit about monstersaurs.”

Hidden Gems in Collection Drawers 
Bolg is a great example of the importance of natural history museum collections,” says co-author Dr. Randy Irmis from the University of Utah. “Although we knew the specimen was significant when it was discovered back in 2005, it took a specialist in lizard evolution like Hank to truly recognize its scientific importance, and take on the task of researching and scientifically describing this new species.”

The new species was identified from an associated skeleton of fragmentary bones: tiny pieces of the skull, vertebrae, girdles, limbs, and the bony armor called osteoderms. 

“What's really interesting about this holotype specimen of Bolg is that it's fragmentary, yes, but we have a broad sample of the skeleton preserved,” Woolley says. “There's no overlapping bones—there's not two left hip bones or anything like that. So we can be confident that these remains likely belonged to a single individual.”

Most of the fossil lizards from the Age of Dinosaurs are even more fragmented—often just single isolated bones or teeth—so despite their fragmentary nature, the parts of Bolg’s skeleton that survived contain a stunning amount of information.

“That means more characteristics are available for us to assess and compare to similar-looking lizards. Importantly, we can use those characteristics to understand this animal's evolutionary relationships and test hypotheses about where it fits on the lizard tree of life,” says Woolley.

Other fossils described in the study, including remarkably armored skull bones, demonstrate that the ancient, seasonally tropical forests of what is now southern Utah, USA hosted at least three different types of large, predatory lizards. “Even though these lizards were large, their skeletons are quite rare, with most of their fossil record based on single bones and teeth,” says co-author Dr. Joe Sertich from the Smithsonian Tropical Research Institute and Colorado State University. “The exceptional record of big lizards from Grand Staircase-Escalante National Monument may prove to be a normal part of dinosaur-dominated ecosystems from North America, filling key roles as smaller predators hunting down eggs and small animals in the forests of Laramidia."

Stairway to Monstersaurs
The Monstersauria are characterized by their large size and distinctive features like pitted, polygonal armor attached to their skulls and sharp, spire-like teeth. They have a roughly 100 million-year history, but their fossil record is largely incomplete, making the discovery of Bolg a big deal for understanding these charismatic lizards, and Bolg would have been a bit of a monster to our eyes.

“Three feet tip to tail, maybe even bigger than that, depending on the length of the tail and torso,” says Woolley. “So by modern lizard standards, a very large animal, similar in size to a Savannah monitor lizard; something that you wouldn’t want to mess around with.”

The identification of a new species of monstersaur highlights the likelihood that there were many more kinds of big lizards in the Late Cretaceous. Additionally, this find shows that unexplored diversity is waiting to be dug up in the field and in paleontology collections.

Bolg’s closest known relative hails from the other side of the planet in the Gobi Desert of Asia. While dinosaurs have long been known to have traveled between the once connected continents during the Late Cretaceous Period, the discovery of Bolg reveals that smaller animals also made the trek, suggesting there were common patterns of biogeography across terrestrial vertebrates during this time.

Dr. Woolley began this research as a PhD student at the Dinosaur Institute and has continued it as a National Science Foundation Postdoctoral Fellow in the department, underscoring the value of funding scientific research and the unique role the Dinosaur Institute plays as a source of mentorship for young paleontologists. 

“The Natural History Museum and Dinosaur Institute has been proud to lead the way in empowering early career scientists,” says Dr. Nathan Smith, co-author and Gretchen Augustyn Director & Curator of the Dinosaur Institute. In addition to Dr. Woolley, co-author Dr. Keegan Melstrom (now an Assistant Professor at the University of Central Oklahoma), was also a Postdoctoral Fellow in the Dinosaur Institute. “The result of that investment and continued growth of programs like the Dinosaur Institute Fellowship Fund is groundbreaking paleontological research and new discoveries that highlight the value of museum collections and expand our knowledge of Earth history.”

Field collection of the specimens described in this study was conducted under paleontological permits issued by the Bureau of Land Management. This study was funded by the Bureau of Land Management, National Science Foundation award 2205564, the NHMLAC Dinosaur Institute, and the University of Utah. 

The rocks where Bolg was discovered, the Kaiparowits Formation of Grand Staircase-Escalante National Monument, have emerged as a paleontological hotspot over the past 25 years, producing one of the most astounding dinosaur-dominated records anywhere in North America with dozens of new species and critical insights into the past. Discoveries like this underscore the importance of protecting public lands in the western USA for science and research.
 

LEFT IMAGE: Publicly available rendered CT scan of Heloderma horridum (UF:Herp:153328) from morphosource.org used for comparison to Bolg amondol, with mound-like, polygonal osteoderms in green. RIGHT IMAGE: Skull of the modern monstersaur Heloderma horridum (LACM 159136), from the herpetology collection at NHMLAC, used for comparison with Bolg amondol.

Identified bones belonging to Bolg amondol (Specimen UMNH VP 16266, Natural History Museum of Utah / Bureau of Land Management). They are not much to look at, but they are jam-packed with valuable information on the anatomy and lifestyle of Bolg amondol.

Size comparison of the holotype specimen of Bolg amondol (UMNH VP 16266, left, Natural History Museum of Utah / Bureau of Land Management) and a modern Gila monster (Heloderma suspectum, right). Gold: preserved skeletal elements. Light gray: reconstructed elements based on preserved material. Dark gray: missing skeletal elements, based on publicly available rendered CT scans of Heloderma horridum available on morphosource.org (UF:Herp 153328). 

 

Credit

Dr. Hank Woolley