Revisiting Wilkinson 1984

In 1984, Gerald Wilkinson published a paper in Nature showing that vampire bats share food in the form of regurgitated blood, within groups that contain both kin and non-kin. This was one of the fi…

Source: Revisiting Wilkinson 1984

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Summer interns for the vampire bat project

Every season, two interns will be assisting the vampire bat food-sharing project at the Smithsonian Tropical Research Institute in Gamboa, Panama. These are our two STRI-funded interns for Summer 2016.

Emily Dong is a major in the Biology and Society, and will be starting her third year at Cornell (my alma mater). Emily is always positive, excited, and enthusiastic. Ever curious, she seems to absorb information like a sponge. She is linking feeding interactions between vampire bats with grooming and food-sharing, and testing whether specific bats follow each other to feeders.

What are your interests?

Beyond scrolling through socialbat.org, my interests revolve around examining relationships, especially friendships that occur across animals, whether it be humans or vampires. I’m intrigued by cooperative bonds, the behaviors that enable (or disable) social bonds, and how specific bonds have become evolutionarily persistent. I also like stories! A lot! Storytelling, from historical narratives to dinner table conversations, is a powerful way to share information and create (or maintain) social order.

What do you hope to gain from working on the vampire bat project?

I’m excited to spend quality time with vampire bats and observe the colony to the point of knowing specific bats’ unique habits. Besides working closely with bats, I’m hoping to see first-hand what the research life entails. (And I’m stoked to hang out with Gerry and other cool bat people, so that I absorb all their knowledge and coolness). 

What are your plans for the future?

My future holds many social bonds and much reciprocity, but whether I’ll partake in them, study them, or both is still undecided!

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Yelitza Garcia is entering her final year at Earlham College driven by a passion for science and research that she has fostered since childhood. As a first-generation college student, she became interested in science after entering a science fair at the age of eight. Yeli is highly-motivated to get research experience and she plans to study animal behavior, evolution or ecology. Armed with valuable combination of being highly-motivated and ambitious without a drop of overconfidence or selfishness. She is working on the sensory basis of roost-finding in vampire bats.

What are your interests?

Like most young, starry-eyed, field ecologists, I have a deep-set admiration for being outdoors and learning about the world around me. I am primarily interested in behavioral ecology and conservation biology, but love to learn and read about vertebrate evolution and bioethics in my spare time. Other than reading, I love spending time outdoors hiking, climbing, and birding, and cooking for my loved ones.

What do you hope to gain from working on the vampire bat project?

In addition to interacting on a daily basis with these adorable flying furballs, I hope to learn as much as I can from this project about research as a career, behavioral ecology, and tropical communities. This is the first opportunity I have to do research full-time, so I want to learn about and contribute to meaningful discussions about reciprocity, sensory ecology and more. I have never felt as grateful or lucky as I am now that I get to wake up every day to do research and discover more about animal behavior.

What are your plans for the future?

After graduating from Earlham College this coming spring, I hope to attend graduate school and work towards a masters, and hopefully a doctoral degree in Ecology and Evolutionary Biology. Through the rest of my education and after, I hope to continue research and conservation work in the tropics. I am incredibly grateful for the opportunity to work on this project because this is the work I plan to dedicate my life to.

emilyyeli

Figure 1. To test the effects of association, we began housing unfamiliar female humans Emily (left) and Yeli (right) together in close proximity within the same roost, and we have already observed cooperative behavior, including huddling and food sharing (shown above).

 

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Social inheritance in vampire food-sharing networks?

We are soon to be wrapping up several analyses and starting some new ones. I want to mention one analysis that never really got off the ground, but it’s a good idea. My intern Jana asked me a great question: Does a female vampire bat inherit some of her food-sharing partners from her mother?

This question has some really interesting theoretical work behind it. I looked into my PhD data a bit, but unfortunately, I don’t have the necessary sample size. Food sharing most often occurs among females, and I have focused my data collecting on females, but there were only 4 females born during my PhD study. And I have only poor data for the 15 males born during my study (I’ll say more about that sex ratio bias in another blogpost).

Anyhow, to look into Jana’s question, I just now measured the average donation rate to bat A from all 37 possible donors, then compared that metric for bat A’s mom. If you just look at whether bats have more similar sharing networks to their moms versus all other bats, you find that females (n=4) do, while the males (n=15) do not. But this does not prove anything. We should expect that females should have more similar connections to all other females just because females are more similar to females in general when it comes to food-sharing. So this might have nothing to do with maternal bonds. What we really want to know is: Is the sharing network of female bat A more similar to bat A’s mother than to the mothers of bats B, C, or D?

Answer: Nope. That was only true in one case. Under perfect social inheritance, the rankings of similarity of the four bats to their own mom should have been all 1st place (out of 4). Instead the rankings were 4th (last), 4th (last), 1st, and 3rd. Clearly, it’s too few observations to draw conclusions, but there’s nothing very striking here.

As we collect more data from more bats, it will be interesting to do a more powerful comparison of the sharing networks of mothers and their adult daughters. With more data, we can also ask whether adult bats with many sharing partners have adult offspring with many sharing partners.

In the next few months, we hope to be looking at how bats form new food-sharing relationships with strangers. And we also have three more new pups.

Here’s a picture of the Rachel Page Bat Lab/Family in Gamboa:

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And here’s a preliminary association network of female (red) and male (blue) frog-eating bats (Trachops cirrhosus) based on 4 years of roost capture data (more on that coming soon!).

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Finally, some results of some recent cool papers:

Are there different cooperative social structure types? Or do animals socialize along a gradient? There are types! “Using phylogenetically informed comparative analyses, we found strong evidence indicating that not all reproductive arrangements within social groups are viable in nature and that in societies with multiple reproductives, selection favours instead taxon-specific patterns of decrease in the proportion of breeders as a function of group size.”

Do bacteria within your own gut cooperate with each other? Yes: “Using in vitro systems and gnotobiotic mouse colonization models, we find that extracellular digestion of inulin increases the fitness of B. ovatus owing to reciprocal benefits when it feeds other gut species such as Bacteroides vulgatus. This is a rare example of naturally-evolved cooperation between microbial species.”

Do individuals choose to cooperate based on expected payoffs?  “We experimentally created a situation of high conflict in communally nursing house mice, by using a genetic tool to create a difference in birth litter sizes. Females in the high conflict situation (unequal litter sizes at birth) showed a reduced propensity to give birth as part of a communal nest, therefore adjusting their cooperativeness to the circumstances.”

Do individuals pay attention and change their behavior depending on their own dominance status relative to that of others? “In this study, it is shown that male mice form linear dominance hierarchies characterized by individuals attacking in bursts. Temporal pairwise-correlation analysis reveals that non-dominant individuals avoid behaving aggressively concurrently with an aggressively behaving alpha male. This anti-correlation is only found with alpha males and is greater for more despotic alpha males. It is concluded that less dominant individuals modulate their aggressive behaviour in response to their social context, resulting in an attentional group structure.”

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New paper on vampire bat communication

Our newest paper is Common vampire bat contact calls attract past food-sharing partners in the journal Animal Behaviour. You can download the paper for free until June 12, 2016 here at this link: http://authors.elsevier.com/a/1SwLKmjLdkSa

It’s a simple playback experiment where we disentangled kinship and food sharing as predictors of a bat’s attraction to calls of different individuals. Subject bats chose between moving towards and spending time near two ultrasonic speakers pretending to be different bats. With the simulated callers were paired by kinship, we found that bats were biased to callers that had fed them more. But when callers were paired by sharing history, bats were not biased towards closer kin. The playback responses suggest that the vampire bats vocally recognized individuals, and this is a further illustration of how food sharing history can overshadow kinship in determining social bonds and behavior.

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Non-maternal allogrooming of pups

We have four new vampire bats. The bats here at the field station have been breeding in captivity, which is a good sign that they are doing well, and it ensures we have some highly related dyads for our experiments. My first intern, Jana, just took this neat video of a mother and her new pup being groomed by another female. I don’t know yet if this is a relative of the mom, but I will after I genotype everyone. This non-maternal (even non-kin) allogrooming of young pups is not uncommon in vampire bats, and I wonder if there’s an analogy to the common phenomenon of infant handling in primates.  I sped up the video except for some of the allogrooming.

The pups are born with open eyes and the ability to scramble around pretty well. The gestation period is about 7 months–pretty long! Pups grow quickly from about ~6 grams at birth to ~12 g in about 3 weeks, and ~24 g in 3 months. But the pups start out with big, almost fully-grown, feet! Adorable.

Pups also depend on their mothers for a longer period than other bat species. And for female pups, maternal care extends into an enduring mother-daughter relationship that can last many years.

In captivity, the mothers will carry their pups around while nursing them for 2 months, when the pups weigh roughly half their mother’s weight. At 4 months, the pups are flying around and drinking blood, and I’ve seen a 5-month-old juvenile regurgitate food to her mother!

Mothers stop providing milk to pups at about 10 months, after they have gradually switched them from milk to regurgitated blood. Wilkinson observed a mother regurgitating blood to her pup within minutes of its birth, which may inoculate the pup’s intestinal microbiome.  Pups are fed with regurgitated blood primarily by their mother, but increasingly also by other groupmates, especially maternal kin. Both Uwe Schmidt and I have even seen vampire bats fed by related males, such as a father or older half-brother.

Allonursing happens in captivity, although Jerry Wilkinson never observed it during his 400 hours of observation in the wild. Uwe Schmidt’s lab observed an orphaned vampire pup that was adopted by a non-lactating female. After a few days, the foster-mother began lactating and raised the adopted bat successfully [1]!

Female bats become sexually mature after their first year [2], and there is no strict reproductive season. A female can produce a new offspring about every 10 months, and individuals can live to be more than 30 years old in captivity [3]. There is even a record of a 37-year-old captive vampire bat [4]. In the wild, there are records of bats surviving at least 15 years [5], and some weaker dental evidence suggesting 18-year longevity [6]. This suggests that a female vampire bat could have have more than 20 descendants in her lifetime despite such an extreme life history strategy of high investment and slow reproduction.

Interestingly, at the captive colony I studied at Organization for Bat Conservation in Michigan, several females had very high reproductive success, while other females had no pups at all during the same 4-year-span. Given the huge geographic range of vampire bats, and given that these low-fitness females tend to come from different source populations, one explanation is that the males and females may have been sourced from wild populations that are too genetically distant.

——–

References

  1. Schmidt, Christel. 1988. Reproduction. In: Natural History of Vampire Bats, edited by Greenhall, AM and Schmidt U. CRC Press. (And refs therein)
  2. Wilkinson, G. 1985. The social organization of vampire bats I and II. Behavioral Ecology and Sociobiology.
  3. Me. 2012. the oldest vampire bat. socialbat.org https://socialbat.org/2012/08/22/the-oldest-vampire-bat/ (and refs therein)
  4. On Oct 1, 2013, Guy Lichty, Curator of Mammals at North Carolina Zoological Park told me that a male vampire bat (#1272) was still alive and “records indicate that he was born 2 March 1976, which means today he is 37 years, 6 months and 29 days old.”
  5. Lord, R. D., Muradali, F., Lazaro, L., 1976. Age composition of vampire bats (Desmodus rotundus) in northern Argentina and southern Brazil. Journal of Mammology. 57, 573-576.
  6. Tschapka, M., Wilkinson, G.S. 1999. Free-ranging vampire bats (Desmodus rotundus, Phyllostomidae) survive 15 years in the wild. Z. Säugetierkunde, 64, 239-240.

 

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Q & A with our new interns

Every season, two volunteer interns will be assisting the vampire bat food-sharing project at the Smithsonian Tropical Research Institute in Gamboa, Panama. These are our first two interns for Spring 2016.

jana

A month ago, eighteen-year-old whiz kid Jana Nowatzki (left) joined our project from Konstanz, Germany. Jana is a surprisingly self-motivated, bright, positive, and precocious student. Armed with an infectious enthusiasm and a curious mind, Jana has undertaken a project that involves focal sampling of bats to construct a social grooming network. While learning how to read scientific papers, she is also helping to translate some of the German literature on vampire bat research into English.

 

What are your interests?

I finished high school in June this year. Of course, I am happy with new possibilities for life experiences. That is why I really enjoy traveling, which allows me a closer look at the way of life in my country compared to other countries. I am really interested in the interface of human beings with their natural environment, the different attitudes of dealing with nature, what influences them, and how they arise. I am absolutely impressed by nature, ecosystems, and evolution.

What do you hope to gain from working on the vampire bat project?

I think my main idea is getting an impression of “the real working life” of research, which is hard to miss in Gamboa. Also, I wish to gain a basic understanding of how animal social structures work, why and how individuals act differently. Finally, I want to learn more about these strange little flying fur balls, and their amazing psychology and physiology.

What are your plans for the future?

I want to study biology. I’ve learned that plans change constantly, and at the moment it is hard to separate plans from dreams.

 

photoNext month, Yesenia Valverde (left) will be join our project from Brown University, where she is pursuing a degree in Conservation Science & Policy with a focus on tropical rainforests. After her volunteer work with vampire bats in Panama, she will serve as a research assistant at the Monteverde Cloud Forest Reserve in Costa Rica studying the ecology of epiphytes in order to predict their vulnerability to climate change.

 

What are your interests?

My entire life, I’ve grown up visiting family in Costa Rica and have always felt myself attracted to tropical rainforests. As I began to learn about current environmental issues, my admiration of nature evolved into a passion as I dedicated myself to learning more in order to join the effort to protect it. I’m especially intrigued by wildlife ecology and love to learn about and see new animals out in the field. When I’m not trying to save the rainforest, I really enjoy spending time with family and am currently really into improving my cooking skills!

What do you hope to gain from working on the vampire bat project?

I’m hoping to walk away from this experience with a lot of new knowledge about bats in general. Despite their bad reputation, bats are incredible animals that are vital to, among other things, the proper ecosystem functioning of tropical systems. I hope to gain valuable research experience during this time, especially learning how to mist net. I plan to mist net birds in Costa Rica for my senior thesis project later this year. I’m excited to further my development as a scientist as well by designing and carrying out behavioral experiments in order to answer fundamental questions.

What are your plans for the future?

After graduation, I plan on seeking field positions in wildlife research as part of my never-ending quest to gain more research experience. I want to explore different opportunities so that I can be sure of what it is that I want for the rest of my career. After that, I plan to go to grad school and pursue a PhD in Conservation Biology.

 

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An essay about caves and the origins of echolocation

Caves and the origins of echolocation

Imagine that you are in a cave, a very familiar cave, but with no light. Do you think you could collect information about your location by shouting or clapping and listening to the echoes? Would a large chamber sound different than a tight passage? Sound ridiculous? Try this experiment. Make a loud constant “shhh” sound as if telling someone to be quiet. Now close your eyes and move your hand back and forth in front of your face as if waving. Hear that? What you are now doing is very simple version of biosonar, or echolocation.

Bats do it way better. Most cavers know that bats use echolocation– a kind of sonar. But perhaps it’s demeaning to call what bats are doing “sonar” (an acronym for the human technology of SOund Navigation And Ranging), because comparing a bat’s echolocation to our military sonar is like comparing human vision to an earthworm avoiding light. Human-engineered sonar can detect submarines, but echolocating bats can detect miniscule flying insects, minnow fins protruding from a water surface, the tiny water ripples made by a calling frog, even the gossamer strands of spider webs. How do they do it?

Most echolocating bats produce a very loud chirp that sweeps across a precise range of frequencies. And when I say loud, I mean loud— louder than a smoke alarm or a circular power saw. Some bats actually disengage their ear bones to avoid deafening themselves. But the pulse is so brief and so high-pitch, your ears cannot perceive it. When the echoes reflect off an object and come back to the bat, she can form an image of what’s in front of her, whether it’s a cave wall or a moth. Note that a cave wall, especially a large flat surface, is much easier to detect than a moth, which is one reason why I think bat echolocation first evolved in caves.
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Right: A bat flies through the dark of a cave with its mouth open. Photo by Brock Fenton.

In many ways, a cave is the perfect environment for a primitive form of echolocation to get started. Many animals take refuge in caves to avoid predation. If you’re a bat from the Eocene, caves are pretty safe and environmentally stable places to, literally, hang out. The problem is that deep inside where it’s safest, it’s also absolutely dark. But solve that problem and you can make a living deeper and deeper in a cave. The hard solid walls of some caves can produce crisp clear echoes compared to a cluttered leafy forest or open air. Actually, the easiest environment for echolocating would be inside a building with flat smooth solid walls. But alas, caves are the closest thing that Mother Nature gave bats. There are only a few natural environments where vision is absolutely useless, and the two that most easily come to mind are underground and under deep or murky water. And echolocation evolved independently in both these places. Bats and toothed whales both “invented” sonar in their own way.

Cave-dwelling animals have also independently evolved the ability to echolocate several times. This happened at least twice in two different groups of birds, oilbirds and swiflets, which both live in caves (Brinklov et al. 2013 Frontiers in Physiology). And as you’ll read below, it happened at least three times in cave-dwelling bats.

Do bats see with their ears?

The story of how people discovered bat echolocation is actually quite interesting. In 1793, an Italian scientist named Spallanzani found that flying owls would crash into obstacles in absolute darkness and they needed a bit of candlelight to avoid them. Bats on the other hand could avoid the same obstacles even in pitch-black darkness. He then did a rather unsavory experiment. He poked out the bats’ eyes and found that they could still somehow magically “see”. Further experiments showed that by plugging their ears, the bats could no longer orient and would crash into objects in total darkness. Spallanzani’s conclusion was simply that bats… somehow… see with their ears. Of course, this seemed ridiculous.

The puzzle was finally solved more than a hundred years later in 1939-1941 by a young biologist named Donald Griffin who realized, using fancy new microphones, that flying bats were making high-frequency ultrasound beyond the range of what a human was capable of hearing. So the story goes that when Griffin first presented his results at a scientific conference, his fellow scientists thought he was crazy (just like Spallanzani). In Griffin’s words,

“One distinguished scientist was so indignantly incredulous that he seized [my colleague] by the shoulders and shook him while complaining that we could not possibly mean such an outrageous suggestion. Radar and sonar were still highly classified developments in military technology, and the notion that bats might do anything even remotely analogous to the latest triumphs of electronic engineering struck most people as not only implausible but emotionally repugnant.” (quote from p. 35, Dawkins 1986 The Blind Watchmaker)

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We now know that there are more than 1,300 bat species, and most of them echolocate. Different bats echolocate in very different ways. Most echolocating bats produce the echolocation sounds with their larynx, just like when you speak or sing. The local bats of the Eastern USA all scream loudly through their tiny mouths, but some bats in other parts of the world also send the signals out through their nose– which is one reason why their faces are often so weird-looking. To see what I mean, just do a Google image search for “bat noseleaf”.

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Left: A bat that echolocates through its nose leaf

Natural selection creates two kinds of bat echolocation

This laryngeal echolocation later evolved into two distinct forms. The original “design” was to produce a series of brief pulses (each about 1/500th of second) separated by much longer pauses. For the bat, this is a bit like a strobe light, except the bat uses the timing of echoes to determine distances. So imagine a strobe light where you see near surfaces in first, then farther surfaces second, than even farther surfaces third and so on. (Now, for all I know, the bats might compile all these images into a single movie image, just like your brain puts together the illusion of a large visual field from a mosaic of your many focused eye movements.)

Some bats evolved a completely different kind of laryngeal echolocation, where they emit a long signal and then use Doppler shift to detect objects. That is, they use changes in frequency, not time, to detect objects and how they are moving. This is similar to how an ambulance that is approaching you sounds different than one that is driving farther away. This kind of bat echolocation is better for longer distances and hunting insects in open environments. Amazingly, it evolved twice independently in two completely unrelated groups of bats. (This is, by the way, not that strange: two different groups of bats also independently “invented” suction cups on their thumbs to cling to smooth surfaces. Google it.).

Anyhow, that’s all laryngeal echolocation. Laryngeal echolocation has become astonishingly sophisticated due to an evolutionary arms race with bat-detecting moths. But that’s a whole other story. Our story continues in a different direction. One group of bats, the Old World fruit bats, stopped living in caves, started hanging in tree branches, developed great vision, and started eating fruit. No one knows in which order those things happened. But they eventually developed big cute eyes and they are often called “flying foxes” because that’s what they look like. In fact, they look and act so different from other bats that biologists once classified them as something else entirely. (It’s possible that the ancestors of flying foxes never had echolocation and that their close relatives evolved it independently, but I think the more simple explanation is that the ancestors of flying foxes simply lost the ability to echolocate. That’s a common theme in evolution: use it or lose it.)

Echolocation by tongue

Ok, now comes a plot twist: one group of these non-echolocating flying foxes eventually moved from trees back into caves and re-evolved the ability to echolocate in a completely different fashion!

Instead of using their larynx, they click with their tongues (like those cool clicking languages used by some human tribes in southern Africa). Researchers use to think that tongue-clicking echolocation in bats was pretty unsophisticated, but work by the biologist Yossi Yovel showed that the bats actually steer and adjust their sonar beams in complex ways much like the more advanced laryngeal echolocators (Yovel et al 2010 Science, 2011 PLOS Biology).

There are some small flying foxes, like the flower-feeding Dawn bat, that don’t seem to echolocate at all, even though they live in caves. Or so we thought.

Echolocation by wing

In 1988, a biologist named Edwin Gould, who was mentored by Donald Griffin (the guy who first discovered echolocation), observed Dawn bats leaving caves in Malaysia and noticed that they made clicking sounds when in total darkness deep inside the caves but not near the lighted entrances. He did some experiments and concluded it was not tongue-clicking but wing-clicking. He then put paint on one wrist of some bats and found that it was transferred to the other wrist, so he hypothesized that the bats were actually clapping their wings together (Gould, 1988, Journal of Mammalogy). Nobody followed up on this work for 25 years, and at least some bat biologists thought this idea was a little crazy (sound familiar?).

temp4Left: A Dawn Bat. Photo by Alyssa Stewart.

Arjan Boonman, a postdoctoral researcher working with Yossi Yovel, now a professor at Tel Aviv University, followed up on Gould’s work and found that yes indeed—the Dawn bat does echolocate with its wings, along with two other supposedly non-echolocating fruit bats they tested (Boonman et al 2014 Current Biology and comment). The team showed that these bats were not actually clapping their wings to produce the sound, but they failed to figure out exactly how the clicks were created. So that’s one puzzle for the future.

But the team did clearly show that this rudimentary form of echolocation is pretty primitive. That is, it’s not great. The bats could detect and land on a flat solid surface that was one meter squared, but they struggled to detect smaller objects and they helplessly crashed into thin wires.

So, in summary, to navigate the pitch-black total darkness of caves, bats have evolved at least three different kinds of echolocation: wing-clicking (bad), tongue-clicking (good), and vocalizing (great).

There is one fact, however, that is a bit inconsistent with my simple story that caves created the necessity for the evolution of bat echolocation. Only the Dawn bat, one of the three wing-clicking bats that Boonman and his team discovered, was a cave dweller. It was the most frequent clicker and the best performer. But the other two species roost in foliage. What does that mean? I’m not sure. It’s still unclear how many kinds of Old Fruit bats can echolocate with their wings. One possibility is that all Old World fruit bats can echolocate with their wings, and that some of them later developed this ability further to varying degrees. Once a bat could do this well, it might be much easier to start echolocating using its mouth, because it’s already able to process the echoes. Boonman and his coauthors point out that primitive echolocation is more easily “evolve-able” than we once thought; it only requires that animals make some kind of sound (like clicks or squeaks) and then can make some sense of the echoes. So even though not all bats have sophisticated echolocation, perhaps many or all have rudimentary echolocation? After Boonman’s recent work, we may have to re-evaluate the whole evolutionary history of echolocation.

Echolocation for all (even humans)

Boonman and Yovel also pointed out that many mammals can probably be trained to echolocate. Humans are a prime example. Indeed, many blind people have learned the ability to echolocate to get around and do all sorts of useful things. Ben Underwood was a blind kid without eyes who could echolocate well enough to ride a bike and shoot a basketball into a hoop. Look it up on Youtube and have your mind blown. Daniel Kish is a blind echolocator who teaches echolocation to other blind people. He started a non-profit called World Access for the Blind and has teamed with researchers studying human echolocation who claim that in just 2 hours a day for 2 weeks a student can learn to detect large objects using tongue clicks, and in another two weeks, they can discriminate trees from walls.

So there you have it. If you ever want to have the ultimate backup to your headlamp, just remember that you can always train yourself to echolocate so you can tongue-click, hand-clap, or yelp your way through a cave without a light. But it might still take a few million years of living and evolving in the darkness before humans can reach bat-level and hear the soft echoes reflecting off a moth or a spider web.

Gerald Carter is a PhD Candidate at The University of Maryland a postdoc working with John Ratcliffe at University of Toronto and Rachel Page the Smithsonian Tropical Research Institute. He is not an expert on echolocation.

Feb 4, 2015. Written for The Speleograph, the publication of District of Columbia Grotto, a local chapter of the National Speleological Society. When I lived in Maryland, I went caving with the DC Grotto of the National Speleogical Society. I wrote an essay about caves and echolocation for their newsletter, the Speleograph, but I don’t think it was ever published. This essay came from a ‘what-if’ conversation I had with someone in a cave about humans using echolocation to navigate caves without a headlamp. I argued that it’s easier for us to hear echoes inside a building with hard flat walls, rather than say in a forest. And since caves were the only ‘indoors’ before humans existed, echolocation probably first evolved as a way for animals to live deeper in caves. So I thought I would write an article about that idea using some neat stories from bats. Now, a year later, I just found it sitting on my hard drive. 

 

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