Mention fungi, and most people will probably think of the mushrooms they pick in fall, or maybe the yeast they add when baking or making wine. Others will perhaps recall last week’s mouldy bread – or cucumbers gone bad in the refrigerator. Indeed, mycologists have studied these fungi as sources of food and fermentation but also decay and disease for centuries.
Sampling soil and sediments for fungal diversity not far from Göteborg, Sweden. Photo by Henrik Nilsson
But while we’re used to thinking of fungi as organisms that form physical structures such as fruiting bodies, or yeast-like life forms that we can grow in our kitchens or laboratories, it is gradually becoming clear that fungi don’t readily assort into only these two groups. DNA sequencing studies of environmental substrates such as soil are finding massive evidence of large groups of fungi that do not seem to form fruiting bodies and that we seem unable to grow in the lab – but that are there nonetheless. These groups are often called “dark fungi,” in analogy with the concept of “dark matter” in astronomy – something we know is out there, but that we cannot directly observe right now.
A new study in MycoKeys contrasts the accumulation of fungal species recovered using traditional mycological approaches with those recovered using environmental DNA sequencing over time. Even when allowing for various kinds of biases, the authors found that species discovery through environmental sequencing vastly outpaces traditional species recovery in a strongly increasing trend over the last five years. The authors conclude that dark fungi form a defining feature of the fungal kingdom.
Field work on the Tibetan Plateau. Photo by Wengang Kang
“And that’s where it gets interesting”, Henrik Nilsson at the University of Gothenburg, Sweden, and the lead author of the study, says. “Under the current rules of nomenclature, these fungi cannot be given scientific names – they cannot be described formally. And species and groups that cannot be named formally, well, they tend to fall between the cracks. They’re typically not considered in nature conservation initiatives. They are often left out from efforts to estimate the evolutionary history of fungi, and their ecological roles and associations are largely overlooked when we try to figure out how mass and energy flow in ecosystems. They’re essentially treated as if they didn’t exist.”
Examining minute fungal fruiting bodies not far from Stockholm, Sweden. Photo by Kristina Stenmarck
Second author Martin Ryberg at the University of Uppsala, Sweden chimes in, “And it’s not like we’re adding the few missing pieces to an otherwise nearly complete jigsaw puzzle. It seems to be the other way around. We’re talking about tens of large groups of fungi – and thousands upon thousands of species, some of which seem to be so common that we have yet to find a soil sample from which they’re absent. Indeed, we’re talking about what could well prove to be the dominant life style in the fungal kingdom.”
The mycological community has been debating whether the rules of fungal nomenclature should be modified to allow formal description of these dark fungi. So far, the matter has not been resolved in the affirmative. “I think our study shows that it’s time to stop that debate, like, right away,” Nilsson says. “What we should be debating is how we should describe them. What criteria must be fulfilled for a dark fungus to be given a formal scientific name? Clearly, formation of a fruiting body or growth in the laboratory can’t be part of those criteria.”
Field work in New Caledonia. Photo by Sten Anslan
Co-author Alice Retter of the University of Vienna, Austria explains, “We figured we’d kickstart the how debate by listing criteria that we think make sense – criteria that would be stringent enough to allow for only particularly well-vetted dark fungi to be described, upholding a high level of scientific rigor and reproducibility in the process. We blended our own observations with suggestions from the mycological community, culled from depositing a preprint of the manuscript at bioRxiv. We’re certainly not claiming that our suggestions form the final word in the debate. It’s more like they’re the first. We’re thinking that the mycological community will jointly be able to come up with a set of sound guiding principles on the matter – and here comes an initial set of well-meaning observations for nucleation.”
Field work in the German Alps. Photo by Vanessa Schulz
The authors advocate gentle modifications to the nomenclatural rules governing the naming of fungi to allow giving formal names to at least the most well-documented species and groups of dark fungi. The suggested modifications would, at present, exclude many rare or otherwise less well-documented dark fungi from formal description.
“But you don’t have to have a theory of everything to have a theory of something,” senior author Kessy Abarenkov of the Tartu Natural History Museum, Estonia asserts. “By establishing rules for what’s needed to describe dark fungi, and specifying when we’ll have to refrain from describing such species at present, mycologists can do what they do best: doggedly gather enough research data to warrant naming of the dark fungi, group by group, and species by species. It’s what mycology has excelled at for hundreds of years. It’s just the setting that’s a bit new.”
Drying soil samples immediately upon collection under field conditions in Norway. Photo by Sten Anslan
Sten Anslan, University of Tartu, continues: “Much is at stake, obviously. The current rules governing the naming of fungi have served mycology well for a long time. We don’t want to upend or overthrow them. But we fear that if they’re not updated in this particular regard, there’s a risk that they grow increasingly obsolete over time. Having a book of rules that govern maybe only some few percent of the organisms it was originally conceived to govern – the fungal kingdom – would seem untenable in the long run.”
Getting ready for DNA extraction from soil samples. Photo by Sten Anslan
Marisol Sanchez-Garcia of the Swedish Agricultural University concludes: “The nomenclatural aspects of dark fungi will presumably be discussed at some length at next year’s international mycological congress in Maastricht, the Netherlands. We’re hopeful that the mycological community will reach meaningful agreement on integration of the dark fungi into the rules of nomenclature. After all, mycologists are used to negotiating and solving non-trivial questions on a day-to-day basis, and this one is hardly any different. Being part of tackling a huge, more or less unknown group of organisms where precious little is set in stone and where the rules will have to be adapted over time for the endeavour to stay attuned to recent developments, well, that’s what makes being a mycologist so interesting and rewarding in my eyes.”
Research article:
Nilsson RH, Ryberg M, Wurzbacher C, Tedersoo L, Anslan S, Põlme S, Spirin V, Mikryukov V, Svantesson S, Hartmann M, Lennartsdotter C, Belford P, Khomich M, Retter A, Corcoll N, Gómez Martinez D, Jansson T, Ghobad-Nejhad M, Vu D, Sanchez-Garcia M, Kristiansson E, Abarenkov K (2023) How, not if, is the question mycologists should be asking about DNA-based typification. MycoKeys 96: 143-157. https://doi.org/10.3897/mycokeys.96.102669
Working with the scientific travel agency Taxon Expeditions, citizen scientists studed specimens, and analyse their DNA in a field camp in the heart of the forest.
Semislugs, or ‘snugs’ as they are affectionately known among mollusc researchers, are like the squatters of the snail world: they do carry a home on their back but it is too small to live in. Still, it offers a sort of protection, while not getting in the way of the worm-like physique of the slug. For reasons unknown, on the island of Borneo, which is shared among the countries of Brunei, Malaysia, and Indonesia, most slugs are of the semislug type. The genus Microparmarion there consists of around 10 semislug species, most of which are found in the cooler forests of the mountains. So, when citizen scientists discovered a Microparmarion in the hot lowland forest of Ulu Temburong National Park, Brunei, as part of their expedition, they were surprised.
A team of citizen scientists in the Brunei forest, searching for slugs and snails.The type locality of the new species.
For the past years, the scientific travel agency Taxon Expeditions, in collaboration with Universiti Brunei Darussalam (UBD) has been organising biodiversity discovery trips for scientists, students, and laypersons to this forest. On the first trip, in 2018, during a night walk, participant Simon Berenyi, who runs an ethical pest control company in the UK, reached up to a dead leaf suspended over the trail. Everybody—the other participants, even the resident snail expert—had ducked and passed underneath this dead leaf without so much as giving it a glance. But something on its surface caught Simon’s eye. “Oi, is that a slug?” he exclaimed, and picked a slimy, well-camouflaged mollusc off it.
Microparmarion sallehi.
At the time, the team’s zoologists already suspected it was a new species – nothing like it had ever been found in this corner of the island. But that single specimen was not enough to publish its description as a new species. Over the years, successive expeditions to the same area came up with several more specimens of the same species, which made it clear that it was really a species never seen before.
On the 2022 expedition, a team composed of UBD students Nurilya Ezzwan and Izzah Hamdani and citizen scientist Harrison Wu from Virginia, USA, finished the description. Using the portable lab that Taxon Expeditions always carries with them, the team studied the animals’ shell, reproductive organs, and DNA, and prepared a paper for the open-access Biodiversity Data Journal, where it was published this week.
As usual on Taxon Expedition trips, on the last night the team voted on the scientific name for the new species. With an overwhelming majority, the ‘snug’ was named after Mr. Md Salleh Abdullah Bat, the field centre supervisor, who would retire just weeks after the team left. Mr. Salleh himself agrees that it is indeed a very fitting farewell gift.
Research article:
Schilthuizen M, Berenyi S, Ezzwan NSMN, Hamdani NIAA, Wu H, De Antoni L, Vincenzi L, de Gier W, van Peursen ADP, Njunjić I, Delledonne M, Slik F, Grafe U, Cicuzza D (2023) A new semi-slug of the genus Microparmarion from Brunei, discovered, described and DNA-barcoded on citizen-science ‘taxon expeditions’ (Gastropoda, Stylommatophora, Ariophantidae). Biodiversity Data Journal 11: e101579. https://doi.org/10.3897/BDJ.11.e101579
With expertise in science communication, dissemination and exploitation, Pensoft is involved in this project set to develop new technologies for monitoring species and their habitats across Europe
With expertise in science communication, dissemination and exploitation, Pensoft became part of this project dedicated to new technologies for species and habitat monitoring across Europe
Background
The European Union puts a great value in monitoring the health of ecosystems, as comprehensive mapping can aid policy makers’ work in adopting appropriate legislation for nature conservation. It allows for understanding the impact of human activities and making informed decisions for effective management of nature’s resources. This is particularly important for the EU, as it has set ambitious goals to halt biodiversity loss and restore degraded ecosystems by 2030, as outlined in the EU Biodiversity Strategy for 2030.
Effective biodiversity monitoring can help the EU track progress towards these goals, assess the effectiveness of conservation policies and initiatives, and identify emerging threats to biodiversity.
Despite this awareness, efforts to monitor animals and plants remain spatially and temporally fragmented. This lack of integration regarding data and methods creates a gap in biodiversity monitoring, which can negatively impact policy-making. Today, modern technologies such as drones, artificial intelligence algorithms, or remote sensing are still not widely used in biodiversity monitoring.
MAMBO project (Modern Approaches to the Monitoring of BiOdiversity) recognises this need and aims to develop, test, and implement enabling tools for monitoring conservation status and ecological requirements of species and habitats. Having started in late 2022, the project is set to run for four years until September 2026.
Pensoft – with its proven expertise in communicating scientific results – is committed to amplifying the impact of MAMBO. Pensoft supports the project through tailored approaches to communication, dissemination and exploitation so as to reach the most appropriate target audience and achieve maximum visibility of the project.
Deep-dive into the project
In order to enrich the biodiversity monitoring landscape, MAMBO will implement a multi-disciplinary approach by utilising the technical expertise in the fields of computer science, remote sensing, and social science expertise on human-technology interactions, environmental economy, and citizen science. This will be combined with knowledge on species, ecology, and conservation biology.
More specifically, the project will develop, evaluate and integrate image and sound recognition-based AI solutions for EU biodiversity monitoring from species to habitats as well as promote the standardised calculation and automated retrieval of habitat data using deep learning and remote sensing.
“Classification algorithms have matured to an extent where it is possible to identify organisms automatically from digital data, such as images or sound,”
“Technical breakthroughs in the realm of high spatial resolution remote sensing set the future of ecological monitoring and can greatly enrich traditional approaches to biodiversity monitoring.”
In order to achieve its goals, the project will test existing tools in combination with MAMBO-developed new technologies at the project’s demonstration sites geographically spread across Europe. This will contribute to an integrated European biodiversity monitoring system with potential for dynamic adaptations.
Pensoft is part of MAMBO’s Work Package 7 (WP7): “Science-policy interface and dissemination”, led by Helmholtz Centre for Environmental Research (UFZ). The work package is dedicated to providing a distinct identity of the project and its services through branding, visualisation and elaborated dissemination and communication strategy.
Within WP7, Pensoft will also be taking care after the launch of an open-science collection of research outputs in the scholarly journal Research Ideas and Outcomes(RIO).
Amongst the tasks of the partners in WP7 is also the development of different pathways for integrating new technologies and innovations into the EU Pollinators Monitoring Scheme (EU PoMS; SPRING).
You can find more about the project on the MAMBO website: mambo-project.eu. Stay up to date with the project’s progress on Twitter (@MAMBO_EU) and Linkedin (/MAMBO Project).
Tetrigidae, commonly known as pygmy grasshoppers, are an ancient and diverse family, currently numbering about 2000 species. As their name suggests, tetrigids are very small; their largest representatives are barely several centimeters long, so they might be difficult to spot on a casual stroll through tropical vegetation. However, when they are spotted, they are immediately recognizable by their elongated pronotum, a hard structure that starts behind the head and covers the entire body like a hood. They come in many shapes and colors and are often exciting to see, but this comes with a price—the taxonomy of Tetrigidae, the way they are organized into natural groups, is a mess. This is where we come in.
In our latest paper, we dealt with Choriphyllini, a small Caribbean tribe that belongs to the subfamily Cladonotinae. This subfamily had been filling up with unrelated but similar-looking tetrigids for more than a century. It had never been clearly defined so almost everything wingless and robust was assigned to Cladonotinae. We decided to put an end to this by slowly removing the superficially similar genera from the subfamily and describing tribes to group the genera that are clearly related to each other. We piloted this system just last year, when we described the tribe Valalyllini from Madagascar, with only two endemic (and endangered) genera and species.
The diversity and the distribution of the tribe Valalyllini, the Malagasy dead-leaf-like Cladonotinae. Both species are endemic to small areas and are likely endangered because of deforestation. Both species most probably inhabit rainforest leaf litter.
Put the species of Choriphyllini and Valalyllini together, mix them up, and try to guess which belongs where—this is no simple task; they are all doing their impressions of dead leaves that our primate brains struggle to differentiate. And there’s more: such leaf-like grasshoppers live in Africa and South East Asia as well, and then there are those that look like twigs and spiky tree bark.
Only now that we have an idea of what the true Cladonotinae are can we be properly amazed by the duality they represent to us. On the one hand, they are incredibly diverse with every species having its own variation on the basic shape. On the other, they are so alike that they either represent the best example of convergent evolution ever documented or they all stem from a common ancestor that is currently supposed to have lived during the Mesozoic. The evolutionary history of Cladonotinae will take many years to unravel, but the work can only begin after we define what to call by that name.
Valalyllum folium, a member of the tribe Valalyllini, subfamily Cladonotinae. This species, endemic to Madagascar, is a relative of Choriphyllini.
It only took 250 years
The first species of Choriphyllini, Phyllotettix rhombeus, was described in 1765 as Cicada rhombea, that is, as a member of an entirely different order of insects. Continuing in this manner, many authors (including the great Linnaeus himself) made many taxonomic and nomenclatural mistakes that compounded over the centuries and made these grasshoppers difficult to identify and refer to. It didn’t help that new species and new records kept being reported without being contextualized by comprehensive literature reviews. Like detectives, we followed the scattered crumbs of data and arrived at a synthesis that will make future research in the region much more pleasant.
Hancock’s plate I from the “Tettigidae of North America” shows leaf-like Caribbean species under the numbers 1), 2) and 7), but has many taxonomic and nomenclatural errors. 1) – Phyllotettix foliatus (= female holotype of Hancock’s Choriphyllum foliatum), 2) – Phyllotettix rhombeus (= Hancock’s Choriphyllum westwoodi), 7) – Choriphyllum saussurei. (= Hancock’s Phyllonotus saussurei). Source: Biodiversity Heritage Library, available at https://www.biodiversitylibrary.org/item/25899#page/10.
This is not where interesting facts about Phyllotettix rhombeus stop. While looking through the literature, we tried to extract the measurements of drawings. Most of the drawings had a scale bar printed next to them, but the archaic usage of “lines” as the standard measurement initially gave us some trouble. That is why at first we doubted one of our most fascinating discoveries: with the pronotal length measuring nearly 3 centimeters, Phyllotettix rhombeus is the largest tetrigid ever recorded! Many, many authors dealt with this species over the last 250 years, but this record was never made explicit.
It should not go unnoticed now that its proposed common name is “Jamaican Colossal Jumping Leaf”. Inspired by this, we took the measurements of the other species as well and made a figure where all the specimens are resized to a common scale, which shows the diversity of both shapes and sizes.
The genera and species of the tribe Choriphyllini. All specimens are drawn to scale.
Besides P. rhombeus, there are three more species in the genus Phyllotettix: P. plagiatus,P. foliatus, and P. compressus. All four of them are known only from Jamaica. P. foliatus and P. compressus are known from the Blue Mountains, but for the other two no precise localities are known; we still don’t know where exactly the largest tetrigid lives. The other genus of the tribe is Choriphyllum, also with four species. Three of them, C. sagrai, C. saussurei, and C. wallaceum live in Cuba, while C. bahamense is all alone on Hummingbird Cay island in the Bahamas. The easiest way to differentiate these two genera is a little strange but practical, the tallest point of the leaf-like crest in Choriphyllum species is in the front, while in Phyllotettix species it is in the back.
A map of all known Choriphyllini records. For three species, not a single precise locality is known.
Some Caribbean leaves dance and jump
For each species, we proposed a common name as a means to give these animals even more character. Names, such as “Jamaican Bitten Jumping Leaf” and “Old Cuban Dancing Leaf” may not be “official”, but they have certainly found their audience. The tweet in which we shared the collage of all the species was viewed over 17000 times; everyone was amazed by the pretty shapes and some even noted that they especially liked the crazy common names. We were very glad to see our scientific and artistic package that is Choriphyllini be so warmly received.
Another hit on Twitter, with over 20000 views, is the post showcasing the newly-described species from Cuba, Choriphyllum wallaceum. The holotype of this species has been awaiting description for a long time. We found it in Museo Nacional de Ciencias Naturales in Madrid, Spain, with a note from Ignacio Bolívar, the father of the Tetrigidae classification system. He referred to it as “Choriphyllum Seoanei” but never managed to publish it.
This “new” species presented us with the perfect opportunity to honor the 200th anniversary of Alfred Russel Wallace’s birth. Wallace is often called the “father of biogeography” but is all too often neglected when discussing the origins of the theory of evolution, with which Charles Darwin is considered synonymous. Wallace, with his independent arrival at the key concepts of the evolutionary theory, his correspondence with Darwin, and his staunch defense of Darwin’s ideas, was (and is) at the very least equal to Darwin and deserves much more recognition than he currently gets.
Choriphyllum wallaceum, a newly-described species from Cuba, named after Alfred Russel Wallace.
This is just the start
Choriphyllini are a pretty package, but one that merely introduces the real problem. The history of this tribe is long, yet we have very few specimens to work with. Although we have an understanding of how morphology varies within species, P. compressus and P. foliatus are not only suspiciously similar to each other, but they also live in the same general area of the Blue Mountains. It remains to be seen if they are in fact a single species.
Much more pressing is that we have only a vague idea of where these animals live and how their populations are impacted by various factors such as human activity and climate change—we do not have a baseline against which to assess their conservation status. Then there is the fact that there are many more islands in the Caribbean, making the possibility of discovering new Choriphyllini species on them real and exciting. We can only guess what the future holds for these neglected animals.
Old Cuban Dancing Leaf (Choriphyllum sagrai) in its natural environment among the leaf litter in Cuba, photographed by Sheyla Yong.
The stage is set; everything we know about this group is laid out in the paper and now there is no path but forward. Research is expensive, dedication to this work takes a certain kind of soul, and everything takes time. It is our sincere hope that someone someday takes this further. The pygmy jumping leaves will wait for as long as they can, on their islands, hopping without a care in the world.
References:
Deranja M, Kasalo N, Adžić K, Franjević D, Skejo J (2022) Lepocranus and Valalyllum gen. nov. (Orthoptera, Tetrigidae, Cladonotinae), endangered Malagasy dead-leaf-like grasshoppers. ZooKeys 1109: 1-15. https://doi.org/10.3897/zookeys.1109.85565
UC Berkeley entomologist Kipling Will discovered a specimen of Bembidion brownorum while sampling for insects near Freshwater Creek on former Gov. Jerry Brown’s ranch.
When University of California, Berkeley, entomologist Kipling Will first heard that former Gov. Jerry Brown was hosting field scientists on his Colusa County ranch, he jumped at the chance to hunt for beetles on the property.
“I reached out and said, ‘Hey, I want to sample your beetles,’” Will said. “And [Brown] was quite game to let me come up there.”
Will, a professor of environmental science, policy and management, has travelled to all corners of California to study carabid beetles, ground beetles that are important predators of other insects. But Will’s repeated visits to Brown’s ranch proved especially fruitful.
While sampling for insects near Freshwater Creek, Will collected a rare species of beetle that had never been named or described — and which, according to records, had not been observed by scientists in over 55 years. The new species will be named Bembidion brownorum, in honor of Brown and his wife, Anne Brown.
Former California Gov. Jerry Brown (right) and his wife, Anne Brown, with their dogs, Colusa and Cali, on the Browns’ Colusa County ranch. Photo courtesy Evan Westrup
“I’m very glad that [my ranch] is advancing science in some interesting and important ways,” said Brown, who has hosted a wild variety of field researchers, including geologists, anthropologists and botanists, on the property. “There are so many undiscovered species. I think it’s very important that we catalog and discover what we have and understand their impact on the environment — how it’s functioning and how it’s changing.”
Brown’s 2,500-acre ranch is about an hour’s drive north of Sacramento, in an agricultural region where most of the land is privately owned and insect biodiversity is historically understudied. For more than two years, Will has regularly sampled for insects on the ranch, sometimes even showing the beetles that he finds to the Browns.
Jerry Brown said his dedication to welcoming researchers onto his land is rooted in the ranch’s history as a stagecoach stop called Mountain House, and in his own interest in climate change and conservation.
“We don’t have stagecoach stop, but we have a place of gathering, of research and collaboration,” said Brown, who is currently chair of the California-China Climate Institute at UC Berkeley.
The location on Freshwater Creek where Kipling Will discovered the Bembidion brownorum beetle. Photo courtesy Kipling Will
After collecting a beetle at the ranch that didn’t resemble any species he was familiar with, Will called up Bembidion expert David Maddison, a professor of integrative biology at Oregon State University, to help identify the specimen. Together, the scientists used morphological and DNA analysis to confirm that the beetle represented a completely new species.
Will then combed through entomology collections at museums throughout California in search of other specimens that may have been unlabeled or misidentified. He found only 21 other specimens of the species, the most recent of which was collected in 1966.
UC Berkeley entomologist Kipling Will discovered a specimen of Bembidion brownorum while sampling for insects near Freshwater Creek on former Gov. Jerry Brown’s ranch. The species had not been observed by scientists in more than 55 years. Photo courtesy David Maddison
The lack of any more recent specimens indicated to him that the species likely collapsed during the second half of the 20th century, driven out of its natural habitat by rapid urbanization and agricultural development across the state.
“The sad truth is, [the species] has probably been in a huge decline. If you look at the places that it was found the ‘20s and ‘30s and ‘40s, almost none of that natural habitat is left,” Will said. “But we don’t know for sure. So, the thing to do is to get it out there, describe it and tell people, ‘Hey, look for this thing,’ because maybe we’ll find some place where it’s doing fine.
“Having access to Jerry’s ranch in Colusa County gives me the opportunity to really spend time sampling, to look for rare things like this.”
Will and Maddison describe Bembidion brownorum in a study published in the journal Zookeys.
Big for a Bembidion
To the naked eye, Bembidion brownorum isn’t particularly remarkable: The diminutive beetle is brown in color and measures around 5 millimeters in length, about the diameter of a standard pencil. But under magnification, it glows with a green and gold metallic shimmer.
The Bembidion brownorum beetle is approximately 5 millimeters long, slightly larger than other Bembidion beetles. The section of the insect behind the head, called the prothorax, is also larger than usual. Photo courtesy David Maddison
It was the unusual shape of the beetle’s prothorax, the segment of the insect that sits right behind its head, that first caught Will’s eye.
“I was looking at this one beetle thinking, ‘It just doesn’t fit any of the ones that I can identify,’” Will said. “The shape of prothorax is just not like any of the others.”
According to Maddison, Bembidion brownorum is also relatively large compared to other Bembidion beetles, which are usually closer to 3 to 4 millimeters in length.
“It’s big for a Bembidion,” Maddison said. “At first glance, it was pretty obvious that it was probably something new.”
With so few examples to study, it’s difficult to describe the lifestyle and behavior of Bembidion brownorum with any certainty, Will said. However, given where the beetle was found on Brown’s ranch — in the vicinity of Freshwater Creek, which occasionally dries into a series of trellis-like pools in the summer months — it is likely that the beetle lives near the edges of bodies of water that periodically flood and then evaporate.
The 21 historical specimens of Bembidion brownorum are housed at either the Essig Museum Entomology Collection at UC Berkeley or at the California Academy of Sciences in San Francisco, which both have insect specimens going back more than 100 years. The discovery highlights the vital importance of maintaining these collections for current and future research, the scientists said.
“One of the things that I find interesting about is that, before Kip found that specimen, there were already specimens in collections — there was this hidden diversity that people didn’t recognize,” Maddison said. “At one point, [the beetle] probably was much more widespread and much more common, and Kip and I have some ideas as to where you would target to try to find more.”
Drawers containing prepared specimens of carabid beetles that Kipling Will collected on Jerry Brown’s ranch. Photo courtesy Kipling Will
Previous specimens were collected at locations throughout the Central Valley and in the Los Angeles Basin, regions that have been transformed over the last century. While the beetle may still survive in some areas, Will said that the patchwork of private landownership may make it difficult to find.
“There is a lot of desire to conserve the environment and combat climate change, but in many cases, we’re not keeping up with the rate of extinction — we’re not able to describe the species that need to be described as fast as things are going extinct,” Will said. “And this certainly is true in California, where there are an awful lot of undescribed insects out there and not a lot being done to get them described. I think that having more knowledge of what they are and where they where they live is really fundamental.”
John S. Sproul of the University of Nebraska Omaha is also a co-author of the study. This research was supported by the Harold E. and Leona M. Rice Endowment Fund at Oregon State University.
Research article:
Maddison DR, Sproul JS, Will K (2023) Re-collected after 55 years: a new species of Bembidion (Coleoptera, Carabidae) from California. ZooKeys 1156: 87-106. https://doi.org/10.3897/zookeys.1156.101072
Press release originally published by Kara Manke, UC Berkeley. Republished with permission.
Pangolins, unique scale-covered mammals, are drastically declining in numbers across Asia and Africa, largely due to illegal trade. Part of the trade, both legal and illegal, supports the traditional Chinese medicine market, which has attracted conservation attention. The level of demand for pangolins and other animals in traditional Chinese medicine, however, hasn’t been thoroughly studied.
In a new study published in the journal Nature Conservation, Dr Yifu Wang, currently a postdoc researcher at the University of Hong Kong, investigated pangolin scale trade in China, interviewing staff in hospitals and pharmaceutical shops in two provinces (Henan and Hainan). Between October 2016 and April 2017, she and her team talked to doctors from 41 hospitals and shop owners and assistants from 134 pharmaceutical shops.
An adult Philippine Pangolin and her pup photographed in the forests of Palawan. Photo by Gregg Yan under a CC-BY-SA-4.0 license
The research found pangolin scales and their derivatives were widely available in hospitals and pharmaceutical shops across Henan and Hainan Provinces. The legislation in place, however, has not been able to prevent ongoing illegal trade in pangolin products. Her team found that 46% of surveyed hospitals and 34% of surveyed pharmaceutical shops were selling pangolin scale products illegally.
“Existing legal trade allows 711 hospitals to sell pangolin products as medicine with regulations on manufacturer, package, and national annual sale quantity,” explains Dr Yifu Wang. “However, we show that pangolin scales are under heavy demand and unpermitted sellers are commonly found illegally selling pangolin products.”
Processed pangolin scales. Photo by Dr. Yifu Wang
“Quantities of products traded by permitted legal sellers are estimated to greatly exceed the supply capacity of legal sources,” she continues.
This widespread illegal trade, coupled with the very limited legal supply capacity compared to market demand, is concerning. The researchers point to the urgent need to reduce demand from traditional Chinese medicine on pangolin scales and revise the current legal pangolin scale trade system.
“We also highlight the importance of incorporating the traditional Chinese medicine sector into combating illegal wildlife trade and species conservation beyond pangolins,” they conclude.
The researchers plan to continue investigating the pangolin scale market in China to understand the trade after COVID-19.
Research article:
Wang Y, Turvey ST, Leader-Williams N (2023) The scale of the problem: understanding the demand for medicinal pangolin products in China. Nature Conservation 52: 47-61. https://doi.org/10.3897/natureconservation.52.95916
The shore is a mutual caress. More than just a place of encounter between land and water, it is one of the physical and imagined thresholds between humans and the other-than-human world. This place of touch - through thoughts, actions, interconnections, and affect - is the inevitable crossing at the beginning and end of every inquiry into the world’s bodies of water.
🥈Second prize for Best Talk Award at #esp22europe goes to @KrBurkhard and her couple of gorgeous co-authors🥰 about her research on co-benefits and risks of carbon sequestration in coastal ecosystems! 🎉 pic.twitter.com/hsQufK9Ncm
I find the idea of improving the understanding of how ecosystems – and more generally the natural world – contribute to our well-being to be very inspiring and refreshing in a high-tech profit-oriented world,
she says to explain how she stays motivated in her research work.
To communicate this knowledge to policy- and decision-makers, as well as the general public is key, especially when we consider the threats of climate change and the fact that our deep dependency on nature seems to be largely undervalued,
she adds.
Kremena highlights the role of nature-based solutions:
When utilising conventional and nature-based solutions, the focus is often on a single benefit that is demanded in a certain area, time and situation.
In contrast to conventional solutions, nature-based solutions provide additional co-benefits. These may include biodiversity protection and other ecosystem services that address broader societal demands and are more sustainable in the long term.
As part of the CDRmare research mission “Marine carbon sinks in decarbonisation pathways” of the German Marine Research Alliance, Kremena’s work on coastal ecosystems aims to develop approaches and methodologies which can be applied in an international context.
Our project sea4soCiety focuses on the carbon storage capacity and co-benefits of four coastal vegetated ecosystems which play a key role as carbon sinks around the world and thus contribute to climate regulation. The analysis and methods developed in the project contribute scientifically to the studied topics and have an international relevance.
The German coast is representative of three coastal ecosystems, namely seagrass, salt marsh and macroalgae. The fourth ecosystem – that of mangrove forests in the tropics, is also investigated within the project as a key ocean carbon sink of global relevance.
But climate regulation is only one of the multiple services that these ecosystems provide. Coastal protection, water purification, food and material provision and recreation are among the key services of coastal ecosystems, the benefits of which are used and highly appreciated by the local communities and have significant role in the local safety, economy and culture.
What are the strategies for mitigating or further analysing the risks of carbon sequestration in coastal ecosystems?
We prioritise conservation and restoration of coastal vegetated ecosystems, which are often heavily degraded, and we identify the most suitable areas for establishment of new ecosystems. This reduces the risk of carbon release and provides additional carbon sink capacity.
Further risks are related to unknown climate change impacts. The sea temperature and hydrodynamics are changing, and we are not sure how those changes of habitat will impact the coastal ecosystems. We are studying their reaction in laboratory environments and in the field, identifying thresholds for their functionality and capacity to supply ecosystem services.
Finally, the identification and mitigation of conflicts with other users of those ecosystems is also key to reduce the social risks for all beneficiaries, including labour, human rights, public health issues, and political uncertainty.
When it comes to stakeholders and non-experts, is science communication around the topic of carbon sequestration in coastal ecosystems effective?
On a national and international level, Germany seems to be on track with setting targets and planning actions to become climate neutral through the Climate Action Programme 2030.
The CDRmare research mission and in particular the sea4soCiety project on carbon sequestration in coastal ecosystems are in a way part of that effort, receiving funding to provide the knowledge base for the action programme. Thus, to some extent, the science communication on that level is working and the action plan is based on scientific knowledge.
The shortcomings are in the implementation phase. Local governments are often lacking established mechanisms that allow and support the implementation of action plans related to the national targets.
Such regulated implementation strategies should operationalise the uptake of scientific knowledge in the management of coastal ecosystems and by the local communities, and also in all fields of policy and management.
Now, we are up to another challenge: we have joined the Advisory Panel appointed by cOAlition S to help further the improvement and development of this important service. The Advisory Panel consists of twelve members (six publishers and six end-users) representing different stakeholders in the scholarly communication ecosystem.
Journal Comparison Service (JSC) is an initiative by cOAlition S aimed to improve transparency and communication regarding publishing costs between publishers and institutions.
It serves to provide the libraries with all the information they need to make informed decisions about whether the fees charged by a particular journal are reasonable and commensurate with the services delivered.
In their turn, the publishers can use it to demonstrate their dedication to fostering an open business culture and to bring awareness of the value of their services.
To facilitate this process, the publishers are advised to submit information about their prices and publishing policies on an annual basis using the JCS Frameworks format.
An Advisory Panel will review the Frameworks and offer suggestions on how to improve them, aiming to make the data collected as valuable as possible to all involved parties. Additionally, the Panel will actively promote the use of JCS among stakeholders.
The panel will meet twice a year, and the first meeting has already been scheduled for May 2023.
We are delighted that we have been able to establish such a high-quality Advisory Panel, representing all the key stakeholders. The primary function of the Panel will make recommendations on how the data collection frameworks might be further developed to ensure that the price and service data is as useful as possible for those who procure publishing services, whilst remaining deliverable by the publishers who are asked to provide these data,
commented Robert Kiley, Head of Strategy at cOAlition S.
Additional information
About JCS:
Journal Comparison Service is a secure, free-of-charge service that enables libraries, library consortia, and funders to better understand if the fees they pay are commensurate with the publication services delivered. Publishers provide information in a standard format, including information about the publication frequency, the peer review process, times from submission to acceptance, the range of list prices for APCs and subscriptions and more.
About cOAlition S:
On 4 September 2018, a group of national research funding organisations, with the support of the European Commission and the European Research Council (ERC), announced the launch of cOAlition S, an initiative to make full and immediate Open Access to research publications a reality. It is built around Plan S, which consists of one target and 10 principles. Read more on the cOAlition S website.
The so-called High Seas Treaty is a legal framework for the protection of marine biodiversity and responsible and equitable use of resources of areas beyond national jurisdiction (BBJN). Its draft, published on the 5th of March 2023, is the outcome of two decades of negotiations, and is part of the international effort to protect a third of the world’s biodiversity by 2030.
An unwavering dedication to the protection and conservation of biodiversity will be required to see the firm landing of this hopeful step.
On this occasion, we look back at some impactful studies published in our journals that have made waves, hopefully in the right direction towards impactful conservation measures and actions.
Following President Barack Obama’s expansion of the largest permanent Marine Protected Area on Earth (Papahānaumokuākea Marine National Monument) in 2016, a new species of coral-reef fish was named in his honour. The fish is the only known coral-reef species to be endemic to the Monument, and, despite its small size, it carries wide-reaching cultural and political significance as a reminder of how politics go hand in hand with science.
Former President of the United States, Barack Obama, arriving on Midway Atoll Midway on September 1, 2016 to commemorate his use of the Antiquities Act to expand the boundaries of the Papahānaumokuākea Marine National Monument. Dr. Sylvia Earle gives President Barack Obama a photograph of Tosanoides obama on Midway Atoll, from the film “Sea of Hope: America’s Underwater Treasures” premiered on National Geographic Channel on January 15, 2017. See also the news story on National Geographic.
Other studies from our flagship zoology journal ZooKeys have focused on the benthic megafauna and abyssal fauna of the Clarion-Clipperton Zone (CCZ) in the Pacific Ocean.
The Clarion-Clipperton Zone, managed by the International Seabed Authority, has been targeted by deep-sea mining interests. In the context of heightened concern over potential biodiversity loss, scientific research is crucial for informing policy-makers and the general public about the risks and outcomes of such initiatives.
The Clarion-Clipperton Zone, central Pacific Ocean (purple box), spanning 6 milllion km2. Knowledge of marine biodiversity in the area is crucial for its protection. Image source: A. Glover at al. (2016).
The rich biodiversity of the deep sea has also been documented in big-scale taxonomic inventories and checklists in the Biodiversity Data Journal.
The echinoderm Amphioplus cf. daleus Lyman, 1879. Image source: A. Glover at al.Hymenopenaeus cf. nereus observed in the UK-1 exploration contract area. Image source: Amon et al. (2017).
Going forward, the expansion of Marine Protected Areas should also ensure the implementation of policies for the methods of resource extraction and their equitable sharing and use among the world’s nations.
Over the next few years, we hope to see an ever increasing interest in biodiversity conservation - from the general public, stakeholders and policy makers, and, of course, research institutions.
We need to love what we protect in order to be able to protect it.
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Costus prancei, one of the 18 newly described species in Costaceae.
I am a retired government bureaucrat who worked for 40 years as an administrator in state and federal taxation. I have absolutely no formal training in botany, but now I find myself as an active participant in a major taxonomic revision and a coauthor in the publication of 18 new species in a plant family called Costaceae. This is the story of how my gardening hobby turned into an avocation and led me to work with some of the premier botanists in the world. It is also the story of how I have met several other plant enthusiasts from countries throughout the tropics who have contributed so very much to our work. I write this story in the hopes of encouraging more professional scientists to incorporate the observations of such “citizen scientists” in their research, and to encourage these enthusiasts to more carefully document their observations and post their photos and notes to resources like Inaturalist.org.
Costus whiskeycola, one of the 18 newly described species in Costaceae.
My story started about 30 years ago when my wife gave me a rhizome of the white butterfly ginger (Hedychium coronarium) as a Christmas present. I became interested in gingers, species of the family Zingiberaceae, but soon my interests began to focus almost exclusively on the closely related “spiral gingers” in the family Costaceae. I loved the architecture of the plants with their spiral staircase of leaves leading up to a variety of shapes and colors of bracts and flowers. I started collecting any cultivated Costus plants I could find in nurseries or mail-order catalogues. Soon, I learned that only a few species can survive outdoors in the winter where I live, so built a greenhouse.
Costus convexus, one of the 18 newly described species in Costaceae.
My serious interest in Costaceae began after I obtained a copy of the 1972 monograph of New World Costaceae by Dr. Paul Maas. It became my bible.
As I studied his descriptions of the species and applied his identification keys to the cultivated plants, I soon realized that many of the popular Costus species in cultivation had been incorrectly identified. I started doing presentations to garden clubs and posting to online groups. I developed a website called “Gingers ‘R’ Us.”
My “real job” had me traveling to Washington, DC periodically and I always tried to carve out time to visit Mike Bordelon at the Smithsonian Greenhouses in Suitland, Maryland. On one of these trips, I met Dr. Chelsea Specht, who was working at the Smithsonian Institution as a postdoctoral fellow.
Chelsea Specht and Mike Bordelon at the Smithsonian Greenhouses in 2004.
She had written what I believe is the first molecular study in Costaceae in 2001.This opened up a whole new world of interest for me as I tried to understand these new-to-me terms, like “clades” and “phylogenetic relationships”. In this paper she introduced the new generic divisions of the family that were solidified five years later in a more complete phylogenetic study . Chelsea very patiently answered my novice questions about phylogenetic trees and how they relate to the taxonomy of the plants.
Reinaldo Aguilar in 2013 at the type locality of Costus maritimus, now a synonym in the Costus comosus complex.
In 2005 I made my first trip to the New World tropics looking for Costus in its native habitat. On the Osa Peninsula of Costa Rica, I was incredibly lucky to meet Reinaldo Aguilar, the world-famous “para-taxonomist” who has studied the plants of the Osa for over 30 years. He is is self-taught like me and does not have a botanical degree, but has coauthored many scientific articles. He worked closely with the late Scott Mori of the New York Botanical Garden and was honored in a 2017 article in NYBG Science Talk.
That first trip to Costa Rica had me hooked. I fell in love with tropical forests and over the next few years made trips to several other Latin American countries as well as back to Costa Rica. Always, my focus was on Costus and the other members of its family.
Along the way, I met several “unsung heroes” in the plant world, like Marco Jiménez Villata, whom I met in the town of Zamora in southern Ecuador. Marco specializes in orchids, but he is also a generalist and knows a lot about the plants of southern Ecuador. He (now retired) was a school administrator and had traveled to many remote villages in the province and was always on the lookout for interesting plants. I have traveled with Marco and his son Marco Jiménez León several other times and we have become good friends.
Marco Jimenez and son Marco with Costus convexus.
In 2015 we went to the type locality of the species Costus zamoranus and took the first photographs of this species. At that trip, Marco showed me an area of high elevation near the Podocarpus National Park, where I found an unusual-looking Costus that we are now describing as Costus oreophilus. He also showed me unexplored places where I found another new species, Costus convexus. I made sure we credited him with his role in the discovery and documentation of those new species in our publication in PhytoKeys.
I have also traveled several times in Panama and Ecuador with another very well known, but non-doctorate plant enthusiast – Carla Black. Carla is the president of the Heliconia Society International, an organization uniting enthusiasts (scientists and non-scientists) in the order Zingiberales.
Carla Black with Juan Carlos Amado on the old Camino Real.
In 2015 we searched for the critically endangered Costus vinosus. We found a few plants growing deep in the forest of the Chagres National Park along an old Spanish trail used to transport gold to the Atlantic coast. There is still a mystery regarding the true form of the flower of C. vinosus, and I am in touch with another Inaturalist observer who has found it (not in flower) in the mountains northeast of Panama City. He will let me know when he finds it in flower!
Costus callosus, one of the 18 newly described species in Costaceae.
In 2019 Carla and I visited the “Willie Mazu” site in Panama to photograph and study the new species Costus callosus, and in Santa Fé de Veraguas, we looked for a species proposed by Dr. Maas that is now described as Costus alleniopsis.
My serious collaboration with Dr. Maas began in 2017, when I was preparing for a trip to Oaxaca in southern Mexico. He asked me to be on the lookout for two species of Costus from that region that he had identified as new based solely on his examination of herbarium specimens, without any good data on the floral parts.
By that time, I was posting my Costus observations on Inaturalist.org and using that resource to look for interesting plants. I also used it to find plant people to contact for local information. For this Mexico trip I found a huge number of observations posted by Manuel Gutiérrez from Oaxaca City.
Manuel Gutiérrez photographing the plant that turned out to be Costus sepacuitensis.
I found that he had extensive knowledge of the Chinantla region in the mountains east of Oaxaca City and had worked with the indigenous tribe there. Together, we explored the indigenous lands of Santa Cruz Tepetotutla.
We found many plants in flower of what Dr. Maas wanted to describe as Costus alticolus. We also found the species he planned to describe as Costus oaxacus, but I later found the same species in Guatemala, already described as Costus sepacuitensis.
Later I learned of the plans to prepare a complete revision to the taxonomy of the New World Costaceae. Together with Paul and Hiltje Maas, we spent several days at the Naturalis Herbarium in Leiden, comparing my photos against the hundreds of Costus herbarium specimens there. I had a long list of species that was curious about, and we were able to get through it and figure out what questions remained, even though we had not come up with all the answers.
Dave Skinner and Paul Maas discussing some Costus spp. in Leiden in 2017.
It was soon apparent that there are major changes needed in the taxonomy and nomenclature of these plants, and that information from the field would be an essential supplement to the observations made from the herbarium specimens.
Paul and Hiltje Maas in Leiden in 2017.
In 2016 I visited the type locality of Costus laevis in central Peru. I was surprised to find that the plants there are nothing at all like the Costus laevis of Central America, but match perfectly to the herbarium specimen that was deposited in Spain over 230 years ago. It was clear to me that the herbarium specimen designated as the type had been misinterpreted. I wrote an article explaining the problem – but I had no idea what the solution might be.
Dr. Maas agreed that there was a problem with that species that we eventually resolved. This resolution will be a part of the forthcoming revision of the New World Costaceae that is in preparation, nearing completion.
Another major problem involved the Costus guanaiensis complex. Paul and Hiltje, along with Chelsea, had visited the New York Botanical Garden Herbarium, where the holotype of that species is held, and realized that it had been misinterpreted due to the lack of a good flower description. What had been identified as Costus guanaiensis in the herbarium was actually a completely different species that Maas had planned to describe as a new species.
Dave Skinner with a plant in cultivation of Costus gibbosus at Rio Palenque Science Center, Ecuador.
The entire C. guanaiensis complex needed name changes and redefinitions of species boundaries, ultimately resulting in the description of Costus gibbosus that is published in PhytoKeys. The resolution of the other members of that complex will be explained in the forthcoming revision. Over the next several years, Paul and I exchanged 1,626 emails (yes, I counted them – with the help of MS Outlook) pounding out the details of the changes needed in the taxonomy of New World Costaceae. In collaboration with him, I made many more field trips to resolve the remaining questions we had.
My extensive collaboration with Paul Maas has been one of the most rewarding experiences of my lifetime. He has taught me so much about the rules of nomenclature and the process of describing a new species. The one thing he could never teach me was his almost uncanny ability to look at a dried herbarium specimen and make a determination of the species. I suppose that only comes from experience as he has examined over 11,000 specimens of Costaceae that will become our list of exxicatae when the full revision is published.
I should not fail to mention my time working with Dr. Thiago André. In 2014 I flew to Rio de Janeiro and then Thi and I, along with his academic advisor and another student, went to the state of Espirito Santo to look for the endangered species Chamaecostus cuspidatus. Thi has been our expert in that genus and has helped with the review of the new species published in PhytoKeys, Chamaecostus manausensis. In 2014 he was still finishing his doctorate and was in process of preparing a molecular phylogeny and morphological study of the species complex of Chamaecostus subsessilis.
Thiago André with Chamaecostus cuspidatus in 2014.
Thi and I have stayed in close contact, and he came to Florida one year to visit in my home and see the Costaceae in my private garden, Le Jardín Ombragé. He is now a professor at the Universidade de Brasília.
Finally, I should discuss my collaboration with Eugenio Valderrama and the other members of the Specht Lab at Cornell University. In 2018 I went to Cornell to visit Eugenio and we discussed the sampling to be used in the molecular phylogeny that will be a very important part of the full revision when it is published.
Eugenio Valderrama and Chelsea Specht with Costus convexus.
At Cornell, Eugenio produced a novel baiting schema for extracting specific genes from across all Costus species and in 2020 published a paper. With further sampling, another paper was published in 2022 to reveal interesting data on a whole package of pollination-related characters, and how they show evidence of convergent evolution. Eugenio’s phylogenies very well support the new species we are publishing in PhytoKeys, and the full molecular phylogeny will be included in our full revision when it is published.
Eugenio checking out a Renealmia sp. Antioquia, Colombia 2022.
Just this past December I went to Colombia to attend the Heliconia Society Conference at Quindío, and Eugenio and I each made presentations there about our work with Costaceae. Then we traveled together to investigate several other interesting species of Costaceae, including the new species Costus antioquiensis, and a strange yellow bracted form of Costus comosus found in the species-rich area of San Juan de Arama in Meta.
How did I know to look there? An observer, a citizen scientist, had posted his records and photos on Inaturalist.org. I have my account set to filter all Costaceae and send me a daily email with all the new postings of the family, and this plant will now be appearing as a sample in a molecular phylogeny and as an observed species in a monograph.
I hope this blog article will provide some background and insight into what I think must be an unusual collaboration between a citizen scientist and the much more qualified lead authors of our PhytoKeys article describing eighteen new species in Costaceae. It has certainly been a rewarding experience for me, and I hope other plant enthusiasts will be encouraged to share their observations on forums like Inaturalist.org, providing detailed and accurate information and photos. At least for the one plant family I have some expertise in, I will continue to monitor and curate those observations on Inaturalist.
André T, Specht CD, Salzman S, Palma-Silva C, Wendt T (2015) Evolution of species diversity in the genus Chamaecostus (Costaceae): Molecular phylogenetics and morphometric approaches. Phytotaxa 204(4): 265-276. https://doi.org/10.11646/phytotaxa.204.4.3
Maas, P. J. M. (1972). Costoideae (Zingiberaceae). Flora Neotropica 8, 1–139. doi: 10.1093/aob/mch177
Maas PJM, Maas-van de Kamer H, André T, Skinner D, Valderrama E, Specht CD (2023) Eighteen new species of Neotropical Costaceae (Zingiberales). PhytoKeys 222: 75-127. https://doi.org/10.3897/phytokeys.222.87779
Salzman S, Driscoll HE, Renner T, André T, Shen S, Specht CD (2015) Spiraling into history: A molecular phylogeny and investigation of biogeographic origins and floral evolution for the genus Costus. Systematic Botany 40(1): 104–115. https://doi.org/10.1600/036364415X686404
Skinner D (2008) Costus of the Golfo Dulce Region. Heliconia Society Bulletin 14(4):1-6
Skinner D and Jiménez M (2015) Costus zamoranus: An endemic species to Zamora-Chinchipe Province in Southeastern Ecuador. Heliconia Society Bulletin 21(3):4-9
Skinner D (2016) Following Ruiz. Heliconia Society Bulletin 22(4): 7–14.
Skinner D and Black C. (2016) Search for the Mysterious Lost Plant (Costus vinosus). Heliconia Society Bulletin 22(3):1-3
Skinner D (2019) A Tale of Two Costus (Costus sepacuitensis) and Costus cupreifolius) Heliconia Society Bulletin 25(1):1-3
Specht CD, Kress WJ, Stevenson DW, DeSalle R (2001) A molecular phylogeny of Costaceae (Zingiberales). Molecular Phylogenetics and Evolution 21(3): 333–345. https://doi.org/10.1006/mpev.2001.1029
Specht CD, Stevenson DW (2006) A new phylogeny-based generic classification of Costaceae (Zingiberales). Taxon 55(1): 153–163. https://doi.org/10.2307/25065537
Valderrama E, Sass C, Pinilla-Vargas M, Skinner D, Maas PJM, Maas-van de Kamer H, Landis JB, Guan CJ, AlmeidaA., Specht CD (2020) Unraveling the spiraling radiation: A phylogenomic analysis of neotropical Costus L. Frontiers in Plant Science 11: 1195. https://doi.org/10.3389/fpls.2020.01195
Valderrama E, Landis JB, Skinner D, Maas PJM, Maas-van de Kamer H, Sass C, Pinilla-Vargas M, Guan CJ, Phillips R, Almeida A, Specht CD (2022) The genetic mechanisms underlying the convergent evolution of pollination syndromes in the Neotropical radiation of Costus L.Frontiers in Plant Science 13: https://doi.org/10.3389/fpls.2022.874322
