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Category Archives: Evolution

Lighter and Louder: The Evolution of Bass Amplification – Premier Guitar

Posted: December 7, 2021 at 5:31 am

Since achieving fame and fortune, Joe has spent most of his time in his various residences across the country. He's got Nerdville in sunny California and Nerdville East in Nashville. Both are almost-literal museums that house Joe's arguably unmatched collection of vintage guitars and amps. You will be very hard pressed to come across a finer collection. Yet in 2019, Joe found himself drawn back to New York City.

Joe Bonamassa - "Time Clocks" - Official Music Video

"It's a mid-life crisis. I always wanted to go back to where I lived 20 years ago, but not have to worry about hustling down sessions and gigs to make the rent every month," he says. "For as cool and exciting of a time as it was, it was also a very stressful time. I had this thing in my mind where I just wanted to be able to enjoy New York City and not have the stress of, 'Oh shit, it's either a subway ticket or ramen noodles tonight.'"

In February 2021, Joe went to Germano Studios in Manhattan's NoHo neighborhood to record Time Clocks, aka "The New York Album." Because of the pandemic, this recording session was like no other Joe had been involved with. Kevin "the Caveman" Shirley, Joe's longtime producer, was stuck in Australia due to strict COVID travel restrictions. The two have had a long working relationship and they've been inseparable for most of Joe's career. Shirley produced the guitarist's fifth studio album, You & Me, and has since produced over 30 of Joe's subsequent projects. So, when it came time to record Time Clockstravel restrictions be damnedthey found a way to work together. "We recorded it virtually with Zoom and some other technology where my producer in Australia could get the actual tracks from the session in real time or with about a second latency," says Joe. "It was totally fine. It was an odd record to make because of what we were doing, but it was also an odd time. So, why should anything be normal anymore?"

Even with the differences in time zones (Sydney, Australia, is 16 hours ahead of New York City), they found a way to make it work. "We would get there about noon. Kevin's an early riser so he would get to his studio, which is by his house in Sydney, by about 2 a.m. So, we would go from about noon to 6 or 7 at night, and he would go from 2 to 9 a.m., and then take naps."

Time Clocks is full of unexpected twists. It was recorded live as a trio with drummer Anton Fig and bassist Steve Mackey, and then other parts were layered in. The album has a diversity of sounds that belie Joe's blues categorization. "Notches" opens with an Ali Farka Tour-inspired 12-string riff, "Time Clocks" has a country-esque, Americana vibe, and "Curtain Call" is an homage to Led Zeppelin. "My ADD transcends into my musical life. It's a very different record for me. It's not a blues record, for sure. I just try to make records that don't bore me all the way throughwe've got this groove covered, we've got that groove covered, let's put a sorbet in, something out of left field," says Joe.

TIDBIT: Bonamassa's longtime producer, Kevin Shirley, had his hands on the wheel again, but remotely. He produced from Sydney, Australia, while the band recorded live on the floor in New York City.

"Some people think that all I do is play blues. I don't just play blues. I play whatever. We've all been in this game for a long time where we can adapt to any musical situation. It's fun. Every once in a while, I'll go sit in with friends. I just sat in with Nir Felder [at NYC's famed 55 Bar]. If I sit in with Nir, I'll be like, 'When these chord changes get too fucking adult for me, I'm bailing.' You just know your strengths and your weaknesses."

For the Time Clocks sessions, Joe used a much leaner gear selection than you might expect, especially given his cavernous collection. "In New York City, you go with what you got. There were only three amps. I had a [JB signature] high-powered Twin sent from our inventory that's still in the spare bedroom in the apartment. I ended up using two Deluxes. I had one Deluxe Reverb and one brown Deluxe," says Joe. We wonder if Joe was referring to a reissue Deluxe Reverb that might have happened to be at the studio, or one of his rare closet classics. He immediately snaps, "Think about that critically and ask yourself again, 'Who are you talking to?'" Point taken, Joe!

Im not going to live my life in indentured servitude to the fucking guitar.

"When I was a kid, I had a black [panel] Deluxe Reverb. I think this is probably one of the ones that I had for 25 years," he clarifies. "The black and brown circuits are totally different, so you get that kind of sweet/salt mix. The brown Deluxe does the real thick midrange stuff and the black does the low and the high. That's kind of always been my M.O. It's never just one amp. You're mid-stacking with amps that don't necessarily do the same thing. The bigger, thicker tones you hear on the solo, that's the high-powered Twin. The more jangly stuff was the Deluxes."

Joe also only used a handful of guitars on Time Clocks. "I don't keep a lot of guitars here," he says. "All I had was a Les Paul, a Strat, a 335, a Rickenbacker 12-string, an old bass from the movie Spinal Tap, the one that Nigel kept telling Rob Reiner not to touchI actually own that one. But it was less than six guitars. I also used a Martin acoustic. All the acoustic stuff was done with one acoustic.

A recent snapshot of Bonamassa's pedalboard.

"In Nashville, I have all my road guitars, but to be honest with you, over the years I've never gotten into this thing where you bring 50 guitars and use five. I did that a long time ago. It's nothing but a photo op. Most of the time I bring a Whitman's Sampler of what I think would work. The days of humping in 50 guitars and setting up six racks of them and going, 'Look what I got'that's a young man's game. That's for somebody in their 30s."

Joe's newfound minimalism goes hand in hand with living in a New York City apartment, where even playing with an amp on 1 will get you the "broomstick on a ceiling" retaliation. "I've been playing guitar and cranking amps for 40 years. Do you know the two things that prompt me to run away? Loud music and crowds. It's a paradox, I know," he says. "So, when I'm at home and I'm enjoying a very quiet Sunday afternoon, I have zero, absolutely zero, desire to crank an amp. And I have zero-plus-5-percent desire to sometimes even practice on a given day. It gives you a break. It's important to be good at your job on a given day. It's important to also step away and give yourself some perspective, so you're not so consumed by it."

Rig Rundown - Joe Bonamassa [2018]

Every aspiring guitaristno matter the genrelongs for their day in the spotlight. They too want to be a guitar god and inspire thousands of players and listeners, just like Joe. But what does it feel like on the other sidewhen you actually win?

The guitarist onstage with his frequent on-tour sparring partner in recent years, bass giant Michael Rhodes.

Photo by Debi Del Grande

"Then a whole 'nother set of circumstances come into play," he says. "It's managing time, managing your energy. It's also trying to keep in perspective what is it that you really do, because sometimes life comes at you twice as fast as it used to. All my energy is dedicated to the fans that keep me in business and come time and time again. That is 100 percent my biggest priority. When distractions and other things come into play that tend to take energy away from what you're supposed to be doing, that's the challenge. I realize I'm a very fortunate person, but I don't make any apologies for it, because, to be honest with you, that's what everybody strives for. Why should I apologize for working hard? I always tell people it's easy to dismiss, hard to replicate. If it was easy, as some claim, then it would be as easy as starting a TikTok. If it was that easy, then anybody could do it. But to be honest with you, anybody can do it. You've got to have the intestinal fortitude and the drive, and the ability to stick it out through thick and thin."

Bonamassa cradles his famed 1959 Les Paul Standard, Lazarus. The guitar was recreated for a limited-edition issue via Epiphone this year.

Older and wiser at 44, Joe, who started performing onstage at age 12, has now found time to explore other things in life besides guitar. He indulges in Law and Order marathons, is on an excruciatingly strict diet with Diet Coke as his only vice, and has found a new passion in cycling. He'll just as likely post details of his Central Park bike excursions on Instagram as he would another guitar-safari vintage find. Even with the potential danger of a career-ending fall, Joe, much like the late Allan Holdsworthwho was also an avid cyclistis willing to chance it. "If I fall and somehow my career ends on that particular day, then so be it," he says. "I'm not going to live my life in indentured servitude to the fucking guitar. If it's over, it's over. You've got to enjoy your life."

Joe Bonamassa is labeled a blues guitarist, but anyone that's heard him knows that he brings a huge diversity of stylistic elements to the genre. In this breathtaking, hyper-speed duo, Joe's right hand is like an unstoppable machine that keeps the intensity of the performance alive for over 6 minutes.

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Massive simulation of the universe probes mystery of ghostly neutrinos – Space.com

Posted: at 5:31 am

Paul M. Sutteris an astrophysicist at SUNY Stony Brook and the Flatiron Institute, host of "Ask a Spaceman" and "Space Radio," and author of "How to Die in Space." Sutter contributed this article to Space.com's Expert Voices: Op-Ed & Insights.

How do you test theories of the universe? By building gigantic supercomputers and simulating the evolution of the cosmos.

A team of Japanese scientists has built the largest-ever cosmic simulation to include tiny "ghost" particles called neutrinos. To explore one of physics' biggest unsolved mysteries, the researchers used a whopping 7 million CPU cores to solve for the evolution of 330 billion particles and a computational grid of 400 trillion units.

By far the most important form of matter in the universe is dark matter. We're not sure what it is or what it's made of, but we do know that there's a lot of it. It makes up about 80% of all matter. Baryonic matter the stuff that makes up stars, planets and the rich variety of the entire periodic table makes up just a small fraction of all the matter in the universe.

Related: Where did all the baryons go?

Dark matter forms the backbone of the cosmos. Billions of years ago, there were no structures in the universe. All of the matter dark or otherwise was smoothly distributed, and not lumpy at all. There simply weren't very many variations in density from place to place. Overall, it was a pretty boring universe.

But with time, the universe became more interesting. There were tiny density differences, seeded from microscopic quantum fluctuations in the early seconds of the Big Bang. Places with slightly higher density had slightly more gravity, and that's where dark matter began to pool together. As those early structures budded, they attracted even more material. Over billions of years, this process emptied out vast regions of the cosmos now known as cosmic voids pulling all the matter into an extensive network of clusters, walls and filaments.

And then there are neutrinos, extremely tiny particles that have barely any mass. Indeed, they make up less than 0.1% of all the mass in the universe. But these minuscule particles have an outsize influence on the evolution of structures. They are fast really fast capable of traveling at nearly the speed of light. This incredible speed dampens the formation of large structures, such as galaxies and clusters.

Whereas dark matter wants to keep piling up through gravity, neutrinos go too fast to settle down in one spot. And although neutrinos have very little mass, they still have some mass. They can use their gravity to weakly influence the behavior of dark matter, thus preventing it from clumping as tightly as it normally would.

In other words, the universe is a little smoother than it would be without neutrinos.

Finding the masses of the three known neutrino "flavors" electron neutrinos, muon neutrinos and tau neutrinos is a major unsolved problem in modern physics. But ironically, we can measure the masses of these tiny particles by mapping the largest structures in the universe.

To try to understand the nature of dark matter and the role of neutrinos in shaping cosmic evolution, cosmologists often turn to computer simulations. If you change the neutrino mass just a bit in the simulations, it will change how the neutrinos influence the formation of structures over billions of years. So by measuring those same structures, you can get an understanding of neutrino mass.

These simulations usually encompass a small fraction of the real universe and start with a set of dark matter "particles," with each particle representing a certain amount of dark matter for example, a single blob with a mass millions of times the mass of the sun. The simulations then position these particles as they would be in the early universe. The simulations track how those particles evolve through their mutual gravity, giving rise to the giant structures we see today.

This is an approximation technique, because the true behavior of dark matter is represented by a limited number of particles, but it works very well for dark matter. Simulating neutrinos is much more difficult because of their ridiculous speed. It's difficult to follow their behavior within the simulation because they can move from one side of the simulation to the other in a short amount of time. So the simulations can't keep up with how the neutrinos are acting and how they're influencing dark matter.

So maybe we shouldn't bother trying to approximate the behavior of neutrinos. To correctly follow the evolution of neutrinos and account for their fast behavior requires solving an incredibly complex equation. Solving this equation called the Vlasov equation, after Russian physicist Anatoly Vlasov however, requires immense computational resources.

So a team of Japanese scientists did just that: They used 7 million processors on the Fugaku supercomputer to trace the evolution of dark matter and the influence of neutrinos on the formation of structures. The researcher used 330 billion particles to represent dark matter and a computational grid of 400 trillion components to represent neutrinos, in the largest simulation of its kind.

And while it may not have solved the mystery of neutrinos' mass, the simulation does pave the way for more of its kind. In essence, this simulation was a proof-of-concept to show that we can now include neutrinos in simulations more accurately than ever before. Armed with this new technology, future simulations will open a window into the role of neutrinos in the universe and perhaps even reveal a key to unlocking their mass.

The team's paper was posted recently on the preprint server arXiv, and you can see the simulation here.

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Women’s FA Cup final: The evolution of women’s football – BBC Sport

Posted: at 5:31 am

On 5 December 1921 the Football Association banned women from playing football on league affiliated grounds, saying it was "most unsuitable" for them to play the game.

At the time the women's game was growing in popularity and attracting crowds of up to 53,000. The decision changed the course of female football forever.

The ban lasted nearly 50 years until an FA Council meeting in January 1970 rescinded the resolution made five decades earlier.

A year later the first women's FA Cup final was held and as the competition celebrates its 50 year anniversary season - and 100 years on from the controversial ban - BBC Sport speaks to female football legends on the evolution of the women's game.

Lesley Lloyd captained Southampton to glory in the first Women's FA Cup final in 1971, beating Scottish side Stewarton & Thistle 4-1 at Crystal Palace National Sports Centre. The Saints went on to win eight of the next 11 cups.

What was it like playing football during that time?

"I used to have a cheese and pickle sandwich before I walked out on to the pitch. We just turned up and that was it. When we went back to the changing rooms after playing we used to have to wash our boots with a pipe. It was crazy! We had no facilities but we just played for the love of it. We loved the game.

What was it like playing during the ban?

"We were laughed at, to be quite honest. We used to hear about this ban and boys used to make fun of us for being women playing football, but in the end I think they could see that we were serious about it."

What do you remember about that first FA Cup final and lifting the trophy?

"I remember getting to the final. We arrived at the pitch and they had forgotten to cut the grass.

"I remember the manager saying to us it was the most important game we were ever going to play so to get out there and do something.

"There was a wooden hut full of supporters. I'll never forget lifting the cup. That was my Wembley. I know now what the ladies have is brilliant, but to us that was it.

"We didn't even take a photograph of the team until the following week. We washed all our kit and the photo was taken a week later, not on the day. It shows how much things have changed."

How do you feel now watching a women's cup final and seeing way the game has developed?

"I think it's brilliant - they have everything. It's great to see what there is now. Women should be entitled to exactly what the men have and I'd love to have been born in this era. But no matter what era you're playing in, the competitive spirit doesn't change.

"I've still got people that come up to me and talk about it now. I'm still a season ticket holder at Southampton and my whole family have been brought up on football. My children and grandchildren love watching women's football. I'm actually bringing my grandson to the final at Wembley. You never lose the love of the game."

Former England captain Gill Coultard had a 25-year playing career from the mid 70s until 2001, winning the FA Cup six times with Doncaster Belles. Having become the first woman to reach 100 England caps, England's first woman to score in a Wembley international and the first to score for England in a women's World Cup, in 2021 she was awarded an MBE for services to football.

What were some of the barriers that you had to overcome as a woman playing football?

"One of the biggest things that stands out for me will always be the comment that I'm a woman, I shouldn't be playing football and that I should be washing up at the sink.

"We'd get comments from the stands about getting married and having kids. That happened quite a lot during my time, especially when I first started. I couldn't get my head around it, I was only 13 and I was playing for Doncaster Belles when I kept hearing people saying this even as I got older and just thought, it's not right."

Have you seen attitudes change?

"They've definitely changed and I think that's because there's much more opportunity now for women and girls to play the game. There's more of a pathway. There are England teams at youth level which weren't there before and it's massively grown, which is where we should be today."

On the 50th anniversary of the women's FA Cup final, how do you feel about being a pioneer in the game?

"There were pioneers before me and we've all paved the way for the next generation. It's great to see the final be played at Wembley again. It's the home of football, it's where everyone wants to play and for the final to be played there, that's how much the women's game has evolved and hopefully that'll be how it is for the rest of my lifetime."

Manchester City and England defender Lucy Bronze is one of the most recognisable names in women's football today. Two-time FA Cup winner with City, Bronze's first FA Cup final came as a 17-year-old with third-tier Sunderland. In 2018 and 2020 she was named BBC Women's Footballer of the Year and in 2019 became the first English footballer to win the Uefa women's player of the year award.

Do you feel you've had to overcome barriers during your time as a professional footballer?

"I think I'm part of the generation in the middle. I remember being in the FA Cup final for Sunderland in 2009 against Arsenal. I was 17 years old at the time and we couldn't even afford the bus to the stadium.

"We had to bag pack in one of the supermarkets to raise money to pay for the bus. We didn't even have a warm-up kit for that game - we had to wear our old away shirt from the season before to warm up in."

How do you think the game has developed?

"In my first final when I was 17 I was a complete nobody - my own team-mates barely knew who I was let alone anyone else. Whereas in the 2017 final, it was off the back of the World Cup where people started to know who I was. The women's game was changing in England, the England international team was changing and that was a really big deal, especially with it being played at Wembley."

Did you ever think you'd be walking out at Wembley in an FA Cup final?

"I never thought we'd be playing at Wembley. It's a stadium with so much history, it's so well known and so many people have a connection to it. It's amazing to get to play there."

What more needs to be done to change attitudes towards women's football?

"It's easy to say that the growth has been good for the sport, because it has. I don't think any other sport has grown at the rate of women's football but that's not to say that we're where we need to be.

"We need to grow, we need to get more girls playing and playing to a higher level. We need more clubs to invest so it's not just a small group of clubs that people talk about winning things and we want to see a bit more competition. I think we need to grow the fanbases of women's clubs too. Personally, I think there is room for improvement in every single area.

"I've experienced both ends of the scale and I love it. It makes me appreciate things more but it also makes me want to fight and keep going and not settle."

How much further on could the game be without the ban all those years ago?

"When you look back at pictures before the ban came in, stadiums are more full then than they are now. It blows my mind to see stadiums that full watching women's football, but it just shows that at one time it was at that same level as the men's game.

"We're chasing after that ban because during that time men's football was constantly growing and women's wasn't. It's a tough task but everyone involved is pushing for it to get back to where it was."

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Map of Hutterite colonies shows religion and evolution – Big Think

Posted: November 28, 2021 at 10:29 pm

Hutterite women and child from the Milford colony in Alberta. The size of the polka dots on the headscarf indicates which specific group the women belong to. (Credit: Eye Ubiquitous / Universal Images Group via Getty Images)

This is a map about religion and evolution. It shows the geographic distribution of Hutterite settlements across North America, but its also a snapshot of the evolutionary concept of speciation at work.

Speciation is the biological process by which a single population evolves into different species. The typical cause is geographic isolation of a subgroup, which then becomes subject to another set of environmental pressures. In combination with genetic drift, this may result in a population with distinct habits and characteristics, which is no longer able to interbreed with the original species in other words, a new species.

Darwin got a first inkling of the concept when he observed how finches and tortoises in the Galpagos archipelago differed from island to island. The archipelago continues to fascinate evolutionary biologists. In 2017, scientists caught a population of Galpagos finches in the act of becoming a separate species the first time speciation has been observed live.

It is tempting to recognize the rudiments of speciation in processes outside of biology for example, the fractious world of religious ideas. Just look at the assorted family trees, showing the diversification over time of biological species and theological concepts. Clearly, something similar is going on here. Except that in religious speciation, geographic isolation typically is the effect, not the cause, which usually is some doctrinal difference of opinion on whether to make the sign of the cross with two fingers or three, for instance.

This map of Hutterite colonies is an interesting snapshot of such religious speciation at work. With a total population of around 50,000, Hutterites are the smallest of the three major branches of the Anabaptist movement. Mennonites are the main branch (pop. 1.5 million worldwide), but everybody knows the Amish around 360,000, mainly in the U.S. set apart by their typical dress and their blanket rejection of modern technology.

Anabaptism emerged as one of the more radical Protestant strains of the 16th-century Reformation of the church. It rejects child baptism, believing instead that candidates should be able to make a free and conscious choice for Christ as adults hence the name for the movement, which in Greek means to baptize again.

Each of the three main groups is named after the religious leader that established it as a separate movement.

Menno Simons (1496-1561) was a Catholic priest from Friesland (northern Netherlands) who became an Anabaptist preacher and leader, instrumental in consolidating and institutionalizing the new faith. His followers were known as Mennonites.

Jakob Amann (1644- c. 1720) was a Swiss Mennonite elder who wanted to preserve what he saw as biblical discipline within the church. In the 1690s, this led to a schism. Amanns followers were called the Amish.

Jakob Hutter (c. 1500-1536) was a hatmaker from Tyrol and an Anabaptist reformer. He led his followers, later called Hutterites, to Moravia, where they adopted the early Christian practice of communal ownership, in addition to traditional Anabaptist practices such as nonviolence and adult baptism.

Anabaptists were considered so radical that they were persecuted by Catholics and mainstream Protestants alike in Western Europe, which is why so many fled, first to Eastern Europe, then eventually to North America.

Many Hutterites in North America can trace their origin to Hutterdorf, a Hutterite colony in Ukraine and the source of a very successful back to basics campaign. This is because, by the mid-19th century, most Hutterites no longer lived in strict community of goods.

In 1859, Michael Waldner was one of the leaders of a congregation in Hutterdorf that reintroduced the practice. The group became known as the Schmiedeleut (people of the blacksmith), after Waldners profession. The next year, another group did the same on the other side of town. Their leader was Darius Walter, and they became the Dariusleut. Around 1875, both groups emigrated to South Dakota. Then, arriving in South Dakota a few years later, a third group of Ukrainian Hutterites under the leadership of teacher Jakob Wipf became known as the Lehrerleut (people of the teacher).

Hutterites established small farming settlements (called colonies), which grew rapidly in number because of the groups high birth rates. In the years immediately after World War I, due to anti-pacifist and anti-German sentiment in the U.S., many Hutterites relocated to Alberta in Canada.

In the following decades, as the animosity wore off, many new Hutterite colonies were established south of the border. As the map shows, Hutterites now live on either side of the U.S.-Canada border. The map also shows how the Hutterite movements three branches cluster together geographically:

As indicated by the map, the Schmiedeleut themselves seem to be in the process of dividing into two distinct groups just like those finches in the Galapagos. In the early 1990s, various disagreements led to a split between a group of Schmiedeleut who were slightly more accommodating of certain innovations (led by Jacob Kleinsasser) and a more conservative group. The Kleinsasser group are known as the Hutterian Brethren, or simply generically as Group 1. They are nicknamed Oilers. The more traditionally minded Group 2 is also known as Committee Hutterites, and nicknamed Gibbs.

Hutterites live in rural communities, speak a German dialect, and dress conservatively, so they are often mistaken for members of the other Anabaptist movements. However, unlike the Amish, they do use electricity and other modern technology. And unlike both the Amish and the Mennonites, they continue to practice communal living.

One way to visually distinguish the various Hutterite subgroups from each other is by the size of the polka dots on the womens scarves. Lehrerleut women have scarves with large dots, Dariusleut scarves have smaller dots, and Schmiedeleut ones have very small dots, or none at all. No wonder they are considered the most progressive of the Hutterites.

Map produced by Alex McPhee, reproduced with kind permission. Mr. McPhee is an independent cartographer based in Grasslands National Park, Canada. Check out his web page and his Twitter.

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Evolutionary change in the construction of the nursery environment when parents are prevented from caring for their young directly – pnas.org

Posted: at 9:53 pm

Significance

Parents can care for offspring directly by giving them food or warmth, for example, or they can help them without direct contact via an extended phenotype by manipulating the nursery environment in which offspring develop. Using an experimental evolution approach, we prevented parents from directly supplying care to their offspring and observed how the extended care phenotype and offspring traits evolved in response. We found that depriving offspring of direct care caused rapid adaptive change in the construction of the nursery environment, which rescued offspring from otherwise poor developmental conditions. Overall, offspring tended to perform better when transplanted to a nursery environment constructed by parents of their own lineage, suggesting that offspring adapt to the evolved extended parental phenotype.

Parental care can be partitioned into traits that involve direct engagement with offspring and traits that are expressed as an extended phenotype and influence the developmental environment, such as constructing a nursery. Here, we use experimental evolution to test whether parents can evolve modifications in nursery construction when they are experimentally prevented from supplying care directly to offspring. We exposed replicate experimental populations of burying beetles (Nicrophorus vespilloides) to different regimes of posthatching care by allowing larvae to develop in the presence (Full Care) or absence of parents (No Care). After only 13 generations of experimental evolution, we found an adaptive evolutionary increase in the pace at which parents in the No Care populations converted a dead body into a carrion nest for larvae. Cross-fostering experiments further revealed that No Care larvae performed better on a carrion nest prepared by No Care parents than did Full Care larvae. We conclude that parents construct the nursery environment in relation to their effectiveness at supplying care directly, after offspring are born. When direct care is prevented entirely, they evolve to make compensatory adjustments to the nursery in which their young will develop. The rapid evolutionary change observed in our experiments suggests there is considerable standing genetic variation for parental care traits in natural burying beetle populationsfor reasons that remain unclear.

Parental care encompasses all parental traits that enhance offspring fitness and that have evolved for this purpose (1). Direct forms of care have been analyzed extensively in previous work. They involve parents engaging directly with their young by defending their offspring from attack, for example, or by brooding them when they are cold or feeding them (2). Yet, parental care can also take the form of an extended phenotype. Before their offspring even exist, parents can manipulate the nursery environment in which their future young will develop by carefully choosing the territory within which the nursery is sited, by constructing a nursery or nest, and by stockpiling it with food for the newly hatched offspring (3). In some species, such as dung beetles, beewolves, skates, and jacky dragons, parents and their offspring never meet again after egg laying. Nevertheless, the extended parental care phenotype in these species endures to influence offspring fitness (47).

Here, we are interested in the evolutionary relationship between the extent of direct care and the extended parental care phenotype and how that, in turn, influences the evolution of offspring traits. Each form of care is understood to generate a fitness benefit for the offspring, usually at some fitness cost to the parent that supplies it (1, 2). Any existing fitness costs limit the supply of care, but the relative benefits derived from each form of care presumably determine the relative level of investment in each of them. If the relative fitness benefits derived from direct care suddenly decline, for example, then we might expect a corresponding adaptive increase in the extended parental care phenotype to compensate for any loss in fitness experienced by the offspring. Previous studies have produced correlational evidence that is consistent with this possibility (e.g., refs. 810). Furthermore, recent work has investigated whether such compensatory changes can be induced via phenotypic plasticity within the lifetime of an individual (11, 12). However, we are unaware of any work that has considered how changes in one form of care cause evolutionary change in other forms of care or how that could causally influence the evolution of offspring traits.

To address this question, we took advantage of the natural variation in parental care found in the burying beetle Nicrophorus vespilloides, which comprises both direct care and an extended parental care phenotype. Burying beetles use small dead vertebrates, such as mice or small birds, to rear their larvae (13, 14). The extended parental care phenotype is expressed when parents transform the carcass into an edible nest. They scissor off the fur or feathers, roll it into a ball, cover the flesh with antimicrobial exudates, and bury it in a shallow grave (1416). Eggs are laid in the soil surrounding the carcass, and when larvae hatch, they crawl to the carcass. Parents assist the offspring in colonizing the carcass by biting small holes in the flesh, which are used by larvae to penetrate the carcass. Parents may stay to supply their offspring with direct care, which involves defending them and feeding them via oral trophallaxis (17). Larvae can also feed themselves and can survive without any posthatching care (17, 18). Approximately 1 wk after hatching, larvae disperse to pupate in the soil. Parental presence during larval development increases larval survival (19), yet the duration of posthatching parental care is highly variable, with a range spanning from no posthatching care at all to the whole period of larval development (13, 2022). Thus, the extent of direct parental care experienced by burying beetle larvae in early life is highly variable.

We used experimental evolution to investigate how a change in the supply of direct parental care affects the evolution of an extended parental care phenotype: that is, construction of the nursery environment through the conversion of the carcass into an edible nest. Both types of care have been shown to improve offspring survival (19, 23) and incur life span costs in burying beetles (2426). We established experimental populations that evolved either with Full Care (FC; i.e., direct care plus extended parental care) or No Care (NC; i.e., only extended parental care but no direct contact with parents).

We have reported some of the outcomes of this experimental evolution work previously. We found that preventing direct posthatching care in experimental NC populations for generation after generation initially resulted in lower breeding success and larval survival (27). However, this was followed by a rapid increase in fitness in subsequent generations so that FC and NC populations had similar measures of fitness by generation 13 (27). We have also investigated how the evolution of larval traits, such as their morphology (22) and social interactions on the carcass (18, 28), contributed to this recovery in fitness in the NC populations. Here, we focus more on the evolution of parental traits in the NC populations by examining how changes in the parental extended phenotype of carcass preparation promote and interact with offspring fitness in the absence of direct care.

To disentangle the fitness consequences of changes in the parental traits from changes in larval traits, we cross-fostered larvae within and between FC and NC after multiple generations of experimental evolution (18, 22, 27, 29). By measuring correlates of larval fitness in the absence of direct care, we further determined whether evolved change in the extended parental care phenotype compensated for the loss of direct parental care (in our laboratory environment). Our results demonstrate that there is rapid adaptive evolution of the extended parental care phenotype when parents are prevented from interacting with their offspring and that offspring adapt rapidly to this changed nursery environment.

The experimental populations were founded from four wild populations of N. vespilloides collected in Cambridgeshire, United Kingdom (Byrons Pool, Gamlingay Woods, Overhall Grove, and Waresley Woods) in the summer of 2014. Further details of these wild populations are given in refs. 22 and 28. The populations were interbred to create a genetically diverse stock laboratory population from which the experimentally evolving populations could be derived. This allowed us to avoid potential confounding effects of inbreeding depression, which is masked by direct parental care (30). Two selective regimes were established, one with full posthatching parental care, FC, and the other without any posthatching parental care, NC. Two independent replicates (hereafter referred to as block 1 and block 2) of the FC and NC regimes were maintained, with block 2 breeding a week after block 1.

At each generation, males and females were paired within each experimental population, excluding sibling and cousin pairings. Each pair was placed in a breeding box (17 12 6 cm) half filled with moistened compost, and a small, thawed dead mouse (8 to 14 g, obtained frozen from LiveFoods Direct) was placed on top of the soil. Breeding boxes were kept in dark cabinets to simulate natural underground conditions. In the FC populations, a minimum of 30 pairs of unrelated beetles were bred at each generation. Parents were allowed to remain in the box throughout larval development and so, were able to provide posthatching care. In the NC populations, we set up a minimum of 50 pairs each generation to compensate for the increased number of failed broods (27), and both parents were removed from the breeding box 53 h after pairing, before larvae started hatching. This allowed parents sufficient time to convert the mouse body into a carrion nest and for females to lay eggs in the surrounding soil, but it deprived larvae of any posthatching care (31, 32).

Eight days after pairing, dispersing larvae were placed into individual cells (2 2 2 cm) in an eclosion box (10 10 2 cm), with one brood per eclosion box. They were covered with moistened peat and left undisturbed to pupate. Newly eclosed adults were then removed and housed individually until breeding a minimum of 17 d after eclosion. All adult beetles were fed raw ground beef twice a week.

In generation 11, we created a third type of experimental population (from here on, known as Nmstanding for No Care, maternal) from each of the two replicates of the NC population. The two replicate Nm populations were bred in parallel with each of the two FC and NC replicates. The Nm population passed through one generation of full posthatching parental care to eliminate potential maternal and/or other environmental effects in the NC population. To create each Nm population, we thus followed the same protocol as for the FC populations and bred an additional 30 pairs of unrelated beetles from each NC replicate population. The Nm populations were discontinued after the experimental analysis described below.

In generation 13 of the FC and NC populations (corresponding to generation 1 of Nm), we collected newly eclosed adults from all three populations. We housed all beetles individually in plastic boxes (12 8 2 cm) until pairing for the experiment described below.

We randomly selected males and females from each population to set up 450 pairs (75 pairs per population per block) and placed them in a breeding box (17 12 6 cm) with a 10- to 12-g thawed mouse carcass. After 53 h, before any larvae hatched, we removed the parents and measured the parents pronotum width, which is standardly used as a proxy for adult body size. As a measure of reproductive investment, we counted the number of eggs visible on the bottom of the breeding boxes, which is a noninvasive accurate method for deducing clutch size (33). Eggs were left in situ, in the soil in the breeding boxes, to hatch into larvae.

Before the eggs hatched, carcasses were swapped between breeding boxes to create a fully factorial 3 3 experimental design, such that larvae from each experimental population were allowed to develop on carcasses prepared by adults either from the larvaes own natal population or from the other two experimental populations (SI Appendix, Fig. S1). Larvae were, therefore, always unrelated to the adults that had prepared the carcass whereupon they developed. Furthermore, since the adults were removed, no broods received any direct parental care. After 8 d, we counted and weighed surviving larvae to derive correlates of offspring fitness on the different carcasses.

Using this experimental design, we were able to separate contributions to larval fitness due to the extended parental care phenotype from any contributions to larval fitness made by larval traits (22, 28).

Burying beetles typically roll the denuded flesh of the carcass into a ball to create a nest for their larvae. The first measure we made of the nursery environment was the roundness of the carrion nest. Although no correlation between development on a rounder carcass and larval mass at dispersal has yet been found, rounder carcasses correlate negatively with paternal life span, indicating a cost to nest construction (25), and are also less hospitable to rival blowfly larvae (34). Burying beetles also smear costly antimicrobial exudates on the carcass surface (35, 36), which improves larval survival (23). A rounder carcass would minimize the surface area to volume ratio, thus potentially reducing defense costs and the possibility of carcass desiccation. This could be particularly advantageous to offspring in the absence of posthatching parental care.

We measured carcass roundness 53 h after pairing adults and presenting them with a dead mouse, following methods described in ref. 25. Briefly, we took two photographs of each carcass from perpendicular positions 30 cm away with identical cameras and settings (Fujifilm av200). When visible, we digitally removed the mouses tail from all photos with GIMP (v. 2.8.16; The GIMP Development Team; https://www.gimp.org/), as the tail strongly influences roundness estimates. We then calculated carcass roundness with a custom-written script (SI Appendix) in ImageJ (1.49v, Wayne Rasband; NIH; https://imagej.nih.gov/ij). By applying this process to a ping-pong ball, we established that a perfect sphere has a roundness score of 0.9. We, therefore, adjusted subsequent measures of roundness by dividing by 0.9, meaning that a value of 1.0 intuitively equaled a sphere.

Second, we recorded whether small holes were visible on the surface of the carcass at 53 h after pairing. There is individual variation in the timing of these holes. In previous work, we found that only 26% of wild-caught N. vespilloides, breeding in laboratory conditions, made a visible hole in the surface of the carcass before larval hatching (22). In the absence of posthatching care, the presence of a hole in the carcass is critical for larval survival (19). We hypothesized that NC populations might make these entrance holes earlier than FC populations. Some of these data, namely the presence of a hole in the carcass in FC and NC populations at 53 h after pairing, were published in ref. 22. Here, we present a different analysis of these data, including examining the fitness consequences of the presence of a hole for different experimental populations, as well as data regarding Nm populations.

Third, we measured antimicrobial activity in male and female anal exudates at 53 h after pairing. Adults deposit these exudates all over the carrion nest during carcass preparation. The presence of these exudates improves larval survival (23) and affects the composition of bacterial communities growing on prepared carcasses (16, 37). Lytic activity is heritable in N. vespilloides (38), with significant positive maternal effects, and therefore, it is feasible that selection could act to increase it in NC and Nm populations, where parents cannot maintain the carcass after larval hatching.

Burying beetles readily produce a redbrown liquid when gently tapped on the back of the abdomen. However, in some cases, individuals did not produce exudates. The total numbers of successfully sampled individuals were 398 males (131 FC, 130 NC, and 137 Nm) and 401 females (133 FC, 129 NC, and 139 Nm). We collected exudates with Pasteur pipettes, stored them in 1.5-mL Eppendorf tubes, and kept them frozen at 20C until further analysis. Lytic activity was measured in an automated plate reader (Biotek ELx808) by a microplate turbidity assay that quantifies the degradation rate of bacterial cell walls (adapted from ref. 23). Briefly, we diluted exudates 25-fold in potassium phosphate buffer (pH 6.4, 0.02 M). We added 10 L of diluted exudates per well to 96-well microtiter plates filled with 100 L per well of a 1.3-mg mL1 suspension of lyophilized Micrococcus luteus (Sigma-Aldrich) in potassium phosphate buffer. Samples were initially incubated in the plate reader at 25C for 30 s with continuous shaking. Absorbance at 450 nm was measured every 10 min for 60 min, with continuous shaking for 10 min at 25C between measurements. We calculated lytic activity as the percentage change in absorbance relative to control wells, with 10 L of potassium phosphate buffer and 100 L of M. luteus suspension. We report here results for change in absorbance at 450 nm after 60 min.

All analyses were performed using the statistical program R version 4.1.1 (39). Mixed effects models were performed with the package lme4 version 1.1-27.1 (40). Seventeen breeding pairs were removed from the analysis either because a parent died or was damaged before adult removal (FC = 2, NC = 3, Nm = 3) or because no eggs were observed in the box upon adult removal (FC = 2, NC = 5, Nm = 2). Therefore, 433 breeding pairs were included in the analysis (FC = 146, NC = 142, Nm = 145).

A further seven breeding pairs were removed from the analysis of carcass preparation traits because we could not measure pronotum width for both parents, and body size is an important explanatory variable for carcass roundness and lytic activity. Therefore, 426 breeding pairs were included in the analysis of carcass preparation traits (FC = 143, NC = 140, Nm = 143). To analyze the presence of a hole in the carcass, we initially fitted a generalized linear mixed model (GLMM) with a binomial distribution and a population per block random effect to account for variation between the independent replicates due to founder effects and asynchronous maintenance. No variance was explained by the random effect, and therefore, we used a generalized linear model (GLM) with a binomial distribution to analyze the presence of a hole. Carcass roundness and lytic activity were analyzed with linear mixed models. Lytic activity was log transformed to ensure that model residuals met the assumptions of normality for regression. In these and subsequent models with mixed effects, we included a population per block random effect.

To compare brood performance between experimental populations, we analyzed breeding success (i.e., survival of at least one larva to dispersal), brood size, and brood mass. Offspring mass is a known correlate of fitness in burying beetles (19, 26). Brood success was initially analyzed with a binomial GLMM. Again, the random effect population per block did not explain any variance, and we, therefore, used a binomial GLM to model brood success. Brood size and brood mass were analyzed with linear mixed models (LMMs). We removed brood failures (i.e., no larvae survived to dispersal) from the analysis of brood size and brood mass. To evaluate the relative contribution of parental and larval traits on larval survival to dispersal, we fitted a linear regression model on brood size, with the number of eggs as a covariate. We then used regression residuals as a measure of offspring survival from egg laying to larval dispersal. We included the residuals as the response variable in a LMM with population, carcass type (i.e., population that prepared the carcass), presence of a hole, female size, and carcass roundness as explanatory factors/covariates. Post hoc comparisons were performed with the package emmeans version 1.6.3 in R (41).

Model selection in all analyses was performed by comparing nested models with the Akaike Information Criterion (AIC) and ANOVA (42). Here, we report the minimal adequate models, where nonsignificant terms (P > 0.05, as reported by ANOVA of nested models) were dropped when this resulted in a decrease of AIC by two units. The significance of interaction terms was determined by performing ANOVAs on nested models (with and without the interaction terms). To validate models, we inspected residuals of all minimal adequate models. Female and male sizes, as well as carcass mass, were included as covariates in all the initial models. For LMMs, we used Satterthwaites approximation to calculate degrees of freedom and P values with the package lmerTest version 3.1-3 (43). To check whether approximating degrees of freedom alters statistical results, we ran all minimal adequate models with and without random effects. All models maintained the same qualitative results, and most variables showed P values and effect sizes of the same magnitude whether we included or excluded random effects. Only in the models for brood mass and offspring mortality did we find lower P values when the random effect population per block was excluded (reported in Results).

We found significant differences in the extended parental care phenotype across the different populations we sampled. Carcasses prepared by NC and Nm beetles were twice as likely to have a hole as carcasses prepared by FC beetles (Table 1). At the time of the removal of parents (53 h after pairing), the percentages of carcasses with a visible hole were 30% in FC carcasses, 61% in NC carcasses, and 59% in Nm carcasses.

Summary of minimal adequate models of carcass preparation traits in FC, NC, and Nm populations

Carcasses prepared by NC and Nm beetles were also rounder (Table 1) (overall effect of population: 2 = 25.80, P = 2.5 106). There was a complex, significant interaction between population type and both female and male body sizes (Fig. 1 and Table 1). In FC lines, carcass roundness was positively associated with both male and female sizes. In NC and Nm lines, the slope of the relationship between carcass roundness and body size was significantly shallower than in FC lines, with beetles across a range of sizes producing similarly well-rounded carcasses.

The predicted partial effects of the experimental population of origin and female (Left) and male (Right) pronotum width on carcass roundness. Each data point represents a carcass prepared by a pair of beetles (n = 426). Lines represent adjusted carcass roundness values predicted by a linear mixed model.

Lytic activity in anal exudates at 53 h was similar across the three populations, for both males and females (Table 1). The main predictors of lytic activity, for both sexes, were the individuals size and the lytic activity of their breeding partner. Larger individuals produced higher lytic activity, and individuals paired with beetles that had higher lytic activity also showed higher lytic activity themselves. Lytic activity did not differ significantly between males and females (ANOVA: F1,722 = 1.245, P = 0.265).

Across all populations, the presence of a hole on the surface of the carcass was the best predictor of breeding success (i.e., whether at least one larva would survive to dispersal; absence of hole: 79% successful [169 of 214]; presence of hole: 96% successful [211 of 219]) (Table 2). The number of successful broods did not differ across the three experimental populations, nor were broods more likely to be successful on FC-, NC-, or Nm-prepared carcasses after the effect of hole presence was controlled for statistically by including it in the model (Table 2).

Summary of minimal adequate models explaining differences in the success, size, and mass of broods from FC, NC, and Nm populations on carcasses prepared by FC, NC, and Nm parents

The number of eggs observed at the bottom of breeding boxes, a proxy measure for clutch size, was significantly larger in the FC populations than in the NC and Nm populations (on average, FC clutches had 4.21 and 4.15 more eggs than NC and Nm clutches, respectively) (SI Appendix, Fig. S2 and Table S1). Brood size varied across populations and also depended on which populations had prepared the carcasses that the larvae developed upon (Table 2). Despite FC females laying larger clutches, FC broods were not larger overall at dispersal. The presence of a hole was, of all the factors considered, the one with the largest positive effect on the number of larvae that survived to dispersal (Fig. 2 and Table 2) across all three populations. There was, however, a significant interaction between population and the presence/absence of a hole. In carcasses where no hole could be seen, FC broods were significantly smaller than both NC and Nm broods (Fig. 2 and SI Appendix, Table S2). There was no effect of carcass roundness on brood size (ANOVA between nested models with and without carcass roundness as a fixed effect: 2 = 2.16, P = 0.14).

Brood size and brood mass of larvae at dispersal from FC, NC, and Nm broods (red, blue, and green box plots, respectively) developing on carcasses with and without a hole. Sample sizes: FC = 125, NC = 127, and Nm = 128. Box plots depict the first quartile, median, and third quartile. Whiskers on the box plots range from the samples lowest to highest value within 1.5 interquartile range. Points depict sample outliers. Significant differences between populations are indicated with asterisks (post hoc analyses with emmeans using the Tukey method; *P < 0.05; **P < 0.01; ***P < 0.001) (SI Appendix, Tables S2 and S4).

A significant interaction was also found between the broods population of origin and the population of origin of the beetles that prepared the carcass (population carcass) (Fig. 3 and Table 2). There was a tendency for broods to be larger at dispersal on carcasses prepared by parents of their own population of origin (Fig. 3 and SI Appendix, Table S3). FC larvae performed significantly better on carcasses prepared by FC parents than on carcasses prepared by NC and Nm parents (post hoc tests and 95% CIs) (SI Appendix, Table S3). NC and Nm larvae showed a nonsignificant tendency to perform better on carcasses prepared by NC and Nm parents, respectively (SI Appendix, Table S3).

Brood size and brood mass of larvae at dispersal from FC, NC, and Nm broods developing on carcasses prepared by FC, NC, or Nm parents (red, blue, and green box plots, respectively). Sample sizes: FC = 125, NC = 127, and Nm = 128. Box plots depict the first quartile, median, and third quartile. Whiskers on the box plots range from the samples lowest to highest value within 1.5 interquartile range. Points depict sample outliers. Significant differences between populations are indicated with asterisks (post hoc analyses with emmeans using the Tukey method; *P < 0.05; **P < 0.01; ***P < 0.001) (SI Appendix, Tables S3 and S5).

Brood mass varied across populations and carcasses prepared by different populations, in a similar way to brood size, with presence of a hole having the largest positive effect (Fig. 2 and Table 2). There was a significant interaction between the presence of a hole and the population of origin, with FC broods having significantly smaller mass than NC and Nm broods in the absence of a hole (post hoc tests and 95% CIs) (SI Appendix, Table S4). Again, there was a tendency for brood mass to be highest in broods reared on carcasses from their own parental population (Fig. 3) (post hoc tests and 95% CIs) (SI Appendix, Table S5). The minimal adequate model for brood mass also included a marginally significant interaction between the presence of a hole and the carcass of origin; post hoc tests suggest that the presence of a hole leads to broods attaining a greater mass at dispersal when reared on NC and Nm carcasses than on FC carcasses (post hoc tests and 95% CIs) (SI Appendix, Table S6). When running the minimal adequate model without random effects, the P values for population were lower than in the mixed model with Satterthwaites approximated degrees of freedom (LMM: P = 0.01 and P = 0.02 for NC and Nm, respectively; LM: P = 0.006 and P = 0.007 for NC and Nm, respectively).

To deduce the extent of larval mortality, we applied a best-fit regression on the entire dataset of brood size on clutch size (F1,378 = 148.5, P < 0.001, R2 = 0.28) (SI Appendix, Fig. S3 and Table S7) and used the residuals as a response variable in a GLMM (Table 3).

Generalized linear mixed model of offspring mortality between egg laying and larval dispersal for FC, NC, and Nm populations reared on FC, NC, and Nm carcasses

Overall, there was a population effect on larval mortality, with FC populations showing higher mortality than NC or Nm populations, thus potentially explaining why FC broods were not on average larger, despite having larger clutch sizes. Again, we found a significant interaction between the larval population of origin and the presence of a hole in the carcass. On carcasses without a hole, FC populations showed greater offspring mortality between egg laying and larval dispersal than NC and Nm populations (Table 3 and SI Appendix, Table S8). An interaction between the larval population of origin and the carcass population of origin (population carcass) (Table 3) was retained in the minimal adequate model, despite being only marginally significant, because removing it did not decrease AIC. Post hoc tests revealed that FC offspring were most likely to die between egg laying and larval dispersal across all types of carcasses (Tables 4 and 5). There was again a tendency for larvae to perform better on carcasses prepared by parents of their own parental population (Fig. 4 and Tables 4 and 5). Offspring were equally likely to survive on FC-prepared carcasses, regardless of their population of origin. However, FC populations were significantly more likely to die between egg laying and larval dispersal than NC and Nm populations when they developed on NC-prepared carcasses. On Nm-prepared carcasses, FC populations differed significantly only from Nm populations in the likelihood that larvae would die between egg laying and larval dispersal (Tables 4 and 5). When running the minimal adequate model without random effects, the P values for population were lower than in the mixed model with approximated degrees of freedom (LMM: P = 0.045 and P = 0.009 for NC and Nm, respectively; LM: P = 0.009 and P = 0.0002 for NC and Nm, respectively).

Variation in offspring mortality between egg laying and larval dispersal due to the interaction between population and carcass type: least-square means (LS) and 95% confidence levels (CL)

Number of larvae that survived from egg laying to larval dispersal in relation to clutch size (estimated from the number of eggs on the bottom of the breeding box). Figures show the FC, NC, and Nm populations (red, blue, and green circles and box plots, respectively) developing on FC- (Top), NC- (Middle), or Nm-prepared carcasses (Bottom). Each data point in Right represents a different brood. Sample sizes: FC = 125, NC = 127, and Nm = 128. A regression model for brood size was fit to all the data (black lines in Left; R2 = 0.28) (SI Appendix, Fig. S2 and Table S7), and the residuals from those lines (Right) were used as response variables in linear mixed models (i.e., this was our measure of relative offspring mortality). Box plots depict the first quartile, median, and third quartile. Whiskers on the box plots range from the samples lowest to highest value within 1.5 interquartile range. Points depict sample outliers. Significant differences between populations (indicated with an asterisk; *P < 0.05; **P < 0.01; ***P < 0.001) were found in NC- and Nm-prepared carcasses but not FC-prepared carcasses. Post hoc analyses were performed with emmeans using the Tukey method (Tables 4 and 5).

Variation in offspring mortality between egg laying and larval dispersal due to the interaction between population and carcass type: estimated differences between populations within carcass types

We found that when experimental populations were prevented from supplying care directly to their young for several generations, individuals adapted by modifying the way they constructed the nursery in which their offspring developed. Even though they could no longer meet their larvae, parents were still able to enhance their offsprings fitness through this modified form of indirect care. Importantly, Nm parents, which were reared with posthatching care, showed the same response as NC parents, which had not experienced parental care themselves. Hence, the differences we observed between FC and NC parents were not a result of phenotypic plasticity in response to their own early life environment but represented evolved divergence in this trait.

We measured change in several facets of nursery construction, and the results offer preliminary insights into the modular nature of carcass preparation and the extent to which the different elements are subject to different selection pressures and are genetically uncorrelated. The prehatching care trait with the strongest fitness consequences to the brood was the timing of insertion of a hole in the carcass. When we tested the adaptive value of the new nursery environment, in a previous study, by inserting a hole in the carcass ourselves and forcing larvae to develop with no posthatching care, we observed increased brood survival, brood size, and mass and more surviving larvae for a given clutch sizeregardless of the population from which larvae were drawn (22). By biting this hole in the carcass before they were removed, parents probably enabled larvae to enter the carcass, to feed upon it, and to take up residence there (19, 22), even in the parents absence. Furthermore, the fitness gained by the evolution of this extended parental care phenotype helped to compensate for the fitness lost from the experimental removal of direct care; by generation 13, parents from the NC and FC populations produced similar numbers of larvae per gram of carrion, as shown in ref. 27.

If the presence of a hole has such strong fitness consequences when parents are absent, why did roughly 70% of FC and 40% of NC and Nm parents still not bite a hole before larval hatching? The results for FC parents are unsurprising because they typically remain with their brood under their selective regime. Unlike NC parents, FC parents had the opportunity to make entrance holes in the carcass after their larvae hatched, as they crawled through the soil to the carcass. The proportion of FC parents making a hole in the carcass before larval hatching is close to the proportion found in wild-caught beetles breeding in the laboratory (26%) (22). As for the NC parents, it is possible that the accompanying evolution of morphological and behavioral larval traits (22, 28) helped to decrease selective pressure on accelerated hole biting by parents, or it may be that this trait was still under selection and spread further through the NC population in subsequent generations. Either possibility can explain why we did not observe all NC parents biting a hole prior to their removal by the 13th generation of experimental evolution.

Focusing on carcass roundness, we found that carcasses prepared by parents from the NC populations were rounder by the time we removed parents. In FC populations, by contrast, only larger parents were able to produce rounder carrion nests. However, just as in a previous study (25), we found no evidence that carcass roundness affected brood performance directly. Why, then, did we observe a change in this trait?

One possibility is that the shape of the carcass itself is not the trait under selection but that it changes as a by-product of selection on the pace of carcass preparation. Our experimental protocol placed NC beetles under selection to complete carcass preparation within 53 h to ensure they had bitten a hole for their larvae before they were removed and perhaps thereby incidentally favored beetles that had prepared rounder nests. FC parents, by contrast, are not under selection to complete this task as quickly. Furthermore, converting the dead body into a carrion nest is likely to be energetically costly because it involves rolling the corpse around and pushing it against the soil. This might explain why smaller beetles, with fewer energy reserves, engaged in carcass preparation less vigorously and, consequently, produced less rounded carcasses in the same timeframe in the FC lines (and also as observed in ref. 25). The strong fitness consequences of the presence of a hole contrast with the seeming lack of fitness consequences of carcass roundness, and suggest that both traits belong to the same behavioral module. Strong selection on the timing of the hole could, therefore, have dragged the rest of the behavioral module with it as a correlated response, resulting in rounder carcasses. It would be interesting to test whether this speculation is correct or whether carcass rolling is in fact a distinct behavioral module.

We found no difference between our experimental populations in a third trait associated with carcass preparationthe lytic activity of the anal exudateswhich is again consistent with the suggestion that carcass preparation comprises distinct behavioral modules. Our results imply that the different modules evolve under different selection pressures and are not strongly genetically correlated, much as has been found in the extended parental phenotypes of bees (44) and mice (45). This could explain why some elements of carcass preparation have evolved in response to the elimination of posthatching care, while others have not.

An additional explanation for the lack of differences in lytic activity between experimental populations, which does not exclude the suggestion of modularity, is that the NC larvae have evolved stronger lytic activity in their antimicrobial exudates (4648). A further explanation is that the lytic activity of the anal exudates is a highly plastic trait (24), and its plasticity prevented any evolutionary change. All these interpretations remain to be tested in future work.

We detected evolved change in the way that parents constructed the nursery environment after only 13 generations of experimental evolution. The most likely explanation is that we selected on existing standing genetic variation in the mix of wild populations that founded the experimental populations. By mixing different wild populations, we intentionally increased the genetic variation in our experimental laboratory populations. We must, therefore, be cautious not to directly extrapolate our results to wild populations and risk overestimating their ability to respond to similar selective pressures. Nevertheless, the question remains of how so much genetic variation is able to persist naturally over relatively small geographic distances. One possibility is that there is genetic differentiation between the different wild populations that we sampled originally (49). We have recently found that even nearby populations can be divergently adapted to the specific local conditions within their woodland (50). It is also possible that there is considerable genetic variation in parental care within populations, which is maintained by variation in key environmental conditions governing each breeding attempt, such as the species of the dead animal, the density of carrion, and the extent of competition within and among species for the opportunity to breed upon it (51).

The extended parental care phenotype was not the only trait to diversify across populations. NC populations had smaller clutch sizes than FC populations. Typically, burying beetles lay more eggs than can successfully be reared on a carcass and then, cull the hatchlings to adjust brood size to the existing resources (52). It is possible that NC populations are selected to lay fewer eggs that more accurately match the existing resources because they are removed before they can cull any hatchlings. This hypothesis is currently under investigation in our laboratory.

Furthermore, the cross-fostering experiment suggests that larvae were divergently and locally adapted to develop under NC vs. FC. FC larvae not only suffered higher mortality in the absence of posthatching care than NC larvae, they were also more likely than NC larvae to die when developing in carcasses prepared by parents from other lineages. NC larvae also tended to perform better in carcasses prepared by their own lineage, although this was not statistically significantperhaps because our experiment did not have enough statistical power to detect very small effects, despite a large sample size. Why is the effect size larger for FC larvae than for NC larvae? One possibility is that NC larvae mostly rely on their own traits to secure access to the carcass, whereas FC larvae are more locally adapted to a particular extended parental phenotype. We know from previous work that NC larvae have evolved relatively larger mandibles (22), a propensity to hatch more synchronously (53), and an inclination to behave more cooperatively toward siblings (28). However, further work is needed to fully understand the extent and mechanisms of local adaptation to the extended parental phenotype in FC vs. NC larvae.

How does carcass preparation in burying beetles compare with other forms of extended parental phenotypes? Despite the widespread occurrence of extended parental phenotypes across taxa, the fitness consequences are likely to depend on the extent of parental presence during offspring development. In species with no direct parental care, such as the mass provisioning beewolves and dung beetles, it is likely that the extended parental phenotype has strong fitness consequences. In dung beetles, the size of the brood mass (the ball of dung with developing larvae) that the mother stockpiles determines many offspring traits (4); in beewolves, the antimicrobial secretions applied by the parent protect the larvae against pathogens during development (5). In other species, direct parental care may mask the adaptive value of extended parental phenotypes. For example, egg size can be considered an extended parental phenotype, yet evidence that avian egg size correlates with offspring survival is inconclusive (reviewed in ref. 54). As pointed out in ref. 54, large eggs can benefit offspring survival in harsh environments, but in good conditions, parental provisioning masks any effect of egg size on survival. Interestingly, the same occurs in burying beetles, where effects of egg size on survival are masked by posthatching care (55). Our findings regarding the timing of carcass preparation parallel results obtained previously for egg size in burying beetles and birds. It suggests that the stability of the environment (whether social or abiotic) affects the evolutionary feedbacks between different forms of parental care. In future work, it will be important to test whether these evolutionary feedbacks exist in natural populations, where the environment is more complex and unpredictable than in the laboratory.

Parental care comprises a suite of traits that are integrated to promote offspring fitness (56). Previous work has emphasized that the wider physical environment and the social environment within the family are each sources of selection on the form and function of parental behavior (5659). Here, we have shown that acts of care are themselves a source of selection on other types of parental traits. Preventing any form of direct care causes parents to evolve modifications in the way in which they construct the nursery environment. Furthermore, coadapted traits in parents and offspring evolved rapidly when we experimentally eliminated direct forms of posthatching care, suggesting that these traits have a high degree of genetic variability in natural populations.

All data, code for image analysis and statistical analysis are available in the SI Appendix.

We thank S. Aspinall and C. Swannack for maintenance of beetle populations and laboratory assistance, J. Troscianko for the ImageJ script, A. Sutter and E. Postma for statistical advice, and B. Kuijper for helpful discussions. We also thank D. Mock and an anonymous reviewer for comments, which greatly improved this manuscript. This work was funded by European Research Council Consolidators Grant 310785 Baldwinian_Beetles (to R.M.K.). R.M.K. was also supported by a Wolfson Merit Award from the Royal Society.

Author contributions: A.D., D.R., and R.M.K. designed research; A.D., D.R., A.C.H., and B.J.M.J. performed research; A.D. and D.R. analyzed data; and A.D., D.R., B.J.M.J., and R.M.K. wrote the paper.

The authors declare no competing interest.

This article is a PNAS Direct Submission.

This article contains supporting information online at https://www.pnas.org/lookup/suppl/doi:10.1073/pnas.2102450118/-/DCSupplemental.

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The Fruit of Evolution Anime Episode 9 Release Date and Time, COUNTDOWN, Where to Watch, News and Everything You Need to Know – Epicstream

Posted: at 9:53 pm

The Fruit of Evolution follows Seiichi Hiragi, who was insulted and bullied because of his appearance. He does not have any friends and was often remarked with harsh words by his classmates. One day, he heard a voice, claiming to be God, saying that he and the rest of his classmates should prepare to be transported to another world.

True enough, Seiichi and his classmates were taken to a fantasy world where stats, levels, and skills are needed to survive. Since he has no one, he was summoned to a different area where he found the Fruit of Evolution that changed his life. The series will follow Seiichis journey in this new world and how he became a champion.

Episode nine of The Fruit of Evolution will be released on November 30, 2021. It is entitled, Black Cat Oliga. It will make its premiere in Japan on Monday at 10:00 PM JST. Of course, for international fans, they can expect the series by Tuesday at 2:30 AM PST.

The series can be streamed on Crunchyroll in English subtitles. For the English dubbed version, fans will have to wait for Cruchyrolls official announcement regarding the matter. While waiting, fans can stream the series on site for free as Crunchyroll has a free trial period for non-members. Members or those who are interested to become members must pay $9.99 per month or $79.99 per year to keep their membership.

Those who signed up can get exclusive perks from the site such as ad-free shows, access to the sites libraries, and simulcasts after Japan. Fans should take note that simulcasts are not available for non-members which means that they must wait for a few days before they can stream The Fruit of Evolution on Crunchyroll, unlike members who will just wait an hour after the episode was released in Japan.

It has been confirmed that the series will have 12 episodes that will run until December 2021, unless there is a delay in production or airing. Those who are curious as to what will happen to Seiichi can stay tuned to find out.

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The evolution of cheating in viruses – Nature.com

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Pendant from 41,500 years ago may have uncovered a ‘step in evolution’ – CNET

Posted: at 9:45 pm

The pendant, seen here in slightly different views, would've been about 4.5 centimeters (1.8 inches) long and 1.5 centimeters (just over half an inch) wide, with a thickness between 0.36 and 0.39 centimeter (1.4 and 1.5 inches).

In 2010, scientists unearthed an ivory pendant from an abandoned Polish cave. Punctured with patterns reminiscent of moon cycles and mathematics, the artifact's origins eluded archaeologists -- until now. An international team of researchers just declared the relic to be 41,500 years old.

That makes it the earliest piece of ornate jewelry ever found in Eurasia, and a wonderful reminder that art is timeless. Images and details of the discovery were published Thursday in the journal Scientific Reports.

But beyond its aesthetic value, this ancient pendant also marks the first proof of post-Neanderthal civilization in the Polish region, enriching our textbooks with new data on the movement of early human populations.

"It was not expected to have any evidence of early Homo sapiens in this cave, because it was sort of seen as a Neanderthal cave," said Sahra Talamo, lead author of the study. "This sort of shows the next step in evolution."

The Polish cave where the pendant was excavated.

The pendant's modern story began 11 years ago, when researchers excavated two fragments of it in Poland's Stajnia Cave. Each piece is made of mammoth bone and has unique looping designs etched with a sequence of dots. There are two holes on the adornment, which presumably were used to attach a string of sorts to make a necklace.

"When I saw it, I was shocked," Talamo said of her first encounter with the item.

The confusion came from its location in a stratigraphic layer of the cave attributed to Neanderthals. Tools, and even teeth, of the human ancestors had already been discovered there. But according to Talamo, it was peculiar to connect the jewelry to the early humans, because of the dotted decorations. Such artwork is typical of Homo sapiens, who lived after the Neanderthal era, and are thought to have engaged in more-complicated creative endeavors.

Neanderthals had their own jewelry, but it wasn't nearly as elaborate as what's on the pendant. Armed with a ton of questions, Talamo decided to investigate.

Talamo is an expert in the field of radiocarbon dating, a method that uses an isotope of carbon to determine the age of organic material. Lo and behold, the pendant didn't originate with the Neanderthals. It likely just migrated to a lower layer of the cave even though it was made by later generations from the Early Upper Paleolithic era.

Given its rarity, she also suggests these relics were probably not commonplace items for the Homo sapiens. Perhaps it could've been a status symbol of sorts?

Interestingly, the pendant's remains were found alongside an ancient awl, or small pointed tool used for piercing holes. Coincidence? Well, probably. The team realized the awl dates back 500 years prior to the pendant's creation, and either way, was likely too "soft" to make the dotted markings in bone. That finding is also a result of Talamo's radiocarbon dating technique.

Typically, radiocarbon dating is seen as invasive because it requires breaking off a physical piece of the object being studied. As Talamo puts it, it's a "destroying method."

But for the purpose of this study, she invented a new way to radiocarbon-date. It requires only a tiny piece of the artifact to arrive at an accurate result while preserving nearly all the delicate art, teeth or tools being examined.

"I tried to develop this method because I want to destroy as little as possible," Talamo said, but "start to combine this puzzle of human evolution with a real piece of the puzzle."

Going forward, Talamo's novel technique could be employed for other fossils or artifacts. She hopes to apply it to ancient jewelry found in France and Germany, for example, but thinks it could help decode any valuable objects worth studying.

Sahra Talamo (left) as she hands her 3D representation of the ancient pendant and accompanying awl back to Polish researchers.

A future queue of such objects could include similar decorated ornaments, then move to figurines and weapons. The possibilities are endless, which Talamo hopes will prove radiocarbon dating can be done in a careful way by any team across the globe -- an important consideration, she says, because archaeology requires a grand perspective.

"You have to have a big team with different minds, different opinions, different disciplines that work together for the same goal," she stressed. "This will make it stronger -- the evidence that we're going to bring to the world."

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Darwinist: Learning to Live with Panpsychism? – Discovery Institute

Posted: at 9:45 pm

Photo: Cat's Eye Nebula, by NASA, ESA, HEIC, and Hubble Heritage Team (STScI/AURA).

Jerry Coyne,a traditional Darwinian evolutionary biologist and author ofWhy Evolution Is True,is having a hard time understanding why anyone would even consider taking panpsychism seriously. His bafflement over the growing acceptance of the idea that every living thing (or everything) is conscious to some extent may shed light on some new features of the changing science landscape.

His jumping off point is a recent three-way debate/discussion, sponsored byMindChat,between panpsychist philosopherPhilip Goff,naturalist theoretical physicistSean Carroll,and physicalist philosopherKeith Frankish,who views the mind as anillusion created bythe brain or, as Coyne puts it, a trick of the biological mind.

Coyne, as a metaphysical naturalist (nature is all there is), is quite sure that panpsychism is bunk and that Carrollwonthe debate:

I watched only until an hour and 45 minutes in, so I cant tell you what happens in the rest of the discussion. But if you watch up to that point, and listen to Seans eloquent and patient explanations, and see the sweating panpsychist professor try to prop up his crumbling ideas, you will not be any more enamored with panpsychism than you were before. In other words, youll see that its a theory without substance.

But speaking of theories without substance, a commenter, Maximillian,writes to notethat Carroll is a proponent of themultiverse,itself acontestedidea, deemed unfalsifiable. The commenter also points out a fact that Coyne appears reluctant to take into consideration:

Maximillian:Integrated information theory (which strictly speaking is not panpsychist in Goffs sense) is currently a leading contender for a theory of consciousness. According to IIT, consciousness is what it feels like when information is processed. If that turns out to be the case, then it is within the realm of possibility that the arrangement of physical matter in patterns unlike the biological brain might lend conscious experience to other entities that current scientific theories do not comprehend. In fact, if IIT is correct, physical fields could in principle be conscious. But that is a big if.

Yes, thats right.Integrated Information Theory(IIT)isa leading contender for a theory of consciousness. Would it likely be a leading theory if cranks, kooks, and charlatans were driving the bus?

Read the rest at Mind Matters News, published by Discovery Institutes Bradley Center for Natural and Artificial Intelligence.

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Stephen Lewis: The continuing evolution of ‘savage’ | News – Traverse City Record Eagle

Posted: at 9:45 pm

My last column on the changing meaning of savage from the 17th century to now invites further development. The word took on a central place in the late 18th on into the 19th century when Romanticism emerged in Europe and here.

The first step in that transformation can be found in the words of the ever-sensible Benjamin Franklin. Writing out of his direct experience dealing with the Indigenous population during the French and Indian War, Franklin declared, Savages we call them, because their manners differ from ours, which we think the perfection of civility; they think the same of theirs.

Romanticism pushed savage from Franklins balanced assessment to a preferred status by positing that human nature is essentially good. Therefore, all the clearly morally reprehensible acts we see daily result from societys corruption of that goodness. Tracking back to the words root as meaning uncivilized, savage individuals were deemed to be noble, because they had not been corrupted by society. The exact origin of this phrase is often ascribed to the French philosopher Jean-Jacque Rosseau, a contemporary of Franklin, who without using the term itself did praise the virtue of the uncorrupted savage. Later usage elevated uncorrupted to noble.

English Romantic poet William Wordsworth opens a famous sonnet by declaring, The world is too much with us late and soon/Getting and spending, we lay waste our powers and in the process lose contact with the essential goodness of nature. The more civilized we become, that is, the more removed we are from this essential goodness, the more corrupted we are.

All of the above is essentially a European concept. American Romantics, such as Emerson and Thoreau generally subscribed to the view of nature as essentially good, the motivation for Thoreau to retreat into the woods surrounding Walden Pond, but real-life exposure to the ongoing conflicts with the Indigenous population as settlers moved across the continent provided a different perspective. One form of this perspective was to respect the pre-contact Indigenous population as an evolutionary step toward a superior white civilization. Such a view is expressed in Henry Wadsworth Longfellows The Song of Hiawatha, which offers a noble savage existence that gives way to the reality of the presence of a superior culture carried here from Europe. Ralph Waldo Emerson protested the removal of the Cherokees but argued that the tribe could raise itself up by becoming more civilized, that is, whiter.

James Fennimore Cooper in his Leatherstocking Tales, novels set in the then-frontier of western upstate New York, describes his main character Natty Bumppo as a being removed from the every-day inducements to err which abound in civilized life, who sees God in the forest a being who finds the impress of the Deity in all the works of nature without the blots produced by the expedients, and passions, and mistakes of man. In short, a man of nature.

Coopers view of the tribes is an interesting attempt at balance. Some of his critics chastised him for presenting the tribes in a too-positive light. That criticism seems to be an indicator of the move toward seeing the tribes as savage Indians, a view that soon dominated their representation in the American imagination. Cooper invented the term of gifts, attributes he associated with each culture, that is, white gifts and red man gifts.

For Cooper, his Natty Bumppo represented the best resolution, one who retained his superior white gifts while acknowledging, and learning from, the red mans gifts. That resolution, of course, did not take hold, giving way to the perceived necessity of cleansing the tribes gifts and replacing them with those of the whites settlers moving across the continent.

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