74 Australian Geographic September . October 75

S E E I N G W I T H T H E I R E A R S

The tiniest bat and mightiest toothed whale navigate and hunt in darkness using echolocation. Even some humans have this remarkable skill.

STORY BY PETER MEREDITH ILLUSTRATIONS BY LEVENT EFE In pitch-dark ocean depths, the sperm whale uses echolocation to find its prey

and is as proficient with it as any bat.

DEEP IN THE ABANDONED mine shaft, bats were on the move. It was dusk – time for insect hunt-ing. At the mine entrance, Kyle Armstrong knew they were coming because his bat detector device was picking up their ultrasonic echolocation

calls, normally inaudible to the human ear.It was the late 1990s. Kyle was working on his zoology

doctorate at the University of Western Australia and wanted to catch some bats to collect non-lethal biopsy samples for a genetic study. The species he was focusing on was the orange leaf-nosed bat; the mine was at Bamboo Creek, in the East Pilbara, in northern WA. He’d set up a fine-mesh mist net across the entrance and was confident he’d catch a few.

The bats arrived from the depths – but refused to f ly into the net. Detecting the filmy material in front of them, they spun around, tried again and spun around again. “They did this for some minutes, backwards and forwards, and then they dis-covered a pre-existing tear in my net, and in the blink of an eye they all zipped out through the hole and were gone,” Kyle tells me.

The experience left a lasting impression. “It brought home to me how clever bats are at processing their ultrasonic echo-location signals at high speed and finding out about their envi-ronment, even something as fine as a mist net,” he says.

For Kyle, it led to a career-long fascination with bats, their echolocation abilities and calls, particularly as a tool for identi-fying species. It’s all part of a branch of science known as bioacoustics. “I’m interested in bat echolocation because it underpins so much of their ecology and evolution,” he says. “You can see bats as similar to rodents, with two extra layers of complexity: they f ly and they echolocate.”

Kyle’s bat research has taken him to many parts of Australia, to Japan’s islands and Papua New Guinea’s rainforests, as an independent zoological consultant or on behalf of institutions such as the University of Adelaide and the South Australian Museum. When I met him in early 2018, he was running a citizen science project for the museum to gather information on microbats in the Murray-Darling Basin.

ECHOLOCATION IS A biological sonar capability that ena-bles some animals to ‘see’ in murky to pitch-dark con-ditions. An echolocating animal emits specialised calls

and detects echoes of those calls ref lecting back from surround-ing objects. From those echoes the animal builds a comprehen-sive, three-dimensional mental image of the world around it.

Nature’s finest echolocators are microbats and toothed whales, which include sperm whales, orcas and dolphins. Two kinds of cave-dwelling birds – swiftlets and oilbirds – also echolocate. So do a few small mammals, such as shrews and shrew-like creatures called tenrecs, although their techniques are more basic. Increasingly, blind people are also learning to use echo-location with remarkable success (see page 82).

There are two main bat types. Megabats include large fruit-eaters and nectar-feeders, such as f lying foxes, and smaller blossom bats. Microbats mostly hunt insects, although some feed on small animals, blood and nectar. Megabats usually don’t echolocate; microbats mostly do, each species in its own way.

Zoologist Kyle Armstrong holds a bat detector, which helps him identify bat species in northern Australia and PNG from their echolocation calls.

As it closes in on a cricket, this California leaf-nosed bat is likely to be using high-speed echolocation pulses that are eff ective at fast-diminishing range.

76 Australian Geographic September . October 77

PHO

TO C

RED

ITS,

PRE

VIO

US

PAG

E: A

LEX

AN

DER

SA

FON

OV;

TH

IS P

AG

E: M

ATT

LO

XTO

N P

HO

TOG

RAPH

Y / N

EWSP

IX; O

PPO

SITE

: MER

LIN

TU

TTLE

. SC

IEN

TIFI

C N

AM

ES, P

REV

IOU

S PA

GE:

Phy

sete

r mac

roce

phal

us; O

PPO

SITE

: Mac

rotu

s ca

lifor

nicu

s

78 Australian Geographic

information on things like texture and distance. The bat will also change the rapidity of its calls; when it’s about to catch an insect, the call’s repetition rate might get up to 200 pulses per second.”

During the search phase, calls can be coupled to breathing and this, in turn, is coupled to wing beats, saving energy.

At the same time, because search-phase calls are so loud – they’re among the loudest sounds in nature – bats momentarily desensitise their hearing so that they don’t deafen themselves and miss the very soft return echoes.

Microbats beam their echolocation calls into the outside world either through their mouths or nostrils. “Some species of mouth-emitting bats have beautiful parabolic lips that open in the shape of a horn to project the call,” Kyle says.

“Nose-emitting bats may have interesting nose specialisations, complex skin structures, called a nose leaf, around the nostrils to project the signal. Every species has a different nose leaf.”

Not all bats generate echolocation calls in their larynxes; some fruit bats use low-frequency tongue clicks for navigating dark caves.

A BAT PRODUCES CALLS in the same way most mammals make sounds – by forcing air through vocal cords in the glottis, part

of the larynx. “A vibrating signal is produced at a particular

audio frequency. It has overtones or harmonics, so it’s rich in various harmonic characteristics,” explains zoologist Kyle Armstrong.

The calls are essentially squeaks or chirps sent out in bursts or pulses at up to 200 per second.

The audio frequencies of microbat echoloca-tion calls generally range from 12kHz to above 200kHz, although Kyle says the highest known in Australia is 167kHz. Because humans can’t hear sounds above 20kHz, nearly all of these calls are inaudible to us and termed ultrasound.

Bats vary their calls to suit their environment and prey. A call can sweep through a broad band of frequencies or stay at one frequency. Gener-ally, low frequencies travel further and are good over longer ranges. High frequencies are good at close range and broadband sweeps are good for discerning detail.

Echolocation calls vary in intensity (from 60 to 140 decibels), duration and pulse interval. Bats alter the characteristics of their calls according to whether they’re navigating in open sky or dense forest, searching for prey, closing in on prey or making the final approach.

On final approach, the call pulses can speed up to what’s called a terminal buzz, allowing the bat to accurately track its target’s movements at a fast-diminishing range.

“Search-phase signals are quite stereotypical and are useful for telling species apart,” Kyle says. “When a bat approaches something interesting or gets too close to a wall of vegetation, it changes the shape of the signal and that gives different

The expression ‘blind as a bat’ is a timeworn slur. Although eyesight in many microbat species isn’t great, visual deficiency is more than offset by echolocation and hearing skills, to the point where a bat on the wing on a pitch-black night can detect and catch a mosquito and perceive something as fine as a human hair.

PRODUCING SIGNALS IS ONE half of echolocation; the other is detecting and analysing returning echoes. These are so faint that not even an electronic bat detector can pick

them up. So it’s hardly surprising bats are gifted with one of nature’s most sensitive hearing systems. And tiny though its brain may be, a bat is capable of analysing vast amounts of information streaming in from its ears at mind-boggling speed.

Delays are important to this process. The nanosecond delay between the call and the echo tells how far away an object is. And, like other animals, if the bat detects an echo arriving in one ear before the other, it can fix the object’s horizontal position.

Internally, the bat’s hearing equipment is much like that of other mammals, except that in the inner ear of some species a section of the cochlea called the basilar membrane is super-sen-sitive to the frequencies of returning echoes.

Signals from the cochlea pass via auditory neurons to an organ called the inferior colliculus, in the midbrain, where ‘delay-tuned’ neurons precisely measure the gap between out-going calls and their echoes. From there, the results go to the auditory cortex, which combines the information with a mass of other data, including memorised material about the envi-ronment garnered during past f lights. Not surprisingly, the bat auditory cortex is relatively large.

With this comprehensive auditory and neural toolkit, a bat can very precisely gauge an object’s position, size, shape, tex-ture, density, movement, speed and direction. Zeroing in on a moth f lying across its path, a bat works out exactly where the insect will be in a moment’s time and aims for that spot so as to capture it with minimal use of energy. It makes these calculations in microseconds.

The image of the outside world that the bat creates in its head through echolocation may be as comprehensive and pre-cise as that created by the visual cortex of a visually guided creature. It’s a remarkable feat for such a small brain. One sci-entific study concluded: “Bats face some of the most difficult neurobiological challenges of any mammal…and solve them with apparent ease and grace.”

IN SUNLESS OCEAN DEPTHS or in murky, silt-laden river shal-lows, toothed whales can’t rely on their eyes to navigate or catch food. Instead, these marine mammals – including

T H E S E C R E T S O F B A T

E C H O L O C A T I O N

Bursts of echolocation calls produced in the larynx are projected from the nose. The nose leaf directs and shapes the signals.

1

1

2

Bat ears come in different shapes and sizes, but all are geared to picking up the often faint echoes of echolocation calls.

Nose leaf

2

2

Outer ears funnel incoming echoes to super-sensitive inner ears, from where they move to the enlarged and highly specialised auditory cortex.

3

3

September . October 79

A pitcher plant in Borneo (left ) lures bats with a specially shaped refl ector to enhance and return their echoes. The bat will roost in the pitcher during the day and leave nutrient-rich droppings for the plant. The nose leaf of echo-locating bats (below) comes in all shapes and sizes.

A bat can very precisely gauge an object’s position, size, shape, texture, density, movement, speed and direction.

PHO

TO C

RED

IT: A

VALO

N/P

HO

TOSH

OT

LIC

ENSE

/ALA

MY

STO

CK

PHO

TOSC

IEN

TIFI

C N

AM

E: R

hino

loph

us m

egap

hyllu

s

PHO

TO C

RED

ITS,

FRO

M L

EFT:

© C

HIE

N C

. LEE

; RIC

K &

NO

RA B

OW

ERS/

ALA

MY

STO

CK

PHO

TO

SCIE

NTI

FIC

NA

MES

, FRO

M L

EFT:

Ker

ivou

la h

ardw

icki

i; M

acro

tus

calif

orni

cus

80 Australian Geographic September . October 81

sperm whales, belugas, narwhals, orcas, porpoises and river dol-phins – echolocate. And they’re every bit as proficient as bats.

However, although there are many similarities in the way they do it, the underwater realm demands some very different sonar tools. Most significantly, water is more dense than air. Sound travels 4–5 times faster in water than in air, as well as much further. Dr Brian Miller, a marine mammal acoustician with the Australian Antarctic Division, says this allows toothed whales to detect prey hundreds of metres away and, in the case of the sperm whale, possibly more than 1km. Contrast this with the bat’s maximum range of 30m.

Water’s density also means echoes from echolocation calls are more stable. Prey such as fish and squid f leeing a predator move very differently from fast-f luttering insects and create their own echo patterns.

Brian says sperm whales and harbour porpoises use mainly clicks to perceive their environment and communicate. “How-ever, dolphins and killer whales, which are taxonomically dol-phins, create sounds beyond echolocation clicks,” he says. “They produce whistling and singing sounds, burst-pulse sounds half-way between a whistle and a click, and quite complex commu-nication sounds beyond that.”

The frequencies of echolocation signals of toothed whales range from three to 150 kilohertz (kHz), so humans can hear a few of them. Like bats, they modify their sonar signals as they detect and approach prey. They adjust the sound’s intensity, auditory sensitivity and pulse rate according to the distance to the target, ending a chase with a high-repetition, low-intensity buzz. And, again like bats, they can widen the echolocation beam at close range to get a wide-angle ‘view’.

Toothed whales generate their echolocation calls in their heads, and the sound streams out through a large blob of fat called the melon that sits on the upper jawbone. This melon accounts for the bulbous shape of all toothed whales’ heads. It acts as a kind of acoustic lens; massive surrounding muscles squeeze and change its shape, organising and focusing the sound vibrations.

The vibrations are generated in the air passage directly beneath the blowhole and behind the melon. Under water, the animal keeps the blowhole closed and can circulate pressurised air through the passage, associated air sacs and what are called phonic lips, setting up sound vibrations that reverberate through surrounding tissue and bounce forward off the front of the skull. In sperm whales, the blowhole is further forward and the anat-omy is different, but the mechanism works similarly.

To detect returning echoes, toothed whales have hearing that’s as sensitive as any bat’s. But, as with sound production, the mechanism is different.

A toothed whale’s external ears are just tiny holes and play no hearing role. Instead, incoming sound is picked up by fatty deposits in the lower jaw and carried by fat-filled channels to the internalised middle and inner ears, housed in two bony capsules, called the auditory bullae, under the skull. There, they are shielded from the intense outgoing calls.

The toothed whale’s cochlea is as specialised and attuned to higher frequencies as the bat’s. The specialisations continue into the auditory cortex, ensuring that some of the biggest creatures on Earth can hear the smallest of sounds.

“Generally, on land, bigger animals hear lower frequencies. That’s a function of bigger mechanical structures being better able to pick up those frequencies,” Brian says. “But in dolphins and larger whales you can still have large hearing structures that are tuned to very high frequencies.”

That bats and toothed whales, in a beautiful case of conver-gent evolution, separately honed echolocation to the same heights of perfection may have something to do with their shared mammal ancestry. Some birds also echolocate, but are

their skills as highly developed? Brian is doubtful. “It does seem there might be something in the mammalian auditory system that makes it better suited to echolocation than the avian audi-tory system,” he says.

BIRDS WEREN’T ON DERMOT SMYTH’S mind when he visited Chillagoe Caves, west of Cairns, while doing his zoology PhD on fruit bats at James Cook University in

1979. “A friend of mine invited me on a field trip to the caves to do a bat inventory,” he said. “We set some nets 30m under-ground in total darkness and to our surprise birds f lew into them, instead of bats. That really got me intrigued.”

The birds were Australian swiftlets. “Echolocation had not been confirmed in those swiftlets at that time,” Dermot says. “It had been discussed as a possibility but this idea of them f lying in total darkness had not been proven. It was known about in other swiftlet species in Asia and the Pacific but not in the Australian species. So I dumped the fruit bats and switched to echolocation in swiftlets.”

Swiftlets are small, fast-f lying relatives of swifts that hunt insects in f light by day, especially at dawn and dusk, and roost during the night in caves or rock shelters. There are 30 species, mostly spread around southern Asia and the Pacific. Only the Australian swiftlet breeds on this continent, in north-eastern Queensland. It’s among 16 swiftlet species where individuals use echolocation to navigate in caves to find their nests among hundreds, even thousands, of others.

Dermot and two colleagues confirmed in a 1976 research paper that the Australian swiftlet uses double clicks to echo-locate. Because the clicks are of low frequency, ranging from one to 10kHz, humans can hear them. The clicks speed up when the bird approaches an obstacle, enters its cave or is close to its nest.

The swiftlet produces the clicks in its syrinx, which corre-sponds to the larynx of mammals. This organ doesn’t seem to be specialised for producing echolocation-specific calls, nor are the ears or brain adapted for receiving or analysing them.

T H E S E C R E T S O F S P E R M W H A L E E C H O L O C A T I O N

A pair of phonic lips generates the initial echolocation clicks. These travel backwards through the wax-filled spermaceti organ to the skull.

Phonic lips

Echoes are detected by fatty structures around the lower jaw and carried by fat-filled channels to the middle and inner ears. From there they travel to the auditory cortex.

The melon (fatty tissue beneath the spermaceti organ) focuses the outgoing signals.

The signals hit the front of the skull and bounce forward off it. The skull’s dished shape is specially adapted for this task.

1

1

1

3

3

3

2

2

4

4

4

THE SPERM WHALE’S echolocation mechanism is every bit as specialised and efficient as the bat’s. A big difference is in hearing. A

whale’s ears are just tiny holes that don’t hear. Instead a fatty structure in the lower jawbone detects sounds.

Sound waves travel further and faster under water than through the air. This allows a sperm whale’s echolocation signals to detect prey as far away as a kilometre.

Marine mammal acoustician Dr Brian Miller launches a sonobuoy, which will monitor whale calls in the Southern Ocean, where

he’s studying blue whale population numbers.

2

PHO

TO C

RED

ITS,

FRO

M L

EFT:

AU

STRA

LIA

N M

ARI

NE

MA

MM

AL

CEN

TRE;

AVA

LON

/PH

OTO

SHO

T LI

CEN

SE/A

LAM

Y ST

OC

K PH

OTO

SC

IEN

TIFI

C N

AM

E: P

hyse

ter m

acro

ceph

alus

82 Australian Geographic September . October 83

In a 1983 study, Dermot and a colleague found Australian swiftlets weren’t able to detect objects smaller than 1–2cm wide. “Echolocation in swiftlets is less sophisticated than in bats, but then it’s not doing the same job. It’s not used for hunting insects; it’s an orientation system,” Dermot says.

Dr Mike Tarburton, a Melbourne-based ornithologist who has been researching and writing about swiftlets for more than 30 years, says swiftlet echolocation may not be on a par with bat echolocation but is very good nonetheless.

A double click gives more information than a single click, Mike says. “It can show whether an object is stationary or moving, whether it’s a rock wall or another swiftlet, or a pred-ator like an owl. It’s good enough to enable the birds to avoid each other in the air or in a cave.”

As well, the Australian swiftlet’s eyesight is exceptionally good. “Their eyes occupy 50 per cent of the skull, more than normal for birds. That shows the eyes are very important. In fact, a lot of the birds don’t echolocate in the entrances to caves they’re familiar with because they can see and also they have a good memory,” Mike says.

Only one other kind of bird echolocates – the oilbird of northern South America, a relative of nightjars. Like them, it’s

nocturnal but, unlike nightjars, the oilbird feeds on fruit rather than insects. It roosts in caves by day and emerges at dusk to f ly to its feeding grounds, which may be up to 120km away. It uses bursts of clicks generated in its syrinx, usually in the frequency range of 1–15kHz, to navigate in the darkness of caves, but relies on its sharp eyesight and sense of smell when f lying in the open and searching for food. Like the swiftlet, the oilbird has no echolocation specialisations.

The oilbird has a wingspan of nearly 1m and is skilled at hovering and twisting in f light. Its common name derives from the fact that young birds carry a lot of fat and Indigenous peo-ple would boil them down for their oil, which they used in food and lamps. The first European to see oilbirds was the Prussian geographer and naturalist Alexander von Humboldt. While exploring what is now Venezuela in 1799, he visited a cave that was home to as many as 18,000 of them.

Some 172 years later, I visited that same cave with my father, a dedicated birder. My journal entry for 15 August 1971 tells how a guide carrying a gas lamp led us deep into the cave, to an enormous chamber where thousands of birds f lew around us, “cackling, twittering, screeching and clicking… More and more came into view. Some were f lying, some sat on the rocks above us… I felt things dropping on my head”.

My dad was spellbound, but I couldn’t share his passion: at that moment, my mind was concentrating not on all that raucous life in the gloom around us but on the precise nature of what was falling on my head.

H U M A N S W H O E C H O L O C A T E

DANIEL KISH, A 52-YEAR-OLD American, navigates by echolocation. Blind since losing his eyes to cancer at 13 months, he

taught himself to use tongue clicks to find his way around as a toddler and now cycles on busy roads, hikes in forests and climbs mountains.

In 2000 he founded World Access for the Blind, a non-profit aimed at spreading the word about human echolocation and teaching blind kids how to use it. Greg Downey, a Macquarie University anthropology professor who studies people with extraordinary skills, is a board member of the body’s Australian sister organisation.

“Echolocation in humans is a behavioural and cultural input that can change the brain,” Greg says. The biggest change in a blind person is that the part of the brain that normally deals with sight can turn to processing sound; the visual cortex begins to help the auditory cortex. Remember that in echolocating animals the auditory cortex does all the echo processing.

“The human brain is usually dominated by sight,” Greg says. “About 80 per cent of electrical activity in the brain is allocated to sight when you’re moving around. If you take that away, you free up an enormous amount of real estate to be reallocated to other tasks.”

Another major change is the treatment of echoes. Sighted humans don’t hear them well, if at all, due to the ‘prece-dence effect’. This focuses on a primary sound and discounts the echo as having no use. In blind people this effect weakens, and echoes become stronger. With training, a blind person can come to detect echoes very clearly. But, Greg says, you have to work hard at it.

However, he’s convinced most people can do it. “My observations are that it’s pretty universal,” he says. “It’s an incredi-bly exciting idea because instead of seeing blind people as disabled, we can see them as people who have a really unusual skill. A lot of folks who develop this skill say it changes them funda-mentally, giving them greater inde-pendence and freedom.”

Greg says many human echolocators don’t tongue click, preferring instead to tap their canes, snap their fingers or listen for the echo of their footsteps.

AG

Swiftlet echolocation may not be on a par with bat echolocation but is very good nonetheless.

An Australian swift let negotiates a tight passage in northern Queensland’s Chillagoe Caves, using echolocating skills. The swift let is one of only two kinds of bird known to echolocate.

1

Made redundant by blindness, the visual cortex of a blind person can restructure itself to process sound and do some of the auditory cortex’s work.

4

2

To generate echolocation signals, humans can tap their canes, click their tongues or snap their fingers.

1

3

Anthropologist Greg Downey studies the eff ects on the brain of acquiring new skills. He believes learning to echolocate can be transformative for a blind person.

Daniel Kish, who lost his sight as a toddler, is dedicated to helping blind people ‘see’ better without their eyes. He refers to human echolocation as “fl ash sonar”.

4

Unlike other echolocating creatures, a human processes incoming echoes not only with the auditory cortex (3) but also with the visual cortex (4).

3

The outgoing signals bounce off an object, return to the outer ear and move to the inner ear.

2

PHO

TO C

RED

IT: G

RAH

AM

AN

DER

SON

SCIE

NTI

FIC

NA

ME:

Aer

odra

mus

terr

aere

gina

e

PHO

TO C

RED

ITS,

FRO

M T

OP:

MA

CQ

UA

RIE

UN

IVER

SITY

; WIK

IMED

IA