Around 2004, a poison dart frog resembling the polka dot poison dart frog Oophaga arborea was smuggled into the European pet trade. The striking animal had yellow dots characteristic of the species and caused a sensation. Part of the excitement is because Oophaga arborea is a critically endangered species is from a very small known distribution where it inhabits bromeliads in the tree canopies. It was thought to be common in the 1980’s but may have declined due to chytridiomycosis. According to the IUCN redlist it was last seen in 2012, but was possibly heard from the Fortuna forest reserve more recently. Could this animal indicate that the polka dot poison dart frog survived the amphibian chytrid epidemic?
Male Oophaga vicentei photographed by Ariel Rodríguez
In June 2022, a population of frogs with yellow spots resembling the polka dot poison frog was discovered in Veraguas, outside the known distribution of the polka dot frog. The researchers collected a few specimens and analyzed them genetically where they most closely matched Vicente’s poison frog Oophaga vicentei. These dart frogs are known to be highly polymorphic, coming in slate gray, metallic blue, yellow or brick red with mottling. The yellow polka dot version, however, has now been confirmed to be Vicente’s poison dart frog as opposed to strawberry poison dart frogs Oophaga pumilio, or Oophaga arborea. Vicente’s poison dart frogs are listed by the IUCN as an endangered species, they can be locally abundant in places they are found and are known from a small area in Panama.
Habitat destruction and disease are both well-documented causes of the decline of amphibians—among the most threatened animals on the planet—but a new paper analyzing two decades’ worth of data from around the world has found that climate change is emerging as one of the biggest threats to frogs, salamanders, and caecilians. The study was published today, Oct. 4, in the scientific journal Nature.
The study, “Ongoing declines for the world’s amphibians in the face of emerging threats,” is based on the second global amphibian assessment, coordinated by the Amphibian Red List Authority, which is a branch of the International Union for Conservation of Nature’s Species Survival Commission’s Amphibian Specialist Group, hosted and managed by Re:wild.
The assessment evaluated the extinction risk of more than 8,000 amphibian species from all over the world, including 2,286 species evaluated for the first time. More than 1,000 experts across the globe contributed their data and expertise, which found that two out of every five amphibians are threatened with extinction. These data will be published on the IUCN Red List of Threatened Species™.
Between 2004 and 2022, a few critical threats have pushed more than 300 amphibians closer to extinction, according to the study. Climate change was the primary threat for 39% of these species. This number is expected to rise as better data and projections on species’ responses to climate change become available. Climate change is especially concerning for amphibians in large part because they are particularly sensitive to changes in their environment.
Geminis’ dart frog is a critically endangered Panamanian amphibian that was only recently described. It is known from an area just 40km 2, males care for the offspring and transport tadpoles on their backs.
“As humans drive changes in the climate and to habitats, amphibians are becoming climate captives, unable to move very far to escape the climate change-induced increase in frequency and intensity of extreme heat, wildfires, drought and hurricanes,” said Jennifer Luedtke Swandby, Re:wild manager of species partnerships, Red List Authority coordinator of the IUCN SSC Amphibian Specialist Group, and one of the lead authors of the study. “Our study shows that we cannot continue to underestimate this threat. Protecting and restoring forests is critical not only to safeguarding biodiversity, but also to tackling climate change.”
Habitat destruction and degradation as the result of agriculture (including crops, livestock like cattle and livestock grazing, and silviculture), infrastructure development and other industries is still the most common threat, according to the paper. Habitat destruction and degradation affect 93% of all threatened amphibian species. Expanded habitat and corridor protection in the places most important for biodiversity is going to continue to be critical.
Disease caused by the chytrid fungus–which has decimated amphibian species in Latin America, Australia and the United States–and overexploitation also continue to cause amphibian declines. Habitat destruction and degradation, disease, and overexploitation are all threats that are exacerbated by the effects of climate change.
The study also found that three out of every five salamander species are threatened with extinction primarily as the result of habitat destruction and climate change, making salamanders the world’s most threatened group of amphibians. North America is home to the most biodiverse community of salamanders in the world, including a group of lungless salamanders abundant in the Appalachian Mountains of the eastern United States. Because of this, conservationists are concerned about a deadly salamander fungus that has been found in Asia and Europe, called Batrachochytrium salamandrivorans (Bsal), entering the Americas.
“Bsal has not yet been detected in the United States, but because humans and other animals can introduce the fungus to new places, it may only be a matter of time before we see the second global amphibian disease pandemic,” said Dede Olson, a research ecologist with the USDA Forest Service, member of the IUCN SSC Amphibian Specialist Group, and co-author on the paper. “It is critical that we continue to implement proactive conservation actions to prevent the spread of Bsal into the United States, including effective biosecurity practices for wild and captive amphibians, and rapid detection and response measures. The North American Bsal Task Force includes a multi-pronged strategic plan that includes: a continental surveillance and monitoring network; research studies identifying high-risk geographies and species; and collaborative partnerships across public, private, and governmental sectors.”
The Nature paper provides an update to the 2004 landmark paper that was based on the first global amphibian assessment for the IUCN Red List, which revealed the unfolding amphibian crisis for the first time and established a baseline for monitoring trends and measuring conservation impact. According to this new study, nearly 41% of all amphibian species that have been assessed are currently globally threatened, considered critically endangered, endangered or vulnerable. This is compared to 26.5% of mammals, 21.4% of reptiles and 12.9% of birds.
Four amphibian species were documented as having gone extinct since 2004—the Chiriquí harlequin toad (Atelopus chiriquiensis) from Costa Rica, the sharp snouted day frog (Taudactylus acutirostris) from Australia, Craugastor myllomyllon and the Jalpa false brook salamander (Pseudoeurycea exspectata), both from Guatemala. Twenty-seven additional critically endangered species are now considered possibly extinct, bringing the total to more than 160 critically endangered amphibians that are considered possibly extinct. The assessment also found that 120 species improved their Red List status since 1980. Of the 63 species that improved as the direct result of conservation action, most improved due to habitat protection and management.
“The history of amphibian conservation itself proves how vital this information is,” said Adam Sweidan, chair and co-founder of Synchronicity Earth. “If the IUCN Red List had been updated on a similar scale in the 1970s that it is today, we could have traced the sweeping amphibian disease pandemic 20 years before it devastated amphibian populations. It isn’t too late–we have this wealth of information, we have the Amphibian Conservation Action Plan, but plans and information are not enough. We need to act. We need to act fast.”
Conservationists will use the information from this study to help inform a global conservation action plan, to prioritize conservation actions at the global level, to seek additional resources, and to influence policy that can help reverse the negative trend for amphibians.
With the formation of the Atelopus Survival Initiative (ASI)–a new alliance of more than 40 organizations from 13 countries–comes a new day for harlequin toads, the jewels of South and Central America’s forests and creeks and a group of amphibians hardest hit by the deadly chytrid fungus Batrachochytrium dendrobatidis (Bd).
While amphibian researchers and conservationists have worked for many years to save harlequin toads (which make up the Atelopus genus) and groups of species in individual countries, the ASI is bringing them together for the first time to pool the resources, decades of experience and knowledge necessary to prevent the extinction of the entire genus of harlequin toads across the region where these species still survive.
“As an incredibly diverse group of amphibians facing a number of threats, harlequin toads require innovative solutions coming from a diverse group of individuals and organizations with different expertise, knowledge and capacities,” said Lina Valencia, ASI founder, co-coordinator of the IUCN SSC Amphibian Specialist Group Atelopus Task Force and Andean countries coordinator for Re:wild, one of the primary ASI conveners. “More than ever before, we need a constellation of champions working together to bring harlequin toads back from the brink of extinction. The ASI underscores the vital need to implement on-the-ground conservation actions that will mitigate the main threats to this beautiful group of amphibians.”
Over the past few decades, many harlequin toad species have suffered severe population declines and extinctions throughout their range. Today, of the 94 harlequin toad species that have been assessed by the IUCN, 83 percent are threatened with extinction, while about 40% of Atelopus species have disappeared from their known homes and have not been seen since the early 2000s, despite great efforts to find them. Four harlequin toad species are already classified as extinct, according to the IUCN Red List of Threatened Species, but this number is likely higher.
The fungus Batrachochytrium dendrobatidis (Bd) causes the lethal disease chytridiomycosis, which has resulted in amphibian declines all around the world, including in South and Central America, Australia and the western United States. Although Bd may likely be the primary driver of these declines, a number of other threats are exacerbating the precipitous drops in population numbers. This includes habit destruction and degradation (as the result of animal agriculture, logging, mining and infrastructure development), the introduction of invasive species such as the rainbow trout that prey on harlequin toad tadpoles, pollution, illegal collection for the pet trade, and the effects of climate change.
The ASI and its members, including governments, local communities and Indigenous peoples, will collaboratively address each of these threats–and new ones as they arise–across the genus’s full range, taking into account the social, political and cultural realities of each of the 11 countries where harlequin toads are found.
“With their beautiful songs and unique lifestyles, amphibians are among the most extraordinary animals on Earth, and among them, harlequin toads stand out for their amazing colors,” said Luis Fernando Marin da Fonte, coordinator of the ASI and director of partnerships and communications for the Amphibian Survival Alliance. “But these colorful and delicate jewels are becoming increasingly rarer. Harlequin toads must be protected not only because of their beauty and uniqueness, but also because of their intrinsic value and biological, ecological and even cultural importance.”
The initiative’s newly developed Harlequin Toad (Atelopus) Conservation Action Plan (HarleCAP) provides the roadmap for conserving and restoring harlequin toads as a genus and their habitat. The action plan’s goals, which ASI aims to achieve by 2041 (the 200th anniversary of the description of the genus Atelopus), include:
protecting and restoring harlequin toads’ forests and watersheds;
creating and maintaining conservation breeding programs;
searching for species that are lost to science and filling in other gaps in scientific knowledge about harlequin toads;
sharing stories that will transform harlequin toads into symbols of hope for the region and the world and a flagship for conservation success, and demonstrate a commitment to the conservation of harlequin toads;
ensuring the Atelopus conservation network has the technical, logistical, and financial support to secure the long-term conservation of harlequin toads
“The establishment of collaborative initiatives at the international and regional level is essential to coordinate efforts and obtain tangible results that have an efficient and real impact on the conservation of an endangered species,” said Gina Della Togna of the Universidad Interamericana de Panamá, Panamá. “The Atelopus Survival Initiative is a concrete example, which not only aims to conserve one species, but an entire genus, perhaps the most threatened by the global amphibian extinction crisis.”
Harlequin toads are found from Costa Rica in the north to Bolivia in the south, and Ecuador in the west and French Guiana to the east. They are known as the jewels of South and Central America in part because of their beautiful and varied colors, which range from orange, green, yellow, brown, black, red, and sometimes even purple. They are celebrated in a number of Latin American cultures, including Indigenous cultures, and across entire countries, like in Panama, where the national animal is the Panamanian golden toad.
Like other amphibians, harlequin toads support healthy ecosystems. Their tadpoles depend on clean water and, because of this, the presence of harlequin toads indicates better quality water in an ecosystem, while their decline or absence is often the first sign of an ecosystem in trouble.
“Protecting and restoring harlequin toads and their habitats will also benefit the species that share the ecosystems in which they live and that provide water to tens of millions of people, and ultimately all life on Earth,” Valencia said. “And we’re hoping that the ASI will be a successful model that conservationists can emulate for other groups of threatened species.”
The Atelopus Survival Initiative includes national and international conservation groups, zoos, captive breeding centers, academic institutions, governments and local communities. Its current members represent the following organizations: Amphibian Ark, Amphibian Survival Alliance, Asociación Pro Fauna Silvestre – Ayacucho, Bioparque Municipal Vesty Pakos, Bolivian Amphibian Initiative, Centre National de la Recherche Scientifique, Centro de Conservación de Anfibios AMARU, Centro Jambatu de Investigación y Conservación de Anfibios/Fundación Jambatu, CORBIDI, DoTS, El Valle Amphibian Conservation Center Foundation, Facultad Latinoamericana de Ciencias Sociales, Florida International University, Fort Worth Zoo, Fundación Atelopus, Fundación Zoológica de Cali, Universidad del Tolima (GHEE), Grupo de Trabajo Atelopus Venezuela, Image Conservation, Instituto Nacional de Pesquisas da Amazônia, Instituto Venezolano de, Investigaciones Científicas, Ministerio del Ambiente de Perú, MUBI (Museo de Biodiversidad del Perú), Parque Explora, Parque Nacional Natural Puracé, Photo Wildlife Tours, Pontificia Universidad Católica del Ecuador, Pontificia Universidad Javeriana, Re:wild, San Diego State University, Smithsonian Tropical Research Institute, Trier University, Universidad de Antioquia, Universidad de Costa Rica, Universidad de los Andes, Universidad del Tolima, Universidad del Magdalena, Universidade Federal do Pará, Universidad Nacional, Universidad Interamericana de Panamá, Universidad Nacional de Colombia, Universidad San Francisco de Quito, Universidade Estadual de Campinas, Universidade Federal do Oeste do Pará, University of Nevada, Reno, University of Notre Dame, University of Pittsburgh, WCS (Wildlife Conservation Society), WCS Colombia, Zoológico Cuenca Bioparque Amaru
An exciting new publication has just been released by the International Union for the Conservation of Nature (IUCN) of best practice guidelines for a wide range of amphibian conservation translocations. The project was many years in development through the coordinated effort of numerous translocation specialists across the globe, but the project the led by the Panama Amphibian Rescue and Conservation Project’s post-doctoral research fellow Dr. Luke Linhoff. The guidelines cover the reasons for conducting amphibian translocations, pre-translocation planning and risk assessment, and also cover important topics such as disease, welfare, human social dimensions, post-release monitoring and reporting results.
Milan Vesely and Abel Batista scientifically described Panama’s seventh harlequin frog species in the journal Zoological Research in April 2021. This beautiful harlequin frog species has been in biological collections for some time, the first museum specimen was first collected by Henri Pittier at Puerto Obaldlia in 1911, but has previously been assumed to be related to other sister species that it closely resembles. Historically collected formalin-preserved specimens were unsuitable for genetic analyses and so the frog remained undescribed.
Vesely and Batista conducted expeditions to the Darien to collect new specimens and this allowed them to conduct genetic analysis showing that this species is most closely related to Atelopus certus and Atelopus glyphus, but is genetically distinct enough to warrant recognition as a species. It also has a slightly different call and morphological characteristics that are also described in the paper.
Atelopus fronterizo Photo by Abel Batista, UNACHI-Fundación Los Naturalistas-SNI (SENACYT)
They named the frog Atelopus fronterizo to refer to borderland inhabitants and the Panamanian border security force who protect the Darien mountain range in NE Panama on the border where this frog is found. Like other harlequin frog species in Panama, even though they live in well-protected habitat, they are Likely Critically Endangered due to the threat of the amphibian chytrid fungus that has caused the declines of other Atelopus species in Panama.
Panamanian toads Rhinella centralis are distinguished by their dorsal skin covered with pointed warts. They are common along the Pacific coastal areas, often in urban areas around Panama City and small towns, and form large choruses on rainy nights. The small but strongly swollen poison glands on their heads secrete a white toxic goop. This effective defense mechanism makes predators spit them out, or froth at the mouth, vomit and it may even kill them if they try to eat the toad.
Scientists working at INDICASAT, the University of Panama and STRI began screening wild frogs for substances with pharmacological potential to treat various tropical diseases. When they analyzed secretions from these toads they discovered a chemical in the poison glands called 19-hydroxy-bufalin. They found that this chemical was very potent at killing the parasites that cause Chagas disease, and that it was not very toxic to cells. Chagas disease is a neglected tropical disease that kills 10,000 people per year, and current medications used to treat the disease are not very effective, particularly in acute cases. The fact that this chemical is quite selective with low cell toxicity means it is a promising compound that make it a candidate compound to further explore as a potential way to treat Chagas disease.
Salamanders are remarkably enigmatic amphibians, both due to their often-cryptic colorations and their extremely secretive lifestyles. They are nocturnal, but even at night, one can hardly call them “active”. Many species are burrowing, and barely ever come out of their moist and dark haven created by earth and fungi. Other species prefer a life in the canopy of mysterious and often nearly inaccessible cloud forests, in a world dominated by bark, moss and lichen.
Panamanian salamanders are no exception to this rule. Herpetologists that set out to find them, regularly return after their tiring night-time missions without seeing even a trace of these wonderful creatures. Local people that work and live in areas where salamanders occur sometimes don’t even know of their existence – which is exactly why a handful of Panamanian institutions decided to organize the first International Festival of the Salamander.
The Festival took place from November 1 to 3 in Boquete, located right at the border of Volcan Barru National Park. Aptly, this is the place to be if one wants to set out on a nocturnal quest to look for them. During the days, there were photographic exhibitions, stands of the organizations involved, an interactive kids corner with movies and drawings for coloring and a small T-shirt shop, hosted by a team of volunteers that where sitting on the edge of their seats to tell you about their beloved salamanders.
But Friday and Saturday night, things even got better: every evening from 6 to 10 p.m., there was a free guided tour by Los Naturalistas in Volcan Barru National Park, with experts knowing where exactly chances of spotting one where highest.
The tour started off at the entrance of Volcan Barru National Park, where we met with a team of tour guides and biologists, led by Dr. Abel Batista. Then we drove on for another 3 km on a bumpy gravel road, until we were at a place known for its substantial salamander population. The first and most important part of the tour consisted of disinfecting boots and equipment, to avoid spreading diseases. After all, one of the major reasons of amphibian declines worldwide is an extremely infectious fungus, Batrachochytrium sp.. While harmless to us, for many amphibian species, it causes severe skin damage, leading to death of susceptible individuals. After sterilizing our shoes, we began measuring about a dozen environmental variables; humidity, temperature, elevation, etc.. In the meantime, the guides explained a few rules: avoid touching the animals, don’t shine too bright lights directly in their eyes and don’t take pictures of them for too long and preferably without flash.
Then, we started searching. The weather was cold, but very dry, and therefore, our subjects of interest would mostly be hiding in moist places, one of the guides explained. We adjusted our strategies accordingly, looking in small holes between rocks and gently turning branches to peer underneath. We slowly walked upwards, and over the course of 20 minutes, we had only progressed about a hundred meters. When we had nearly given up, one of the guides suddenly called us further on.
We hurried on, nearly running uphill. But with a great reward waiting for us: a nearly 20 cm long, female magnificent web-footed salamander (Bolitoglossa magnifica) was cautiously watching us from underneath her hiding place – a thick, rotting branch covered with lichen, about a meter and a half away from the road. All excited, we had to take turns to take a look from the right angle, so you could see her. We all took a few pictures, then we stopped bothering her with our flashlights and left her in peace.
We started hiking down again, all the while scanning every hole and crevice. We passed the cars, and soon after, someone else already shouted he had found another one. This one was more difficult to see, hiding in a small hole in between rocks and loose earth. We had barely gotten to the second, when a third one was found, and soon after even a fourth and fifth!
Biologist taking photographs of a salamander (left), while a guide in the back continues searching in other holes. (right) A magnificent web-footed salamander (Bolitoglossa magnifica) that found a hiding place behind a mushroom.
Despite the unfavorable weather conditions, our evening excursion turned out to be a great success, spotting 5 individuals of the endangered magnificent web-footed salamander in less than an hour and a half. We returned tired and quite hungry, but fully satisfied nonetheless.
Los Naturalistas are currently working on a salamander-focused guided tour, which will roughly follow a similar structure as the tour we could enjoy that night. They hope to organize a lot of similar events like the International Festival of the Salamander, to further raise awareness among tourists as well as Panamanians concerning these mysterious and intriguing animals. Salamanders are facing many threats, but in protected places like Volcan Barru National Park, they are thriving.
By Leni Lamens Herpetologist, biologist and intern at the Panama Amphibian Research and Conservation center, in collaboration with the Smithsonian Tropical Research Institute and Smithsonian National Zoological Park.
By Leni Lammens
I would like to express my gratitude towards all organizing parties involved in the International Festival of the Salamander:
Action Hub+, Bioguias Panamá, Los Naturalistas, Universidad Autónoma de Chiriquí (UNACHI), Vicerrectoria de Investigación y Posgrado (VIP)
as well as to all sponsors, without which the Festival would not have been possible.
A new research paper published on strawberry poison dart frogs in Bocas del Torro found that one of the reasons we have polymorphism or so many different color forms within one species of frog. Female tadpoles prefer to mate with males that have the same color as their parents (sexual imprinting), and males defend their territories more vigorously from other males that are the same color as their parents (rival imprinting). The researchers demonstrated this experimentally by using foster parents of different color forms to raise offspring, and then tested mate or rival preference of the adult offspring.
This process of sexual selection can lead to sexual isolation even in populations that live in the same places. From an evolutionary perspective this would be a rare example of sympatric speciation, or the evolution through natural selection without geographical isolation.
An international study led by The Australian National University (ANU) has found that a fungal disease has caused dramatic population declines in at least 501 amphibian species, including 90 extinctions, over the past 50 years. The study involved collaborations with 41 different amphibian and wildlife disease experts from around the world. Smithsonian scientists contributed data from Panama—one of the worst-hit areas of the world by the disease—for the study.
Of the 90 confirmed extinctions of frogs across the globe, eight of those species were from Panama. Another 52 species of frogs in Panama have experienced more than a 90 percent decline.
“This study confirms that we are not dealing with a unique problem in Panama,” said Brian Gratwicke, amphibian biologist, international coordinator of the Panama Amphibian Rescue and Conservation Project, and one of the co-authors of the study who provided data. “If we or anyone does find a solution or cure for chytrid, it will likely have global implications.”
Collaborators like Smithsonian scientists allowed the lead researchers from ANU to get a first-hand insight into the conditions on-the-ground in countries around the world.
Chytridiomycosis, which eats away at the skin of amphibians, has completely wiped out some species, while causing more sporadic deaths among other species. Amphibians, which commonly live part of their life in water and the other part on land, mainly consist of frogs, toads and salamanders.
The deadly disease is present in more than 60 countries – the worst affected parts of the world are Australia, Central America and South America. The researchers found that chytridiomycosis is responsible for the greatest loss of biodiversity due to a disease.
The disease is caused by chytrid fungus, which likely originated in Asia where local amphibians appear to have resistance to the disease.
The unprecedented number of declines places chytrid fungus among the most damaging of invasive species worldwide, threatening similar numbers of species as rats and cats.
Lead researcher Ben Scheele, Fenner School of Environment and Society at ANU, said highly virulent wildlife diseases, including chytridiomycosis, were contributing to the Earth’s sixth mass extinction.
“The disease we studied has caused mass amphibian extinctions worldwide. We’ve lost some really amazing species,” said Scheele.
He said more than 40 frog species in Australia had declined due to this disease during the past 30 years, including seven species that had become extinct.
“Globalisation and wildlife trade are the main causes of this global pandemic and are enabling disease spread to continue,” said Scheele. “Humans are moving plants and animals around the world at an increasingly rapid rate, introducing pathogens into new areas.”
Scheele said improved biosecurity and wildlife trade regulation were urgently needed to prevent any more extinctions around the world.
“We’ve got to do everything possible to stop future pandemics, by having better control over wildlife trade around the world.”
Scheele said the team’s work identified that many impacted species were still at high risk of extinction over the next 10–20 years from chytridiomycosis due to ongoing declines.
“Knowing what species are at risk can help target future research to develop conservation actions to prevent extinctions.”
Scheele said conservation programs in Australia had prevented the extinction of frog species and developed new reintroduction techniques to save some amphibian species.
“It’s really hard to remove chytrid fungus from an ecosystem – if it is in an ecosystem, it’s pretty much there to stay unfortunately. This is partly because some species aren’t killed by the disease,” he said.
“On the one hand, it’s lucky that some species are resistant to chytrid fungus; but on the other hand, it means that these species carry the fungus and act as a reservoir for it so there’s a constant source of the fungus in the environment.
Video by Katie Garrett and Jonathan Kolby
Citation to the Paper: Scheele, B.C., Pasmans, F., Skerratt, L.F., Berger, L., Martel, A., Beukema, W., Acevedo, A.A., Burrowes, P.A., Carvalho, T., Catenazzi, A., De la Riva, I., Fisher, M.C., Flechas, S. V, Foster, C.N., Frías-Álvarez, P., Garner, T.W.J., Gratwicke, B., Guayasamin, J.M., Hirschfeld, M., Kolby, J.E., Kosch, T.A., La Marca, E., Lindenmayer, D.B., Lips, K.R., Longo, A. V, Maneyro, R., McDonald, C.A., Mendelson, J., Palacios-Rodriguez, P., Parra-Olea, G., Richards-Zawacki, C.L., Rödel, M.-O., Rovito, S.M., Soto-Azat, C., Toledo, L.F., Voyles, J., Weldon, C., Whitfield, S.M., Wilkinson, M., Zamudio, K.R., Canessa, S., 2019. Amphibian fungal panzootic causes catastrophic and ongoing loss of biodiversity. Science (80-. ). 363, 1459 LP-1463. https://doi.org/10.1126/science.aav0379